Mycosphere 9(2): 271–430 (2018) www.mycosphere.org
ISSN 2077 7019
Article
Doi 10.5943/mycosphere/9/2/8
Copyright © Guizhou Academy of Agricultural Sciences
Mycosphere notes 169–224
Hyde KD1,2, Chaiwan N2, Norphanphoun C2,6, Boonmee S2, Camporesi E3,4,
Chethana KWT2,13, Dayarathne MC1,2, de Silva NI1,2,8, Dissanayake AJ2,
Ekanayaka AH2, Hongsanan S2, Huang SK1,2,6, Jayasiri SC1,2, Jayawardena RS2,
Jiang HB1,2, Karunarathna A1,2,12, Lin CG2, Liu JK7,16, Liu NG2,15,16, Lu YZ2,6,
Luo ZL2,11, Maharachchimbura SSN14, Manawasinghe IS2,13, Pem D2, Perera
RH2,16, Phukhamsakda C2, Samarakoon MC2,8, Senwanna C2,12, Shang QJ2,
Tennakoon DS1,2,17, Thambugala KM2, Tibpromma, S2, Wanasinghe DN1,2, Xiao
YP2,6, Yang J2,16, Zeng XY2,6, Zhang JF2,15, Zhang SN2,12,16, Bulgakov TS18, Bhat
DJ20, Cheewangkoon R12, Goh TK17, Jones EBG21, Kang JC6, Jeewon R19, Liu
ZY16, Lumyong S8,9, Kuo CH17, McKenzie EHC10, Wen TC6, Yan JY13, Zhao Q2
1
Key Laboratory for Plant Biodiversity and Biogeography of East Asia (KLPB), Kunming Institute of Botany, Chinese
Academy of Science, Kunming 650201, Yunnan, P.R. China
2
Center of Excellence in Fungal Research, Mae Fah Luang University, Chiang Rai 57100, Thailand
3
A.M.B. Gruppo Micologico Forlivese ‘‘Antonio Cicognani’’, Via Roma 18, Forlı`, Italy
4
A.M.B. Circolo Micologico ‘‘Giovanni Carini’’, C.P. 314, Brescia, Italy
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
Engineering and Research Center for Southwest Bio-Pharmaceutical Resources of national education Ministry of
Education, Guizhou University, Guiyang, Guizhou Province 550025, P.R. China
7
Department of Entomology and Plant Pathology, Faculty of Agriculture, Chiang Mai University, Chiang Mai 50200,
Thailand.
8
Department of Biology, Faculty of Science, Chiang Mai University, Chiang Mai, 50200, Thailand
9
Center of Excellence in Bioresources for Agriculture, Industry and Medicine, Faculty of Science, Chiang Mai
University, Chiang Mai, 50200, Thailand
10
Landcare Research Manaaki Whenua, Private Bag 92170, Auckland, New Zealand
11
College of Agriculture & Biological Sciences, Dali University, Dali 671003, Yunnan, P.R. China
12
Department of Entomology and Plant Pathology, Faculty of Agriculture, Chiang Mai University, Chiang Mai 50200,
Thailand
13
Beijing Municipal Key Laboratory of Environmental Friendly Management on Fruits Pests in North China, Institute
of Plant and Environment Protection, Beijing Academy of Agriculture and Forestry Sciences, Beijing 100097, PR
China
14
Department of Crop Sciences, College of Agricultural and Marine Sciences, Sultan Qaboos University, P.O. Box 34,
123 Al-Khoud, Oman
15
Faculty of Agriculture, Natural Resources and Environment, Naresuan University, Phitsanulok, 65000, Thailand
16
Guizhou Key Laboratory of Agricultural Biotechnology, Guizhou Academy of Agricultural Sciences, Guiyang,
550006, P.R. China
17
Department of Plant Medicine, National Chiayi University, 300 Syuefu Road, Chiayi City 60004, Taiwan
18
Russian Research Institute of Floriculture and Subtropical Crops, 2/28 Yana Fabritsiusa Street, Sochi 354002,
Krasnodar region, Russia
19
Dept of Health Sciences, Faculty of Science, University of Mauritius, Reduit, Mauritius
20
No. 128/1-J, Azad Housing Society, Curca, Goa Velha-403108, India
21
No. 33 B St. Edwards Road Southsea Hants. PO5 3DH, UK
Hyde KD, Chaiwan N, Norphanphoun C, Boonmee S, Camporesi E, Chethana KWT, Dayarathne
MC, de Silva NI, Dissanayake AJ, Ekanayaka AH, Hongsanan S, Huang SK, Jayasiri SC,
Submitted 1 April 2018, Accepted 29 April 2018, Published 30 April 2018
Corresponding Author: K.D. Hyde – e-mail – kdhyde3@gmail.com
271
Jayawardena R, Jiang HB, Karunarathna A, Lin CG, Liu JK, Liu NG, Lu YZ, Luo ZL,
Maharachchimbura SSN, Manawasinghe IS, Pem D, Perera RH, Phukhamsakda C, Samarakoon
MC, Senwanna C, Shang QJ, Tennakoon DS, Thambugala KM, Tibpromma, S, Wanasinghe DN,
Xiao YP, Yang J, Zeng XY, Zhang JF, Zhang SN, Bulgakov TS, Bhat DJ, Cheewangkoon R, Goh
TK, Jones EBG, Kang JC, Jeewon R, Liu ZY, Lumyong S, Kuo CH, McKenzie EHC, Wen TC,
Yan JY, Zhao Q 2018 – Mycosphere notes 169–224. Mycosphere 9(2), 271–430, Doi
10.5943/mycosphere/9/2/8
Abstract
This is the fourth in a series of Mycosphere notes wherein we provide notes on various fungal
genera. In this set of notes, we introduce Phaeoseptaceae as a new family, Pseudobyssosphaeria
(Melanommataceae) as a new genus, 40 new species, 11 new host or country records, one reference
specimen, one new combination and provide a description of the holotype of Uleodothis balansiana
(Dothideaceae). The new species are Acrospermum longisporium (Acrospermaceae), Ascitendus
aquaticus (Annulatascaceae), Ascochyta clinopodiicola (Didymellaceae), Asterina magnoliae
(Asterinaceae), Barbatosphaeria aquatica (Barbatosphaeriaceae), Camarosporidiella populina
(Camarosporidiellaceae),
Chaetosphaeria
mangrovei
(Chaetosphaeriaceae),
Cytospora
predappioensis, Cytospora prunicola (Cytosporaceae), Dendryphiella phitsanulokensis
(Dictyosporiaceae), Diaporthe subcylindrospora, Diaporthe subellipicola (Diaporthaceae),
Diplodia arengae (Botryosphaeriaceae), Discosia querci (Sporocadaceae), Dyfrolomyces sinensis
(Pleurotremataceae), Gliocladiopsis aquaticus (Nectriaceae), Hysterographium didymosporum
(Pleosporomycetidae
genera,
incertae
sedis),
Kirschsteiniothelia
phoenicis
(Kirschsteiniotheliaceae), Leptogium thailandicum (Collemataceae), Lophodermium thailandicum
(Rhytismataceae), Medicopsis chiangmaiensis (Neohendersoniaceae), Neocamarosporium
phragmitis (Neocamarosporiaceae), Neodidymelliopsis negundinis (Didymellaceae), Neomassarina
pandanicola (Sporormiaceae), Neooccultibambusa pandanicola (Occultibambusaceae),
Neophaeosphaeria phragmiticola (Neophaeosphaeriaceae), Neosetophoma guiyangensis
(Phaeosphaeriaceae), Neosetophoma shoemakeri (Phaeosphaeriaceae), Neosetophoma xingrensis
(Phaeosphaeriaceae),
Ophiocordyceps
cylindrospora
(Ophiocordycipitaceae),
Otidea
pseudoformicarum (Otideaceae), Periconia elaeidis (Periconiaceae), Phaeoisaria guttulata,
Pleurotheciella krabiensis, Pleurotheciella tropica (Pleurotheciaceae), Pteridiospora bambusae
(Astrosphaeriellaceae), Phaeoseptum terricola (Phaeoseptaceae), Poaceascoma taiwanense
(Lentitheciaceae), Pseudobyssosphaeria bambusae (Melanommataceae) and Roussoella mangrovei
(Roussoellaceae). The new host records or new country records are provided for Alfaria terrestris
(Stachybotryaceae),
Arthrinium
phragmites
(Apiosporaceae),
Bertiella
ellipsoidea
(Melanommataceae), Brevicollum hyalosporum (Neohendersoniaceae), Byssosphaeria siamensis
(Melanommataceae), Cerothallia subluteoalba (Teloschistaceae), Cryptophiale hamulata
(Chaetosphaeriaceae), Didymella aliena (Didymellaceae), Epicoccum nigrum (Didymellaceae),
Periconia pseudobyssoides (Periconiaceae) and Truncatella angustata (Sporocadaceae). We
provide new molecular data for 52 species and updated phylogenetic trees for 15 orders
(Acrospermales,
Amphisphaeriales,
Annulatascales,
Asterinales,
Botryosphaeriales,
Chaetosphaeriales, Diaporthales, Dyfrolomycetales, Hypocreales, Kirschsteiniotheliales,
Peltigerales, Pleosporales, Pleurotheciales, Rhytismatales and Teloschistales) and 35 families
(Acrospermaceae, Annulatascaceae, Apiosporaceae, Asterinaceae, Astrosphaeriellaceae,
Barbatosphaeriaceae,
Botryosphaeriaceae,
Camarosporidiellaceae,
Chaetosphaeriaceae,
Collemataceae,
Cytosporaceae,
Diaporthaceae,
Dictyosporiaceae,
Didymellaceae,
Kirschsteiniotheliaceae,
Lentitheciaceae,
Melanommataceae,
Neocamarosporiaceae,
Neohendersoniaceae,
Neophaeosphaeriaceae,
Nectriaceae,
Occultibambusaceae,
Ophiocordycipitaceae,
Otideaceae,
Periconiaceae,
Phaeoseptaceae,
Phaeosphaeriaceae,
Pleurotheciaceae,
Pleurotremataceae,
Rhytismataceae,
Roussoellaceae,
Sporocadaceae,
Sporormiaceae, Stachybotryaceae and Teloschistaceae) and 45 genera (Acrospermum, Alfaria,
Arthrinium, Ascitendus, Ascochyta, Asterina, Barbatosphaeria, Bertiella, Brevicollum,
272
Byssosphaeria, Camarosporidiella, Cerothallia, Chaetosphaeria, Cryptophiale, Cytospora,
Dendryphiella, Diaporthe, Didymella, Diplodia, Discosia, Dyfrolomyces, Epicoccum,
Gliocladiopsis, Hysterographium, Kirschsteiniothelia, Leptogium, Lophodermium, Medicopsis,
Neocamarosporium, Neodidymelliopsis, Neooccultibambusa, Neomassarina, Neophaeosphaeria,
Neosetophoma, Ophiocordyceps, Otidea, Periconia, Phaeoisaria, Phaeoseptum, Pleurotheciella,
Poaceascoma, Pseudobyssosphaeria, Pteridiospora, Roussoella and Truncatella). A reference
specimen is provided for Periconia cookei (Periconiaceae). A new combination is proposed for
Seimatosporium ciliata (Sporocadaceae).
Key words – 42 new taxa – Ascomycota – Description of type species of genus –
Dothideomycetes– Lecanoromycetes – Leotiomycetes – Molecular phylogeny – New combinations
– New family – New records – New species – Pezizomycetes – Phylogenetic – Reference
specimens – Sordariomycetes – Taxonomy
Table of Contents
The numbers of taxa in this study are a continuation from previous papers (Thambugala et al.
2017, 1–50, Boonmee et al. 2017, 51–101, Jayawardena et al. 2018, 102–168) and organized as in
the “Outline of Ascomycetes” (Wijayawardene et al. 2018).
Phylum Ascomycota
Class Dothideomycetes
For recent treatments of Dothideomycetes we follow Liu et al. (2017a) and Wijayawardene et
al. (2018).
Subclass Dothideomycetidae
Dothideales Lindau
Dothideaceae Chevall.
169. Uleodothis balansiana (Sacc., Roum. & Berl.) Theiss. & Syd., Annls mycol. 13(3/4): 305
(1915), description of type species of genus
Subclass Pleosporomycetidae
Pleosporales Luttrell ex M.E. Barr
Astrosphaeriellaceae Phook. et al.
170. Pteridiospora bambusae Tennakoon, C.H. Kuo & K.D. Hyde, in Mycosphere 9(2): 281
(2018), new species
Camarosporidiellaceae Wanas. et al.
171. Camarosporidiella populina Chethana, Bulgakov & K.D. Hyde, in Mycosphere 9(2): 284
(2018), new species
Dictyosporiaceae Boonmee & K.D. Hyde
172. Dendryphiella phitsanulokensis N.G. Liu & K.D. Hyde, in Mycosphere 9(2): 287 (2018), new
species
Didymellaceae Gruyter et al.
173. Ascochyta clinopodiicola Pem, Camporesi & K.D. Hyde, in Mycosphere 9(2): 289 (2018),
new species
174. Didymella aliena (Fr.) Q. Chen & L. Cai, Stud. Mycol. 82: 173 (2015), new record
175. Epicoccum nigrum Link, Mag. Gesell. naturf. Freunde, Berlin 7: 32 (1816), new record
176. Neodidymelliopsis negundinis Manawasinghe, Bulgakov & K.D. Hyde, in Mycosphere 9(2):
293 (2018), new species
273
Lentitheciaceae Yin. Zhang et al.
177. Poaceascoma taiwanense Tennakoon, C.H Kuo & K.D. Hyde, in Mycosphere 9(2): 297
(2018), new species
Melanommataceae G. Winter
178. Bertiella ellipsoidea Ekanayaka, Q. Zhao & K.D. Hyde, Fungal Diversity 80: 79 (2016), new
record
179. Byssosphaeria siamensis Boonmee, Q. Tian & K.D. Hyde, Fungal Diversity 74: 283 (2015),
new record
180. Pseudobyssosphaeria H.B. Jiang & K.D. Hyde, in Mycosphere 9(2): 303 (2018), new genera
181. Pseudobyssosphaeria bambusae H.B. Jiang & K.D. Hyde, in Mycosphere 9(2): 306 (2018),
new species
Neocamarosporiaceae Wanas. et al.
182. Neocamarosporium phragmitis Wanas., E.B.G. Jones & K.D. Hyde, in Mycosphere 9(2):
306 (2018), new species
Neohendersoniaceae Giraldo & Crous
183. Brevicollum hyalosporum Kaz. Tanaka & Toy. Sato, Mycologia 109: 4 (2017), new host
record
184. Medicopsis chiangmaiensis Q.J. Shang & K.D. Hyde, in Mycosphere 9(2): 313 (2018), new
species
Neophaeosphaeriaceae Ariyaw. & K.D. Hyde
185. Neophaeosphaeria phragmiticola A. Karunarathna & K.D. Hyde, in Mycosphere 9(2): 316
(2018), new species
Occultibambusaceae D.Q. Dai & K.D. Hyde
186. Neooccultibambusa pandanicola Tibpromma, D.J. Bhat & K.D. Hyde, in Mycosphere 9(2):
317 (2018), new species
Periconiaceae Nann.
187. Periconia cookei E.W. Mason & M.B. Ellis, Mycol. Pap. 56: 72 (1953), reference specimen
188. Periconia elaeidis T. Sunpapao & K.D. Hyde, in Mycosphere 9(2): 323 (2018), new species
189. Periconia pseudobyssoides S. Markovskaja & A. Kačergius, Mycological Progress 13 (2):
293 (2014), new record
Phaeoseptaceae S. Boonmee, Thambugala & K.D. Hyde
190. Phaeoseptaceae S. Boonmee, Thambugala & K.D. Hyde, in Mycosphere 9(2): 323 (2018),
new family
191. Phaeoseptum terricola S. Boonmee & K.D. Hyde, in Mycosphere 9(2): 326 (2018), new
species
Phaeosphaeriaceae M.E. Barr
192. Neosetophoma guiyangensis J.F. Zhang, J.K. Liu, K.D. Hyde & Z.Y. Liu, in Mycosphere
9(2): 331 (2018), new species
193. Neosetophoma shoemakeri Senwanna, Wanas., Bulgakov, E.B.G. Jones & K.D. Hyde, in
Mycosphere 9(2): 332 (2018), new species
194. Neosetophoma xingrensis J.F. Zhang, J.K. Liu, K.D. Hyde & Z.Y. Liu, in Mycosphere 9(2):
335 (2018), new species
274
Sporormiaceae Munk
195. Neomassarina pandanicola Tibpromma & K.D. Hyde, in Mycosphere 9(2): 336 (2018), new
species
Roussoellaceae J.K. Liu et al.
196. Roussoella mangrovei C. Phukhamsakda & K.D. Hyde, in Mycosphere 9(2): 339 (2018),
new species
Pleosporomycetidae genera, incertae sedis
197. Hysterographium didymosporum S. Boonmee & K.D. Hyde, in Mycosphere 9(2): 342
(2018), new species
Dothideomycetes orders, incertae sedis
Acrospermales Minter et al.
Acrospermaceae Fuckel
198. Acrospermum longisporium Jayasiri, E.B.G. Jones & K.D. Hyde, in Mycosphere 9(2): 347
(2018), new species
Asterinales M.E. Barr ex D. Hawksw. & O.E. Erikss.
Asterinaceae Hansf. (= Lembosiaceae Hosag.)
199. Asterina magnoliae X.Y. Zeng, T.C. Wen & K.D. Hyde, in Mycosphere 9(2): 349 (2018),
new species
Botryosphaeriales C.L. Schoch et al.
Botryosphaeriaceae Theiss. & H. Syd.
200. Diplodia arengae R.H. Perera, Wanas. & K.D. Hyde, in Mycosphere 9(2): 352 (2018), new
species
Dyfrolomycetales K.L. Pang et al.
Pleurotremataceae Walt. Watson
201. Dyfrolomyces sinensis Samarak., Tennakoon & K.D. Hyde, in Mycosphere 9(2): 354 (2018),
new species
Kirschsteiniotheliales Hern.-Restr. et al.
Kirschsteiniotheliaceae Boonmee & K.D. Hyde
202. Kirschsteiniothelia phoenicis S.N. Zhang & K.D. Hyde, in Mycosphere 9(2): 357 (2018),
new species
Class Lecanoromycetes O.E. Erikss. & Winka
Subclass Acarosporomycetidae Reeb et al.
Teloschistales D. Hawksw. & O.E. Erikss.
Teloschistaceae Zahlbr.
203. Cerothallia subluteoalba (S.Y. Kondr. & Kärnefelt) Arup, Frödén & Søchting, Nordic Jl Bot.
31(1): 40 (2013), new record
Subclass Lecanoromycetidae P.M. Kirk et al. ex Miadl. et al.
Peltigerales W. Watson
Collemataceae Zenker
204. Leptogium thailandicum Ekanayaka, Zhao & K.D. Hyde, in Mycosphere 9(2): 361 (2018),
new species
275
Class Pezizomycetes O.E. Erikss. & Winka
Pezizales J. Schröt.
Otideaceae Eckblad
205. Otidea pseudoformicarum Ekanayaka, Q. Zhao and K.D. Hyde, in Mycosphere 9(2): 364
(2018), new species
Class Leotiomycetes O.E. Erikss. & Winka
Rhytismatales M.E. Barr ex Minter
Rhytismataceae Chevall.
206. Lophodermium thailandicum N.I. de Silva & K.D. Hyde, in Mycosphere 9(2): 369 (2018),
new species
Class Sordariomycetes O.E. Erikss. & Winka
Subclass Diaporthomycetidae Senan. et al.
Annulatascales D’souza et al.
Annulatascaceae S.W. Wong et al.
207. Ascitendus aquaticus Dayarathne, Fryar, K.D. Hyde, in Mycosphere 9(2): 372 (2018), new
species
Diaporthales Nannf.
Cytosporaceae Fr.
208. Cytospora predappioensis Q.J. Shang, Norphanph., E. Camporesi & K.D. Hyde, in
Mycosphere 9(2): 376 (2018), new species
209. Cytospora prunicola Norphanph., Camporesi, T.C. Wen & K.D. Hyde, in Mycosphere 9(2):
378 (2018), new species
Diaporthaceae Höhn. ex Wehm.
210. Diaporthe subellipicola S.K. Huang, T.C. Wen & K.D. Hyde, in Mycosphere 9(2): 381 (2018),
new species
211. Diaporthe subcylindrospora S.K. Huang, T.C. Wen & K.D. Hyde, in Mycosphere 9(2): 381
(2018), new species
Diaporthomycetidae families, incertae sedis
Barbatosphaeriaceae H. Zhang et al.
212. Barbatosphaeria aquatica N.G. Liu & K.D. Hyde, in Mycosphere 9(2): 384 (2018), new
species
Subclass Hypocreomycetidae O.E. Erikss. & Winka
Hypocreales Lindau
Nectriaceae Tul. & C. Tul.
213. Gliocladiopsis aquaticus Y.Z. Lu, R.H. Perera & K.D. Hyde, in Mycosphere 9(2): 387
(2018), new species
Ophiocordycipitaceae G.H. Sung et al.
214. Ophiocordyceps cylindrospora Y.P. Xiao, T.C. Wen & K.D. Hyde, in Mycosphere 9(2): 391
(2018), new species
Stachybotryaceae L. Lombard & Crous
215. Alfaria terrestris L. Lombard & Crous, in Lombard, Houbraken, Decock, Samson, Meijer,
Réblová, Groenewald & Crous, Persoonia 36: 181 (2016), new record
276
Subclass Sordariomycetidae O.E. Erikss & Winka
Chaetosphaeriales Huhndorf et al.
Chaetosphaeriaceae
216. Chaetosphaeria mangrovei Dayarathne, E.B.G. Jones & K.D. Hyde, in Mycosphere 9(2):
395 (2018), new species
217. Cryptophiale hamulata Whitton, K.D. Hyde & McKenzie, in Whitton, McKenzie & Hyde,
Fungal Diversity Res. Ser. 21: 174 (2012), new record
Subclass Savoryellomycetidae Hongsanan et al.
Pleurotheciales Réblová & Seifert
Pleurotheciaceae Réblová & Seifert
218. Phaeoisaria guttulata J. Yang & K.D. Hyde, in Mycosphere 9(2): 401 (2018), new species
219. Pleurotheciella krabiensis J. Yang & K.D. Hyde, in Mycosphere 9(2): 403 (2018), new
species
220. Pleurotheciella tropica J. Yang & K.D. Hyde, in Mycosphere 9(2): 407 (2018), new species
Subclass Xylariomycetidae O.E. Erikss & Winka
Amphisphaeriales D. Hawksw. & O.E. Erikss.
Apiosporaceae K.D. Hyde et al.
221. Arthrinium phragmites Crous, new host record
Sporocadaceae Corda
222. Discosia querci Jayasiri, E.B.G. Jones & K.D. Hyde, in Mycosphere 9(2): 410 (2018), new
species
223. Seimatosporium ciliata (Petr.) Hongsanan & K.D. Hyde, new combination
224. Truncatella angustata (Pers.) S. Hughes, Can. J. Bot. 36: 822 (1958), new host record
Taxonomic treatment
Phylum Ascomycota
Class Dothideomycetes
For recent arrangements of Dothideomycetes we follow Liu et al. (2017a) and
Wijayawardene et al. (2018).
Subclass Dothideomycetidae P.M. Kirk et al.
Dothideales Lindau
Dothideaceae Chevall.
The family is characterized by immersed to erumpent or superficial, uni to multiloculate
ascostromata, 8- or poly-spored, bitunicate asci and hyaline or brown ascospores, with transversely
septate, or muriform ascospores (Thambugala et al. 2014). The family was recently revised by
Thambugala et al. (2014). In this paper, Uleodothis is accommodated within Dothideaceae as a
sister genus to Plowrightia based on the type material of Uleodothis balansiana.
Uleodothis Theiss. & Syd.
Plowrightia was introduced in the family Dothideaceae to accommodate twelve species, with
P. ribesia as the type species (Saccardo 1883). Theissen and Sydow (1915) synonymized P.
balansiana Sacc. et al., P. rhynchosporae (Rehm) Höhn and Polystomella aphanes Rehm under a
new genus Uleodothis in Dothideales based on morphological characters. Uleodothis was originally
placed in Dothioraceae Theiss. & Syd. (Dothideales) by Stevens (1920), while Müller and von Arx
(1962) accepted this genus in Mycosphaerellaceae Lindau (Pseudosphaeriales). Subsequently, the
genus was referred to Venturiaceae E. Müll. et al. (Venturiales) by Luttrell (1973). However,
Zhang et al. (2011) excluded Uleodothis from Venturiales and treated it as genus in
277
Dothideomycetes incertae sedis based on its immersed ascomata in ascostromata, persistent
pseudoparaphyses, cylindrical asci, with 1-septate ascospores and this scheme was followed by
Wijayawardene et al. (2014, 2017a, 2018). Uleodothis is somewhat similar to Plowrightia in
having semi-immersed, pulvinate, multi-loculate, dark brown to black ascostromata and 1-septate
ascospores. However, Plowrightia has thick-walled locules developing in the upper part of stroma,
lacks pseudoparaphyses and ascospores are constricted at the septum (Barr 1987, Thambugala et al.
2014), while Uleodothis has thin-walled locules developing in the middle of stroma, persistent
pseudoparaphyses and ascospores not or slightly constricted at the septum (Theissen & Sydow
1915). Uleodothis is also similar to Aplosporella Speg. (Aplosporellaceae, Botryosphaeriales),
however, Aplosporella has a wide and irregular opening, thick-walled locules, lacks
pseudoparaphyses, usually clavate asci with short pedicels and hyaline to dark brown, uniseptate
ascospores (Liu et al. 2012, Thambugala et al. 2014, Ekanayaka et al. 2016).
Morphological similarity and the relationships between Plowrightia, Uleodothis and other
relatives are unresolved due to lack of molecular data. No DNA sequence data is available for
Uleodothis. Recent phylogenetic studies indicate that Plowrightia is a member of Dothideaceae
(Winton et al. 2007, Thambugala et al. 2014). Therefore, we place Uleodothis in Dothideaceae as a
sister genus to Plowrightia based on morphological evidence. Fresh collections and sequence data
of Plowrightia and Uleodothis-like taxa are needed to further clarify their phylogenetic placement
and relationships.
Uleodothis balansiana (Sacc., Roum. & Berl.) Theiss. & Syd., Annls mycol. 13(3/4): 305 (1915)
Fig. 1
Index Fungorum number: IF153174; Facesoffungi number: FoF04463
Epiphytes on the lower surface of leaves of Bignonia sp. Sexual morph Ascostromata
0.33−0.47 mm high × 1−2.6 mm diameter ( = 3.6 × 2.2 mm, n = 3), scattered, semi-immersed in
host tissue, pulvinate, subglobose to globose, dark brown to black, with branched, projected-ostiole,
surface verrucose. Locules 177−197 µm high × 114−162 µm diameter ( = 182 × 145 µm, n = 10),
completely immersed in the middle of the ascostromata, composed of thin, flattened cells of textura
angularis, pale brown to hyaline. Hamathecium of 2−2.5 µm wide, hyaline, aseptate,
pseudoparaphyses, embedded in a gelatinous matrix. Peridium 40−50 µm, comprising two-layers,
outer layer: yellowish-brown to dark brown, cells of textura angularis, inner layer: thin, hyaline,
flattened cells of textura angularis. Asci 92−98 × 12−15 µm ( = 96 × 13 µm, n = 10), 8-spored,
bitunicate, cylindrical to oblong, with slightly long pedicel. Ascospores 11−13 × 4−5 µm ( = 12 ×
4.5 µm, n = 20), uni to biseriate in ascus, uniseptate or swollen at the center, oval to ellipsoid,
generally straight, rounded at both ends, not constricted, hyaline, verrucose, embedded in
gelatinous matrix. Asexual morph Undetermined.
Material examined – BRAZIL, Santos, on lower surface of leaves of Bignonia
(Bignoniaceae), June 1884, J. Roumeguère no. BT8164 (BR 5020077187720, holotype).
Notes – Uleodothis balansiana was described as having uniseptate ascospores (Saccardo &
Berlese 1885). However, we re-examined the type specimen of this species and found that
ascospores are generally swollen in the middle of the ascospores, which makes them appear as 2septate. The ascospore septation of U. balansiana is unclear, thus, more specimens are needed to
check its septation and other characters (Fig. 1).
Order Pleosporales Luttrell ex M.E. Barr
Astrosphaeriellaceae Phookamsak & K.D. Hyde
The family Astrosphaeriellaceae was introduced by Phookamsak et al. (2015b) to
accommodate two genera viz. Astrosphaeriella and Pteridiospora, which have distinct trabeculae
(sensu Liew et al. 2001). Members of Astrosphaeriellaceae are characterized by large, conical,
carbonaceous ascostromata, with ruptured, reflexed, stellate, host remnants, fusiform or obclavate
ascospores and a coelomycetous asexual morph (Phookamsak et al. 2015b).
278
Figure 1 – Uleodothis balansiana (BR 5020077187720, holotype). a Habit and ascostromata on
substrate. b Section through ascostroma. c Locule. d Pseudoparaphyses. e Peridium of locules. f−g
Asci. h Ascus in Melzer’s reagent. i Projecting ostiole. j−k Ascospores. l Ascospore in Melzer’s
reagent. Scale bars: b, c = 100 μm, d, e = 20 μm, f–i = 50 μm, j–l = 10 μm.
279
Figure 2 – Phylogram generated from maximum likelihood analysis based on combined LSU,
SSU, and TEF sequenced data of Astrosphaeriellaceae. Related sequences were obtained from
Phookamsak et al. (2015b) and Wanasinghe et al. (2017a). Forty-nine strains are included in the
combined sequence analyses, which comprise 2935 characters with gaps. Lophiotrema lignicola
280
(CBS 122364) is used as the outgroup taxa. Tree topology of the ML analysis was similar to the BI.
The best scoring RAxML tree with a final likelihood value of -13717.595943 is presented. The
matrix had 912 distinct alignment patterns, with 22.65% of undetermined characters or gaps.
Estimated base frequencies were as follows; A = 0.245984, C = 0.246474, G= 0.282945, T =
0.224598; substitution rates AC = 0.896188, AG = 2.815720, AT = 0.826212, CG = 1.104916, CT
= 8.845426, GT = 1.000000; gamma distribution shape parameter α = 0.769779. Bootstrap support
values for ML equal to or greater than 60% and BYPP equal to or greater than 0.95 are given above
the nodes respectively. Newly generated sequences are in red.
Pteridiospora Penz. & Sacc.
Pteridiospora was established by Penzig and Saccardo (1897) to accommodate species
having mammiform, carbonaceous ascostromata with obclavate to ellipsoidal, or subfusoid
ascospores, with a thick, distinct, mucilaginous sheath (Penzig & Saccardo 1897, Phookamsak et al.
2014a, 2015b). Currently, seven Pteridiospora species have been recorded viz. P. chiangraiensis
Phook. & K.D. Hyde, P. chochrjakovii Hüseyın, P. curreyi (Tul. & C. Tul.) E. Müll., P. javanica
Penz. & Sacc. (Type species), P. munkii Subhedar & V.G. Rao, P. scoriadea (Fr.) Dennis and P.
spinosispora Filer (Index Fungorum 2018). In this study, we illustrate Pteridiospora bambusae as a
new species from Taiwan.
Pteridiospora bambusae Tennakoon, C.H Kuo & K.D. Hyde, sp. nov.
Fig. 3
Index Fungorum number: IF554199; Facesoffungi number: FoF04083
Etymology – the specific epithet bambusae was given after the host bamboo on which the
fungus was collected.
Holotype – MFLU 18-0071
Saprobic on Bamboo. Sexual morph Ascostromata 270–350 μm high, 380–530 μm diameter,
dark opaque, black, cone-like, gregarious, erumpent to superficial, stellate, host remnants around
the base, uni-loculate, glabrous, brittle, carbonaceous, ostiolate. Ostiole central, with pore-like
opening. Peridium 25–50 μm wide, of unequal thickness, poorly developed at the base, composed
of small dark, dark brown to black, pseudoparenchymatous cells of textura angularis, base and
corners comprising a mixture of host and fungal cells. Hamathecium composed of 1.5–2 μm wide,
filiform, trabeculate pseudoparaphyses, anastomosing at the apex, embedded in a gelatinous matrix.
Asci (140–)145–180(–190) × (13–)14–17(–18) μm ( = 168.6 × 15.5 μm, n = 25), 8-spored,
bitunicate, fissitunicate, cylindrical to cylindric-clavate, with short obtuse pedicel, apically rounded
with well-developed ocular chamber. Ascospores (31–)32–36(–38)×6–8 μm ( = 34.5 × 7.5 μm, n
= 25), overlapping, uni-to bi-seriate, initially hyaline, becoming brown when mature, subfusoid,
with rounded to acute ends, narrow towards the apex, 1-septate, constricted at the septum, upper
cell shorter than lower cell, surrounded by an irregular, distinct mucilaginous sheath, with winglike appendage extending from the lower cell. Asexual morph Undetermined.
Material examined – TAIWAN, Chiayi, Shihnong Forest, dead stems of Bamboo sp.
(Poaceae), 25 June 2017, D.S. Tennakoon, DTW 020 (MFLU 18-0071, holotype).
GenBank numbers – LSU: MG831565, SSU: MG831566, TEF1: MG833012.
Notes – The morphological characters of Pteridiospora bambusae fit in to the generic
concept of Pteridiospora in having mammiform, carbonaceous ascostromata with obclavate to
ellipsoidal, or subfusoid ascospores with a thick, distinct mucilaginous sheath (Penzig & Saccardo
1897, Phookamsak et al. 2015b). Phylogenetic analysis of combined LSU, SSU and TEF sequence
data indicates that P. bambusae is closely related to P. chiangraiensis with high bootstrap support
(92% ML, 0.97 BYPP) (Fig. 2). Pteridiospora bambusae shares some similar characters with P.
chiangraiensis and P. javanica in having erumpent to superficial, carbonaceous ascomata, short
pedicellate, cylindrical-clavate asci and sub-fusoid ascospores, with a thick, distinct mucilaginous
sheath and wing-like appendage extending from lower cell (Phookamsak et al. 2015b).
Pteridiospora bambusae differs from P. javanica, in having brown ascospores (initially hyaline),
while P. javanica has distinct hyaline ascospores. In particular, P. bambusae can be distinguished
281
from P. chiangraiensis by using ascospores characters. Ascospores of Pteridiospora bambusae
have wing-like appendages which extend from lower long cell, whereas P. chiangraiensis
appendages extend from lower, short cell. Additionally, P. bambusae has a broad mucilaginous
sheath around the upper cell (clearly visible with Indian ink), whereas P. chiangraiensis lacks a
broad mucilaginous sheath. A comparison of the 871 nucleotides across the TEF gene region of P.
bambusae and P. chiangraiensis (MFLU 11-0198) reveals 29 base pair differences (a difference of
3.3% which supports our establishment of the new taxon as recommended by Jeewon & Hyde
(2016)).
Figure 3 – Pteridiospora bambusae (MFLU 18-0071 holotype). a Ascostromata on host. b Closeup of ascostroma. c Vertical section of ascostroma. d Section of peridium. e Pseudoparaphyses. f–j
Asci. k–n Ascospores. o, p Ascospores stained in Indian ink. Scale bars: c = 75 µm, d = 50 µm, e–j
= 50 µm, k–p = 20 µm.
Camarosporidiellaceae Wanas. et al.
Camarosporidiellaceae was introduced by Wanasinghe et al. (2017b) to accommodate species
with conidial morphology resembling Camarosporium sensu stricto and other camarosporium-like
genera. The family includes a single genus Camarosporidiella. The family comprises endophytes,
plant pathogens and saprobes (Wanasinghe et al. 2017b). An updated phylogenetic tree for the
family is presented in Fig. 4 and we introduce a new species in the genus Camarosporidiella.
282
Figure 4 – Phylogenetic tree generated by maximum likelihood analysis of combined LSU, SSU and
ITS sequence data of Camarosporidiella species. Related sequences were obtained from GenBank.
Thirty-six strains are included in the analyses, which comprise 2403 characters including gaps. Tree
was rooted with Staurosphaeria rhamnicola (MFLUCC 17-0814). Tree topology of the ML analysis
was similar to the MP and BI. The best scoring RAxML tree with a final likelihood value of 4430.059713 is presented. The matrix had 223 distinct alignment patterns, with 12.01% of
undetermined characters or gaps. Estimated base frequencies were as follows; A = 0.251012, C =
0.214068, G = 0.274213, T = 0.260707, substitution rates AC = 1.402990, AG = 3.311951, AT = 1.
880669, CG = 0.441830, CT = 4.042681, GT = 1.000000; gamma distribution shape parameter α =
283
0.020013. The maximum parsimonious dataset consisted of constant 2283, 50 parsimony-informative
and 70 parsimony-uninformative characters. The parsimony analysis of the data matrix resulted two
equally most parsimonious trees with a tree length of 166 steps (CI = 0.783, RI = 0.829, RC = 0.649,
HI = 0.217) in the first tree. RAxML and maximum parsimony bootstrap support values ≥ 50% are
shown respectively near the nodes. Bayesian posterior probabilities ≥ 0.95 (PP) indicated as
thickened black branches. The scale bar indicates 0.01 changes. The ex-type strains are in bold and
new isolates in blue.
Camarosporidiella Wanas. et al.
Camarosporidiella was introduced by Wanasinghe et al. (2017b) with Camarosporidiella
caraganicola (Phukhams. et al.) Phukhams. et al. as the type species (Wanasinghe et al. 2017b). This
genus includes camarosporium-like asexual morph and cucurbitaria-like sexual morph.
Camarosporidiella populina Chethana, Bulgakov & K.D. Hyde, sp. nov.
Fig. 5
Index Fungorum number: IF553974; Facesoffungi number: FoF03871
Etymology – the specific epithet ‘populina’ was given after the host genus Populus, from
which the fungus was collected.
Holotype – MFLU 16-1722
Necrotrophic or saprobic on dead and dying branches of Populus nigra L. var. italica
Münchh. Sexual morph Undetermined. Asexual morph Conidiomata 0.2–0.36 mm ( = 0.27 mm,
n = 10) diameter, pycnidial, solitary, scattered, superficial to semi-immersed, unilocular, globose,
black. Pycnidial wall multi-layered, 25 µm wide at the base, 16–20 µm wide in sides, outer layer
composed of 3–5 layers of thick, brown cells, inner 4–5 layers of hyaline cells of textura angularis,
cells towards inner layer becomes lighter. Conidiophores reduced to conidiogenous cells.
Conidiogenous cells 7–16 × 2–4 µm ( = 10.2 × 3.6 µm, n = 20), enteroblastic, phialidic,
doliiform, hyaline, smooth-walled, formed from the inner most layer of pycnidial wall. Conidia 11–
17 × 4–7 µm ( = 13.9 × 5.2 µm, n = 30), oblong, rarely allantoid, straight or slightly curved,
initially aseptate, hyaline, becoming 1–3 transverse septate, pale to dark brown at maturity, rarely a
longitudinal septum in the middle cell, smooth-walled, rounded at both ends.
Culture characteristics – Colonies on PDA, circular, fimbriate, rough margin, with both
surfaces grayish-white in the margin and grey olivaceous towards the center, slow growing, reach 3
cm diameter after 7 days at 20 °C.
Material examined – RUSSIA, Rostov region, Krasnosulinsky District, Donskoye forestry,
bottomland shrubbery near Kundryuchya River, on dead and dying branches of Populus nigra L.
var. italica Münchh. (Salicaceae), 6 April 2016, Timur S. Bulgakov TB(F) 1428 (MFLU 16-1722,
holotype), ex-type living culture, JZB31170001, JZB31170002, MFLUCC 18-0087, KUMCC 170324; ibid. (HKAS 101462, isotype).
GenBank numbers – MFLUCC 18-0087 – ITS: MG571222, LSU: MG571224, SSU:
MG571226; JZB31170002 – LSU: ITS: MG571223, MG571225, SSU: MG571227.
Notes – Morphological characters indicate that our species belongs to Camarosporidiellaceae
and phylogenetic analysis reveal that C. populina constitutes a strongly supported independent
lineage (93% MLBT /1.00 PP) in Pleosporinae, Pleosporales. In our phylogenetic analyses of
combined LSU, SSU and ITS sequence data of Camarosporidiella (Fig. 4), C. populina strains
cluster together sister to C. celtidis (MFLUCC 15-0444) and an unidentified Camarosporidiella
species. Camarosporidiella populina is distinct in having superficial to semi-immersed, smaller
conidiomata (0.2–0.36 mm), phialidic, conidiogenous cells and 1–3 transversely septate, larger
conidia (11–17 × 4–7 µm), in contrast to immersed, large conidiomata (0.35−0.45 mm), annellidic,
integrated, conidiogenous cells, smaller conidia (15−20 × 6−8 μm) with 2−3-transverse septa and a
longitudinal septum in C. celtidis (Wanasinghe et al. 2017b). Our species also shares some
morphological characters with C. populinum Maubl. Since, this species was introduced in 1923,
DNA sequences are unavailable in databases. Our species exhibits conidial characters similar to C.
populinum, while all other morphological characters were found to be different (Henkel 1923).
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Figure 5 – Camarosporidiella populina (MFLUCC 18-0087, holotype). a Appearance of
conidiomata on dead branch of Populus nigra var. italica. b Horizontal section of a conidioma. c
Longitudinal section of a conidioma. d–e Longitudinal section of conidioma wall showing cell
organization. f–g Conidiogenous cells with developing conidia. h–i Different septation patterns on
conidia. j–k Upper view (j) and the reverse view (k) of the colony on PDA. Scale bars: a = 1 mm, b
= 10 μm, c = 50 μm, d–e = 20 μm, f– i = 10 μm.
Dictyosporiaceae Boonmee & K.D. Hyde
The family Dictyosporiaceae was first mentioned in Liu et al. (2015) which was referred as
Dictyosporiaceae. Tanaka et al. (2015) included Dictyosporiaceae in the order Pleosporales within
the suborder Massarineae. Boonmee et al. (2016) validated this family typified by the genus
Dictyosporium Corda, along with nine genera named Aquaticheirospora, Cheirosporium,
Dendryphiella,
Dictyocheirospora,
Digitodesmium,
Gregarithecium,
Jalapriya,
Pseudocoleophoma, Pseudodictyosporium and an undetermined genus based on both morphology
and phylogeny. In this paper, we introduce a new species of Dendryphiella and provide an updated
backbone tree for the family.
285
Figure 6 – Maximum likelihood (RAxML) tree based on analysis of a combined dataset of ITS and
LSU sequence data representing Dictyosporiaceae. Related sequences are obtained from GenBank.
Thirty-one stains are included in the analyses, which comprise 1892 characters including gaps. Tree
topologies are similar between ML and BI analyses. The tree is rooted with Paradendryphiella
arenariae (CBS 181.58) and P. salina (CBS 142.60). The best scoring RAxML tree with a final
likelihood value of -8400.855146 is presented. The matrix had 527 distinct alignment patterns, with
23.99% of undetermined characters or gaps. Estimated base frequencies were as follows; A =
286
0.244437, C = 0.231831, G = 0.277299, T = 0.246433; substitution rates AC = 1.481839, AG =
2.484138, AT = 2.286573, CG = 0.384699, CT = 6.114449, GT = 1.000000; gamma distribution
shape parameter α = 0.145556. Bootstrap support values for ML greater than 60% and Bayesian
posterior probabilities greater than 0.95 are given near nodes respectively. The scale bar indicates
0.03 changes. The genetic types are indicated in bold and new isolate in bold and blue.
Dendryphiella Bubák & Ranoj.
The genus Dendryphiella was established by Ranojevic (1914), with the type species D.
interseminata (Berk. & Ravenel) Bubák. The genus is characterized by macronematous
conidiophores with polytretic, integrated conidiogenous cells at the swollen apices and intercalary
swellings and catenate or solitary conidia (Ellis 1971, Matsushima 1971, Rai & Kamal 1986, Guo
& Zhang 1999, Crous et al. 2014, Crous et al. 2016, Liu et al. 2017b). Conidiogenesis in
Dendryphiella is distinct from other genera residing in Dictyosporiaceae. Historically,
Dendryphiella was transferred to Dendryphion and Scolecobasidium based on morphology. Liu et
al. (2017b) updated the taxonomy of this genus with twelve species and a key.
Dendryphiella phitsanulokensis N.G. Liu & K.D. Hyde, sp. nov.
Fig. 7
Index Fungorum number: IF554049; Facesoffungi number: FoF03897
Etymology – name reflects the location where the specimen was collected.
Holotype – MFLU 17-2651
Saprobic on decaying wood. Sexual morph Undetermined. Asexual morph Colonies on
natural substrate superficial, effuse, greyish-brown. Mycelium partly immersed, composed of
septate, branched, brown, guttulate, 1.5−3 μm wide hyphae. Conidiophores 130−260 μm long,
macronematous, mononematous, occasionally fasciculate, dark brown at base, paler towards the
apex, thick-walled, erect, straight or slightly flexuous, minutely verruculose, septate, unbranched,
wider at the sub-section. Conidiogenous cells 15−28 μm long ( = 22.35 μm, n = 15), polytretic,
terminal and intercalary, proliferating asymmetrically, integrated, brown, minutely verrucose,
enlarged at vertex. Conidia 16−26 × 2.5−4.4 μm ( = 12.09 × 3.59 μm, n = 25), solitary to catenate,
when catenate in acropetal chains, fusiform to ellipsoidal, rounded at apex, truncate at the base,
pale brown, aseptate when young, brown or dark brown, 3(4)-septate when mature, slightly
constricted at septa, thick-walled, verrucose.
Culture characteristics – Conidia germinated on WA (Water Agar) within 24 hours. One or
two germ tubes produced from both ends. Colonies reached about 5 cm diameter after 2 weeks on
PDA at 25°C. Mycelia are superficial, with entire edge, floccose at the center, white to light brown
from above and light brown from below.
Material examined – THAILAND, Phitsanulok Province, on decaying wood, 10 October
2016, Ningguo Liu, J4 (MFLU 17-2651, holotype); ex-type living culture, MFLUCC 17-2513.
GenBank numbers – ITS: MG754400, LSU: MG754401, SSU: MG754402.
Notes – Dendryphiella phitsanulokensis and D. fasciculata share similar conidial characters
(pale brown, aseptate when young, brown, 3-septate when mature). The appearance of most solitary
conidiophores makes D. phitsanulokensis different from D. broussonetiae, D. fasciculata and D.
lycopersicifolia which have fasciculate conidiophores. Dendryphiella phitsanulokensis resembles
D. aspera in occasionally having fasciculate conidiophores. However, D. phitsanulokensis has
shorter conidiophores and smaller conidia than those of D. aspera (100−150 vs. up to 136−544 μm
and 9−14 × 2.5−4.4 vs. 10−22 × 4−6 μm). Moreover, D. phitsanulokensis differs from D. eucalypti
in having rough conidia, while the latter have smooth conidia. Phylogenetic analysis indicates a
close affinity of D. phitsanulokensis to D. vinosa and D. eucalyptorum without much resolution
(Fig. 6). This could be an artifact of taxon sampling (e.g Huang et al 2017) and problems associated
with only the rDNA sequence data analysed (Liu et al. 2017b). However, D. phitsanulokensis
differs from D. vinosa and D. eucalyptorum in terms of shorter and unbranched conidiophores.
Moreover, examination of the 500 nucleotides across the ITS regions reveals 12 bp (2.4%)
differences between D. phitsanulokensis and D. eucalyptorum (KJ869139).
287
Figure 7 – Dendryphiella phitsanulokensis (MFLU 17-2651, holotype). a, b Colonies on host. c, d
Conidiophores. e–g Conidiogenous cells and conidia. h–m Conidia. Scale bars: c, d = 25 μm, e–m
= 10 μm.
Didymellaceae Gruyter et al.
Didymellaceae was introduced by de Gruyter et al. (2009) to accommodate Ascochyta,
Didymella, Phoma and phoma-like genera. The family comprises numerous endophytic, pathogenic
288
and saprobic species associated with a wide range of hosts worldwide (Chen et al. 2015, Hyde et al.
2016). In this paper, we provide an updated phylogeny for selected genera (Ascochyta, Didymella,
Epicoccum and Neodidymelliopsis) in Didymellaceae (Fig. 8). We also introduce the new species,
Ascochyta nepetigena and Neodidymelliopsis negundinis and provide new host records for
Didymella aliena and Epicoccum nigrum.
Ascochyta Lib.
The genus Ascochyta was described and introduced by Libert in 1830 typified by A. pisi
(=Didymella pisi Chilvers, J.D. Rogers & Peever 2009). Ascochyta is characterized by globose
locules with perithecial protuberances immersed in the stroma. Most species in Ascochyta are
endophytic, pathogenic and saprobic and associated with a number of hosts worldwide
(Wijayawardene et al. 2017a). Ascochyta was accepted as a genus in Didymellaceae (Hyde et al.
2013, Kirk et al. 2013). The asexual morph of Ascochyta is coelomycetous (Chen et al. 2015).
Currently, around 400 species are accomodated in this genus (Wijayawardene et al. 2017a),
however, the genus Ascochyta needs generic revision as more than 1000 records are available. In
this paper, we introduce a new species in the genus Ascochyta which was isolated from the stem of
Clinopodium nepeta in Italy.
Ascochyta clinopodiicola D. Pem, Camporesi & K.D. Hyde, sp. nov.
Fig. 9
Index Fungorum number: IF554300; Facesoffungi number: FoF04374
Etymology – referring to the host from which the specimen was isolated.
Holotype – MFLU 17-1034
Saprobic on Clinopodium nepeta. Sexual morph Ascomata 16−33 µm diameter, 15−33 µm
high ( = 19.3 × 25.4 µm, n = 20), pseudothecia, solitary or gregarious, semi-immersed to
erumpent, or rarely superficial, black, papillate, ostiole filled with hyaline cells. Peridium 8–9 µm
wide at the base, 15–20 µm wide at the sides, comprising two layers cells of textura angularis,
innermost layer pale brown to darker brown at outer surface, thin-walled. Hamathecium lacking
pseudoparaphyses. Asci 25−34 × 5−7 µm ( = 28.6 × 6.5 µm, n = 10), 8-spored, bitunicate,
fissitunicate, cylindric-clavate to clavate, short-pedicellate, apically rounded, with an ocular
chamber. Ascospores 8−11 × 2−3 µm ( = 9.5 × 2.5 µm, n = 10), overlapping uniseriate or
biseriate, hyaline, ovoid to ellipsoidal, asymmetrical, upper cell wider than lower cell, 1-septate,
slightly constricted at the septum, smooth-walled. Asexual morph Undetermined.
Culture characteristics – Ascospores germinating on MEA within 24 hours. Colonies growing
on MEA, reaching 2 cm diameter in 1 week at 16°C. Mycelium superficial, surface smooth,
irregular, slightly raised, edge crenate, velutinous, from above white to pale yellow, reverse yellowbrown to orange-brown.
Material examined – ITALY, Province of Forlì-Cesena [FC], Spinello - Santa Sofia, on dead
aerial stem of Clinopodium nepeta (L.) Kuntze (Lamiaceae), 18 May 2017, Erio Camporesi IT
3353 (MFLU 17-1034, holotype); ex-type living culture, MFLUCC 18-0344, ICMP; ibid. (HKAS
97475, isotype).
GenBank numbers – ITS: MH017431, LSU: MH017429, SSU: MH017430.
Notes – During our investigation on the diversity of microfungi in Italy, a fungal isolate was
collected from a dead stem of Clinopodium nepeta in the Province of Forlì-Cesena. Morphological
characters, namely erumpent, papillate ascomata, bitunicate, subcylindrical to subclavate,
somewhat curved, slightly pedicellate asci and ovoid to ellipsoidal, hyaline, 1-septate ascospores
(Fig. 9), fit well within the species concept of Ascochyta (Kaiser et al. 1997, Chilvers et al. 2009).
DNA sequence analyses from the ribosomal genes confirm that our new taxon belongs to the genus
Ascochyta and is closely related to A. medicaginicola and A. premilcurensis (Fig. 8). However, our
new taxon A. clinopodiicola differs from A. medicaginicola by its smaller ascomata (16−33 µm
diameter, 15−33 µm high vs. 165−190 μm high, 170−210 μm wide), shorter asci (25−34 × 5−7 µm
vs. 50−84 × 8−14 μm) and narrower ascospores (8−11 × 2−3 µm vs. 12−16 × 3−5 μm) (Jayasiri et
al. 2017).
289
290
Figure 8 – Phylogram generated from maximum likelihood analysis of a combined LSU, ITS,
RPB2 and β-tubulin DNA sequence data of Didymellaceae. Related sequences were obtained from
Hyde et al. (2017a, b) and Chen et al. (2017). One hundred and thirty-three strains are included in
the combined dataset, which comprise 2863 characters with gaps. Single gene analyses were
carried out and compared with each species, to compare the topology of the tree and clade stability.
291
Leptosphaerulina australis (CBS 317.83) and Leptosphaerulina trifolii (CBS 235.58) are used as
the outgroup taxa. Tree topology of the ML analysis was similar to the BI. The best scoring
RAxML tree with a final likelihood value of -22539.886013 is presented. The matrix had 748
distinct alignment patterns, with 17.94% of undetermined characters or gaps. Estimated base
frequencies were as follows; A = 1.000000, C = 0.242053, G= 0.239955, T = 0.274733;
substitution rates AC = 1.818252, AG = 1.330539, AT = 5.943005, CG = 1.868178, CT =
0.883503, GT = 11.631688; gamma distribution shape parameter α = 0.788126. Bootstrap support
values for ML equal to or greater than 65% and BYPP equal to or greater than 0.90 are given above
the nodes respectively. Ex-type strains and reference strains are in bold. Newly generated
sequences are in red.
Figure 9 – Ascochyta clinopodiicola (MFLU 17-1034, holotype) a, b Ascomata on host surface. c
Section of ascoma. d Peridium. e Asci. f–h Developmental stages of Asci. i–k Ascospores. l
Germinated Ascospores. m, n Culture characters on MEA (m: above view, n: reverse view). Scale
bars: a, b = 500 µm, c = 15 µm, d–h = 10 µm, i–k = 5 µm, l = 10 µm.
292
Didymella Sacc.
Didymella was described by Saccardo in 1880 with the description of Didymella exigua
(Niessl) Sacc. (Corlett 1981, Chen et al. 2015). This genus has received much attention in recent
studies and several new species has been introduced (Wijayawardene et al. 2017a, Thambugala et
al. 2017, Chen et al. 2017, Valenzuela-Lopez et al. 2018).
Didymella aliena (Fr.) Q. Chen & L. Cai, Stud. Mycol. 82: 173 (2015)
Fig. 10
Facesoffungi number: FoF03870
Basionym – Sphaeria aliena Fr., Syst. Mycol. 2(2): 502 (1823).
Opportunistic pathogenic on dead and dying branch of Malus domestica Borkh. Sexual
morph Undetermined. Asexual morph Coelomycetous. Conidiomata on host 0.17–0.33 mm ( =
0.23 mm, n = 10) diameter, pycnidial, solitary, scattered, semi-immersed, unilocular, globose,
black; in culture: 0.12–0.25 mm ( = 0.17 mm, n = 10) diameter, pycnidial, solitary, scattered,
superficial, globose, black. Pycnidial wall 9–17 µm, multi-layered, outer layer composed of 3–4
layers of thick, dark brown cells, inner 2–3 layers of hyaline cells of textura angularis.
Conidiophores reduced to conidiogenous cells. Conidiogenous cells on host 4–8 × 1–3 µm ( = 5.6
× 2.1 µm, n = 20), enteroblastic, phialidic, globose to bottle-shaped, hyaline, smooth-walled,
formed from the inner most layer of pycnidial wall. Conidia on host 6–10 × 3–5 µm ( = 8.2 × 4.2
µm, n = 30), ellipsoid, straight, initially aseptate, hyaline, becoming 1-septate, pale brown at
maturity, smooth-walled, acutely-rounded apex; in culture: 3–7.5 × 2–4 µm ( = 5.8 × 3.1 µm, n =
30), ellipsoid to slightly ovoid, initially hyaline, becoming light brown at maturity, aseptate,
smooth-walled, with acutely-rounded ends, with or without some small guttules at the ends.
Culture characteristics – Colonies on PDA reach 45 mm diameter, after 3 days at 25°C,
floccose to woolly, white to grey olivaceous aerial mycelium, reverse olivaceous grey to dark iron
grey, with vinaceous buff margin.
Material examined – RUSSIA, Rostov region, Shakhty City, on dead and dying branch of
Malus domestica Borkh. (Rosaceae), 1 March 2016, Timur S. Bulgakov T-1235 (MFLU 16-1529);
living cultures, JZB380010, MFLUCC 18-0086, KUMCC 17-0323.
GenBank numbers – β-tubulin: MG571228, ITS: MG571229, LSU: MG571230, RPB2:
MG571231.
Notes – Didymella aliena has been reported from France, Italy, Netherlands and Poland on
several hosts including Berberis sp., Buxus sp., Cotoneaster sp., Euonymus europaeus, Mahonia
aquifolium and Pyrus calleryana (Boerema et al. 2004, Chen et al. 2015, Farr & Rossman 2017).
Based on our phylogenetic analysis of combined LSU, ITS, RPB2, and β-tubulin sequence data of
Didymellaceae species (Fig. 8), our strain (MFLUCC 18-0086) clusters with the ex-type strain of
Didymella aliena (CBS 379.93) with high bootstrap support (100% MP/1.00 PP respectively). The
holotype of the Sphaeria aliena (basionym of Didymella aliena) is not known to exist. Therefore, a
neotype was designated for the species (de Gruyter et al. 1998). Our isolate is similar to the neotype
of D. aliena (CBS 379.93). However, the description only includes several morphological
characters on different kinds of agar. In this paper we provide a full description of the species on
the host, as well as on media.
Epicoccum Link
The genus Epicoccum has an estimated over 30 species (Hyde et al. 2017b, Wijayawardene et
al. 2017a, Wanasinghe et al. 2018). Epicoccum has a hyphomycetous asexual morph and the sexual
morph is presently unknown. Epicoccum has a cosmopolitan distribution and comprises saprobes in
terrestrial habitats (Wijayawardene et al. 2017a).
Epicoccum nigrum Link, Mag. Gesell. naturf. Freunde, Berlin 7: 32 (1816) [1815]
Fig. 11
Facesoffungi number: FoF02685
Saprobic on dead leaves of Sasa nipponica. Sexual morph Undetermined. Asexual morph
293
Figure 10 – Didymella aliena (MFLUCC 18-0086). a, b Semi-immersed conidiomata on the host
surface. c Superficial pycnidia in agar. d Longitudinal section of a conidioma. e Conidioma wall
showing cell organization in host. f Sections of the pycnidial wall from culture. g Conidiogenous
cells from host. h Conidiogenous cells from culture. i Conidia from host. j Conidia from culture. k–
l Upper view (k) and the reverse view (l) of the colony on PDA. Scale bars: a = 2 mm, b, c = 500
μm, d = 50 μm, e, f = 20 μm, g = 10 μm, h = 20 μm, i–j = 20 μm.
294
Colonies effuse, dark brown to black with occasional bright orange anterior margins. Mycelium
mostly immersed, with dematiaceous hyphae. Conidiophores 3–6 × 3–4 μm (x̅ = 5 × 3 μm, n = 10),
micronematous, mononemous, closely packed, hyaline, thin-walled, aseptate, smooth.
Conidiogenous cells 3–4 × 4–7 μm (x̅ = 3 × 5 μm, n = 10) integrated. Conidia 10–21 × 8–21 μm (x̅
= 15 × 15 μm, n = 40), solitary, sphaerical or pyriform, dark golden brown, verrucose, muriform,
slightly constricted at septa, sometimes with septa obscured at maturity by rough wall, sometimes
remaining attached with a pale protuberant basal stalk cell.
Culture characteristics – Conidia germinating on PDA, within 12 hours. Colonies growing on
PDA, cottony, bright yellow to bright orange reaching 5 mm in 10 days at 25°C, mycelium
superficial, effuse, with regular edge, hyphae pale to bright orange, producing orange pigments in
PDA
Material examined – CHINA, Yunnan Province, Kunming Institute of Botany, Botanical
Garden, on dead leaves of Sasa nipponica, 5 October 2016, A. Karunarathna, AKKIB 10 (MFLU
17-0359, HKAS 97353); living culture MFLUCC 17-1371, KUMCC 16-0221.
GenBank numbers – ITS: MG822858, LSU: MG822859.
Notes – Pairwise comparison of DNA sequences of ITS regions between our strain KUMCC
16-0221 and the type strain CBS 173.73 revealed very minor differences. Based on Jeewon & Hyde
(2016) and morphological similarities, we treat both as the same species. Phylogeny (Fig 8) also
supports this identification. Thus, this fresh collection is identified as Epicoccum nigrum.
Description and illustrations are provided which will facilitate identification of Epicoccum nigrum
on Sasa nipponica (Poaceae) as the first report from China.
Neodidymelliopsis Q. Chen & L. Cai.
Neodidymelliopsis was introduced by Chen et al. (2015), with Neodidymelliopsis cannabis
(G. Winter) Q. Chen & L. Cai as the type species (Chen et al. 2015, Hyde et al. 2016, Thambugala
et al. 2016a).
Neodidymelliopsis negundinis Manawasinghe, Bulgakov & K.D. Hyde, sp. nov.
Fig. 12
Index Fungorum number: IF554044; Facesoffungi number: FoF03891
Etymology – name reflects the host genus.
Holotype – MFLU 16-1733
Saprobic or necrotrophic on dead and dying twigs and branches of Acer negundo L. Sexual
morph Undetermined. Asexual morph Coelomycetous. Conidiomata on host 255–559 µm diameter
( = 395.4 µm, n = 10), pycnidial, solitary, scattered, globose to subglobose, black, semi-immersed
to immersed, sometimes erumpent; Conidiophores not observed. Conidiogenous cells not observed.
Conidia on host 4–10 × 1.5–2 µm diameter ( = 5.5 × 1.8 µm, n = 40), oblong to ellipsoid, hyaline,
aseptate, smooth-walled; in culture, 4.6–6.7 × 1.6–2.4 µm diameter ( = 5.5 × 2 µm, n = 40),
straight. Conidial exudates not observed.
Culture characteristics – Colonies on PDA reach 80 mm diameter after 7 days at 25 °C, with
undulate edge, with grey olivaceous aerial mycelium, surface floccose to woolly
Material examined – RUSSIA, Rostov region, Shakhty City, urban artificial forest, on dead
and dying twigs and branches of Acer negundo (Sapindaceae), 6 April 2016, Timur S. Bulgakov
(MFLU 16-1733, holotype), ex-type living culture, MFLUCC 18-0083, KUMCC 18-0006.
GenBank numbers – β-tubulin: MG564164, ITS: MG564165, LSU: MG564163, RPB2
MG564166.
Notes – Neodidymelliopsis negundinis was collected from Acer negundo in the Rostov region
(European part of Russia). Morphological characters such as spores and colony characters (Fig. 12),
fit well within the species concept of Neodidymelliopsis (Chen et al. 2015). DNA sequence
analyses from the ITS, LSU, RPB2 and β-tubulin genes confirm that our new taxon belongs in
Neodidymelliopsis (Fig. 8). Neodidymelliopsis negundinis has a particular neighbour relationship
with N. longicolla and N. polemonii, but this was not supported in our phylogeny analyses (Fig. 8).
Neodidymelliopsis negundinis differs by developing smaller-sized (4–10 × 1.5–2.45 conidia in both
295
culture and on the host compared to other species in this genus (7.5–10(–18) × 2–3.5(–5) (Chen et
al. 2015, Hyde et al. 2017b). In addition, this is the first report of Neodidymelliopsis reported on
Acer species (Farr & Rossman 2018).
Figure 11 – Epicoccum nigrum (MFLU 17-0359). a Appearance of synnemata on host substrate. b
Longitudinal section of synnema. c–d Attachment of conidia to the conidiophores. e–g Conidia. h
Germinated conidium. i–j Culture characteristics on PDA (i = from above, j = from below). Scale
bars: b = 50 µm, c–f = 10 µm, g–j = 10 µm.
Lentitheciaceae Yin. Zhang et al.
The family Lentitheciaceae was established by Zhang et al. (2012) to accommodate
massarina-like species in the suborder Massarineae and is typified by Lentithecium fluviatile
(Aptroot & Van Ryck.) K.D. Hyde et al. (Zhang et al. 2009, 2012, Hyde et al. 2013, 2016,
Tibpromma et al. 2017). Ten genera have been reported in the family, viz. Darksidea (Knapp et al.
2015), Katumota, Keissleriella, Lentithecium, Murilentithecium (Wanasinghe et al. 2014),
Neoophiosphaerella (Tanaka et al. 2015), Phragmocamarosporium (Wijayawardene et al. 2015),
Poaceascoma (Phookamsak et al. 2015a), Setoseptoria (Tanaka et al. 2015) and Tingoldiago (Hyde
et al. 2013, Knapp et al. 2015, Phookamsak et al. 2015a, Tanaka et al. 2015, Wanasinghe et al.
2018). In this paper, we introduce Poaceascoma taiwanense as a new species form Taiwan.
296
Figure 12 – Neodidymelliopsis negundinis (MFLU 16-1733, holotype) a Appearance of conidia on
dead branch of Acer negundo. b Submerged conidiomata on the host surface. c Conidia on host. d,
e Conidia on host in agar. e Upper view of 7 days old culture on PDA. f Reverse view of 7 days old
culture on PDA. f Upper side of colony on PDA. Scale bars: a = 2000 µm, b = 200 µm, c, d, e = 20
µm.
Poaceascoma Phookamsak & K.D. Hyde
Poaceascoma was introduced by Phookamsak et al. (2015a) to accommodate a
dothideomycete species associated with Poaceae which forms setose ascoma with filiform
ascospores and is typified by Poaceascoma helicoides Phookamsak & K.D. Hyde. Currently, three
species have been recognized in the genus viz. Poaceascoma aquaticum Z.L. Luo & K.D. Hyde, P.
halophila Dayarathne & K.D. Hyde and P. helicoides Phook. & K.D. Hyde (Index Fungorum 2018)
Poaceascoma taiwanense Tennakoon, C.H. Kuo & K.D. Hyde, sp. nov.
Fig. 14
Index Fungorum number: IF554200; Facesoffungi number: FoF04084
Etymology – named after the country where this fungus was collected, Taiwan.
Holotype – MFLU 18-0083
Saprobic on dead stem of Panicum virgatum L. Sexual morph Ascomata 140–180 µm high,
150–220 µm diameter, semi-immersed to erumpent, solitary, scattered or sometimes grouped
beneath the host tissues, globose, dark brown to black, coriaceous, ostiolate, papillate. Peridium
40–50 μm wide, with thick walls, composed of several layers of pseudoparenchymatous cells, outer
layer comprising several layers of dark brown cells, arranged in a textura angularis, inner layer
comprising several layers of hyaline, flattened cells, arranged in a textura angularis to textura
prismatica. Hamathecium composed of numerous, 2−3.5 μm wide, filamentous, broad, cellular
pseudoparaphyses, with distinct septa, embedded in a mucilaginous matrix. Asci (63–)65–80(–82) ×
6–8(–8.6) μm ( = 73 × 7.2 μm, n = 30), 4-spored, bitunicate, fissitunicate, cylindrical to cylindricclavate, short pedicellate, apically rounded with an ocular chamber. Ascospores 50–60 × 2–3 μm
( = 55.6 × 2.6 μm, n = 30), fasciculate, scolecosporous, hyaline, elongate, filiform, tapering
towards the rounded ends, slightly curved, 15–17-septate, slightly constricted at septa, smoothwalled. Asexual morph Undetermined.
297
Figure 13 – Phylogram generated from maximum likelihood analysis based on combined LSU,
SSU, ITS and TEF sequenced data of Lentitheciaceae. Related sequences were obtained from Luo
298
et al. (2016) and Hyde et al. (2017b). Sixty-six strains are included in the combined sequence
analyses, which comprise 3421 characters with gaps. Massarina eburnea (H 3953, CBS 47364) and
Massarina cisti (CBS 26662) are used as the outgroup taxa. Tree topology of the ML analysis was
similar to the BI. The best scoring RAxML tree with a final likelihood value of -15553.519520 is
presented. The matrix had 847 distinct alignment patterns, with 22.85% of undetermined characters
or gaps. Estimated base frequencies were as follows; A = 0.239923, C = 0.248830, G= 0.272362, T
= 0.238884; substitution rates AC = 1.408036, AG = 2.598862, AT = 1.789820, CG = 1.645928,
CT = 8.851789, GT = 1.000000; gamma distribution shape parameter α = 0.497322. Bootstrap
support values for ML equal to or greater than 60% and BYPP equal to or greater than 0.95 are
given above the nodes respectively. Newly generated sequences are in red.
Figure 14 – Poaceascoma taiwanense (MFLU 18-0083, holotype) a Ascomata on host. b Close-up
of ascomata. c Vertical section of ascoma. d Section of peridium. e Pseudoparaphyses. f–i Asci. j–o
Ascospores. Scale bars: c = 50 µm, d–i = 20 µm, j–o = 18 µm.
Material examined – TAIWAN, Chiayi, Shihnong Forest, dead stems of Panicum virgatum L.
(Poaceae), 25 June 2017, D.S. Tennakoon, DTW 030 (MFLU 18-0083, holotype).
299
GenBank numbers – ITS: MG831569, LSU: MG831567, SSU: MG831568.
Notes – Morphological characters of Poaceascoma taiwanense fit into the generic concept of
Poaceascoma in having filiform, multi-septate ascospores (Phookamsak et al. 2015a, Luo et al.
2016, Hyde et al. 2017b). Poaceascoma taiwanense differs from other Poaceascoma species in
having unique morphological characters, such as 4-spored asci and 15–17-septate ascospores. The
main morphological differences across similar species are shown in Table 1.
Table 1 Synopsis of morphological similarities and differences among Poaceascoma species.
Poaceascoma
species
P. aquaticum
P. halophila
P. helicoides
P. taiwanense
Ascomata
Size (μm)
Asci
Ascospores
280–380 ×
310–500
184.5–210.5
× 12–16
214–231 ×
4.2–5.2
260–315 ×
270–350
270–360 ×
320–450
140–180×
150–220
248.5–300 ×
11–14
160–185 ×
8.5–10
191–230 ×
2.5–3
150–185 ×
2–2.5
65–80× 6–8
50–60 × 2–3
Ascospores
Septation Colour
Pale
brown to
21–36
brown
References
Luo et al. 2016
20–25
Hyaline
29–33
Hyaline
Hyde et al.
2017b
Phookamsak et
al. 2015a
15–17
Hyaline
This study
Melanommataceae G. Winter
The family Melanommataceae was established by Winter (1885), which is typified by
Melanomma pulvis-pyrius. Tian et al. (2015) accepted 20 genera in Melanommataceae based on
morphological and phylogenetic evidence. Recent taxonomic surveys have also revealed the family
to be highly diverse with many new species (Hyde et al. 2016, Li et al. 2016b, 2017, Almeida et al.
2017, Jaklitsch & Voglmayr 2017). Hashimoto et al. (2017) reported that Melanommataceae should
be restricted to the type genus Melanomma. In addition, they have introduced Pseudodidymellaceae
to accommodate four genera viz. Mycodidymella, Petrakia, Pseudodidymella, and Xenostigmina.
However, in a recent study, Wanasinghe et al. (2018) considered that the new family
Pseudodidymellaceae was untenable as it resulted in more intergeneric taxonomic confusion and
the family Melanommataceae was already a well established family with strong phylogenetic
support. Besides, they introduced five new genera in the family Melanommataceae, viz.
Melanocucurbitaria, Marjia, Melanodiplodia, Monoseptella, Uzbekistanica. In this paper, we
introduce Pseudobyssosphaeria gen. nov with Pseudobyssosphaeria bambusae sp. nov, two new
records, Bertiella ellipsoidea and Byssosphaeria siamensis, our conclusions based on multi-gene
analyses and morphological characters (Fig. 15).
Bertiella (Sacc.) Sacc. & P. Syd.
The genus Bertiella was introduced by Saccardo and Sydow (1899) and it is typified by
Bertiella macrospora (Sacc.) Sacc. & Traverso. Recent phylogenetic studies support its placement
within Melanommataceae close to Byssosphaeria (Mugambi & Huhndorf 2009, Tian et al. 2015,
Hyde et al. 2016, Li et al. 2016b, 2017). Previously, Bertiella was characterized by black,
superficial, subglobose ascomata with a carbonaceous peridium. However, we considered that a
carbonaceous peridium is not an absolute characteristic to identify this genus. In this paper, we
introduce a new collection of Bertiella ellipsoidea from submerged wood in a freshwater stream in
northern Thailand.
Bertiella ellipsoidea Ekanayaka, Q. Zhao & K.D. Hyde, Fungal Diversity 80: 79 (2016)
Fig. 16
Index Fungorum number: IF552201; Facesoffungi number: FoF03894
Saprobic on woody substrata in a freshwater stream. Sexual morph Ascomata 170–375 high ×
165–345 μm diameter ( = 285 × 214 μm, n = 15), superficial, solitary or scattered, sessile,
300
Figure 15 – Phylogram generated from maximum likelihood analysis based on combined ITS,
LSU, SSU and TEF1α sequence data for species of Melanommataceae. The newly generated
nucleotide sequences were compared against the GenBank database using the Mega BLAST
program. Sequences that relate were obtained from GenBank (http://www.ncbi.nlm.nih.gov/).
Fifty-five strains are included in the combined sequence analyses which comprise 3,894 characters
with gaps (539 characters for ITS, 1308 for LSU, 1061 for SSU and 986 for TEF1α). Cyclothyriella
rubronotata TR9 and Cyclothyriella rubronotata TR (Cyclothyriellaceae) are used as the outgroup
taxa. Tree topology of the maximum-likelihood analysis is similar to the maximum parsimony
analysis and the Bayesian analysis. The best scoring RAxML tree with a final likelihood value of 16110.950054 is presented. RAxML bootstrap support values equal to or greater than 70% are
given before the forward slash. Bayesian posterior probabilities equal to or higher than 0.95 are
given after the forward slash. Hyphen (“–”) indicates a value lower than 70% for RAxML and a
posterior probabilities lower than 0.95 for Bayesian analysis. Newly generated sequences are in red.
301
globose to subglobose, dark brown, setose, coriaceous. Ostiole single, central. Setae 160–330(–
398) × 3–4 μm, covering the whole ascoma, brown to dark brown, unbranched, apically rounded,
thick-walled. Peridium 27–40 μm thick, 2-layered, outer layer comprising pale brown to dark
brown cells of textura angularis, inner layer comprising hyaline cells of textura angularis.
Hamathecium comprising numerous, 1.5–2.5 μm wide, hyaline, long, filiform, aseptate, branched
pseudoparaphyses, anastomosing between and above the asci. Asci 100–150 × 8–12 μm ( = 120 ×
10 μm, n = 20), 8-spored, bitunicate, cylindrical to clavate, short pedicellate, apically rounded, with
an ocular chamber. Ascospores 16–21 × 6–8 μm ( = 18 × 7 μm, n = 50), 1–2-seriate, ellipsoid to
fusiform, hyaline to pale brown, 1-septate, constricted at septum, acute at the apex, smooth and
thick-walled, with 2–4 globules. Asexual morph Undetermined.
Culture characteristics – Ascospores germinating on water agar (WA) within 48 hours and
germ tubes produced from ascospores. Colonies growing slowly on malt extract agar (MEA),
circular, with flat surface, edge undulate, reaching 6 mm in 2 weeks at 28°C, white to brown in
MEA media. Mycelium superficial and partially immersed, branched, septate, hyaline to pale brown
(Figs 16t, 16u).
Material examined – THAILAND, Chiang Rai, Muang, Ban Nang Lae Nai, on submerged
decaying wood in a freshwater stream, 31 December 2016, Yong-Zhong Lu, CR10 (MFLU 171092, HKAS 100762); living culture, MFLUCC 17-2015, TBRC.
GenBank numbers – ITS: MG543922, LSU: MG543913, RPB2: MG547224, TEF1:
MG547226.
Notes – A new collection of Bertiella ellipsoidea (MFLU 17-1092) was found on submerged
wood in a freshwater stream from northern Thailand and it is the first report of B. ellipsoidea from
aquatic habitats. Phylogenetic analyses placed our new strain in a clade with B. ellipsoidea (MFLU
16-0583) (Fig. 15). We compared our new taxon with B. ellipsoidea based on base pair differences
of ribosomal and protein loci. There was only one base pair difference in LSU, and there were no
differences in SSU and TEF1α, which provide further evidence to support our new strain as
Bertiella ellipsoidea. Morphologically there are two different features between our new collection
and the holotype of B. ellipsoidea (MFLU 16-0583) (Hyde et al. 2016): viz. i) there are abundantly
long setae covering the whole ascoma and ii) coriaceous ascomata (carbonaceous in the holotype).
Thus, we name our strain as B. ellipsoidea (MFLUCC 17-2015) as the first record from aquatic
habitats. Moreover, we also provide their culture characteristics which were obtained from single
spore isolations.
Byssosphaeria Cooke
Byssosphaeria was introduced by Cooke & Plowright (1879) and is typified by B. keithii
based on its superficial ascomata seated on a “tomentose subiculum of interwoven threads” (Cooke
& Plowright 1879). Tian et al. (2015) introduced a new Byssosphaeria species and reported that
Byssosphaeria belongs to Melanommataceae based on multigene phylogenetic analyses. Index
Fungorum (2018) lists 44 species epithets under Byssosphaeria, but only seven species have been
confirmed by molecular data, viz. B. jamaicana (Sivan.) M.E. Barr, B. musae Phook. & K.D. Hyde,
B. rhodomphala (Berk.) Cooke, B. salebrosa (Sacc.) M.E. Barr, B. schiedermayeriana (Fuckel)
M.E. Barr, Byssosphaeria siamensis Boonmee, Q. Tian & K.D. Hyde, Byssosphaeria villosa
(Samuels & E. Müll.) Boise. In this study, we report Byssosphaeria siamensis with new
morphological data.
Byssosphaeria siamensis Boonmee, Q. Tian & K.D. Hyde, Fungal Diversity 74: 283 (2015)
Fig. 17
Index Fungorum number: IF551430; Facesoffungi number: FoF03895
Saprobic on woody substrata in a freshwater stream. Sexual morph Ascomata 445–475 μm
high × 430–555 μm diameter ( = 460 × 490 μm, n = 5), superficial, globose to subglobose,
solitary to gregarious, dark brown to black, setose, coriaceous, apically ostiolate. Ostiole single,
central, with a pore-like opening, orange to yellow around pore, apapillate. Setae 325–400 × 4.5–6
302
μm, covering the whole ascoma, dark brown to black, unbranched, thick-walled. Peridium 50–70
μm wide, thickest at the apex, 2-layered, outer layer comprising pale brown to dark brown cells of
textura angularis, inner layer comprising hyaline to pale brown cells of textura prismatica.
Hamathecium 1–2 μm wide, comprising hyaline, dense, filiform, septate pseudoparaphyses,
anastomosing and branching between and above the asci. Asci 110–150 × 12–14 μm ( = 131 × 13
μm, n = 20), 8-spored, bitunicate, cylindrical to clavate or saccate, pedicellate, with a 29–45 μm
long pedicel, apically rounded with an ocular chamber. Ascospores 28–33 × 5–7 μm ( = 30 × 6
μm, n = 50), 1–2-seriate, hyaline to pale brown, ellipsoid to fusiform, 1-septate, constricted at
septum, slightly curved, with 1–2 globules, acute at the apex, smooth and thick-walled. Asexual
morph undetermined.
Culture characteristics – Ascospores germinating on water agar within 12 hours and germ
tubes produced from ascospores. Colonies growing slowly on MEA, circular, with flat surface,
edge undulate, reaching 31 mm in 3 weeks at 28°C, pale brown to yellow in MEA media. Mycelium
superficial and partially immersed, branched, septate, hyaline to pale brown.
Material examined – THAILAND, Chiang Rai, Muang, Ban Nang Lae Nai, on submerged
decaying wood in a freshwater stream, 31 December 2016, Yong-Zhong Lu, CR25 (MFLU 171104, HKAS 100774); living culture, MFLUCC 17-1800.
GenBank numbers – ITS: MG543923, LSU: MG543914, SSU: MG543917, RPB2:
MG547225.
Notes – Our new collection shares a sister relationship to Byssosphaeria siamensis
(MFLUCC 10-1099). We compared our new taxon with B. siamensis based on base pair
differences. There were 3, 2, 1 and 3 base pair differences in ITS, LSU, SSU and RPB2
respectively, which confirmed they should belong to same species (Jeewon & Hyde 2016).
Morphologically there are four different features between our new collection and the holotype of B.
siamensis (MFLU 10-0029) (Tian et al. 2015): 1), smaller ascomata (445–475 × 430–555 μm vs.
501–692 × 561–720 μm); 2) longer pedicel (29–45 μm vs. 14.5–24 μm); 3) smaller ascospores (28–
33 × 5–7 μm vs. 40.5–50 × 7–11 μm); 4) ascospores surrounded by a gelatinous and fusiform
sheath in the holotype, but our new collection lacks these.
Pseudobyssosphaeria H.B. Jiang & K.D. Hyde, gen. nov.
Index Fungorum number – IF554471
Etymology – refers to the morphologically being similar to Byssosphaeria.
Saprobic on dead bamboo culms. Sexual morph Ascomata superficial, solitary, scattered or
gregarious, subglobose, blackened, with a central ostiole, ascomata covered with outwardly brown,
septate, branched hyphae. Setae covering the whole ascoma, dark brown, unbranched, septate.
Peridium multi-layered, outer layer composed of thick-walled, dark brown cells of textura
angularis; inner layer composed of thin-walled, hyaline cells of textura angularis. Hamathecium
comprising septate, branched, anastomosing pseudoparaphyses embedded in a gelatinous matrix.
Asci 8-spored, bitunicate, cylindrical to clavate, short pedicellate with furcate ends, apically
rounded. Ascospores overlapping bi-seriate, fusiform, 1-septate, constricted at the septum, hyaline.
Asexual morph Undetermined.
Type species – Pseudobyssosphaeria bambusae H.B. Jiang & K.D. Hyde
Notes – Although Pseudobyssosphaeria grouped with Bertiella macrospora (strains GKM
L122N and SMH 3953) and three collections of B. macrospora (IL 5005, GKM L122N and SMH
3953), they clustered into two groups in our multi-gene phylogenetic tree. Bertiella macrospora
strain IL 5005 has LSU sequence data only and strains GKM L122N and SMH 3953 have TEF1alpha sequences only. Pseudobyssosphaeria however, forms a basal clade to Bertiella and is
morphologically distinct.
303
Figure 16 – Bertiella ellipsoidea (MFLU 17-1092). a, b Superficial ascomata on substrate. Note
ascomata surrounded by brown setae. c, d Vertical section of ascoma. e Peridium. f Ostiole g
Hamathecium. h, i Setae. j–n Asci. o–r Ascospore. s Hyphae in culture. t, u Colony on PDA from
above and below. Scale bars: a = 500 µm, c, d = 100 µm, e, f, h–n = 50 µm, g = 20 µm, o–r = 10
µm, s = 50 µm, t, u = 20 mm.
304
Figure 17 – Byssosphaeria siamensis (MFLU 17-1104). a, b Superficial ascomata on substrate.
Note ascomata surrounded by dark brown setae. c Ascoma. d Ostiole. e Peridium. f Setae. g Asci
with hamathecium. h Hamathecium. i–k Ascus. l–o Ascospore. p Germinating ascospore. q, r
Colony on PDA from above and below. Scale bars: a = 500 µm, b, c = 200 µm, d–f = 100 µm, g =
50 µm, h–k = 20 µm, l–p = 10 µm, q, r = 20 mm.
305
Pseudobyssosphaeria bambusae H.B. Jiang & K.D. Hyde, sp. nov.
Fig. 18
Index Fungorum Number: IF554470; Facesoffungi number: FoF 03921
Etymology – in reference to the host genus.
Holotype – MFLU 18-0151
Saprobic on rotting bamboo culms in terrestrial habitats. Sexual morph Ascomata 530–570
μm high, 470–525 μm diam. ( = 550 × 497 µm, n = 10), superficial, solitary, scattered or
gregarious, subglobose, black, with a central ostiole, ascomata covered with outwardly brown,
septate, branched hyphae. Setae 100–260 × 4.5–6 μm, dense, covering the whole ascoma, dark
brown, unbranched, septate. Peridium 65–85 μm wide, multi-layered, outer layer composed of
thick-walled, dark brown cells of textura angularis; inner layer composed of thin-walled, hyaline
cells of textura angularis. Hamathecium composed of 1–2 μm wide, septate, branched,
anastomosing pseudoparaphyses, embedded in a gelatinous matrix. Asci 125–160 × 8–11 μm ( =
143.5 × 9.5 μm, n = 20), 8-spored, unitunicate, cylindrical to clavate, short pedicellate, with furcate
to obtuse ends, apically rounded. Ascospores 29.5–32.7 × 4.6–5.6 μm ( = 31.1 × 5.1 μm, n = 20),
overlapping bi-seriate, fusiform with obtuse ends, 1-septate, constricted at the septum, hyaline,
normally 4-guttulate, straight, smooth-walled. Asexual morph Undetermined.
Culture characteristics – Ascospores germinating on PDA within 24 hours and germ tubes
developing from one end. Colonies slowly growing on PDA, reaching 20 mm in 4 weeks at 23 °C,
under 12 h light/12 h dark, flocculent, brown from above and below.
Material examined – THAILAND, Chiang Rai, near Fathai market, on dead bamboo culms,
11 March 2017, Hong-Bo Jiang, Fathai005 (MFLU 18-0151, holotype), ex-type living cultures,
MFLUCC 18-0145, KUMCC 17-0327; ibid. (HKAS 101463, isotype).
GenBank numbers – ITS: MG737555, LSU: MG737556, TEF: MG737557.
Notes – Although a LSU NCBI blast search places Pseudobyssosphaeria bambusae close to
Bertiella macrospora (IL 5005), the former differs from Bertiella species in having hyaline
ascospores. Pseudobyssosphaeria bambusae also has setose ascomata and forms a basal clade to
Bertiella sensu stricto and is thus cosidered a new genus and species.
Neocamarosporiaceae Wanas. et al.
Neocamarosporiaceae was introduced by Wanasinghe et al. (2017b) to accommodate
Dimorphosporicola and Neocamarosporium. We followed the classification of Wanasinghe et al.
(2017b) for species delineations. In this paper we introduce Neocamarosporium phragmitis as a
new species which was collected from the UK.
Neocamarosporium Crous & M.J. Wingf.
The genus Neocamarosporium was introduced based on Neocamarosporium goegapense
from South Africa, which is morphologically similar to the genus Camarosporium (Wanasinghe et
al. 2017b). In a recent study, Wanasinghe et al. (2017b) amended this genus to accommodate its
sexual morphs. Currently there are 14 accepted species in Neocamarosporium and we introduce the
fifteenth species herein.
Neocamarosporium phragmitis Wanas., E.B.G. Jones & K.D. Hyde, sp. nov.
Fig. 20
Index Fungorum number: IF554240; Facesoffungi number: FoF04099
Etymology − name reflect the host genus, Phragmites.
Saprobic on culms of Phragmites australis. Sexual morph Ascomata 150−200 μm high,
120−180 μm diameter ( = 180.6 × 147.6 µm, n = 10), black, semi-immersed, confluent,
gregarious, cupulate when dry, globose, uniloculate, with an apapillate ostiole. Ostiole central,
short, slightly sunken, minute and inconspicuous at the surface, smooth, ostiolar canal filled with
brown to hyaline cells. Peridium 10–25 µm wide at the base, 30–40 µm wide in sides, comprising
4–5 layers of reddish-brown to brown, cells, lighter and flattened towards the inside, of thin-walled
cells of textura angularis. Hamathecium comprising numerous, 2–3 µm (n = 40) wide,
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Figure 18 – Pseudobyssosphaeria bambusae (MFLU 18-0151, holotype). a–c Ascomata on
bamboo host. d Brown mycelium (surrounding ascomata). e Ascomata. f Peridium. g Ostiole. h
Pseudoparaphyses. i–k Asci. m, n Culture characters on PDA (note n reverse). o, p Ascospores. q
Germinating ascospores. Scale bars: b = 1000 μm, c = 500 μm, e = 150 μm, d, f, g = 50 μm, h–l =
30 μm, q = 20 μm, o, p = 10 μm.
307
Figure 19 – Phylogram generated from maximum likelihood analysis based on combined LSU,
SSU, ITS and TEF partial sequence data. Related sequences were obtained from Wanasinghe et al.
(2017b). Twenty-eight strains are included in the combined LSU, SSU, ITS and TEF sequence
analyses, which comprise 3,382 characters with gaps (857 for LSU, 982 for SSU, 572 for ITS and
953 for TEF). Single gene analyses were carried out and compared with each species, to compare
the topology of the tree and clade stability. Staurosphaeria rhamnicola (Coniothyriaceae) is used as
the outgroup taxon. Tree topology of the maximum-likelihood analysis was similar to the Bayesian
analysis. The best scoring RAxML tree with a final likelihood value of -7076.920933 is presented.
The matrix had 327 distinct alignment patterns, with 39.76% of undetermined characters or gaps.
Estimated base frequencies were as follows; A = 0.244371, C = 0.240432, G = 0.267282, T =
0.247915; substitution rates AC = 1.075762, AG = 2.163702, AT = 1.116814, CG = 0.758517, CT
= 5.979769, GT = 1.000000; gamma distribution shape parameter α = 0.756726. Bootstrap support
values for ML (first set) and BYPP equal to or greater than 0.95 are given above the nodes. The
newly generated sequence is in blue.
filamentous, branched, septate, pseudoparaphyses. Asci 60–90 × 14–17 µm ( = 75.9 × 15.4 µm, n
= 40), 8-spored, bitunicate, fissitunicate, cylindrical-clavate to cylindrical, short pedicellate (8–12
µm long), apex rounded with a minute ocular chamber. Ascospores 12–16 × 6–8 µm ( = 14.9 ×
7.2 µm, n = 50), overlapping biseriate, muriform, mostly ellipsoidal, with 3 transverse septa and
one longitudinal septum, deeply constricted at the middle septum, slightly constricted at remaining
septa, initially hyaline, becoming pale brown at maturity, upper part wider than lower part, slightly
paler, rounded at both ends, conical at the lower end. Asexual morph Undetermined.
308
Culture characteristics − Colonies on PDA reaching 2 cm diameter after 4 weeks at 16 °C,
later with dense mycelium, circular, rough margin, white, reverse cream-grey, flat on the surface,
without aerial mycelium. Hyphae septate, branched, hyaline, thin-walled.
Material examined − UK, Hampshire, Southsea, Eastney shore, on a washed-up culm of
Phragmites australis (Poaceae), 9 November 2015, EBG Jones GJ212 (MFLU 17-0585, holotype),
ex-type living culture MFLUCC 17-0756.
GenBank numbers − ITS: MG844345, LSU: MG844347, SSU: MG844349, TEF:
MG844351.
Notes − Neocamarosporium phragmitis is similar to N. lamiacearum in having globose
ascomata with an apapillate ostiole, cylindrical-clavate asci and muriform, ellipsoidal, with 3transversely septate, ascospores with one longitudinal septum, which are rounded at upper end and
conical at lower end. However, N. lamiacearum has thin peridium and larger asci comparatively to
N. phragmitis. To further support the establishment of the new taxon as proposed by Jeewon &
Hyde (2016) a comparison of the 522 ITS (+5.8S) nucleotides reveals 32 (6.13%) differences that
justify they are different taxa. We therefore identify our isolate (MFLUCC 17-0756) as
Neocamarosporium phragmitis sp. nov.
Figure 20 – Neocamarosporium phragmitis (MFLU 17-0585, holotype). a, b Ascomata on host
substrate. c Section of ascoma. d Pseudoparaphyses. e–g Asci. h–j Ascospores. Scale bars: a = 1
mm, b = 200 µm, c = 100 µm, d = 5 µm, e–g = 20 µm, h–i = 10 µm.
309
Neohendersoniaceae Giraldo & Crous
The family Neohendersoniaceae was introduced by Giraldo et al. (2017) with
Neohendersonia as the type genus. Tanaka et al. (2017) listed Brevicollum, Crassiparies,
Medicopsis and Neohendersonia as members in this family based on multi-gene phylogenetic
analysis. In this paper, a new host and country record for Brevicollum hyalosporum and a new
taxon, Medicopsis chiangmaiensis, are described, illustrated and compared with related taxa, based
on the evidence from morphology and phylogenetic analyses of combined LSU, RPB2, SSU and
TEF1 sequence data.
Figure 21 – Phylogram generated from maximum likelihood analysis based on combined LSU,
RPB2, SSU and TEF1 partial sequence data. Related sequences were obtained from Hyde et al.
(2016). Fifty-four strains are included in the combined LSU, RPB2, SSU and TEF1 sequence
analyses, which comprise 3,769 characters with gaps (986 for LSU, 973 for SSU, 907 for TEF and
310
903 for RPB2). Cyclothyriella rubronotata is used as the outgroup taxon. The best scoring RAxML
tree with a final likelihood value of -31102.781686 is presented. The matrix had 1716 distinct
alignment patterns, with 43.56% of undetermined characters or gaps. Estimated base frequencies
were as follows; A = 0.250598, C = 0.237272, G= 0.273530, T = 0.238600; substitution rates AC =
1.408199, AG = 4.058254, AT = 1.480490, CG = 1.063567, CT = 7.536065, GT = 1.000000;
gamma distribution shape parameter α = 0.204321. Bootstrap support values for ML (first set)
equal to or greater than 60% are given above/below the nodes. The newly generated sequences are
in red.
Brevicollum Kaz. Tanaka et al.
Tanaka et al. (2017) introduced Brevicollum to accommodate B. hyalosporum, characterized
by immersed ascomata with short ostiolar necks, a thin ascomatal wall, clavate asci with a shallow
ocular chamber and broadly fusiform, 3–5-septate ascospores. Currently, two species have been
reported, viz. Brevicollum hyalosporum and B. versicolor (Tanaka et al. 2017). Brevicollum is
morphologically similar to Crassiparies in ascospore characteristics. They are however, different,
as Brevicollum has 8-spored asci, while Crassiparies has 4-spored asci. They are also
phylogenetically apart (Tanaka et al. 2017). The ML tree generated based on analysis of the
combined LSU, RPB2, SSU and TEF1 sequence data indicates that our isolate of Brevicollum
hyalosporum, B. hyalosporum type (MAFF 243400) and B. versicolor group as a monophyletic
subclade sister to Amarenographium solium with moderate support (63% ML) and nested in
between Crassiparies and Neohendersonia (Fig. 21). Unfortunately, no sexual morph has been
reported for A. solium (Hodhod et al. 2012).
Fig. 22
Brevicollum hyalosporum Kaz. Tanaka & Toy. Sato, Mycologia 109: 4 (2017)
Facesoffungi number: FoF 03899
Saprobic branch of Hevea brasiliensis Müll.Arg. Sexual morph Ascomata 200–300 µm
diameter, 100–250 µm high, scattered, sometimes clustered, immersed, globose to subglobose,
ostiole central. Peridium 13–45 μm wide, composed of two type of cell layers, outer layer
comprising 2–4 layers of thick-walled, dark brown to black cells of textura angularis, inner layer
comprising 2–3 layers of thin-walled, hyaline cells of textura angularis. Hamathecium 2–3 μm
wide, comprising dense, hyaline, septate, branched, filamentous pseudoparaphyses, slightly
constricted at septa. Asci (90–)92–120(–155) × 17–22(–25) µm (x̅ = 110 × 21 μm, n = 10), 8spored, bitunicate, fissitunicate, cylindrical to clavate, subsessile to short pedicellate,apically
rounded, with indistinct ocular chamber. Ascospores (23–)29–43(–44) × (3–)7–12(–13) µm (x̅ =
38.4 × 10.5 μm, n = 30), uniseriate at the lower part and overlapping biseriate at the upper part of
the ascus, hyaline to sub-hyaline, broadly fusiform, thick-walled, straight to straightly curved,
initially aseptate with small guttules, becoming 3–5-septate, deeply constricted at the middle
septum, slightly constricted at the remaining septa, median cell slightly broader, each cell
containing distinct large guttule, smooth-walled, with a mucilaginous sheath. Asexual morph
Undetermined.
Culture characteristics – Ascospores germinated on MEA within 24 hours and germ-tube
produced from one or both sides of the ascospore. Colonies on MEA reaching 2 cm diameter after
21 days at 25°C, colonies circular, medium dense, surface smooth with edge entire, velvety to
woolly, colony from above white at the margin, with white-greyish to grey in the centre; from
below: white brown to light brown at the margin, white-greyish to grey in the middle, dark grey to
blackish at the centre; not producing pigmentation in agar.
Material examined – THAILAND, Chiang Rai Province, Wiang Chiang Rung District, on
branch of Hevea brasiliensis (Euphorbiaceae), 1 November 2016, Chanokned Senwanna,
RBCR012 (MFLU 17-1978), living culture, MFLUCC 17-0071.
GenBank numbers – ITS: MG602204, LSU: MG602200, SSU: MG602202, TEF1:
MG739516.
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Figure 22 – Brevicollum hyalosporum (MFLU 17-1978, new host record). a, b Longitudinal
sections through ascomata. c Peridium. d Paraphyses. e–g Asci. h Germinated ascospore. i, j
Culture characteristic on MEA after 21 days (i = colony from above, j = colony from below). k
Ascospores. l Arrowheads indicate gelatinous sheath surrounding ascospores after treating with
India ink. Scale bars: a, b = 200, c–h, k, l = 30 µm.
312
Notes – During our investigation on diversity of microfungi in Thailand, an isolate
(MFLUCC 17-0071) was recovered from Hevea brasiliensis in Chiang Rai Province. The
phylogeny inferred using analysis of LSU, RPB2, SSU and TEF1 sequence data showed that our
isolate clustered with Brevicollum hyalosporum with strong bootstrap support (100% ML, Fig. 21).
The comparisons of ITS sequence data show 100% similarity with Brevicollum hyalosporum
(Tanaka et al. 2017). Morphological characters of our collection (Fig. 22) also resembles those in
Tanaka et al. (2017). Brevicollum hyalosporum was introduced from Japan on dead twigs of
Syzygium samarangense (Myrtaceae) (Tanaka et al. 2017). Therefore, we consider the new
collection as a new host record and new record for Thailand.
Medicopsis Gruyter
Medicopsis is a monotypic genus introduced to accommodate Pyrenochaeta romeroi by de
Gruyter et al. (2013), although it was originally established for a coelomycetous taxon. Multigene
phylogeny from Tanaka et al. (2017) indicates that Medicopsis belongs to Neohendersoniaceae in
Pleosporales. Only one species Medicopsis romeroi is included in Medicopsis (Index Fungorum
2018). Members of this genus can be found as a pathogen of humans and also can occur on plants
(Badali et al. 2010, Khan et al. 2011, de Gruyter et al. 2013, Ahmed et al. 2014).
Medicopsis chiangmaiensis Q.J. Shang & K.D. Hyde, sp. nov.
Fig. 23
Index Fungorum number: IF553978; Facesoffungi number: FoF 03875
Etymology – the specific epithet “chiangmaina’’ refers to the province in Thailand from
which the holotype was collected.
Holotype –MFLU 17-1960
Saprobic on bark. Sexual morph Stromata with poorly developed interior, scattered or in
groups, immersed to erumpent from the bark by ostiolar canal, circular to irregular in shape.
Ascomata 295–530 µm high, 410–770 µm diameter ( = 410 × 590 µm, n = 25), perithecial,
immersed in a stroma, solitary or gregarious, dark brown to black, globose to subglobose, ostiolate,
with cylindrical necks, periphysate. Peridium 30–55 µm wide, composed of two section layers,
outer section comprising 5–10 layers of relatively small, dark brown to black, thick-walled cells,
arranged in textura globulosa to textura angularis, inner part comprising 5–8 layers of hyaline cells
of textura angularis. Hamathecium of 1–2 µm wide, dense, filamentous, septate, hyaline,
pseudoparaphyses which branch and anastomose frequently between and above asci. Asci (59–)90–
137(–178) × (10.5–)12–16(–20) µm ( = 115 × 14 µm, n = 50), 8-spored, bitunicate, fissitunicate,
clavate, with long pedicel and indistinct ocular chamber. Ascospores (16–)18–21(–23.5) × (6–)7.5–
9.5(–10) µm ( = 20 × 8.5 µm, n = 145), obliquely uniseriate and partially overlapping to biseriate,
fusoid to fusoid-ellipsoidal, with narrowly rounded ends, brown when mature, some lightly curved,
1- to 3-septate, with dark bands at the septa, constricted at the median septum, the upper cell often
broader than the lower one, minutely verrucose, guttulate, smooth-walled, without a mucilaginous
sheath. Asexual morph Undetermined.
Culture characteristics – Ascospores germinating on PDA within 24 hours. Germ tubes
produced from all sides. Colonies on PDA reaching 5–6.5 cm diameter after 7 days at room
temperature, colonies circular to irregular, medium dense, flat or effuse, slightly raised, with edge
fimbriate, fluffy to fairy fluffy, white from above and below; not producing pigments in agar.
Material examined – THAILAND, Chiang Mai Province, on undetermined dead wood, 27
January 2017, Qiuju Shang, M7P4-15 (MFLU 17-1960, holotype), ex-type living culture,
MFLUCC 17-2457, KUMCC 17-0327; ibid. (HKAS 100867, isotype).
GenBank numbers – ITS: MG873485, LSU: MG873481, SSU: MG873483.
Notes – Medicopsis chiangmaiensis fits well with the classical description of Pleosporales
and is characterized by pseudoparaphyses, ostiolate ascomata with bitunicate asci and single-celled
ascospores (Zhang et al. 2012). We cannot compare our sexual morphs with other species of
Medicopsis as this genus lacks a known sexual morph (de Gruyter et al. 2013). However,
313
Figure 23 – Medicopsis chiangmaiensis (MFLU 17-1960, holotype). a Host. b Stroma on host
surface. c Section of stroma. d Section of ostiole. e Section of peridium. f–i Asci. j
Pseudoparaphyses. k–q Ascospores. r Germinating ascospore. Scale bars: c = 200 μm, d = 50 μm,
e, l–r = 10 μm, f–k = 20 μm.
314
phylogenetic analyses based on the combined LSU, SSU, TEF1 and RPB2 sequence data (Fig. 21)
show that M. chiangmaiensis (MFLUCC 17-2457) forms a distinct lineage from M. romeroi with
moderate support (60% in ML).
Neophaeosphaeriaceae Ariyaw. & K.D. Hyde
Neophaeosphaeriaceae was introduced by Ariyawansa et al. (2015) to accommodate
Neophaeosphaeria and its allied species in the suborder Pleosporineae, order Pleosporales,
Dothideomycetes (Ariyawansa et al. 2015). Currently, Neophaeosphaeria is the only genus
accommodated in this family (Wijayawardene et al. 2018). Detailed taxonomic notes and revisions
are provided in Ariyawansa et al. (2015) for this family. Herein, an updated tree for the family is
provided (Fig. 24) and a new species, Neophaeosphaeria phragmiticola, is introduced.
Figure 24 – Phylogram generated from maximum likelihood analysis based on combined LSU,
SSU, ITS and TEF sequenced data of Neophaeosphaeriaceae. Related sequences were obtained
from Ariyawansa et al. (2015). 23 strains are included in the combined sequence analyses, which
comprise 3256 characters with gaps. Single gene analyses were carried out and compared with each
species, to compare the topology of the tree and clade stability. Shiraia bambusicola (NBRC
30753) and Shiraia bambusicola (NBRC 30772) are used as the outgroup taxa. Tree topology of
the ML analysis was similar to the BI. The best scoring RAxML tree with a final likelihood value
of -8537.368649 is presented. The matrix had 490 distinct alignment patterns, with 32.23%
undetermined characters or gaps. Estimated base frequencies were as follows; A = 1.000000, C =
0.244434, G= 0.235548, T = 0.267643; substitution rates AC = 0.503206, AG = 1.237779, AT =
2.388948, CG = 2.835816, CT = 0.417870, GT = 9.060268; gamma distribution shape parameter α
= 0.512703. Bootstrap support values for ML equal to or greater than 65% and BYPP equal to or
greater than 0.90 are given above the nodes respectively. Ex-type strains and reference strains are in
bold, newly generated sequences are in red.
315
Neophaeosphaeria M.P.S. Câmara et al.
Neophaeosphaeria was introduced by Câmara et al. (2003) to accommodate N. filamentosa,
N. barrii, N. conglomerata and N. quadriseptata. Currently, five species are known
(Wijayawardene et al. 2017a). The asexual morphs are coelomycetous and species are pathogenic
or saprobic in terrestrial habitats (Wijayawardene et al. 2017a).
Neophaeosphaeria phragmiticola A. Karunarathna & K.D. Hyde, sp. nov.
Fig. 25
Index Fungorum number: IF554111; Facesoffungi number: FoF 03942
Etymology – named after Phragmites, the host genus from which it was collected.
Holotype – HKAS 97353
Saprobic on dead stems of Phragmites australis (Cav.) Trin. ex Steud. Sexual morph
Undetermined. Asexual morph Coelomycetous, Conidiomata 69–76 µm high × 74–79 µm diameter
( = 73.3 × 76.4 µm, n = 5), solitary, immersed to slightly erumpent, dark brown to black, globose
to subglobose, slightly depressed, unilocular, lacking ostiole. Conidiomatal wall 12–14 μm wide,
composed of 3–4 cell layers of thin-walled, brown to dark brown pseudoparenchymatous cells of
textura angularis, hyaline towards the inner layers. Conidiophores reduced to conidiogenous cells.
Conidiogenous cells 15–13 µm high × 1–1.6 µm wide ( = 14 × 1.3 µm, n = 10), enteroblastic,
phialidic, integrated to discrete, branched, doliiform, lageniform or cylindrical, smooth, hyaline,
thin-walled, arising from the basal cavity. Conidia 32–37 × 17–20 µm ( = 34 × 18 µm, n = 40),
ellipsoidal to obovoid, truncate at the base, obtuse at the apex, muriform at maturity, hyaline when
immature and brown to dark brown at maturity, with 4 transverse septa and 1 longitudinal septum
between 1–2 and 2–3, continuous or constricted at the septa, rough-walled.
Culture characteristics – Conidia germinating on PDA, within 14 hours. Colonies growing on
PDA, cottony, white to gray reaching 5 mm in 20 days at 25ºC, mycelium superficial, effuse, with
regular edge, hyphae pale yellow to white.
Material examined – CHINA, Yunnan Province, Erhai Lake, on dead stems of Phragmites
australis (Cav.) Trin. ex Steud, 5 October 2016, K.D Hyde, AKDF 05 (HKAS 97353, holotype),
ex-type living culture, MFLUCC 18-0449, KUMCC 16–0216.
GenBank numbers – LSU: MG837009, RPB2: MG838021, SSU: MG837008, TEF1:
MG838020.
Notes – The phylogenetic affinities of Neophaeosphaeria phragmiticola to other species is
not well-resolved. A close relationship to N. quadriseptata and N. barrii is observed but without
support (Fig. 24). However, this is the first time muriform spores are reported in Neophaeosphaeria
and hence we describe our taxon as a new species.
Occultibambusaceae D.Q. Dai & K.D. Hyde
The family Occultibambusaceae was introduced by Dai et al. (2016) with Occultibambusa as
the type genus. The family is characterized by immersed, solitary to gregarious ascomata, with
black ostioles, broadly cylindrical to clavate, bitunicate asci, and broad-fusiform, hyaline to dark
brown ascospores (sexual morph) and asexual morphs are morphologically diverse (Dai et al.
2017). This family comprises four genera including Seriascoma, Occultibambusa (Dai et al. 2017),
Versicolorisporium (Hatakeyama et al. 2008) and Neooccultibambusa (Doilom et al. 2017). In this
paper, we introduce a new Neooccultibambusa species from Pandanaceae habitats with support
from combined multi-loci phylogenetic analyses (Fig. 26).
Neooccultibambusa Doilom & K.D. Hyde
The genus Neooccultibambusa is typified by N. chiangraiensis which is characterized by
sexual morphs having cylindrical to subcylindrical asci, and ascospores with 1–3 transverse septa.
The asexual morph of this genus produces chlamydospores in culture (Doilom et al. 2017) and it
has circular, floccose, dark brown to black colonies, with subglobose to globose, dark brown
conidia (Jayasiri et al. 2016). In this study, we introduce a new species Neooccultibambusa
pandanicola which is the hyphomycetous asexual morph.
316
Figure 25 – Neophaeosphaeria phragmiticola (HKAS 97353, holotype). a, b Appearance of
conidiomata on host. c Longitudinal section of conidioma. d Conidioma wall. e–f Conidiogenous
cells and developing conidia. g–j Conidia. k Germinated conidium. l–m Culture characteristics on
PDA (l = from above, m = from below). Scale bars: c = 50 µm, d = 10 µm, e–f = 10 µm, g–k = 5
µm.
Neooccultibambusa pandanicola Tibpromma, D.J. Bhat & K.D. Hyde, sp. nov.
Fig. 27
Index Fungorum number: IF553942; Facesoffungi number: FoF03814
Etymology – refers to the name of the host plant which the fungus was collected.
Holotype – HKAS 99623
Saprobic on dead leaves of Pandanus utilis Bory. Colonies dark olive-green, erumpent.
Mycelium comprising hyaline to pale brown, branched, septate, thin-walled hyphae. Sexual morph
Undetermined. Asexual morph Hyphomycetous. Conidiophores 13–71 µm high × 3.5–7 wide µm
( = 36.8 × 5.4 µm, n = 10) diameter, macronematous, mononematous, solitary, pale-brown to
brown, cylindrical, 3–5-septate, straight or slightly flexuous, erect, guttulate, smooth-walled.
Conidiogenous cells 2.5–5.5 × 4–5.5 µm ( = 3.1 × 3.6 µm, n = 10), holoblastic, monoblastic,
integrated, terminal, brown, determinate, cylindrical. Conidia 28–150 × 7–21 µm ( = 86.2 × 12.8
µm, n = 10), acrogenous, solitary, olivaceous brown to mid-brown, obclavate, rostrate at apex,
straight or curved, 7–17-euseptate, wide at broadest middle part, truncate at the base, some with
basal scar, without with a mucilaginous sheath.
Culture characteristics – Conidia germinating on PDA within 24 hours. Germ tubes
produced from both ends. Colonies on PDA reaching 9 cm diameter after 2 weeks at room
temperature, greyish-green on the surface, with dense, fluffy mycelium, circular colony with entire,
raised on the surface, dark green in reverse, with smooth margin.
317
Figure 26 – Phylogram generated from maximum likelihood analysis based on combined LSU,
SSU, TEF1 and RPB2 partial sequence data. Related sequences were obtained from Zhang et al.
(2017a). Fifty-four strains are included in the combined LSU, SSU, TEF1 and RPB2 sequence
analyses, which comprise 3,808 characters with gaps (848 for LSU, 1019 for SSU, 935 for TEF and
1006 for RPB2). Single gene analyses were carried out and compared with each species, to
compare the topology of the tree and clade stability. Melanomma pulvis-pyrius is used as the
outgroup taxon. Tree topology of the maximum-likelihood analysis was similar to the Bayesian
analysis. The best scoring RAxML tree with a final likelihood value of -21677.689087 is presented.
The matrix had 1126 distinct alignment patterns, with 35.23% of undetermined characters or gaps.
Estimated base frequencies were as follows; A = 0.251517, C = 0.243434, G= 0.276276, T =
0.228772; substitution rates AC = 1.477547, AG = 5.073666, AT = 1.504758, CG = 1.082277, CT
= 9.409453, GT = 1.000000; gamma distribution shape parameter α = 0.159759. Bootstrap support
values for ML (first set) and BYPP equal to or greater than 0.80 are given above the nodes. The
newly generated sequence is in red.
318
Figure 27 – Neooccultibambusa pandanicola (HKAS99623, holotype). a Colonies on the
substratum. b, c Conidiophores and conidia. d–f Conidia. g Germinating conidium. h, i Cultures on
PDA, h from above, i from below. Scale bars: b–g = 10 μm.
319
Material examined – CHINA, Yunnan, Xishuangbanna Tropical Botanical Garden, on fallen
dead and decaying leaves of Pandanus utilis Bory. (Pandanaceae), 28 April 2017, Rungtiwa
Phookamsak and Nimali Indeewari de Silva, XTBG15 (HKAS 99623, holotype), extype living
culture, KMUCC 17-0179, MFLUCC 17-2265; ibid. (MFLU, isotype).
GenBank numbers – ITS: MG298941, LSU: MG298940, RPB2: MG298944, SSU:
MG298942, TEF1: MG298943.
Notes – The new species, Neooccultibambusa pandanicola, is introduced based on multigene
analysis of LSU, SSU, TEF1 and RPB2 sequence data. The new species is phylogenetically closely
related to N. jonesii Jayasiri et al. with moderate support (74% in ML, 0.87 in PP) (Fig. 26). We
also compared the morphology of the asexual morph of our new taxon with N. jonesii which has
subglobose to globose, unicellular, dark brown conidia (Jayasiri et al. 2016).Our new taxon is
different in having obclavate, olivaceous brown to mid-brown, 3–17-euseptate conidia, which are
widest at the middle part.
Periconiaceae (Sacc.) Nann.
The family Periconiaceae was introduced by Nannizzi (1934) with Periconia as the type
genus. Periconiaceae has long been ignored in modern fungal systematics and Tanaka et al. (2015)
reinstated Periconiaceae in the suborder Massarineae based on phylogenetic analyses. In this paper,
we introduce one new species, Periconia elaeidis, designate a reference specimen for Periconia
cookei E.W. Mason & M.B. Ellis and report Periconia pseudobyssoides S. Markovskaja & A.
Kačergius from China for the first time.
Periconia Tode
The genus Periconia was introduced by Tode (1791) with P. lichenoides Tode as the type
species. There are 187 species epithets in Index Fungorum (2018) with more than 20 transferred to
other genera, but currently only ca. 50 species are recognized as genuinely belonging to Periconia
(Carmarán & Novas 2003, Kirk et al. 2008, Seifert et al. 2011, Markovskaja & Kačergius 2014,
Tanaka et al. 2015, Chuaseeharonnachai et al. 2016, Hyde et al. 2017b, Liu et al. 2017a,
Thambugala et al. 2017).
Periconia cookei E.W. Mason & M.B. Ellis, Mycol. Pap. 56: 72 (1953)
Fig. 29
Facesoffungi number: FoF 03856
Saprobic on submerged decaying wood. Sexual morph Undetermined. Asexual morph
Colonies on natural substrate effuse, scattered, hairy, brown to dark brown. Mycelium mostly
immersed, branched, septate, smooth, brown. Conidiophores mononematous, macronematous,
cylindrical, erect, straight or flexuous, septate, dark brown at the base and gradually paler towards
the apex, unbranched, 240–372 μm ( = 306 μm, SD = 66, n = 10) long, 15.5–19.5 μm ( = 17.5
μm, SD = 2, n = 10) wide. Conidiogenous cells polyblastic, discrete, sphaerical, terminal, brown.
Conidia catenate, verrucose, brown, mostly sphaerical, dry, thin-walled, 12–15 μm ( = 13.5 μm,
SD = 1.5, n = 35) diameter.
Material examined – CHINA, Yunnan Province, saprobic on decaying wood submerged in
stream in Cangshan Mountain, July 2016, S.M. Tang, S-835 (MFLU 17-1969, reference specimen
designated here); living culture, MFLUCC 17-1679, ICMP; S-812 (MFLU 17-1972); living culture,
MFLUCC 17-1399, ICMP.
GenBank numbers – MFLUCC 17-1679 – LSU: MG333492, TEF1: MG438278; MFLUCC
17-1399 – ITS: MG333490, LSU: MG333493, TEF1: MG438279.
Notes – Periconia cookei was introduced by Mason & Ellis (1953) and is characterized by
unbranched, septate, pale brown to dark brown, polyblastic, conidiophores, catenate, verrucose,
brown, mostly sphaerical conidia, mostly 13–16 μm diameter (Mason & Ellis 1953). Two fresh
isolates were obtained from submerged wood in a stream in Cangshan Mountain during our study
of lignicolous freshwater fungi in Yunnan Province, China. The morphological characters of our
fresh collection especially the shape and diameter of conidia fit well with the description of P.
320
cookei. Thus, we identified our fresh isolate as P. cookei and provided sequence data for this
species. Ariyawansa et al. (2014) suggested that if some of morphological characters of the fungus
being studied differ from those in the original description, the original material exists but cannot be
examined, or its location is different, or the host differs from the holotype, or if no sequences can
be obtained from an otherwise satisfactory existing type material, we can designate a reference
specimen (RefSpec) in order to clarify the placement of the species using morphology coupled with
molecular data. The morphological characters of our new collections fit well with the holotype of
Periconia cookei Mason & Ellis (1953), and since we could not obtain material from the same host
or location, a reference specimen is designated here.
Figure 28 – Phylogram generated from maximum likelihood analysis based on combined ITS, LSU
and TEF sequence data from selected species of Periconiaceae, Massarinaceae,
Didymosphaeriaceae and Lentitheciaceae. Maximum likelihood and maximum parsimony
bootstrap support values greater than 75% are shown in above and below and branches, while
Bayesian posterior probabilities greater than 0.95 are in bold. The new isolates are in red. The tree
is rooted with Morosphaeria ramunculicola (KH 220) and M. velatispora (KH 221). Type, ex-type,
epitype and reference strains are in bold.
321
Figure 29 – Periconia cookei (MFLU 17-1969, reference specimen) a Colonies on wood. b
Conidiophore with conidia. c Conidiogenesis cells and conidia. d–i Conidia. j Germinating
conidium. g, k Culture on PDA, g from above, k from below. Scale bars: b = 100 μm, c = 30 μm,
d–j = 15 μm.
322
Periconia elaeidis T. Sunpapao & K.D. Hyde, sp. nov.
Fig. 30
Index Fungorum number: IF554399; Facesoffungi number: FoF04473
Etymology – name reflects the host genus Elaeis.
Holotype – MFLU 18−0626
Saprobic on dead oil palm leaves. Asexual morph Colonies on substrate numerous, effuse,
dark brown to black. Conidiophores 200−400 μm long × 8−10 μm wide, macronematous,
mononematous, unbranched, erect, straight or slightly flexuous, single, grayish-brown to dark
brown, 4−7-septate, smooth to minutely verruculose, thick-walled. Conidiogenous cells polyblastic,
proliferating, terminal, integrated or discrete, ovoid to globose, pale brown, smooth, collapsing
when old. Conidia 4.5−6.5 μm diameter ( = 5.5 μm, n = 30), solitary, subglobose to globose,
subhyaline to pale brown, verruculose, aseptate. Sexual morph Undetermined.
Culture characters – Conidia germinating on PDA within 24 hours. Colonies reaching about 6
cm diameter in 5 days on PDA with sparse white mycelia on the surface. The reverse of colony
with a dark in the center with a white margin.
Material examined – THAILAND, Phrae, on decaying oil palm, 28 January 2017, Thanawat
Sunpapao ESL-1 (MFLU 18−0626, holotype), ex-type living culture MFLUCC 17−0087.
GenBank number – ITS: MG742713, LSU: MH108552, SSU: MH108551.
Notes – Periconia elaeidis resembles P. cookei in having macronematous, mononematous,
unbranched, erect conidiophores, polyblastic, terminal conidiogenous cells and verruculose conidia.
However, Periconia elaeidis differs from P. cookei in having thinner conidiophores (8−10 μm vs
15.5−19.5 μm) and smaller conidia (4.5−6.5 μm vs 12−15 μm). The phylogenetic analysis based on
combined ITS, LSU and TEF1 sequence data also indicate that they are distinct species.
Periconia pseudobyssoides S. Markovskaja & A. Kačergius, Mycological Progress 13 (2): 293
(2014)
Fig. 31
Facesoffungi number: FoF 03857
Saprobic on submerged decaying wood. Sexual morph Undetermined. Asexual morph
Colonies effuse, scattered, hairy, dark brown to black. Mycelium mostly immersed but sometimes
partly superficial, septate. Conidiophores mononematous, macronematous, straight or flexuous,
solitary, unbranched, often verruculose, cylindrical, brown, 277–411 μm ( = 344 μm, SD = 67, n
= 10) long, 9–13 μm ( = 11 μm, SD = 2, n = 10) wide. Conidiogenous cells polyblastic,
holoblastic, determinate, integrated, brown, cylindrical. Conidia sphaerical or subsphaerical, golden
yellow to golden brown, verruculose, 5.5–7.5 μm ( = 6.5 μm, SD = 1, n = 35) diameter
Material examined – CHINA, Yunnan Province, saprobic on decaying wood submerged in
stream in Cangshan Mountain, September 2016, S.M. Tang, S-850 (MFLU 17-1970); living
culture, DLUCC 0850.
GenBank numbers – ITS: MG333491, LSU: MG333494, TEF1: MG438280.
Notes – Periconia pseudobyssoides was introduced by Markovskaja and Kačergius (2014)
and is characterized by macronematous, often verruculose, brown to reddish-brown conidiophores,
sphaerical, golden yellow to golden brown or reddish-brown conidia (Markovskaja & Kačergius
2014). A fresh isolate was obtained from submerged wood in a stream in Cangshan Mountain
during our study of lignicolous freshwater fungi in Yunnan Province, China. The morphological
characters of our fresh collection fit well with the description of P. pseudobyssoides. Phylogenetic
analysis also indictes that our strain clusters with P. pseudobyssoides (MAFF 243868, MAFF
243874) with strong support (93 ML/1.00 PP). Thus, we identified our fresh isolate as P.
pseudobyssoides and provide sequence data for this species. This is the first record of this fungus
for China.
Phaeoseptaceae S. Boonmee, Thambugala & K.D. Hyde, fam. nov.
Index Fungorum number: IF 554385; Facesoffungi number: FoF04462
Saprobic on dead or decaying wood in terrestrial and freshwater habitats. Sexual morph
323
Figure 30 – Periconia elaeidis (MFLU 18−0626, holotype). a Substrate. b Conidiophores on
substrate. c–e Conidiophores. f–i Conidia. j Germinated conidia. k Colony on MEA. Scale bars: b =
100 μm, c–e = 50 μm, f–i = 5 μm, j = 10 μm.
Ascomata immersed in host tissues, erumpent when mature, visible as black spots, solitary,
scattered, globose to subglobose, dark brown, with or without a pseudoclypeus, short papillate, with
an apical ostiole. Peridium comprising several layers, outer layers dark brown to black, of
somewhat flattened cells of textura angularis; inner layers hyaline to lightly pigmented cells of
textura angularis. Hamathecium comprising cylindrical, branched, septate, anastomosed,
324
Figure 31 – Periconia pseudobyssoides (MFLU 17-1970). a, b Colonies on wood. c, d
Conidiophore with conidia. e, f Conidiogenous cells and conidia. g–i Conidia. j Germinating
conidium. Scale bars: c, d = 100 μm, e–f = 30 μm, g–m = 15 μm.
325
pseudoparaphyses. Asci 8-spored, bitunicate, cylindrical-clavate, long pedicellate, with a small
ocular chamber. Ascospores 2–(-3)-seriate, light brown, muriform, allantoid, cylindrical, broadly
fusoid to broadly tapering towards the rounded ends, slightly curved, ends asymmetrical, slightly
wider at median part, rounded at both ends, with multi-transverse septa, and 1 longitudinal septum
in each cell, sometimes with 2 longitudinal septa, constricted and darkened at the septa, smoothwalled. Asexual morph Undetermined.
Family type – Phaeoseptum Ying Zhang, J. Fourn. & K.D. Hyde
Notes – The new family Phaeoseptaceae is introduced to accommodate the genera
Phaeoseptum, Lignosphaeria Boonmee et al. and Neolophiostoma Boonmee & K.D. Hyde. In the
present phylogenetic analysis of LSU, SSU, RPB2 and TEF1α sequence data Phaeoseptum,
Lignosphaeria and Neolophiostoma form a well-supported clade (Fig. 32) together with putatively
named strains of Decaisnella formosa (BCC 25616 and BCC 25617) and Thyridaria
macrostomoides (GKM 1033 and GKM 1159), sister to the family Halotthiaceae (Fig. 32).
However, Decaisnella formosa and Thyridaria macrostomoides need to be recollected and
epitypified with DNA sequence data in order to ensure correct placement (Abdel-Wahab and Jones,
2003, Mugambi and Huhndorf 2009). Lignosphaeria and Neolophiostoma are characterized by
phragmosporous, elongate, cylindrical to fusiform and hyaline ascospores and Phaeoseptum mainly
differs from Lignosphaeria and Neolophiostoma in having muriform, allantoid, cylindrical,
pigmented ascospores. All of the species of these genera are wood inhabiting taxa collected in
Thailand (Zhang et al. 2013, Ariyawansa et al. 2015, Thambugala et al. 2015).
Phaeoseptum Ying Zhang et al.
Phaeoseptum was introduced by Zhang et al. (2013) to accommodate a single species P.
aquaticum Ying Zhang et al. Zhang et al. (2013) placed this genus in Halotthiaceae based on
analysis of LSU sequence data. The genus is characterized immersed ascomata with a black
pseudoclypeus, bitunicate, broadly fusoid asci with broadly rounded ends and multi-transversally
septate, brown ascospores, which are darkly pigmented at the septa. In this study, we provide an
updated phylogenetic tree for the new family Phaeoseptaceae and introduce a new species of
Phaeoseptum (Fig. 32).
Phaeoseptum terricola S. Boonmee & K.D. Hyde, sp. nov.
Figs 33, 34
Index Fungorum number – IF554376, Facesoffungi number: FoF 04384
Etymology – the epithet terricola (Latin) means living on terrestrial ecosystems.
Holotype – MFLU 10-0032
Saprobic on dead wood. Sexual morph Ascomata 137–214 µm high × 155–224 µm diameter
( = 172 × 183.5 µm, n = 5), initially immersed and erumpent when mature, visible as black spots,
solitary, scattered, globose to subglobose, dark brown, short papillate, with an apical ostiole.
Peridium 14–24 µm wide, comprising several layers, outer layers dark brown to black, somewhat
flattened cells of textura angularis; inner layers hyaline to lightly pigmented cells of textura
angularis. Hamathecium comprising 0.5–1 µm wide, cylindrical, branched, septate, anastomosed,
pseudoparaphyses. Asci 64–90 × 13–17.5 µm ( = 79 × 16 µm, n = 15), 8-spored, bitunicate,
fissitunicate, cylindrical-clavate, long pedicellate, with a small ocular chamber. Ascospores 19–25
× 5–7 µm ( = 22 × 6 µm, n = 20), 2-seriate, light brown, muriform, allantoid, cylindrical, tapering
towards the ends, slightly curved, ends asymmetrical, slightly wider at median part, rounded at both
ends, with 9–10(–11) transverse septa, and 1 longitudinal septum in each cell, sometimes with 2
longitudinal septa, constricted and darkened at the septa, smooth-walled. Asexual morph
Undetermined.
Culture characteristics – Ascospores germinating on MEA within 12 hours and germ tubes
produced from each cell. Colonies slow growing on MEA, reaching less than 8 mm in 7 days at
28°C, slightly convex, undulating to raised, dentate, with slightly radial striations and lobate edges,
brown.
326
Figure 32 – RAxML phylogenetic tree generated from LSU, SSU and TEF1 sequence data of
Phaeoseptum terricola (strain MFLUCC 10–0102) and related genera of Pleosporales. The tree is
rooted with Melanomma pulvis-pyrius CBS 124080 (Melanommataceae, Pleosporales). RAxML
bootstrap support values ≥ 50% (BT) and Bayesian posterior probabilities ≥ 95% (PP) are given
above the nodes. The ex-type strains are in bold and new strain in blue.
327
Figure 33 – Phaeoseptum terricola (MFLU 10–0032, holotype). a Herbarium material. b
Appearance of ascomata on host. c Section of ascoma. d Peridium. e Pseudoparaphyses. f–i
Immature and mature asci. j–m Ascospores. Scale bars: a = Material, b = 500 µm, c = 100 µm, d–i
= 20 µm, j–m = 10 µm.
Material examined – THAILAND, Chiang Mai, Doi Inthanon, Jom Thong, elev. 800–1000
msl., N18°31.576' E 98°29.790', on dead wood, 5 September 2009, S. Boonmee, ITN01 (MFLU10–
0032, holotype), ex-type living culture: MFLUCC 10–0102, IFRDCC 2183, BCC 52150.
GenBank numbers – ITS – MH105778, LSU – MH105779, RPB2 – MH105782, SSU –
MH105780, TEF1 – MH105781.
Notes – Phaeoseptum terricola is introduced as a new species in the genus Phaeoseptum
based on morphology and phylogenetic evidence. In the present multi-gene phylogenetic analysis
328
(LSU, SSU and TEF1 sequence data), Phaeoseptum terricola clusters with the ex-type strain of P.
aquaticum with strong support (100% MLBT and 1.00 BYPP, Fig. 32). Phaeoseptum terricola
shares common features of the genus, such as dark pigmented and immersed ascomata, cylindricalclavate, pedicellate asci and muriform, brown ascospores. However, Phaeoseptum aquaticum
differs from P. terricola in having elongate, immersed ascomata with a pseudoclypeus, clearly
fissitunicate asci and allantoid, cylindrical ascospores, with multi-transverse septa and a single
longitudinal septum in each cell.
Figure 34 – Phaeoseptum terricola (MFLUCC 10-0102, ex-type culture). a Germinating spore. b, c
Colonies cultures on MEA from surface and reverse. d Aerial hyphae in culture. e, f The formation
of chlamydospore-like structures. Scale bars: a = 10 µm, b, c = 10 mm, d–f = 20 µm.
Phaeosphaeriaceae M.E. Barr.
The family Phaeosphaeriaceae was introduced by Barr (1979) with the type species
Phaeosphaeria oryzae. Members of this family are characterized by immersed to superficial,
globose to subglobose ascomata, bitunicate asci and hyaline, yellowish or brown, fusiform to
ellipsoidal, muriform, 1 to multi-septate ascospores, and holoblastic or enteroblastic conidiogenesis
with aseptate or septate conidia (Barr 1979, Zhang et al. 2012, Hyde et al. 2013, Phookamsak et al.
2014b, Tennakoon et al. 2016, Tibpromma et al. 2017). Phaeosphaeriaceae is recognized as a
heterogeneous group, comprising plant pathogens, saprobes and endophytes associated with a wide
range of host plants (Zhang et al. 2012, Hyde et al. 2013, Phookamsak et al. 2014b, 2017,
Tibpromma et al. 2017). According to recent publications, 52 genera are accepted within this
family (Phookamsak et al. 2014b, 2017, Hyde et al. 2016, Tibpromma et al. 2017, Wijayawardene
et al. 2018). In the present study, three novel species are assigned to this family and placed in the
genus Neosetophoma on account of their morphological and phylogenetic relationships.
Descriptions of new taxa and the comparisons with reported species are also provided.
Neosetophoma Gruyter et al.
Neosetophoma, is characterized by globose to irregular conidiomata, with papillate ostioles,
and with yellowish conidia that are attenuate at one end (de Gruyter et al. 2010, Liu et al. 2015,
Wijayawardene et al. 2016), which has been often reported as a pathogen causing leaf spots of
various hosts (Phookamsak et al. 2014b). In recent studies, Tibpromma et al. (2017) introduced N.
329
garethjonesii as the first report of the sexual morph of Neosetophoma and Wanasinghe et al. (2018)
introduced another three taxa to Neosetophoma viz. N. rosae, N. rosarum and N. rosigena. In this
study, we introduce another new species Neosetophoma shoemakeri, with both its sexual and
asexual morphs.
Figure 35 – Phylogram generated from maximum likelihood analysis based on combined LSU,
SSU, ITS and TEF sequence data of Phaeosphaeriaceae. Related sequences were obtained from
Hyde et al. (2017b) and Phookamsak et al. (2017). 132 strains are included in the combined
sequence analyses, which comprise 3510 characters with gaps. Leptosphaeria doliolum (CBS
505.75) and Paraleptosphaeria dryadis (CBS 643.86) are used as the outgroup taxa. Tree topology
of the ML analysis was similar to the BI. The best scoring RAxML tree with a final likelihood
value of -29771.43732 is presented. The matrix had 1473 distinct alignment patterns, with 26.99%
of undetermined characters or gaps. Estimated base frequencies were as follows; A = 0.246460, C =
0.234850, G = 0.265591, T = 0.253100; substitution rates AC = 1.126918, AG = 2.548533, AT =
1.959684, CG = 0.714174, CT = 5.985048, GT = 1.000000; gamma distribution shape parameter α
= 0.478992. Bootstrap support values for ML equal to or greater than 60% and BYPP equal to or
greater than 0.90 are given above the nodes respectively. Newly generated sequences are in red.
330
Figure 35 – Continued.
Neosetophoma guiyangensis J.F. Zhang, J.K. Liu, K.D. Hyde & Z.Y. Liu, sp. nov.
Fig. 36
Index Fungorum number: IF554080; Facesoffungi number: FoF 04378
Etymology – in reference to the collection site where the holotype was collected, Guiyang
City, China.
Holotype – GZAAS 18–0101
Saprobic on unidentified dead branch, visible as raised, black, globular structures on host
surface. Sexual morph Ascomata immersed under epidermis to superficial, globose to subglobose,
dark brown to black, solitary to gregarious, coriaceous, ostiolate. Peridium up to 13–18.5 µm wide,
comprising two strata, the outer stratum composed of thick-walled, black, pseudoparenchymatous
cells, fusing at the outside with the host tissue, inner stratum composed of slightly pigmented, thinwalled cells of textura angularis. Hamathecium comprising numerous, cellular pseudoparaphyses,
anastomosing among and above the asci, embedded in a gelatinous matrix. Asci (52–)55–69.5(–
75.5) × 8–11.5 µm ( = 64.5 × 9 µm, n = 25), 8-spored, bitunicate, fissitunicate, cylindric-clavate,
short pedicellate with a furcate pedicel, apically rounded, with a minute ocular chamber.
Ascospores (17–)20–25.5(–28) × 3.5–4.5 µm ( = 22 × 3.9 µm, n = 25), overlapping 1–2-seriate,
fusiform, slightly curved, with acute ends, 1–3(–5)-septate, hyaline when young, becoming pale
brown at maturity, not constricted at the septum, smooth-walled, without any appendages. Asexual
morph Undetermined.
331
Culture characteristics – Ascospores germinating on WA within 12 hours. Colony reaching
up to 23 mm on PDA after 10 days at 25°C, circular, surface flat or effuse, slightly raised at center,
fluffy, greenish-grey from above, and yellowish-grey from below.
Material examined – CHINA, Guizhou Province, Guiyang City, on dead branch of
undetermined host, 12 May 2016, J.F. Zhang, GZ-13 (GZAAS 18-0101, holotype), ex-type living
culture, GZCC 18-0111.
GenBank numbers – ITS: MH018134, LSU: MH018132, SSU: MH018136.
Notes – Neosetophoma guiyangensis is phylogenetically close to N. xingrensis with high
bootstrap support (99% BP and 1.00 PP) (Fig. 35). However, it can be distinguished from the latter
in its longer asci (52–75.5 µm vs. 40–64 µm). The most conspicuous difference between these two
taxa, is that N. guiyangensis has a furcate pedicel, while the pedicel in N. xingrensis is sessile.
Neosetophoma shoemakeri Senwanna, Wanas., Bulgakov, E.B.G. Jones & K.D. Hyde, sp. nov.
Figs 37, 38
Index Fungorum number: IF554067; Facesoffungi number: FoF03920
Etymology – in honour of Shoemaker R.A., for his immense contribution to mycology.
Holotype – MFLU 16-1606
Saprobic on Cirsium sp. (Asteraceae) and Malva sp. (Malvaceae) in terrestrial habitats.
Sexual morph Ascomata 170–230 µm high, 170–200 µm diameter, solitary, sometimes clustered,
immersed to slightly erumpent, visible as slightly raised, small, black dots on the host surface,
uniloculate, globose to subglobose, dark brown to black, ostiolate. Peridium 15–40 µm wide, thick
at the apex, thinner at the base and side, composed of 2–4 layers of dark brown to black cells of
textual angularis to textura prismatica. Hamathecium comprising 1.5–3 µm wide, numerous,
filamentous, septate, pseudoparaphyses. Asci 60–100 × 6–11 µm (x̅ = 83.7 × 9.4 μm, n = 40), 8spored, bitunicate, fissitunicate, cylindrical to clavate, short pedicellate, apically rounded with an
indistinct ocular chamber. Ascospores 15–27 × 3–6 µm (x̅ = 20.4 × 4.6 μm, n = 70), 1–2 seriate,
partially overlapping, fusiform, initially hyaline to subhyaline with small guttules, becoming
yellowish-brown to brown at maturity, 3-septate, enlarged at the second cell below the apex, slightly
curved, constricted at the septa, smooth-walled. Asexual morph coelomycetous. Conidiomata 120–
180 µm high, 120–160 µm diameter (x̅ = 164.7 × 142.9 µm, n = 5), pycnidial, separate, dark brown,
globose, subepidermal, unilocular, thin-walled, papillate. Conidiomatal wall 5–15 μm wide,
composed of 2–4 layers, lightly pigmented to brown, thick-walled, cells of textura angularis.
Conidiophores reduced to conidiogenous cells. Conidiogenous cells 4–5 μm long × 2.5–4 μm wide,
enteroblastic, phialidic, doliiform to ampulliform, determinate, hyaline, smooth-walled. Conidia
7.5–10.5 × 2.5–3 µm (x̅ = 9.1 × 2.6 µm, n = 30), subcylindrical, fusiform, or ellipsoid to fusiform,
individually hyaline, olivaceous-brown at maturity as a mass, smooth-walled, 1-septate, usually
attenuate at one end, thin and smooth-walled, with minute granules.
Culture characteristics – Colonies on PDA reaching 2 cm diameter after 21 days at 16 °C,
colonies medium dense, irregular, flat or effuse, slightly raised, margin filiform, fluffy, white from
above, yellowish in media from below, not produced pigmentation on PDA media. Mycelium
immersed in and superficial on the media.
Material examined – RUSSIA, Rostov region, Shakhty City, Alexandrovsky (Central) Park,
on dead stem of Cirsium arvense (L.) Scop. (Asteraceae), 14 March 2016, Timur S. Bulgakov, T1312 (MFLU 16-1606, holotype); ex-type living culture, MFLUCC 17-2510; United Kingdom,
Hampshire, Southsea, Eastney shore, on stems of Malva sp., (Malvaceae), 17 April 2016, EBG
Jones GJ268 (MFLU 17-0607); living culture, MFLUCC 17-0780, ICMP.
GenBank numbers – MFLUCC 17-2510 – ITS: MG602203, LSU: MG602199, SSU:
MG602201, TEF: MG739515; MFLUCC 17-0780 – ITS: MG844346, LSU: MG844348, SSU:
MG844350, TEF: MG844352.
Notes – During our investigations on diversity of microfungi in Russia and UK, two fresh
collections (MFLU 16-1606 and MFLU 17-0607) were made from Cirsium arvense and Malva sp.
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Figure 36 – Neosetophoma guiyangensis (GZAAS 18-0101, holotype) a Appearance of ascomata
on host surface. b Vertical section through ascoma. c Section of peridium. d Pseudoparaphyses. e–g
Asci. h–l Immature and mature ascospores. Scale bars: b = 50 µm, c, h–l = 5 µm, d = 20 µm, e–g =
10 µm.
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Figure 37 – Sexual morph of Neosetophoma shoemakeri (MFLU 16-1606, holotype). a, b
Appearance of ascomata on host surface. c Section through ascoma. d Ostiole. e Peridium. f
Pseudoparaphyses. g Germinated spores. h–n Immature and mature asci. o–u Immature and mature
ascospores. Scale bars: b = 500 µm, c = 100 µm, d–g = 20 µm, h–n = 30 µm, o–u = 15 µm.
in the Rostov region of European Russia and Eastney shore of Hampshire in the United Kingdom,
respectively. MFLUCC 17-2510 was derived from the Russian collection (ascomycetous fungal
species), while MFLUCC 17-0780 was obtained from the UK collection (coelomycetous asexual
morph). In our combined LSU, SSU, ITS and TEF sequenced data analyses, these two new isolates
share a monophyletic relationship within Neosetophoma in Phaeosphaeriaceae. A comparison of
334
the 574 ITS (+5.8S) nucleotides with these two new strains reveals only two base-pair differences
and a comparison of the 898 TEF nucleotides were identical within both of these strains. This
justifies both strains as the sexual and asexual morphs of the same taxon. The asexual morph
morphologically fits well within the generic concept of Neosetophoma in having hyaline, phialidic
conidiogenous cells and slightly yellowish, ellipsoidal to cylindrical, conidia which are usually
attenuate at one end (de Gruyter et al. 2010). There are no previously published literature records of
the sexual morph in this genus. The sexual morph of our new isolate is morphologically similar to
Phaeosphaeria oryzae in having phragmosporous, narrowly fusiform, pale brown to brown or
yellowish-brown, straight or slightly curved, 3-septate ascospores. However, they are
phylogenetically not closely related (Fig. 35).
Figure 38 – Asexual morph of Neosetophoma shoemakeri (MFLU 17-0607). a, b Conidiomata on
host substrate. c Vertical section through conidioma. d, e Conidia attached to conidiogenous cells. f
Conidia. Scale bars: a = 1 mm, b = 200 μm, c = 100 μm, d, f = 10 μm, e = 5 μm.
Neosetophoma xingrensis J.F. Zhang, J.K. Liu, K.D. Hyde & Z.Y. Liu, sp. nov.
Fig. 39
Index Fungorum number: IF554079; Facesoffungi number: FoF04379
Etymology – in reference to the collection site where the fungus was collected, Xingren
County, China.
Holotype – GZAAS 18–0100
Saprobic on decaying wood, visible as raised, shiny-black spots on host surface. Sexual
morph Ascomata 112–195 µm high, 120–193 µm diameter, globose to subglobose, dark brown to
black, solitary to gregarious, immersed to erumpent through the host surface, coriaceous, ostiolate.
Peridium 11.5–21 µm wide, composed of two strata, an outer stratum comprising dark brown to
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black, thick-walled, pseudoparenchymatous cells, and an inner stratum composed of hyaline to
slightly pigmented, thin-walled cells of textura angularis. Hamathecium comprising numerous, 2–3
µm wide, cellular pseudoparaphyses, anastomosing among and above asci, embedded in a
gelatinous matrix. Asci (40–)51–59(–64) × 8–11 µm ( = 52 × 9 µm, n = 20), 8-spored, bitunicate,
fissitunicate, cylindric-clavate, with a very short pedicel, apically rounded with an ocular chamber.
Ascospores (15–)21–28 × 3–6 µm ( = 22 × 5 µm, n = 30), 1–3-seriate, fusiform, straight to
slightly curved, hyaline when young, and becoming pale brown at maturity, normally 3-septate, not
constricted at the septum, smooth-walled, guttulate, without any mucilaginous sheath and
appendages. Asexual morph Undetermined.
Material examined – CHINA, Guizhou Province, Xingren County, on decaying wood, 2 June
2015, J.F. Zhang, XY 5–7 (GZAAS 18–0100, holotype), ex-type living culture, GZCC 18–0110.
GenBank numbers – ITS: MH018135, LSU: MH018133.
Culture characteristics – Ascospores germinating on WA within 12 hours and germ tubes
produced from one or both ends. Colonies growing on PDA, reaching a diameter of 31 mm after 15
days at 25°C, flat, circular, with entire edge, grey-greenish from above, moderately dense, and
slightly yellowish from below.
Notes – In the present analysis (Fig. 35), the new taxon is placed in Neosetophoma and
clusters with N. guiyangensis with high bootstrap support. However, it differs from other
Neosetophoma species by its sessile asci.
Sporormiaceae Munk
The family Sporormiaceae was introduced by Munk (1957) with Sporormia as the type genus
and S. fimetaria (Rabenh.) De Not as the type species. Species of this family occur worldwide,
mostly as saprobes on various substrates, including dung, plant debris, soil and wood (Cain 1961,
Dissing 1992, Kruys et al. 2006, Sue et al. 2014). Sexual morphs of this family are dark brown
ascospores which are strongly constricted at septa and often fragment into part-spores at maturity
(Barr 2000) and asexual morphs are coelomycetous or hyphomycetous (Phukhamsakda et al. 2016).
In this study, we introduce a new species of Neomassarina based on morphological and molecular
data and include this genus in Sporormiaceae.
Neomassarina Phook. et al.
Neomassarina was introduced by Hyde et al. (2016) with N. thailandica as the type species.
Earlier this monospecific genus did not cluster with members of the main families of
Dothideomycetes in phylogenetic analyses. Therefore, it was placed in Pleosporales genera
incertae sedis. The morphological features of the species in this genus are massarina-like and
according to our multi-gene phylogenetic analyses in this study, Neomassarina clusters in
Sporormiaceae with high bootstrap support (Fig. 21). In this study, we introduce a second species
of Neomassarina on a dead leaf of Pandanus sp. from Thailand.
Neomassarina pandanicola Tibpromma & K.D. Hyde, sp. nov.
Fig. 40
Index Fungorum number: IF553941; Facesoffungi number: FoF03813
Etymology – refers to the name of the host plant which the fungus was collected.
Holotype – MFLU 16-0554
Saprobic on dead leaf of Pandanus sp. Sexual morph Ascomata 140–160 µm high × 130–180
µm diameter ( = 152 × 156.4 µm, n = 5), semi-immersed to erumpent through host surface,
globose to subglobose, as small black dots on the host surface, solitary, without papilla, ostiole
central, black, smooth-walled. Peridium 16–18.5 µm, thin-walled, composed two layers, the outer
layer composed of thick-walled, dark brown to black cells of textura angularis; the inner layer
composed of hyaline to light brown cells of textura angularis. Hamathecium comprising 1.5–2.5
µm wide, hyaline, septate, cylindrical pseudoparaphyses. Asci 50–80 × 6.5–10 µm ( = 67.7 × 8.9
µm, n = 20), 6–8-spored, bitunicate, fissitunicate, cylindrical to cylindric-clavate, with short, club-
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Figure 39 – Neosetophoma xingrensis (GZAAS 18-0100, holotype). a, b Appearance of ascomata
on host surface. c Section of peridium. d Vertical section through ascoma. e Pseudoparaphyses. f–i
Immature and mature asci. j–m Immature and mature ascospores. Scale bars: a = 200 µm, b = 100
µm, c = 5 µm, d = 30 µm, e = 20 µm, f–m = 10 µm.
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Figure 40 – Neomassarina pandanicola (holotype). a Colony on dead leaves of Pandanus sp. b
Section of ascoma. c Peridium. d Hamathecium. e–g Asci. h, i Ascospores. j Ascospore stained
with India ink. k Germinating ascospore. l, m Culture characters on MEA (note m reverse). Scale
bars: a = 200 µm, b = 50 µm, c = 10 µm, d = 5 µm, e–g = 20 µm, h–k = 5 µm.
338
shaped pedicel, apically rounded with an ocular chamber. Ascospores 11–20 × 3–5 µm ( = 15.2 ×
3.8 µm, n = 20), overlapping uni- to bi-seriate, hyaline, fusiform, 1-septate, constricted at the
septum, narrowly rounded both end, ends cone-shaped, smooth-walled, guttulate, with appendages
at both ends (3.5–6 µm long). Asexual morph Undetermined.
Culture characteristics – Colonies on MEA reaching 9 cm diameter after 4 weeks at room
temperature, colony from above, white to yellow-white; from below, white to yellow-white at the
margin, dark-brown at the centre; circular, with entire edge, raised on media surface, flossy,
velvety.
Material examined – THAILAND, Prachuap Khiri Khan Province, Sai Khu Waterfall, Bang
Saphan District, on dead leaf of Pandanus sp. (Pandanaceae), 30 July 2015, Saowaluck
Tibpromma, SF15-032 (MFLU 16-0554, holotype); ex-type living culture, MFLUCC 16-0270,
KUMCC; ibid. (HKAS100837bis, paratypes).
GenBank numbers – ITS: MG298946, LSU: MG298945, SSU: MG298947.
Notes – Neomassarina pandanicola was collected on dead leaves of a Pandanus sp. from
southern Thailand. Based on phylogenetic analysis, N. pandanicola clusters with N. thailandica
with strong bootstrap support (100% in ML, Fig. 21). Moreover, we compared the culture and
morphological characteristics of N. pandanicola with N. thailandica and found that N. thailandica
differs from N. pandanicola (Hyde et al. 2016) in having ascospores with a distinct mucilaginous
sheath and being pale brown at maturity, whereas N. pandanicola has ascospores with appendages
at both ends of the ascospores and are not pale brown at maturity. We also compared the DNA
sequences of N. pandanicola and N. thailandica and found that there are nine base pair differences
in ITS and eleven base pair differences in both LSU and SSU regions. Thus, based on both
morphological and phylogenetic support (Jeewon & Hyde 2016), Neomassarina pandanicola is
introduced here as a new species.
Roussoellaceae J.K. Liu et al.
Although Jaklisch et al. (2016) consider Roussoellaceae to be a synonym of Thyridariaceae,
this has not been followed by numerous subsequent authors (Liu et al. 2017a, Tibpromma et al.
2017, Wanasinghe et al. 2018, Wijayawardene et al. 2017a, 2018). Thus, we accept Roussoellaceae
as a distinct family. A new species of Roussoella, R. mangrovei is described here based on
morphological characters and phylogenetic analysis.
Roussoella Sacc.
The genus Roussoella was described by Saccardo & Paoletti (1888) with R. nitidula as the
type species. A distinctive character of this genus is the trabeculate pseudoparaphyses, although
Liew et al. (2001) has shown this characters occurs across the Dothideomycetes (e.g. also in some
genera of Melanommataceae, Tian et al. 2015). Roussoella occurs mostly on monocotyledons
except Roussoella mexicana Crous & Yáñez-Moral. and R. solani Crous & M.J. Wingf. (Crous et
al. 2016). Currently, 33 species epithets are listed in Index Fungorum (2018).
Roussoella mangrovei Phukhamsakda & K.D. Hyde, sp. nov.
Fig. 42
Index Fungorum number: IF554073; Facesoffungi number: FoF 03923
Holotype – MFLU 17-1542
Saprobic on dead branches of Rhizophora L., forming raised, dark brown areas with black
dots on the host surface. Sexual morph Ascomata 352–476 μm high × 290–400 μm wide ( = 434
× 340 μm, n = 10) diameter, on the surface of the host, solitary, gregarious, erumpent through host
surface by papilla, globose to subglobose, coriaceous, ostioles central, black to dark brown, smooth
to papillate, ostiolate with periphysoids. Peridium 12–43(–58 at apex) μm wide, multi-layered,
outer layer composed of 7–12 layers of dark, brown to light brown cells of textura angularis, the
inner layer comprising hyaline thin-walled cells. Hamathecium of dense, 1–1.6 μm ( = 1.3 μm, n
= 20), filiform, branches, anastomosing, transverse septate, trabeculate psedoparaphyses. Asci
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Figure 41 – The best scoring RAxML tree based on combined partial LSU, SSU, ITS, and TEF1-α
gene datasets. Bootstrap values ≥70% from the maximum likelihood analysis are followed by
Bayesian posterior probabilities (PP) values ≥ 0.90 at the nodes. The tree is rooted with Torula
herbarum (CPC 24114). The species determined in this study indicated in blue. The ex-type and
references strains are indicated in black bold. Hyphen (-) represents support values ≤ 70%/0.90.
82–115 × 12–19 μm ( = 117 × 15 μm, n = 20), 8-spored, bitunicate, fisitunicate, broad-cylindrical
to clavate, apically round with an ocular chamber. Ascospores 15–20 × 7–11 μm ( = 16 × 9 μm, n
= 50), uni-seriate to partial overlapping, oval with round ends, pale-brown to brown, uni-septate,
constricted at septum, granulate in each cell well visible when young, rough-walled, surrounded
with mucilaginous sheath. Asexual morph Undetermined.
Culture characteristics – Colonies on MEA, with dense mycelium, flat on the surface circular,
umbonate, upper part white, become cream after 4 weeks, without aerial mycelium. Reverse brown
in the middle, mycelia radiating outwards with cream, hyphae septate branched.
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Figure 42 – Roussoella mangrovei (MFLU 17-1542, holotype) a Substrate. b Ascomata on
Rhizophora sp. c Vertical section of ascoma. d Ostiole filled with periphysoids. e Partial part of
peridium. The peridium comprising textura angularis. f Hyaline pseudoparaphyses. g–i Developing
stages of asci. j–n Developing ascospores. Scale bars: b = 500 µm, c = 200 µm.
341
Material examined – THAILAND, Krabi Province, on dead branches of Rhizophora sp.
(Rhizophoraceae), 16 December 2015, C. Phukhamsakda (MFLU 17-1542, holotype), ex-type
living culture, MFLUCC 16-0424, ICMP.
GenBank numbers – ITS: MH025951, LSU: MH023318, RPB2: MH028250, TEF1:
MH028246.
Notes – Roussoella mangrovei (MFLUCC 16-0424) is described herein as a novel species in
Roussoella occurring on an intertidal mangrove branch in Thailand. The strain shares a close
relationship with Roussoella mukdahanensis Phook. et al. (90% ML/1.00 PP), which was reported
from bamboo (Dai et al. 2016b). Roussoella mangrovei can be distinguished by its relatively larger
solitary ascomata, a thick peridium, cylindrical to clavate asci and pale-brown to brown, 1-septate
ascospores, with clearly visible granules.
Pleosporomycetidae genera, incertae sedis
The class Dothideomycetes comprises two subclasses which are Dothideomycetidae and
Pleosporomycetidae (Hyde et al 2013, Liu et al. 2017). Pleosporomycetidae includes the orders
Pleosporales, Mytilinidiales and Hysteriales (Boehm et al. 2009, Hyde et al. 2013; Wijayawardene
et al. 2014, Liu et al. 2017). In this study, we provide an updated phylogenetic tree for the subclass
Pleosporomycetidae and introduce a new species of Hysterographium, Pleosporomycetidae genera,
incertae sedis.
Hysterographium Corda
Hysterographium is a genus in the subclass Pleosporomycetidae, and is typified by H. fraxini
(Pers.) De Not (Boehm et al. 2009a, b). Currently, there are 142 epithets listed under this genus
(Index Fungorum 2018). However, only one species has molecular data and other species are
synonymised under different genera. In this study, we describe and introduce a new species
Hysterographium didymosporum based on morphology and phylogenetic support.
Hysterographium didymosporum S. Boonmee & K.D. Hyde, sp. nov.
Figs 44, 45
Index Fungorum number: IF554386; Facesoffungi number: FoF04107
Etymology – the epithet “didymosporum” refers to two celled ascospores.
Holotype – MFLU 10–0031
Saprobic on dead branches of dicotyledonous plans. Sexual morph Ascomata 179–242 µm
high × 145–167 µm diameter ( = 216 × 155 µm), hysterothecial, completely immersed, becoming
erumpent at maturity, with longitudinal slit, dark brown, grouped to scattered, straight to slightly
curved, subglobose to obovoid in vertical section. Peridium 27–30 µm wide, composing several
layers of dark brown cells of textura angularis. Hamathecium comprising numerous, 1–2(–2.5) µm
wide, cylindrical, septate, branched, anastomosed, pseudoparaphyses, embedded within a
gelatinous matrix. Asci 60–99 × 10–14 µm ( = 85 × 12 µm, n = 15), 8-spored, bitunicate,
cylindrical-clavate, sessile or with short pedicel, with a small ocular chamber. Ascospores 13–16 ×
6–8 µm ( = 15 × 7 µm, n = 20), biseriate, conglobate, ellipsoid, upper cell wider, ends rounded, 1septate, constricted at the septum, hyaline, smooth-walled. Asexual morph coelomycetous.
Conidiomata 28–42 × 28–52 µm, pycnidial, clustered, globose-subglobose, dark brown, invisible
conidiophores and conidiogenous cells, surrounded by aerial hyphae, septate, branched, white when
viewed with the unaided eye, and under microscopic hyaline. Conidia 7–11 × 2–4 µm ( = 8 × 3
µm, n = 20), oblong, slightly curved, aseptate, with or without guttules, hyaline, smooth-walled.
Cultural characteristics – Ascospores germinating on MEA within 12 hours and germ tubes
produced at both ends. Colonies slow growing on MEA, reaching less than 5 mm in 7 days at 28 C,
slightly convex, undulating to raised, dentate, with slightly radial striations and lobate edges,
brown. Asexual morph produced in culture.
Material examined – THAILAND, Lam Pang, Wang Neua, Wat Pa Putta Sri, elev. ca. 800–
900 msl., on dead branches of an unidentified host, 7 January 2011, S. Boonmee, LP01 (MFLU 10–
0031, holotype), ex-type culture: MFLUCC 10–0101, IFRDCC 2187, BCC 52329.
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Figure 43 – RAxML phylogenetic tree generated from LSU, RPB2, SSU and TEF1 sequence data
of Hysterographium didymosporum MFLUCC 10–0101 with related orders of subclass
Pleosporomycetidae. RAxML bootstrap support values ≥ 50% (BT) and Bayesian posterior
probabilities ≥ 95% (PP) are given above the nodes. The ex-type strains are in bold and new strain
in blue.
343
Figure 43 – Continued.
344
GenBank numbers – LSU – MH105775, RPB2 – MH105777, SSU – MH105776.
Notes – Hysterographium didymosporum is introduced as a new species with both sexual and
asexual morphs. The asexual morph developed in culture grown on MEA (Fig. 45). In our
phylogenetic analysis of combined LSU, RPB2, SSU and TEF1 sequence data, Hysterographium
didymosporum constitutes an independent lineage with high statistical support (97% MLBT and
1.00 BYPP, Fig. 43) and appears to be phylogenetically distinct from H. fraxini (type species).
Hysterographium didymosporum shares common features of dark brown, hysterothecial, immersed
to erumpent ascomata, with a longitudinal slit, but differs from taxa in Hysterographium in
ascomata lacking carbonaceous wall and having 1-septate and hyaline ascospores.
Figure 44 – Hysterographium didymosporum (MFLU 10–0031, holotype). a, b Appearance of
ascomata on dead branches of an unidentified host. c Section of ascoma. d Peridium. e
Pseudoparaphyses. f–h Asci (Figs. g, h stained in cotton blue reagent). i–l Ascospores. Scale bars: b
= 500 µm, c = 100 µm, d = 50 µm, e = 5 µm, f–h = 20 µm, i–l = 10 µm.
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Figure 45 – Hysterographium didymosporum (MFLU 10–0031, ex-type culture). a Germinating
spore. b, c Colonies on MEA from surface and reverse. d Developing conidiomata. e Aerial hyphae
in culture. f Squash mount of conidiomata. g Conidia. Scale bars: a, g = 10 µm, b–c = 10 mm, d =
100 µm, e = 5 µm, f = 50 µm.
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Dothideomycetes orders, incertae sedis
Acrospermales Minter et al.
Acrospermaceae Fuckel
Minter et al. (2007) introduced the order Acrospermales to accommodate the family
Acrospermaceae, which is typified by Acrospermum compressum Tode. Lumbsch & Huhndorf
(2010) placed the Acrospermales in Dothideomycetes, order incertae sedis. The family
Acrospermaceae presently includes two genera Acrospermum and Oomyces (Wijayawardene et al.
2018). The asexual morphs of this family include members of Dactylaria and Gonatophragmium
which are linked to asexual Acrospermum (Wijayawardene et al. 2012, 2017b).
Figure 46 – Simplified phylogram showing the best RAxML maximum likelihood tree obtained
from a combined multigene (LSU and SSU) matrix including species in Acrospermaceae and
related families. Related sequences were obtained from GenBank. Twenty strains are included in
the combined sequence analyses, which comprise 2774 characters with gaps. The best scoring
RAxML tree with a final likelihood value of -9380.272776 is presented. The matrix had 611
distinct alignment patterns, with 48.85% of undetermined characters or gaps. Estimated base
frequencies were as follows; A = 0.248348, C = 0.230480, G = 0.299990, T = 0.221182. ML values
above 70% are given at each branch. MLBS values above 70% are given at each branch. The tree is
rooted with Tubeufia javanica (MFLUCC 120545) (Tubeufiaceae). New isolates are in bold and
red.
Acrospermum longisporium Jayasiri, E.B.G. Jones & K.D. Hyde, sp. nov.
Fig. 47
Index Fungorum number: IF554046; Facesoffungi number: FoF03887
Etymology – with reference to the long ascospores
Holotype – MFLU 17-2849
Saprobic on Urtica dioica (Urticaceae). Sexual morph Ascomata 1.5–2 mm high × 0.4–0.5
mm diameter ( = 1.8 × 0.45 mm), solitary or in groups, superficial, club-shaped to conoid, erect,
uni-locular, brown to blackish when dry, with a short pedicel or sessile, flattened when dry,
swelling when moist, ostiole large, apex rounded. Peridium in horizontal section comprising three
layers, an outer layer comprising dark brown cells of textura angularis, a central thick layer,
comprising pale brown tissue of gelatinized hyphae with elongated cells, and an inner layer
comprising dense tissue of small, light brown cells. Hamathecium of narrow, long,
347
pseudoparaphyses. Asci 300–370 × 3–5 μm ( = 350 × 4 µm), 8-spored, bitunicate, narrowly
cylindrical, pedicellate, with an ocular chamber. Ascospores 150–170 × 0.5–1 μm ( = 160 × 0.75
µm), fasciculate, filiform, hyaline, multi-septate, almost long as the asci, smooth-walled. Asexual
morph Undetermined.
Material examined – United Kingdom, England, Hampshire, Botley wood, decaying branch
of Urtica dioica (Urticaceae), 25 May 2016, E.B.G. Jones GJ 293 (MFLU 17-2849, holotype);
ibid., (PDD, isotype).
GenBank numbers – LSU: MG815827, SSU: MG815828.
Notes – Acrospermum longisporium fits well with the generic concept of Acrospermum in
having club-shaped, brown, uni-locular ascomata and narrow, long paraphyses which resemble
ascospores (Hyde et al. 2013). Acrospermum longisporium is related to Acrospermum graminum
(77% ML support). However the latter differs from Acrospermum longisporium in having platted
ascoma and unequally thick peridia.
Figure 47 – Acrospermum longisporium. a, b Ascomata. c, d Sections through ascomata. e–g Asci.
h, i Ascospores. Scale bars: a, b = 500 μm, c = 200 μm, d = 100 μm, e–i = 50 μm.
Asterinales M.E. Barr ex D. Hawksw. & O.E. Erikss.
Asterinaceae Hansf.
Asterinaceae was established as a member of Myriangiales by Hansford (1946). Species of
Asterinaceae are characterized by dark brown hyphae with hyphopodia, dark brown
thyriotheciawith stellate dehiscence, and dark brown ascospores with a single septum. Phylogenetic
studies have resulted in several different interpretations of this family (Hongsanan et al. 2014,
Guatimosim et al. 2015, Liu et al. 2017). In this study, we introduce a new species of Asterina
based on morphological and molecular data.
348
Asterina Lév.
Asterina is the type genus of the family Asterinaceae, which was introduced by Léveillé in
1845. Asterina is the largest genus in Asterinaceae and has a cosmopolitan distribution in tropical
and subtropical regions (Hongsanan et al. 2014).
Figure 48 – Phylogram generated from Bayesian analysis based on LSU sequence data from
species of Asterinales. Maximum parsimony/likelihood bootstrap support values greater than
50%/70% are shown above the nodes, while Bayesian posterior probabilities greater than 0.9 are
displayed below the nodes. Sequences from type specimens are in bold and new sequences are in
blue bold.
Asterina magnoliae X.Y. Zeng, T.C. Wen & K.D. Hyde, sp. nov.
Fig. 49
Index Fungorum number: IF554238; Facesoffungi number: FoF04089
Etymology – referring to the host genus Magnolia.
Holotype – MFLU 16-0072
Colonies epiphyllous or hypophyllous, scattered. Hyphae superficial, brown, septate,
reticulate, with hyphopodia. Hyphopodia subglobose, unicellular, alternate. Sexual morph
Thyriothecia 88–144(-152) μm (x̅ = 115 µm, n = 20) in diameter, dense, circular, flattened, dark
brown, with stellate dehiscence. Upper wall brown, comprising radial arrangement of septate cells
of textura prismatica. Asci 40–48 μm (x̅ = 45 µm, n = 20) in diameter, 8-spored, bitunicate, globose
to subglobose. Ascospores 25–28 × 11–12 μm (x̅ = 26.5 × 11.5 µm, n = 20), cylindrical, 1-septate,
slightly constricted at the septum, hyaline when young, becoming brown when mature, with a
darkened band and a large guttule in the middle of each cell. Asexual morph Conidia ovoid,
aseptate, hyaline when young, become brown when mature, with two-layered cell wall.
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Material examined – THAILAND, Chiang Mai, Mae Taeng, Pa Pae, Bahn Pa Deng, 128 Moo
3, Mushroom Research Centre, on living leaves of Magnolia odora (Magnoliaceae), 8 July 2015,
Xiang-Yu Zeng (MFLU 16-0072, holotype).
GenBank numbers – LSU: MG844186.
Notes – There was no report of Asterina species found on Magnolia (Zeng et al. 2017) and
our new taxon is different from other Asterina species in having a coloured band at center of each
cell of the ascospores. The phylogeny herein also positions A. magnolia in an independent lineage
basal to other Asterina species analysed (Fig 48).
Figure 49 – Asterina magnoliae. a Host leaves. b, c Colony on host surface. d–g Ascus from young
state to mature state. h Upper wall of thyriothecium. i Squash mount of thyriothecium. j–m
Ascospore from young state to mature state. n–o Conidia. Scale bars: b–c = 200 μm, d–g = 20 μm,
h–I = 50 μm, j–o = 10 μm.
Botryosphaeriales C.L. Schoch et al.
Botryosphaeriaceae Theiss. & H. Syd.
Taxa of the family Botryosphaeriaceae are endophytes, plant pathogens and saprobes
(Phillips et al. 2008, 2013, Liu et al. 2012, Dissanayake et al. 2016). Currently the family comprises
23 accepted genera (Dissanayake et al. 2016). Morphological characters alone are inadequate to
define genera or identify species in the family (Phillips et al. 2013). Hence SSU, ITS, LSU, EF1-α
and β-tubulin sequence data are needed when defining a new species and genus level identifications
(Phillips et al. 2013). In this study, we introduce a new Diplodia species based on molecular and
morphological evidence.
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Figure 50 – Phylogenetic tree generated by maximum likelihood analysis of combined ITS and
TEF-α sequence data of Diplodia species. Related sequences were obtained from GenBank. Fortysix strains are included in the analyses, which comprise 820 characters including gaps. The tree is
rooted with Lasiodiplodia theobromae (CBS 164.96). Tree topology of the ML analysis was similar
to the MP and BI. The best scoring RAxML tree with a final likelihood value of -3144.936756 is
presented. The matrix had 49 isolates, 254 distinct alignment patterns, with 4.02% of undetermined
characters or gaps. Estimated base frequencies were as follows; A = 0.208206, C = 0.208206, G =
0.261341, T = 0.233093; substitution rates AC = 1.031688, AG = 3.931659, AT = 0.914981, CG =
1.704693, CT = 5.063628, GT = 1.000000; gamma distribution shape parameter α = 0.662619. The
maximum parsimonious dataset consisted of constant 634, 138 parsimony-informative and 48
parsimony-uninformative characters. The parsimony analysis of the data matrix resulted in the
351
maximum of two equally most parsimonious trees with a length of 361 steps (CI = 0.657, RI =
0.857, RC = 0.563, HI = 0.343) in the first tree. RAxML and maximum parsimony bootstrap
support values ≥ 50% (BT) are shown respectively near the nodes. The scale bar indicates 0.04
changes. The ex-type strains are in bold and new isolates in blue.
Diplodia Fr.
The genus Diplodia was introduced based on D. mutila (Fr.) Mont. by Montagne (1834). The
genus comprises 31 species with sequence data (Dissanayake et al. 2016). Species of Diplodia, are
pathogens, endophytes or saprobes on a wide range of woody hosts (Crous et al. 2006, Slippers &
Wingfield 2007, Phillips et al. 2012). Two types of conidia can be observed in Diplodia species
(Phillips et al. 2005, 2012). Conidia remain hyaline for a long time before they become brown and
1-septate, but in some species, conidia become pigmented before discharge from the conidiomata
and mostly remain aseptate (Phillips et al. 2005, 2012).
Diplodia arengae R.H. Perera, Wanas. & K.D. Hyde, sp. nov.
Fig. 51
Index Fungorum number: IF554057; Facesoffungi number: FoF03908
Etymology – name reflects the host genus Arenga.
Holotype – MFLU 17-2769
Saprobic on dead leaves of Arenga hookeriana (Becc.) Whitmore. Sexual morph Ascomata
240–280 μm high, 250–300 μm diameter ( = 262.9 × 271.3 µm, n = 5), scattered, immersed,
globose to subglobose, glabrous, brown to dark brown, ostiolate. Ostiole 60–90 µm long 55–75 µm
diameter, short papillate, black, smooth, filled with hyaline cells. Peridium 30–40 µm wide at the
base, 35–60 µm wide at the sides, comprising 4–5 layers, outer layer pigmented, comprising
reddish-brown to dark brown, thin-walled cells of textura angularis, inner layer composed of
hyaline, thin-walled cells of textura angularis. Hamathecium comprising numerous, 3.5–4.5 µm
wide, filamentous, branched, septate, pseudoparaphyses. Asci 130–160 × 25–35 µm ( = 144.6 ×
30.5 µm, n = 30), 8-spored, bitunicate, fissitunicate, cylindric-clavate to clavate, with a pedicel 20–
30 μm long, apically rounded, with broad ocular chamber. Ascospores 25–35 × 10–12 µm ( =
30.8 × 11.3 µm, n = 40), overlapping biseriate to triseriate, ellipsoidal, hyaline, aseptate, smoothwalled, guttulate, ends remaining cone-shaped, with pointed ends, not surrounded by a
mucilaginous sheath. Asexual morph Undetermined.
Culture characteristics – Colonies on PDA reaching 3 cm diameter after 30 days at 25°C,
circular, smooth margin white at first, dark coffee after 4 weeks, flat on the surface, without aerial
mycelium, reverse black. Hyphae septate branched, hyaline, thin, smooth-walled. We could not
manage to maintain a living culture as subsequent attempts to subculture failed and hence a living
culture is unavailable.
Material examined – CHINA, Mengla, Menglun, Xishuangbanna Tropical Botanical Garden,
on dead leaf of Arenga hookeriana (Arecaceae), 21 December 2013, D.N. Wanasinghe, XTBG28
(MFLU 17-2769, holotype).
GenBank numbers – β-tubulin: MG783039, ITS: MG762771, LSU: MG762772, SSU:
MG762773, TEF: MG762774.
Notes – Diplodia arengae forms a well-supported independent lineage sister to D. corticola,
D. gallae and D. quercivora. Diplodia arengae differs from D. corticola by possessing unilocular,
smaller ascomata (vs. multiloculate, 1 mm diameter), longer asci (vs. 160–250 µm) and narrower
ascospores (vs. 14.6–15.3 µm) (Alves et al. 2004). There is no sexual morph reported for D.
quercivora for morphological comparison with D. arengae (Linaldeddu et al. 2013, Phillips et al.
2013). However, those two species are phylogenetically distant in our analysis (Fig. 50).
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Figure 51 – Diplodia arengae (MFLU 17-2769, holotype). a, b Appearance of ascomata on dead
leaf of Arenga hookeriana. c Section of ascoma. d Close up of ostiole. e Peridium. f
Pseudoparaphyses. g, h Asci. i–l Ascospores. Scale bars: c = 100 µm, d, e, g, h = 20 µm, f, i–l = 10
µm.
353
Dyfrolomycetales K.L. Pang et al.
Pleurotremataceae Walt. Watson
The family Pleurotremataceae was re-established after several contradictions with reexamination of the isotype of Pleurotrema polysemum, and exclusion of Dyfrolomycetaceae from
the class Sordariomycetes. However, sequence data for the type of Pleurotrema is still lacking
(Maharachchikumbura et al. 2016a). Recently, studies have revealed that the family
Pleurotremataceae is phylogenetically close to Acrospermaceae (Dothideomycetes), and comprises
Dyfrolomyces, Melomastia and Pleurotrema (Pang et al. 2013, Norphanphoun et al. 2017b, Zhang
et al. 2017b). In this study, a new Dyfrolomyces species collected from Camellia sinensis is
introduced with detailed morphological information and support from combined multi-loci
phylogenetic analysis.
Dyfrolomyces K.D. Hyde et al.
Pang et al. (2013) introduced Dyfrolomyces, with species characterized by a clypeus on the
substrate, immersed ascomata and multi-septate ascospores with/without a sheath in
bitunicate/fissitunicate asci, to accommodate D. tiomanensis. Currently, the genus consists of eight
species, which have mostly been collected from marine habitats on mangrove wood (Hyde 1992,
Pang et al. 2013, Hyde et al. 2017b, Norphanphoun et al. 2017b, Zhang et al. 2017b).
Figure 52 – Phylogram generated from maximum likelihood (RAxML) analysis of combined SSU
and LSU partial sequence data. The best scoring RAxML tree with a final likelihood value of 4848.762798 is presented with Dendryphiopsis atra and Kirschsteiniothelia aethiops
(Kirschsteiniotheliaceae) as the outgroup taxa. The best scoring RAxML tree with a final likelihood
value of -4848.762798 is presented. The matrix had 222 distinct alignment patterns, with 0% of
undetermined characters or gaps. Estimated base frequencies were as follows; A = 0.240566, C =
0.238552, G = 0.309758, T = 0.211124; substitution rates AC = 1.010469, AG = 3.557274, AT =
1.030465, CG = 1.606891, CT = 10.067712, GT = 1.000000; gamma distribution shape parameter
α = 0.292895. Bootstrap values for ML (≥60) and Bayesian posterior probabilities (≥0.95) are
shown near the nodes. The newly generated sequence is in blue, and the ex-type strains are in bold.
The scale bar represents the expected number of nucleotide substitutions per site.
Dyfrolomyces sinensis Samarak., Tennakoon & K.D. Hyde, sp. nov.
Index Fungorum number: IF554110; Facesoffungi number: FoF03935
Fig. 53
354
Etymology – named sinensis, after the host species from which it was collected.
Holotype – MFLU 17-0777
Saprobic on Camellia sinensis (L.) Kuntze (Theaceae). Sexual morph Ascomata 450–600 μm
diameter × 350–450 μm high ( = 510 × 397 μm, n = 10), solitary, superficial, clypeate, globose to
subglobose, dark brown to black, ostiolate. Ostiole centric to eccentric, apapillate. Peridium 55–75
μm wide ( = 65.1 μm, n = 10), 10–35 μm wide at base ( = 21.8 μm, n = 10), comprising an
outer layer of dark brown cells of textura angularis, becoming lighter inwardly. Hamathecium
comprising numerous, 2.4–5 μm wide ( = 3.5 μm, n = 20), septate pseudoparaphyses, embedded
in a gelatinous matrix. Asci 160–220 × 8–10 μm ( = 187.5 × 8.5 μm, n = 12), 8-spored, bitunicate,
cylindrical, short-pedicellate, with a thickened apex. Ascospores 18–30 × 5– 8 μm ( = 23.1 × 6.2
μm, n = 25), L/W ratio 3.7, uniseriate, hyaline, cylindrical, mostly 6–7-septate, often similar width
of cells with several small guttules, slightly constricted at the septum. Asexual morph
Undetermined.
Culture characteristics – Colonies on PDA at 25–28°C, reaching 33–35 mm in 8 days, surface
smooth, effuse, undulate to slightly with radially striated lobate edges. Colonies from above site are
greyish to dirty white at the margin, whitish sparse mycelia in the centre; reverse yellowish-brown
to light yellow at the margin. Odour not pronounced.
Material examined – THAILAND, Chiang Rai, Mae Fah Luang University, on stems of
Camellia sinensis (Theaceae), 25 March 2017, Danushka S. Tennakoon, MFLU 001 (MFLU 170777, holotype), ex-type living culture, MFLUCC 17-1344.
GenBank numbers – LSU: MG836699, SSU: MG836700.
Notes – The new taxon, Dyfrolomyces sinensis, shares similar morphological characters with
other Dyfrolomyces species (Pang et al. 2013, Norphanphoun et al. 2017b, Zhang et al. 2017b).
However, it differs from otherspecies (viz. D. aquatica, D. mangrovei, D. marinospora, D.
rhizophorae and D. thailandica) by lacking a gelatinous sheath around the ascospores (Hyde 1992,
Tsui et al. 1998, Pang et al. 2013). Dyfrolomyces phetchaburiensis and D. thamplaensis also have
similar characters and lack a gelatinous sheath and cluster with D. sinensis in the multi-loci
phylogenetic analyses. However, D. sinensis differs from D. thamplaensis based on number of
septa (6–7, 3) and L/W ratio (3.7, 4) of the ascospores. In addition, both D. phetchaburiensis has
been reported on submerged wood of Rhizophora apiculata in marine habitats and D. thamplaensis
on corticated dead branch in Karst habitats (Hyde et al. 2017b, Zhang et al. 2017b). Therefore D.
sinensis is introduced as a new species on dead stem of Camellia sinensis from terrestrial habitats
(Fig. 52).
Kirschsteiniotheliales Hern.-Restr. et al.
Kirschsteiniotheliaceae Boonmee & K.D. Hyde
The family Kirschsteiniotheliaceae Boonmee & K.D. Hyde was proposed by Boonmee et al.
(2012), to accommodate the genus Kirschsteiniothelia D. Hawksw. (Hawksworth 1985), and its
asexual morph Dendryphiopsis S. Hughes (1953). Members of Kirschsteiniotheliaceae are
distinguished by superficial, globose to subglobose ascomata, ellipsoidal, septate, coloured
ascospores and a Dendryphiopsis asexual morph (Boonmee et al. 2012, Hyde et al. 2013, 2017b, Li
et al. 2016a, Mehrabi et al. 2017). However, Kirschsteiniothelia arasbaranica and K. thujina
(Hawksworth 1985), as well as K. phoenicis, which we introduce here, lacks any asexual morph,
but groups within this family supported by molecular data. Based on phylogenetic analysis,
Kirschsteiniotheliaceae was assigned to the order Kirschsteiniotheliales (Hernández-Restrepo et al.
2017, Wijayawardene et al. 2018). In the study, we provide an updated phylogenetic tree (Fig. 54)
and introduce a new species, Kirschsteiniothelia phoenicis.
Kirschsteiniothelia D. Hawksw.
Kirschsteiniothelia was introduced by Hawksworth (1985) and typified by K. aethiops (Sacc.)
D. Hawksw based on Sphaeria aethiops Berk. & M.A. Curtis. The ascomycetous genus comprises
21 species according to Index Fungorum (2018) and is characterized by superficial, erumpent,
355
globose to subglobose, dark brown to black, membranaceous, scattered or loosely aggregated
ascomata with or without a central papilla; a thick pseudoparenchymatous peridium consisting of
thick-walled cells arranged in a pallisadic configuration at the base angles (textura angularis); a
hamathecium comprising numerous, filiform, hyaline, pseudoparaphyses; bitunicate, fissitunicate,
cylindrical-clavate, 8-spored asci with a long pedicel and an ocular apical chamber and bi- or triseriate ascospores, ellipsoidal, slightly curved, smooth-walled, olive brown to dark brown, 1-2septate with a median or submedian septum, with or without a mucilaginous sheath (Boonmee et al.
2012, Hawksworth 1985, Hyde et al. 2013, Mehrabi et al. 2017).
Figure 53 – Dyfrolomyces sinensis (MFLU 17-0777, holotype). a, b Appearance of ascomata on
host surface. c Vertical section through ascoma. d Peridium. e Pseudoparaphyses. f–i Asci. j–l
Ascospores. m Germinating ascospore. n, o Culture on PDA (n. upper, o. lower). Scale bars: c, e–i
= 50 μm, d = 20 μm, j–l = 10 μm.
356
The asexual morph of Kirschsteiniothelia is Dendryphiopsis (Wijayawardene et al. 2017b),
which is supported by molecular data (Boonmee et al. 2012, Hawksworth 1985, Hyde et al. 2013,
Schoch et al. 2006). Wijayawardene et al. (2014) proposed to use Kirschsteiniothelia over
Dendryphiopsis and corrected the name D. atra as Kirschsteiniothelia atra (Corda) D. Hawksw.
The asexual morph is macronematous, mononematous, erect, branched or unbranched, septate,
brown to dark brown, smooth-walled conidiophores; monoblastic, terminal, with delimited
conidiogenous cells, constricted at the septa and apically produced, broadly ellipsoid-obovoid, 1–
2(–3)-septate, light brown, red brown to dark brown and smooth-walled conidia (Hawksworth
1985, Hyde et al. 2013, Su et al. 2016).
The genus Kirschsteiniothelia was originally assigned in Pleosporaceae (Hawksworth 1985,
Barr 1987), but later Barr (1993) reported that Kirschsteiniothelia belongs to the Pleomassariaceae
based on host, morphology and anamorph characters. With the support of molecular data, Schoch et
al. (2006) showed that the type species, K. aethiops, was not phylogenetically related to the
Pleosporaceae, and suggested that Kirschsteiniothelia should be transferred to a separate family. In
further phylogenetic analysis, K. elaterascus Shearer grouped within Morosphaeriaceae, while K.
maritima (Linder) D. Hawksw. clustered with Mytilinidion spp. in the Mytilinidiaceae clade
(Schoch et al. 2009, Shearer et al. 2009, Suetrong et al. 2009). The taxa which are related to K.
aethiops are included in Kirschsteiniotheliaceae (Boonmee et al. 2012).
Kirschsteiniothelia phoenicis S.N. Zhang & K.D. Hyde, sp.nov.
Fig. 55
Index Fungorum number: IF554239; Facesoffungi number: FoF04093
Etymology – name reflects the host genus Phoenix.
Holotype: MFLU 18-0153.
Saprobic on Phoenix paludosa Roxb. Sexual morph Ascomata 135–160 μm high, 183–235
μm diameter ( = 143.8 × 201.7 µm, n = 10), black, scattered, subglobose to globose or slightly
conical in appearance, superficial with apical papilla or plane, base remaining immersed, usually
flattened. Peridium laterally 24–34 µm wide, composed of 5-6 layers of textura angularis, with
dark brown outer layer cells and inner layers pale brown, cells around the ostiole with small
lumina, reduced at the base, of 1–2 layers of cells. Pseudoparaphyses 0.8–1.5 µm wide, branched,
embedded in a gelatinous matrix. Asci 70–112 × 14–24 µm ( = 84.7 × 19.4 µm, n = 15), 8-spored,
bi-or tri- seriate in the middle of the ascus, cylindrical-clavate, straight or slightly curved, apically
rounded, with a short pedicellate up to 5 µm long. Ascospores 18–27 × 5–7.5 µm ( = 21.3 × 6.6
µm, n = 36), brown, ellipsoid, rounded or slightly pointed at the ends, 1-septate, septum submedian
and constricted, upper cell broader than the lower cell, each cell containing a distinct large guttule,
smooth-walled, with a mucilaginous sheath. Asexual morph Undetermined.
Culture characteristics – Ascospores germinating on PDA within 24 hours. Colonies growing
on PDA and MEA dense, reaching 1–1.5 cm diameter after two weeks growing on PDA at 25°C,
the surface covered with a white mat of aerial hyphae in first week, then becoming grayish-blue
and dark bluish, composed of brown to dark brown, septate, smooth or verrucose hyphae 1.5–2.5
µm wide and up to 5 µm wide.
Material examined – THAILAND, Ranong, Ranong Mangrove Forest Research Center, on
rachis of Phoenix paludosa (Arecaceae), that was probably intertidal as it was immersed in
mangrove mud with water, 7 December 2016, S.N. Zhang, SNT76 (MFLU 18-0153, holotype); extype living culture, MFLUCC 18-0216, TBRC; ibid. (BBH 43493, isotype).
GenBank numbers – ITS: MG859978, LSU: MG860484, SSU: MG859979 (RPB2:
MG994912, TEF1: MG994911, are submitted but not included in this analysis).
Notes – Kirschsteiniothelia phoenicis is morphologically simiar to the type species K.
aethiops and K. reticulata C.Y. Chen et al. in ascospore width. However, K. phoenicis differs from
K. aethiops as it has smaller ascospores, and a mucilaginous sheath. The asexucal morph was not
determined. Kirschsteiniothelia phoenicis can easily be distinguished from K. reticulata as the
ascospores lack any reticulate ornamentation. The phylogeny based on multigene analysis herein,
357
also indicates that K. phoenicis is distinct from other species, given that it constitutes a strongly
supported independent lineage, basal to other Kirschsteiniothelia species (Fig. 54).
Figure 54 – RAxML tree based on analysis of combined ITS, LSU and SSU rDNA sequence data.
Bootstrap values for ML and MP equal to or greater than 75% are placed above and below the
branches respectively. Branches with Bayesian posterior probabilities (PP) from MCMC analysis
equal or greater than 0.95 are in bold. The ex-type strains are in bold and newly generated
sequences are indicated in red. The tree is rooted with Phyllobathelium anomalum and
Flavobathelium epiphyllum. The final dataset consisted of 27 taxa (25 ingroup) with a total 2726
characters after alignment. 1673 characters were constant and 741 characters were parsimony
informative, while 312 variable characters were parsimony uninformative. After a heuristic search
using PAUP, one equally most parsimonious tree was obtained (tree length = 2195 steps, CI =
0.678, RI = 0.767, RC = 0.520, HI = 0.322).
358
Figure 55 – Kirschsteiniothelia phoenicis (MFLU 18-0153, holotype). a Appearance of ascomata
on host surface with ostiole. b Vertical section through the ascoma. c, d Structure of peridium. e
Pseudoparaphyses. f, g Asci. h–k Ascospores. l Ascospore in India ink showing sheath. m–p
Colony and mycelia growing on PDA. Scale bars: a = 100 μm, b = 50 μm, c, d, f, g, o, p = 20 μm,
e, h–l = 10 μm, n = 1 mm.
359
Class Lecanoromycetes O.E. Erikss. & Winka
Subclass Acarosporomycetidae Reeb et al.
Teloschistales D. Hawksw. & O.E. Erikss.
Teloschistaceae Zahlbr.
Taxa are lichenized with photobiont member Trebouxia. Ascomata are apothecial with welldeveloped thallin margins (Gaya et al. 2008). Most Teloschistaceae species produce anthraquinone
pigments in the cortex which provide yellow to orange colours to their apothecia (Arup et al. 2013).
Paraphyses are unbranched to slightly branched and usually slightly capitate. Asci are
semifissitunicate, with apical tholus and distinct ocular chamber. Ascospores are mostly 1–3septate and ellipsoid. The asexual morph is pycnidial (Jaklitsch et al. 2016a). In this study, we
illustrate Cerothallia subluteoalba collected from Australia.
Cerothallia Arup et al.
Cerothallia is a small genus in the family Teloschistaceae with four species (Lücking et al.
2016). They form crustose lichenized thalli with trebouxia-like photobionts. This genus is
geographically restricted in temperate regions (Nimis & Martellos 2017).
Figure 56 – Phylogenetic tree generated by maximum likelihood analysis of combined ITS
sequence data of Caloplaca and Cerothallia species. Related sequences were obtained from
GenBank. Twenty-one strains are included in the analyses, which comprise 605 characters
including gaps. The tree is rooted with Lecanora contractuala (AFTOL ID 877). The best scoring
RAxML tree with a final likelihood value of -4428.138068 is presented. The matrix had 344
distinct alignment patterns, with 9.93% of undetermined characters or gaps. Estimated base
frequencies were as follows; A = 0.200210, C = 0.293629, G= 0.276064, T = 0.230097;
substitution rates AC = 1.227023, AG = 3.402371, AT = 1.963531, CG = 1.415528, CT =
6.016852, GT = 1.000000; gamma distribution shape parameter α = 1.734937. RAxML bootstrap
support values ≥ 50% (BT) are shown respectively near the nodes. The scale bar indicates 0.07
changes. The new isolates are in blue.
Cerothallia subluteoalba (S.Y. Kondr. & Kärnefelt) Arup, Frödén & Søchting (2013)
Facesoffungi number: FoF04096
Fig. 57
360
Lichenized on dead stem. Sexual morph 350–400 × 134–178 µm ( = 373.5 × 152 µm, n =
10). Apothecia scattered on the host thallus, sessile, erumpent from the substrate, yellow when both
dry and moist conditions. Hypothecium pulvinate, disc convex, margins slightly elevated, disc and
the margins yellow. Exciple 26–42 µm ( = 34.3 µm, n = 10), yellowish-brown in water, becoming
reddish in KOH, laterally consisting of thick-walled, isodiametric cells (mainly in the upper part),
basally of conglutinated, thick-walled, hyaline, more or less isodiametric cells. Hymenium upper
part yellowish-brown and lower part is hyaline, in KOH upper part becoming reddish, embedded in
a thick gelatinous matrix. Paraphyses 0.5–1 µm wide ( = 0.8 µm, n = 20), numerous, filiform,
aseptate, capitate or enlarged and pigmented at the apex, exceeding asci in length, apices are glued
together to develop epithecium. Asci 28–35 × 8–12 µm ( = 33.6 × 10 µm, n = 30), 8-spored,
sessile, arising from croziers, cylindric–clavate, rounded at the apex, amyloid ring absent at the
ascus apex. Ascospores 8–10 × 4–6 µm ( = 9.6 × 5.3 µm, n = 40), multiseriate, 1-celled, ovoid,
hyaline to reddish, smooth, thick-walled. Asexual morph Undetermined.
Material examined – AUSTRALIA, Peninsula, Mornington, on the dead stem, 10 March
2015, E.B. Gareth Jones, GJ103 (MFLU 16-0561).
GenBank number –ITS: MG820705, LSU: MH216681.
Notes – Cerothallia subluteoalba is characterized by pulvinate apothecia, a yellowish-brown
excipulum which turns red in KOH, filiform paraphyses swollen at the apex, short sessile asci and
ellipsoid ascospores (Kantvilas 2016).
Subclass Lecanoromycetidae P.M. Kirk et al. ex Miadl. et al.
Peltigerales W. Watson
Collemataceae Zenker
Taxa form gelatinous lichen thalli with cyanobacteria of the genus Nostoc (Otálora et al.
2014). Thalli lobes are small to large, leathery to papery when dry and swell to becomes extremely
gelatinous when wet, lower side glabrous or tomentose. They are mostly epiphytic on bark, on
rocks, soil and between bryophytes (Otálora et al. 2013, 2014). Ascomata are apothecial. The
hamathecium comprises amyloid, unbranched paraphyses and asci are mostly 8-spored or rarely 4spored, semi-fissitunicate, clavate to cylindrical and amyloid. Ascospores are transversely septate
to muriform, fusiform to ellipsoid and hyaline. Asexual morphs are pycnidial (Otálora et al. 2013,
2014, Jaklitsch et al. 2016a). In the study, we introduce Leptogium thailandicum sp. nov. from
Thailand.
Leptogium (Ach.) Gray
Leptogium is a widely distributed genus which includes around 250 species (Aragón et al.
2005). More studies are required on this genus as there are many species complexes and groups
which need to be resolved (Otálora et al. 2008).
Leptogium thailandicum Ekanayaka & K.D. Hyde, sp. nov.
Fig. 59
Index Fungorum number: IF554068; Facesoffungi number: FoF04094
Etymology – with reference to the country where the holotype was collected.
Holotype – MFLU 16-0586
Lichenized on dead stems. Sexual morph Apothecia 1125–1828 × 251–655 µm ( = 1525.5 ×
425.7 µm, n = 10), arising in small groups, sessile to sub-stipitate on the thallus. Hypothecium
cupulate, external surface greenish. Disc flat to slightly concave, light brown to dark brown when
fresh. Margins light green. Hymenium hyaline, enclosed in a thick gelatinous matrix. Ectal
excipulum 17.7–37.5 µm ( = 25.4 µm, n = 10), composed of small, thin-walled, hyaline cells of
textura angularis. Medullary excipulum composed of hyaline cells of textura intricata. Paraphyses
1.4–2.4 µm wide ( = 1.9 µm, n = 20), numerous, filiform, obtuse at the apex, aseptate,
unbranched. Asci 98.6–123.4 × 14.9–26.9 µm ( = 112.9 × 18 µm, n = 30), 8-spored, medium to
long pedicellate, cylindric–clavate, rounded at the apex, amyloid ring absent at the ascus
361
Figure 57 – Cerothallia subluteoalba (MFLU 16-0561). a Substrate. b Ascomata on wood. c
Ascoma on wood. d Cross section of an ascoma. e Close up of a vertical section of the ascoma at
the margin. f Aseptate paraphyses. g Arrangement of asci and paraphyses in the hymenium layer.
h–k Short sessile asci (f–k mounted in KOH). l–o Ovoid ascospores. Scale bars: b = 500 µm, d =
200 µm, c = 100 µm, e = 40 µm, f = 20 µm, g−k = 15 µm, l−o = 5 µm.
362
Figure 58 – Phylogenetic tree generated by maximum likelihood analysis of LSU sequence data of
Leptogium species. Related sequences were obtained from GenBank. Seventy-four strains are
included in the analyses, which comprise 1243 characters including gaps. The tree is rooted with
363
Lecanora contractula (AFTOL ID 877). The best scoring RAxML tree with a final likelihood value
of -6512.383381 is presented. The matrix had 499 distinct alignment patterns, with 26.01% of
undetermined characters or gaps. Estimated base frequencies were as follows; A = 0.239, C =
0.252, G= 0.323, T = 0.185; substitution rates AC = 1.384799, AG = 2.394283, AT = 0.793732,
CG = 1.334471, CT = 8.586646, GT = 1.000000; gamma distribution shape parameter α =
0.198923. Maximum likelihood bootstrap values ≥50% are given near the nodes. The scale bar
indicates 0.06 changes. Strain/culture numbers are given after the taxon names. Newly generated
sequences are in blue bold.
apex. Ascospores 18.6–24.2× 6.5–11.6 µm ( = 20.1 × 8.7 µm, n = 40), partially biseriate, lower
spores are uniseriate, hyaline, smooth, muriform, 3–5-transverse septate, with 1-longitudinal septa,
ellipsoid to subfusiform with pointed apices in mature spores. Asexual morph Undetermined.
Material examined – THAILAND, Chiang Rai Province, Doi Mae Salong, on dead stems, 22
June 2015, A.H. Ekanayaka, HD025 (MFLU 17-2769, holotype); ibid. (HKAS, isotype).
GenBank number – LSU: MG820706, ITS: MH216041.
Notes – Leptogium thailandicum is phylogenetically close to L. phyllocarpum. They
however, differ by 22 base pairs in the LSU region. Moreover, L. phyllocarpum has highly
wrinkled margins, while L. thailandicum has smooth margins and ascospores with pointed apices
(Nash et al. 2004). Leptogium corticola is morphologically similar to L. thailandicum, but differs in
having larger ascospores (30–56 × 10–20 µm) (Sierk 1964, Brodo et al. 2001).
Class Pezizomycetes O.E. Erikss. & Winka
Pezizales J. Schröt.
Otideaceae Eckblad
This family contains eight genera and around 70 species (Jaklitsch et al. 2016a, Ekanayaka et
al. 2018). Most Otideaceae taxa are characterized by cupulate to ear-shaped, brownish, and sessile
to sub-stipitate apothecia. Some form cleistothecia (Ekanayaka et al. 2018). The ectal excipulum is
composed of textura angularis and medullary excipulum of textura intricata. Paraphyses are
straight or apically curved and asci are 8-spored, cylindric-clavate with ellipsoid ascospores (Liu &
Zhuang 2006, Leuchtmann & Clémençon 2012, Hansen et al. 2013). In this study, we introduce
Otidea pseudoformicarum sp. nov. from China.
Otidea (Pers.) Bonord.
The genus Otidea was established by Bonorden (1851). The recent comprehensive study on
this genus is by Olariaga et al. (2015). The species of this are restricted to the Northern Hemisphere
(Olariaga et al. 2015).
Otidea pseudoformicarum Ekanayaka, Q. Zhao and K.D. Hyde, sp. nov.
Fig. 61
Index Fungorum number: IF554071; Facesoffungi number: FoF04095
Etymology – the specific epithet pseudoformicarum refers to the morphological similarity of
this species to O. formicarum.
Holotype – HKAS 101386
Saprobic on soil. Sexual morph Apothecia 2–5 cm high, 2–7 cm in diameter, broadly earshaped to cupulate or flattened, sub-stipitate, split, hymenium surface yellowish-brown when dry.
Receptacle concave, yellowish-brown dry, with pruinose outer surface. Disc concolorous to
receptacle. Ectal excipulum composed of smooth, thin-walled hyaline cells of textura angularis to
globulosa with yellowish to brownish exudates. Medullary excipulum composed of loosely
arranged, hyaline, non-gelatinized, septate, smooth, thin-walled hyphae of textura intricata.
Paraphyses 1.5–2.3 µm ( = 1.9 µm, n = 20) wide at the middle, filiform, smooth and thin-walled,
septate, hyaline. Asci 115–150 × 7–10 µm ( = 131.4 × 9 µm, n = 15), 8-spored, unitunicate,
subcylindrical, short pedicellate, arising from croziers, non-amyloid, ascospores released from an
eccentric split at the apical apex. Ascospores 8–10 × 5–7 µm ( = 9 × 5.8 µm, Qm= 1.6–1.4, n =
364
25), ellipsoid, with 2 guttules, sometimes multi-guttulate, smooth-walled, uniseriate. Asexual
morph Undetermined.
Material examined – CHINA, Yunnan Province, Lijiang, Yushuizhai, 7 September 2014, Qi
Zhao N243 (HKAS 101386, holotype).
GenBank numbers – ITS: KY498601, LSU: KY498606.
Notes – Otidea pseudoformicarum is phylogenetically close to O. formicarum and O.
subformicarum. However, O. pseudoformicarum separates from them in a single lineage with
100% bootstrap support (Fig. 60). Otidea pseudoformicarum is morphologically similar to O.
formicarum in having broadly ear-shaped apothecia, an ectal excipulum with exudates, however, O.
pseudoformicarum differs in having shorter asci arising from croziers and smaller ascospores
(Olariaga et al. 2015).
Figure 59 – Leptogium thailandicum (MFLU 16-0586, holotype). a Substrate. b Apothecia on
wood. c Apothecium on wood. d Cross section of an apothecium. e Close up of a vertical section of
the apothecium at the margin. f Septate paraphyses. g–j Long pedicellate asci. k Non-amyloid
apical apex. l–o Ovoid ascospores. Scale bars: b = 1000 µm, d = 800 µm, c = 500 µm, e = 200 µm,
f = 70 µm, g−j = 50 µm, k = 20 µm, l−o = 10 µm.
365
Figure 60 – Phylogenetic tree generated by maximum likelihood analysis of combined ITS and
LSU sequence data of Otidea species. Related sequences were obtained from GenBank. Seventyfour strains are included in the analyses, which comprise 1598 characters including gaps. Single
gene analyses were carried out and compared with each species, to compare the topology of the tree
and clade stability. The tree is rooted with Monascella botryosa (CBS 233.85). The best scoring
RAxML tree with a final likelihood value of -15790.294971 is presented. The matrix had 895
distinct alignment patterns, with 22.64% of undetermined characters or gaps. Estimated base
366
frequencies were as follows; A = 0.248, C = 0.206, G= 0.275, T = 0.270; substitution rates AC =
1.117181, AG = 2.684224, AT = 1.368399, CG = 0.761371, CT = 5.232977, GT = 1.000000;
gamma distribution shape parameter α = 0.366724. RAxML bootstrap support values ≥ 50% (BT)
are shown respectively near the nodes. The scale bar indicates 0.06 changes. The new isolates are in
blue bold.
Figure 61 – Otidea pseudoformicarum (HKAS 101386, holotype). a Habit of fresh apothecia. b
Habit of dried apothecia. c–f Ellipsoid ascospores. g Cylindrical paraphyses. h, i Cylindrical asci.
Scale bars: h, i = 50 μm, g = 25 μm, c–f = 10 μm.
Class Leotiomycetes O.E. Erikss. & Winka
Rhytismatales M.E. Barr ex Minter
Rhytismataceae Chevall.
Rhytismataceae is a large family in Rhytismatales (Leotiomycetidae, Leotiomycetes,
Pezizomycotina, Ascomycota) and includes 44 genera and 400 species (Lumbsch & Huhndorf
2010, Lantz et al. 2011, Hyde et al. 2016). Taxa in Rhytismataceae can be endophytes, pathogens
and saprobes (Lantz et al. 2011, Hyde et al. 2016).
367
Figure 62 – The maximum parsimony (MP) tree based on analysis of ITS sequences of
Lophodermium species. Related sequences were obtained from Li et al. (2016c). Forty-nine strains
368
are included in the combined sequence analyses, which comprise 1773 characters with gaps.
Bisporella citrina and Neofabraea malicorticis are used as the outgroup taxa. The matrix had 1089
distinct alignment patterns, with 35% of undetermined characters or gaps. Bootstrap support values
higher than 50% and Bayesian posterior probabilities ≥ 0.95 (PP) are defined above the nodes
respectively. Newly generated sequences are in red bold. Ex-type strains are shown in black bold.
Lophodermium Chevall.
Lophodermium is the largest genus in family Rhytismataceae with more than 100 species
(Lantz et al. 2011, Hyde et al. 2016). They are economically important parasites, particularly on
conifers (Lantz et al. 2011).
Lophodermium thailandicum N.I. de Silva & K.D. Hyde, sp. nov.
Fig. 63
Index Fungorum number: IF554201; Facesoffungi number: FoF04081
Etymology – specific epithet refers to the country that specimen collected.
Holotype – MFLU 17-0673
Saprobic on dead leaf of unknown host species. Sexual morph Apothecia 350–420 µm high ×
750–1000 µm diameter ( = 400 × 880 µm, n = 10), rounded, scattered, sessile, erumpent, discoid.
Receptacle concave, dark brown to black. Margins concolorous to receptacle. Disc olive green,
concave. Ectal excipulum 26–33 µm, composed of cells of textura globulosa to angularis.
Medullary excipulum 35–40 µm, composed of cells of textura porrecta. Paraphyses 2–3 µm wide,
cylindrical, hyaline, embedded in common gelatinous matrix with asci. Asci 120–200 × 4–8 μm (
= 180 × 7 µm, n = 25), 8-spored, cylindrical, with a short pedicel. Ascospores 80–160 × 1–2 µm (
= 130 × 1.5 µm, n = 25), hyaline, elongated, filiform, sometimes curved and semi-circular. Asexual
morph Undetermined.
Material examined – THAILAND, Chiang Mai Province, Muang District, on dead leaf of
unknown host species, 27 September 2016, N. I de Silva, NI101 (MFLU 17-0673, holotype).
GenBank number – ITS: MG818852.
Notes – The phylogenetic analysis of ITS data indicates that our taxon belongs to
Lophodermium. We tried to isolate this taxon, unfortunately its ascospores failed to germinate.
Therefore, we obtained sequence data directly from the fruiting bodies. The phylogeny depicts an
interesting finding to support establishment of our new taxon. Lophodermium thailandicum is
positioned in a moderately supported independent lineage and nested inbetween other
Lophodermium species analysed but with poor support (Fig 62). The new taxon differs from
Lophodermium seditiosum in having rounded apothecia. Apothecia of Lophodermium seditiosum
are oval to elliptical and open by a split in the middle (Minter et al. 1978). Both Lophodermium
seditiosum and our strain have similar ascus morphology and ascospore length and width range
(Minter et al. 1978). Lophodermium seditiosum was found on pine needles in the USA (Minter et
al. 1978), but our strain was found on dead leaves of unknown host plant in Thailand. Therefore,
we introduce a new species, Lophodermium thailandicum, considering its phylogeny, morphology
and biogeography.
Class Sordariomycetes O.E. Erikss. & Winka
Subclass Diaporthomycetidae Senan. et al.
Annulatascales D’souza et al.
Annulatascaceae S.W. Wong et al.
Annulatascaceae comprises the genera Annulatascus, Annulusmagnus, Ascitendus and
Submersisphaeria, which are freshwater ascomycetes growing on submerged wood (Zhang et al.
2017a). Based on molecular data, Campbell and Shearer (2004) considered that taxa assigned to
Annulatascaceae were polyphyletic, and this was subsequently confirmed by various researchers.
(Abdel-Wahab et al. 2011, Boonyuen et al. 2012, Dayarathne et al. 2016) However, the recent
study of Zhang et al. (2017a) resolved taxonomic relationships among members of this family by
excluding several genera from Annulatascaceae based on analyses of a combined analysis of LSU,
369
SSU, ITS and RPB2 sequence data. In this study, we introduce a novel Ascitendus species from a
decaying wood submerged in the Mulgrave River, Australia.
Ascitendus J. Campb. & Shearer
The genus Ascitendus was established for Ascolacicola austriacus, by Campbell & Shearer
(2004) as a monotypic genus. Ascitendus comprises saprobic, freshwater or terrestrial ascomycetes
found from Austria and its sexual morph is unknown (Campbell & Shearer 2004, Wijayawardene at
al. 2017a).
Figure 63 – Lophodermium thailandicum (MFLU 17-0673, holotype). a, b Appearance of
ascomata on host. c, d Vertical section of ascoma. e Peridium. f, g Paraphyses. h–j Asci. k, l
Ascospores. Scale bars c–e = 80 μm, f = 30 μm, g–k = 50 μm, l = 30 μm.
370
Figure 64 – Phylogram generated from maximum likelihood analysis based on combined LSU and
SSU sequence data of selected taxa. Related sequences were obtained from GenBank. Twenty-nine
strains are included in the analyses, which comprise 1785 characters including gaps. Single gene
analyses were carried out and compared with each species, to compare the topology of the tree and
clade stability. The tree is rooted with Papulosa amerospora (AFTOL ID 748), Ascobrunneispora
aquatica (HKUCC 3708) and Fluminicola coronata (HKUCC 3717). The tree topology of the ML
analysis was similar to the MP and BI. The best scoring RAxML tree with a final likelihood value
of -7506.256928 is presented. The matrix had 483 distinct alignment patterns, with 47.67% of
undetermined characters or gaps. Estimated base frequencies were as follows; A = 0.253321, C =
0.232540, G = 0.300821, T = 0.213318; substitution rates AC = 0.627195, AG = 1.154324, AT =
0.764306, CG = 0.897430, CT = 3.282253, GT = 1.000000; gamma distribution shape parameter α
= 0.910487. The maximum parsimonious dataset consisted of constant 1204, 200 parsimonyinformative and 381 parsimony-uninformative characters. The parsimony analysis of the data
matrix resulted in the maximum of two equally most parsimonious trees with a length of 1002 steps
(CI = 0.736, RI = 0.625, RC = 0.459, HI = 0.264) in the first tree Maximum parsimony bootstrap
(MPBT, black) values > 60%, Bayesian posterior probabilities (PP, blue) > 0.80% and maximum
likelihood bootstrap (ML, green) values > 60%) are given above the nodes. The scale bar indicates
0.04 changes. The ex-type strains are in bold and new isolates in blue bold.
371
Fig. 65
Ascitendus aquaticus Dayar, Fryar & K.D. Hyde, sp. nov.
Index Fungorum number: IF554115; Facesoffungi number: FoF03926
Etymology – name reflects the host habitat aquatic environment.
Holotype – MFLU 18-0143
Saprobic on decaying wood submerged in freshwater. Sexual morph Ascomata 160–280 μm
high, 135–190 μm diameter, solitary to gregarious, semi-immersed to superficial, black, coriaceous,
globose to subglobose, ostiolate with a prominent neck, venter 150–230 × 195–240 μm. Peridium
25–45 μm wide, comprised of three layers: outer layer of pseudoparenchyma cells incorporated
with dark brown amorphous material; middle layer of brown, thin-walled, laterally compressed,
elongated cells; inner layer of hyaline, laterally compressed, elongated, hyaline cells. Neck 85–300
× 50–140 μm, central, cylindrical, periphysate, black at wall 2-layered, outer layer of dark brown,
laterally compressed thin-walled cells incorporated with brown amorphous material, inner layer of
laterally compressed thin-walled hyaline cells. Hamathecium paraphysate; paraphyses longer than
asci, 160–210 μm long, 4–10 μm broad at base, septate. Asci 150–200 × 5–12 μm ( = 175.5 × 8
μm, n = 20), 8-spored, unitunicate, cylindrical, long pedicellate, with a prominent cylindrical to
flaring apical ring, 2.5–3 × 3–3.5 μm wide at apex, 1–3 μm wide at the base, J-. Ascospores 15–25
× 4–8 μm ( = 20 × 6 μm, n = 20), uniseriate, hyaline when immature, becoming brown, fusoid,
straight to curved, 3-septate when mature, each cell with a single large guttule, slightly/not
constricted at septa; septa prominent, smooth-walled, without appendages or a sheath. Asexual
morph Undetermined.
Material examined – AUSTRALIA, North Queensland, Mulgrave River, S 17.17724, E
145.72374, on decaying wood submerged in a river, 14 April 2015, Sally Fryar and Ben Cawson,
MR150 (MFLU 18-0143, holotype); ibid. (AD279865, isotype).
GenBank numbers – LSU: MG813820, SSU: MG813821.
Notes – The new species, Ascitendus aquaticus, best fits the genus Ascitendus in both
morphology and receives phylogenetic support. It resembles Ascitendus by having globose to
subglobose, ostiolate ascomata with a prominent neck, cylindrical to flaring apical ring and brown
ascospores, with 3-prominent septa and each cell having a single large guttule. Our phylogenetic
analysis indicates that A. aquaticus is closely related to A. austriacus, but with moderate support
(78% ML, 70% MP, 1.00 PP) forming a separate lineage basal to A. austriacus. Ascitendus
aquaticus can be clearly distinguished from A. austriacus by having smooth-walled ascospores,
where the ascospore wall of A. austriacus are roughened in a striated pattern and ascospore end
cells have small refractive dots at the ends. Annulatascus velatisporus, Annulusmagnus triseptatus,
and Ascolacicola aquatica are also morphologically similar to Ascitendus aquaticus. However, A.
aquaticus can be clearly distinguished from all these species by asci, ascospore nature and
measurements as listed in Table 2.
Diaporthales Nannf.
For treatment of Diaporthales we follow Senanayake et al. (2017).
Cytosporaceae Fr.
Cytosporaceae was introduced by Fries (1823) as a family in Diaporthales (Wehmeyer 1975,
Barr 1978, Eriksson 2001, Castlebury et al. 2002). Maharachchikumbura et al. (2015, 2016a) listed
13 genera under Cytosporaceae. Recently, Senanayake et al. (2017) excluded seven genera and
accepted only five genera in Cytosporaceae as Cytospora, Paravalsa, Pachytrype, Waydora and
Xenotypa based on morphological characteristics. In the study, we provide an updated tree and
introduced two new species of Cytospora (Fig. 66).
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Figure 65 – Ascitendus aquaticus (MFLU 18-0143, holotype). a Ascomata on submerged wood. b,
c Sections of ascomata. d Section through neck region. e Section through peridium. f Paraphyses g
Part of an immature ascus. h Apical ring. i Apical ring in Melzer’s reagent. J Mature ascus. k–o
Ascospores. Scale bars: b, c = 50 μm, c–d, f, g, j, k = 20 μm, e, h, i, l–o = 10 μm.
373
Table 2 – Synopsis of the characteristics of Ascitendus aquatica and other related species.
Nature of
end cells
Taxon
Ascomata
Asci
Apical ring
Ascospores
Reference
Annulatascus
velatisporus
Globose to
subglobose,
445–500 ×
280–450 μm
J-, 6–8 × 2–5
μm
Aseptate,
fusiform,
hyaline, thick,
19.5–28 × 8–
12 μm
-
Dayarathne et
al. (2016)
Annulusmagnus
triseptatus
Oblate to
suboblate to
globose,
venter 375–
660 × 400–
700 µm
Unitunicate,
cylindrical,
apically
rounded, 224
–300 300 ×
10.5 10.5–14
μm
Cylindrical,
138–283 ×
7–14 μm
J-, 2–4 × 3–5
µm, tapering
1–2 µm from
top to
bottom
-
Campbell &
Shearer,
(2004)
Ascitendus
austriacus
Globose to
subglobose,
venter 350–
585 × 295–
505 µm
Cylindrical,
100–160 ×
4–6 μm
J-, cylindrical
to flaring,
2.5–3.5
μm long,
3.0–3.8 μm
With
small
refractive
dots at
ends
Campbell &
Shearer,
(2004)
Ascitendus
aquaticus
Globose to
subglobose,
160–280 ×
135–190 μm
Cylindrical,
150–200 ×
5–12 μm
J-,
cylindrical
Without
small
refractive
dots at
ends
In this study
Ascolacicola
aquatica
Subglobose,
250–375 µm
high, 225–
275 µm
Cylindrical,
125–163 ×
10–12.5 μm
J-, discoid,
apical ring,
2.5–3.7 μm
high, 6.2 μm
3 septate,
hyaline,
multiguttulate,
becoming pale
straw-colored
or pale brown
with age, 16–
37 × 5–10 μm
3-septate,
middle cells
darker than
outer cells,
each cell with
a single large
guttule, wall
roughened in
a striated
pattern, 4–27
× 4–9 μm
Hyaline when
immature
becoming 3septate at
maturity, each
cell with a
single large
guttule,
smooth
walled, 15–25
× 4–8 μm
3-septate,
brown 12.5–
16. 4–7.5 μm
With
hyaline to
pale
brown
end cells
Ranghoo &
Hyde (1998)
Cytospora Ehrenb.
Cytospora was introduced by Ehrenberg (1818), which causes canker and dieback disease on
branches of a wide range of host and has a worldwide distribution (Adams et al. 2005, 2006,
Norphanphoun et al. 2017a). Currently, there are 611 epithets for Cytospora (Index Fungorum
2018, 16 April 2018) with an estimated 110 species in Kirk et al. (2008). Norphanphoun et al.
(2017a), Tibpromma et al. (2017), Senanayake et al. (2017) and Hyde et al. (2016, 2017b) provided
a recent account of the genus with several new taxa.
374
Figure 66 – Phylogram generated from maximum parsimony analysis based on combined ITS,
LSU, RPB2 and ACT sequence data of Cytospora isolates with Phomopsis vaccinii (ATCC 18451)
as the outgroup taxon. The newly generated nucleotide sequences were compared against the
GenBank database using the Mega BLAST program. Related sequences were obtained from
GenBank (http://www.ncbi.nlm.nih.gov/). Taxa selection was based on Norphanphoun et al.
(2017a). Maximum parsimony and maximum likelihood bootstrap values ≥50%, Bayesian posterior
probabilities ≥0.90 (MPBS/MLBS/PP) are given at the nodes. The species obtained in this study are
in blue and ex-types from the study are in blue bold. Ex-type taxa from other studies are in black
bold.
375
Figure 66 – Continued.
Cytospora predappioensis Q.J. Shang, Norphanph., Camporesi & K.D. Hyde, sp. nov.
Fig. 67
Index Fungorum number: IF554083; Facesoffungi number: FoF03936
Etymology – the specific epithet “predappioensis’’ refers to the town of Predappio in Italy
from which the holotype was collected.
Holotype – MFLU 17-0323
Saprobic on dead branch of Platanus hybrida. Sexual morph Stromata 875–2685 µm wide,
with poorly developed interior, solitary to gregarious, immersed, becoming raised to erumpent by
ostiolar canal, dark brown to black, glabrous, circular to irregular in shape, arranged with
conspicuous, clustered, roundish to cylindrical prominent ostioles in the centre. Ascomata
(excluding necks) 240–480 μm high, 450–680 µm diameter ( = 365×567 μm, n = 10), perithecial,
immersed in a stroma, dark brown to black, globose to subglobose, glabrous, with individual
ostiolar neck. Ostiole 70–520 μm high, 100–150 μm diameter ( = 444 × 124 μm, n = 6),
cylindrical, sulcate, periphysate. Peridium 25–45 μm wide, composed of two section layers, outer
section comprising 5–7 layers, of relatively small, brown to dark brown, thick-walled cells,
arranged in textura angularis, inner part comprising 3–5 layers of hyaline cells of textura
angularis. Hamathecium 2.5–4 μm wide, dense, cylindrical, septate, hyaline, paraphyses. Asci (25–
)32–42(–54) × (4.5–)5.5–8(–9.8) μm ( = 37 × 7.7 μm, n = 50), 8-spored, unitunicate, clavate,
without stalks, apically rounded to truncate, with a J- apical ring. Ascospores (6.5–)8–10(–11) × (1–
)1.5–3(–3.5) μm ( = 9 × 2 μm, n = 110), biseriate, hyaline, fusiform to oblong, aseptate, smoothwalled. Asexual morph Undetermined.
Culture characteristics – Ascospores germinating on PDA within 12 hours. Germ tubes
produced from all sides. Colonies on PDA reaching 2–2.5 cm diameter after 3 days at room
temperature, colonies circular to irregular, medium dense, flat or effuse, slightly raised, with edge
fimbriate, fluffy to fairy fluffy, white to gray from above, light yellow to green from below; not
producing pigments in agar.
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Figure 67 – Cytospora predappioensis (MFLU 17-0323, holotype). a Stromata on substrate. b
Surface of fruiting bodies showing the black ostioles. c Transverse sections through stroma to show
distribution of locules. d, e Longitudinal sections through stroma to show distribution of locules
and ostiolar canal. f Peridium. g Ostiolar neck. h Paraphyses. i–m Asci. n Ascospores. o
Germinating ascospore. p, q Culture characteristic on PDA after 7 days (p = colony from above, q =
colony from below). Scale bars: d, g = 50 µm, e, f, o = 20 µm, h–m = 10 µm, n = 5 µm.
377
Material examined – ITALY, Province of Forlì-Cesena [FC], Massera, Predappio, on dead
aerial branch of Platanus hybrida, 27 January 2017, E. Camporesi, IT 3225 (MFLU 17-0323,
holotype), ex-type living culture, MFLUCC 17-2458, KUMCC 17-0328; ibid. (KUN-HKAS
100896, isotype).
GenBank numbers – MFLUCC 17-2458, ITS: MG873484, LSU: MG873480, SSU:
MG873482.
Notes – Based on phylogenetic analyses and morphological comparison, our isolates belong
to Cytosporaceae. Phylogenetic analyses of a combined sequence data, show that our taxon,
Cytospora predappioensis (MFLUCC 17-2458 and MFLUCC 17-0327), forms a distinct lineage
and shares a close relationship to Cytospora sacculus (Schwein.) Gvrit. (HMBF281 and
HMBF282), with good-support (ML 93%, MP 73% and 0.98 PP) (Fig. 66). The morphology of
stromata, ascomata, asci and ascospores of our isolate (Fig. 67), fits well with the description of the
sexual morph of Cytospora (Adams et al. 2005, Fan et al. 2015, Norphanphoun et al. 2017a).
Compared with C. ceratosperma, the sexual morph of Cytospora sacculus, the morphological
characters of C. predappioensis are similar except for the ascospores sizes (Spielman 1985, 3–12 ×
1–3 μm versus (6.5–)8–10(–11) × (1–)1.5–3(–3.5) μm, this study). However, given that our two
isolates cluster together in a highly supported subclade and segregated from C. sacculus, there is
ample evidence to justify a new species.
Cytospora prunicola Norphanph., Camporesi, T.C. Wen & K.D. Hyde, sp. nov.
Fig. 68
Index Fungorum number: IF554078; Facesoffungi number: FoF04097
Etymology – named after the host, Prunus from which it was isolated.
Holotype – MFLU 17-0995
Associated with twigs and branches of Prunus sp. Sexual morph Undetermined. Asexual
morph Conidiomata 500–1000 × 450–500 µm diameter, immersed in host tissue, scattered,
erumpent, discoid, circular, with 2–4 locules, ostiolate. Ostioles 170–180 µm diameter, at the same
level, with flattened top. Peridium comprising a few to several layers of cells of textura angularis,
inner layer thick, brown, outer later dark brown. Conidiophores branched, reduced to
conidiogenous cells. Conidiogenous cells blastic, enteroblastic, phialidic, formed from the inner
most layer of pycnidial wall, hyaline, smooth-walled. Conidia (4–)5.2–6.6 × 1.1–1.3(–1.6) µm (x̅ =
5.5 × 1.3 µm, n = 30), unicellular, allantoid, slightly curved ends, hyaline, smooth-walled.
Material examined – ITALY, Province of Forlì-Cesena [FC]), Pieve di Rivoschio - Bagno di
Romagna, on dead land branch of Prunus sp., 30 April 2017, E. Camporesi, IT3337 (MFLU 170995, holotype); ibid. (PDD, isotype).
GenBank numbers – ACT: MG742353, ITS: MG742350, LSU: MG742351, RPB2:
MG742352, TEF1: MG742354.
Notes – Cytospora species are important plant pathogens causing dieback and canker diseases
on a wide range of hosts, such as Prunus sp. Norphanphoun et al. (2017a) found eleven species of
Cytospora on Prunus sp. with species identifications confirmed with molecular data. In this study,
the new species has immersed, 2–4-loculate conidiomata with ostioles and unicellular conidia.
Cytospora prunicola is most similar to C. sorbicola Norphanph. et al. and C. ulmi Norphanph. et al.
in conidial size (5.6 × 1.5, 5.4 × 1.4 respectively) (Norphanphoun et al. 2017a). However, C.
sorbicola differs from C. prunicola in having 1–2 loculate conidiomata with unbranched
conidiophores (Norphanphoun et al. 2017a). Phylogenetic analyses using a combined ITS, LSU,
RPB2 and ACT sequence dataset indicate that C. prunicola constitutes a moderately supported
independent lineage (Fig 66) and hence can be distinguished from C. sorbicola and C. ulmi and
other species. Therefore, we introduce C. prunicola as a new species from Prunus in Italy.
Diaporthaceae Höhn. ex Wehm.
Diaporthaceae was established by Wehmeyer (1926). This family is characterized by globose
or compressed, coriaceous, black ascomata with 0–1-septate, ellipsoid to fusoid ascospores and
378
Figure 68 – Cytospora prunicola (MFLU 17-0995, holotype). 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 Conidia attached to
conidiogenous cells. i Morphology drawing. j Conidia. Scale bars: d = 400 µm, f = 100 µm, e, g–h
= 10 µm, j = 5 µm.
aseptate, variously coloured conidia (Maharachchikumbura et al. 2016a, Gao et al. 2017).
Diaporthaceae was transferred to Valsaceae as a synonym (Barr 1978). However, Castlebury et al.
(2002) showed that Diaporthaceae taxa could be distinguished from others in Diaporthales based on
phylogenetic (2002) showed that Diaporthaceae taxa could be distinguished from others in
Diaporthales based on phylogenetic analyses of rDNA LSU sequence data. Senanayake et al.
(2017) listed Allantoporthe,Apioporthella, Chaetoconis, Chiangraiomyces, Diaporthe,
Hyaliappendispora,
Leucodiaporthe,
Mazzantia,
Ophiodiaporthe,
Paradiaporthe,
379
Phaeocytostroma, Phaeodiaporthe, Pustulomyces and Stenocarpella as members in this family
based on multi-gene analysis and morphology. In this study, we introduce two new species,
Diaporthe subellipicola and D. subcylindrospora, collected in China. The novel species are
described morphologically and selected phylogenetic datasets from Dissanayake et al. (2017) and
Gao et al. (2017) are used to determine its phylogenetic affinity with other species.
Figure 69 – Maximum likelihood phylogenetic tree generated from analysis of a combined ITS,
TEF and TUB sequences dataset for 47 taxa of Diaporthe with Diaporthe elaeagni-glabrae and D.
stictica as the outgroup taxa. ML support values greater than 50% and a best scoring tree with a
final optimization likelihood value of -7531.270123 is indicated above the nodes. The strain
numbers are noted after the species names. Ex-type strains are indicated in bold.
Diaporthe Nitschke
Diaporthe was introduced by Nitschke (1870) with Diaporthe eres as the type species.
Species of Diaporthe are well-known as pathogens, endophytes or saprobes on a diverse range of
host plants (Uecker 1988, Santos et al. 2011, Hyde et al. 2014, Udayanga et al. 2014a, b, 2015). The
380
genus is characterized by black to brown ascomata with hyaline to brown, ovoid to ellipsoid
ascospores and hyaline conidia (Dissanayake et al. 2017, Gao et al. 2017, Senanayake et al. 2017).
Diaporthe subellipicola S.K. Huang, T.C. Wen & K.D. Hyde, sp. nov.
Fig. 70
Index Fungorum number: IF554076; Facesoffungi number: FoF03855
Etymology – the name subellipicola refers to the ellipsoid spores.
Holotype – MFLU 17-1197
Saprobic on dead straw. Sexual morph Ascomata 245–280 μm diameter, scattered, immersed,
eventually the neck erumpent, unilocular, globose to subglobose, black. Neck about 2 mm long,
central, lined with periphyses. Peridium 15–25 μm diameter, membranaceous, composed of brown
to hyaline cells of textura angularis. Asci 48–65 × 10–15 μm ( = 57 × 13 μm, n = 20), 8-spored,
unitunicate, clavate, without pedicel, rounded at the apex, with a conspicuous refractive apical ring.
Ascospores 10–18 × 2–6 μm ( = 13 × 4 μm, n = 50), ellipsoid, hyaline, 0–1-septate, the apical cell
swellom, slightly constricted at the septum at maturity, smooth-walled. Asexual morph
Undetermined.
Culture characteristics – Colonies on PDA reaching 2 cm diameter after 7 days at room
temperature (~ 25 °C), circular, brown from above and reverse, with filamentous mycelium,
filiform at the margin, with rough surface and raised elevation.
Material examined – CHINA, Yunnan Province, Songming City, on dead wood, 21
December 2016, S.K. Huang (MFLU 17-1197, holotype), ex-type living culture, KUMCC 17-0153;
ibid. (HKAS99560, isotype).
GenBank numbers – β-tubulin: MG746634, ITS: MG746632, TEF1: MG746633.
Notes – Phylogenetic analysis of ITS, TEF and TUB sequence dataset indicates that
Diaporthe subellipicola belongs to Diaporthe (Fig. 69). Diaporthe subellipicola is closely related to
D. endophytica, an asexual species (Thompson et al. 2011, Gomes et al. 2013), but with weak
bootstrap support. Unfortunately, we could not obtain the asexual morph of D. subellipicola for any
further morphological comparison. The endophytic lifestyle of D. endophytica from Maytenus spp.
and Schinus spp., distributed in Brazil (Gomes et al. 2013) is different from our species, which is a
saprobe. Futher, comparison of the 544 bp across the ITS1-5.8S-ITS2 regions also reveals that there
are 5 bp (1%) differences when compared to D. endophytica (LGMF911). In the same way,
comparison of 712 bp of TUB region reveals 8 bp (1.2%) difference compared to D. endophytica
(LGMF911).
Diaporthe subcylindrospora S.K. Huang, T.C. Wen & K.D. Hyde, sp. nov.
Fig. 71
Index Fungorum number: IF554077; Facesoffungi number: FoF03854
Etymology – the name subfusispora refers to the subcylindrospora spores.
Holotype – MFLU 17-1195
Saprobic on dead branch of Salix. Sexual morph Ascomata 390–395 μm diameter, scattered
to gregarious, immersed, eventually the neck erumpent, unilocular, globose to subglobose, black.
Neck about 1 mm long, central, lined with periphyses. Peridium 45–65 μm diameter,
membranaceous, composed of brown to hyaline cells of textura angularis. Asci 52–55 × 7–10 μm
( = 55 × 9 μm, n = 20), 8-spored, unitunicate, cylindrical, without pedicellate, rounded at the
apex, with a conspicuous refractive apical ring. Ascospores 10–15 × 2–4 μm ( = 12 × 3 μm, n =
50), fusiform, hyaline, 0–1-septate, slightly constricted at the septum at maturity, smooth-walled.
Asexual morph Undetermined.
Culture characteristics – Colonies on PDA reaching 2 cm diameter after 7 days at room
temperature (25 °C), circular, pale brown from above and reverse, with filamentous mycelium,
filiform at margin, with rough surface and raised elevation.
Material examined – China, Yunnan Province, Dali City, on dead branch of Salix, 21
December 2016, S.K. Huang (MFLU 17-1195, holotype), ex-type living culture KUMCC 17-0151;
ibid. (HKAS 99547, isotype).
GenBank numbers – β-tubulin: MG746631, ITS: MG746629, TEF: MG746630.
381
Figure 70 – Diaporthe subellipicola (MFLU 17-1197, holotype). a Material. b Ascomata on host. c
Appearance of ascomata on host. d Ascoma in vertical section. e Squashed neck with light brown
apex. f Peridium. g–j Asci. k–n Ascospores. Scale bars: d–e = 100 µm, f = 50 µm, g–j = 20 µm, k–
n = 10 µm.
382
Figure 71 – Diaporthe subcylindrospora (MFLU 17-1195, holotype). a Herbarium material. b
Ascomata on host. c Appearance of ascomata on host. d Ascoma in vertical section. e Squashed
neck with light brown apex. f Peridium of neck with periphyses. g Peridium. h–l Asci (l: stained in
Melzer’s reagent). m–s Ascospores. t Germinating ascospores. Scale bars: e = 500 µm, d, f = 100
µm, g–l, t = 20 µm, m–s = 5 µm.
383
Notes – Phylogeny positions our new taxon in a moderately supported subclade (74%) with
D. salicicola, an asexual species (Fig. 69). A close phylogenetic affinity to D. cynaroidis is also
noted. The asexual morph of D. cynaroidis was isolated from Protea in South Africa (Gomes et al.
2013) and D. salicicola was isolated from Salix in Australia (Tan et al. 2013). However, the sexual
morph of Diaporthe subcylindrospora was found from Salix in China. We noted a 6 bp (1.1%)
difference across the 560 bp nucleotides of the ITS regions and 8 bp (1.7%) difference across the
484 bp of TUB region compared to D. salicicola. Diaporthe subcylindrospora is introduced as a
new species.
Diaporthomycetidae families incertae sedis
Barbatosphaeriaceae H. Zhang et al.
The family was established by Zhang et al. (2017a) with the type genus Barbatosphaeria
Réblová. It currently comprises three genera, namely Barbatosphaeria, Ceratostomella and
Xylomelasma, which share similar characters in having dark, long-necked, astromatic ascomata,
mostly surrounded by sparse mycelia, clavate asci, arising from ascogenous hyphae and mostly
ellipsoidal ascospores (Zhang et al. 2017a). Based on phylogenetic analysis, Zhang et al. (2017a)
placed this family in Diaporthomycetidae families incertae sedis. In the study, we provide an
updated tree for the family and introduce a new species, Barbatosphaeria aquatica, characterised
by different asexual form.
Barbatosphaeria Réblová
The genus Barbatosphaeria was introduced by Réblová (2007) for Calosphaeria barbirostris
(Fr.) Ellis & Everh. based on cultivation experiments, revision of the herbarium material and
phylogenetic analysis. It is characterized by dark ascomata with long decumbent necks and
ellipsoid to oblong, hyaline, septate ascospores in unitunicate, clavate asci with a non-amyloid
apical ring (Réblová 2007, Réblová et al. 2015, Zhang et al. 2017a). Réblová (2007) placed the
genus in the Sordariomycetes incertae sedis based on LSU sequence data. Réblová et al. (2015)
revised taxonomy of the genus and expanded it to include nine species. Zhang et al. (2017a) studied
Annulatascaceae-like taxa. Barbatosphaeria was accommodated in a new family
Barbatosphaeriaceae, along with Ceratostomella and Xylomelasma. The known asexual morphs are
Ramichloridium- and Sporothrix-like taxa (Réblová 2007, Réblová et al. 2015, Zhang et al. 2017a).
Barbatosphaeria aquatica N.G. Liu & K.D. Hyde, sp. nov.
Fig. 73
Index Fungorum number: IF554085; Facesoffungi number: FoF 03937
Etymology – with reference to the habitat of this fungus.
Holotype – MFLU 18-0040
Saprobic on decaying wood. Sexual morph Undetermined. Asexual morph Colonies on
natural substrate superficial, effuse, dark brown to black, velvety. Mycelium immersed, composed
of septate, branched, medium brown, 1.5–3 μm wide hyphae. Conidiophores 70–100 μm long, 4–6
μm wide at base, 1.5–2.2 μm wide at apex, mononematous, erect, straight or broadly curved,
greenish-brown to brown, smooth-walled, 4–6-septate, thick-walled, unbranched. Conidiogenous
cells 10–15 μm long, monoblastic, terminal, integrated, medium brown, smooth-walled. Conidia
20–27 × 10–17 μm ( = 22.22 × 13.29 μm, n = 13), solitary, dry, clavate, green to greenish-brown,
with 3–4 transverse septa, deeply constricted at the septa, distinctly verruculose, with 1–2 basal
cells subhyaline to pale brown, smooth-walled and appressed at the upper half.
Culture characteristics – Conidia germinated on WA (Water Agar) within 24 hours. One
germ tube produced from the basal cell. Colonies reaching about 5 cm diameter after 10 days on
PDA at 25 °C. Mycelia superficial, initially white, later becoming brown to dark brown, hairy,
effuse with the entire edge; reverse dark brown at center, light brown towards margin.
Material examined – THAILAND, Chiang Rai Province, Muang, Ban Nang Lae Nai, on
decaying wood submerged in a freshwater stream, 31 December 2016, Ningguo Liu, CR004
(MFLU 18-0040, holotype); ex-type living culture MFLUCC 18-0356.
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Figure 72 – Maximum likelihood (RAxML) tree based on analysis of a combined dataset of ITS
and LSU sequence data representing four families in Sordariomycetes. Related sequences were
obtained from GenBank. Twenty-seven stains are included in the analyses, which comprise 1526
characters including gaps. Tree topologies are similar between ML and BI analyses. The tree is
rooted with Catabotrys deciduum SMH3436. The best scoring RAxML tree with a final likelihood
value of -7652.009089 is presented. The matrix had 565 distinct alignment patterns, with 24.34% of
undetermined characters or gaps. Estimated base frequencies were as follows; A = 0.244210, C =
0.246103, G = 0.294155, T = 0.215533; substitution rates AC = 0.912854, AG = 1.825402, AT =
1.503492, CG = 1.089820, CT = 5.670850, GT = 1.000000; gamma distribution shape parameter α
= 0.470150. Bootstrap support values for ML greater than 70% and Bayesian posterior probabilities
greater than 0.80 are given near nodes. The scale bar indicates 0.05 changes. The ex-type strains are
indicated in bold and the new isolate is in bold and blue.
385
Figure 73 – Barbatosphaeria aquatica (MFLU 18-0040, holotype). a Specimen. b, c Colonies on
substrate. d, e Conidiophores and conidia. f Conidiogenous cell and conidium. g Germinated spore.
h–m Conidia. n Colony on PDA media. Scale bars: b = 100 μm, c–e = 15 μm, f, g = 10 μm, h–m =
5 μm.
386
GenBank numbers – ITS: MG835711, LSU: MG835712.
Notes – The asexual morphs of this genus are dematiaceous hyphomycetes with blastic
denticulate conidiogenesis and resemble Ramichloridium and Sporothrix (Réblová 2007). In this
study, a new asexual morph is accommodated in this genus. Unlike the other morphs, which have
ellipsoidal conidia with a hilum in Ramichloridium (Arzanlou et al. 2007) and tear-shaped conidia
on small, clustered denticles in Sporothrix (Zhou et al. 2014), the present asexual morph has
clavate, green to greenish-brown conidia with several transverse and longitudinal septa. Despite the
differences in morphology, the new strain formed a distinct clade in the genus Barbatosphaeria
with 79% ML bootstrap and 0.89 Bayesian PP support. Thus, a new species is introduced based on
both morphology and phylogenetic studies.
Subclass Hypocreomycetidae O.E. Erikss. & Winka
Hypocreales Lindau
Nectriaceae Tul. & C. Tul.
The family Nectriaceae is characterized by uniloculate ascomata that are white, yellow,
orange-red or purple. They are associated with phialidic asexual morphs producing amerosporous
to phragmosporous conidia (Rossman et al. 1999, Rossman 2000). Nectriaceae includes 55 genera
and approximately 900 species (Index Fungorum 2018, Wijayawardene et al. 2018). The majority
of these species are soil-borne saprobes or weak to virulent, facultative or obligate plant pathogens,
while some are facultatively fungicolous or insecticolous (Rossman et al. 1999, Rossman 2000,
Schroers et al. 2011). Several species have been reported as important opportunistic pathogens of
humans (Chang et al. 2006, Guarro 2013), while others produce mycotoxins of medical concern
(Rossman 1996). In this study, we introduce a new Gliocladiopsis species based on phylogenetic
and morphological evidence.
Gliocladiopsis S.B. Saksena
The genus Gliocladiopsis was introduced by Saksena (1954) based on the type species G.
sagariensis S.B. Saksena that had penicillate conidiophores resembling Penicillium and
Gliocladium. Crous & Wingfield (1993) resurrected Gliocladiopsis to accommodate species
characterized by dense, penicillate conidiophores, which unlike Cylindrocladiella and Calonectria,
lacked sterile stipe extensions. The taxonomic status of Gliocladiopsis was re-evaluated by
Lombard & Crous (2012) based on multi-gene phylogeny. Liu & Cai (2013) introduced a new
Gliocladiopsis species, G. guangdongensis, which was the first isolation from a freshwater habitat.
Species of Gliocladiopsis are difficult to distinguish morphologically as the branching structure of
conidiophores as well as the size and shape of conidia, are similar between species (Liu & Cai
2013, Parkinson et al. 2017). Currently Gliocladiopsis contains 13 species, namely, G. curvata, G.
elghollii, G. forsbergii, G. guangdongensis, G. indonesiensis, G. irregularis, G. mexicana, G.
peggii, G. pseudotenuis, G. sagariensis, G. sumatrensis, G. tenuis and G. whileyi (Lombard &
Crous 2012, Liu & Cai 2013, Parkinson et al. 2017). In this paper, we introduce a new
Gliocladiopsis species, G. aquaticus, based on multi-gene phylogenetic analyses, which is the
second taxon from freshwater habitat.
Gliocladiopsis aquaticus Y.Z. Lu, R.H. Perera & K.D. Hyde, sp. nov.
Fig. 75
Index Fungorum number: IF554048; Facesoffungi number: FoF03893
Etymology – ‘aquaticus’ referring to aquatic habitat of this fungus.
Holotype – MFLU 17-1976
Saprobic on submerged decaying wood in a lake. Sexual morph Undetermined. Asexual morph
appearing as white masses on the substrate, becoming yellowish with age. Conidiophores
penicillate, 100–140 × 4–6.5 μm, without stipe extensions and terminal vesicles. Conidiogenous
apparatus with several series of hyaline branches: primary branches 19–26 × 3.5–4.5 μm, aseptate;
secondary branches 15–25 × 3–4 μm, aseptate; tertiary branches 8–15 × 2–3.5 μm, aseptate;
phialides doliiform to cymbiform to cylindrical, 11–19 × 2–3 μm, arranged in terminal whorls of 2–
387
Figure 74 – Phylogenetic tree generated by maximum likelihood analysis of the combined ITS, βtub and H3 dataset of Gliocladiopsis species. Related sequences were obtained from GenBank.
388
Fourty-two strains are included in the analyses, which comprises 1697 characters including gaps.
The tree is rooted with Cylindrocladiella parva (ATCC 28272). Tree topology of the ML analysis
was similar to the BI. The best scoring RAxML tree with a final likelihood value of - 5668.766010
is presented. The matrix had 381 distinct alignment patterns, with 5.77% of undetermined
characters or gaps. Estimated base frequencies were as follows: A = 0.219613, C = 0.322518, G=
0.235367, T = 0.222502; substitution rates AC = 0.926729, AG = 1.749796, AT = 0.794257, CG =
0.329025, CT = 3.846114, GT = 1.000000; gamma distribution shape parameter α = 0.177707.
RAxML bootstrap support values greater than 50% (BT) (before the forward slash) and Bayesian
posterior probabilities greater than 0.90 (PP) (after the forward slash) are shown near the nodes.
The ex-type strains are in bold and new isolates in blue.
6 per branch, with minute collarettes, central phialide frequently extending above the rest. Conidia
cylindrical, 16.5–21 × 2–3 μm ( = 19 × 2.5 μm, n = 50), hyaline, smooth with rounded ends,
straight, aseptate to 1-septate.
Culture characteristics – Conidia germinating on water agar (WA) within 8 hours. Colonies
growing on PDA, circular, with flat surface, edge entire, reaching 36 mm within 2 weeks at 28 °C,
white to pale brown, mycelium superficial and partially immersed, branched, septate, hyaline to
pale brown, smooth-walled.
Material examined – THAILAND, Chiang Rai Province, Mae Fah Luang University, on
submerged decaying wood in a freshwater lake, 19 February 2017, Yong-Zhong Lu, MFU07
(MFLU 17-1976, holotype); ex-type living culture, MFLUCC 17-1811, TBRC; MFU08 (MFLU
17-1977, paratype); living culture, MFLUCC 17-2028.
GenBank numbers – MFLUCC 17-1811 – β-tubulin: MG574421, Histone3: MG734182, ITS:
MG543924, LSU: MG543915, SSU: MG543918; MFLUCC 17-2028 – β-tubulin: MG574422,
Histone3: MG734183, ITS: MG543925, LSU: MG543916, SSU: MG543919.
Notes – Gliocladiopsis aquaticus is morphologically similar to Gliocladiopsis elghollii in
conidiophores and conidia but can be distinguished from G. elghollii by its smaller conidia (16.5–
21 × 2–3 vs. 19–23 × 2–4 μm) and different series of conidiogenous apparatus branches (3–4
branches vs. 5 branches). Phylogenetically, two isolates of G. aquaticus formed one subclade with
good support (1.00 PP, 100% ML) and shares a sister relationship to G. elghollii.
Ophiocordycipitaceae G.H. Sung et al.
The family Ophiocordycipitaceae (order: Hypocreales) was introduced by Sung et al. (2007)
based on phylogenetic analyses and later listed by Kirk et al. (2013) and Quandt et al. (2014). Kirk
et al. (2013) accepted eleven genera in Ophiocordycipitaceae, but Quandt et al. (2014) refined and
proposed six genera, Drechmeria, Harposporium, Ophiocordyceps, Polycephalomyces,
Purpureocillium and Tolypocladium to comprise the family. Matočec et al. (2014) introduced
Perennicordyceps under Ophiocordycipitaceae. Maharachchikumbura et al. (2015) confirmed this
system and Spatafora et al. (2015) introduced some necessary species combinations based on this
classification. Simmons et al. (2015) refined the genus Hirsutella under Ophiocordycipitaceae.
Most species of Ophiocordycipitaceae produce darkly pigmented, tough to pliant stromata that
often possess aperithecial apices (Sung et al. 2007). The main distinguishing characters of this
genus are that the ascospores do usually not break into part-spores at maturity and asci have thin
apical caps (Petch 1931, 1932). In the study, we introduce one new species Ophiocordyceps
cylindrospora based on phylogeny and morphology.
Ophiocordyceps Petch
Ophiocordyceps was introduced by Petch (1931) and revised as the type genus under the
family Ophiocordycipitaceae by Sung et al. (2007), with the type species Ophiocordyceps blattae
(Petch) Petch, which was a cockroach pathogen. This genus includes most of the species in the
family and some of the species are host-specific insect pathogens (Quandt et al. 2014).
389
Figure 75 – Gliocladiopsis aquaticus (MFLU 17-1976, holotype). a, b Colonies on substrate. c, d
Conidiophores. e, f Conidiogenous cells with attached conidia. g Conidia. h, i Colony on PDA from
above and below. Scale bars: a = 200 µm, b = 500 µm, c, d, g = 20 µm, e, f = 10 µm, h, i = 20 mm.
390
Figure 76 – Phylogram of Ophiocordyceps cylindrospora generated from maximum likelihood
analysis of ITS, SSU, LSU and RPB2 sequence data. Tolypocladium inflatum (OSC 71235) and
Tolypocladium ophioglossoides (NBRC 106332) were used as outgroup taxa. The tree topology of
the ML was similar to the ML and BI analysis. The best scoring RAxML tree with a final
likelihood value of -22779.351826 is presented. The matrix had 1558 distinct alignment patterns,
with 45.49% of undetermined characters or gaps. Estimated base frequencies were as follows; A =
0.259547, C = 0.235029, G = 0.269944, T = 0.235480; substitution rates AC = 1.192392, AG =
3.825106, AT = 1.220677, CG = 0.895628, CT = 6.933089, GT = 1.000000; gamma distribution
shape parameter α = 0.202843. Maximum likelihood bootstrap values greater than 75% and
Bayesian posterior probabilities over 0.95 are indicated above the nodes. The scale bar indicates
0.04 changes. The new species was in red.
Ophiocordyceps cylindrospora Y.P. Xiao, T.C. Wen & K.D. Hyde, sp. nov.
Fig. 77
Index Fungorum number: IF553983; Facesoffungi number: FoF03878
Etymology – the specific epithet refers to Thanathon in north Thailand, the collection
location.
Holotype – MFLU 17-1961
Parasitic on wasps (Hymenoptera), collected from the underside of leaf litter. Sexual morph
Host 11 mm long, 3–5 mm wide, black wasp without hyphae on the surface. Stromata 50–90 mm
long, 0.5–1.5 mm diameter, mostly single, stipitate, rarely double, arising from between the head
and thorax of adult wasps, pale yellow to yellow. Stipe 25–27 mm long, 0.5–2 mm diameter,
391
clavate, with a fertile apex, becoming pale to yellow. Fertile head 3–3.2 mm wide, 1–1.2 mm
diameter, fusiform, yellow, upper surface roughened, lateral surface, ridged and furrowed.
Ascomata 551–638 × 261–327 μm ( = 594 × 294 µm, n = 30), immersed, oblique flask-shaped,
thin-walled. Peridium 20–25 mm ( = 23 µm, n = 60) wide. Asci 248–313 × 5–7 μm ( = 480 × 6
µm, n = 60), 8-spored, hyaline, narrow cylindrical, with a thickened apex. Apical cap 5–12.5 × 8.5–
10.4 μm ( = 8.7 × 9.5 µm, n = 60), with a small channel in the center. Ascospores easily breaking
into part-spores, filiform. Secondary ascospores 3.1–3.9 × 1.6–2 μm ( = 3.5 × 1.8 µm, n = 90)
cylindrical, truncated at both ends, straight, hyaline, smooth. Asexual morph Undetermined.
Material examined – THAILAND, Chiang Mai Province, Mushroom Research Centre, on
dead wasp, 19 July 2015, YuanPin Xiao, MSCM15071905 (MFLU 17-1961, holotype); ibid.
(MFLU 17-1962, MFLU 17-1963 isotypes).
GenBank numbers – ITS: MG553635, LSU: MG553652, RPB2: MG647029, SSU:
MG553651.
Notes – Ophiocordyceps cylindrospora is a pathogen of wasps and was collected from
Chiang Mai, Thailand. The morphology of the new species is similar to Ophiocordyceps
sphecocephala, O. myrmecophila, O. pseudolloydii, O. hemisphaerica, O. irangiensi, which are
shown in Table 3. The morphology of Ophiocordyceps cylindrospora differs from other species in
the genus Ophiocordyceps in having cylindrical and shorter secondary ascospores. Phylogenetic
analyses of combined SSU, LSU, ITS, RPB1 and RPB2 sequence data also support it as a new
species in the genus (Fig. 76).
Table 3 Synopsis of Ophiocordyceps species discussed in the paper.
Species
Host
Stromata
(mm)
Ascomata
(μm)
Asci
(μm)
Ascospore
s (μm)
O. cylindrospora
Wasp
50–90 ×
0.5–1.5
551–638 ×
261–327
248–313 ×
5–7
As long as
asci
Secondary
ascospores
(μm)
3.1–3.9 ×
1.6–2
O. myrmecophila
Ant
10–40 ×
0.2–0.5
600–890 ×
180–275
460–630 ×
4–6,5
310–390 ×
1–2
8–10 ×
1.5–2.3
cylindrical
O. sphecocephala
Wasp
45 × 1.4–
1.8
880–1000
× 200–260
700 ×7
As long as
asci
10–14 ×
1.5–2.5
O. irangiensi
Ant
25–60 ×
0.2–0.7
1000 ×
150–200
900 × 6–8
700 × 1.3–
2.3
8.5–12.5 ×
1.3–2.3
fusoid
O. hemisphaerica
Fly
12–20
×0.8–1
780–860 ×
220–290
500–640 ×
5–6
-
-
O. pseudolloydii
Ant
2–6 ×
0.5–2
380–550 ×
140–240
240–485 ×
12–16
160–470 ×
5–6
4.5–6.3 ×
1–2.5
cylindrical
Reference
This study
Mains
1940,
LuangsaArd et al.
2008
HywelJones
1995
HywelJones1996
Hyde et al.
2016
Evans &
Samson
1984
Stachybotryaceae Crous
The family Stachybotryaceae was introduced by Crous et al. (2014) within the order
Hypocreales (Hypocreomycetidae, Sordariomycetes). Thirty-four genera are accepted in
Stachybotryaceae based on morphology and multi-gene phylogeny and the family is typified by
Stachybotrys Corda (Crous et al. 2014, Lombard et al. 2016, Gordillo & Decock 2017,
Wijayawardene et al. 2016, 2018). The species are saprobes or pathogens and some may cause risk
to human health (Lombard et al. 2016). In the study, we introduce a saprobic Alfaria terrestris as a
new record in Thailand from dead grass leaves.
392
Figure 77 – Ophiocordyceps cylindrospora (MFLU 17-1961, holotype). a Habitat of
Ophiocordyceps cylindrospora. b Ascostroma emerging from infected wasps. c Overview of host. d
Fertile head of ascostroma. e, f Perithecia. g Peridium. h–j Immature to mature asci. k Part of the
ascospores. l Apical cap of asci. m, n Secondary ascospores. Scale Bars: d, e = 500 µm, f = 200
µm, g = 20 µm, h–k = 100 µm, l = 10 µm, m, n = 2 µm.
393
Alfaria Crous et al.
The genus Alfaria belongs in the family Stachybotryaceae (Crous et al. 2014,
Maharachchikumbura et al. 2015, 2016a, Wijayawardene et al. 2016, 2017a). Alfaria was
introduced to accommodate pathogenic A. cyperi-esculenti Crous, N.J. Montaño-Mata & GarcíaJim (Crous et al. 2014). Nine species are currently accepted in Alfaria (Lin et al. 2017).
Figure 78 – Phylogram generated from maximum likelihood analysis based on combined LSU and
ITS sequenced data of Alfaria species. Related sequences were obtained from Lin et al. (2017).
Fifteen strains were included in the combined sequence analyses, which comprise 1424 characters
with gaps. Myrothecium simplex (CBS 582.93) was used as the outgroup taxa. Tree topology of the
ML analysis was similar to the BI. The best scoring RAxML tree with a final likelihood value of 3363.437417 is presented. The matrix had 290 distinct alignment patterns, with 20% of
undetermined characters or gaps. Estimated base frequencies were as follows; A = 0.242693, C =
0.256230, G= 0.288945, T = 0.212132; substitution rates AC = 1.096356, AG = 0.714721, AT =
0.772509, CG = 0.707572, CT = 2.464121, GT = 1.000000; gamma distribution shape parameter α
= 0.451441. Bootstrap support values for ML equal to or greater than 50% and BYPP equal to or
greater than 0.90 are given above the nodes respectively. Newly generated sequences are in red
bold. The ex-type strains are in black bold.
394
Alfaria terrestris L. Lombard & Crous, Persoonia 36:181 (2016)
Fig. 79
Facesoffungi number: FoF04082
Saprobic on dead leaves of Grass (Poaceae) species. Sexual morph Undetermined. Asexual
morph Conidiomata 250–400 µm ( = 310 µm, n = 10) diameter, superficial, solitary, black, oval
to elongate or irregular and a green agglutinated mass of conidia surrounded by setae. Setae straight
or bent, dark brown, arising from the basal of conidiomata. Conidiophores unbranched, green,
smooth, up to 11 µm long. Conidiogenous cells 4–7 × 1–2 µm ( = 5.1 × 1.2 µm, n = 15),
phialidic, cylindrical, hyaline, smooth-walled, cylindrical. Conidia 11–14 × 2–3 µm ( = 12 × 2
µm), smooth, light green to green, ellipsoidal and straight.
Material examined – THAILAND, Chiang Rai Province, Amphur Muang, on leaves of dead
grass species, 23 September 2016, N. I de Silva, NI113 (MFLU 17-0679).
GenBank numbers – ITS: MG818852, LSU: MG821635.
Notes – Our strain clusters with the type Alfaria terrestris (CBS 477.91) in the combined
LSU and ITS phylogenetic analysis. We were unable to get the culture from conidia. Therefore, we
obtain sequence data directly from fruiting bodies. Alfaria terrestris has been isolated from soil in
Turkey and the USA and leaf litter in Spain. The current collection of A. terrestris was from leaves
of dead grass species in Chiang Rai, Thailand. Our strain has aseptate, smooth-walled, ellipsoidal
conidia that are similar to type collection. However, our strain has light green to green, 11–14 × 2–
3 µm (x = 12 × 2 µm) conidia and the type specimen had hyaline (4–)5–7 × 2–3 µm (av. 6 × 2 µm)
conidia (Lombard et al. 2016), which suggests this may be a species complex. Our strain produces
setae that arise from the base of conidiomata that has previously not been observed in the type.
Subclass Sordariomycetidae O.E. Erikss & Winka
Chaetosphaeriales Huhndorf et al.
Chaetosphaeriaceae Réblová et al.
The family Chaetosphaeriaceae was introduced by Locquin (1984) to accommodate
Chaetosphaeria Tul. & C. Tul., Loramyces W. Weston, Niesslia Auersw., Rhagadostoma Körb.
and Zignoëlla Sacc., but it was not validly published (Réblová et al. 1999). Réblová et al. (1999) redescribed and validated Chaetosphaeriaceae and accepted Ascocodinaea, Chaetosphaeria,
Melanochaeta, Melanopsammella, Porosphaerella, Porosphaerellopsis and Striatosphaeria.
Presently, 38 genera were accepted within the family Chaetosphaeriaceae (Maharachchikumbura et
al. 2016a, Wijayawardene et al. 2018). This family comprises the asexual morph genera
Catenularia, Cylindrotrichum, Chalara, Chloridium, Custingophora, Dictyochaeta, Menispora,
Phialophora and Zanclospora (Réblová et al. 1999, Ho et al. 2001, Huhndorf et al. 2001,
Fernández & Huhndorf 2005, Fernández et al. 2006, Atkinson et al. 2007, Liu et al. 2016, Perera et
al. 2016, Wijayawardene et a. 2017a, 2018). In this study, we introduce Chaetosphaeria mangrovei
sp. nov. from an intertidal mangrove host, based on both morphological and phylogenetic analyses
and designate an epitype for Cryptophiale hamulata.
Chaetosphaeria Tul. & C. Tul.
Chaetosphaeria is placed in Chaetosphaeriaceae (Chaetosphaeriales) (Réblová et al. 1999,
Huhndorf et al. 2004, Maharachchikumbura et al. 2015, 2016a) based on molecular data, although
it previously placed in Lasiosphaeriaceae by Barr (1990). Morphological characters of the sexual
morphs are simple and difficult to differentiate, while the asexual morphs characters are considered
as distinguishable (Gams & Holubová-Jechová 1976, Huhndorf et al. 2004). Taxa that belong to
this genus are reported from decayed plant material in terrestrial, freshwater and marine habitats,
worldwide and comprise more than 100 species (Ho et al. 2001, Huhndorf et al. 2001, Fernández &
Huhndorf 2005, Fernández et al. 2006, Atkinson et al. 2007, Liu et al. 2016, Perera et al. 2016,
Wijayawardene et al. 2017a).
Chaetosphaeria mangrovei Dayar., E.B.G. Jones & K.D. Hyde, sp. nov.
Index Fungorum number: IF554116; Facesoffungi number: FoF03927
Fig. 81
395
Figure 79 – Alfaria terrestris (MFLU 17-0679). a–c Appearance of conidiomata on host. d, e
Squash mounts of conidiomata. f Conidiogenous cells. g Setae arising from base of conidiomata. h,
i Conidia. Scale bars d, e = 50 μm, f = 5 μm, g = 20 μm h, i = 5 μm.
Etymology – name referring the host, of which the species was collected.
Holotype – MFLU 18-0146
Saprobic on wood. Sexual morph Ascomata 250–350 μm diameter, 300–450 μm high,
superficial, arranged in clusters, ovoid, dark brown, surface rough, ostiolate. Ostiole periphysate.
Peridium composed of dark brown cells of textura angularis in surface view, 15–20 μm thick in
longitudinal section, 2-layered, inner layer 3–5 cells thick, composed of hyaline elongate cells of
textura angularis, with outer layer 6–10 cells thick, composed of brown to dark brown, cell layers
of textura angularis. Paraphyses 3–5 μm wide, sparse, simple, septate. Asci 75–80 × 8–10.5 ( =
82 × 10 μm, n = 20) μm, 8-spored, unitunicate, cylindrical-clavate, short-pedicellate, with a
conspicuous, refractive, J-, apical ring. Ascospores 16–18.5 × 3–4.2 ( = 18 × 3.5 μm, n = 20) μm,
overlapping biseriate, hyaline, ovoid-fusiform, 3–5-septate, straight, with small guttules. Asexual
morph Undetermined.
396
397
Figure 80 – Phylogram generated from maximum likelihood analysis based on combined LSU, and
ITS sequenced data. One hundred strains are included in the analyses, which comprise 1470
characters including gaps. The tree is rooted to Gelasinospora tetrasperma (CBS 178.33) and
Sordaria fimicola (CBS 508.50). Tree topology of the ML analysis was similar to the BI. The best
scoring RAxML tree with a final likelihood value of -17707.998520 is presented. The matrix had
792 distinct alignment patterns, with 36.43% of undetermined characters or gaps. Estimated base
frequencies were as follows; A = 0.233261, C = 0.262150, G = 0.300958, T = 0.203631;
substitution rates AC = 1.405832, AG = 2.262138, AT = 2.187639, CG = 0.916401, CT =
7.644284, GT = 1.000000; gamma distribution shape parameter α = 0.264850. Maximum
likelihood bootstrap (ML, black) values > 60% and Bayesian posterior probabilities (PP, blue) >
0.80% are given above the nodes.
Culture characteristics – Colonies on MDA reaching 10 mm diameter after 2 weeks at 20–
25°C, dense, circular, slightly raised, surface smooth with even edge, cottony, colony from above:
white at the margin, grayish at the centre; from below: white at the margin, brownish-yellow at the
centre; not producing pigmentation on PDA media.
Material examined – THAILAND, Ranong Province, Amphoe Muang, Mu 4 Tambol Ngao,
Ranong Mangrove Research Center (GPS: 9°43' to 9°57'N; 98°29' to 98°39'E) on decaying wood of
mangrove species, 7 December 2016, Monika C. Dayarathne, MCD 069 (MFLU 18-0146,
holotype); ex-type living culture, MFLUCC 18-0144, TBRC.
GenBank numbers – LSU: MG813818, SSU: MG813819.
Notes – Chaetosphaeria mangrovei, the second Chaetosphaeria species reported from a
marine based habitat is unique in having fusiform, 3–5-septate, straight ascospores with small
guttules. However, Jones et al. (2014) reported that previously reported marine Chaetosphaeria
species, C. chaetosa formed a monophyletic group with Juncigena adarca, Moheitospora fruticosa
and two Fulvocentrum species, with high bootstrap support in a new family Juncigenaceae and this
was followed in Jones et al. (2015). Hence, this the only record of marine Chaetosphaeria species
at present. However, we did not observe the setae, enclosing the ascomata which is a characteristic
feature of this genus (Fernández & Huhndorf 2005, Perera eta al. 2016). Maximum likelihood and
Bayesian analyses of combined LSU and ITS sequence data indicate that C. mangrovei is a
moderately supported lineage nested in between C. innumera Berk. & Broome ex Tul. & C. Tul.
(SMH 2748), C. lentomita W. Gams & Hol.-Jech. (MR 1265), C. lignicola (Munk) Tomilin (CBS
143.54) and C. talbotii S. Hughes, W.B. Kendr. & Shoemaker (EXP0560F) (Fig. 80), which is
suggestive of a new species status. Chaetosphaeria innumera is different from our species in
having smaller ascospores (3.5–4.5 × 1.3–1.7 μm; Fernández & Huhndorf 2005). Chaetosphaeria
lentomita is also morphologically similar to C. mangrovei, but C. lentomita possesses fairly longstalked asci with an obtuse apex holding an inconspicuous apical ring and bi-seriately arranged, 1septate ascospores with two guttules (Réblová et al. 1999).
Cryptophiale Piroz.
The genus Cryptophiale was established by Pirozynski (1968) to accommodate C.
kakombensis Piroz. and C. udagawae Piroz. & Ichinoe. Presently, 20 species are accepted in the
genus (Whitton et al. 2012). Each is characterized by having unbranched or apically dichotomous
or verticillate, setiform conidiophores with a subapical or apical fertile region that produce hyaline,
unicellular to multiseptate conidia (Seifert et al. 2011). The conidiogenous cells are in rows
obscured by a shield of sterile cells (Seifert et al. 2011). The conidia are produced in slimy masses
on one side of the conidiophore (Seifert et al. 2011).
Cryptophiale hamulata Whitton, K.D. Hyde & McKenzie, in Whitton, McKenzie & Hyde, Fungal
Diversity Res. Ser. 21: 174 (2012)
Fig. 83
Index Fungorum number: IF554379; Facesoffungi number: FoF04383
398
Figure 81 – Chaetosphaeria mangrovei (MFLU 18-0146, holotype). a Specimen. b Appearance of
superficial ascomata on host surface. c Vertical section of an ascoma. d Section through peridium. e
Paraphyses f–h Asci. i Close up of an apical ring of an asci. j–k Ascospores. l Germinating
ascospore. m–n Culture on MEA (m-upper, n-lower). Scale bars: b = 200 μm, c = 100 μm, d–e = 20
μm, f–h = 50 μm, i–l = 10 μm.
399
Figure 82 – Phylogenetic tree generated from maximum likelihood (ML) analysis based on
combined LSU and ITS rDNA sequence data for selected genera within the family
Chaetosphaeriaceae. Bootstrap support values for maximum likelihood and maximum parsimony
greater than 50% are indicated above or below the nodes as ML/MP. Ex-type strains are in bold,
the new isolate is in red. The tree is rooted with Lasiosphaeria ovina (SMH 4605) and
Endophragmiella dimorphospora (FMR 12150).
400
Saprobic on decaying leaves. Asexual morph Colonies effuse, hairy, consisting of tall,
scattered individual and fertile conidiophores. Mycelium predominantly immersed, partially
superficial. Separate setae absent. Conidiophores macronematous, mononematous, solitary,
scattered, erect, straight or flexuous, subulate, dark brown, smooth, thick-walled, multiseptate, base
swollen, tapering to the apex, with the upper part sterile, setiform, simple, sometimes with a single
dichotomously branching tip, 170–300 μm ( = 225 μm, n = 20) long, 5.5–10.5 μm ( = 8.2 μm, n
= 20) wide just above the base, 4–6.5 μm ( = 5.4 μm, n = 20) wide below the fertile region.
Fertile region on the sub-apex of conidiophores, septate, dark brown, cylindrical, 21.5–81.5 μm (
= 59.8 μm, n = 20) long, 9.5–15 μm ( = 12.6 μm, n = 20) wide. Conidiogenous cells
monophialidic, discrete, in rows obscured by the shield of sterile cells, determinate, subsphaerical
or lageniform. Conidia aggregated in slimy masses on one side of the conidiophore, simple,
hyaline, smooth-walled, 0(–1)-septate, falcate, elongate to nearly filiform, lower end truncate to
rounded, upper end appendiculate, 13–23 μm ( = 18.67 μm, n = 55) long, 1.6–3.4 μm ( = 2.42
μm, n = 55) wide. Sexual morph Undetermined.
Culture characteristics – Conidia germinating on PDA within 12 hours. Colonies on PDA
effuse, dark brown from above, dark from below, reaching a diameter of 3–5 cm in 10 days at
25°C.
Material examined – THAILAND, Chiang Mai Province, Mae Taeng, Ban Pa Deng,
Mushroom Research Centre, on decaying leaf, 24 August 2016, Chuan-Gen Lin, MRC 11-1
(MFLU 17-1263, epitype designated here), ex-epitype living culture, MFLUCC 18-0098, KUMCC;
ibid. (HKAS 101821)
GenBank numbers – LSU: MG386756, SSU: MG386757.
Notes – The combined LSU and ITS phylogenetic analysis showed that Cryptophiale clusters
together with Morrisiella and Stanjehughesia (Fig. 82) but with weak support but these taxa are
morphologically distinct. Keys and synopses to the genus Cryptophiale were provided by Whitton
et al. (2012) and Goh & Hyde (1996). The specimen observed in this study agrees with the original
description given in Whitton et al. (2012), except that sometimes conidia are 1-septate. This study
is the first report of C. hamulata in Thailand which was previously known from the Philippines. We
designate an epitype for the species (sensu Ariyawansa et al. 2014). This is the first study yielding
DNA sequence data for the genus Cryptophiale.
Subclass Savoryellomycetidae Hongsanan et al.
Pleurotheciales Réblová & Seifert
Pleurotheciaceae Réblová & Seifert
Réblová et al. (2016) introduced the monotypic order Pleurotheciales based on morphological
characters and phylogenetic analyses. Pleurotheciaceae was typified by Pleurothecium with an
asexual species Pleurothecium recurvatum (Morgan) Höhn. as the type species. We introduce two
new species of Pleurotheciella and a new species of Phaeoisaria. An updated tree for the order
Pleurotheciales is provided in Fig. 84.
Phaeoisaria Höhn.
Phaeoisaria was established by Höhnel (1909) with P. bambusae Höhn. (now known as P.
clematidis (Fuckel) S. Hughes) as the type species. The genus is characterized by indeterminate
synnemata (except P. fasciculata Réblová & Seifert) with septate or aseptate ellipsoidal, obovoidal,
fusiform-cylindrical or falcate conidia, formed on a sympodially extending rachis (Ruiz et al. 2002,
Réblová et al. 2016). Presently, 23 species are accepted in the genus.
Phaeoisaria guttulata J. Yang & K.D. Hyde, sp. nov.
Index Fungorum number: IF554233; Facesoffungi number: FoF04086
Etymology – referring to the guttulate conidia.
Holotype – MFLU 18-0139
Fig. 85
401
Figure 83 – Cryptophiale hamulata (MFLU 17-1263, epitype). a, b Host material. c, d
Conidiophores on leaf surface. e, f Conidiophores, conidiogenous cells and conidia. g
Conidiogenous cells. h–j Conidia. Scale bars: c, d = 100 μm, e, f = 50 μm, g–j = 10 μm.
Saprobic on submerged decaying wood. Sexual morph Undetermined. Asexual morph
Colonies effuse, scattered, brown, hairy, visible as solitary, dark brown conidiophores with white
mass of conidia on the upper part. Mycelium partly immersed, partly superficial. Conidiophores
macronematous, synnematous, erect, septate, smooth, mid brown to dark brown, 480–700 µm long,
2–5 µm wide. Synnemata erect, rigid, dark brown to black, velvety, smooth, composed of compact
and parallel adpressed conidiophores, with flared conidiogenous cells in the above half.
Conidiogenous cells integrated, terminal, polyblastic, pale brown to hyaline, sympodial, splaying
out with one to several denticulate conidiogenous cells loci. Conidia globose to obovoid, hyaline,
smooth-walled, guttulate, aseptate, 3.5–5.5 × 2.5–4.8 µm ( = 4.5 × 3.5 µm, n = 20).
Culture characteristics – Conidia germinating on PDA within 24 hours. Germ tubes produced
from both ends. Colonies on PDA, reaching 5–10 mm diameter after two weeks at
25°C in natural light, with greyish-green aerial mycelium and brown mycelium on the
surface, dark brown in reverse, margin filamentous.
402
Material examined – CHINA, Guizhou Province, Anshun City, Gaodang Village, 26°4′16″N,
105°41′53″E, on decaying wood submerged in Suoluo River, 19 October 2016, Jing Yang, GD 1-1
(MFLU 18-0139, holotype); ex-type living cultures, MFLUCC 17-1965, ICMP.
GenBank numbers – ITS: MG837021, LSU: MG837016, SSU: MG837026.
Notes – Phylogenetic analyses indicated Phaeoisaria guttulata belongs to Phaeoisaria and
clusters with P. aquatica with little support (Fig. 84). Phaeoisaria guttulata can easily be
distinguished from previously described species of Phaeoisaria by its morphology. Among the
species sampled in our molecular dataset, P. guttulata is the only one which is characterized by
globose to obovoid conidia, while others have clavate, narrowly ellipsoidal to obovoid conidia.
Phaeoisaria guttulata is most similar to P. clavulata (Grove) E.W. Mason & S. Hughes in conidial
shape, however, the conidia of P. guttulata (3.5–5.5 × 2.5–4.8 µm) are larger than P. clavulata (1–2
µm diam) (Révay 1985).
Pleurotheciella Réblová et al.
The holomorph genus Pleurotheciella was introduced by Réblová et al. (2012) with two
species, Pl. rivularia Réblová et al. and Pl. centenaria Réblová et al., which have astromatic
perithecia, unitunicate asci, persistent paraphyses and hyaline, septate ascospores and holoblastic,
denticulate conidiogenous cells and hyaline, septate conidia. Based on morphological characters
and phylogenetic analyses, Dactylaria uniseptata Matsush. was transferred to Pleurotheciella as Pl.
uniseptata (Matsush.) Seifert (Réblová et al. 2016). Recently, Luo et al. (2017) introduced six new
species from freshwater habitats. Thus, nine species from water-saturated decaying wood are
accepted in Pleurotheciella with good descriptions and sequence data (Réblová et al. 2012, 2016,
Luo et al. 2017).
Pleurotheciella krabiensis J. Yang & K.D. Hyde, sp. nov.
Fig. 86
Index Fungorum number: IF554234; Facesoffungi number: FoF04087
Etymology – referring to the collection site.
Holotype – MFLU 18-0140
Saprobic on submerged decaying wood. Sexual morph Undetermined. Asexual morph
Colonies effuse, scattered, brown, hairy, upper part covered with bright white mass of conidia.
Mycelium partly immersed, partly superficial. Conidiophores macronematous, synnematous,
septate, smooth, brown, becoming paler towards the apex, straight or slightly curved, splaying out
at the apex, 240–390 × 3.3–4.8 µm ( = 310 × 4.3 µm, n = 15). Conidiogenous cells integrated,
terminal, polyblastic, sympodial, with cylindrical or tapering apex, pale brown or hyaline,
denticulate, denticles conspicuously cylindrical. Conidia fusiform, subcylindrical to obovoidsubclavate, hyaline, smooth-walled, often guttulate, straight or slightly curved, with a single
median septum, 19–25 × 4.5–6 µm ( = 22 × 5.3 µm, n = 20), obtuse at the apex, pointed at the
base.
Culture characteristics – Conidia germinating on PDA within 24 hours. Germ tubes produced
from both ends. Colonies on PDA slow growing, reaching 7–10 mm diameter after one month at 25
°C in natural light, circular, with yellowish-brown, dense aerial mycelium on the central surface
and white, sparse mycelium on the entire margin, yellowish-brown in reverse.
Material examined – THAILAND, Krabi Province, on decaying wood submerged in a
freshwater stream near Morakot Lake, 15 December 2015, S. Tibpromma, Site 6-6-3 (MFLU 180140, holotype); ex-type living cultures, MFLUCC 16-0852, ICMP.
GenBank numbers – ITS: MG837018, LSU: MG837013, SSU: MG837023.
Notes – The combined LSU, SSU and ITS phylogenetic analyses indicate that Pleurotheciella
krabiensis nested within Pleurotheciella as a phylogenetically distinct species. Pleurotheciella
krabiensis and Pl. guttulata Z.L. Luo et al. differ from other species in the genus by their
synnematous conidiophores, and the brown conidiophores in Pl. krabiensis are distinguished from
the hyaline conidiophores in Pl. guttulata. The occurrence of Pl. krabiensis on water saturated
403
decayed wood is consistent with the ecology of the other nine species in the genus (Réblová et al.
2012, 2016, Luo et al. 2017).
Figure 84 – Maximum likelihood majority rule consensus tree for the analyzed
Hypocreomycetidae isolates based on a dataset of combined LSU, SSU and ITS sequence data.
High branch support is shown at the nodes, maximum likelihood bootstrap support (ML BS) and
maximum parsimony (MP BS) ≥ 70% and Bayesian posterior probability (PP) ≥ 0.95. The scale bar
represents the expected number of changes per site. The tree is rooted with Canalisporium
grenadoideum. The original isolate numbers are noted after the species names. The newly
generated strains are in red, while new species are in red bold. Branches with 100% ML BS, 100%
MP BS and 1.0 PP are noted with black node. Monotypic clades are indicated as coloured blocks.
404
Figure 85 – Phaeoisaria guttulata (MFLU 18-0139, holotype) a Colonies on natural substrate. b
Colonies on PDA medium. c, d Reproduced conidiophores with conidia. e, g, h Conidiogenous
cells with conidia from culture. f Conidiogenous cells growing on natural substrate. i, j Brown
globose cells from culture. k Germinated conidium on PDA medium. m, n Culture, m from above,
n from below. Scale bars: a = 50 μm, b, f = 30 μm, c, j = 20 μm, d, e = 100 μm, g–i = 10 μm.
405
Figure 86 – Pleurotheciella krabiensis (MFLU 18-0140, holotype) a Colonies on natural substrate.
b, c Conidiophores with conidia. d, e Conidiophores. f–i Conidia. j Germinated conidium on PDA
medium. k, l Culture, k from above, l from below. Scale bars: a = 50 μm, b, f = 30 μm, c, j = 20
μm, d, e = 100 μm, g–i = 10 μm.
406
Pleurotheciella tropica J. Yang & K.D. Hyde, sp. nov.
Fig. 87
Index Fungorum number: IF554235; Facesoffungi number: FoF04088
Etymology – referring to the collecting site in tropical country, Thailand.
Holotype – MFLU 18-0141
Saprobic on submerged decaying wood. Sexual morph Undetermined. Asexual morph
Colonies effuse, scattered, brown, hairy, visible as solitary, dark brown conidiophores with bright
white mass of conidia. Mycelium partly immersed, partly superficial. Conidiophores
macronematous, mononematous, erect, septate, smooth, cylindrical, dark brown at the base,
becoming paler towards the apex, straight or slightly curved, 100–250 µm long, 4–4.8 µm wide.
Conidiogenous cells integrated, terminal, polyblastic, cylindrical, pale brown to hyaline, forming
conidia sympodially on cylindrical denticles. Conidia narrowly obovoid or subclavate, hyaline,
smooth-walled, guttulate, straight, uniseptate, 16–21 × 5.5–7 µm ( = 18.5 × 6.3 µm, n = 20),
obtuse at the apex, pointed at the base.
Culture characteristics – Conidia germinating on PDA within 24 hours. Germ tubes
produced from both ends. Colonies on PDA slow growing, reaching 7–10 mm diameter after one
month at 25 °C in natural light, circular, wrinkled, with sparse aerial white mycelium in the center,
yellowish-brown on the middle ring and sparse white mycelium at the entire margin, yellowishbrown in reverse.
Material examined – THAILAND, Phang Nga Province, Bann Tom Thong Khang, on
decaying wood submerged in a freshwater stream, 17 December 2015, J Yang, Site 7-21-4 (MFLU
18-0141, holotype); ex-type living cultures, MFLUCC 16-0867, ICMP.
GenBank numbers – ITS: MG837020, LSU: MG837015, SSU: MG837025.
Notes – Pleurotheciella tropica is phylogenetically close to Pl. krabiensis, but its mononematous
conidiophores differ from the synnematous conidiophores of Pl. krabiensis. Morphologically, Pl.
tropica resembles Pl. submersa Z.L. Luo & K.D. Hyde and Pl. uniseptata (Matsush.) Seifert, in
having brown, mononematous conidiophores, terminal conidiogenous cells and hyaline straight
conidia. However, conidiophores of Pl. tropica (100–250 µm long) are longer than those in Pl.
submersa (113–174 μm long) and Pl. uniseptata (100–150 µm long). Among them, Pl. submersa
has larger conidia (25–28 × 5.5–6.5 µm) than in Pl. tropica (16–21 × 5.5–7 µm) and Pl. uniseptata
(12.5–16.5 × 2–4 µm) (Réblová et al. 2016, Luo et al. 2017). In addition, the obovoid conidia with
a pointed base in Pl. tropica are different from the subcylindrical conidia with rounded apex and
tapering base in Pl. submersa and Pl. uniseptata.
Subclass Xylariomycetidae O.E. Erikss & Winka
Amphisphaeriales D. Hawksw. & O.E. Erikss.
Apiosporaceae K.D. Hyde et al.
The family Apiosporaceae was introduced by Hyde et al. (1998) with the type genus
Arthrinium Kunze ex Fr. (sexual morph: Apiospora Sacc.). Currently, the family comprises
Appendicospora, Arthrinium, Dictyoarthrinium, Endocalyx, Scyphospora and Spegazzinia
(Lumbsch & Huhndorf 2010, Kirk et al. 2013, Maharachchikumbura et al. 2016a, Hongsanan et al.
2017, Wijayawardene et al. 2017a, 2018). In this study, we record Arundo plinii as a new host for
Arthrinium phragmites.
Arthrinium Kunze
Arthrinium was established by Kunze & Schmidt in 1817 with A. caricicola Kunze & J.C.
Schmidt as the type species (Wijayawardene et al. 2016). Currently the genus comprises 64
accepted species, with saprobic, pathogenic or endophytic nutritional modes (Wijayawardene et al.
2017a). Crous & Groenewald (2013) provided a phylogeny for this genus and the older name
Arthrinium was conserved as it is a more commonly encountered name and more frequently used in
the literature. Recently, Dai et al. (2016a) provided an updated phylogenetic analysis with two new
Arthrinium species on bamboo from China.
407
Figure 87 – Pleurotheciella tropica (MFLU 18-0141, holotype). a Colonies on natural substrate. b,
c Conidiophores and conidiogenous cells. d Conidia. e Germinated conidium on PDA medium. f, g
Culture, f from above, g from below. Scale bars: a, c = 50 μm, b = 30 μm, d, e = 20 μm.
Arthrinium phragmites Crous [as 'phragmites'], in Crous & Groenewald, IMA Fungus 4(1): 147
(2013)
Fig. 89
Index Fungorum number: IF545031; Facesoffungi number: FoF03934
Saprobic on dead culms of Arundo plinii Turra. Sexual morph Stromata 1.8–3.2 mm long,
0.23–0.35 mm wide, 188–282 μm high, brown to black, scattered to gregarious, immersed,
becoming erumpent, splitting along its length, revealing a row of separate, subglobose ascomata.
Ascomata 170–210 μm high, 160–200 μm diameter, arranged in rows, clustered, gregarious, with
3–6 perithecia forming groups immersed in stromata, to erumpent through host surface, ellipsoidal
to subglobose, dark brown, membranous, ostiolate. Ostiole raised from centre of ascomata,
internally lined with periphyses. Peridium 6.5–12.5 μm ( = 9.6 μm, n = 15) wide, comprising 3–4
layers of dark brown or reddish-brown brown or reddish-brown to lightly pigmented cells, of
textura angularis. Hamathecium composed of dense 3–5 μm ( = 3.9 μm, n = 25) wide, hyaline,
filamentous, branched, septate paraphyses. Asci 90–130 × 15–20 μm ( = 107.5 × 16.9 μm, n =
15), 8-spored, unitunicate, clavate, with short basal pedicel, with obtusely rounded apex, lacking an
apical ring. Ascospores 20–26 × 8–10 μm ( = 24.2 × 9.2 μm, n = 25), 2–3-seriate, 1-septate,
hyaline, apiosporous, straight to curved, ellipsoid to reniform, with a gelatinous mucoid sheath.
Asexual morph Undetermined.
408
409
Figure 88 – Phylogram generated from Bayesian analysis based on ITS sequence data. Related
sequences were obtained from GenBank following Dai et al. (2016a). A total of 56 strains are
included in the analyses. Seiridium phylicae (Sporocadaceae) is used as the outgroup taxon. Tree
topology of the Bayesian analysis was similar to the maximum-likelihood analysis. The best
scoring RAxML tree with a final likelihood value of -3843.7723 is presented. The matrix had 315
distinct alignment patterns, with 17.11% of undetermined characters or gaps. Estimated base
frequencies were as follows; A = 0.230383, C = 0.251582, G = 0.231302, T = 0.286733;
substitution rates AC = 1.068770, AG = 2.218132, AT = 1.683025, CG = 0.944962, CT =
3.903708, GT = 1.000000; gamma distribution shape parameter α = 0.396682. Bootstrap values for
ML (≥60) and Bayesian posterior probabilities (≥0.95) are given at the nodes. The newly generated
sequence is in blue. The ex-type strains are in bold. The scale bar represents the expected number
of nucleotide substitutions per site.
Culture characteristics – Colonies cottony, flat, spreading, with moderate aerial mycelium.
On PDA surface dirty white, zonate, with patches of pale luteous, reverse luteous.
Material examined – ITALY, Province of Forlì-Cesena [FC], near Meldola, on dead aerial
stem of Arundo plinii Turra (Poaceae), 16 January 2017, E. Camporesi, IT3211 (MFLU 17-0310);
living culture MFLUCC 18-0099.
GenBank numbers – ITS: MG836697, LSU: MG836698.
Notes – Arthrinium phragmites Crous was introduced from Italy on culms of Phragmites
australis (Crous & Groenewald 2013). The sexual morph and the culture characteristics of the
taxon from a dead culm of Arundo plinii are similar to Arthrinium phragmites. Crous &
Groenewald (2013) described some ascospores with remnants of mucoid sheath. The ascospores
observed in this study clearly have a mucoid sheath seen following Indian ink treatment, while the
asci and ascospore lengths and widths are slightly different (asci CBS H-21267: 70–110 × 17–25
μm, MFLU 17-0310: 90–130 × 15–20 μm; ascospores CBS H-21267: 22–25 × 7–9 μm, MFLU 170310: 20–26 × 8–10 μm). The ITS based phylogenetic analyses (100% ITS sequence similarity)
also revealed a high support (100%/0.99PP) for this clade and this is a new host record as the
Arthrinium phragmites on Arundo plinii from Italy (Fig. 88).
Sporocadaceae Corda
Jaklitsch et al. (2016a, b) treated Bartaliniaceae, Discosiaceae, Pestalotiopsidaceae and
Robillardaceae as synonyms of Sporocadaceae and this is followed in Wijayawardene et al. (2018).
This family comprises the genera Annellolacinia, Bartalinia, Broomella, Ciliochorella, Discosia,
Doliomyces, Dyrithiopsis, Hyalotiella, Immersidiscosia, Monochaetia, Morinia, Monochaetinula,
Neopestalotiopsis, Neotruncatella, Pestalotiopsis, Pseudopestalotiopsis, Robillarda, Sarcostroma,
Seimatosporium, Seiridium, Strickeria, Truncatella and Zetiasplozna (Wijayawardene et al. 2018).
In the present paper, we introduce a new species of genus Discosia and provide an updated tree for
the genus.
Discosia Lib.
Libert (1837) introduced Discosia with D. strobilina as the type species. Wijayawardene et
al. (2017a) treated Adisciso as a synonym of Discosia agreeing with the ‘one fungus one name’
concept (Hawksworth 2012). Senanayake et al. (2015) showed that Discosia sensu stricto groups
with Adisciso, Discostroma, Sarcostroma and Seimatosporium as a distinct clade in
Amphisphaeriales, and Senanayake et al. (2015) introduced Discosiaceae to accommodate these
genera, which is now considered a synonym of Sporocadaceae.
Discosia querci Jayasiri, E.B.G. Jones & K.D. Hyde, sp. nov.
Index fungorum number: IF554075; Facesoffungi number: FoF03925
Holotype: MFLU 17-2848
Fig. 91
410
Figure 89 – Arthrinium phragmites (MFLU 17-0310). a–c Pseudostromata on the host surface. d
Vertical section through pseudostroma. e Ostiole with periphyses. f Peridium. g, h Asci. i–n
Ascospores. o Sheath surrounding ascospores after treatment in India ink. p Germinating ascospore.
q Upper view of the colony. r Reverse view of the colony. Scale bars: a = 1000 µm, b, c = 500 µm,
d = 50µm, e = 20 µm, f–h, o, p = 10 µm, i–n = 5 µm.
Saprobic on dead seeds of Quercus sp. fruits. Sexual morph Undetermined. Asexual morph
coelomycetous. Conidiomata 110–170 μm diameter, 50–80 μm high, ( = 150 × 70 μm) pycnidial,
stromatic, solitary, scattered to gregarious or confluent, partly immersed or superficial, rounded in
outline, glabrous, unilocular, textura porrecta, ostiole absence. Peridium 10–15 μm thick,
composed of 5–6 layers of cells, with outer 5–6 layers dark brown, inner 1–2 layers colourless,
comprising thin-walled cells of textura angularis. Conidiophores reduced to conidiogenous cells,
arising from the upper cells of the basal stroma. Conidiogenous cells 4–8 × 1–3.5 ( = 6 × 2) μm,
holoblastic, smooth, restricted to the basal conidiomata wall, each producing a single conidium,
integrated, hyaline, smooth. Conidia 13–20 × 2.5–3 μm ( = 17 × 2.8 μm, n = 50), hyaline,
fusiform to cylindrical, thick-walled, smooth, straight or slightly curved, 3-septate, slightly
constricted at one septum at apex, with cells of equal width, basal cell obconic, with a truncate
base, apical cell subconical with a rounded apex, unbranched, filiform, flexuous or straight
appendage; presence of both ends.
411
Figure 90 – Simplified phylogram showing the best RAxML maximum likelihood tree obtained
from the combined LSU and ITS matrix of species from the genus Discosia. Related sequences
were obtained from GenBank (http://www.ncbi.nlm.nih.gov/). Twenty-eight strains are included in
the combined sequence analyses, which comprise 1824 characters with gaps. The best scoring
RAxML tree with a final likelihood value of -4959.075067 is presented. The matrix had 262
distinct alignment patterns, with 13.81% of undetermined characters or gaps. Estimated base
frequencies were as follows; A = 0.247425, C = 0.233400, G = 0.257560, T = 0.261614. MLBS
above 70% are given at each branch. The tree is rooted with Pestalotiopsis uvicola (UCD2465TX).
New isolates are in bold and green.
Material examined – UK, Hampshire, Swanick Lakes, on dead seed of Quercus sp.
(Fagaceae) fruits, 28 September 2015, E.B. Gareth Jones, GJ 195 (MFLU 17-2848, holotype), extype living culture, MFUCC 16-0642, BCC; ibid. (PDD, isotype).
GenBank numbers – ITS: MG815829, LSU: MG815830, SSU: MG815831.
Notes – Tanaka et al. (2011) introduced four strains of Discosia aff. brasiliensis (KT2193,
KT2190, KT2194 and MAFF237018) from Japan based on morphology and phylogenetic data. Our
isolate, clusters with Discosia aff. brasiliensis (KT2190) with high statistical support (81 ML), but
other strains of Discosia brasiliensis cluster separately from our isolate and KT 2190. Therefore, in
this study, we introduce our strain as a new species of Discosia. Discosia querci shares similar
conidiomata, conidiogenous cells and conidial morphology with Discosia brasiliensis (Nag Raj
1993, Tangthirasunun et al. 2015). However, Discosia querci differs from Discosia brasiliensis in
having wider conidiomata (110–170 μm vs. 363–390 μm) and unilocular conidiomtaa. The type
412
species of Discosia brasiliensis was identified from a Bignoniaceae host. However, Discosia aff.
brasiliensis (KT2193, KT2190, KT2194 and MAFF237018) and Discosia brasiliensis (MFLUCC
12-0429) were isolated from decayed leaves of unknown hosts. Discosia querci was collected from
seeds of Quercus sp. from the UK and in the USDA fungal database, Discosia sp. are listed from
different Quercus sp. in different localities. Discosia artocreas has been reported from Quercus sp.
in the UK, but in multigene phylogenetic analysis our isolate is in a distinct clade to Discosia
artocreas. We were unable to loan the type of Discosia quercicola De Not. and even if we could it
is unlikely we could obtain sequence data, and thus would be unable to use this name (Liu et al.
2016).
Figure 91 – Discosia querci (MFUCC 16-0642, holotype). a Seed samples. b, c Black conidiomata
on the host surface. d Vertical section of conidioma. e, f, h Conidiogenous cells and developing
conidia. g Section of peridium. i, j Conidia. Scale bars: a = 1 cm, b = 500 μm, c = 200 μm, d = 50
μm, e, f = 30 μm, g–m = 10 μm.
Seimatosporium Corda
The genus was introduced by Corda (1833), the type species is S. rosae Corda. Phylogenetic
studies indicated that Seimatosporium should be placed in Discosiaceae, Amphisphaeriales
(Senanayake et al. 2015, Norphanphoun et al. 2015, Wijayawardene et al. 2016). However,
Discosiaceae was synonymized under Sporocadaceae with other families Bartaliniaceae,
Pestalotiopsidaceae and Robillardaceae based on morphology and phylogeny (Jaklitsch et al. 2016).
The asexual/sexual morph connection between Discostroma and Seimatosporium was confirmed by
using a molecular approach (Jeewon et al. 2002, Norphanphoun et al. 2015). Rossman et al. (2016)
concluded that the name Seimatosporium should be used as it is the oldest name and is most
commonly used.
413
Seimatosporium ciliata (Petr.) Hongsanan & K.D. Hyde, comb. nov.
Fig. 92
Index Fungorum number: IF554378; Facesoffungi number: FoF04465
≡ Neobroomella ciliata Petr., Sydowia 1(1-3): 5 (1947)
Epiphyte on surface of dead stems of Phlomis brevilabris Ehrenb. ex Boiss. Sexual morph
Ascomata 197−220 µm high × 190−240 µm diameter ( = 210 × 200 µm, n = 5), scattered, semiimmersed in host tissue, subglobose to globose or ellipsoid, membranaceous, dark brown to black,
with long papilla at the center. Neck 130 µm high × 95 µm diameter, papillate, composed of thin,
flattened cells of textura angularis, hyaline to pale brown. Peridium 10−15 µm ( = 13 µm, n =
20), two-layered, comprising outer, dark brown to black, cells of textura angularis and inner, thin,
hyaline, flattened cells of textura angularis. Hamathecium of 1 µm, pseudoparenchyma, aseptate,
embedded in a gelatinous matrix. Asci 85−97 × 12−15 µm ( = 95 × 14 µm, n = 20), numerous, 8spored, unitunicate, cylindrical to oblong, with a short pedicel. Ascospores 20−24 × 7−9 µm ( =
22 × 8 µm, n = 30), biseriate in ascus, 2-layered, 3-septate, oval to ellipsoid, generally straight,
rounded at both ends, not constricted at the septum, hyaline, verrucose surface, embedded in
gelatinous matrix. Asexual morph Undetermined.
Material examined – SYRIA, Lebanon, Jugi Sanin, in subalpine region, 1700-1900 m., on
dead stems of Phlomis brevilabris (Lamiaceae), 21 July 1897, J. Bornmüller, (W no. 11029,
holotype).
Notes – Neobroomella was introduced in Sphaeriales by Petrak (1947) with N. ciliata Petr.
as the type species. The genus was accepted as Xylariomycetidae genera incertae sedis (Lumbsch
& Huhndorf 2010, Kirk et al. 2013, Wijayawardene et al. 2017a). Senanayake et al. (2015) and
Maharachchikumbura et al. (2015, 2016a) accepted Neobroomella in Phlogicylindriaceae,
Amphisphaeriales. Sequence data is unavailable. Eriksson (1980) re-studied the herbarium
specimen from W (no. 11029) and noted that Neobroomella ciliata was in good condition and we
made same observation. Neobroomella ciliata is identical to species in Discostroma (current name
Seimatosporium) in having semi-immersed ascomata, with a central papilla, unitunicate asci and
uniseriate, ellipsoid, transversely septate, hyaline ascospores (Paulus et al. 2006). However, the
subapical ring in the ascus apex was not J+ (Fig. 92). Culture and molecular data for Neobroomella
are unavailable. Therefore, we synonymize N. ciliata under the genus Seimatosporium based on
morphological characters (Fig. 92).
Truncatella Steyaert
The genus Truncatella was established by Steyaert (1949) to accommodate T. truncata which
had distinct 3-septate, verruculose, pigmented conidia (Steyaert 1949, Maharachchikumbura et al.
2015). This genus was previously placed in Amphisphaeriaceae (Jeewon et al. 2002, 2003) and
Bartaliniaceae (Senanayake et al. 2015). Jaklitsch et al. (2016a) placed this genus in the family
Sporocadaceae. There are 22 species epithets in Index Fungorum (2018), however, there are only
eleven estimated species (Wijayawardene et al. 2017a). Species of this genus are saprobes and have
a wide terrestrial distribution (Wijayawardene et al. 2017a).
Truncatella angustata (Pers.) S. Hughes, Can. J. Bot. 36: 822 (1958)
Fig. 94
≡ Stilbospora angustata Pers., Syn. meth. fung. (Göttingen) 1: 96 (1801)
Index Fungorum number: IF307155; Facesoffungi number: FoF04092
Saprobic on dead aerial branch of Alnus glutinosa (L.) Gaertn. Sexual morph not observed
(after 5 weeks). Asexual morph Conidiomata 0.2–0.35 mm diameter ( = 0.24 mm, n = 10),
pycnidial, semi-immersed to superficial, becoming erumpent at maturity, globose with a flattened
base, solitary to aggregated, black. Conidiomata wall consisting of outer light brown to hyaline
cells of textura angularis to textura globulosa. Conidiophores 23–26 × 2–4.5 µm ( = 23× 2 µm, n
= 10), hyaline, cylindrical, simple or branched. Conidiogenous cells 6–9 µm long, hyaline, simple,
cylindrical, integrated, smooth. Conidia 18–20 × 7–8 µm ( = 19 × 6 µm, n = 40), fusiform,
straight, with 3 transverse septa, with a truncate base, age absent.
414
Figure 92 – Seimatosporium ciliata (W no. 11029, holotype). a, b Habit and ascoma on substrate.
c, d, f Section of ascoma. e Peridium. g, h Pseudoparaphyses. i Ascus. j Ascus in Melzer’s reagent.
k, l Asci in cotton blue reagent. m Apex of ascus. n, p Ascospores. o Ascospores in Melzer’s
reagent. q Ascospore in cotton blue reagent. Scale bars: d, f = 50 μm, e, g, h, n−q = 10 μm, i–l = 20
μm, m = 5 μm.
415
Figure 93 – Phylogenetic tree generated from maximum parsimony analysis based on ITS
sequence data of Truncatella species. Related sequences were obtained from GenBank. Eleven
strains are included in the analyses, which comprise 542 characters including gaps. Tree topology
of the MP analysis was similar to the ML and BI. The maximum parsimonious dataset consisted of
constant 456, 27 parsimony-informative and 59 parsimony-uninformative characters. The
parsimony analysis of the data matrix resulted in the maximum of three equally most parsimonious
trees with a length of 102 steps (CI = 0.922, RI = 0.896, RC = 0.826, HI = 0.078) in the first tree.
The best scoring RAxML tree with a final likelihood value of -1262.909320 is presented. The
matrix had 76 distinct alignment patterns, with 5.32% of undetermined characters or gaps.
Estimated base frequencies were as follows; A = 0.261470, C = 0.234898, G= 0.200531, T =
0.303100; substitution rates AC = 1.928476, AG = 6.544930, AT = 1.879121, CG = 1.234597, CT
= 6.560020, GT = 1.000000; gamma distribution shape parameter α = 0.188065. Maximum
parsimony bootstrap, maximum likelihood support values higher than 70%, shown respectively
near the nodes. Bayesian posterior probabilities ≥ 0.95 (PP) indicated as thickened black branches.
The scale bar indicates 0.04 changes. The ex-type strains are in bold and new isolates in blue. The
new isolate is shown in blue bold and ex-type strains in black bold. The strain introduced in this
study is in blue. The tree is rooted with Pseudopestalotiopsis theae (MFLUCC 12-0055).
Culture characteristics – On PDA, fast growing attaining 40 mm diameter after 7 days at 25
°C, serrate margin, white from above, reverse greyish-white. Black pycnidia were observed in the
center after 7 days.
Material examined – ITALY, Province of Forli-Cesena [FC], San Benedetto in Alpe, on dead
and aerial branch of Alnus glutinosa (Betulaceae), 18 April 2016, Erio Camporesi, IT2935 (MFLU
16-1271); living culture MFLUCC 17-1283.
GenBank number – ITS: MG835271.
Material examined – ITALY, Province of Forli-Cesena [FC], San Benedetto in Alpe, on dead
and aerial branch of Alnus glutinosa (Betulaceae), 18 April 2016, Erio Camporesi, IT2935 (MFLU
16-1271); living culture MFLUCC 17-1283.
GenBank number – ITS: MG835271.
Notes – Phylogeny inferred using sequence data of ITS regions from the isolates obtained in
this study with other known isolates of T. angustata and other Truncatella species in Tibpromma et
al. (2017) clustered our isolates with T. angustata (100% bootstrap support value and 1.00
416
Bayesian posterior probability) (Fig. 93). The morphology of our isolates arein full agreement with
the description for T. angustata (Sutton 1980). Truncatella angustata has a wide distribution and
this is the first report of this species from Alnus glutinosa from Italy.
Figure 94 – Morphology of Truncatella angustata. a Conidiomata on host tissue. b–d
Conidiophores and conidiogenous cells. e Conidia. f Immature conidia. g Branched apical
appendages. h Above view of the culture after 7 days. i Reverse view of the culture after 7 days.
Scale bars: b–d = 20 µm, e–g = 10 µm.
Acknowledgements
Kevin D. Hyde would like to thank 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
417
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 “Biodiversity, phylogeny and role of fungal endophytes of Pandanaceae” (grant number:
592010200112), ‘‘Diseases of mangrove trees and maintenance of good forestry practice’’ (grant
number: 60201000201) and Taxonomy diversity, phylogeny and evolution of fungi in Capnodiales
(grant no: 666713), for supporting this study. S. Boonmee would like to thank the National
Research Council of Thailand (projects No. 61215320013 and No. 61215320023). This work was
jointly supported by the National Natural Science Foundation of China (No. 31460012 &
31760014), the Science and Technology Foundation of Guizhou Province (No. [2016]2863). Shaun
Pennycook is thanked for checking and suggesting corrections to the Latin names. R. Phookamsak
expresses appreciation to The CAS President’s International Fellowship for Postdoctoral
Researchers, project number 2017PB0072, the Research Fund from China Postdoctoral Science
Foundation (grant no. Y71B283261) and Chiang Mai University for financial support. We
appreciate the staff at Ranong Mangrove Forest Research Center for giving us knowledge and
granting us permission to study and collect specimens. Shi-Ke Huang is particularly grateful to Dr.
Qi Zhao for his invaluable suggestions. The Department of Plant Medicine, National Chiayi
University (NCYU), Taiwan is thanked for providing facilities for DNA molecular sequencing.
Saowaluck Tibpromma thanks the Mushroom Research Foundation (MRF), Chiang Rai, Thailand
for financial support. MC Dayarathne thanks the Thailand Research Fund grant no. MRG6080089.
Rajesh Jeewon thanks Mae Fah Luang University and University of Mauritius for research support.
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