Fungal Diversity (2016) 80:1–270
DOI 10.1007/s13225-016-0373-x
Fungal diversity notes 367–490: taxonomic and phylogenetic
contributions to fungal taxa
Kevin D. Hyde1,2,3 • Sinang Hongsanan3,32 • Rajesh Jeewon55 • D. Jayarama Bhat12,13 •
Eric H. C. McKenzie20 • E. B. Gareth Jones5 • Rungtiwa Phookamsak15 •
Hiran A. Ariyawansa9 • Saranyaphat Boonmee3 • Qi Zhao1,2,28 • Faten Awad Abdel-Aziz6 •
Mohamed A. Abdel-Wahab5,6 • Supharat Banmai3 • Putarak Chomnunti10 •
Bao-Kai Cui38 • Dinushani A. Daranagama3,4 • Kanad Das26 • Monika C. Dayarathne2,3 •
Nimali I. de Silva2,3,14,15 • Asha J. Dissanayake3,30 • Mingkwan Doilom3 •
Anusha H. Ekanayaka2,3,15 • Tatiana Baptista Gibertoni56 • Aristóteles Góes-Neto50 •
Shi-Ke Huang2,3 • Subashini C. Jayasiri3,29 • Ruvishika S. Jayawardena3,30 •
Sirinapa Konta3 • Hyang Burm Lee39 • Wen-Jing Li3 • Chuan-Gen Lin3,11 •
Jian-Kui Liu3,9 • Yong-Zhong Lu3,29 • Zong-Long Luo3,10,31 • Ishara S. Manawasinghe3,30 •
Patinjareveettil Manimohan58 • Ausana Mapook3 • Tuula Niskanen49 •
Chada Norphanphoun3 • Moslem Papizadeh51 • Rekhani H. Perera3,9 •
Chayanard Phukhamsakda2,3 • Christian Richter17,18 • André L. C. M. de A. Santiago40 •
E. Ricardo Drechsler-Santos42 • Indunil C. Senanayake2,3,15 • Kazuaki Tanaka7,8 •
Received: 27 July 2016 / Accepted: 30 August 2016 / Published online: 28 September 2016
Ó School of Science 2016
Electronic supplementary material The online version of this
article (doi:10.1007/s13225-016-0373-x) contains supplementary
material, which is available to authorized users.
& Qi Zhao
zhaoqi@mail.kib.ac.cn
Kevin D. Hyde
kdhyde3@gmail.com
1
2
3
4
Institute of Biotechnology and Germplasm Resources;
Key Lab of Southwestern Crop Gene Resources
and Germplasm Innovation Ministry of Agriculture;
Yunnan Provincial Key Laboratory of Agricultural
Biotechnology, Yunnan Academy of Agricultural Sciences,
Kunming 650223, Yunnan,
People’s Republic of China
Key Laboratory of Plant Diversity and Biogeography of East
Asia, Kunming Institute of Botany, Chinese Academy of
Sciences, Kunming 650201, Yunnan,
People’s Republic of China
Center of Excellence in Fungal Research,
Mae Fah Luang University, 333 M. 1, T. Thasud,
Muang, Chiang Rai 57100, Thailand
State Key Laboratory of Mycology, Institute of
Microbiology, Chinese Academy of Sciences,
No 3 1st West Beichen Road, Chaoyang District,
Beijing 100101, People’s Republic of China
5
Department of Botany and Microbiology, College of Science,
King Saud University, P.O. Box: 2455, Riyadh, 1145,
Saudi Arabia
6
Department of Botany and Microbiology, Faculty of Science,
Sohag University, Sohag 82524, Egypt
7
Faculty of Agriculture and Life Sciences, Hirosaki
University, 3 Bunkyo-cho, Hirosaki, Aomori 036-8561, Japan
8
The United Graduate School of Agricultural Sciences, Iwate
University, 18-8 Ueda 3 Chome, Morioka 020-8550, Japan
9
Guizhou
Guizhou
Guiyang
People’s
10
School of Science, Mae Fah Luang University,
Chiang Rai 57100, Thailand
11
Department of Plant Pathology, College of Agriculture,
Guizhou University, Guiyang 550025, Guizhou,
People’s Republic of China
12
Department of Botany, Goa University, Taleigão, Goa, India
13
No. 128/1–J, Azad Housing Society, Curca,
Goa Velha 403108, India
Key Laboratory of Agricultural Biotechnology,
Academy of Agricultural Sciences, Xiaohe District,
City, Guizhou Province 550006,
Republic of China
123
2
Fungal Diversity (2016) 80:1–270
T. M. D. S. Tennakoon2,3,15 • Kasun M. Thambugala3,9 • Qing Tian2,3,10,15,16 •
Saowaluck Tibpromma3,15,32 • Benjarong Thongbai3 • Alfredo Vizzini35 •
Dhanushka N. Wanasinghe2,3 • Nalin N. Wijayawardene3 • Hai-Xia Wu43 •
Jing Yang3 • Xiang-Yu Zeng3,29 • Huang Zhang33 • Jin-Feng Zhang9 •
Timur S. Bulgakov19 • Erio Camporesi21,22,23 •
Ali H. Bahkali5 • Mohammad A. Amoozegar54 • Lidia Silva Araujo-Neta56 •
Joseph F. Ammirati48 • Abhishek Baghela27 • R. P. Bhatt24 • Dimitar Bojantchev47 •
Bart Buyck25 • Gladstone Alves da Silva56 • Catarina Letı́cia Ferreira de Lima40 •
Rafael José Vilela de Oliveira40 • Carlos Alberto Fragoso de Souza40 •
Yu-Cheng Dai38 • Bálint Dima45,46 • Tham Thi Duong39 • Enrico Ercole35 •
Fernando Mafalda-Freire42 • Aniket Ghosh24 • Akira Hashimoto7,8 •
Sutakorn Kamolhan3 • Ji-Chuan Kang29 • Samantha C. Karunarathna2,3,15,16,32 •
Paul M. Kirk57 • Ilkka Kytövuori44 • Angela Lantieri36 • Kare Liimatainen45,49 •
Zuo-Yi Liu9 • Xing-Zhong Liu4 • Robert Lücking34 • Gianfranco Medardi37 •
Peter E. Mortimer2,15,16 • Thi Thuong Thuong Nguyen39 •
Itthayakorn Promputtha14 • K. N. Anil Raj58 • Mateus A. Reck41 • Saisamorn Lumyong14 •
Seyed Abolhassan Shahzadeh-Fazeli51,53 • Marc Stadler17,18 • Mohammad Reza Soudi52 •
Hong-Yan Su31 • Takumasa Takahashi7 • Narumon Tangthirasunun3,59 •
Priyanka Uniyal24 • Yong Wang11 • Ting-Chi Wen29 • Jian-Chu Xu2,15,16 •
Zhong-Kai Zhang1 • Yong-Chang Zhao1 • Jun-Liang Zhou38 • Lin Zhu38
Abstract This is a continuity of a series of taxonomic
papers where materials are examined, described and novel
combinations are proposed where necessary to improve our
traditional species concepts and provide updates on their
classification. In addition to extensive morphological
descriptions and appropriate asexual and sexual
14
Department of Biology, Faculty of Science, Chiang Mai
University, Chiang Mai 50200, Thailand
25
15
World Agroforestry Centre, East and Central Asia,
Kunming 650201, Yunnan, People’s Republic of China
Muséum National d’Histoire Naturelle, Département
Systématique et Evolution, CP 39, ISYEB, UMR 7205 CNRS
MNHN UPMC EPHE, 12 Rue Buffon, 75005 Paris, France
26
Cryptogamic Unit, Botanical Survey of India, P.O. Botanic
Garden, Howrah 711103, India
27
National Fungal Culture Collection of India (NFCCI),
Biodiversity and Palaeobiology Group, MACS - Agharkar
Research Institute, G.G. Agarkar Road, Pune 411004, India
28
Institute of Edible Fungi, Yunnan Agricultural University,
Kunming 650201, Yunnan, People’s Republic of China
29
Engineering Research Center of Southwest BioPharmaceutical Resources, Ministry of Education, Guizhou
University, Guiyang 550025, Guizhou Province,
People’s Republic of China
30
Institute of Plant and Environment Protection, Beijing
Academy of Agriculture and Forestry Sciences,
Beijing 100097, People’s Republic of China
31
College of Agriculture & Biological Sciences, Dali
University, Dali 671003, Yunnan,
People’s Republic of China
32
Mushroom Research Foundation, 128 M.3 Ban Pa Deng T.
Pa Pae, A. Mae Taeng, Chiang Mai 50150, Thailand
33
Yunnan Institute of Food Safety, Kunming University of
Science & Technology, Kunming 650500, Yunnan,
People’s Republic of China
16
17
18
Centre of Mountain Ecosystem Studies, Kunming Institute of
Botany, Chinese Academy of Sciences,
Kunming 650201, Yunnan, People’s Republic of China
Department of Microbial Drugs, Helmholtz Centre for
Infection Research, Brunswick, Germany
German Centre for Infection Research (DZIF), Partner Site
Hannover/Braunschweig, Inhoffenstrasse 7,
38124 Brunswick, Germany
19
Academy of Biology and Biotechnology, Southern Federal
University, Rostov-on-Don 344090, Rostov Region, Russia
20
Landcare Research Manaaki Whenua,
Private Bag 92170, Auckland, New Zealand
21
Gruppo Micologico Forlivese ‘‘Antonio Cicognani’’, Via
Roma 18, Forlı̀, Italy
22
Circolo Micologico ‘‘Giovanni Carini’’, C.P. 314, Brescia,
Italy
23
Società per gli Studi Naturalistici della Romagna,
C.P. 144, Bagnacavallo, RA, Italy
24
Department of Botany & Microbiology, H.N.B. Garhwal
University, Srinagar, Garhwal, Uttarakhand 246174, India
123
Fungal Diversity (2016) 80:1–270
3
connections, DNA sequence data are also analysed from
concatenated datasets (rDNA, TEF-a, RBP2 and b-Tubulin) to infer phylogenetic relationships and substantiate
systematic position of taxa within appropriate ranks.
Wherever new species or combinations are being proposed,
we apply an integrative approach (morphological and
molecular data as well as ecological features wherever
applicable). Notes on 125 fungal taxa are compiled in this
paper, including eight new genera, 101 new species, two
new combinations, one neotype, four reference specimens,
new host or distribution records for eight species and one
alternative morphs. The new genera introduced in this
paper are Alloarthopyrenia, Arundellina, Camarosporioides, Neomassaria, Neomassarina, Neotruncatella, Paracapsulospora and Pseudophaeosphaeria. The new species
are Alfaria spartii, Alloarthopyrenia italica, Anthostomella
ravenna, An. thailandica, Arthrinium paraphaeospermum,
Arundellina typhae, Aspergillus koreanus, Asterina
cynometrae, Bertiella ellipsoidea, Blastophorum aquaticum, Cainia globosa, Camarosporioides phragmitis,
Ceramothyrium menglunense, Chaetosphaeronema achilleae, Chlamydotubeufia helicospora, Ciliochorella
phanericola, Clavulinopsis aurantiaca, Colletotrichum
insertae, Comoclathris italica, Coronophora myricoides,
Cortinarius fulvescentoideus, Co. nymphatus, Co. pseudobulliardioides, Co. tenuifulvescens, Cunninghamella
gigacellularis, Cyathus pyristriatus, Cytospora cotini,
Dematiopleospora alliariae, De. cirsii, Diaporthe aseana,
Di. garethjonesii, Distoseptispora multiseptata, Dis. tectonae, Dis. tectonigena, Dothiora buxi, Emericellopsis
persica, Gloniopsis calami, Helicoma guttulatum, Helvella
floriforma, H. oblongispora, Hermatomyces subiculosa,
Juncaceicola italica, Lactarius dirkii, Lentithecium unicellulare, Le. voraginesporum, Leptosphaeria cirsii, Leptosphaeria irregularis, Leptospora galii, Le. thailandica,
Lindgomyces pseudomadisonensis, Lophiotrema bambusae, Lo. fallopiae, Meliola citri-maximae, Minimelanolocus submersus, Montagnula cirsii, Mortierella
fluviae, Muriphaeosphaeria ambrosiae, Neodidymelliopsis
ranunculi, Neomassaria fabacearum, Neomassarina thailandica, Neomicrosphaeropsis cytisi, Neo. cytisinus, Neo.
minima, Neopestalotiopsis cocoës, Neopestalotiopsis
musae, Neoroussoella lenispora, Neotorula submersa,
Neotruncatella endophytica, Nodulosphaeria italica, Occultibambusa aquatica, Oc. chiangraiensis, Ophiocordyceps hemisphaerica, Op. lacrimoidis, Paracapsulospora
metroxyli, Pestalotiopsis sequoiae, Peziza fruticosa, Pleurotrema thailandica, Poaceicola arundinis, Polyporus
mangshanensis, Pseudocoleophoma typhicola, Pseudodictyosporium thailandica, Pseudophaeosphaeria rubi, Purpureocillium sodanum, Ramariopsis atlantica, Rhodocybe
griseoaurantia, Rh. indica, Rh. luteobrunnea, Russula
indoalba, Ru. pseudoamoenicolor, Sporidesmium aquaticivaginatum, Sp. olivaceoconidium, Sp. pyriformatum,
34
Botanic Garden and Botanical Museum, Königin-LuiseStrasse 6-8, 14195 Berlin, Germany
43
35
Department of Life Sciences and Systems Biology,
University of Torino, Viale P.A. Mattioli 25, 10125 Turin,
Italy
International Fungal Research and Development Centre, Key
Laboratory of Resource Insect Cultivation & Utilization State
Forestry Administration, The Research Institute of Resource
Insects, Chinese Academy of Forestry, Kunming 650224,
People’s Republic of China
36
Via Novaluce 38, 95125 Tremestieri Etneo, Catania, Italy
44
37
Via Giuseppe Mazzini 21, 25086 Rezzato, Brescia, Italy
Botanical Museum, University of Helsinki, P.O. Box 7,
00014 Helsinki, Finland
45
38
Institute of Microbiology, Beijing Forestry University,
Beijing 100083, People’s Republic of China
Department of Biosciences, Plant Biology, University of
Helsinki, P.O. Box 65, 00014 Helsinki, Finland
46
39
Division of Food Technology, Biotechnology &
Agrochemistry, College of Agriculture and Life Sciences,
Chonnam National University, Gwangju 61186,
Republic of Korea
Department of Plant Anatomy, Institute of Biology, Eötvös
Loránd University, Pázmány Péter sétány 1/c,
1117 Budapest, Hungary
47
MushroomHobby.com, 345 Shipwatch Lane, Hercules,
CA 94547, USA
48
Department of Biology, University of Washington,
Box 351800, Seattle, WA 98195-1800, USA
49
Jodrell Laboratory, Royal Botanic Gardens, Kew TW9 3AB,
UK
50
Laboratório de Biologia Molecular e Computacional de
Fungos (LBMCF), Departamento de Microbiologia,
Universidade Federal de Minas Gerais, Av. Antônio Carlos,
6627, Pampulha, Belo Horizonte, MG CEP: 31270-901,
Brazil
51
Microorganisms Bank, Iranian Biological Resource Center
(IBRC), Academic Center for Education, Culture & Research
(ACECR), Tehran, Iran
40
41
42
Programa de Pós-Graduação em Biologia de Fungos,
Departamento de Micologia, Universidade Federal de
Pernambuco, Av. Prof. Moraes Rego, s/n, Recife,
PE 50670-420, Brazil
Departamento de Biologia Campus Universitário,
Universidade Estadual de Maringá, Av. Colombo
5790, Jardim Universitario, Maringá,
PR CEP 87020-900, Brazil
Micolab, Programa de Pós-Graduação em Biologia de
Fungos, Algas e Plantas, Departamento de Botânica,
Universidade Federal de Santa Catarina, Campus
Universitário Trindade, Florianópolis, SC CEP: 88040-900,
Brazil
123
4
Fungal Diversity (2016) 80:1–270
Stagonospora forlicesenensis, Stagonosporopsis centaureae, Terriera thailandica, Tremateia arundicola, Tr.
guiyangensis, Trichomerium bambusae, Tubeufia hyalospora, Tu. roseohelicospora and Wojnowicia italica. New
combinations are given for Hermatomyces mirum and
Pallidocercospora thailandica. A neotype is proposed for
Cortinarius fulvescens. Reference specimens are given for
Aquaphila albicans, Leptospora rubella, Platychora ulmi
and Meliola pseudosasae, while new host or distribution
records are provided for Diaporthe eres, Di. siamensis, Di.
foeniculina, Dothiorella iranica, Do. sarmentorum, Do.
vidmadera, Helvella tinta and Vaginatispora fuckelii, with
full taxonomic details. An asexual state is also reported for
the first time in Neoacanthostigma septoconstrictum. This
paper contributes to a more comprehensive update and
improved identification of many ascomycetes and
basiodiomycetes.
367. Asterina cynometrae Hongsanan & K.D. Hyde, in
Fungal Diversity 81: 12 (2016), new species
Keywords Ascomycota Basidiomycota
Dothideomycetes Mortierellomycotina
Mucoromycotina Pezizomycetes Phylogeny
Taxonomy New genus New species New host records
371. Pallidocercospora thailandica (Crous et al.)
Phookamsak, Wulandari & K.D. Hyde, in Fungal
Diversity 81: 21 (2016), new combination
Table of contents
Ascomycota
Dothideomycetes
Asterinales M.E. Barr ex D. Hawksw. & O.E. Erikss.
Asterinaceae Hansf.
52
National Laboratory of Industrial Microbiology, Department
of Microbiology, Faculty of Biological Sciences, Alzahra
University, Tehran, Iran
53
Department of Molecular and Cellular Biology, Faculty of
Basic Sciences and Advanced Technologies in Biology,
University of Science and Culture, Tehran, Iran
54
Extremophiles Laboratory, Department of Microbiology,
Faculty of Biology and Center of Excellence in Phylogeny of
Living Organisms, College of Science, University of Tehran,
Tehran, Iran
55
Department of Health Sciences, Faculty of Science,
University of Mauritius, Reduit, Mauritius
56
Departamento de Micologia Avenida da Engenharia, Centro
de Ciências Biológicas (CCB), Universidade Federal de
Pernambuco (UFPE), S/N - Cidade Universitária, Recife,
PE 50740-600, Brazil
57
Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3DS,
UK
58
Department of Botany, University of Calicut, Malappuram,
Kerala 673 635, India
59
Laboratoire Interdisciplinaire des Energies de Demain,
Université Paris Diderot, 35, Rue Hélène Brion, 75205 Paris,
France
123
Botryosphaeriales C.L. Schoch et al.
Botryosphaeriaceae Theiss. & Syd.
368. Dothiorella iranica Abdollahz. et al., in Abdollahzadeh et al., Persoonia, Mol. Phyl. Evol. Fungi 32:
4 (2014), new host record, p 15
369. Dothiorella sarmentorum (Fr.) A.J.L. Phillips et al.,
Mycologia 97: 522. 2005, new host record, p 17
370. Dothiorella vidmadera W.M. Pitt et al., Fungal
Diversity 61 (1): 216 (2013), new host record, p 17
Capnodiales Woron.
Mycosphaerellaceae Lindau
Dothideales Lindau
Dothideaceae Chevall.
372. Dothiora buxi Jayasiri, Camporesi & K.D. Hyde, in
Fungal Diversity 81: 30 (2016), new species
Hysteriales Lindau
Hysteriaceae Chevall.
373. Gloniopsis calami Konta. & K.D. Hyde, in Fungal
Diversity 81: 34 (2016), new species
Dictyosporiaceae Boonmee & K.D. Hyde
374. Pseudocoleophoma typhicola E.B.G. Jones, Kamolhan, Boonmee & K.D. Hyde, in Fungal Diversity 81:
34 (2016), new species
375. Pseudodictyosporium thailandica C.G. Lin, Yong
Wang bis & K.D. Hyde, in Fungal Diversity 81: 37
(2016), new species
Pleosporales Luttr. ex M.E. Barr
Didymellaceae Gruyter et al.
376. Neomicrosphaeropsis cytisi W.J. Li, Camporesi & K.D.
Hyde, in Fungal Diversity 81: 38 (2016), new species
377. Neomicrosphaeropsis cytisinus Tennakoon, Camporesi & K.D. Hyde, in Fungal Diversity 81: 39
(2016), new species
378. Neomicrosphaeropsis minima W.J. Li, Camporesi & K.D.
Hyde, in Fungal Diversity 81: 39 (2016), new species
Fungal Diversity (2016) 80:1–270
379. Neodidymelliopsis ranunculi W.J. Li, Camporesi &
K.D. Hyde, in Fungal Diversity 81: 41 (2016), new
species
380. Platychora ulmi (J. Schröt.) Petr., Annls mycol. 23(1/2):
103 (1925), reference specimen, p 41
381. Stagonosporopsis centaureae Tennakoon, Camporesi
& K.D. Hyde, in Fungal Diversity 81: 43 (2016), new
species
Didymosphaeriaceae Munk
382. Montagnula cirsii Qing Tian, Camporesi & K.D.
Hyde, in Fungal Diversity 81: 43 (2016), new species
383. Tremateia arundicola Wanasinghe, E.B.G. Jones &
K.D. Hyde, in Fungal Diversity 81: 45 (2016), new
species
384. Tremateia guiyangensis J.F. Zhang, J.K. Liu, K.D.
Hyde & Z.Y. Liu in Fungal Diversity 81: 48 (2016),
new species
Lentitheciaceae Y. Zhang et al.
5
Lophiotremataceae K. Hiray. & Kaz. Tanaka
393. Hermatomyces mirum (Starbäck) C.G. Lin, Yong
Wang bis & K.D. Hyde, in Fungal Diversity 81: 69
(2016), new combination
394. Hermatomyces subiculosa C.G. Lin, Yong Wang bis
& K.D. Hyde, in Fungal Diversity 81: 73 (2016), new
species
395. Lophiotrema bambusae Phookamsak, S.C. Karunarathana & K.D. Hyde, in Fungal Diversity 81: 73
(2016), new species
396. Lophiotrema fallopiae A. Hashim. & Kaz. Tanaka, in
Fungal Diversity 81: 74 (2016), new species
Massariaceae Nitschke
397. Neomassaria Mapook, Camporesi & K.D. Hyde, in
Fungal Diversity 81: 77 (2016), new genus
398. Neomassaria fabacearum Mapook, Camporesi &
K.D. Hyde, in Fungal Diversity 81: 77 (2016), new
species
385. Lentithecium unicellulare Abdel-Aziz, in Fungal
Diversity 81: 53 (2016), new species
386. Lentithecium voraginesporum Abdel-Wahab, Bahkali & E.B.G. Jones, in Fungal Diversity 81: 53
(2016), new species
399. Stagonospora forlicesenensis Phukhamsakda, Camporesi & K.D. Hyde, in Fungal Diversity 81: 77
(2016), new species
Leptosphaeriaceae M.E. Barr
Melanommataceae G. Winter
387. Leptosphaeria cirsii Jayasiri, Camporesi & K.D.
Hyde, in Fungal Diversity 81: 55 (2016), new species
388. Leptosphaeria irregularis R.H. Perera, E.B.G. Jones
& K.D. Hyde, in Fungal Diversity 81: 59 (2016), new
species
400. Bertiella ellipsoidea Ekanayaka, Q. Zhao & K.D.
Hyde, in Fungal Diversity 81: 79 (2016), new species
Lindgomycetaceae K. Hiray. et al.
389. Arundellina Wanasinghe, E.B.G. Jones & K.D. Hyde,
in Fungal Diversity 81: 59 (2016), new genus
390. Arundellina typhae Wanasinghe, E.B.G. Jones &
K.D. Hyde, in Fungal Diversity 81: 61 (2016), new
species
391. Lindgomyces pseudomadisonensis Tak. Takah. &
Kaz. Tanaka, in Fungal Diversity 81: 61 (2016), new
species
Lophiostomataceae Sacc.
392. Vaginatispora fuckelii (Sacc.) Thambugala, Wanasinghe, Kaz. Tanaka & K.D. Hyde, Fungal Diversity
74: 242. 2015, new host record, p 62
Massarinaceae Munk
Occultabambusaceae Dai et al.
401. Occultibambusa aquatica Huang Zhang & K.D.
Hyde, in Fungal Diversity 81: 81 (2016), new species
402. Occultibambusa chiangraiensis Phukhamsakda &
K.D. Hyde, in Fungal Diversity 81: 81 (2016), new
species
Phaeosphaeriaceae M.E. Barr
403. Camarosporioides W.J. Li & K.D. Hyde, in Fungal
Diversity 81: 83 (2016), new genus
404. Camarosporioides phragmitis W.J. Li & K.D. Hyde,
in Fungal Diversity 81: 85 (2016), new species
405. Chaetosphaeronema achilleae S.K. Huang & K.D.
Hyde, in Fungal Diversity 81: 85 (2016), new species
406. Dematiopleospora alliariae Thambugala, Camporesi
& K.D. Hyde, in Fungal Diversity 81: 89 (2016), new
species
123
6
Fungal Diversity (2016) 80:1–270
407. Dematiopleospora cirsii Wanasinghe, Camporesi,
E.B.G. Jones & K.D. Hyde, in Fungal Diversity 81:
89 (2016), new species
408. Juncaceicola italica Tibpromma, Camporesi & K.D.
Hyde, in Fungal Diversity 81: 93 (2016), new species
409. Leptospora rubella (Pers.) Rabenh., Klotzschii Herb.
Viv. Mycol., Edn 2: no. 532 (1857), reference specimen, p 93
410. Leptospora galii de Silva & K.D. Hyde, in Fungal
Diversity 81: 96 (2016), new species
411. Leptospora thailandica Phukhamsakda & K.D. Hyde,
in Fungal Diversity 81: 100 (2016), new species
412. Muriphaeosphaeria ambrosiae S.K. Huang & K.D.
Hyde, in Fungal Diversity 81: 104 (2016), new
species
413. Nodulosphaeria italica Phookamsak, Camporesi &
K.D. Hyde, in Fungal Diversity 81: 106 (2016), new
species
414. Poaceicola arundinis W.J. Li, Camporesi, D.J. Bhat
& K.D. Hyde, in Fungal Diversity 81: 111 (2016),
new species
415. Pseudophaeosphaeria Jayasiri, Camporesi & K.D.
Hyde, in Fungal Diversity 81: 111 (2016), new genus
416. Pseudophaeosphaeria rubi Jayasiri, Camporesi &
K.D. Hyde, in Fungal Diversity 81: 112 (2016), new
species
417. Wojnowicia italica Qing Tian, Camporesi & K.D.
Hyde, in Fungal Diversity 81: 112 (2016), new species
423. Helicoma guttulatum Y.Z. Lu, Boonmee & K.D. Hyde,
in Fungal Diversity 81: 125 (2016), new species
424. Neoacanthostigma septoconstrictum (Promp. & A.N.
Mill.) S. Boonmee & K.D. Hyde, Fungal Diversity
68(1): 279 (2014), reference specimen, 125
425. Tubeufia hyalospora Y.Z. Lu, Boonmee & K.D.
Hyde, in Fungal Diversity 81: 126 (2016), new
species
426. Tubeufia roseohelicospora Y.Z. Lu, Boonmee &
K.D. Hyde, in Fungal Diversity 81: 128 (2016), new
species
Pleosporaceae Nitschke
418. Comoclathris italica Tibpromma, Camporesi & K.D.
Hyde, in Fungal Diversity 81: 117 (2016), new species
430. Neomassarina Phookamsak & K.D. Hyde, in Fungal
Diversity 81: 136 (2016), new genus
431. Neomassarina thailandica Phookamsak & K.D. Hyde,
in Fungal Diversity 81: 138 (2016), new species
Roussoellaceae J.K. Liu et al.
Eurotiomycetes
419. Neoroussoella lenispora J.F. Zhang, J.K. Liu, K.D.
Hyde & Z.Y. Liu, in Fungal Diversity 81: 119 (2016),
new species
Eurotiales G.W. Martin ex Benny & Kimbr.
Dothideomycetes family, incertae sedis
Pleurotremataceae K.D. Hyde et al.
427. Pleurotrema thailandica Dayarathne, Jones E.B.G. &
K.D. Hyde, in Fungal Diversity 81: 131 (2016), new
species
Trypetheliaceae Eschw.
428. Alloarthopyrenia Phukhamsakda, Lücking & K.D.
Hyde, in Fungal Diversity 81: 131 (2016), new genus
429. Alloarthopyrenia italica Phukhamsakda, Camporesi,
Ariyawansa & K.D. Hyde, in Fungal Diversity 81:
135 (2016), new species
Pleosporales genera incetae sedis
Torulaceae Corda
432. Aspergillus koreanus Hyang B. Lee, T.T. Duong &
T.T.T. Nguyen, in Fungal Diversity 81: 142 (2016),
new species
420. Neotorula submersa Z.L. Luo, H.Y. Su & K.D. Hyde,
in Fungal Diversity 81: 121 (2016), new species
Chaetothyriales M.E. Barr
Chaetothyriaceae Hansf. ex M.E. Barr
Tubeufiales Boonmee & K.D. Hyde
Tubeufiaceae M.E. Barr
433. Ceramothyrium menglunense Mapook, J.F. Li & K.D.
Hyde, in Fungal Diversity 81: 142 (2016), new species
421. Aquaphila albicans Goh, K.D. Hyde & W.H. Ho, Mycol.
Res. 102(5): 588 (1998), reference specimen, p 121
422. Chlamydotubeufia helicospora Boonmee, Y. Z. Lu &
K.D. Hyde, in Fungal Diversity 81: 123 (2016), new
species
Herpotrichiellaceae
123
434. Minimelanolocus submersus Z.L. Luo, H.Y. Su &
K.D. Hyde, in Fungal Diversity 81: 143 (2016), new
species
Fungal Diversity (2016) 80:1–270
7
Trichomeriaceae Chomnunti & K.D. Hyde
Valsaceae Tul. & C. Tul.
435. Trichomerium bambusae Hongsanan & K.D. Hyde,
in Fungal Diversity 81: 145 (2016), new species
447. Cytospora cotini Norphanphoun, Bulgakov & K.D.
Hyde, in Fungal Diversity 81: 176 (2016), new species
Leotiomycetes
Rhytismatales M.E. Barr ex Minter
Rhytismataceae Chevall.
Glomerellales Chadef. ex Réblová et al.
Glomerellaceae Locq.
436. Terriera thailandica Jayasiri & K.D. Hyde, in Fungal
Diversity 81: 146 (2016), new species
Pezizomycetes
Pezialaes J. Schrot.
Helvelaceae Fr.
437. Helvella tinta Q. Zhao, B. Feng & K.D. Hyde, in
Fungal Diversity 81: 149 (2016), new species
438. Helvella floriforma Q. Zhao & K.D. Hyde, in Fungal
Diversity 81: 154 (2016), new species
439. Helvella oblongispora Harmaja, Karstenia 18(2): 57
(1978), new distribution record, p 157
Pezizaceae Dumort.
440. Peziza fruticosa Lantieri, Medardi & Vizzini, in
Fungal Diversity 81: 157 (2016), new species
Sordariomycetes
Coronophorales Nannf.
Coronophoraceae Höhn.
448. Colletotrichum insertae Jayawardena, Bulgakov &
K.D. Hyde, in Fungal Diversity 81: 176 (2016), new
species
Reticulascaceae Réblová & W. Gams
449. Blastophorum aquaticum Z.L. Luo, Bhat, H.Y. Su &
K.D. Hyde, in Fungal Diversity 81: 177 (2016), new
species
Hypocreales Lindau
Ophiocordycipitaceae G.H. Sung et al.
450. Ophiocordyceps hemisphaerica Mafalda-Freire, Reck &
Drechsler-Santos, in Fungal Diversity 81: 181 (2016),
new species
451. Ophiocordyceps lacrimoidis Mafalda-Freire, Reck &
Drechsler-Santos, in Fungal Diversity 81: 186 (2016),
new species
452. Purpureocillium sodanum Papizadeh, Soudi, Wijayaw.,
Shahz.-Faz. & K.D. Hyde, in Fungal Diversity 81: 186
(2016), new species
Hypocreales genus incertae sedis
441. Coronophora myricoides H.X. Wu & K.D. Hyde, in
Fungal Diversity 81: 164 (2016), new species
Diaporthales Nannf.
Diaporthaceae Höhn. ex Wehm.
Diaporthe Nitschke
442. Diaporthe aseana Dissanayake, Tangthirasunun &
K.D. Hyde, in Fungal Diversity 81: 167 (2016), new
species
443. Diaporthe eres Nitschke, Pyrenomycetes Germanici
2: 245 (1870), new host record, 167
444. Diaporthe foeniculina Niessl, in von Thümen, Contr.
Ad. Fl. Myc. Lusit. 2: 30. 1880. new record, p 169
445. Diaporthe garethjonesii Dissanayake, Tangthirasunun
& K.D. Hyde, in Fungal Diversity 81: 171 (2016),
new species
446. Diaporthe siamensis Udayanga et al., Cryptogamie
Mycologie 33: 298 (2012), new host record, p 171
453. Alfaria spartii Senan., Camporesi & K.D. Hyde, in
Fungal Diversity 81: 187 (2016), new species
Bionectriaceae Samuels & Rossman
454. Emericellopsis persica Papizadeh, Wijayaw, Soudi &
K.D. Hyde, in Fungal Diversity 81: 191 (2016), new
species
Meliolales Gäum. ex D. Hawksw. & O.E. Erikss.
Meliolaceae G.W. Martin ex Hansf.
455. Meliola citri-maximae X.Y. Zeng, K.D. Hyde & T.C.
Wen, in Fungal Diversity 81: 192 (2016), new species
456. Meliola pseudosasae I. Hino, Bull. Faculty of Agriculture, Yamaguchi University 9: 882 (1958), reference specimen, p 196
123
8
Xylariales Nannf.
Apiosporaceae K.D. Hyde et al.
457. Arthrinium paraphaeospermum Senan. & K.D. Hyde,
in Fungal Diversity 81: 198 (2016), new species
Bartaliniaceae Wijayaw. et al.
458. Neotruncatella Hyang B. Lee & T.T.T. Nguyen, in
Fungal Diversity 81: 198 (2016), new genus
459. Neotruncatella endophytica Hyang B. Lee, P.M.
Kirk, K.D. Hyde, S.S.N. Maharachch, & T.T.T.
Nguyen, in Fungal Diversity 81: 198 (2016), new
species
Fungal Diversity (2016) 80:1–270
Distoseptisporaceae Su et al.
470. Distoseptispora multiseptata J. Yang & K.D. Hyde, in
Fungal Diversity 81: 220 (2016), new species
471. Distoseptispora tectonae Doilom & K.D. Hyde, in
Fungal Diversity 81: 222 (2016), new species
472. Distoseptispora tectonigena Doilom & K.D. Hyde, in
Fungal Diversity 81: 222 (2016), new species
Sordariomycetidae, Incertae sedis
473. Paracapsulospora Konta & K.D. Hyde, in Fungal
Diversity 81: 223 (2016), new genus
474. Paracapsulospora metroxyli Konta & K.D. Hyde, in
Fungal Diversity 81: 223 (2016), new species
Cainiaceae J.C. Krug
460. Cainia globosa Senan., Camporesi & K.D. Hyde, in
Fungal Diversity 81: 201 (2016), new species
Basidiomycota
Agaricomycetes
Agaricales Underw.
Clavariaceae Chevall.
Pestalotiopsidaceae Maharachch. & K.D. Hyde
461. Ciliochorella phanericola Norphanphoun, T.C. Wen
& K.D. Hyde, in Fungal Diversity 81: 207 (2016),
new species
462. Neopestalotiopsis cocoës Norphanphoun, T.C. Wen &
K.D. Hyde, in Fungal Diversity 81: 207 (2016), new
species
463. Neopestalotiopsis musae Norphanphoun, T.C. Wen &
K.D. Hyde, in Fungal Diversity 81: 209 (2016)
464. Pestalotiopsis sequoiae W.J. Li, Camporesi & K.D.
Hyde, in Fungal Diversity 81: 210 (2016), new species
Xylariaceae Tul. & C. Tul.
465. Anthostomella ravennica Daranagama, Camporesi &
K. D. Hyde, in Fungal Diversity 81: 210 (2016), new
species
466. Anthostomella thailandica Daranagama & K.D.
Hyde, in Fungal Diversity 81: 213 (2016)
Sordariomycetes incertae sedis
Sporidesmiaceae Fr.
467. Sporidesmium pyriformatum J. Yang & K.D. Hyde,
in Fungal Diversity 81: 215 (2016), new species
468. Sporidesmium aquaticivaginatum J. Yang & K.D.
Hyde, in Fungal Diversity 81: 217 (2016), new
species
469. Sporidesmium olivaceoconidium J. Yang & K.D.
Hyde, in Fungal Diversity 81: 220 (2016), new
species
123
475. Clavulinopsis aurantiaca Araujo-Neta, Silva & Gibertoni, in Fungal Diversity 81: 225 (2016), new
species
476. Ramariopsis atlantica Araujo-Neta, Silva & Gibertoni, in Fungal Diversity 81: 226 (2016), new species
Cortinariaceae R. Heim ex Pouzar
477. Cortinarius fulvescens Fr., Epicr. syst. mycol. (Upsaliae): 311 (1838), neotype, 227
478. Cortinarius fulvescentoideus Kytöv., Niskanen &
Liimat., in Fungal Diversity 81: 230 (2016), new
species
479. Cortinarius nymphatus Kytöv., Niskanen, Liimat. &
Bojantchev, in Fungal Diversity 81: 230 (2016), new
species
480. Cortinarius pseudobulliardioides Kytöv., Niskanen,
Liimat. & Ammirati, in Fungal Diversity 81: 232
(2016), new species
481. Cortinarius tenuifulvescens Kytöv., Niskanen &
Liimat., in Fungal Diversity 81: 232 (2016), new
species
Entolomataceae Kotl. & Pouzar
482. Rhodocybe indica K.N.A. Raj & Manim., in Fungal
Diversity 81: 236 (2016), new species
483. Rhodocybe luteobrunnea K.N.A. Raj & Manim., in
Fungal Diversity 81: 241 (2016), new species
484. Rhodocybe griseoaurantia K.N.A. Raj & Manim., in
Fungal Diversity 81: 242 (2016), new species
Fungal Diversity (2016) 80:1–270
Agaricaceae Chevall.
485. Cyathus pyristriatus B. Thongbai, C. Richt. & M. Stadler,
in Fungal Diversity 81: 244 (2016), new species
Polyporales Gäum.
Polyporaceae Fr. ex Corda
486. Polyporus mangshanensis B.K. Cui, J.L. Zhou &
Y.C. Dai, in Fungal Diversity 81: 249 (2016), new
species
Russulales Kreisel ex P.M. Kirk et al.
Russulaceae Lotsy
487. Russula indoalba A. Ghosh, Buyck, A. Baghela, K. Das
& R.P. Bhatt, in Fungal Diversity 81: 250 (2016), new
species
488. Russula pseudoamoenicolor A. Ghosh, Buyck, K.
Das, A. Baghela & R.P. Bhatt, in Fungal Diversity
81: 251 (2016)
489. Lactarius dirkii Uniyal, K. Das, A. Baghela & R.P.
Bhatt, in Fungal Diversity 81: 252 (2016), new species
Mortierellomycotina Kerst. Hoffm. et al.
Mortierellales Caval.-Sm.
Mortierellaceae A. Fisch.
Mortierella fluviae Hyang B. Lee, K. Voigt & T.T.T.
Nguyen, in Fungal Diversity 81: 254 (2016), new species
Mucorales Dumort.
Cunninghamellaceae Naumov ex R.K. Benj.
490. Cunninghamella gigacellularis A.L. Santiago, C.L.
Lima & C.A. de Souza, in Fungal Diversity 81: 255
(2016), new species
Materials and methods
Sampling, isolation and identification
Specimens from a number of plants examined in this
study were collected from at least 15 countries around the
world, including Brazil, China, Egypt, Germany, Iran,
India, Italy, Japan, Philippines, Republic of Korea, Russia, Saudi Arabia, Thailand and the UK. Soil samples
collected from the Atlantic rainforest in Recife, Brazil to
isolate Cunninghamella gigacellularis followed methods
outlined by Benny (2008). The color designation of
C. gigacellularis colonies was established according to
Maerz and Paul (1950). Samples were observed under the
microscope; macro-morphological (e.g. from basidiomycetes) and micro-morphological characters (e.g.
9
ascomata sections, peridium structures, asci and ascospores) were examined, described and photographed. Most
measurements (e.g. from basidiospores and ascospores)
were taken from at least 20 representatives and both the
mean and the standard deviation for both the length and
the width, together with the range of spore quotient (Q,
the length/width ratio) and its mean value (Qm) are given.
Axenic cultures (following single spore isolations) and
specimens with duplicates have been deposited in Culture
Collections and Herbaria where appropriate and accession
numbers are provided in the taxonomic descriptions.
Faces of Fungi and Index Fungorum numbers are given as
outlined by Jayasiri et al. (2015) and Index Fungorum
(2016). Wherever possible, appropriate techniques were
employed to induce the formation of the asexual morph in
culture using sterilized pieces of plant materials. Colony
characters were observed and recorded. We have tried to
maintain consistency in terminology, however this may
not always be possible as different authors prefer to use
different terms. For example the use of ascostromata is
variable and often difficult to interpretate and is left to
each author’s discretion.
DNA extraction, PCR amplification and sequencing
For most fungal samples (especially ascomycetes), total
genomic DNA was extracted from fresh fungal mycelium
grown on appropriate media agar at room temperature with
appropriate Genomic DNA Extraction Kit purchased from
countries where samples were collected following manufacturer’s instructions. Under circumstances where fungi
fail to grow in culture, DNA was extracted directly from
fruiting bodies using aseptic techniques. The procedure
described by Izumitsu et al. (2012) was employed for
extracting genomic DNA from dried Rhodocybe specimens. For Cunninghamella isolates, genomic DNA
extraction was carried out with macerated material
according to Góes-Neto et al. (2005).
DNA amplification for most samples was performed by
polymerase chain reaction (PCR) using universally standard primers such as LROR and LR5 (Vilgalys and Hester
1990) for the partial large subunit nuclear rDNA (28S,
LSU); NS1 and NS4 (White et al. 1990) for the small
subunit nuclear rDNA (18S, SSU); ITS4 and ITS5 (White
et al. 1990) for the internal transcribed spacers (5.8S, ITS);
EF1-983F and EF1-2218R (Rehner 2001) for the translation elongation factor 1-alpha gene (TEF1a); and fRPB25F and fRPB2-7cR (Liu et al. 1999) for the partial RNA
polymerase second largest subunit (RPB2). Primer pair
LR1/LSU2 were used to amplify LSU region of Cunninghamella isolates (van Tuinen et al. 1998) while LROR/
LR7 for (Vilgalys and Hester 1990) and b6Fb7.1R
123
10
Fungal Diversity (2016) 80:1–270
Abrothallales
Abrothallus parmotrematis AB1
Melaspileopsis diplasiospora Ertz 16247
88/1.0
Melaspileopsis sp. Ertz 17904
100/1.0 Stictographa lentiginosa Ertz 17570
100/1.0
Stictographa lentiginosa Ertz 17447
66/0.9
Karschia talcophila 16749
100/1.0
Karschia cezannei Cezanne-Eichler 7453
95/1.0 100/1.0 Buelliella physciicola Ertz 19173
Buelliella physciicola Ertz 18113
/0.9
Labrocarpon canariense 16907
99/1.0
Labrocarpon canariense 16308
Asterina zanthoxyli TH 561
100/1.0
Asterina fuchsiae TH 590
Asterinales
Asterina weinmanniae TH 592
Asterina sp. MFU13-0619
94/1.0
Asterina cestricola TH 591
50/
Asterina phenacis TH 589
64/0.89
Asterina siphocampyli ppMP 1324
66/0.9
Asterina cynometae MFLU 13–0373
100/1.0
Inocyclus angularis VIC 39747
Lembosia albersii MFU13-0377
100/1.0
Lembosia xyliae MFLU14-0004
100/1.0
Buelliella poetschii Ertz 18115
100/1.0
Buelliella poetschii Ertz 18116
Asterotexis cucurbitacearum PMA M-0141224
Parmularia styracis VIC 42587
100/1.0
Parmularia styracis VIC 424477
Prillieuxina baccharidincola VIC 42817
98/1.0
Incertae sedis clade
Asterina melastomatis VIC 42822
100/1.0
Asterina chrysophylli VIC 42823
96/1.0 Lembosia abaxialis VIC 42825
99/1.0
54/0.9
Batistinula sp. MFLU 13-0514
51/0.89
Batistinula gallesiae VIC 42514
Coniosporium uncinatum CBS 100212
Capronia munkii AFTOL-ID 656
0.1
Fig. 1 Phylogram generated from maximum likelihood and Bayesian
analyses based on LSU sequence data from species of Asterinales and
Asterotexales. The first set of numbers above the nodes are RAxML
bootstrap value expressed from 1000 repetitions with values above
50 % shown. The second set of numbers above the nodes are
Bayesian posterior probabilities, with values above 0.85 shown. The
new isolates are in blue bold and other ex-type strains are in bold. The
tree is rooted with Capronia munkii Unter
(Matheny 2005) for RPB2 region were used for Rhodocybe
specimens. Thermal cycle program for ITS, LSU and
TEF1a amplification were as follows: initial 94 °C for
3 mins, followed by 35 cycles of denaturation at 94 °C for
30 s, annealing at 55 °C for 50 s, elongation at 72 °C for 1
mins, and final extension at 72 °C for 10 mins. For ascomycetes, the thermal cycle program for RPB2 was as follows: initial 95 °C for 5 mins, followed by 40 cycles of
denaturation at 95 °C for 1 mins, annealing at 52 °C for 2
mins, elongation at 72 °C for 90 s, and final extension at
72 °C for 10 mins. Thermal cycle program for Cunninghamella isolates were follows: 95 °C for 5 mins, followed
by 39 cycles of denaturation at 94 °C for 45 s, annealing at
60 °C for 1 min, elongation at 72 °C for 1 min, and final
extension at 72 °C for 7 mins. For Rhodocybe specimens,
thermal profiling and amplification reactions of ITS, nLSU
and RPB2 regions were performed following Latha et al.
(2015). Prior to sequencing, quality of PCR amplicons
were checked and purified with appropriate purification
kits as per manufacturer’s guidelines before being subjected to automated DNA sequencing using the same primers used for PCR.
Sequence alignment and phylogenetic analyses
123
A careful verification of all sequences obtained were done
especially with appropriate reference sequences following
a Blast search in GenBank to ensure that no erroneous
sequences are used in further analyses and then submitted
to GenBank. Following sequence verification and Blast
search, DNA sequences from appropriate taxonomic ranks
were downloaded to construct datasets for phylogenetic
analyses. BioEdit sequence alignment editor (Hall 1999),
AliView v.1.17 (Larsson 2014), CLUSTALX (Larkin et al.
2007), Mega 6.0.5 (Tamura et al. 2013) and MAFFT:
multiple sequence alignment software version 7.215
(Katoh et al. 2002) were used for alignment purposes.
Under most circumstances, concatenated DNA datasets are
analyzed to generate gene trees but in cases of limited
availability of DNA sequences from respective gene
regions, phylogenies are inferred from single or two genes
datasets and either a consensus of these gene phylogenies
or one of the most parsimonious phylogeny is used to infer
phylogenetic relationships across taxa sampled. Selection
of outgroup (s) for rooting purposes was based on
Fungal Diversity (2016) 80:1–270
11
Fig. 2 Asterina cynometrae (holotype). a Appearance of thyriothecia on leaf. b, c Thyriothecia with star-like opening when viewed in
squash mounts. d Superficial hyphae with hyphopodia. e Upper wall
of thyriothecium, f Ascus in Melzer’s reagent. g Ascus at maturity. h,
i Ascospores. Scale bars b, c = 100 lm, d, e = 20 lm, f–i = 10 lm
knowledge of potential common ancestor to our in-group
as well as taxon sampling from previously published
studies. Phylogenetic analyses were performed by maximum parsimony (MP), maximum likelihood (RAxML) and
Bayesian inference (BI) analyses.
A maximum parsimony (MP) analysis was performed
with stepwise additions of sequences by using PAUP v.
4.0b10 (Swofford 2002). The heuristic search option with
1000 random sequences addition and tree-bisection
reconnection (TBR) of branch-swapping algorithm were
123
12
performed. Maxtrees were setup at 5000, branches of zero
length were collapsed and gaps were treated as missing
data. Tree length (TL), consistency index (CI), retention
index (RI), rescaled consistency index (RC) and homoplasy
index (HI) were calculated for trees generated under different optimality criteria. The robustness of the most parsimonious trees was evaluated by 1000 bootstrap
replications resulting from maximum parsimony analysis
with each 100 replicates of random stepwise addition of
taxa (Liu et al. 2011, 2012).
Maximum likelihood (ML) analysis was performed
using RAxML v.8.0.26 (Stamatakis 2014) with 1000 rapid
ML bootstrap replicates. The available substitution models
comprised a generalized time reversible (GTR) for
nucleotides was applied with a discrete gamma distribution
(Silvestro and Michalak 2012). A discrete GAMMA (Yang
1994) was complemented for each substitution model.
Rapid bootstrap analysis (Stamatakis et al. 2008) and
search for a best-scoring ML tree were applied (Silvestro
and Michalak 2012). The parameters were followed; run
mode = ML ? rapid bootstrap, replicates = 1000, BS
brL = selected and Model = GTRGAMMA.
Bayesian analysis was performed by MrBayes v. 3.0b4
(Ronquist and Huelsenbeck 2003) with the best-fit model
of sequence evolution estimated with MrModeltest 2.2
(Nylander et al. 2008). Markov Chain Monte Carlo sampling (BMCMC) was used to determine the posterior
probabilities (PP) (Rannala and Yang 1996; Zhaxybayeva
and Gogarten 2002) in MrBayes v. 3.0b4 (Huelsenbeck and
Ronquist 2001). Six simultaneous Markov chains were run
for 1,000,000 generations sampling one tree every 100th
generations of trees (resulting 10,001 total trees). The burnin (first 2000 trees) which represented the phase of the
analysis were discarded and the remaining 8000 trees were
used to build a majority rule consensus tree (Liu et al.
2011, 2012) with posterior probabilities (PP).
Phylograms were visualized in Treeview (Page 2001) or
FigTree 1.4.2 (Rambaut 2014) with bootstrap values above
or below the nodes. All the sequences generated in this
study have been deposited in GenBank and accession
numbers provided where appropriate.
Dothideomycetes
We follow Hyde et al. (2013) and Wijayawardene et al.
(2014a) for the latest arrangement of this class.
Asterinales M.E. Barr ex D. Hawksw. & O.E. Erikss.
The order was monographed by Hongsanan et al.
(2014). Guatimosim et al. (2015) provided sequence data
for Asterina and Parmularia directly from ascomata, but
did not include other sequence data used in Hofmann
(2009). Ertz and Diederich (2015) provided sequence data
for taxa of Melaspileaceae and placed them in Asterinales
123
Fungal Diversity (2016) 80:1–270
based on their phylogenetic data. However, Ertz et al.
(2016) indicated that Asterina species are segregated in two
unrelated clades. Asterotexis cucurbitacearum and Inocyclus angularia (Parmulariaceae) clustered with strains of
Asterina provided by Hofmann (2009) and Hongsanan
et al. (2014), and formed a sister group to Melaspileaceae
(Ertz et al. 2016). Thus, these Asterinales strains were
considered to represent the order Asterotexiales based on
the type species of Asterotexis cucurbitacearum (Ertz et al.
2016). In the present study we treat the main Asterinales
clade, which includes Asterotexis cucurbitacearum, as
Asterinales sensu stricto because most of the Asterinales
strains, from both Asterina and Lembosia cluster in this
clade and also because the large clade supporting Asterinales as circumscribed by Ertz et al. (2016) does not have
any phylogenetic support. We therefore synonymize the
younger Asterotexiales (in December, 2015) under Asterinales (in 1986). It is questionable that the other clade,
which contains the putatively named, type species of Asterina is actually Asterinales as most strains in this order
cluster in Asterinales sensu stricto. It may be that DNA was
amplified from other taxa in the black mildew colonies.
Since Parmularia represents a distinct monophyletic clade
with high support outside Asterinales sensu stricto, we
reinstate Parmulariaceae to accommodate this clade
(Fig. 1).
Asterinaceae Hansf.
The family Asterinaceae was established in Myriangiales by Hansford (1946). Several studies placed Asterinaceae in an uncertain position in the Dothideomycetes
incertae sedis (Cannon and Kirk 2007; Kirk et al. 2008).
Phylogenies of Hongsanan et al. (2014) place Asterinaceae
within Asterinales in Dothideomycetes. They also accepted
17 genera in the family based mainly on morphology. In
this paper, we introduce a new species, Asterina cynometrae with morphological details and molecular data
(Figs. 1, 2).
Asterina Lév.
The genus Asterina is the type genus of the family Asterinaceae which was introduced by Léveillé (1845). Asterina is the largest genus in Asterinaceae, and has a
worldwide distribution in tropical and subtropical regions
(Hongsanan et al. 2014). According to Ertz et al. (2016)
Asterina species cluster in two well-supported distinct
clades, which are placed in the order Asterinales and in the
incertae sedis clade in this study (Fig. 1).
Asterina cynometrae Hongsanan & K.D. Hyde, sp. nov.
Index Fungorum number: IF552216; Facesoffungi
number: FoF02430, Fig. 2
Holotype: MFLU 13-0373.
Fungal Diversity (2016) 80:1–270
13
Fig. 3 One of 309 most parsimonious trees obtained from combined
ITS and EF-1a sequence data, for all ex-types from species in
Dothiorella. Isolate numbers of new host records are in blue.
Maximum parsimony bootstrap values ([70 %) and Bayesian inference values ([0.9) are given on the nodes. The tree is rooted with
Spencermartinsia viticola
Epiphytes on the upper surface of leaves. Superficial
hyphae branched, septate, darker at the septum, brown,
with hyphopodia. Hyphopodia 11–13 lm high 9 4–8 lm
wide (
x = 12 9 6 lm, n = 20) capitate, alternate, rarely
opposite on hyphae, near to hyphal septum, 1-celled, with 2
branches at the apex, brown. Sexual morph Thyriothecia
150–200 lm diam. (
x = 160 lm, n = 10), superficial on
the surface of host, solitary to gregarious, circular, flattened, with star-like opening, sometimes variously shaped.
Upper wall comprising parallel arrangement of cells radiating from the center, base poorly developed. Hamathecium not observed. Asci up to 40 lm diam., vertically
arranged within thyriothecium cavity, 6-spored, bitunicate,
fissitunicate, subglobose to globose, short pedicellate,
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Fungal Diversity (2016) 80:1–270
Fig. 4 Dothiorella iranica (MFLU 15-1402). a–c Conidiomata on host surface. d Section through the conidioma. e Conidioma wall. f–
i Conidiophore and conidiogenous cells. j–m Conidia. Scale bars d = 50 lm, e = 20 lm, f–m = 10 lm
without an ocular chamber, with evanescent wall. Ascospores 18–22 lm 9 10–12 lm (
x = 20 9 11 lm,
n = 15), 2–3-seriate, oblong, hyaline to dark brown,
uniseptate at the middle, constricted and with dark band at
the septum, smooth-walled, ends rounded. Asexual morph
Undetermined.
Material examined: PHILIPPINES, Luzon, Laguna
Province, Mount Makiling, on living leaves of Cynometra
sp. (Fabaceae), February 2012, Pamela Alva (MFLU
13-0373, holotype).
Notes: Asterina cynometrae was collected on living
leaves of Cynometra sp. (Fabaceae) and is most similar to
A. trachycarpa Syd. & P. Syd. in the shape and size of
ascospores, the latter species being found on Derris atroviolacea Elmer (Fabaceae) in the Philippines. However, it
differs from A. trachycarpa in having a thick and dark band
at the septum of ascospores, with 2–3 branching
hyphopodia. We therefore introduce A. cynometrae as a
new species based on host and morphology (Fig. 2).
123
Furthermore, we place our new species within Asterinales
based on the phylogeny of LSU sequence data (Fig. 1). As
we were unable to isolate the new taxon because of its
obligate parasitic habitat, sequence data was prepared
directly from thyriothecia and ascospores.
Botryosphaeriales C.L. Schoch et al.
The order Botryosphaeriales has undergone significant
taxonomic changes during the past decade by addition of
several new families. Currently, there are seven families,
Aplosporellaceae, Botryosphaeriaceae, Melanopsaceae,
Planistromellaceae, Phyllostictaceae, Saccharataceae and
Septorioideaceae (Schoch et al. 2006; Minnis et al. 2012;
Wikee et al. 2013; Slippers et al. 2013; Wyka and Broders
2016) and several new genera (Liu et al. 2012; Crous et al.
2015b). Botryosphaeriales is a diverse order with a
worldwide distribution, comprising species that vary from
endophytes to pathogens (Slippers and Wingfield 2007)
and occurring on a wide range of monocotyledonous,
Fungal Diversity (2016) 80:1–270
15
Fig. 5 Dothiorella sarmentorum (MFLU 16-1274). a Conidiomata
on Morus alba. b Vertical section through a conidioma. c Peridium of
conidioma. d–g Conidia attached to conidiogenous cells. h–l Mature
and immature conidia. Scale bars a = 500 lm, b = 100 lm,
c = 50 lm, d = 20 lm, e–l = 10 lm
dicotyledonous, and gymnosperm hosts (Liu et al. 2012;
Crous et al. 2015b) and lichens (Barr 1987; von Arx 1987).
Many are considered as pathogens that cause disease on a
wide range of economically and ecologically significant
plants (Slippers et al. 2013).
is almost impossible without the support of molecular data
(Slippers et al. 2013). We provide an updated tree for the
genus (Fig. 3).
Dothiorella Sacc.
The species in the genus Dothiorella (Botryosphaeriaceae, Botryosphaeriales, Dothideomycetes) is characterized based on conidia that become pigmented and 1-septate
while they are still attached to the conidiogenous cells
(Phillips et al. 2013). Due to wide host ranges and morphological plasticity, identification of species in this genus
Dothiorella iranica Abdollahz. et al., in Abdollahzadeh
et al., Persoonia, Mol. Phyl. Evol. Fungi 32: 4 (2014)
Facesoffungi number: FoF02202, Fig. 4
Saprobic on Paliurus bark. Asexual morph Conidiomata 280–305 lm high, 275–310 lm diam. (
x=
290 9 295 lm, n = 5), acervular, solitary to gregarious,
superficial to immersed, unilocular, globose to subglobose,
dark brown to black. Conidiomata wall 30–40 lm wide,
composed of brown, thin or thick-walled cells of textura
123
16
Fig. 6 Dothiorella vidmadera (MFLU 16-1273). a Robinia pseudoacacia with fungus. b Conidiomata on host substrate. c Vertical
section through a conidioma. d Peridium of conidioma. e Conidium
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Fungal Diversity (2016) 80:1–270
attached to conidiogenous cell. f–i Mature and immature conidia.
Scale bars b = 500 lm, c = 100 lm, d = 20 lm, e–i = 10 lm
Fungal Diversity (2016) 80:1–270
angularis, apex and base thicker than middle, with setae.
Conidiophores 7–9 9 2–3 lm (
x = 8 9 2.5 lm, n = 10),
cylindrical, filiform, septate, branched, hyaline. Conidiogenous cells 9–17 9 4–6 lm, (
x = 12 9 5 lm, n = 20),
holoblastic, annellidic, integrated or discrete, hyaline,
determinate. Conidia 20–25 9 8–11 lm (
x = 22 9 9 lm,
n = 30), cylindrical, oval or ellipsoid, 1-septate, hyaline
when immature, brown to dark brown when mature. Sexual
morph Undetermined.
Culture characteristics: Ascospores germinating on
MEA within 36 h. Colonies growing on MEA attaining
2 cm diam. in 1 week at 28 °C. Mycelium superficial,
felted, gummy, dark brown to black. Asexual structures not
formed in culture.
Material examined: ITALY, Province of Forlı̀-Cesena,
Monte Pallareto—Meldola, on Paliurus bark (Rhamnaceae), 1 January 2012, Erio Camporesi IT 962 (MFLU
15-1402, KUN, new host record), living culture,
MFLUCC 15-0656, KUNCC
Notes: Phylogenetically this species resides in a distinct
subclade in Dothiorella with high support and closely
related to D. ulmacea (Fig. 3). The conidia of D. iranica
are longer than those of all other Dothiorella species except
D. casuarini J. de Wet et al. (27 9 11 lm). Our isolate is
morphologically and phylogenetically similar to Dothiorella iranica (strain IRAN 1587C), but associated with a
different host. Dothiorella iranica (type) was recorded on
Olea europaea L. (Oleaceae) and our collection was found
on Paliurus (Rhamnaceae) bark. Host distribution is poorly
known in this genus, but according to Dissanayake et al.
(2016), Dothiorella species may be host specific, although
Dothiorella iberica A.J.L. Phillips et al., D. sarmentorum
(Fr.) A.J.L. Phillips et al. and D. symphoricarposicola W.J.
Li occur on many host families and orders.
Dothiorella sarmentorum (Fr.) A.J.L. Phillips, J. Luque &
A. Alves, Mycologia 97: 522. 2005
: Sphaeria sarmentorum Fr., K. svenska VetenskAcad. Handl. 39: 107. 1818.
: Diplodia sarmentorum (Fr.) Fr., Summ. veg. Scand.
(Stockholm) 2: 417. 1849.
= Botryosphaeria sarmentorum A.J.L. Phillips, J. Luque
& A. Alves, Mycologia 97: 522. 2005.
Facesoffungi number: FoF02148, Fig. 5
Saprobic on Celtis occidentalis L. Sexual morph
Undetermined. Asexual morph Conidiomata 140–240 lm
high 9 175–300 lm diam. (
x = 230 9 180 lm, n = 10),
stromatic, solitary or scattered in small groups, immersed,
uni or biloculate, black, globose to subglobose, ostiolate.
Conidiomatal wall 25–45 lm (
x = 35 lm, n = 15), comprising several layers; outer layers comprising thick-walled, dark brown, somewhat flattened cells of textura
angularis and inner layers of larger, thin-walled, lightly
17
pigmented or hyaline cells. Conidiophores reduced to
conidiogenous cells. Conidiogenous cells 6–12 9 2–4 lm
(
x = 8.6 9 3.3 lm, n = 15), lining the conidiomatal cavity, holoblastic, hyaline, subcylindrical, proliferating at the
same level giving rise to periclinal thickenings. Conidia
17.8–22.4 9 8.3–11 lm (
x = 20.2 9 9.9 lm, n = 30),
ovoid, with a broadly rounded apex and truncate base,
initially hyaline to lightly pigmented and aseptate,
becoming dark brown and 1-septate, slightly constricted at
the septum, smooth-walled.
Culture characteristics: Conidia germinating on PDA
within 18 h and germ tubes produced from one end or both
cells. Colonies on PDA at 25 °C, covering 9 cm Petridishes in few days, circular, flat, dense, surface initially
white, becoming grey with reverse black, smooth surface
with entire to slightly undulate edge.
Material examined: RUSSIA, Rostov Region, Shakhty
City, former Shakhty forestry, on Celtis occidentalis L.
(Cannabaceae), 26 February 2014, Timur Bulgakov T 35
(MFLU 16-1008, new host record), living culture
MFLUCC 15-0443. RUSSIA, Rostov region, Shakhty city,
Central Park (47.7055886°E, 40.2059913°N), on Morus
alba (Moraceae), 12 March 2014, Timur S. Bulgakov T 07
(MFLU 16-1274, new host record), living culture,
MFLUCC 14-0889, MUCL.
Notes: Dothiorella sarmentorum was introduced by
Phillips et al. (2005) based on the asexual morph of
Botryosphaeria sarmentorum A.J.L. Phillips et al. This
species has been recorded from 34 different host species
(Phillips et al. 2005, 2013). The sexual morph of D. sarmentorum is characterized by partially erumpent ascomata
with papillate ostiole, 4–6(–8)-spored asci and oblong to
ovate, (0–)1-septate, finely verruculose ascospores, which
are widest in the middle part (Phillips et al. 2013). As the
morphological information does not provide any clear
difference, we would like to report our collections as new
host records of D. sarmentorum. This is the first report of
Dothiorella sarmentorum on Celtis occidentalis and Morus
alba from Russia. ITS and EF-1a based phylogenies also
depict that all D. sarmentorum isolates cluster together and
is phylogenetically related to D. americana (Fig. 3)
Dothiorella vidmadera W.M. Pitt et al., Fungal Diversity
61: 216 (2013)
Facesoffungi number: FoF02206, Fig. 6
Saprobic or weak pathogenic on twigs of Robinia
pseudoacacia L. Sexual morph Undetermined. Asexual
morph Conidiomata 240–280 lm high 9 280–320 lm
diam. (
x = 263 9 298 lm, n = 10), pycnidial, stromatic,
mostly solitary, semi-immersed to immersed in the host,
globose, dark brown to black, ostiolate, apapillate.
Peridium 20–30 lm wide at the base, 25–35 lm wide at
the side, comprising 5–6 layers, heavily pigmented, thick-
123
18
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Fungal Diversity (2016) 80:1–270
Fungal Diversity (2016) 80:1–270
b Fig. 7 Phylogram generated from maximum likelihood analysis
(RAxML) based on combined ITS and LSU sequence data of
respective genera in Mycosphaerellaceae. Bootstrap support values
for maximum likelihood (ML, left) and maximum parsimony (MP,
right) equal to or greater than 50 % are given above the nodes. The
values of the Bayesian posterior probabilities from MCMC analyses
(BYPP) equal or higher than 95 % are given below the nodes. The
tree is rooted with Lecanosticta acicola (CBS 871.95). Ex-type and
ex-epitype strains are in bold. The generated sequences in this study
are indicated in blue
walled, blackish to dark brown, angular cells, becoming
flattened towards the outer layers. Conidiogenous cells 5–
9 lm high 9 2–4 lm wide, holoblastic, cylindrical to
subcylindrical, hyaline, the first conidium produced
holoblastically and subsequent conidia enteroblastically
forming typical phialides with periclinal thickenings,
swollen at the base, discrete, producing a single conidium
at the apex. Conidia 18–22 9 8–10 lm (
x = 19.9 9
8.6 lm, n = 30), initially hyaline, unicellular, becoming
dark brown and 1-septate while still attached to conidiogenous cells; detached conidia, hyaline, sepia or blackishbrown, unicellular or 1-septate, moderately thick-walled,
wall externally smooth, roughened on the inner surface,
oval to ovoid, widest in the center, apex obtuse, base
truncate or rounded, guttulate when young.
Culture characteristics: Colonies on MEA reaching
5 cm diam. after 30 days at 16 °C, circular, smooth margin, greyish-green to blackish-green after 28 days flat on
the surface, without aerial mycelium, reverse greyishbrown to black. Hyphae septate branched, hyaline, thin,
smooth-walled.
Material examined: RUSSIA, Rostov region, Rostov-naDonu city, Botanical garden of Southern Federal University,
Higher
Park,
underwood
(47.2389405°E,
39.6484137°N), on Robinia pseudoacacia (Fabaceae), 26
March 2014, Timur S. Bulgakov T 06 (MFLU 16-1273,
new host record), living culture, MFLUCC 14-0888,
MUCL.
Notes: In the combined phylogenetic analysis (ITS and
EF1-a), MFLUCC 14-0888 strain is phylogenetically most
closely related to D. vidmadera (Fig. 3). Our isolate
resembles D. vidmadera in the shape and size of the
conidia but our collection (MFLU 16-1273) has darker
conidia than DAR78992 of Dothiorella vidmadera (Pitt
et al. 2013). As the morphological information does not
provide any clear divergence, we report our collection as a
new host record of D. vidmadera. This species has been
previously recorded only from Vitis vinifera and Fraxinus
19
ornus. This is the first record of Dothiorella vidmadera on
Robinia species.
Capnodiales Woron.
The order Capnodiales comprises human pathogens,
plant saprotrophs and rock-inhabiting species. This order
was reviewed by Chomnunti et al. (2011) and in this paper
we follow the recent publication of Hyde et al. (2013).
Mycosphaerellaceae Lindau
The family Mycosphaerellaceae was introduced by
Lindau (1897) and typified by Mycosphaerella, with
M. punctiformis (Pers.) Starbäck as the type species. The
family was designated to accommodate Dothideomycete
species having small ascomata and often ascostromata
forming on various hosts, mostly parasitic, but also
saprobic on dead plants (von Arx and Müller 1975; Hyde
et al. 2013). Species in Mycosphaeriallaceae lack pseudoparaphyses and ascospores are often 2-celled, oblong to
clavate, or ellipsoidal (Hyde et al. 2013). Based on these
characters, the family was initially placed in the order
Dothideales (von Arx and Müller 1975; Hawksworth et al.
1995; Hyde et al. 2013; Liu et al. 2015a). Kirk et al. (2001)
treated the family in a separate order—Mycosphaerellales.
Schoch et al. (2006) assigned Mycosphaerellaceae to
Capnodiales based on phylogenetic support and this was
followed by various mycologists (Crous et al. 2007, 2009;
Kirk et al. 2008; Hyde et al. 2013; Wijayawardene et al.
2014a; Liu et al. 2015a). Recently, more than 50 sexual and
asexual genera have been accommodated in Mycosphaerellaceae (Wijayawardene et al. 2014a).
Pallidocercospora Crous et al.
Pallidocercospora was introduced by Crous et al. (2013)
to accommodate cercospora-like species, but not congeneric with Cercospora and is typified by P. heimii
(Crous) Crous. Crous et al. (2013) designated the genus
based on its pale brown cercosporoid conidia, which are
generally referred to as the Mycosphaerella heimii complex
(Crous et al. 2004a, 2013). Seven species were initially
included in the genus based on multi-gene phylogenetic
analyses (Crous et al. 2013). However, they did not synonymize Pseudocercospora colombiensis and P. thailandica under Pallidocercospora, even though these two
species clustered with other Pallidocercospora species
(Crous et al. 2013). Subsequently, these two species have
been stated as Pallidocercospora colombiensis and
P. thailandica (Crous et al. 2013; Pérez et al. 2013;
Quaedvlieg et al. 2014), although, the species combinations
have not been formally established. Therefore the sexual
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20
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Fungal Diversity (2016) 80:1–270
Fungal Diversity (2016) 80:1–270
21
morph, Mycosphaerella thailandica is transferred to Pallidocercospora in this study and this is congruent with our
rDNA based phylogenies (Fig. 7).
b Fig. 8 Pallidocercospora thailandica (MFLU 11-0170, MFLU
11-0177). a Herbarium material with leaf spots. b Appearance of
ascomata on the host surface (a = from MFLU 11-0177, b = MFLU110170). c Close up of ascomata on the host (MFLU 11-0170).
d Section through the ascomata (MFLU 11-0170). e Section through
peridium (MFLU 11-0170). f–i Asci (MFLU 11-0170). j–n Ascospores (MFLU 11-0170). o–p Culture characteristics (MFLU
11-0170; o = from above, p = from below). Scale bars
d = 20 lm, e = 10 lm, f–i = 5 lm, j–n = 2 lm
Pallidocercospora thailandica (Crous et al.) Phookamsak,
Wulandari & K.D. Hyde, comb. nov.
= Mycosphaerella thailandica Crous et al., in Crous
et al., Stud. Mycol. 50(2): 465 (2004)
Elsinoe phaseoli CBS 150.27
Neocylindroseptoria
N
Ne
ocylindr
d oseeptoriaa pistaciae
pis
i taaciae CBS
S 471.69
Out group
Neophaeocryptopus
Doth
t ide
d a hippophaeos
hiipppoopphaaeos CBS
CBS 188.58
1888.58
Dothidea
Dothidea
Doth
t iddeea berberidis
berrberridi
dis C
CBS
BS 18
186.58
86.58
97
Dothidea
Doth
thiddea muelleri
muuelleeri C
CBS
BS 191
191.58
1.58
Plowrightia
Plowrigh
i htia ribesia
riibesia MF
MFLU
FLU 14-004
14-0040
40
79
Plowrightia
P
lowriightiaa ribesia MFLUCC
MFLUCC 13-0670
Plowrightia sensu stricto
Stylodothis
Stylodothis
Sty
t loddotthhiiss puccinioides
puuccinioiddes CBS
S 193.58
Dothidea
Dot
thid
idea sambuci
sam
mbuci DAOM
DAO
OM 231303
2313003
55
5
95
9
5
Dothidea
Doth
thide
d a insculpta
inscuulp
l ta CBS
S 189.58
Dothidea sensu stricto
Dothidea
Do
oth
t ide
dea insculpta
inssculp
l ta MFUCC
MF
FUCC 13-0686
Neophaeocryptopus
N
eop
o haeoocry
ryptop
o us cytisi
cytis
cy
i i MFLUCC
MFL
LUC
CC 14-0970
Neophaeocryptopus
Dothiora
Doth
t iora elli
elliptica
ipticaa CBS 736
736.71
6.71
62
6
2
Dothiora
Dothiora
Do
othiora bux
buxi
u i MFLU
MFLU 15-3
15-3404
3404
Dothiora
Doth
t ioraa cann
cannabinae
nabin
inae CBS 7737.71
37.71
50
Cylindroseptoria
Cy
C
ylindr
d osseepptooriaa ceratoniae
cerratonniae CBS
CB
BS 477.69
Cylindroseptoria
Coleophoma
C
Co
leop
ophomaa oleae
oleaae CBS 615.72
60
0
Cylindroseptoria
Coleophoma
C
Co
leop
ophomaa crateriformis
cratterifo
f rmi
r is CBS 473.69
Endoconidioma
E
ndo
d conniddiomaa po
populi
opuli U
UAMH
AMH
H 10902
Endoconidioma
Plowrightia
P
lowrrigh
i tia periclymeni
perricly
l meni 178096
69
Rhizosphaera
Rhi
izosp
s haera ka
kkalkhoffii
llkkhoffii
o ATCC
ATC
CC 26605
63
6
3
Phaeocryptopus
Plow
wrigh
i htia abietis
is ATCC 24339
244339
Plowrightia
Rhizosphaera
Rhiz
izosp
s haera oudemansii
oude
demansii 184813
1844813
75
7
5
Phaeocryptopus
Pha
haeoccry
ryptop
o us nnudus
udus CBS
S 268.37
96
Rhiz
i ossphaeeraa pin
ni 64367
6436
367
Rhizosphaera
pini
Delphinella
Delp
l hinnellla
l sstrobiligena
t obilig
tr
igena
n CB
CBS
BS 735.71
59
Delphinella
CBS
116.29
SSydowia
ydow
wiaa ppolyspora
oly
lyspoora C
BS 116
6.229
0.1
Fig. 9 RAxML maximum likelihood phylogenetic tree based on a
LSU and ITS sequence data from species of Dothideaceae. Maximum
likelihood bootstrap support values greater than 50 % are shown in
above. The new isolates are in blue and other extype strains in bold.
The tree is rooted with Elsino/ phaseoli
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Fungal Diversity (2016) 80:1–270
Fig. 10 Dothiora buxi (holotype). a, b View of ascostromata on host surface. c Section through the ascostroma. d Peridium. e–g Asci. h–
k Ascospores. Scale bars c = 100 lm, d = 50 lm, e–g = 30 lm, h = 20 lm, i–k = 10 lm
: Pseudocercospora thailandica Crous et al., in Crous
et al., Stud. Mycol. 50(2): 465 (2004)
Index Fungorum number: IF552204; Facesoffungi
number: FoF02258, Fig. 8
Biotrophic, hemibiotrophic, or saprotrophic on various
hosts, leaf spots on the margins of leaves, causing tip
123
blight, or lesions or lesions initially start from the tip of
leaves, irregular in shape, dried, pale brown to brown at the
middle, and reddish-brown to dark brown at margin of the
lesions. Sexual morph Ascomata 45–70 lm high, 45–
80 lm diam., as small black dots on the host surface,
scattered, sometimes clustered, gregarious, immersed to
Fungal Diversity (2016) 80:1–270
89
62
71
23
Hysterium hyalinum CBS 237.34
100 Hysterographium fraxini CBS 109.43
Hysterographium fraxini CBS 242.34
Rhytidhysteron opuntiae GKM 1190
Hysterodifractum partisporum CCMB 252/2012
Rhytidhysteron rufulum MFLUCC 12-0013
100
Rhytidhysteron rufulum EB 0383
Rhytidhysteron rufulum EB 0382
Rhytidhysteron rufulum EB 0384
Rhytidhysteron neorufulum HUEFS 192194
Rhytidhysteron_neorufulum GKM 361A
Rhytidhysteron neorufulum MFLUCC 12-0529
Rhytidhysteron neorufulum EB 0381
80
Rhytidhysteron neorufulum MFLUCC 12-0567
Rhytidhysteron neorufulum MFLUCC 12-0011
Rhytidhysteron neorufulum MFLUCC 12-0569
Rhytidhysteron neorufulum MFLUCC 12-0528
Rhytidhysteron neorufulum CBS 306.38
72
Rhytidhysteron hysterinum EB 0351
Rhytidhysteron thailandicum MFLUCC 12-0530
99
_Gloniopsis subrugosa GKM 1214
Gloniopsis subrugosa SMH 557
Gloniopsis arciformis GKM L166A
Gloniopsis praelonga CBS 123337
100
95 Gloniopsis praelonga CMW 19983
Gloniopsis praelonga CBS 112415
Gloniopsis calami MFLUCC 15–0739
Gloniopsis subrugosa GKM 1010
Gloniopsis subrugosa CBS 123346
Graphyllium caracolinensis HUEFS 42838
84
Oedohysterium sinense EB 0339
79 74
Oedohysterium insidens CBS 238.34
100
Oedohysterium insidens ANM 1443
Oedohysterium sinense CBS 123345
Hysterobrevium mori GKM 1013
Ostreichnion sassafras CBS 322.34
Hysterobrevium constrictum SMH 5211.1
97
Hysterobrevium constrictum GKM 426N
Hysterobrevium smilacis CMW 18053
Hysterobrevium mori CBS 123564
Hysterobrevium mori CBS 123336
93
89
Hysterobrevium mori CBS 123563
Hysterobrevium mori CBS 123335
Hysterobrevium mori SMH 5273
96 Hysterobrevium smilacis CBS 200.34
Hysterobrevium smilacis SMH 5280
Hysterobrevium smilacis CBS 114601
Hysterobrevium mori SMH 5286
Hysterographium flexuosum GKM 1262c
78
100
Ostreichnion centramurum MFLUCC 12-0802
99
Ostreichnion curtisii CBS 198.34
Hysterium barrianum ANM 1495
100
Hysterium barrianum ANM 1442
Hysterium vermiforme GKM 1234
85
100 Hysterium pulicare ANM 1455
Hysterium pulicare ANM 85
99
Hysterium angustatum CMW 20409
70
Hysterium angustatum GKM5211
Hysterium angustatum SMH 5216
Hysterium angustatum MFLUCC 15-0950
Hysterium angustatum GKM 243A
Hysterium angustatum CBS 236.34
100
Hysterium angustatum CBS 123334
Psiloglonium clavisporum CBS 123338
Psiloglonium clavisporum CBS 123341
100
Psiloglonium clavisporum CBS 123340
Psiloglonium clavisporum GKM 344A
Psiloglonium clavisporum GKM L172A
Psiloglonium araucanum CMW 17941
99
Psiloglonium araucanum CMW 18760
94
87
Psiloglonium araucanum CBS 112412
100 Psilogloniu simulans CBS 206.34
Psiloglonium simulans ANM 1557
Delitschia winteri CBS 225.62
0.02
Fig. 11 Phylogram generated from maximum parsimony analysis
based on LSU sequence data from Hysteriaceae. Maximum likelihood bootstrap support values greater than 50 % is shown above
nodes. The tree is rooted with Delitschia winteri. The new isolates are
in blue and other ex-type strains are in bold
semi-immersed, with protruding papilla, globose to subglobose, glabrous, ostiole central, with minute papilla. Peridium
5–10 lm wide, thin-walled, composed of 2–3 cell layers of
brown to dark brown, pseudoparenchymatous cells, arranged
in a textura angularis. Hamathecium lacking pseudoparaphyses. Asci (23–)25–35(–37.5) 9 7–9 lm (
x = 29 9
7.9 lm, n = 30), 8-spored, bitunicate, fissitunicate, obclavate, rarely ovoid, subsessile, apically rounded, with welldeveloped ocular chamber. Ascospores (8–)9–12 9 2.5–
3.5 lm (
x = 10.8 9 3.1 lm, n = 30), overlapping uni- to
tri-seriate, clavate, hyaline to subhyaline, 1-septate, not
contricted at the septum, smooth-walled, upper cell wider
and shorter than lower cell. Asexual morph Hyphomycetous, pseudocercospora-like (see notes).
Culture characteristics: Colonies on PDA reaching
23.5–30 mm diam. after 4 weeks at 25–30 °C; colony from
above, dark greenish at the margin, paler greenish hair-like
at the center; from below, dark greenish to black; dense,
irregular, flattened to raised, with undulate edge, with
entire margin, surface smooth, velvety to cottony; not
producing pigmentation in agar.
Material examined: THAILAND, Phrae, Rongkwang
District, Maejo University Phrae campus grounds, on dead
leaves of Dracaena loureiri Gagnep (Ruscaceae), 20
August 2010, R. Phookamsak, RP0050 (MFLU 11-0170),
living culture, MFLUCC 11-0134, KUMCC; ibid. Chiang
Rai, Muang District, Mae Fah Luang University campus
grounds, on living leaves of Rhapis sp. (Arecaceae), 4
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24
Fig. 12 Gloniopsis calami (holotype). a Appearance of hysterothecia on host. b Close up hysterothecia. c Section of hysteriothecium
d Pseudoparaphyses. e–h. Asci i–p. Ascospores. q Geminated
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ascospore. r Culture characters on MEA. Scale bars a = 500 lm,
b = 200 lm, c, e–h = 50 lm, d = 10 lm, i–q = 5 lm
Fungal Diversity (2016) 80:1–270
Fig. 13 Phylogenetic tree generated from maximum parsimony (MP)
analysis based on combined ITS and LSU sequence data of genera of
the family Dictyosporiaceae. Bootstrap support values for maximum
parsimony (MP) and maximum likelihood (ML) greater than 50 %
25
and Bayesian posterior probabilities greater than 0.80 are indicated
above or below the nodes as MPBS/MLBS/PP. The ex-type strains
are in bold and the new isolates are in red bold. The tree is rooted with
Letendraea helminthicola
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26
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b Fig. 14 Pseudocoleophoma typhicola (holotype). a Appearance of
conidiomata on host substrate. b Close up of conidioma. c Vertical
section through conidiomata .d Conidia attached to conidiogenous
cells. e–l Immature to mature conidia. m Germinated conidium. Scale
bars a = 1 mm b = 100 lm, c = 50 lm, d–l = 5 lm, m = 10 lm
August 2010, N.F. Wulandari, RP0057 (MFLU 11-0177),
living culture, MFLUCC 11-0141, KUMCC.
Notes: Pallidocercospora thailandica was introduced as
Mycosphaerella thailandica by Crous et al. (2004b) who
Fig. 15 Pseudocoleophoma typhicola (ex-type culture). a, b Culture on PDA (note b reverse). c Conidioma on PDA. d Vertical
section through conidioma. e Peridium. f Conidia attached to
27
noted its asexual morph as Pseudocercospora thailandica
Crous et al. (2004b). The asexual morph was described
with ‘‘mycelium composed of medium brown, branched,
septate, and smooth hyphae; conidiophores dense, pale
brown, subcylindrical, unbranched, 0–2-septate, straight to
curved, smooth-walled, arising from the upper cells of the
stroma; conidiogenous cells terminal, pale brown, subcylindrical, tapering to flat tipped apical loci, proliferating
sympodially; conidia solitary, pale brown, narrowly
obclavate to subcylindrical, subobtuse at the apex, with
conidiogenous cells. g–k Mature and immature conidia. Scale bars
d = 1 mm e = 50 lm, f–k = 20 lm
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Fungal Diversity (2016) 80:1–270
Fig. 16 Pseudodictyosporium thailandica (holotype). a Host (decaying bamboo). b, c Conidiophores on the host surface. d, e Conidiophores
and conidia. f Conidiogenous cells and conidia. g–j Conidia. Scale bars b = 200 lm, c = 100 lm, d–e, k = 20 lm, f–j = 10 lm
long obconically subtruncate at the base, 3–6-septate,
smooth-walled, and guttulate’’ (Crous et al. 2004b).
Based on phylogenetic analysis, Mycosphaerella thailandica clustered with Pallidocercospora species (Crous
et al. 2013; Quaedvlieg et al. 2014). However, Crous et al.
123
(2013) mentioned that Pseudocercospora colombiensis and
Ps. thailandica were typical members of Pseudocercospora sensu stricto based on its morphological features.
Therefore, these taxa were not synonymized under the
genus Pallidocercospora when Crous et al. (2013)
Fungal Diversity (2016) 80:1–270
29
Stagonosporopsis chrysanthemi CBS 137.96
95/1
Stagonosporopsis chrysanthemi CBS 500.63
97/-
Stagonosporopsis inoxydabilis CBS 425.90
100/1
Stagonosporopsis tanaceti CBS 131484
Stagonosporopsis rudbeckiae CBS 109180
91/0.99
Stagonosporopsis artemisiicola CBS 102636
97/1
94/0.97
97/1
Stagonosporopsis trachelii CBS 379.91
Stagonosporopsis centaureae MFLUCC 16-0787
Stagonosporopsis oculihominis CBS 634.92
66/1 100/1
Stagonosporopsis actaeae CBS 114303
Stagonosporopsis actaeae CBS 106.96
99/1
Stagonosporopsis dennisii CBS 631.68
100/1
Stagonosporopsis cucurbitacearum CBS 133.96
Stagonosporopsis
Stagonosporopsis caricae CBS 248.90
Stagonosporopsis helianthi CBS 200.87
100/1
Stagonosporopsis heliopsidis CBS 109182
100/1
Stagonosporopsis andigena CBS 269.80
Stagonosporopsis andigena CBS 101.80
100/1
Stagonosporopsis crystalliniformis CBS 713.85
99/0.98
100/1
Stagonosporopsis hortensis CBS 104.42
Stagonosporopsis hortensis CBS 572.85
93/1
Stagonosporopsis astragali CBS 178.25
96/0.98
Stagonosporopsis lupini CBS 101494
52/0.96
Stagonosporopsis ajacis CBS 177.93
92/0.98
Stagonosporopsis valerianellae CBS 329.67
Stagonosporopsis dorenboschii CBS 426.90
98/1
Stagonosporopsis loticola CBS 562.81
Allophoma zantedeschiae CBS 131.93
100/1
97/-
Allophoma zantedeschiae CBS 229.32
100/1
Allophoma
Allophoma labilis CBS 124.93
Heterophoma sylvatica CBS 874.97
83/0.99
57/-
Heterophoma nobilis CBS 507.91
84/1
Heterophoma
Heterophoma adonidis CBS 114309
76/0.97
Neodidymella thailandicum MFLUCC 11-0140
100/1
Epicoccum nigrum CBS 173.73
95/1
65/0.96
Neodidymella
Epicoccum nigrum CBS 125.82
Epicoccum
Epicoccum pimprinum CBS 246.60
100/1
Didymella acetosellae CBS 179.97
Didymella rumicicola CBS 683.79
98/1
100/1
Didymella exigua CBS 183.55
Didymella
Didymella calidophila CBS 448.83
Macroventuria anomochaeta CBS 502.72
61/-
93/1
Macroventuria anomochaeta CBS 525.71
Macroventuria
Macroventuria wentii CBS 526.71
99/0.98
Paraboeremia adianticola CBS 187.83
Paraboeremia adianticola CBS 260.92
96/0.98
Paraboeremia putaminum CBS 130.69
Paraboeremia
Phoma selaginellicola CBS 122.93
Fig. 17 Phylogenetic tree inferred from a maximum Likelihood
analysis based on a concatenated alignment of LSU, ITS, RPB2,
TUB2 sequence data representing Didymellaceae and allied families.
The RAxML bootstrap support values (MLBS) greater than 50 % and
Bayesian posterior probabilities (BPP) greater than 0.95 are given at
the nodes (MLBS/BPP). The ex-type strains are in bold and the new
isolates are in blue. The tree is rooted with Leptosphaeria doliolum
introduced this genus. However, the name ‘‘Pallidocercospora thailandica’’ has been used instead of ‘‘Pseudocercospora thailandica’’ (Crous et al. 2013; Quaedvlieg
et al. 2014), but the name ‘‘Pallidocercospora thailandica’’
was not formally synonymized and thus, this name was
invalid.
In this study, two isolates were collected from dead
leaves of Dracaena loureiri (Ruscaceae) and living leaves
of Rhapis sp (Arecaceae). Combined ITS and LSU phylogenetic analyses show that these isolates grouped with
Pseudocercospora colombiensis and Ps. thailandica, and
clustered with Pallidocercospora species, with high bootstrap support (97 % ML, 100 % MP, 1.00 PP, Fig. 7).
Furthermore, Pseudocercospora colombiensis and Ps.
thailandica form a distinct clade with Pseudocercospora
sensu stricto which is congruent to Crous et al. (2013) and
Quaedvlieg et al. (2014). Therefore, we transfer the species
Mycosphaerella
thailandica
to
the
genus
Pallidocercospora.
Pallidocercospora thailandica is morphologically distinct from P. colombiensis, but it is difficult to distinguish
these two species based on phylogenetic analyses
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Neomicrosphaeropsis italica MFLUCC 15–0484
Neomicrosphaeropsis italica MFLUCC 15–0485
83/1
Neomicrosphaeropsis italica MFLUCC 16–0284
Neomicrosphaeropsis italica MFLUCC 15–0487
51/-
Neomicrosphaeropsis tamaricicola MFLUCC 14–0443
Neomicrosphaeropsis tamaricicola MFLUCC 14-0602
Neomicrosphaeropsis
Neomicrosphaeropsis tamaricicola MFLUCC 14–0439
77/1
Neomicrosphaeropsis novorossica MFLUCC 14–0578
Neomicrosphaeropsis rossica MFLUCC 14–0586
74/0.95
87/1
Neomicrosphaeropsis minima MFLUCC 13–0394
Neomicrosphaeropsis cytisi MFLUCC 13–0396
98/1
Neomicrosphaeropsis cytisinus MFLUCC 16-0790
Ascochyta fabae CBS 649.71
77/1
Ascochyta fabae CBS 524.77
92/0.99
Ascochyta
Ascochyta viciae CBS 451.68
97/1
Ascochyta pisi CBS 122785
56/0.95
99/0.98
Ascochyta pisi CBS 126.54
Phomatodes aubrietiae CBS 383.67
100/1
Phomatodes
Phomatodes aubrietiae CBS 627.97
Calophoma vodakii CBS 173.53
100/1
Calophoma
Coniothyrium clematidisrectae CBS 507.63
100/1
Coniothyrium clematidina CBS 108.79
75/-
Phoma herbarum CBS 615.75
Phoma herbarum CBS 274.37
Phoma /Platychora
Platychora ulmi MFLUCC 14–1189
53/-
Platychora ulmi CBS 361.52
62/-
Phoma herbarum UAMH 10909
Phoma dutchum CBS 134.96
100/1
90/-
Neodidymelliopsis xanthina CBS 383.68
Neodidymelliopsis xanthina CBS 168.70
91/100/1
Neodidymelliopsis sp. 1 CBS 256.77
Neodidymelliopsis polemonii CBS 109181
98/1
Neodidymelliopsis
Neodidymelliopsis ranunculi MFLUCC 13-0490
94/1
Neodidymelliopsis sp. 2 CBS 382.96
Neodidymelliopsis urticicola CBS 121.75
72/-
100/1
Xenodidymella applanata CBS 115577
Xenodidymella applanata CBS 195.36
99/1
96/1
Xenodidymella
Xenodidymella catariae CBS 102635
Xenodidymella asphodeli CBS 375.62
100/0.95
Neoascochyta europaea CBS 820.84
100/1
Neoascochyta graminicola CBS 301.69
Neoascochyta
Neoascochyta exitialis CBS 118.40
Leptosphaerulina australis CBS 317.83
97/100/1
Leptosphaerulina trifolii CBS 235.58
Leptosphaerulina
Leptosphaerulina arachidicola CBS 275.59
Nothophoma infossa CBS 123395
100/1
Nothophoma arachidishypogaeae CBS 125.93
89/1
Nothophoma
Nothophoma anigozanthi CBS 381.91
96/0.99
Microsphaeropsis olivacea CBS 432.71
Microsphaeropsis olivacea CBS 233.77
70/-
Microsphaeropsidaceae
Microsphaeropsis proteae CPC 1425
Leptosphaeria doliolum CBS 505.75
Leptosphaeriaceae
0.07
Fig. 17 continued
(Quaedvlieg et al. 2014). However, Quaedvlieg et al.
(2014) applied the pairwise homoplasy index (PHI) test
with the GCPSR and CSC concepts and proposed that these
two species are different taxa.
Pallidocercospora thailandica has been collected from
various hosts in Australia, Laos, Thailand and West Indies
(Acacia mangium Willd., Eucalyptus camaldulensis
Dehnh., Musa sp.) and is mostly found as a pathogen on the
hosts (Crous et al. 2004b, 2013; Arzanlou et al. 2008;
Cheewangkoon et al. 2008). In this study, this species was
associated with leaf spots on Rhapis sp., and as a saprobe
on Dracaena loureiri; both are new hosts for
P. thailandica.
Dothideales Lindau
For Dothideales, we follow Li et al. (2016).
123
Dothideaceae Chevall.
The family Dothideaceae was introduced by Chevallier
(1826) as ‘Dothideae’, and later Fuckel (1869) established
this family with Dothidea as the type genus and D. gibberulosa as the type species. Thambugala et al. (2014)
treated the family Dothideaceae with 15 genera. Dothideaceae is characterized by ‘immersed to erumpent or
superficial, uni or multi-loculate ascostromata, 8- or
polyspored, bitunicate asci and hyaline or brown, transversely septate, sometimes muriform ascospores’ (Thambugala et al. 2014). We provide an updated phylogeny in
Fig. 9.
Dothiora buxi Jayasiri, Camporesi & K.D. Hyde, sp. nov.
Index Fungorum number: IF552173; Facesoffungi
number: FoF02223, Fig. 10
Fungal Diversity (2016) 80:1–270
Fig. 18 Neomicrosphaeropsis cytisi (holotype). a Herbarium specimen. b, c Appearance of brown coniodiomata on the host. d Wall of
conidiomata. e Vertical section of conidiomata. f–h Conidiophores,
31
conidiogenous cells and developing conidia. j Germinating spore. i, k,
l Conidia. m Culture. Scale bars b = 500 lm, c = 200 lm, d,
j = 10 lm, e = 20 lm, f–h, i, k–n = 5 lm, n = 25 mm
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32
Etymology: The specific epithet buxi is based on the host
genus from which the taxon was collected.
Holotype: MFLU 15-3404.
Saprobic on Buxus sempervirens L. Sexual morph
Ascostromata 500–1000 lm long 9 220–250 lm high,
320–340 lm diam., erumpent through the epidermis, solitary or clustered, globose, brown to black, with single
locules, with a central longitudinal slit-like opening.
Peridium 32–83 lm wide, two-layered, outer layer composed of dark brown or brown, thick-walled cells of textura
angularis, inner layer composed of hyaline, thin-walled
cells of textura angularis. Hamathecium lacking pseudoparaphyses. Asci 100–115 9 14–21 lm (
x = 102 9
17 lm, n = 20), 32-spored, bitunicate, fissitunicate,
cylindro-clavate, short pedicellate, apically rounded, with a
small ocular chamber. Ascospores 11–15 9 5.4–7 lm
(
x = 13 9 6 lm, n = 30), bi-seriate to multi-seriate,
hyaline to very pale brown, aseptate, fusoid to ovoid, one
end narrower than other, smooth-walled with granular
contents, with a thin mucilaginous sheath.
Material examined: ITALY, Province of Forlı̀-Cesena
[FC]), near Passo delle Forche—Galeata on dead branch of
Buxus sempervirens (Buxaceae), 17 November 2014, E.
Camporesi, IT 2284 (MFLU 15-3404, holotype, KUN,
isotype).
Notes: Dothiora was introduced by Fries (1849) with
D. pyrenophora (Fr.) Fr. as the type species. Our isolate
shares common characters with the genus Dothiora. DNA
was extracted from fruiting bodies and multi-gene phylogenetic analysis placed Dothiora buxi as a sister taxon to
Fig. 19 Neomicrosphaeropsis cytisinus (holotype). a Appearance of
conidiomata on host. b Close-up of conidiomata. c Section of
conidioma. d Section of peridium. e Conidiogenous cells. f–i Conidia.
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Fig. 20 Neomicrosphaeropsis minima (holotype). a Herbarium c
specimen. b, c Appearance of black coniodiomata on the host. d, e
Vertical section of conidiomata. f Wall of conidiomata. g–k Conidiophores, conidiogenous cells and developing conidia. l Germinating
spore. m, n Conidia. o Culture. Scale bars b, c = 200 lm, d,
e = 50 lm, f = 15 lm, g–n = 5 lm, o = 50 mm
Dothiora elliptica (Fig. 9). The latter is similar in having
epidermal erumpent, hysteriiforme ascostromata and hyaline ascospores but differs from Dothiora buxi in having
32-spored asci and aseptate, fusoid to ovoid, ascospores
narrowed at one end (Saccardo 1889). Therefore, we
introduce Dothiora buxi as a new species.
Hysteriales Lindau
For Hysteriales, we follow Hyde et al. (2013).
Hysteriaceae Chevall.
Chevallier (1826) introduced the family Hysteriaceae as
‘Hysterineae’ and this family has been treated with different genera by various authors (Zogg 1962; von Arx and
Müller 1975; Kirk et al. 2001; Lumbsch and Huhndorf
2010). Recent multi-gene phylogenetic studies placed
Hysteriaceae in Hysteriales, Pleosporomycetidae (Boehm
et al. 2009a, b; Hyde et al. 2013; Wijayawardene et al.
2014a, Thambugala et al. 2016b). Hyde et al. (2013) and
Wijayawardene et al. (2014a) accepted 13 genera, while de
Almeida et al. (2014) introduced a new genus Hysterodifractum in this family. The family now contains 14 genera.
A phylogenetic tree for the family is presented in Thambugala et al. (2016b) (Fig. 11).
j. Germinated conidia. l Colony from above. m Colony from below.
Scale bars c = 50 lm, d = 20 lm, e = 5 lm, f–i = 2 lm,
j = 10 lm
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33
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34
Gloniopsis De Not.
The genus Gloniopsis is typified by Gloniopsis praelonga (Schwein.) Underw. & Earle [as ‘praelongum’].
Based on morphology and molecular phylogenetic analyses
Gloniopsis is placed in Hysteriaceae (Boehm et al. 2009b;
Wijayawardene et al. 2014a; Thambugala et al. 2016b).
The genus now contains 65 epithets listed in Index Fungorum (2016). The genus Gloniopsis is characterized by
hyaline to yellow dictyospores, often inequilateral, curved,
multi-septate, with one or more longitudinal septa, constricted at the first-formed septum, sometimes constricted
at additional septa, and usually surrounded by a gelatinous
sheath which may dissipate with age (Zogg 1962).
Gloniopsis calami Konta & K.D. Hyde. sp. nov.
Index Fungorum number: IF552234; Facesoffungi
number: FoF02366, Fig. 12
Etymology: Name reflects the host genus Calamus.
Holotype: MFLU 15-1470.
Saprobic on dead Calamus sp. Sexual morph Hysterothecia 195–215 lm high 9 160–170 lm wide, erumpent
to superficial, solitary to gregarious, scattered, dark, straight to
flexuous. Peridium 37–47 lm wide, carbonaceous, thickwalled, not having distinct layers, relatively smooth on the
outer surface. Hamathecium 1.2–2 lm wide, composed of
dense, branched, hyaline, septate, pseudoparaphyses. Asci 60–
80 9 15–21 lm (
x = 71 9 17 lm, n = 10), 8-spored,
bitunicate, fissitunicate, cylindrical to cylindric-clavate, short
pedicellate, with knob-like pedicel, apically rounded, with a
well-developed ocular chamber, external layer easily broken.
Ascospores 17–20 9 6–8 lm (
x = 19 9 7 lm, n = 10),
dictyosporous, overlapping 1–2-seriate, fusiform, slightly
curved to straight, 4–6-trans-septate and with 2–4 vertical
septa, reddish-brown to brown, constricted at the septa,
smooth-walled. Asexual morph Undetermined.
Culture characteristics: Ascospores germinated on
MEA within 24 h and germ tubes produced from all cells.
Colonies on MEA 7–7.5 cm diam. after 2 weeks at 25 °C,
grey to dark green, outwardly with strongly radiating colony. After 1 month of incubation, colonies irregular, convex, spongy, medium dense, margin undulate.
Material examined: THAILAND, Phang-Nga Province,
on dead Calamus sp. (Arecaceae), 6 December 2014, S.
Konta, DNH07i (MFLU 15-1470, holotype, isotype HKAS
95030); ex-type living culture, MFLUCC 15-0739.
Notes: Molecular analyses indicate that the new species
belongs to the genus Gloniopsis in the Hysteriaceae clade
and is nested in between G. praelonga (type) and G. subrugosa and appear to be phylogenetically distinct (Fig. 11).
The ascospores of G. calami are similar in shape, but
smaller in size and have a different septation to G. praelonga (type) and G. subrugosa. However, G. calami is
distinct from G. arciformis in its ascospores being
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constricted at the septa, while in G. arciformis they are not
constricted.
Pleosporales Luttr. ex M.E. Barr
We follow Tanaka et al. (2015).
Dictyosporiaceae Boonmee & K.D. Hyde
The family Dictyosporiaceae was introduced by Boonmee et al. (2016) to accommodate Aquaticheirospora,
Cheirosporium, Dictyocheirospora, Dictyopalmispora,
Dictyosporium, Digitodesmium, Pseudocoleophoma and
Pseudodictyosporium. The asexual morphs of the family
Dictyosporiaceae are hyphomycetous with brown, multiseptate, cheirosporous conidia (Boonmee et al. 2016). In
this study we provide an updated backbone tree for Dictyosporiaceae (Fig. 13) and introduce the new species
Pseudodictyosporium thailandica and Pseudocoleophoma
typhicola.
Pseudocoleophoma typhicola E.B.G. Jones, Kamolhan,
Boonmee & K.D. Hyde, sp. nov.
Index Fungorum number: IF552326; Facesoffungi
number: FoF02444, Figs. 14 and 15
Etymology: Referring to the host plant Typha latifolia.
Holotype: MFLU 16-0966.
Saprobic on submerged stems in freshwater. Sexual
morph Undetermined. Asexual morph Conidiomata
forming as dark spots on the host surface, 140–150 lm
high 9 60–100 lm diam. (
x = 143 9 80 lm, n = 15),
semi-erumpent in the host tissue, uniloculate solitary to
scattered, subglobose, brown to black. Peridium 40–45 lm
at base, 40–45 lm at sides, comprising 4–5 layers, hyaline
to dark brown, thick-walled cells of textura angularis.
Conidiophores reduced to conidiogenous cells. Conidiogenous cells 2–5 9 2–5 lm (
x = 2 9 3 lm, n = 15),
enteroblastic, smooth-walled, hyaline. Conidia 9–11 9 2–
3 lm (
x = 10 9 3 lm, n = 20), hyaline, oblong to
cylindrical, with rounded or obtuse ends, 1-euseptate,
smooth, thin-walled, guttulate.
Culture characteristics: Colonies on PDA 3 cm diam.
after 4 weeks at 16 °C, dirty white to pale brown at the
margin, pale brown to creamy at the center; reverse iron,
thin, curled, flat.
Material examined: UK, Hampshire, Swanick Lakes, on
submerged stems of Typha latifolia (Typhaceae) in freshwater, 28 August 2015, E.B.G. Jones, GJ190 (MFLU
16–0966, holotype, HKAS94520 isotype), ex-type living
culture, MFLUCC 16–0123, KUMCC 16-0007.
Notes: Pseudocoleophoma typhicola is different from
P. polygonicola and P. calamagrostidis in having conidiomata without neck and with a wide peridium (40–45 lm
at base, 40–45 lm at sides), enteroblastic, 3-septate conidia
while P. polygonicola and P. calamagrostidis have
Fungal Diversity (2016) 80:1–270
35
Fig. 21 Neodidymelliopsis ranunculi (holotype). a Herbarium specimen. b, c Appearance of black coniodiomata on the host. d Vertical
section of conidioma. e, f Vertical section of peridium. g–j Conidiogenous cells and developing conidia. k Germinating conidium. l–
n Conidia. o Culture on PDA. Scale bars b = 200 lm, c–
d = 100 lm, e = 50 lm, f = 20 lm, g–i = 10 lm. k–p = 5 lm,
q, r = 20 mm
conidiomata with a long neck, phialidic, aseptate conidia.
Phylogenetic analyses indicate that Pseudocoleophoma
typhicola is related to P. polygonicola and
P. calamagrostidis with high support, but it can be recognized as a new species as it stands on its own with relatively good support (Fig. 13).
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b Fig. 22 Platychora ulmi (MFLU 16-1972). a, b Ascostromata on
substrate. c, d Cross section of ascostromata. e Peridium f, i. Asci j–
l. Ascospores. m Conidiomata on MEA. o Cross section of
conidiomata. n, p, q Conidia attached to possible conidiogenous
cell. r Conidia. Scale bars a = 500 lm, b, c, m, o = 100 lm,
d = 50 lm, e = 20 lm, f–i = 10 lm, j–l, n, p–r = 5 lm
Pseudodictyosporium Matsush.
The genus Pseudodictyosporium was established by
Kobayasi (1971) and is typified by P. wauense Matsush.
Three species are accepted in this genus, P. wauense,
P. elegans and P. indicum (Boonmee et al. 2016). Molecular phylogenetic analyses place Pseudodictyosporium
within the family Dictyosporiaceae (Pleosporales) (Tanaka
et al. 2015; Boonmee et al. 2016). In this study we introduce P. thailandica as a new species.
Pseudodictyosporium thailandica C.G. Lin, Yong Wang
bis & K.D. Hyde, sp. nov.
Index Fungorum number IF552165; Facesoffungi number: FoF02227, Fig. 16
Etymology: Referring to the country where the fungus
was first collected.
Holotype: MFLU 16-1301.
Saprobic on decaying bamboo stem. Sexual morph
Undetermined. Asexual morph Conidiophores macronematous, mononematous, scattered or caespitose, erect,
flexuous, irregularly branched, smooth, septate, slightly
constricted at septa, hyaline to brown, often geniculate, 23–
305 lm long (
x = 77 lm, n = 27), 2.5–7.9 lm wide
(
x = 5 lm, n = 96). Conidiogenous cells holoblastic,
polyblastic, discrete, determinate or sympodial, terminal
and intercalary. Conidia solitary, acropleurogenous, dry,
cheiroid, ellipsoidal, ovoid, smooth, multi-septate, subhyaline to grey-brown, 14–31 lm long (
x = 20 lm,
n = 70), 12–26.5 lm wide (
x = 17 lm, n = 70) at the
widest point.
Culture characteristics: Conidia germinating on PDA
within 36 h. Colonies on MEA reaching 20–35 mm diam.
after 4 months at room temperature (25 °C), effuse, hairy,
grey-white from above, brown at the center, yellowishwhite at margin from below.
Material examined: THAILAND, Phetchaburi, Cha-am
District, on decaying bamboo stem, 28 July 2015, ChuanGen Lin, KNP 5-3 (MFLU 16-1301, holotype; HKAS
95053, isotype), ex-type living culture, MFLUCC
16-0029.
Notes: Phylogenetic analysis of combined ITS, LSU,
SSU and TEF sequence data indicate that our species
belongs in the genus Pseudodictyosporium (Fig. 13) with
100 % MP bootstrap support, 100 % ML bootstrap support
and 100 % Bayesian posterior probabilities, and forms a
separate clade within Pseudodictyosporium.
37
The conidiophores of P. thailandica are longer than those
of P. wauense (up to 100 lm) and P. elegans (10–56 lm), but
shorter than those of P. indicum (284–630 lm). In addition,
the conidia of our species (12–26.5 lm) are wider than earlier
described species (P. elegans 9–16.5 lm, P. indicum
12.5–16 lm and P. wauense 12–19 lm) (Rao and Subhedar
1976; Tzean and Chen 1990; Kirschner et al. 2013).
Didymellaceae Gruyter et al.
The family Didymellaceae was introduced by De
Gruyter et al. (2009), with type species Didymella exigua
(Niessl) Sacc., to accommodate most species in Phoma
sensu lato and allied genera. The family contains numerous
plants pathogenic, saprobic and endophytic species associated with a wide range of hosts (Aveskamp et al. 2010;
Chen et al. 2015a). Aveskamp et al. (2010) revised the
taxonomy of Didymellaceae based on multi-gene analyses
and included eleven genera in the family, i.e. Ascochyta,
Boeremia, Chaetasbolisia, Didymella, Epicoccum, Leptosphaerulina, Macroventuria, Microsphaeropsis, Peyronellaea, Phoma and Stagonosporopsis. Subsequently,
more genera and information were added (Wijayawardene
et al. 2012; Zhang et al. 2012a; Hyde et al. 2013; Ariyawansa et al. 2015a). Chen et al. (2015a) utilized the
RPB2 gene combined with ITS, LSU as well as tub2 to
distinguish Phoma and related genera and accepted 17
genera in Didymellaceae. However Microsphaeropsis
was excluded from the Didymellaceae, and a new family
Microsphaeropsidaceae was proposed to accommodate
these Microsphaeropsis species. Thambugala et al.
(2016a) included an additional genus Neomicrosphaeropsis due to its morphological similarity with
Microsphaeropsis species, but this is phylogenetically
closely related to Didymellaceae. In this study, three new
species are added in Neomicrosphaeropsis based on both
morphology and phylogeny. Moreover, a new collection
of Platychora ulmi (J. Schröt.) Petr. MFLUCC 14-1189
together with another strain CBS 361.52 from GenBank
clustered with the Phoma group (Fig. 17), but LSU gene
data are only available for these two strains. LSU and
SSU sequence data does not provide sufficient phylogenetic information to distinguish closely related genera or
species (Aveskamp et al. 2009, 2010; Chen et al. 2015a).
Thus, the genus Platychora should be re-evaluated based
on additional genes. Didymellocamarasporium was
introduced in Didymellaceae based on LSU and SSU
sequence data by Wijayawardene et al. (2016), but they
cannot be well separated from other genera in present
study (data not shown). Hence, this genus also needs
additional genes to confirm its placement in Didymellaceae. To date, 21 genera are included in
Didymellaceae.
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Fig. 23 Stagonosporopsis centaureae (holotype). a Appearance of
ascomata on host. b Close-up of ascoma. c Section of ascoma.
d Section of peridium. e Pseudoparaphyses. f–i Asci. j–l Ascospores.
m Germinated ascospore. n Colony from above. o Colony from
below. Scale bars c = 20 lm, d, e = 10 lm, f–i = 20 lm, j–
m = 5 lm
Neomicrosphaeropsis Thambugala et al.
Thambugala et al. (2016a) included Neomicrosphaeropsis in Didymellaceae. In this study, three new
species are added in Neomicrosphaeropsis based on both
morphology and phylogeny.
Etymology: Named after the host genus Cytisus.
Holotype: MFLU 16-1871
Saprobic on dead stem of Cytisus sp. (Fabaceae),
forming numerous, conspicuous, oval, dark brown, conidiomata. Sexual morph Undetermined. Asexual morph
Coelomycetous. Conidiomata 75–155 lm diam. 9 75–
130 lm high, dark brown, solitary to gregarious or confluent, pycnidial, globose to subglobose, immersed,
unilocular, thick-walled, smooth, ostiolate. Ostiole single,
short, with acute apex, centrally located. Wall of
Neomicrosphaeropsis cytisi W.J. Li, Camporesi & K.D.
Hyde, sp. nov.
Index Fungorum number: IF552212; Facesoffungi
number: FoF02347, Fig. 18
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conidiomata 10–24 lm wide, composed of thick-walled,
brown to hyaline cells of textura angularis. Conidiophores
reduced to conidiogenous cells. Conidiogenous cells 2–
4 lm long 9 3.5–6 lm wide, hyaline, enteroblastic, phialidic, doliiform to ampulliform, determinate, discrete,
glabrous. Conidia 4.5–7 9 3–5 lm (
x = 5.7 9 3.9,
n = 30), initially hyaline, becoming pale brown to dark
brown at maturity, globose to obovate, ellipsoidal to subcylindrical, rounded at both ends, unicellular, thick-walled,
smooth.
Culture characteristics: Colonies on PDA attaining
30–40 mm diam. after 4 weeks at 20–25 °C, with circular
margin, dark jacinth to orange red to dark olivaceous,
flattened, dense, aerial mycelium on the surface, reverse
similar in colour.
Material examined: ITALY, Province of Arezzo [AR],
Bagno di Cetica, on dead stem of Cytisus sp. (Fabaceae), 7
October 2012, Erio Camporesi, IT-784 (MFLU 16-1871,
holotype); ex-type living culture, MFLUCC 13-0396,
ICMP; ibid. IT-784B (HKAS 93585, isotype); living culture, KUMCC 16-0026.
Notes: Neomicrosphaeropsis cytisi differs from
N. cytisinus in the form of the conidiomata. Neomicrosphaeropsis cytisi has immersed, ostiolate conidiomata
that are smaller than those of N. cytisinus which are semiimmersed when immature, and become erumpent at
maturity (190–220 lm high 9 210–250 lm diam.)
Neomicrosphaeropsis cytisinus Tennakoon, Camporesi &
K.D. Hyde, sp. nov.
Index Fungorum number: IF552262; Facesoffungi
number: FoF02396, Fig. 19
Etymology: Name reflects the host genus Centaurea,
from which the holotype was collected.
Holotype: MFLU 16-1364.
Saprobic on Cytisus sp. Sexual morph Undetermined.
Asexual morph Coelomycetous Conidiomata 190–220 lm
high 9 210–250 lm diam. (
x = 232 9 204, n = 10),
stromatic, solitary, immersed to semi-immersed when
immature, becoming erumpent at maturity, globose to
subglobose, uniloculate, black, dehiscing by an irregular
split of the host epidermis. Peridium 20–25 lm wide,
composed of 4–5 layers of light brown cells of textura
angularis to textura prismatica. Conidiogenous cells 1–
2 lm wide, phialidic, hyaline, thin-walled, smooth, integrated, producing a single conidium at the apex. Conidia
5–7 9 3–5 lm (
x = 67 9 4.2, n = 30), initially hyaline,
becoming light brown, moderately thick-walled, smooth,
aseptate, ovoid, obtuse at apex, truncate or rounded at base.
Culture characteristics: Colonies on PDA reaching
25–30 mm diam. after 8 days at 20–25 °C, medium sparse,
circular, flat, slightly rough at surface with entire edge,
with a well-defined margin, cottony to fairly fluffy with
39
sparse mycelium; from above: white to cream at the margin, white to yellowish at the centre; from below, light
yellow to light brown at the margin, yellowish at the
centre; mycelium white to cream with tufting; not producing pigments in PDA medium.
Material examined: ITALY, Province of Arezzo [AR],
near Croce di Pratomagno, on dead stem of branch of
Cytisus scoparius L. (Fabaceae), 24 June 2012, E. Camporesi, IT 472 (MFLU 16-1364, holotype; HKAS 93703,
isotype), ex-type living cultures, MFLUCC 16-0790,
KUMCC 15-0557.
Notes: Neomicrosphaeropsis cytisinus resembles N. cytisi
in sharing the size range of conidiophores and conidiogenous
cells, but differs in the size of conidiomata (75–130 9
74–157 lm) and thickness of peridium (10–24 lm). Neomicrosphaeropsis cytisinus differs from Neomicrosphaeropsis
minima in the size of conidiomata (3–5.5 9 2–4 lm),
conidiophores and host. Phylogenetic analyses show that
they are distinct with high bootstrap support (98 % ML, 1.00
BYPP, Fig. 17).
Neomicrosphaeropsis minima W.J. Li, Camporesi & K.D.
Hyde, sp. nov.
Index Fungorum number: IF552213; Facesoffungi
number: FoF02348, Fig. 20
Etymology: Named for the small conidiomata.
Holotype: MFLU 16-1490
Saprobic on dead stems of Verbascum sp. (Scrophulariaceae), forming numerous, conspicuous, rounded,
black conidiomata. Sexual morph Undetermined. Asexual
morph Coelomycetous. Conidiomata 60–80 lm diam.,
60–95 lm high, black, solitary to gregarious or confluent,
pycnidial, globose to subglobose, immersed or semi-immersed, unilocular, thick-walled, smooth, ostiolate. Ostiole
single, short, centrally located. Wall of conidiomata 8–
15 lm wide, composed of thick-walled, brown to hyaline
cells of textura angularis. Conidiophores 2.7–5.5 lm
long 9 3.2–5.5 lm wide, occasionally present, hyaline,
doliiform to ampulliform, arising from inner layers of the
pycnidial wall. Conidiogenous cells 2.6–5.5 lm long 9 2–
3.5 lm wide, hyaline, enteroblastic, phialidic, doliiform or
cylindrical to ampulliform, with a periclinal wall thickening at the tip, smooth. Conidia 2.8–5.4 9 2–3.6 lm
(
x = 4.1 9 2.8, n = 30), hyaline when young, becoming
brown at maturity, oval, rounded at both ends, unicellular,
thick-walled, smooth, guttulate.
Culture characteristics: Colonies on PDA attaining
15–20 mm diam. after 7d at 20–25 °C with circular margins, white to dark brown, flattened, with felt-like, dense,
aerial mycelium on the surface, reverse with dark brown in
the central zone, white on the edge, sporulating.
Material examined: ITALY, Province of Arezzo [AR],
near Montemignaio, on dead stems of of Verbascum sp.
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b Fig. 24 RAxML tree based on a combined dataset of LSU, SSU, ITS
and TEF partial sequences of Didymosphaeriaceae. Bootstrap support
values for maximum likelihood (ML) higher than 60 % and Bayesian
posterior probabilities (BYPP) greater than 0.90 are defined as above
the nodes respectively. The tree is rooted to Pleospora herbarum. The
ex-type strains are in bold; the new isolates are in blue
(Scrophulariaceae), 1 October 2012, Erio Camporesi, IT765 (MFLU 16-1490, holotype); ex-type living culture,
MFLUCC 13-0394; ibid. IT-765B (HKAS 95027, isotype);
living culture, KUMCC 16-0024.
Notes: Phylogenetic tree based on multi-gene (LSU,
ITS, RPB2 and b-tubulin) shows that the three collections
(MFLUCC 13-0394, MFLUCC 13-0396, KUMCC 15-0557)
cluster with newly introduced genus Neomicrosphaeropsis
(Thambugala et al. 2016a). However, these strains formed
a separate branch, basal to N. rossica Thambugala, Bulgakov & K.D. Hyde. Morphologically, they share similar
conidia characters with the type species N. italica.
Neomicrosphaeropsis minima is closely related to N. cytisi
in our phylogenetic tree (Fig. 17), but can be easily distinguished by the dimensions of the conidiomata. Neomicrosphaeropsis minima has immersed to semi-immersed
conidiomata smaller than these of N. cytisi which has
conidiomata 75–155 lm diam. 9 75–130 lm high. In
addition, N. minima has conidiogenous cell walls, thickened at the apex, while this character is not observed in
N. cytisi.
Neodidymelliopsis Q. Chen & L. Cai
Neodidymelliopsis was introduced to accommodate
Neodidymelliopsis cannabis (G. Winter) Q. Chen & L. Cai.
as the type species, N. polemonii (Cooke) Q. Chen & L.
Cai, N. xanthine (Sacc.) Q. Chen & L. Cai, and two
unidentified species (Chen et al. 2015b). The conidia of this
genus are variable in shape: ovoid to ellipsoidal, cylindrical, allantoid, hyaline to pale brown or pale yellowish, 0–1septate.
Neodidymelliopsis ranunculi W.J. Li, Camporesi & K.D.
Hyde, sp. nov.
Index Fungorum number: IF552214; Facesoffungi
number: FoF02349, Fig. 21
Etymology: Named after host genus Ranunculus.
Holotype: MFLU 16-1870
Saprobic on dead stem of Ranunculus sp. (Ranunculaceae), Sexual morph undetermined. Asexual morph
Coelomycetes. Conidiomata 100–120 lm high, 95–
110 lm diam., pycnidial, solitary, globose to subglobose,
black, immersed to semi-immersed, unilocular, ostiolate,
wall 14–32 lm wide, composed of dark brown to light
brown, thick-walled cells of textura angularis. Ostiole
single, short, eccentric. Conidiophores reduced to conidiogenous cells. Conidiogenous cells 3–5 lm long 9 4.5–
41
10 lm wide, hyaline, enteroblastic, phialidic, determinate,
discrete, doliiform, with a small collarette, smooth, arising
from inner layers of conidiomata. Conidia 3–5 9 7.5–
10 lm (
x = 4.3 9 8.7 lm, n = 30), initially hyaline,
becoming pale brown at maturity, ellipsoidal, rounded at
both ends or 1-septate, straight or slightly curved, guttulate,
smooth-walled.
Culture characteristics: Colonies on PDA attaining
40–50 mm diam. after 7d at 20–25 °C, margins circular,
floccose, white, pale grey to olivaceous near the center;
reverse with white margin, dark brown at central zone,
sporulating after 4 weeks.
Material examined: ITALY, Province of Forlı̀-Cesena
[FC], Castrocaro Terme e Terra del Sole, near Converselle,
on dead stem of Ranunculus sp. (Ranunculaceae), 2
December 2012, Erio Camporesi, IT-936 (MFLU 16-1870,
holotype); ex-type living culture, MFLUCC 13–0490,
ICMP; ibid. (KUN, HKAS 95028, isotype), living culture,
KUMCC 16-0025.
Notes: Neodidymelliopsis ranunculi fits well within the
morphological concept for Neodidymelliopsis. Based on
both morphology and phylogeny, N. ranunculi is introduced as a new species in the genus.
Platychora Petr.
The genus Platychora is typified by P. ulmi (J. Schröt.)
Petr. and comprises the type species and P. alni (Peck)
Petr. (Index Fungorum 2016). This genus is characterized
by multi-loculate stromatic tissues and it produces
apiospores.
Platychora ulmi (J. Schröt.) Petr., Annls mycol. 23(1/2):
103 (1925)
= Sphaeria ulmi Schleich. ex Fr., Observ. mycol.
(Havniae): 173 (1815)
Index Fungorum number: IF280985; Facesoffungi
number: FoF02431, Fig. 22
Reference specimen: MFLU 16-1972
Saprobic on dead leaves of Ulmus sp. Sexual morph
Stromata up to 1 mm broad, sub-epidermal, solitary to
aggregated, superficial, cushion-shaped, multi-loculate,
black. Locules 100–130 lm high 9 120–170 lm diam.
(
x = 216 9 250 lm, n = 10), immersed in stromatic tissues, globose to subglobose, coriaceous, papillate, ostiolate.
Peridium 13–16 lm wide (
x = 15 lm, n = 10), comprising
thick-walled, dark brown cells of textura angularis. Hamathecium comprising 3.5–5.5 lm wide (
x = 5 lm,
n = 10), filiform, hyaline, branched pseudoparaphyses. Asci
50–100 9 8.5–10.5 lm (
x = 76 9 10 lm, n = 10), 8spored, bitunicate, cylindrical, short pedicellate. Ascospores
10–15 9 3–6 lm (
x = 13 9 5 lm, n = 10), obliquely uniseriate, apiosporous, hyaline, ovoid, with a transverse septum near the lower end, not constricted at the septum.
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42
Fig. 25 Montagnula cirsii (holotype). a Herbarium material. b, c
Ascomata semi-immersed in the stem. d Vertical hand section of
ascoma. e Ostiole. f Vertical hand section of peridium. g Immature
asci. h, i, k Asci with ascospores. Note the bitunicate asci.
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j Pseudoparaphyses. n Germinating ascospore. o–s Ascospores.
l Colony on PDA from above. m Colony on PDA from below. Scale
bars b = 500 lm, c = 100 lm, d = 50 lm, e–f, n = 20 lm, g–
j = 10 lm, k, o–s = 5 lm
Fungal Diversity (2016) 80:1–270
Asexual morph Conidiomata 130–140 lm high 9 190–
210 lm diam. (
x = 138 9 200 lm, n = 10), subcuticular,
acervular, brown to black, applanate to pulvinate. Conidiophores reduced to conidiogenous cells. Conidiogenous cells
10–15 lm high 9 2–4 lm wide (
x = 138 9 200 lm,
n = 10), solitary, septate at the base, olivaceous, smooth,
cylindrical, straight, holoblastic, annellidic. Conidia 3–
5.5 9 2.5–3 lm (
x = 5 9 2.7 lm, n = 10), oval, pale
brown, aseptate, thin-walled, smooth, ornamented.
Culture characteristics: Colonies growing on MEA,
becoming 1 cm within 7 days at 18 °C, circular, flat,
irregular margin, with sparse aerial mycelium, white.
Material examined: ITALY, Province of Forlı̀-Cesena
[FC], Isola di Santa Sofia, on dead leaves of Ulmus sp.
(Ulmaceae), 20 January 2014, Erio Camporesi, IT 1670
(MFLU 16-1972, reference specimen designated here),
living culture, MFLUCC 14-1186.
Notes: The phylogenetic placement of Platychora ulmi
is not clear. Most phylogenetic studies placed this genus in
Pleosporales without assignment to any family (Winton
et al. 2007). However Hyde et al. (2013) included Platychora within Didymellaceae and combined ITS, LSU,
RPB2 and Beta tubulin gene analysis in this study (Fig. 17)
support the taxonomic placement of Platychora ulmi in
Didymellaceae. We also illustrate the asexual morph of
Platychora ulmi from culture.
Stagonosporopsis Died.
Stagonosporopsis was introduced by Diedicke (1912).
This genus is characterised by having conidia with one, two
or occasionally three septa. Jaczewski (1917) treated
Stagonosporopsis as a subgenus of Ascochyta, whereas
Petrak (1925) reduced this genus to synonymy with
Ascochyta.
Stagonosporopsis centaureae Tennakoon, Camporesi &
K.D. Hyde, sp. nov.
Index Fungorum number: IF552261; Facesoffungi
number: FoF02397, Fig. 23
Etymology: Name reflects the host genus Centaurea,
from which the species was collected.
Holotype: MFLU 16-1365
Saprobic on dead stem of Centaurea cyanus. Sexual
morph Ascomata 110–120 lm high 120–130 lm diam.,
solitary, scattered, partly immersed to superficial, raised,
dark brown to black, globose to subglobose, glabrous, uniloculate, with centrally located ostiole with minute papilla.
Peridium 22–27 lm wide, thin-walled, with unequal
thickness, composed of 4–5 layers of dark brown to black
pseudoparenchymatous cells, arranged in textura angularis
to textura prismatica. Hamathecium composed of numerous, 2–2.5 lm wide, filamentous, distinctly septate, cellular pseudoparaphyses, non-constricted at the septum,
43
anastomosing at the apex, embedded in a hyaline gelatinous matrix. Asci 35–48 9 7.2–8.2 lm (
x = 42.5 9 8.1
lm, n = 30), 8-spored, bitunicate, fissitunicate, cylindrical
or clavate, short pedicellate, with furcate pedicel, apically
rounded with an indistinct ocular chamber. Ascospores 13–
15 9 3–5 lm (
x = 14.3 9 4.3 lm, n = 30), overlapping,
uni- to bi-seriate, hyaline, ellipsoidal to clavate or fusiform
with rounded ends, 1-septate, slightly constricted at the
septum, echinulate, straight to slightly curved. Asexual
morph Undetermined.
Culture characteristics: Colonies on PDA reaching
25–30 mm diam. after 8 days at 20–25 °C, colonies medium sparse, circular, flat, slightly rough surface with entire
edge, well defined margin, cottony to fairly fluffy with
sparse mycelium; from above: dark brown to yellowish at
the margin, dark brown to black at the centre; from below:
dark brown to yellowish-brown at the margin, dark brown
at the centre; mycelium light brown to yellowish with
tufting; not producing pigments in PDA medium.
Material examined: ITALY, Province of Arezzo [AR],
near Quota, on dead stem of Centaurea cyanus (Asteraceae), 24 June 2015, E. Camporesi, IT 2548 (MFLU
16-1365 holotype; HKAS 93705 isotype), ex-type living
cultures, MFLUCC 16-0787, KUMCC 15-0559.
Notes: Stagonosporopsis centaureae clusters with
Stagonosporopsis trachelii with high support (93 % ML,
0.99 BYPP, Fig. 17). It can be easily distinguished from
S. trachelii which has smaller ascomata (60–80 lm diam.)
and smaller ascospores (4–6 9 1.5–2.5 lm).
Didymosphaeriaceae Munk
Details of this family can be seen in Ariyawansa et al.
(2014a), Li et al. (2016) and Wanasinghe et al. (2016).
Montagnula Berl.
The genus Montagnula was introduced by Berlese
(1896) to accommodate M. infernalis (Niessl) Berl. and
M. gigantea (Mont.) Berl. Ariyawansa et al. (2014c) placed
Montagnula in the family Didymosphaeriaceae including
some phragmosporous and didymosporous species, based
on morphology and available phylogenetic analysis. Montagnula was accepted as a distinct genus in the family
Didymosphaeriaceae in several recent studies (Ariyawansa
et al. 2014c; Hongsanan et al. 2015a; Li et al. 2016).
Presently, there are 28 species in this genus including two
recently described species, M. bellevaliae Wanasinghe
et al. and M. scabiosae Wanasinghe et al. (Hongsanan et al.
2015a).
Montagnula cirsii Qing Tian, Camporesi & K.D. Hyde, sp.
nov.
Index Fungorum number: IF552184; Facesoffungi
number: FoF02255, Fig. 25
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b Fig. 26 Tremateia arundicola (holotype). a Appearance of asco-
mata on host substrate. b Section of ascoma. c Peridium. d Pseudoparaphyses. e–g Asci. h–k Ascospores. Scale bars b, c = 50 lm,
d = 10 lm, e–g = 20 lm, h–k = 10 lm
Etymology: In reference to the host genus Cirsium, from
which this holotype was collected.
Holotype: MFLU 14-0730.
Saprobic under periderm or semi-immersed in woody
plant substrates. Sexual morph Ascomata 385–415 lm
diam. 9 510–525 lm high (
x = 400 9 515.5 lm, n =
10), semi-immersed to erumpent, solitary, scattered, or
sometimes gregarious, globose, black, smooth-walled, neck
long, with a small, flattened, ellipsoid ostiole at the apex.
Peridium 41–58.5 lm (
x = 43.3 lm, n = 6), 2-layered,
the outer layer composed of irregular, thick-walled, brown
cells of textura angularis; the inner layer composed of
hyaline, smaller cells of textura angularis. Hamathecium
comprising 1–2 lm broad, septate, long, colourless, branched or simple pseudoparaphyses, surrounding the asci.
Asci 84.5–119.5 9 10.5–13.5 lm (
x = 101 9 12 lm,
n = 10), 8-spored, bitunicate, clavate, long pedicellate,
apically rounded with an ocular chamber. Ascospores 18–
23.5 9 6.5–9.5 lm (
x = 21.5 9 8 lm, n = 10), overlapping uni-seriate or uni-seriate, 3-septate, ellipsoid to fusiform, curved, yellow to brown, constricted at the septa,
broader at the middle two cells, tapering or obtuse at both
ends, smooth-walled. Asexual morph Undetermined.
Culture characteristics: Ascospore germinating on PDA
within 12 h. Colonies on PDA, reaching 8 mm diam. in
7 days at 25 °C. Mycelium superficial, hyaline, hairy, with
entire edge, floccose at the center, drift white from above
and light brown at the center from below.
Material examined: ITALY, Province of Forlı̀-Cesena
[FC], Balze-Verghereto, on dead stem of Cirsium sp.
(Compositae), 21 May 2013, Erio Camporesi (MFLU
14-0730, holotype); ibid., (HKAS 94523, isotype); ex-type
living culture, MFLUCC 13-0680, KUMCC 16-0018.
Notes: Montagnula cirsii is morphologically similar to
M. bellevaliae and M. scabiosae (Hongsanan et al. 2015a).
However, they can be distinguished by various characters,
such as host, shape of ascomata and different sizes of asci.
Montagnula bellevaliae occurs on dead stems of Bellevalia
romana, M. scabiosae on dead stems of Scabiosa sp. and
M. cirsii on dead stems of Cirsium sp. The orientation of
ascomata is also different. Montagnula bellevaliae has an
eccentric papilla, while the other two species have regular
papilla. The size of asci is also quite different; Montagnula
cirsii is smaller than M. scabiosae (84.5–119.5 9
10.5–13.5 lm vs. 110–130 9 14–20 lm) but larger than
M. bellevaliae which is 70–100 lm in length, 9–12 lm in
width.). Based on phylogenetic analysis (Fig. 24), our
strain clusters with M. scabiosae in Montagnula with
45
relatively high support (99 % MP /1.00 PP). We therefore
introduce a new species based both on morphology and
phylogeny.
Tremateia Kohlm.
Facesoffungi number: FoF00223
Tremateia was introduced as a facultative marine genus,
characterized by ‘depressed globose, immersed ascomata,
numerous cellular pseudoparaphyses, fissitunicate and
clavate asci, ellipsoid muriform ascospores, and a Phomalike asexual morph’ (Kohlmeyer et al. 1995; Ariyawansa
et al. 2014a). Earlier the genus was identified as similar to
Lewia and Diademosa and placed in Pleosporaceae
(Kohlmeyer et al. 1995). DNA sequences based studies
have revealed that T. halophila groups in Didymosphaeriaceae (
x = Montagnulaceae), sister to Bimuria novae-zelandiae (Schoch et al. 2009; Suetrong et al. 2009;
Ariyawansa et al. 2014a, 2015a, b, c; Liu et al. 2015a; this
study). Ariyawansa et al. (2014a) confirmed the familial
status of Tremateia in Didymosphaeriaceae.
In this study the novel isolates of Tremateia arundicola
and T. guiyangensis grouped with T. halophila sister to
Bimuria novae-zelandiae. However these Tremateia
members are separated from Bimuria novae-zelandiae with
high statistical support in combined (LSU, SSU, ITS and
TEF) gene analyses (98 % ML and 1.00 BYPP, Fig. 24)
Tremateia arundicola Wanasinghe, E.B.G. Jones & K.D.
Hyde, sp. nov.
Index Fungorum number: IF552134; Facesoffungi
number: FoF02210, Fig. 26
Etymology: Name reflects the place Arun River, from
which the species was collected.
Holotype: MFLU 16-1275
Saprobic on dead herbaceous stems. Sexual morph
Ascomata 200–300 lm high 250–350 lm diam. (
x=
259.3 9 295.1 lm, n = 5) immersed to semi-erumpent,
solitary, scattered, broadly oblong to sub globose and
flattened, dark brown to black, coriaceous, ostiolate. Ostiole 40–60 lm high 50–70 lm diam. papillate, black,
smooth, filled with hyaline cells. Peridium 10–20 lm wide
at the base, 15–25 lm wide in sides, thick, with 4–5 layers,
outer layer heavily pigmented, thick-walled, comprising
reddish to dark brown cells of textura angularis, inner layer
composed of hyaline thin-walled cells of textura angularis.
Hamathecium comprising numerous, 3–4 lm wide, filamentous, branched, septate, pseudoparaphyses. Asci 170–
200 9 15–20 lm (
x = 183.3 9 18.4 lm, n = 20), 8spored, bitunicate, fissitunicate, cylindrical to cylindricclavate, pedicellate, thick-walled at the apex, with minute
ocular chamber. Ascospores 20–30 9 6–9 lm (
x=
25.9 9 6.6 lm, n = 50), overlapping uni-seriate, muriform, ellipsoidal to subfusiform, slightly curved, upper part
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Fig. 27 Tremateia guiyangensis (holotype). a, b Appearance of
ascomata immersed on host surface. c Vertical section of ascoma.
d Section of peridium. e Pseudoparaphyses. f–i Cylindrical-clavate
asci. j–o Ascospores. Scale bars a = 500 lm, b = 100 lm,
c = 50 lm, d, j–o = 10 lm, e–i = 20 lm
wider than the lower part, 3–6 transversely septate, with 1
vertical septum, constricted at the septa, initially hyaline,
becoming pale-brown at maturity, ends remaining coneshaped, with pointed upper ends and rounded lower ends,
lacking a mucilaginous sheath or disappear with maturity.
Asexual morph Undetermined.
Culture characteristics: Colonies on MEA reaching
2 cm diam. after 30 days at 18 °C, circular, smooth margin
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Fig. 28 Phylogram generated from maximum Likelihood analysis
(MEGA6) based on combined dataset of SSU and LSU rDNA
sequences of the two Lentithecium strains and related taxa in the
47
family Lentitheciaceae. The tree is rooted to Dothidea sambuci.
Maximum Likelihood bootstrap support values greater than 50 % are
indicated. The new species are in blue and ex-types in bold
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Fig. 29 Lentithecium unicellulare (CBS H-22674, holotype). a, b Vertical sections of conidiomata. c Peridium. d–g Developing stages of
conidia. h–k Conidia. Scale bars a, b = 50 lm, c = 15 lm, d–m = 10 lm
white at first, dirty white after 4 weeks flat on the surface,
without aerial mycelium, reverse greenish-grey. Hyphae
septate branched, hyaline, thin, smooth-walled.
Material examined: UK, England, Arun River, 6 April
2015, on herbaceous stem, E.B.G. Jones, GJ126 (MFLU
16-1275, holotype); (isotype in BBH)
Notes: Tremateia arundicola resembles T. guiyangensis
and T. halophila in having globose to subglobose ascomata
with a thin peridium comprising cells of textura angularis
and brown muriform ascospores. Both T. guiyangensis and
T. halophila have comparatively shorter (
x = 150 lm)
asci. They also have ascospores with more than 6 transverse septa and 2 vertical septa, while T. arundicola
ascospores have less than 6 transverse septa and only one
vertical septum.
Tremateia guiyangensis J.F. Zhang, J.K. Liu, K.D. Hyde
& Z.Y. Liu, sp. nov.
123
Index Fungorum number: IF552160; Facesoffungi
number: FoF02235, Fig. 27
Etymology: Name reflects the place Guiyang, where the
holotype was collected.
Holotype: MFLU 16-1299
Saprobic on dead herbaceous stems. Sexual morph
Ascomata 130–280 lm high, 190–400 lm diam., scattered
to clustered, subglobose to ovoid, with flattened base,
coriaceous, immersed to semi-immersed or breaking
through the host epidermis, dark brown to black, ostiolate,
minutely papillate. Peridium up to 9–16 lm, composed of
several layers of light to brown, thick-walled cells of textura angularis, becoming thin-walled and hyaline towards
the centrum. Hamathecium comprising numerous, 2.3–
5.1 lm wide, hypha-like, septate pseudoparaphyses,
tapering towards the terminal cells, and intermingled
among asci, embedded in a gelatinous matrix. Asci 152–
160 9 21–27 lm (
x = 149 9 24 lm, n = 20), 8-spored,
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49
Fig. 30 Lentithecium
voraginesporum (CBS
H-22560, holotype). a Vertical
section of ascoma. b Magnified
part of the vertical section of the
ascoma showing the peridium
structure. c, d Mature asci. e,
f Ascospores. Scale bars
a = 50 lm, b–d = 10 lm, e,
f = 5 lm
bitunicate, fissitunicate, cylindrical-clavate, with a furcate
pedicel, apically rounded, with a minute ocular chamber.
Ascospores
20–28 9 9–12 lm
(
x = 25 9 11 lm,
n = 30), muriform, normally overlapping 1–2-seriate,
ellipsoid to broad fusiform, both parts of the spore ±equal
in size, with 3–5 transverse septa, and 1 longitudinal septum in each row, obviously constricted in the central septum and slightly constricted at other septa, initially hyaline,
becoming golden-brownish to brown when mature, rounded at both ends and lacking any gelatinous sheath or
appendages. Asexual morph Undetermined.
Culture characteristics: Colonies on PDA reaching
3.5 cm diam. after 12 days at 25 °C, circular, raised at
center, margin smooth, white at first, palely pigmented
after 10 days at the center, reverse yellowish to reddish.
Hyphae septate, branched, hyaline, thin, smooth-walled.
Material examined: CHINA, Guizhou, Guiyang, around
Guizhou Academy of Agriculture Sciences, on dead
herbaceous stems, 17 August 2015, J.F. Zhang, GZ-01
(MFLU 16-1299, holotype), ex-type living culture,
MFLUCC.
Notes: Tremateia guiyangensis is phylogenetically close
to T. arundicola and T. halophila, and also morphologically similar in having globose to subglobose or flattened,
immersed to semi-immersed ascomata, with clavate asci
and muriform ascospores. However, T. guiyangensis was
collected from terrestrial habitat, while the other two species were collected from aquatic habitats. In addition, the
asci of T. guiyangensis are shorter than T. arundicola
(
x = 149 vs. 185 lm) and the ascospores are smaller than
T. halophila (
x = 26 9 6.5 lm vs. 30 9 16 lm). There
are more transverse and vertical septa in T. guiyangensis
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than in T. arundicola, and the ascospores of T. arundicola
are paler than the other two species.
Lentitheciaceae Yin. Zhang et al.
Zhang et al. (2009b) established Lentitheciaceae to
accommodate the saprobic genera Katumotoa, Keissleriella, Lentithecium and Tingoldiago described from
aquatic and terrestrial habitats. The type genus of the
family is the aquatic genus Lentithecium. Several new
genera and species from different habitats have since been
introduced in Lentitheciaceae based on molecular data.
Quaedvlieg et al. (2013) established Setoseptoria to
accommodate saprobic coelomycetes that are Septoria-like
but with setose conidiomata and belong to Lentitheciaceae.
Knapp et al. (2015) described the endophytic genus,
Darksidea from semiarid sandy grasslands and multi-genes
phylogenetic analyses placed Darksidea in Lentitheciaceae. Phookamsak et al. (2015) introduced Poaceascoma
to accommodate a species with setose ascomata and filiform ascospores. Wijayawardene et al. (2015) introduced
Fig. 31 RAxML maximum
likelihood phylogenetic tree
based on a LSU and ITS
sequence data from species of
Leptosphaeriaceae. Maximum
likelihood bootstrap support
values greater than 50 % are
shown above the nodes. The extype strains are in bold and the
new isolates are in red. The tree
is rooted with Alternariaster
helianthi
Fig. 32 Leptosphaeria cirsii (holotype). a, b Ascomata on host. c
c Section through the ascoma. d Peridium. e Pseudoparaphyses.
f Mature ascus. g Immature ascus. h, i Ascospores. j Germinated
ascospore. k Ascospores stained in Indian ink. l, m Culture from
above (l) and below (m). n Fruiting body in culture. o Vertical section
of conidioma. p Conidioma wall. q, r Conidiogenous cells and
conidia s Conidia. Scale bars a = 500 lm, b = 200 lm,
c = 100 lm, d, e–g, o, p = 30 lm, h–k = 10 lm, l–m = 4 cm, q–
s = 5 lm
Phragmocamarosporium Wijayawardene et al. to accommodate two coelomycetous species with brown phragmospores and phialidic conidiogenesis. The family currently
includes 12 genera (Tanaka et al. 2015). In this paper we
introduce two new species of Lentithecium and two new
species of Poascoma and provide a new phylogenetic tree
for Lentitheciaceae (Fig. 28).
Lentithecium K.D. Hyde et al.
The genus Lentithecium was established to accommodate Massarina arundinacea (Sowerby) Leuchtm.,
Leptosphaeria doliolum CBS 125979
Leptosphaeria doliolum CBS 541 66
54 Leptosphaeria doliolum CBS 505 75
Leptosphaeria doliolum MFLUCC 15-1875
Leptosphaeria veronicae CBS 126583
90
Leptosphaeria veronicae CBS 145 84
Leptosphaeria doliolum CBS 130000
Leptosphaeria doliolum CBS 155 94
62
71
Leptosphaeria errabunda CBS 617 75
errabunda CBS 125978
Leptosphaeria
Leptosphaeria macrocapsa CBS 640.93
sensu stricto
79
Leptosphaeria sydowii CBS 385 80
50
99 Leptosphaeria sydowii CBS 125976
Leptosphaeria ebuli MFLUCC 14-0828
Leptosphaeria cichorium MFLUCC 14-1063
61
Leptosphaeria cirsii MFLUCC 14 1170
Leptosphaeria italica MFLUCC 15-0174
96 Leptosphaeria sclerotioides CBS 148 84
Leptosphaeria sclerotioides CBS 144 84
100 Leptosphaeria
conoidea CBS 616 75
Leptosphaeria conoidea CBS 125977
98Leptosphaeria
100
82
Leptosphaeria slovacica CBS 125975
Leptosphaeria slovacica CBS 389 80
Leptosphaeria irregularis MFLUCC 15 1118
99
Leptosphaeria pedicularis CBS 126582
80
Leptosphaeria pedicularis CBS 390 80
Alloleptosphaeria italica MFLUCC 14-0934
Pseudoleptosphaeria etheridgei CBS 125980
100 Neoleptosphaeria rubefaciens CBS 387 80
Neoleptosphaeria rubefaciens CBS 223.77
Alternariaster helianthi CBS 327.69
100
0.1
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b Fig. 33 Leptosphaeria irregularis (holotype). a, b Appearance of
ascomata on host substrate. c Vertical section through ascoma.
d Close up of the peridium. e Close up of the ostiole. f Pseudoparaphyses. g, h Asci. i–k Immature and mature ascospores. l Germinating ascospore. m, n Colonies on MEA. Scale bars b = 1 mm,
c = 200 lm, d–h = 50 lm, i–i = 20 lm
M. fluviatilis Aptroot & Van Ryck. and Keissleriella linearis E. Müll. ex Dennis. The genus currently contains
seven species that were described from aquatic (Suetrong
et al. 2009; Tanaka et al. 2015), or L. arundinaceum
(Sowerby) K.D. Hyde et al. and L. rarum (Kohlm. et al.)
Suetrong et al. from marine habitats. Lentithecium species
have been described from decayed submerged parts of
Phragmites (three species), Juncus (one species), and
submerged wood (three species) (Zhang et al. 2009b;
Suetrong et al. 2009; Tanaka et al. 2015).
Lentithecium unicellulare Abdel-Aziz, sp. nov.
Index Fungorum number: IF552267; Facesoffungi
number: FoF02433, Fig. 29
Etymology: In reference to the unicellular conidia.
Holotype: CBS H-22674.
Saprobic on decayed wood in freshwater habitats. Sexual morph Undetermined. Asexual morph Conidiomata
115–235 lm high 9 140–235 wide lm (
x = 176.6 9
252.5 lm, n = 6), pycnidial, solitary or aggregated,
uniloculate, immersed, erumpent to superficial, globose,
subglobose, ovate, elongated, papillate, ostiolate, dark
brown to black, surrounded by brown, septate, thick-walled
hyphae, 3–4 lm wide. Pycnidial wall 18–25 lm thick,
composed of 6–8 layers of yellow-brown to black brown
cells of textura angularis, hyaline inner layer lining bearing
conidiogenous cells. Conidiophores reduced to conidiogenous cells. Conidiogenous cells 9–14 9 8–12 lm
(
x = 11.8 9 10.3 lm, n = 12), holoblastic, determinate,
smooth-walled, hyaline, globose, subglobose to pearshaped with rounded or truncate base, sometimes proliferating once and each conidiogenous cell producing one to
two conidia. Conidia 6–9 9 4–5 lm (
x = 8.1 9 4.3 lm,
n = 50), subglobose, ovate, clavate, ellipsoid, allantoid,
rectangular or irregular, with or without truncate base,
hyaline, unicellular, smooth-walled.
Culture characteristics: Colonies on PDA 45–50 mm
diam. after 10 days at 23 °C, dark grey with irregular
margins, reverse black, dense growth with aerial and submerged mycelia without diffusible pigments. Pycnidia
were produced in pure cultures after 4 weeks. Dimensions
of pycnidia and conidia were similar to those recorded
from natural substrates.
Material examined: EGYPT, Sohag City, on decayed
wood submerged in the River Nile, 14 August 2014, F.
53
A. Abdel-Aziz (CBS H-22674, holotype), ex-type living
culture MD 6004.
Notes: Asexual genera reported so far under Lentitheciaceae are coelomycetous with variable morphology and
these include: Phragmocamarosporium, Pleurophoma, Setoseptoria and Stagonospora macropycnidia (Tanaka et al.
2015). Lentithecium unicellulare differs from Stagonospora
species in having globose, unicellular, holoblastic, determinate conidia that are smaller. Species of Stagonospora
have cylindrical, multi-septate conidia, while conidiogenesis
is holoblastic, occasionally annellidic, with single proliferation, discrete and indeterminate (Sutton 1980). The genus
Stagonospora is polyphyletic and considered as the asexual
morph of Phaeosphaeria (Leuchtmann 1984). Pleurophoma
species differ from L. unicellulare in having long, filiform,
septate, branched conidiophores and enteroblastic, phialidic,
integrated, determinate conidiogenous cells (Sutton 1980).
Setoseptoria species differ from L. unicellulare in having
subcylindrical, transversely euseptate conidia, becoming
constricted at septa and disarticulating into phragmospores
when old. Conidiogenesis in Setoseptoria rarely has percurrent proliferations (Quaedvlieg et al. 2013).
Lentithecium voraginesporum Abdel-Wahab, Bahkali &
E.B.G. Jones, sp. nov.
Index Fungorum number: IF 552266; Facesoffungi
number: FoF02432, Fig. 30
Etymology: After the Latin word ‘‘Voraginem’’, meaning ‘‘gulf’’, where the fungus was recorded.
Holotype: CBS H-22560.
Saprobic on submerged wood in mangroves. Sexual
morph Ascomata 125–215 lm diam., globose to subglobose, immersed to erumpent, ostiolate, papillate, coriaceous. Peridium 25–30 lm thick at the upper part around
the ascomatal venter, two-layered, outer layer with yellowbrown cells forming a textura-angularis, inner-layer comprising thin-walled, hyaline, flattened cells; 11–17 lm
thick at the basal part of the ascomata. Asci 38–50 3 8–
10 lm (
x = 44 3 9.3 lm, n = 15), 8-spored, bitunicate,
clavate, short pedicellate, developing at the base of venter.
Ascospores
15–21 3 5–6 lm
(
x = 18.8 3 5.3 lm,
n = 50), bi-seriate, yellow-brown to reddish-brown, ellipsoidal with rounded ends, 1-septate, septum sub-median,
upper cell is longer and wider, with roughened surface.
Asexual morph Undetermined.
Culture characteristics: Colonies on PDA 20–25 mm
diam. after 10 days at 23 °C, brown to dark-brown aerial
and immersed mycelia with dark-brown reverse with circular margins, without diffusible pigments.
Material examined: SAUDI ARABIA, Arabian Gulf,
Tarut mangroves, on submerged, decayed Phragmites
australis (Cav.) Trin. ex Steud. (Poaceae), stem inside the
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b Fig. 34 Phylogram generated from maximum likelihood analysis
based on combined LSU and SSU sequence data for species of
Lindgomycetaceae. Maximum likelihood, Maximum parsimony bootstrap support values greater than 50 % and Bayesian posterior
probabilities greater than 0.90 are near the nodes. The new isolates are
in blue bold, and other ex-type strains are in black bold. The scale bar
indicates 0.006 changes. The tree is rooted with Aigialus mangrovis
and A. grandis
mangrove stand, 28 March 2013, M.A. Abdel-Wahab (CBS
H-22560, holotype), ex-type living culture MD 1342.
Notes: Lentithecium voraginesporum is the third marine
species in the genus and differs from the other seven
described Lentithecium species in having small ascomata,
asci and ascospores. Ascospores of the new species are
characterized by their brown colour, roughened surface and
the absence of a gelatinous sheath. The phylogenetic analyses of both SSU and LSU sequence data place L. voraginesporum within the family Lentitheciaceae in a wellsupported clade with the recently described species,
L. cangshanense Z.L. Luo et al. and L. unicellulare AbdelAziz. Both species were described from freshwater habitats
(Su et al. 2016b; this article). Lentithecium voraginesporum
differs from L. cangshanense in having smaller ascomata
(125–215 lm vs. 210–320 lm) and asci (38–50 3 8–10 lm
vs. 65–78 3 11–13 lm), longer and narrower ascospores
with a rough surface (15–21 3 5–6 lm vs. 16.5–
17.5 3 6–7 lm) and its marine habitat. Lentithecium unicellulare produces dark brown to black pycnidia and unicellular, hyaline conidia and its sexual morph is unknown.
Molecular data delimit L. voraginesporum from L. cangshanense and L. unicellulare (Fig. 28). Lentithecium fluviatile differs from L. voraginesporum in having hyaline,
longer and wider ascospores (24–31 3 7–10 lm vs.
15–21 3 5–6 lm) that are 2–3-septate and surrounded by a
wide expanding mucilaginous sheath (van Ryckegem and
Aptroot 2001). Lentithecium clioninum and L. pseudocloninum have larger asci and ascospores than those reported
for L. voraginesporum. Ascospores of the latter two species
are hyaline and surrounded by gelatinous sheath (Tanaka
et al. 2015).
Leptosphaeriaceae M.E. Barr
The family Leptosphaeriaceae was established by Barr
(1987) in the order Pleosporales and is typified by Leptosphaeria. Species of Leptosphaeriaceae can be saprobic,
hemibiotrophic or parasitic on stems and leaves of herbaceous or woody plants in terrestrial habitats (Hyde et al.
2013). Members of this family are characterized by single,
papillate, immersed or erumpent, perithecial ascomata,
with relatively thick peridia, bitunicate cylindrical asci and
hyaline to brown, transversely septate ascospores (Hyde
et al. 2013). The asexual morphs of the family
55
Leptosphaeriaceae can be coelomycetous or hyphomycetous
(Alves et al. 2013; De Gruyter et al. 2013; Hyde et al. 2013;
Zhang et al. 2012b). In recent classifications Alternariaster,
Heterospora, Leptosphaeria, Neophaeosphaeria, Paraleptosphaeria, Plenodomus, and Subplenodomus were included in
the family (Ariyawansa et al. 2015b). In this study we introduce
two new species within the genus Leptosphaeria.
Leptosphaeria Ces. & De Not.
For classification of Leptosphaeriaceae we follow
Ariyawansa et al. (2015b) and provide an updated tree
(Fig. 31).
Leptosphaeria cirsii Jayasiri, Camporesi & K.D. Hyde, sp.
nov.
Index Fungorum number: IF552155; Facesoffungi
number: FoF02220, Fig. 32
Etymology: The specific epithet circii is based on the
host genus from which the taxon was collected.
Holotype: MFLU 15-1072
Saprobic on dead stem of Cirsium sp. Sexual morph
Ascomata 285–315 9 335–360 lm (
x = 304 9 347 lm,
n = 10), solitary or scattered, superficial, globose to subglobose, broadly or narrowly conical, coriaceous, smoothwalled, ostiolate. Ostiole usually papillate, darkened at the
base. Peridium 22–27 lm (
x = 24 lm, n = 20) wide,
comprising two types of cells, outer cells of 1–2 layers of
heavily pigmented cells of textura angularis, inner layer
composed of small, light brown to hyaline cells of textura
angularis. Hamathecium of septate, long, hyaline, cellular
pseudoparaphyses, between and above the asci. Asci 75–
110 9 10–12 lm (
x = 95 9 11 lm, n = 20), 8-spored,
bitunicate, fissitunicate, cylindro-clavate, with a short, broad
pedicel, thickened and rounded at the apex, with a distinct
ocular. Ascospores 19–22 9 4–4.5 lm (
x = 20 9 4.2 lm,
n = 40), overlapping uni-seriate, hyaline to light brown
when immature, becoming brown to chestnut brown when
mature, ellipsoidal with narrowly rounded ends, 3-septate,
smooth-walled with mucilaginous sheath. Asexual morph
Coelomycetous on MEA. Conidiomata 150–200 9 195–
220 lm (
x = 190 9 205 lm, n = 10), pycnidial, superficial, immersed in media, globose to subglobose, black,
without an ostiole. Conidiomata wall 15–25 lm wide,
composed cells of textura angularis in multiple layers,
pale yellowish-brown, remaining hyaline in inner layer.
Conidiogenous cells 2–5 9 2–4 lm (
x = 3.5 9 2.5 lm,
n = 30), enteroblastic, phialidic, determinate integrated,
subglobose to short conical. Conidia 3–6 9 1–3 lm
(
x = 3.8 9 1.5 lm, n = 50), hyaline, aseptate, oblong to
cylindrical, thin-walled, smooth, guttulate, rounded at
both ends.
Culture characteristics: Colonies on MEA 60 mm diam.
after 4 weeks at 18 °C, white on the top and reverse,
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Fig. 35 Arundellina typhae (holotype). a Appearance of immersed ascomata on host substrate. b Section of ascoma. c Peridium.
d Pseudoparaphyses. e–h Asci. i–m Ascospores. Scale bars b = 50 lm, c, d = 10 lm, e–h = 20 lm, i–m = 10 lm
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Fig. 36 Lindgomyces pseudomadisonensis (holotype). a, b Appearance of ascomata on substrate. c Ascoma in longitudinal section. d Peridium in longitudinal section. e Asci. f Pseudoparaphyses.
57
g–j Ascospores. k Germinating ascospore. Scale bars a = 400 lm,
b = 200 lm, c = 50 lm, d = 25 lm, e = 10 lm, f–k = 5 lm
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b Fig. 37 RAxML tree based on analysis of a combined dataset of ITS,
LSU, SSU and TEF partial sequences of Lophiostomataceae.
Bootstrap support values for maximum likelihood (ML, black) higher
than 50 % and Bayesian posterior probabilities (BYPP, red) greater
than 0.95 are defined as above the nodes. The tree is rooted to
Melanomma pulvis-pyrius. All type strains are in bold. New strains
are given in blue
reverse yellow to brown near the middle, some area dense,
circular, regular margin, without diffusible pigments.
Material examined: ITALY, Province of Trento [TN],
near Vermiglio—Val di Sole, on dead stem of Cirsium sp.
(Asteraceae), 9 August 2014, E. Camporesi IT 2044,
(MFLU 15-1072, holotype); (KUN, isotype); ex-type living culture, MFLUCC 14-1170, KUNCC
Notes: Leptosphaeria cirsii is introduced here based on
both morphology and phylogeny. Leptosphaeria cirsii is
typical of Leptosphaeria in having a peridium of
plectenchymatous cells and reddish to yellowish-brown,
fusoid, 3-septate ascospores. Phylogenetically, L. cirsii
clusters with L. cichorium with relatively moderate support
(61 % ML) and also shares affinities to L. italica. Morphologically L. cirsii is similar to L. cichorium in having
similar asci, ascospores, cellular pseudoparaphyses and
asexual state morphology, but differs in having, superficial,
globose to subglobose, broadly or narrowly conical ascomata and ascospores with a mucilaginous sheath (Ariyawansa et al. 2015b). Leptosphaeria cirsii resembles
L. italica but differs as L. cirsii has long and narrow
ascospores and lacks an ostiolar canal (Dayarathne et al.
2015).
Leptosphaeria irregularis R.H. Perera, E.B.G. Jones &
K.D. Hyde, sp. nov.
Index Fungorum number: IF552172, Facesoffungi
number: FoF02234, Fig. 33
Etymology: Refers to the irregular shape of the fruiting
body.
Holotype: MFLU 16-1091
Saprobic on unidentified plant stem. Sexual morph
Ascomata 245–370 lm high, 180–368 lm diam.
(
x = 285 9 255 lm, n = 10), superficial, appearing as
black raised spots on the host, solitary or arranged in
groups of 2–5, globose to irregularly globose, uniloculate,
black, ostiolate. Ostiole aperiphysate. Peridium 30–67 lm
wide, composed of 8–16 rows of scleroplectenchymatous
cells, outer layer of amorphous black cells, inner layers
composed of hyaline to pale brown cells of textura angularis. Hamathecium comprising numerous, 1.5–2.2 lm
(n = 25) wide, filamentous, branched, septate, pseudoparaphyses. Asci 110–147 9 6–8.5 lm (
x = 130 9 8 lm,
n = 30), 8-spored, bitunicate, long-cylindrical, with a short
furcate pedicel, rounded at the apex. Ascospores 23.5–
29 9 5–7 lm (
x = 27 9 6 lm, n = 40), uni-seriate to
59
overlapping uni-seriate, fusiform, 3-septate, constricted at
the septa, initially hyaline, becoming reddish to yellowishbrown at maturity, conical at the ends, without a
mucilaginous sheath. Asexual morph Undetermined.
Culture characters: Colonies growing on MEA, reaching 4 cm diam. in 21 days at 16 °C, white, dense, moderate
aerial mycelium on the surface, underneath pale yellow,
margins even.
Material examined: UK, Hampshire, Swanick Wood, on
unidentified plant stem, 28 September 2015, E.B. Gareth
Jones, GJ 199 (MFLU 16-1091, holotype), (isotype in
HKAS); ex-type living cultures MFLUCC 15-1118, GAAS).
Notes: Leptosphaeria irregularis is typical of Leptosphaeria in having a peridium of scleroplectenchymatous
cells and 3-septate, reddish to yellowish-brown, fusiform
ascospores. Phylogenies herein show that L. irregularis is a
sister taxon to L. slovacica and can be considered distinct
to other known Leptosphaeria species (Fig. 31). Morphologically L. irregularis is similar to L. slovacica, but differs
in its larger (23.5–29 lm vs. 18–22 lm) ascospores.
Lindgomycetaceae K. Hirayama et al.
The family Lindgomycetaceae was introduced by Hirayama et al. (2010) for the freshwater genus Lindgomyces
(L. breviappendiculatus, L. cinctosporae, L. ingoldianus
and L. rotundatus) and its sister taxon, Massariosphaeria
typhicola. Molecular phylogenetic studies using ribosomal
sequence data from different lineages of Dothideomycetes
also showed that Lindgomycetaceae is a unique lineage
among the Pleosporales (Zhang et al. 2012b; Hyde et al.
2013; Wijayawardene et al. 2014b; Ariyawansa et al.
2015c; Wanasinghe et al. 2015). Currently there are 12
species included the genus Lindgomyces including the
novel taxon introduced below. Multi-gene phylogenetic
analyses also placed Arundellina typhae, Clohesyomyces
aquaticus, Hongkongmyces pedis, Lolia aquatica, Massariosphaeria typhicola, Phyllosticta flevolandica and
Trematosphaeria hydrela in Lindgomycetaceae (AbdelAziz and Abdel-Wahab 2010; Zhang et al. 2012b; Tsang
et al. 2014; this study). Most of the Lindgomycetaceae
members have been recorded from freshwater habitats, but
Hongkongmyces is associated with IgG4-related sclerosing
disease of humans (Tsang et al. 2014).
Arundellina Wanasinghe, E.B.G. Jones & K.D. Hyde, gen.
nov.
Index Fungorum number: IF552132; Facesoffungi
number: FoF02208.
Etymology: Name reflects the Arun River, and town
Arundel, from where the holotype was collected.
Saprobic on dead stem, sheaths and leaves. Sexual
morph Ascomata immersed, solitary, scattered, globose,
dark brown to black, coriaceous, ostiolate. Ostiole
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Fungal Diversity (2016) 80:1–270
b Fig. 38 Vaginatispora fuckelii (HKAS 92495). a Herbarium mate-
rial. b Appearance of ascomata on host. c Vertical section through
ascoma. d Peridium. e–g Asci. h Pseudoparaphyses. i–k Ascospores.
l Ascospore stained in Indian ink. m Germinating ascospore. n Upper
view of culture on PDA. o Lower view of culture on PDA. Scale bars
c = 50 lm, d, e–g, m = 20 lm i–l = 10 lm
papillate, black, smooth, filled with hyaline to pale brown
cells. Peridium thin, with 4–5 layers, outer layer heavily
pigmented, thick-walled, comprising reddish to dark brown
cells of textura angularis, inner layer composed of hyaline
thin-walled cells of textura angularis. Hamathecium comprising numerous, filamentous, branched, septate pseudoparaphyses. Asci 8-spored, bitunicate, fissitunicate,
cylindrical to cylindric-clavate, pedicellate, thick-walled at
the apex, with an ocular chamber. Ascospores overlapping
1–2-seriate, fusiform, slightly curved, widest in the center,
3–4 transversely septate, without vertical septa, constricted
at the septa, initially hyaline, becoming golden-pale brown
at maturity, ends remaining cone-shaped, with pointed
ends, without a mucilaginous sheath. Asexual morph
Undetermined.
Type species: Arundellina typhae Wanasinghe, E.B.G.
Jones & K.D. Hyde
Notes: The genus Arundellina is characterized by
immersed, globose, ascomata, papillate ostiole, a thin
peridium composed of cells of textura angularis, cylindrical to cylindric-clavate asci with a short pedicel and
thick-walled apex and fusiform, golden-pale brown ascospores with 3–4 transverse septa and cone-shape pointed
ends. The globose ascomata, cylindrical to cylindric-clavate asci and fusiform ascospores with 3–4 transverse septa
of Arundellina resemble those of Equiseticola, Galiicola,
Loratospora, Paraleptosphaeria and Phaeosphaeria in
Phaeosphaeriaceae, Neolophiostoma in Halotthiaceae and
Mytilinidion in Mytilinidiaceae. LSU and SSU combined
gene phylogenetic analyses indicate that Arundellina
belongs in Lindgomycetaceae, but is distinct from other
genera in the family (Fig. 34).
Arundellina typhae Wanasinghe, E.B.G. Jones & K.D.
Hyde, sp. nov.
Index Fungorum number: IF552133; Facesoffungi
number: FoF02209, Fig. 35
Etymology: Name reflects the host genus Typha.
Holotype: MFLU 16-1276
Saprobic on submerged stem and sheaths of Typhaceae
sp. Sexual morph Ascomata 250–300 lm high 9 200–
250 lm diam. (
x = 272.1 9 229.5 lm, n = 5), immersed,
solitary, scattered, globose, dark brown to black, coriaceous, ostiolate. Ostiole 70–90 lm high, 40–60 lm diam.
(
x = 80.2 9 51.4 lm, n = 5), papillate, black, smooth,
filled with hyaline to pale brown cells. Peridium 18–22 lm
61
wide at the base, 20–30 lm wide in sides, with 4–5 layers,
outer layer heavily pigmented, thick-walled, comprising
reddish to dark brown cells of textura angularis, inner layer
composed of hyaline, thin-walled cells of textura angularis. Hamathecium comprising numerous, 2.5–3.5 lm
wide, filamentous, branched, septate, pseudoparaphyses.
Asci 110–115 9 15–20 lm (
x = 112.4 9 17.5 lm, n =
20), 8-spored, bitunicate, fissitunicate, cylindrical to
cylindric-clavate, pedicellate, thick-walled at the apex,
with an ocular chamber. Ascospores 30–40 9 5–7 lm
(
x = 34.2 9 6.1 lm, n = 30), overlapping 1–2-seriate,
fusiform, slightly curved, widest in the center, 3–4 transversely septate, without vertical septa, constricted at the
septa, initially hyaline, becoming golden-pale brown at
maturity, ends remaining cone-shaped, with pointed ends,
without a mucilaginous sheath. Asexual morph
Undetermined.
Culture characteristics: Colonies on MEA reaching
2 cm diam. after 30 days at 16 °C, circular, margin
smooth, dirty white at first, greenish dark brown after
4 weeks flat on the surface, without aerial mycelium,
reverse blackish-brown. Hyphae septate branched, hyaline,
thin, smooth-walled.
Material examined: UK, England, Arun River, on dead
submerged stem of Typha sp (Typhaceae), 6 April 2015,
E.B.G. Jones, GJ122 (MFLU 16-1276, holotype); (isotype
in BBH), ex-type culture, MFLUCC 16-0310, MUCL. UK,
Dorset, West Lulworth, Lulworth Cove, 24 February 2015,
on Typha sp., E.B.G. Jones, GJ096 (MFLU 16-1277,
paratype); ex-paratype culture, MFLUCC 16-0309,
MUCL.
Lindgomyces pseudomadisonensis Tak. Takah. & Kaz.
Tanaka, sp. nov.
MycoBank number: MB817598; Facesoffungi number:
FoF02434, Fig. 36
Etymology: Referring to its resemblance to Lindgomyces
madisonensis.
Holotype: HHUF 30513
Saprobic on submerged wood. Sexual morph Ascomata
190–240 lm high, 230–360 lm diam., globose to subglobose, black, scattered to grouped, immersed to erumpent.
Ostiolar neck 90–100 lm long, 40–55 lm wide, papillate,
central. Peridium 50–55 lm thick, composed of an inner
layer of polygonal to subglobose, hyaline to pale brown
cells and an outer layer of brown cells. Pseudoparaphyses
cellular, numerous, 1.5–2.5 lm wide, anastomosed, branched. Asci 92.5–130 9 13.5–17.5 lm (
x = 108.3 9
15.6 lm, n = 8), fissitunicate, clavate, rounded at the apex,
with an apical chamber, with eight overlapping bi-seriate
ascospores. Ascospores 28–41(–45.5) 9 (5.5–)7–10 lm
(
x = 36.3 9 7.9 lm, n = 63), l/w 3.7–6.1 (
x = 4.6,
n = 63), fusiform with acute ends, straight or slightly
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62
curved, with the primary septum almost supramedian
(0.40–)0.42–0.51 (
x = 0.47, n = 62), slightly constricted
at the primary septum, with a broad upper cell, hyaline,
smooth, becoming 3-septate and pale brown with age,
surrounded by an entire gelatinous sheath ca. 2 lm wide.
Asexual morph Undetermined.
Material examined: JAPAN, Aomori, Nishimeya, Seisyu trail, Ooshirosawa River, on submerged dead twigs of
woody plant, 28 August 2010, K. Tanaka et al., KT 2742
(HHUF 30513, holotype); ex-type living culture, MAFF
245610.
Notes: Lindgomyces pseudomadisonensis is morphologically similar to L. madisonensis and phylogeny also
support a close association between them (Fig. 34). However, the former has shorter ascospores (
x = 39 ± 2 9
7 ± 1 lm). In addition, the ascospores of L. pseudomadisonensis have an entire gelatinous sheath. The identities of
ITS sequences between L. pseudomadisonensis and
L. madisonensis are rather low [GenBank KT207819;
Identities = 541/564 (95.9 %), Gaps = 9/564 (1.5 %)].
Lophiostomataceae Sacc.
The family Lophiostomataceae was introduced by
Nitschke (1869) with Lophiostoma macrostomum (Tode)
Ces. & De Not. as the type species (Eriksson 1981;
Mugambi and Huhndorf 2009; Thambugala et al. 2015b).
Most species in this family are widely distributed on twigs,
stems, or bark of various woody plants and herbaceous
plants in terrestrial and aquatic environments (Ellis and
Ellis 1985; Mugambi and Huhndorf 2009; Zhang et al.
2009b; Hirayama and Tanaka 2011; Hyde et al. 2013;
Thambugala et al. 2015b).
Vaginatispora fuckelii (Sacc.) Thambugala, Wanasinghe,
Kaz. Tanaka & K.D. Hyde, Fungal Diversity 74: 242. 2015.
Index Fungorum Number: IF551535; Facesoffungi
number: FoF00829
Basionym: Lophiostoma fuckelii Sacc., Michelia 1(no.
3): 336 (1878)
= Lophiostoma pulveraceum Sacc., Michelia 1: 336,
1878
= Didymosphaeria lophospora Sacc. & Speg., Michelia
1: 376, 1878
= Lophiosphaera mendax Rehm, Ann. Myc. 5: 544,
1907
Saprobic on dead branch of Rosa sp. Sexual morph
Ascomata 145–205 lm high, 140–258 lm diam., solitary,
scattered to clustered, immersed to semi-immersed or
erumpent, globose to subglobose, glabrous, uniloculate,
rarely bi-loculate, papillate, visible as raised, black spots
on host surface. Peridium 18–26 lm wide, thin-walled,
unevenly thickened, two layered; inner layer comprising 3–
5 cell layers of flattened, hyaline cells of textura
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Fig. 39 Phylogram generated from maximum parsimony analysis c
based on combined LSU, SSU and TEF1a sequence data of
Lophiotremataceae. Maximum likelihood (left)/ parsimony (right)
bootstrap support values greater than 50 % are shown above the
nodes. The ex-type strains are in bold and the new species are
indicated in blue. The tree is rooted to Dothidotthia symphoricarpi
and D. aspera
prismatica; outer layer comprising several layers of dark
brown to black cells of textura angularis. Hamathecium
composed of dense, 1.7–2.5 lm wide, filamentous, indistinctly septate, cellular pseudoparaphyses, anastomosing at
the apex, embedded in a hyaline gelatinous matrix. Asci
43–72 9 4.6–6.8 lm (
x = 57 9 5.6 lm, n = 20), 8spored, bitunicate, fissitunicate, cylindric-clavate, short
pedicel with furcate to obtuse ends, apically rounded with
an indistinct ocular chamber. Ascospores 15–23 9 3–
4.7 lm (
x = 19.7 9 3.8 lm, n = 30), overlapping bi-seriate, hyaline, subfusoid, with rounded or obtuse ends,
initially aseptate, becoming 1-septate at maturity, strongly
constricted at the septum, enlarged near the septum at the
upper cell, smooth-walled, guttulate, surrounded by thin
distinctive sheath when immature, invisible at maturity,
bearing appendages at both ends. Asexual morph
Undetermined.
Culture characteristics: Colonies on PDA, 33–35 mm
diam. after 3 weeks, colonies medium dense, irregular, flat,
slightly raised, surface smooth with crenate edge, fluffy to
velvety with smooth aspects, zonate with different sector
yellowish-grey to yellowish-brown at the margin whitishgrey to brownish-grey at the centre; reverse whitish-grey at
margin, yellowish-brown with fimbriate aspect at the
midlle, black at the centre, no pigmentation produced in
media.
Material examined: CHINA, Yunnan Province, Kunming, on dead branch of Rosa sp. (Rosaceae), 9 June 2015,
Wen Jing Li, NI005 (HKAS 92495); living culture
KUMCC 15-0523.
Notes: This is the first record of Vaginatispora fuckelii
on dead branch of Rosa species (Rosaceae) in China. The
current maximum likelihood analysis shows that the new
strain (KUMCC 15-0523) clusters with other Vaginatispora fuckelii strains with low bootstrap support
(Fig. 37). The new strain, however, differs from the others
in asci and ascospores size. Vaginatispora fuckelii has
60–90 9 9–12 lm asci and 15–17 9 4–5 lm ascospores
(Thambugala et al. 2015b), whereas the new strain has
43–72 9 4.6–6.8 lm asci and 15–23 9 3–4.7 lm ascospores. Vaginatispora fuckelii has been recorded from various dead herbaceous twigs including Vitis coignetiae
(Thambugala et al. 2015b) and also occurs on leaves of
Mangifera indica (Wang and Lin 2004). The known distribution of this species is Sweden, UK, Germany,
Fungal Diversity (2016) 80:1–270
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b Fig. 40 Hermatomyces subiculosa (holotype). a Host (decaying
wood). b, c Subiculum, conidiophores and conidia on the host surface,
(1) subiculum (2) conidia. d Infertile hyphae. e Conidiophores.
f Conidiophores, conidiogenous cells and conidia, (3) parts of the
conidiogenous cells, (4) conidiophores. f–h Conidia. l, m 20-day old
colonies on MEA, l from above, m from below. Scale bars
b = 500 lm, c = 200 lm, d–f = 20 lm, g–k = 10 lm
Switzerland, Taiwan and Japan (Wang and Lin 2004,
Thambugala et al., 2015b). Vaginatispora fuckelii is distinct from other Vaginatispora species in having 1-septate,
Fig. 41 Lophiotrema bambusae (MFLU 11-0150, holotype). a Appearance of ascostromata on the host surface. b Section through the
ascostroma. c Section through peridium. d Pseudoparaphyses stained
in cotton blue. e–h Asci. i Ocular chamber stained in Melzer’s
65
guttulate, ascospores with a sheath and appendages at both
ends (Fig. 38).
Lophiotremataceae K. Hiray. & Kaz. Tanaka
The family Lophiotremataceae was introduced by Hirayama and Tanaka (2011) to accommodate a single genus
Lophiotrema, typified by L. nucula (Fr.) Sacc. Lophiotrema
shares morphological characters with Lophiostoma due to
its carbonaceous ascomata with compressed, crest-like
apex and was previously regarded as a synonym of
Lophiostoma (Chesters and Bell 1970; Zhang et al.
reagent. j–l Ascospores. m Ascospores stained in Indian ink. n, o
Culture characteristics (n = from above, o = from below). Scale
bars b = 100 lm, c–h = 20 lm, i = 10 lm, j–m = 5 lm
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66
Fig. 42 Lophiotrema fallopiae. a, b Appearance of ascomata on
substrate. c Ascoma in longitudinal section. d Peridium of ascoma. e,
f Ascus. g Ascus apex. h Pseudoparaphyses. i–k Ascospores.
l Ascospore with gelatinous sheath (in Indian ink). m Germinating
ascospore. n, o Conidiomata in culture. p Conidioma in longitudinal
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section. q Peridium of conidioma. r, s Conidiogenous cells. t, u
Conidia. v Germinating conidium. a–m From HHUF 30506 (holotype); n–v from MAFF 245612 (ex-type culture). Scale bars
a = 1 mm, b, n, o = 250 lm, c, p = 20 lm, d–f, q = 10 lm, g–m,
r–v = 5 lm
Fungal Diversity (2016) 80:1–270
67
Fig. 43 Phylogram generated from RAxML analysis of combined
LSU, SSU and TEF sequence data of Massariaceae. Maximum
Likelihood (ML) bootstrap support values greater than 50 % and
Bayesian posterior probabilities (PP) greater than 0.95 are shown
above and below branches. The ex-type strains are in bold and the
new isolate is in blue. The tree is rooted with Acrospermum
gramineum and A. compressum
2009b, 2012b; Hirayama and Tanaka 2011; Hyde et al.
2013). However, these two genera can be distinguished by
their peridial structure and shape of ascus (Zhang et al.
2009a, b; Hirayama and Tanaka 2011, Hyde et al. 2013).
The peridial structure of Lophiotrema is usually thin, of
equal thickness, composed of cells of textura angularis to
textura globulosa and asci are often oblong to cylindrical
with hyaline ascospores (Zhang et al. 2009a, b; Hirayama
and Tanaka 2011; Hyde et al. 2013). In Lophiostoma the
peridium is of unequal thickness and is usually broader
near the base, while asci are mostly clavate (Zhang et al.
2009b; Hirayama and Tanaka 2011). Doilom et al. (2016)
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68
Fig. 44 Neomassaria fabacearum (holotype). a, b Appearance of
ascomata on substrate. c Section through ascoma. d Ostiole. e Peridium. f Pseudoparaphyses. g, h Asci. i–k Ascospores. l Ascospores
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Fungal Diversity (2016) 80:1–270
surrounded by hyaline gelatinous sheath in Indian ink. Scale bars
a = 500 lm, b = 200 lm, c, g–h = 50 lm, d = 20 lm, e, f, i–
l = 10 lm
Fungal Diversity (2016) 80:1–270
69
Fig. 45 Phylogram generated from maximum likelihood analysis
based on combined LSU, ITS, SSU, and TEF1-a sequence data from
species of Massarinaceae. Maximum likelihood bootstrap support
values greater than 50 % and Bayesian posterior probabilities greater
than 0.90 are shown above the nodes. Ex-type strains are in bold and
the new isolate is in blue. The tree is rooted with Periconia digitata
accepted two other genera, Hermatomyces and Aquasubmersa in the Lophiotremataceae as in the phylogenetic analyses these two genera are related to
Lophiotrema.
Species in Lophiotrema are not well-studied and lack
modern taxonomic treatments and molecular data. There
are 159 epithets recorded in Index Fungorum (2016), but
only 96 sequences from six species in GenBank. In this
study, we introduce two new taxa, Lophiotrema bambusae
and L. fallopiae. We also provide a new combination for
Scyphostroma mirum under Hermatomyces mirum and
introduce H. subiculosa sp. nov.
= Subicularium reticulatum M.L. Farr & Goos, Mem.
N. Y. bot. Gdn 49: 66 (1989)
Index Fungorum number: IF552284; Facesoffungi
number: FoF02435
Notes: Farr and Goos (1989) introduced a monotypic
genus, Subicularium, with S. reticulatum M.L. Farr & Goos
as type, and provisionally placed the genus in the order
Agonomycetales of Deuteromycotina. The Dictionary of
Fungi (Kirk et al. 2008) recorded S. reticulatum under an
older name Scyphostroma mirum Starbäck (Starbäck 1899),
apparently based on their similar morphology, while Seifert
et al. (2011) treated them differently. Incidentally, the
genus Scyphostroma Starbäck, typified by Sc. mirum
Starbäck, was found to be wrongly described by the original author (Starbäck 1899). The description ran as follows:
‘‘Subiculum dark-brown, with reticulately branched
hyphae, forming a cup-like nest with a distinct, thick
Hermatomyces mirum (Starbäck) C.G. Lin, D.J. Bhat,
Yong Wang bis & K.D. Hyde, comb. nov.
Basionym: Scyphostroma mirum Starbäck, Bih. K.
svenska Vetensk Akad. Handl., Afd. 3 25(no. 1): 23 (1899)
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b Fig. 46 Stagonospora forlicesenensis (holotype). a Appearance of
conidiomata on host surface. b Conidioma on host surface. c Vertical
section of conidioma. d Basal mycelium. e Setae at side of conidioma.
f Pycnidial walls. g–i Developing stages of conidia. j, l Conidia with
campanulate appendage. k Conidia stained with cotton blue reagent.
m, n Culture characters on PDA. Scale bars b = 200 lm,
c = 100 lm, d, e, h, i, j = 20 lm, f = 50 lm, g, j, k = 10 lm
Fig. 47 Phylogenetic tree generated by maximum likelihood
(RAxML) analysis of combined LSU, SSU, RPB2 and EF-1a
sequence data from species of Melanommataceae. Bootstrap support
values for maximum likelihood equal or greater than 50 % are given
above the nodes. Bayesian posterior probabilities equal or greater than
71
margin. Perithecia minute, sphaerical, with distinct stalk,
perched in the dense stromatic cup’’. Starbäck (1899)
mistook the conidia for perithecia. The species description
indicated that the cup-like conidiomata measured
0.5–4 mm diam. and conidia (wrongly referred as
0.90 are given below the nodes. Ex-type strains, reference strains and
new isolates are in bold. Newly generated sequences are in blue. The
tree is rooted with Hysterium angustatum CBS 123334 and
H. angustatum
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Fig. 48 Bertiella ellipsoidea (holotype). a Herbarium material.
b Ascomata on wood. c Ascoma on wood. d, e Cross section of an
ascoma. f Vertical section of the ascoma at margin. g Septate hairs.
h Aseptate, branched pseudoparaphyses. i–l Cylindrical asci. m Apical
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apex. n–q Ovoid ascospores. Scale bars b = 500 lm, c, h = 100 lm,
d = 400 lm, e = 200 lm, f = 70 lm, g = 50 lm, i–l = 40 lm,
m = 25 lm, n–q = 10 lm
Fungal Diversity (2016) 80:1–270
‘perithecia’) measured 40–50 lm diam. It is clear that
Hermatomyces is the same as Scyphostroma.
Although Scyphostroma is the first name for the genus,
we preferred to use Hermatomyces because the latter name
is presently in use and the earliest name was wrongly
described in the literature and has never been referred to
any taxonomic discourses of the genus complex. Accordingly, in this study, we propose the synonymy of Scyphostroma mirum and Subicularium reticulatum, under the
new combination Hermatomyces mirum, based on their
morphological similarities and phylogenetic analysis
(Fig. 39).
Hermatomyces subiculosa C.G. Lin, D.J. Bhat, Yong
Wang bis & K.D. Hyde, sp. nov.
Index Fungorum number: IF552285, Facesoffungi
number: FoF02436, Fig. 40
Etymology: Referring to the conspicuous subiculum on
the host surface.
Holotype: MFLU 16-1300.
Saprobic on decaying wood. Sexual morph Undetermined. Asexual morph Colonies subiculate on natural
substrate, superficial, effuse, floccose, greyish to dark
brown. Mycelium superficial, composed of septate, flexuous, repeatedly branched, pale to brown, 2–4 lm wide
infertile hyphae. Conidiophores micronematous or semimacronematous, mononematous, brown, smooth, confined
to the center and surrounded by infertile hyphae, 3–5.5 lm
wide. Conidiogenous cells holoblastic, subhyaline, 3–
5.5 lm wide. Conidia solitary, dry, acrogenous, muriform,
deeply constricted at septa, globose, oblong, ellipsoidal,
pyriform, verruculose, subhyaline to pale brown when
young, dark brown when mature, 15–35 lm
(
x = 26.23 lm, n = 39) long, 18–30 lm (
x = 24.28 lm,
n = 39) wide at the broadest part, sometimes with part of
conidiogenous cell attached.
Culture characteristics: Colonies on MEA attaining a
diam. of 2.5–3.5 cm at room temperature (25 °C) in
3 weeks, effuse, hairy, grey above, pale brown at the
margins, dark brown at the center from below.
Material examined: THAILAND, Chiang Rai, Mae Sai
District, Ang Kep Nam Wat Tham Khao Hin Phayanak
(Wat Tham Sao Hin Payanak), 20°190 16.5800 –
20°190 30.1200 N, 99°510 40.7200 –99°510 54.5000 E, on decaying
wood, 19 June 2015, Chuan-Gen Lin, WTSP 1-1 (MFLU
16-1300, holotype; HKAS 95052, isotype), ex-type living
culture MFLUCC 15-0843.
Notes: We found that our new species forms a separate
well-supported clade within the genus Hermatomyces,
sister to H. thailandica (MFLUCC 14-1143, MFLUCC
14-1145 and MFLUCC 14-1144) and H. tectonae
(MFLUCC 14-1140 and MFLUCC 14-1141) in the family
Lophiotremataceae. The new species is morphologically
73
similar to H. mirum in having a conspicuous subiculum and
the conidia of these two species are hyaline to pale brown
when young, and dark brown when mature. However, the
conidia in our new species (18–30 lm) are smaller than
H. mirum (40–50 lm) verruculose and, in addition, the
subiculum of H. mirum are more or less disc-shaped, with
individual discs 500–4000 lm in diam., while the subiculum in the new species are effuse on the natural substrate.
Therefore, based on differences in morphology and
molecular data, we introduce the new species, Hermatomyces subiculosa.
Lophiotrema bambusae Phookamsak, S.C. Karunarathana
& K.D. Hyde, sp. nov.
Index Fungorum number: IF552203; Facesoffungi
number: FoF02257, Fig. 41
Etymology: The specific epithet ‘‘bambusae’’ refers to
the host.
Holotype: MFLU 11-0150.
Saprobic on bamboo. Sexual morph Ascostromata 140–
270 lm high, 250–370 lm diam., dark brown to black,
scattered, gregarious, immersed to semi-immersed in host
cortex, raised, conical to quadrilateral, or irregular in
shape, uni-loculate, glabrous, coriaceous, ostioles central
or lateral, with rim-like opening. Peridium 20–50 lm wide,
thin- to thick-walled, of unequal thickness, slightly thick at
the sides towards apex, composed of several layers of
small, dark brown to black, pseudoparenchymatous cells,
with host cells plus fungal tissue, arranged in a textura
angularis to textura epidermoidea. Hamathecium composed of dense, 2–3 lm wide, cellular pseudoparaphyses,
distinctly septate, anastomosing among the asci, embedded
in a hyaline gelatinous matrix. Asci 65–100 9 8–10 lm
(
x = 83.3 9 8.7 lm, n = 20), 8-spored, bitunicate, fissitunicate, cylindrical, short pedicellate, apically rounded
with an ocular chamber. Ascospores 18–25 9 4–5 lm
(
x = 21.3 9 4.4 lm, n = 25), overlapping bi-seriate,
hyaline, fusiform, with rounded ends, 1-septate, rarely 3–5septate, upper cell larger that lower cell, constricted at the
central septum, smooth-walled, with guttules, surrounded
by a mucilaginous sheath. Asexual morph Undetermined.
Culture characteristics: Colonies on PDA reaching
39–45 mm diam. after 4 weeks at 25–30 °C; colony from
above, dark green, with black concentric ring at the margin,
black in the centre, with white turfs; from below, dark
green, with black concentric ring at the margin, dark green
to black at the centre, slightly radiating; medium dense,
irregular, flattened, slightly raised at the middle, edge
undulate, with entire margin, surface smooth, slightly
rough, with white tufts, or black granules, woolly; not
producing pigmentation in agar.
Material examined: THAILAND, Chiang Rai, Mae Fah
Luang District, Doi Tung, on dead stem of bamboo, 28
123
74
April 2010, S.C. Karunarathana, RP0030 (MFLU 11-0150,
holotype), ex-type living cultures, MFLUCC 10-0558,
BBC.
Notes: Lophiotrema bambusae shares a size range of
asci and ascospores, including number of ascospore septa with
many Lophiotrema species, such as L. alpinum (Ellis &
Everh.) M.E. Barr, L. arundinariae Rehm, L. culmifragum
Speg., L. incisum Ellis & Everh., L. paspalicola Speg. and
L. radicans (Ellis & Everh.) Sacc. However, the concept of
Lophiotrema was unclear, when many of these species were
introduced or transferred to the genus and they need
restudying. The ascospore septation of L. bambusae is typically 1-septate (rarely 3–5-septate), similar to L. alpinum,
L. culmifragum and L. incisum, while, L. arundinariae,
L. paspalicola and L. radicans usually have 3-septate ascospores. Nevertheless, L. bambusae differs from L. alpinum and
L. incisum based on host, with L. bambusae collected from
bamboo, and the others collected from conifer and Ribes.
Lophiotrema bambusae is most similar to L. culmifragum
given its host and habitat. However, L. bambusae has slightly
larger asci and mature ascospores have more septa. Phylogenetic analyses show that L. bambusae is related to L. lignicola
and distinct from other taxa in Lophiotrema (Fig. 39).
Therefore, the new species is established.
Lophiotrema fallopiae A. Hashim. & Kaz. Tanaka, sp.
nov.
MycoBank number: MB817597; Facesoffungi number:
FoF02475, Fig. 42
Etymology: Referring to the generic name of host plant.
Holotype: HHUF 30506.
Saprobic on dead stem of Fallopia japonica. Sexual
morph Ascomata 200–300 lm high, 180–250 lm diam.,
scattered, immersed, erumpent at the neck, subglobose in
section. Ostiolar neck up to 40 lm high, compressed, composed of carbonaceous, black, thick-walled cells, without
clypeus. Peridium 17.5–25.5 lm thick, 5–8 layers, composed of polygonal to elongate, thin-walled, 8–13 9 2.5–
3 lm cells. Hamathecium comprises branched pseudoparaphyses numerous, trabeculate, 1–1.5 lm wide, septate. Asci
(77–)93–125 9 (5.5–)7–9 lm
(
x = 105.8 9 7.6 lm,
n = 10), 8-spored, numerous, bitunicate, fissitunicate,
cylindrical, with a short stipe (5–11 lm long, x = 7.4 lm,
n = 10), apically rounded with an ocular chamber. Ascospores
19–24 9 4.5–6 lm
(
x = 21.7 9 5.2 lm,
n = 50), l/w 3.7–5.3 (
x = 4.2, n = 50), fusiform with
rounded ends, straight, 1-septate, constricted, with a primary
septum nearly median [0.45–0.55, x = 0.51, n = 50], hyaline, smooth, guttulate when young, with an entire gelatinous
sheath (2.5–6 lm wide at sides). Asexual morph
123
Fungal Diversity (2016) 80:1–270
Coelomycetes. Conidiomata pycnidial, globose to subglobose, up to 200 lm high in section, 120–180 lm diam.,
scattered, semi-immersed, solitary, black. Peridium 10–
14 lm wide; outer layers composed of 7.5–10.5 9 4–
7.5 lm, subglobose, brown cells; inner layers composed of
5–7.5 9 0.5–1.5 lm, ‘‘porrecta’’, hyaline cells. Conidiophores reduced. Conidiogenous cells enteroblastic, phialidic, 7–9 9 2–4 lm, lageniform, hyaline, smooth. Conidia
ellipsoidal with rounded ends, 2–3.5 9 1–2 lm
(
x = 2.7 9 1.5 lm, n = 60), l/w 1.2–2.5 (
x = 1.8,
n = 60), hyaline, aseptate, smooth.
Culture characteristics: Colonies on PDA attaining
12–16 mm diam. within 21 days at 20 °C in the dark.
Material examined: JAPAN, Aomori, Hirakawa, Kuzukawa, near Aseishi River, on dead stem of Fallopia japonica,
5 September 2010, K. Tanaka, KT 2748 (HHUF 30506,
holotype); ex-holotype living culture, MAFF 245612.
Notes: The characters of L. fallopiae fit into the generic
concept of Lophiotrema in having compressed ostiolar
necks, a peridium composed of textura angularis, and
cylindrical asci with short stipes. This species is similar to
L. nucula, the type species of Lophiotrema, but L. nucula
has shorter ascospores [17–21(–25) 9 (4–)5–6.5 lm;
Zhang et al. 2009a]. Lophiotrema fallopiae is phylogenetically close to L. vagabundum (Fig. 39), but L. vagabundum has slightly narrower ascospores (20–26 9 4–5.5 lm,
l/w 4.3–5.9; Tanaka and Harada 2003).
Massariaceae Nitschke
The family Massariaceae was introduced by Nitschke
(1869) to accommodate the genus Massaria with M. inquinans (Tode) De Not. as the type species. The family is
characterized by immersed globose, subglobose to pyriform, ascomata, with a thin-walled peridium comprising
cells of textura angularis, oblong to cylindrical asci with a
wide ocular chamber and refractive ring, and large, oblong
to ellipsoidal ascospores, surrounded by a gelatinous sheath
(Hyde et al. 2013). Seven new species were introduce in
Massaria with molecular analyses of combined LSU, SSU,
RPB2 and TEF1 sequence data. In this study, the new
genus Neomassaria is introduced based on molecular and
morphological comparison with descriptions and illustrations. The phylogenetic tree, based on combined LSU, SSU
and TEF1 sequence data for the new taxon is presented in
Fig. 43.
Neomassaria Mapook, Camporesi & K.D. Hyde, gen. nov.
Index Fungorum number: IF552273; Facesoffungi
number: FoF02437
Etymology: The generic epithet refers to a new genus in
the family Massariaceae.
Fungal Diversity (2016) 80:1–270
75
Neotrematosphaeria biappendiculata KTC 1124
Coelodictyosporium muriforme MFLUCC 13-0351
Lophiopoacea paramacrostoma MFLUCC 11-0463
Sigarispora ravennica MFLUCC 14-0005
Lophiohelichrysum helichrysi MFLUCC 15-0701
Guttulispora crataegi MFLUCC 13-0442
Lophiostomataceae
Platystomum crataegi MFLUCC 14-0925e
Lophiostoma compressum IFRD 2014
Lophiostoma macrostomum KT508
Biappendiculispora japonica MAFF 239452
79
Capulatispora sagittiformis JCM 15100
93
Platystomum scabridisporum BCC 22835
Trematosphaeria terricola SC 12
85
96
Paucispora quadrispora MAFF 239455
Dimorphiopsis brachystegiae CPC 22679
Vaginatispora aquatic MFLUCC 11-0083
98 Angustimassarina populi MFLUCC 13-0034
75 Massarina corticola CBS 154.93
Amoroslaceae
100 Angustimassarina quercicola MFLUCC 14-0506
Angustimassarina acerina MFLUCC 14-0505
Exosporium stylobatum CBS 160.30
Camarographium carpini CBS 128781
99 Preussia lignicola CBS 264.69
94
Sporormia lignicola CBS 363.69
Preussia minima AFTOL ID1256
90 Preussia funiculate 2577
100
Sporormiaceae
Preussia funiculate CBS 659.74
Preussia terricola AFTOL ID 282
Westerdykella
dispersa
CBS
50875
74
Westerdykella ornate CBS 379 55
98
97 Eremodothis angulate CBS 610.74
Westerdykella cylindrical CBS 454.72
Sporormia fimetaria Gr.81.194
100 Decaisnella formosa BCC 25617
97
Decaisnella formosa BCC 25616
100
Lignosphaeria thailandica MFLUCC 11-0376
Lignosphaeria fusispora MFLUCC 11-0377
100
Berkleasmium nigroapicale BCC 8220
Bahusandhika indica GUFCC 18001
100
Misturatosphaeria kenyensis GKM L100Na
Pseudoaurantiascoma kenyense GKM 1195
95
Misturatosphaeria minima ANM 933
Misturatosphaeria minima GKM 169N
Curreya grandicipis CBS 114272
Misturatosphaeria claviformis GKM 1210
Macrodiplodiopsis desmazieri MFLUCC 12-0088
6
Floricolaceae
Floricola striata JK 5678I
Floricola striata JK 5603K
Misturatosphaeria tennesseensis ANM 911
Pseudomisturatosphaeria cruciformis SMH 5151
Floricola viticola IT 2178
Misturatosphaeria aurantonotata GKM 1238
Misturatosphaeria radicans ATCC 42522
Ramusculicola thailandica MFLUCC 13-0284
Neomassarina thailandica MFLUCC 10 0552
93
Pontoporeia biturbinata BBH 23338
4x
100
Halotthia posidoniae BBH 22481
Mauritiana rhizophorae BCC 28866
Halotthiaceae
Sulcosporium thailandica MFLUCC 12-0004
Neolophiostoma pigmentatum MFLUCC 10-0129
Brunneoclavispora bambusae MFLUCC 11-0177
Roussoellopsis tosaensis KT 1659
Roussoella pustulans KT 1709
Roussoella hysterioides HH 26988
95 Arthopyrenia salicis CBS 368.94
Roussoellaceae
Arthopyrenia salicis CBMAI1330
74
Roussoella angustior MFLUCC 15-0186
Roussoella magnatum MFLUCC 15-0185
Neoroussoella bambusae MFLUCC 11-0124
Sporidesmium_australiense
100 Torula herbarum CBS 111855
98
Torula herbarum CBS 379.58
Torula herbarum CBS 220.69
Torulaceae
100
Dendryphion europaeum CPC 22943
Dendryphion europaeum CPC 23231
100
Parameliola acaciae MFLU 15-0378
Parameliola dimocarpi MFLU15-0045
100 Pseudocoleodictyospora tectonae MFLUCC 12-0387
72
Pseudocoleodictyospora_tectonae MFLUCC 12-0385
Pseudocoleodictyospora thailandica MFLUCC 12-0565
100
Pseudocoleodictyospora sukhothaiensis MFLUCC 12-0554 Pseudocoleodictyosporaceae
Subglobosporium tectonae MFLUCC 12-0393
100
Subglobosporium tectonae MFLUCC 12-0390
100
98
100
97
98
Fig. 49 RAxML Maximum lkelihood phylogenetic tree based on a
LSU and SSU sequence data from species of order Pleosporales.
Maximum likelihood bootstrap support values greater than 50 % are
shown on near the nodes. Some branches were shortened to fit the
page—these are indicated by two diagonal lines with the number of
times a branch was shortened indicated next to the lines. The new
isolates are in red. The tree is rooted with Hysterium angustatum
123
76
Fungal Diversity (2016) 80:1–270
84
Occultibambusa bambusae MFLUCC 13-0855
Occultibambusa bambusae MFLUCC 11-0394
Occultibambusa chiangraiensis MFLUCC 16-0380
Occultibambusa aquatica MFLUCC11-0006
Occultibambusa pustula MFLUCC 11-0502
Occultibambusaceae
Versicolorisporium triseptatum JCM 14775
Occultibambusa fusispora MFLUCC 11-0127
100
Seriascoma didymospora MFLUCC 11-0179
Seriascoma didymospora MFLUCC 11-0194
Neooccultibambusa chiangraiensis MFLUCC 12-0559
94 _Biatriospora mackinnonii CBS 674.75
100
Nigrograna mackinnonii E5202H
Biatriosporaceae
Biatriospora marina CY 1228
100
Paradictyoarthriinium diffractum MFLUCC 12-0557
Paradictyoarthriiniaceae
Paradictyoarthriinium tectonicola MFLUCC 13-465
Stagonospora pseudocaricis HKUCC 10833
Neottiosporina paspali CBS 331.37
Corynespora leucadendri CBS 135133
Massarinaceae
Corynespora olivacea CBS 114450
Massarina eburnea CBS 473.64
Pseudodidymosphaeria phlei MFLUCC14-1061
94
Deniquelata barringtoniae MFLUCC 11-0422
Kalmusia brevispora KT 1466
85
Montagnula aloes CPC 19671
Didymosphaeriaceae
100 Pseudocamarosporium pini MFLUCC 14-1091
Didymosphaeria rubi-ulmifolii MFLUCC 14-0023
96
Bambusicola splendida MFLUCC 11-0439
Bambusicolaceae
Bambusicola massarinia MFLUCC 11-0389
100
Latorua caligans CBS 576.65
Latoruaceae
Latorua grootfonteinensis CBS 369.72
86
Katumotoa bambusicola KT 1517a
Lentitheciaceae
Lentithecium fluviatile CBS 122367
Pseudomonodictys tectonae MFLUCC 12-0552
79
Trematosphaeria pertusa CBS 122371
Trematosphaeriaceae
Falciformispora lignatilis BCC 21118
Macrodiplodiopsis desmazieri CPC 24971
99 Macrodiplodiopsis desmazieri CBS 125026
Macrodiplodiopsisceae
Macrodiplodiopsis desmazieri CPC 24648
100
Moroshperia ramunculicola BCC 18404
85
Moroshperia ramunculicola BCC 18405
Moroshperiaceae
Helicascus nypae BCC 36752
Longiostiolum tectonae MFLUCC 12 0562
Thyridaria rubronotata CBS 385.39
Thyridariaceae
Massariosphaeria phaeospora CBS 611.86
100
Halojulella avicenniaea BCC 18422
Halojulellaceae
Halojulella avicenniae BCC 20173
100 Cucurbitaria berberidis CBS 363.93
82
Cucurbitaria berberidis CBS 394.84
Cucurbitariaceae
Pyrenochaeta nobilis CBS 407.76
89 Camarosporium quaternatum DQ377884
Camarosporium aloes CPC 21572
Phaeosphaeria oryzae CBS 110110
94
79
Ophiosphaerella herpotricha AFTOL-ID 1569
Phaeosphaeriaceae
Paraphoma radicina CBS 111.79
100 Neophaeosphaeria filamentosa CBS 102203
100
Neophaeosphaeria filamentosa CBS 102202
Neophaeosphaeriaceae
Neophaeosphaeria agaves CPC 21264
83
Dothidotthia symphoricarpi CPC 12929
Dothidotthiaceae
Dothidotthia aspera CPC 12933
100 Coniothyrium palmarum CBS 758.73
Coniothyriaceae
Coniothyrium palmarum CBS 400.71
100 Alternariaster helianthi CBS 134020
Leptosphaeriaceae
Alternariaster helianthi CBS 327.69
Leptosphaeria maculans AFTOL-ID 277
75
Pleospora herbarum CBS 191.86
100
Alternaria alternata AFTOL-ID 1610
Pleosporaceae
Pyrenophora phaeocomes AFTOL-ID 283
72
Phoma exigua CBS 431.74
97
Didymellaceae
Leptosphaerulina australis CBS 317.83
Didymella exigua CBS 183.55
100 Ascocylindrica marina MD6011
Ascocylindrica mari MD6012
97
Corynespora smithii CABI 5649b
Corynespora cassiicola CBS 100822
72
Beverwykella pulmonaria CBS 283.53
100
Melanomma pulvis-pyrius CBS 371.75
Melanommataceae
Byssosphaeria jamaicana SMH 1403
77
Lindgomyces ingoldianus ATCC 200398
73
Lindgomyces rotundatus KH 114
Lindgomycetaceae
91
Massariosphaeria typhicola MAFF 239218
Lolia aquatic MF644
97 Anguillospora longissimi CCMF 10304
Anguillospora longissima CS869-1D
Amniculicola lignicola Ying01
Anguillospora longissima F00980
Amniculicola parva CBS 123092
Amniculicolaceae
100
Spirosphaera cupreorufescens A20
Amniculicola immersa CBS 123083
100
Repetophragma ontariense HKUCC 10830
96
Massariosphaeria grandispora CBS 613 86
70
Pseudomassariosphaeria bromicola MFLUCC 15-0031
Murispora rubicunda IFRD 2017
Hysterium angustatum CBS 236.34
Out group
87
98
0.05
Fig. 49 continued
123
Fungal Diversity (2016) 80:1–270
Saprobic on dead branches. Sexual morph Ascomata
immersed, solitary or scattered, coriaceous, globose to subglobose, brown to dark brown. Ostiole central. Peridium
comprising light brown cells of textura angularis. Hamathecium comprising cylindrical to filiform, septate, branched,
pseudoparaphyses. Asci 8-spored, bitunicate, oblong to cylindrical, short pedicellate, with wide ocular chamber. Ascospores
overlapping 1–2-seriate, hyaline, ellipsoid to broadly fusiform,
1-septate, constricted at the septa, surrounded by gelatinous
sheath. Asexual morph Undetermined.
Type species: Neomassaria fabacearum Mapook,
Camporesi & K.D. Hyde, sp. nov.
Notes: Neomassaria fabacearum was collected from a dead
branch of Hippocrepis emerus (L.) Lassen. Molecular data
places Neomassaria in the family Massariaceae with high
bootstrap support (91, ML with 0.99, PP) along with other
species of Massaria. However, Neomassaria is distinct in
morphology of ascomata, asci and ascospores. Phylogenies
also reveal that Neomassaria can be considered as distinct
genus given that it did not cluster with any Massaria species.
Instead it is basal to them with high support (Fig. 43).
Neomassaria fabacearum Mapook, Camporesi & K.D.
Hyde, sp. nov.
Index Fungorum number: IF552274; Facesoffungi
number: FoF02438, Fig. 44
Etymology: Name reflects the host family Fabaceae,
from which this holotype was collected.
Holotype: MFLU 16-1875.
Saprobic on a dead branch of Hippocrepis emerus.
Sexual morph Ascomata (150–)200–220 lm high 9
(100–)130–150 lm diam. (
x = 190 9 130 lm, n = 5),
immersed, solitary or scattered, coriaceous, globose to
subglobose, brown to dark brown, Ostiole central. Peridium 10–20 lm wide, comprising light brown cells of textura angularis. Hamathecium comprising 1–2 lm wide,
cylindrical to filiform, septate, branched, pseudoparaphyses. Asci 65–75(–85) 9 10–15 lm (
x = 75 9 12 lm,
n = 5), 8-spored, bitunicate, oblong to cylindrical, short
pedicellate, with wide ocular chamber. Ascospores 18–
20 9 (4–)5–6 lm (
x = 19 9 5 lm, n = 15), overlapping
1–2-seriate, hyaline, ellipsoid to broadly fusiform, 1-septate, constricted at the septum, surrounded by hyaline
gelatinous sheath observed clearly when mounted in Indian
ink. Asexual morph Undetermined.
Material examined: ITALY, Forlı̀-Cesena, CusercoliCivitella di Romagna, on dead branch of Hippocrepis
emerus (L.) Lassen (Fabaceae), 11 September 2014, E.
Camporesi (MFLU 16-1875, holotype), ex-type culture
MFLUCC 14-1117, (isotype as HKAS 95078 in HKAS).
77
Massarinaceae Munk
The family Massarinaceae was introduced by Munk
(1956) and is typified by Massarina with M. eburnea (Tul.
& C. Tul.) Sacc. as the type species. Currently Byssothecium, Corynespora, Helminthosporium, Massarina, Pseudodidymosphaeria, Pseudosplanchnonema, Stagonospora,
and Suttonomyces are included in Massarinaceae (Chethana et al. 2015; Tanaka et al. 2015; Thambugala et al.
2015a; Fig. 45). Corynespora leucadendri Quaedvl. et al.
and C. olivacea (Wallr.) M.B. Ellis cluster within Massarinaceae, however the generic type of Corynespora,
C. cassiicola (Berk. & M.A. Curtis) C.T. Wei clusters in
Corynesporaceae, which is a distinct family. Therefore,
these two Corynespora species should be renamed. In this
study we introduce a new species of Stagonospora.
Stagonospora (Sacc.) Sacc.
Stagonospora is typified by S. paludosa (Sacc. & Speg.)
Sacc., a species from Carex pseudocyparus. Quaedvlieg
et al. (2013), reassembled septoria-like genera and introduced Stagonospora sensu stricto in Massarinaceae
because of its pycnidial conidiomata, immersed, globose,
ostiolate, conidiophores reduced to conidiogenous cells,
with holoblastic, percurrent proliferations, and doliiform,
cylindrical to ellipsoid, hyaline, guttulate conidia. Tanaka
et al. (2015) revised Massarinaceae and included twelve
species in Stagonospora.
Stagonospora forlicesenensis Phukhamsakda, Camporesi
& K.D. Hyde, sp. nov.
Index Fungorum number: IF552238; Facesoffungi
number: FoF02384, Fig. 46
Etymology: In reference to the location where the fungus
was collected.
Holotype: MFLU 16-1337
Saprobic on dead branches of Phragmites australis
(Cav.) Trin. ex Steud. Sexual morph Undetermined.
Asexual morph Conidiomata 70–210 lm high 9 70–
160 lm diam. (
x = 141 9 117 lm, n = 10), pycnidial,
solitary, uniloculate, scattered, immersed in the host, dark
brown to black, globose, containing basal mycelium, ostiole central. Ostiole 22–72 lm high 9 27–62 lm diam.
(
x = 53 9 46 lm, n = 5), papillate, dark brown. Setae
48 9 3 lm, on the side of the pycnidial walls, acicular,
curved, constricted at the septate, light brown to brown.
Pycnidial wall 8–28 lm (–31 lm at apex), composed of 4–
5 layers of brown-walled cells of textura angularis, two
hyaline inner layers, lining bearing conidiogenous cells.
Conidiophores reduced to conidiogenous cells with one
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b Fig. 50 Occultibambusa aquatica (holotype). a Appearance of
ascomata on host surface. b Vertical section through ascoma.
c Peridium. d–f Asci. g, h Pseudoparaphyses. i, j Ascospores.
k Ascospore stained with Indian ink. l Germinating ascospore. m, n
Culture characters on PDA (m = from above, n = from below).
Scale bars b = 100 lm, c = 40 lm, d, e, l = 20 lm, f = 30 lm, g,
h, i–k = 10 lm
supporting cell. Conidiogenous cells 4–10 9 2–6 lm,
(
x = 7 9 4 lm, n = 30), holoblastic, determinate, discrete, ampulliform to clavate, hyaline, smooth, formed
from the inner cells of the pycnidial wall. Conidia 25–
34 9 6–11 lm (
x = 30 9 8 lm, n = 50), broad fusiform
to oblong, obtuse at both ends, thick-walled, with minute
and large guttules in each cell, 3 transverse eusepta, rarely
2-septate, sometimes slightly constricted at the septa,
hyaline; apical appendage, infundibuliform to campanulate-like, gelatinous, 7–8 lm wide.
Culture characteristics: Colonies on PDA, reaching
90 mm diam. after 14 days at 16 °C, colonies covering surface, sparse, cream, with abundant aerial mycelium, margins
lobate; reverse white at the edges, cream at the center, radiating, circular, flattened, margin rough, not pigmented.
Material examined: ITALY, Province of Forlı̀-Cesena,
Pian di Spino—Meldola, on dead and stem of Phragmites
australis (Poaceae), 22 December 2014, E. Camporesi, IT
2306 (MFLU 16-1337, holotype), isotype in HKAS
94613, ex-type living culture, MFLUCC 15-0054,
KUMCC 16-0028.
Notes: Based on a morphological comparison, Stagonospora forlicesenensis is similar to Stagonospora (Neottiosporina) paspali G.F. Atk. B. Sutton & Alcorn (CBS
331.37), a species introduced from Paspalum laeve
Michaux (Poaceae). In our study, sequence data also
reveals a close phylogenetic affinity of Stagonospora forlicesenensis to Stagonospora (Neottiosporina) paspali
(Fig. 45). Therefore we compare the morphology with
Neottiosporina. Sutton (1980) illustrated seven species of
Neottiosporina. Neottiosporina australiensis B. Sutton &
Alcorn and N. clavata B. Sutton are also reported from
Phragmites australis (Poaceae) (Sutton and Alcorn 1974;
Sutton 1980). They are phenotypically similar in having
thin pycnidial walls, central ostioles, and holoblastic,
determinate, 3-septate, hyaline conidia with infundibuliform apical appendage (Sutton and Alcorn 1974; Sutton
1980). Stagonospora forlicesenensis differs from N. australiensis and N. clavata in having immersed and smaller
conidiomata, with brown setae on the pycnidial walls, and
conidia which are obtuse at both ends. Therefore Tanaka
et al. (2015) suggested these species should be treated
under Stagonospora based on morphology and phylogenetic analysis. However, sequence data for the type species
are not available and therefore the position remains
unconfirmed.
79
Melanommataceae G. Winter
Tian et al. (2015) accepted 20 genera in this family
based on morphology and phylogenetic analyses. In this
paper, we introduce a new species of Bertiella, and an
updated phylogenetic tree for the family (Fig. 47).
Bertiella (Sacc.) Sacc. & P. Syd.
The genus Bertiella is typified by B. macrospora (Sacc.)
Sacc. & Traverso. Previously this genus was assigned to
Massarina (as M. macrospora (Sacc.) O.E. Erikss. & J.Z.
Yue) (Lumbsch and Huhndorf 2010). However, recent
phylogenetic studies support its placement within Melanommataceae close to Byssosphaeria (Tian et al. 2015).
Taxa are mainly saprobes on woody hosts and characterized by black, superficial, subglobose ascomata with carbonaceous peridium, composed of few layers of pigmented
cells (Tian et al. 2015).
Bertiella ellipsoidea Ekanayaka, Q. Zhao & K.D. Hyde, sp.
nov.
Index Fungorum number: IF552201; Facesoffungi
number: FoF02229, Fig. 48
Etymology: The specific epithet ellipsoidea is refers to
the shape of the ascospores.
Holotype: MFLU 16-0583
Saprobic on dead stems. Sexual morph Ascomata 170–
280 9 200–250 lm (
x = 255 9 236 lm, n = 10), arising
in small groups, sessile, erumpent from the substrate,
subglobose with a flattened base, black, carbonaceous,
ostiole area black, ascomata covered with outwardly projecting, long hairs. Peridium 50–60 lm (
x = 56.6 lm,
n = 10) wide, outer part composed of thick-walled,
strongly melanized cells of textura angularis and inner part
filled with elongate, hyaline cells of textura angularis.
Hamathecium comprising numerous, 1.3–1.8 lm wide
(
x = 1.5 lm, n = 20), long, filiform, aseptate, trabeculate
pseudoparaphyses, branching and anastomosing between
and above the asci. Asci 112–160 9 10–12 lm
(
x = 131 9 11.4 lm, n = 30), 8-spored, bitunicate,
cylindric–clavate, short pedicellate, rounded at the apex,
with an ocular chamber. Ascospores 14–18 9 6–7 lm
(
x = 16.3 9 6.7 lm, n = 40), 1–2 seriate, ellipsoid,
greenish, 1-septate, constricted at septum, with 1–2 globules, acute at the apex, smooth, thick-walled. Asexual
morph Undetermined.
Material examined: THAILAND, Chiang Rai, Mae Fah
Luang University, on dead stems, 17 October 2015, A.H.
Ekanayaka (MFLU 16-0583, holotype).
Notes: Bertiella ellipsoidea is well distinguished by
ascomata arising in small groups, sessile, erumpent from
the substrate, covered with outwardly projecting, long
hairs, trabeculate pseudoparaphyses, branching and
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b Fig. 51 Occultibambusa chiangraiensis (holotype). a Appearance
of ascostromata on host substrate. b, c Close up of ascostromata on
host. d Vertical section of ascostroma. e Section through peridium.
f Cellular pseudoparaphyses. g–i Developmental stages of asci. j–
o Developmental stages of ascospores. p Germinated ascospore. q, r
Culture characters. Scale bars b, c = 500 lm, d = 100 lm,
e = 50 lm, f–i = 20 lm, j–p = 10 lm
anastomosing between and above the asci, bitunicate,
8-spored, cylindric-clavate, asci with an ocular chamber
and (14–18 9 6–7 lm) ellipsoid, 1–2 seriate, greenish,
1-septate ascospores.
Sequence data obtained directly from the type material
clustered within the family Melanommataceae, close to the
type species B. macrospora (Sacc.) Sacc. & Traverso
(Fig. 47). Bertiella ellipsoidea has ellipsoid ascospores,
while in other species in the genus they are fusiform.
Furthermore, B. ellipsoidea has greenish, 1-septate ascospores, while in B. macrospora ascospores are brownish and
3-septate. Byssosphaeria villosa (Samuels & E. Müll.)
Boise is similar to Bertiella ellipsoidea but differs in
having ascospores with a gelatinous sheath (Mugambi and
Huhndorf 2009).
Occultabambusaceae Dai et al.
Dai et al. (2016) introduced Occultabambusaceae to
accommodate two pleomorphic genera Occultibambusa
and Seriascoma, a coelomycetous genus Versicolorisporium and Neooccultibambusa. In their study, the morphology of sexual morphs and asexual morphs were
discussed. In this paper, we introduce two new species,
Occultibambusa aquatica and O. chiangraiensis and provide an updated tree (Fig. 49).
Occultibambusa aquatica Huang Zhang & K.D. Hyde, sp.
nov.
Index Fungorum number: IF552366; Facesoffungi
number: FoF02439, Fig. 50
Etymology: Refers to its aquatic habitat.
Holotype: MFLU 11-1141.
Saprobic on submerged wood in freshwater stream.
Sexual morph Ascomata perithecioid, 100–250 lm high,
180–280 lm diam., scattered or in small groups, immersed
to semi-immersed, subglobose, with a flattened base, brown
to dark brown, with minute ostiolate, central papilla, with
rounded slot. Papilla up to 40 lm long, 50 lm diam., dark
brown. Peridium carbonaceous and fragile, unequal in
thickness, 35–45 lm thick at the sides, 25–35 lm thick at the
base and near ostiole, composed of 5–7 9 1.5–2.5 lm,
rectangular to polygonal celadon cells. Hamathecium comprises 2–3 lm wide, septate, hypha-like, numerous, branched, anastomosing, pseudoparaphyses embedded in
mucilage. Asci 73–86 9 9–13 lm (
x = 78.2 9 11.1 lm,
81
n = 10), 8-spored, bitunicate, fissitunicate, clavate, with a
short furcate pedicel (5–8 lm), which elongates after discharge (up to 55 lm), apically rounded with a small ocular
chamber (1–3 9 4–4.5 lm). Ascospores 19–25 9 3.5–
6.5 lm (
x = 22.2 9 5 lm, n = 20), L/W 3.8–4.8 (mostly
4.5), mostly 2-seriate, narrowly fusiform with acute ends,
1-septate, not constricted at the septum, septum mostly
median, upper cell slightly broader than lower cell, slightly
swollen near the septum, straight to curved, brownish, with
one large guttule in each cell, smooth-walled, thin-walled,
surrounded by a 5–10 lm thick sheath. Germ tube mainly
formed from both end cells. Asexual morph Undetermined.
Culture characteristics: Ascospores germinating on WA
within 12–24 h. Colonies on PDA, dense, dark grey,
reaching up to 1 cm diameter at after 13 days at 25–28 °C,
raised, central embossing, velvety, aerial mycelium and
entire edge smooth, clear.
Material examined: THAILAND, Chiang Rai, Hui Kang
Pla Waterfall, on submerged bamboo, 16 November 2010,
Huang Zhang a50 (MFLU 11-1141, holotype), ex-type
living culture, MFLUCC 11-0006.
Notes: Occultibambusa aquatica is similar to O. bambusae in having clavate asci and 1-septate, fusiform, brown
ascospores with a mucilaginous sheath. However, O. bambusae has larger ascomata (400–550 lm diam.) and larger
ascospores (23.5–27.5 9 4.5–7 lm). A close phylogenetic
relatedness is noted between O. aquatica and O. pustula in
a monophyletic subclade basal to other Occultibambusa
species (Fig. 49) (99 % MLBP, PP greater than 0.90).
Occultibambusa aquatica has no statistical support to
O. pustula and is similar in having fusiform, 1-septate
ascospores, but the latter has raised and darker ascomata
and hyaline to pale brown ascospores. Based on morphological characters and multi-gene phylogenetic analyses,
we introduce a novel species in the genus Occultibambusa.
Occultibambusa chiangraiensis Phukhamsakda & K.D.
Hyde, sp. nov.
Index Fungorum number: IF552240; Facesoffungi
number: FoF02382, Fig. 51
Etymology: The epithet ‘‘chiangraiensis’’ refers to Chiang Rai Province where the holotype was collected.
Holotype: MFLU 16-1334
Saprobic on dead stem of Bambusoideae. Sexual
morph Ascostromata 195–295 lm high 9 352–520 lm
diam. (
x = 264 9 460 lm, n = 5), erumpent, solitary,
scattered, depressed globose to subglobose, flattened at the
base, brown to light brown. Ostioles central, with slit-like
opening. Peridium (8–)12–34 lm wide, of unequal thickness, thickened at sides, with an outer layer of 7–9 layers of
brown to dark brown, heavily pigment cells, arranged in a
textura angularis and textura prismatica, and polygonal at
the base. Hamathecium composed of dense, 1.6–4(–4.7 in
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b Fig. 52 RAxML tree based on analysis of a combined dataset of ITS,
LSU and SSU partial sequences. Bootstrap support values for
maximum likelihood (ML, black) higher than 50 % and Bayesian
posterior probabilities (BYPP, Red) greater than 0.95 are defined as
above the nodes. The tree is rooted to Didymella exigua. All type
strains are in bold. New stains are given in blue
enlarged cells) lm wide (n = 30), transversely septate,
branched, cellular pseudoparaphyses. Asci 47–92 9 12–
16 lm (
x = 69 9 14 lm, n = 20), 8-spored, bitunicate,
clavate-oblong, with a short pedicel, apically obuse, with
an ocular chamber, clearly visible when immature. Ascospores 16–24 9 5–7 lm (
x = 22 9 6 lm, n = 30),
overlapping bi-seriate, hyaline when immature, pale brown
to red-brown at maturity, fusiform, tapering towards the
ends, guttulate in each cell, (1–)3-septate, strongly contricted at the median septum, sometimes the cells above
median septum wider, smooth-walled, lacking a mucilaginous sheath. Asexual morph Undetermined.
Culture characteristics: Colonies on PDA, reaching
50 mm diam. after 4 weeks at 25 °C, dark brown to black
at the edge, surface covered with grey mycelium, rarely
with aerial mycelium, margins lobate, reverse black radiating, dense, umbonate, margin uneven, producing reddish
pigment in agar.
Material examined: Thailand, Chiang Rai Province, on
dead stem of Bambusoideae sp. (Poaceae), 16 April 2015,
C. Phukhamsakda, CP012 (MFLU 16-1334, holotype), extype living culture, MFLUCC 16-0380, KUMCC 16-0027;
ibid. (KUN; HKAS 94617, isotype).
Notes: Occultibambusa chiangraiensis is closely related
to O. bambusae Dai & K.D. Hyde, the type species of
Occultibambusa, based on maximum likelihood analysis
(87 % ML). However, in O. chiangraiensis the ascostromata are erumpent, solitary, with a mixing of peridium cell
types, while in O. bambusae the ascostromata and peridium
cells are only textura angularis. The asci of O. bambusae
are oblong-clavate and ascospores 3-septate without a
sheath (Dai et al. 2016). Occultibambusa chiangraiensis
also shares similarity with O. fusispora Phookamsak et al.,
but the latter has smaller, solitary ascostromata
(135–185 lm high 9 240–275 diam.), a thick-walled
peridium (up to 60 lm wide), and narrowly fusiform
ascospores, acutely tapering at the ends, and not constricted
at the septa (Dai et al. 2016). Doilom et al. (2016) introduced Neooccultibambusa chiangraiensis from Tectona
grandis. Even though ascospores are similar, N. chiangraiensis has immersed ascomata, with obvious ostioles,
larger asci (128 9 20 lm diam.), and cylindrical to subcylindrical, larger ascospores (37 9 10 lm diam.) surrounded by a mucilaginous sheath.
83
Phaeosphaeriaceae M.E. Barr
The family Phaeosphaeriaceae was introduced by Barr
(1979) to accommodate Dothideomycete species, mostly
on monocotyledons and some dicotyledons (Shoemaker
and Babcock 1989; Schoch et al. 2006, 2009; Zhang et al.
2009a, 2012a; de Gruyter et al. 2010; Hyde et al. 2013;
Phookamsak et al. 2014; Wijayawardene et al. 2014b;
Ariyawansa et al. 2015c). Species in Phaeosphaeriaceae
vary in morphological characters and comprise more than
35 sexual and asexual genera (Phookamsak et al. 2014;
Wijayawardene et al. 2014a; Ariyawansa et al. 2015c).
Species in Phaeosphaeriaceae have often been confused
with taxa in Leptosphaeriaceae (Zhang et al. 2012a; Hyde
et al. 2013; Phookamsak et al. 2014; Ariyawansa et al.
2015b; Tennakoon et al. 2016). Based on phylogenetic
analyses, Phaeosphaeriaceae is a heterogeneous group of
taxa with shared similar morphology, but is phylogenetically distinct from species of Leptosphaeriaceae and
Phaeosphaeria sensu lato (Zhang et al. 2012b; Hyde et al.
2013; Phookamsak et al. 2014; Ariyawansa et al. 2015b;
Liu et al. 2015a; Tennakoon et al. 2016). Therefore, several
genera have been introduced to accommodate ambiguous
phaeosphaeriaceous taxa (Ariyawansa et al. 2015b; Li et al.
2015; Liu et al. 2015a; Phukhamsakda et al. 2015; Tibpromma et al. 2015; Tennakoon et al. 2016). In this study
we introduce the new genera Camarosporioides with a
single species, Ca. phragmitis, and Pseudophaeosphaeria,
with Ps. Rubi. We also introduce the new species, Chaetosphaeronema achilleae, Dematiopleospora alliariae, De.
cirsii, Juncaceicola italica, Leptospora galii, Le. aquatica,
Le. thailandica, Muriphaeosphaeria ambrosiae, Neodidymelliopsis ranunculi, Nodulosphaeria italica, Poaceicola arundinis and Wojnowicia italica and provide an
updated tree (Fig. 52).
Camarosporioides W.J. Li & K.D. Hyde, gen. nov.
Index Fungorum number: IF552209; Facesoffungi
number: FoF02350
Etymology: Morphologically resembling the genus Camarosporium, but phylogenetically distinct.
Saprobic on dead stems of Poaceae. Sexual morph
Undetermined. Asexual morph Conidiomata yellowish to
brown, separate or aggregated, pycnidial, obpyriform,
immersed, unilocular, thick-walled, smooth, ostiolate. Ostiole single, circular, with hyaline periphyses, centrally
located, papillate. Periphyses hyaline, hyphae-like, smooth,
subcylindrical, with obtuse apex, unbranched, septate. Wall
of conidiomata composed of thick-walled, brown cells of
textura angularis. Conidiophores reduced to conidiogenous
cells. Conidiogenous cells hyaline to pale brown,
holoblastic, long lageniform, swollen at the base, discrete,
determinate, formed from the cells lining the inner wall of
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b Fig. 53 Camarosporioides phragmitis (holotype). a Herbarium
specimen. b Appearance of black coniodiomata on the host. c Vertical
section of conidioma. d Ostiole. e, f Section of peridium. g–
j Conidiophores, conidiogenous cells and developing conidia. k, l
Germinated conidia. m–p Conidia. q Culture on PDA. Scale bars
c = 100 lm, d–e = 50 lm, f, k–1 = 20 lm, g–j, m–p = 5 lm,
q = 25 mm
the conidioma. Conidia pale brown to brown, finely
roughened, ellipsoidal to oval, obtuse at the apex, slightly
truncate at base, with primary transverse septa, with a
longitudinal septum, constricted at septa, verruculose,
thick-walled.
Type species: Camarosporioides phragmitis W.J. Li &
K.D. Hyde, sp. nov.
Notes: Combined rDNA gene sequence data reveals that
our new genus belongs to the Phaeosphaeriaceae and in
particular is close to Stagnospora neglecta and S. foliicola
(Fig. 52). Morphologically, Camarosporioides shares
similar conidial morphology with Phragmocamarosporium
(Massarinaceae), but Phragmocamarosporium lack periphyses. In addition, both genera can be distinguished by the
form of conidiomata. Camarosporioides species have yellowish to brown, obpyriform, immersed conidiomata,
while Phragmocamarosporium species have black, globose
to subglobose conidiomata (Wijayawardene et al. 2016). In
combining both morphology and phylogeny, we introduce
Camarosporioides as a new genus.
Camarosporioides phragmitis W.J. Li & K.D. Hyde, sp.
nov.
Index Fungorum number: IF552210; Facesoffungi
number: FoF02351, Fig. 53
Etymology: Named after the host genus Phragmites.
Holotype: MFLU 16-1488.
Saprobic on dead stem of Phragmites australis (Cav.)
Trin. ex Steud. (Poaceae). Sexual morph Undetermined.
Asexual morph Coelomycetous. Conidiomata 143–170 lm
diam., 160–177 lm high, yellowish to brown, separate or
aggregated, pycnidial, obpyriform, immersed, unilocular,
thick-walled, smooth, ostiolate. Ostiole 30–70 9 33–
70 lm, single, circular, with hyaline periphyses, centrally
located, papillate. Periphyses hyaline, hyphae-like, smooth,
subcylindrical, with obtuse apex, septate, unbranched. Wall
of conidiomata composed of thick-walled, brown cells of
textura angularis. Conidiophores reduced to conidiogenous
cells. Conidiogenous cells 3.5–10 lm long 9 3–7 lm wide,
hyaline to pale brown, holoblastic, long lageniform, swollen
at the base, discrete, determinate, formed from the cells
lining the inner wall of the pycnidium. Conidia 11–18 9 5–
8 lm (
x = 14 9 6 lm, n = 30), pale brown to brown,
finely roughened, ellipsoidal to oval, obtuse at the apex,
slightly truncate at base, with 1–3 primary transverse septa,
85
and occasionally 1–2 longitudinal septum, constricted at
septa, verruculose, thick-walled.
Culture characteristics: Colony on PDA, reaching
40–50 mm diam. in 7 days, with circular margin, white and
fluffy, dense, aerial mycelium on the surface, reverse
similar in colour.
Material examined: GERMANY, wet meadow, on dead
stem of Phragmites australis (Poaceae), 6 April 2012,
René K. Schumacher, G3 (MFLU 16-1488, holotype); extype living culture, MFLUCC 13-0365 (HKAS 95025,
isotype); living culture, KUMCC 15-0599.
Chaetosphaeronema Moesz
The genus Chaetosphaeronema was introduced by
Moesz (1915) and is represented by C. hispidulum (Corda)
Moesz (type) and C. herbarum (Hollós) Moesz. In this
genus the conidiomata are pycnidial and globose and have
setae. De Gruyter et al. (2009, 2010) used 18S rDNA and
28S rDNA data to show that Chaetosphaeronema was
related to Phaeosphaeriaceae and Pleosporaceae and
treated
Chaetosphaeronema
sensu
stricto
in
Phaeosphaeriaceae. Petrak (1944) and Zhang et al.
(2009b) suspected that Chaetosphaeronema may be asexual morph of Ophiobolus. Phookamsak et al. (2014) confirmed the taxonomic position of Chaetosphaeronema in
Phaeosphaeriaceae.
Chaetosphaeronema achilleae S.K. Huang & K.D. Hyde,
sp. nov.
Index Fungorum number: IF552176; Facesoffungi
number: FoF02240, Fig. 54
Etymology: Named after the host genus Achillea.
Holotype: MFU 15-1922.
Saprobic on dead cone of Achillea nobilis L. Sexual
morph Undetermined. Asexual morph Conidiomata 200–
290 lm high 9 180–245 wide lm (
x = 240 9 200 lm,
n = 5) diam., pycnidial, solitary, uniloculate, immersed,
subglobose, brown to dark brown, with minute papilla.
Ostiole central, short, lined with periphyses. Wall of
conidiomata 25–35 lm, membranaceous, composed of
dark brown, or brown to hyaline cells of textura angularis.
Conidiophores reduced to conidiogenous cells. Conidiogenous cells 7–12 9 1–2 lm (
x = 10 9 1.5 lm, n = 20),
enteroblastic, phialidic, cylindrical to subcylindrical,
smooth-walled, hyaline, arising from the inner layers of
conidioma. Conidia 9–16 9 1.5–3 lm (
x = 12 9 2 lm,
n = 50), cylindrical to subcylindrical, slightly curved,
hyaline, aseptate to 1-septate, smooth-walled.
Culture characteristics: Colonies on PDA reaching
10 mm diam. after 2 weeks at 16 °C, white at the margin,
dark brown to green at the center; reverse cream to white at
the margins, dark brown to black at the center, curled,
circular, umbonate, without diffusible pigments.
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b Fig. 54 Chaetosphaeronema achilleae (holotype). a Herbarium
specimen. b Conidiomata on host. c Immersed conidioma. d Conidioma in vertical section. e Wall of conidioma. f, g Conidiophores and
conidia. h–j Conidia. k, l Culture characters on PDA. Notes g stained
in lactophenol cotton blue. Scale bars b = 500 lm, c = 200 lm,
d = 100 lm, e = 50 lm, f–g = 10 lm, h–j = 2 lm
Material examined: RUSSIA, Rostov Region,
Oktyabrsky District, southern outskirts of Persianovky
settlement, Khoruli gully (rus. balka Khoruli), on dead
stems of Achillea nobilis (Asteraceae), 28 April 2015, T.S.
Bulgakov (MFLU 15-1922, holotype, GZU 16022408,
isotype), ex-type living culture, MFLUCC 16-0476,
GZUCC 16022408.
87
Notes: Chaetosphaeronema achilleae is morphologically similar to C. hispidulum (type species of Chaetosphaeronema) in having globose pycnidial, cylindrical
conidiogenous cells, and cylindrical to subcylindrical,
slightly curved, and hyaline conidia. However, C. hispidulum is distinct from C. achilleae in having comparatively
large conidiomata (up to 450 lm diam.) with numerous
setae, and a pycnidial wall comprising cells of textura
prismatica, while C. achilleae has smaller conidiomata,
lacking setae and the pycnidial wall comprises cells of
textura angularis (Sutton 1980). Phylogenetic data also
indicate C. achilleae is closely to C. hispidulum, but
morphological differences stated above support our establishment of a new species (Fig. 52).
Fig. 55 Dematiopleospora
alliariae (holotype).
a Appearance of ascomata on
host surface. b, c Vertical
sections through ascomata.
d Peridium.
e Pseudoparaphyses. f, g Asci.
h Ascospores. i Germinating
ascospore. Scale bars b,
c = 100 lm, d–h = 50 lm,
i = 25 lm
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89
Dematiopleospora Wanasinghe et al.
Wanasinghe et al. (2014) introduced Dematiopleospora
as a monotypic genus in the family Phaeosphaeriaceae to
accommodate D. mariae Wanasinghe et al. Subsequently, a
second species D. luzulae Wanasinghe et al., was added by
Ariyawansa et al. (2015a). The genus is characterized by
thick, brown, periphyses in the ostiole, immersed to
superficial ascomata and yellowish-brown to brown,
muriform ascospores with light end cells. Two new additions are proposed in the genus.
Colonies on PDA, reaching 10–12 mm diam. after 7 days at
25 °C, surface dirty white to buff, spreading with moderate
aerial mycelium, and even, smooth and entire margins.
Material examined: ITALY, Forlı̀-Cesena [FC] Province, Ridracoli—Bagno di Romagna, on dead stem of
Alliaria petiolata (M. Bieb.) Cavara & Grande (Brassicaceae), 18 November 2012, Erio Camporesi IT 915
(MFLU 15-0545, holotype); ex-type living culture,
MFLUCC 13-0070, ICMP.
Notes: Dematiopleospora alliariae is introduced as the
third species of Dematiopleospora. Dematiopleospora
alliariae mainly differs from the other two species in having
ascomata with a crest-like ostiole and uni-seriate ascospores
with 3(–4) transverse septa and 2–4 vertical septa. There is
also support herein from our molecular data analysis that
D. alliariae warrants new species status as it did not cluster
with other Dematiopleospora species (Fig. 52).
Dematiopleospora alliariae Thambugala, Camporesi &
K.D. Hyde, sp. nov.
Index Fungorum Number: IF552265; Facesoffungi
number: FoF02440, Fig. 55
Etymology: The specific epithet alliariae refers to the
host genus on which the fungus occurs.
Holotype: MFLU 15-0545.
Saprobic on Alliaria petiolata (M. Bieb.) Cavara &
Grande. Sexual morph Ascomata 210–350 lm
high 9 175–300 lm diam. (
x = 270 9 224 lm, n = 6),
solitary to gregarious, scattered, immersed, slightly erumpent, coriaceous, dark brown to black, subglobose to
pyriform, thickened at the apex, ostiolate. Ostiole central,
papillate, with a pore-like opening. Peridium up to 60 lm
wide, thin at the sides, broad at the apex, comprising two
strata, outer stratum composed of small, brown to dark
brown, somewhat flattened, thick-walled cells of textura
angularis, fusing and indistinguishable from the host tissues, inner stratum composed of few layers of lightly
pigmented to hyaline cells of textura angularis. Hamathecium comprising 1–2 lm wide, septate, cellular pseudoparaphyses, embedded in a gelatinous matrix. Asci 100–
125 9 10–12 lm (
x = 110 9 10.9 lm, n = 15), 8spored, bitunicate, fissitunicate, cylindrical, pedicellate,
rounded at the apex, with a distinct ocular chamber. Ascospores 14–17.5 9 6.4–8.4 lm (
x = 15.5 9 7.5 lm,
n = 35), uniseriate, partially overlapping, initially hyaline,
becoming yellowish-brown at maturity, ellipsoidal to
fusiform, muriform, transversely 3-septate, with 0–3(–4)
vertical septa, slightly curved, deeply constricted at the
central septum, rounded at the ends, smooth-walled, lacking a mucilaginous sheath. Asexual morph Undetermined.
Culture characteristics: Ascospores germinating on PDA
within 24 h and producing germ tubes from several cells.
Dematiopleospora cirsii Wanasinghe, Camporesi, E.B.G.
Jones & K.D. Hyde, sp. nov.
Index Fungorum number: IF552135; Facesoffungi
number: FoF02205, Fig. 56
Etymology: Name reflects the host genus Cirsium, from
which the species was collected.
Holotype: MFLU 16-0145.
Saprobic on dead and hanging branches of Cirsium sp.
Sexual morph Ascomata 250–300 lm high 250–350 lm
diam. (
x = 271.1 9 299.5 lm, n = 5), immersed to semierumpent, solitary, scattered, broadly oblong and flattened,
dark brown to black, coriaceous, cupulate when dry, ostiolate. Ostiole 50–65 lm high 40–50 lm diam.
(
x = 58.2 9 43.4 lm, n = 5), papillate, black, smooth,
with short and light brown setae. Peridium 10–20 lm wide at
the base, 15–30 lm wide in sides, thick, with 4–5 layers,
outer layer heavily pigmented, thick-walled, comprising
reddish to dark brown cells of textura angularis, inner layer
composed of hyaline thin-walled cells of textura angularis.
Hamathecium comprising numerous, 1.5–2 lm wide, filamentous, branched, septate, pseudoparaphyses. Asci 80–
120 9 10–14 lm (
x = 100.9 9 12.3 lm, n = 30), 8spored, bitunicate, fissitunicate, cylindrical to cylindricclavate, pedicellate, thick-walled at the apex, with minute
ocular
chamber.
Ascospores
20–30 9 6–9 lm
(
x = 25.9 9 6.6 lm, n = 50), overlapping uni-seriate,
muriform, ellipsoidal to subfusiform, slightly curved, upper
part wider than the lower part, 6–7 transversely septate, with
1 vertical septum, deeply constricted at the central septum,
initially hyaline, becoming golden-brown at maturity, ends
remaining cone-shaped, with rounded ends, without a
mucilaginous sheath. Asexual morph Undetermined.
Culture characteristics: Colonies on MEA, reaching
1.5 cm diam. after 30 days at 18 °C, circular, smooth
b Fig. 56 Dematiopleospora cirsii (holotype). a Appearance of asco-
mata immersed beneath host substrate. b, c Sections of ascomata.
d Close up of ostiole with short, light brown, setose hyphae.
e Peridium. f Pseudoparaphyses. g–j Asci. k–n Ascospores. o Germinated ascospore. p, q Culture on PDA (note q reverse). Scale bars
a = 500 lm, b = 100 lm, c, d = 50 lm, e, k–o = 10 lm,
f = 5 lm, g–j = 20 lm
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margin white at first, greenish-grey after 4 weeks, flat on
the surface, without aerial mycelium, reverse blackishgreen. Hyphae septate, branched, hyaline, thin, smoothwalled.
Material examined: ITALY, Arezzo [AR] Province,
Papiano, dead herbaceous branches of Cirsium sp.
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Fig. 58 Leptospora rubella (reference specimen). a Appearance of c
ascomata on host. b Close up of an ascoma. c, d Vertical section of
ascoma. e Pseudoparaphyses. f, g Asci. h Ascospores. i Germinated
ascospore. j Mycelium. k, l Culture on PDA (note l reverse). Scale
bars a = 1 mm b = 200 lm, c = 100 lm, d = 50 lm, e = 5 lm, f–
j = 20 lm
Fig. 57 Juncaceicola italica (holotype). a Appearance on the host substrate. b Section of ascoma. c Section of peridium. d Pseudoparaphyses.
e–h Ascus. i–m Ascospores. Scale bars a = 100 lm, b = 20 lm, c = 5 lm, d = 2 lm, e–h = 10 lm, i–m = 5 lm
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b Fig. 59 Leptospora galii (holotype). a, b Appearance of ascomata
on host. c Vertical section of ascoma. d, e Peridium. f Pseudoparaphyses. g–i Asci. j–m Ascospores. n Germinating ascospore. o Upper
view of culture on PDA. p Lower view of culture on PDA. Scale bars
c, g, h, n = 20 lm, d, e, j–m = 10 lm
(Asteraceae), 13 June 2013, E. Camporesi (MFLU
16-0145, holotype); (isotype in BBH), ex-type living
culture, MFLUCC 13-0615
Notes: In this study, our strain MFLUCC 13-0615
groups with Dematiopleospora mariae (type strain of
Dematiopleospora) with very high bootstrap and Bayesian
support (Fig. 52). Dematiopleospora cirsii resembles
D. mariae in having brown periphyses in the ostiole, a
peridium comprising brown to dark brown cells of textura
angularis and multi-septate ascospores, but differs in
having immersed ascomata, ascospores with 7 transverse
septa and much longer lower part than upper part, while
D. mariae has superficial ascomata, ascospores with 9–10
transverse septa, and upper and lower parts of equal
length.
Juncaceicola Tennakoon et al.
The genus Juncaceicola (Phaeosphaeriaceae) was
introduced by Tennakoon et al. (2016), with J. luzulae as
the type species.
Juncaceicola italica Tibpromma, Camporesi & K.D.
Hyde, sp. nov.
Index Fungorum number IF552246; Facesoffungi number: FoF02388, Fig. 57
Etymology: Refers to the name of the country where the
holotype was collected
Holotype: MFLU 14-0685.
Saprobic on dead stem of Dactylis glomerata L. Sexual
morph Ascomata 115–130 lm high 9 130–140 lm diam.
(
x = 125 9 134 lm, n = 4), superficial, solitary, scattered,
globose to subglobose, with a flattened base, easy to remove
from the host, shiny, ostiole central, black, smooth-walled.
Peridium 5–8 lm, yellow-brown, a single stratum comprised of 2–3 cell layers of textura prismatica. Hamathecium
comprising numerous 1.3–3 lm wide, filamentous, guttulate, branched anastomosing, septate pseudoparaphyses.
Asci 62–88 9 11–20 lm (
x = 77 9 17 lm, n = 10), 6–8spored, bitunicate, clavate to cylindrical, thin-walled, short
pedicellate, apex rounded, ocular chamber not well-developed. Ascospores 18–21 9 6–8 lm (
x = 18 9 7 lm,
n = 10), fusiform with broadly to narrowly rounded ends,
3-septate, constricted at septa, enlarged at the second cell,
green-yellow, guttulate, surrounded by mucilaginous sheath.
Asexual morph Undetermined.
Culture characteristics: on PDA reaching 2 cm diam.
after 2 weeks at 16 °C, later with dense mycelium, with
circular, rough margin, flattened; upper surface white at
93
first, dark brown after 4 weeks, producing pigments in
PDA media agar; hyphae septate, branched, thick-walled.
Material examined: ITALY, Forlı̀-Cesena Province,
Monte Falco, on dead stem of Dactylis glomerata (Poaceae), 15 July 2013, Erio Camporesi, IT1378 (MFLU
14-0685, holotype); ex-type living culture, MFLUCC
13-0750; ibid. (HKAS94558 bis, paratypes).
Notes: Phylogenetic results show Juncaceicola italica to
be related to J. dactylidis; however, we found our new
taxon is well-differentiated from J. dactylidis based on
morphological differences. Our new taxon is characterized
by superficial ascomata, a peridium of textura prismatica,
6–8-spored asci and 3-septate ascospores, while J. dactylidis has immersed or erumpent ascomata, a peridium of
textura angularis, 8-spored asci and 4-septate ascospores
(Tennakoon et al. 2016).
Leptospora Rabenh.
Leptospora is typified by L. rubella (Pers.) Rabenh.
Persoon (1801) originally assigned the genus to Sphaeria
rubella Pers., however, Rabenhorst (1858) transferred it to
Dothideomycetes under Leptospora. Leptospora frequently stains the surface of host tissues red to purple and
the red colour is also present at the apical part of ostiolar
canal. The taxa has multi-septate ascospores. Leptospora
is similar to Ophiobolus Riess base on their ascospore
morphology (Shoemaker 1976; Crous et al. 2006).
Therefore, in this study we reconstruct the phylogenetic
analysis (Fig. 52) for Phaeosphaeriaceae and introduce
three new species, L. aquatica, L. galii, L. thailandica
with provide a reference specimen for L. rubella
(96 %ML, 1.00 PP).
Leptospora rubella (Pers.) Rabenh., Klotzschii Herb. Viv.
Mycol., Edn 2: no. 532 (1857)
Index Fungorum number: IF552325; Facesoffungi
number: FoF02442, Fig. 58
Saprobic on decaying wood submerged in freshwater.
Sexual morph Ascomata 250–260 lm high, 185–190 lm
diam., solitary or aggregated, immersed to semi-immersed,
erumpent, globose to subglobose, coriaceous, black,
apapillate, ostiolate. Ostiole central, ostiolar canal filled
with hyaline to brown cells. Peridium 8–9 lm at base, 20–
24 lm at sides comprising 3–4 layers of pale brown to
brown, inwardly lighter, thin-walled cells of textura
angularis. Hamathecium comprising numerous, 2–3 lm
wide, branched, cellular, hyaline pseudoparaphyses. Asci
100–150 9 5–9 lm (
x = 107 9 6 lm, n = 10), 8-spored,
bitunicate, long-cylindrical, short pedicellate, apex rounded, with minute ocular chamber. Ascospores 90–
125 9 2–3 lm (
x = 115 9 2.5 lm, n = 10), arranged
spirally in the ascus, initially hyaline becoming pale brown
at maturity, filiform, ends narrowly rounded, straight or
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Fig. 60 Leptospora thailandica (holotype). a Specimens. b Appearance of ascomata on host surface. c Vertical section of ascoma.
d Ostiolar canal. e Section through peridium. f Cellular
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pseudoparaphyses. g–i Developing state of asci. j–l Developing
stages of ascospores. m Ascospore stained with Indian ink to show
septa. Scale bars c = 100 lm, d, g–m = 50 lm, e, f = 10 lm
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95
Fig. 61 Muriphaeosphaeria ambrosiae (holotype). a Herbarium
packet. b Herbarium material. c Conidiomata on host. d Close up
of conidiomata, superficial on host. e Conidioma in vertical
section. f Wall of conidioma. g–i Conidiophores and conidia. j–
m Conidia. Notes h–i, m stained in lactophenol cotton blue, g stained
in Indian ink. Scale bars c = 1 mm, d = 200 lm, e = 100 lm, f–
i = 10 lm, j–m = 5 lm
curved, thick-walled, smooth-walled, ascospores are
sometimes longer than asci. Asexual morph
Undetermined.
Culture characteristics: Colonies on PDA 4 cm diam.
after 4 weeks at 16 °C, dirty white to creamy at the margins, grey to pale-brown at the center; reverse grey to pale
brown and cream at the center, gauzy, curled, flat.
Material examined: UK, Hampshire, Swanick Lakes, on
decaying wood submerged in freshwater, 28 August 2015,
E.B.G. Jones, GJ187 (MFLU 16-0965, reference specimen designated here, HKAS94519), living culture,
MFLUCC 16-0122, KUMCC 16-0006.
Notes: The strain MFLU 16-0965 has morphologically
identical to Leptospora rubella, although, it differs from
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96
the type of L. rubella as the ostiole lacks a distinct papilla.
In the phylogenetic analysis (Fig. 52), our collection
grouped with the Leptospora rubella (CPC 11006), thus
they are not phylogenetically distinct (Fig. 52).
Leptospora galii de Silva & K.D. Hyde, sp. nov.
Index Fungorum number: IF552317; Facesoffungi
number: FoF02441, Fig. 59
Etymology: Name reflects the host genus Galium.
Holotype: HKAS 92493a.
Saprobic on dead branch of Galium sp. Sexual morph
Ascomata 105–226 lm high, 109–188 lm diam., solitary,
Fig. 62 Nodulosphaeria italica (MFLU 16-1359, holotype). a Appearance of ascomata on the host surface. b Section through
ascoma. c Section through neck. d Section through peridium.
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Fig. 63 Poaceicola arundinis (holotype). a Appearance of ascoma c
on host surface. b Vertical section of ascoma. c Ostiole. d Section through peridium. e Cellular pseudoparaphyses. f Immature ascus
with ocular chamber. g Mature ascus. h–k Developing state of
ascospores. l Ascospore stained with Indian ink to show sheath.
m Germinated ascospore. n, o Culture character on PDA from surface
and reverse. Scale bars a, b = 100 lm, c, d = 50 lm, e–g,
m = 20 lm, h–l = 10 lm
scattered to clustered, immersed to semi-immersed to
erumpent, globose to subglobose, glabrous, visible as
raised, black spots on host surface. Peridium 18–25 lm
wide, unevenly thickened, two-layered; inner layer comprising 3–4 cell layers of hyaline cells of textura angularis
e Pseudoparaphyses. f, g Asci. h–j Ascospores. Scale bars
b = 100 lm, c = 50 lm, d–j = 20 lm
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or 3–4 cell layers of flattened, hyaline cells of textura
prismatica; outer layer comprising several layers of brown
to dark brown cells of textura angularis. Hamathecium
composed of dense, 1.5–2.5 lm wide, filamentous, septate,
cellular pseudoparaphyses, anastomosing at the apex. Asci
Fig. 65 Wojnowicia italica (holotype). a Herbarium material. b, c c
Appearance of ascomata semi-immersed in the host. d–f Hand sections of
ascomata showing peridia. g–j Asci with ascospores. k–o Ascospores.
p Pseudoparaphyses. q Germination of spore. r Colony on PDA from
above. s Colony on PDA from below. Scale bars b–c = 200 lm, d–
e = 50 lm, f = 20 lm, g–j, p–q = 10 lm, k–o = 5 lm
Fig. 64 Pseudophaeosphaeria rubi (holotype). a, b Appearance of
ascomata on host surface. c, d Sections through the ascomata. e Papilla.
f Peridium. g Pseudoparaphyses. h Asci with pseudoparaphyses in
gelatinous matrix. i–h Asci. m–p Ascospores. Scale bars c, d = 50 lm,
e = 20 lm, f, g–l = 20 lm, m–p = 5 lm
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99
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100
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55–87 9 3.7–6.9 lm (
x = 70 9 5.2 lm, n = 30), 8spored, bitunicate, cylindric-clavate, short pedicellate, with
an obtuse to slightly furcate end, apically rounded, with
ocular chamber when immature and indistinct at maturity.
Ascospores 17–32 9 2.1–3.3 lm (
x = 23.2 9 2.7 lm,
n = 30), overlapping bi-seriate, hyaline, elongate fusiform
with obtuse ends, and sometimes curved at ends, 1-septate
at maturity, guttulate. Asexual morph Undetermined.
Culture characteristics: Colonies on PDA 32–35 mm
diam. after 3 weeks, colonies sparse, circular, flat, surface
smooth with entire edge, velvety with smooth aspects,
zonate with different sector light brown to brown at the
margin whitish-grey at the centre; reverse light brown at
margin, brown at the middle, dark brown at the centre.
Material examined: ITALY, Forlı̀-Cesena Province,
Monte Mirabello-Predappio, on dead branch of Galium sp.
(Rubiaceae), 11 May 2015, E. Camporesi, IT2478 (HKAS
92493, holotype), (MFLU 15-1230, isotype); ex-type living culture KUMCC 15-0521, MFLUCC.
Notes: Our new species clearly fits into Leptospora
where all species constitute a monophyletic clade with
high support (Fig. 52). In addition, phylogenies also
reveal that L. galli is phylogenetically distinct as it
branches basal to other species with high support.
Leptospora galii differs from other Leptospora species
by having hyaline, elongate, fusiform, rarely light
yellow, mature, 1-septate ascospores, 17–32 lm long,
while other Leptospora species in the same clade have
brown, filiform, multi-septate ascospores more than
60 lm long.
Fig. 66 Phylogram generated from maximum likelihood analysis
based on combined LSU, ITS and SSU sequence data of Pleosporaceae. Maximum likelihood bootstrap support values greater than
60 % are shown above nodes. The ex-type strains are in bold and the
new isolate is in blue. The tree is rooted with Leptosphaeria doliolum
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Leptospora thailandica Phukhamsakda & K.D. Hyde, sp.
nov.
Index Fungorum number: IF552239; Facesoffungi
number: FoF02381, Fig. 60
Etymology: The name reflects the country, where the
holotype was collected.
Holotype: MFLU 16-1335
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101
Fig. 67 Comoclathris italica (holotype). a Appearance of ascomata
on host substrate. b Section of ascoma. c Section of peridium.
d Pseudoparaphyses. e, f Asci. g–i Ascospores. j Germinating
ascospore. Scale bars b = 50 lm, c = 5 lm, d = 2 lm, e,
f = 20 lm, g–j = 10 lm
Saprobic on dead branches of Duranta sp. Sexual
morph Ascomata 188–207 lm high 9 112–170 lm wide
diam. (
x = 197 9 141 lm, n = 10), immersed to
erumpent through host tissue with papilla, solitary, scattered, globose, smooth, brown to dark brown, ostiole central. Ostioles 66–115 lm high 9 53–102 lm diam.
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102
(
x = 79 9 83 lm, n = 10), papillate, dark brown to light
brown, heavily pigmented at outer layer, smooth, filled
with periphyses, orange around pore. Peridium (5–)10–
24(–27) lm wide, up to 30 lm wide at the apex, thickwalled, brown to dark brown, pseudoparenchymatous cells,
composed of 5–7 layers of textura angularis, in hyaline
gelatinous layers, thin. Hamathecium comprising numerous, filamentous, 1.7–4 lm (n = 50) wide, broad, branched, transversely septate, cellular pseudoparaphyses. Asci
68–114 9 7–13 lm (
x = 94 9 10 lm, n = 35), 8-spored,
bitunicate, cylindrical to cylindrical-clavate, with short
furcate pedicel, apically rounded, ocular chamber visible
when immature. Ascospores 63–89 9 1.8–3.8 lm
Fig. 68 Maximum Parsimony
tree obtained from a heuristic
search with 1000 random taxon
additions of combined LSU and
ITS sequence data using PAUP
v. 4.0b10. Bootstrap support
values of more than 50 % are
indicated above the nodes, and
Bayesian posterior probabilities
more than 0.9 are indicated in
bold. The isolate from this study
is shown in blue background,
and Biatriospora marina as the
out group taxon, and the scale
bar shows 10 changes. The extype strains are in bold
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Fig. 69 Neoroussoella lenispora (holotype). a, b Appearance of c
ascostromata on the host surface. c Pseudoparaphyses. d Vertical
section through ascostroma. e Section of peridium. f–i Long cylindrical asci with ascospores. j–m Ascospores. Scale bars a = 500 lm,
b = 100 lm, c = 30 lm, d = 50 lm, e = 10 lm, f–i = 20 lm, j–
m = 5 lm
(
x = 76 9 3 lm, n = 40), fasciculate, scolecosporous,
filiform, tapering towards the ends, minute guttules in each
cell, hyaline when immature, pale brown at maturity, (14–)
20–22-septate, contricted at the septa. Asexual morph
Undetermined.
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Culture characteristics: Colonies on PDA, reaching
20 mm diam. after 4 weeks at 16 °C, surface cream, dense,
circular, convex with moderate aerial mycelium, downy,
slightly irregular at margins; reverse cream at the edges,
brown at the center, dense, margin rough, not pigmented.
Material examined: THAILAND, Chiang Rai, on dead
branches of Duranta sp. (Verbenaceae), 12 June 2015, C.
Phukhamsakda, CP014 (MFLU 16-1335, holotype), isotype in HKAS 94616, ex-type living culture, MFLUCC
16-0385, KUMCC 16-0030.
Notes: Leptospora thailandica forms a sister clade
(99 % ML, 100 %PP) with L. aquatica, L. galii (this
study), and L. rubella (Pers.) Rabenh., the type species of
Leptospora. Leptospora thailandica is similar to Leptospora in staining the host substrate pinkish-red and
having remarkable reddish colour at the apical part of
ostiole canal, with pale brown and cylindrical ascospores
(Crous et al. 2006).
Muriphaeosphaeria C. Phukhamsakda et al.
Muriphaeosphaeria was introduced by Phukhamsakda
et al. (2015) as a monotypic genus with M. galatellae
Phukhamsakda et al. as the type species. The genus is
characterized by superficial ascomata, muriform ascospores and conidiomata with cylindrical to subclavate, 1–3-
Fig. 70 Phylogram generated from maximum likelihood analysis
(RAxML) based on combined ITS and LSU sequence data. Bootstrap
support values for maximum likelihood (ML, red) and maximum
parsimony (MP, green) equal to or greater than 50 % are given above
the nodes. The values of the Bayesian posterior probabilities from
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Fig. 71 Neotorula submersa (HKAS 92660, holotype). a Colonies c
on the substrate. b–d Branched chains of conidia with conidiophores.
e–j Conidia. k Germinating conidium. l, m Cultures on PDA. Scale
bars b = 30 lm, c, d = 20 lm, e, h–k = 15 lm, f, g = 10 lm
septate and brown conidia. This genus was accepted in
Phaeosphaeriaceae based on combined ITS, 18S and 28S
sequence data (Fig. 52).
Muriphaeosphaeria ambrosiae S.K. Huang & K.D. Hyde,
sp. nov.
Index Fungorum number: IF552177; Facesoffungi
number: FoF02241, Fig. 61
Etymology: The specific epithet ambrosiae is based on
the host genus from which the holotype was collected.
Holotype: MFU 15-1971
Saprobic on dead branch of Ambrosia artemisiifolia L.
Sexual morph Undetermined. Asexual morph Conidiomata 228–328 lm high 9 224–268 wide lm
(
x = 281 9 249 lm, n = 5), globose, cleistothecial, solitary, uniloculate, black, superficial, globose to subglobose.
Wall of conidiomata 10–14 lm diam., membranaceous,
composed of dark brown, or hyaline to brown cells of textura
angularis. Conidiophores reduced to conidiogenous cells.
Conidiogenous cells 7–12 9 1.5–3 lm (
x = 9 9 2 lm,
MCMC analyses (BYPP, blue) equal or higher than 95 % are given
below the nodes. The tree was rooted to Melanomma pulvis-pyrius.
Newly generated sequences are indicated in red, and other ex-type
strains are in bold
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n = 20), enteroblastic, phialidic, oblong to cylindrical,
smooth-walled, hyaline. Conidia 9.5–11.5 9 3–4 lm
(
x = 10 9 3.5 lm, n = 50), oval, fusiform to oblong,
1-septate, initially hyaline, pale brown at maturity, smoothwalled, with three inconspicuous and filiform, apical
appendages at the apex.
Material examined: RUSSIA, Rostov Region, Shakhty
City, Cotton Fabric Urban Microdistrict, Grushevka steppe
slopes, near Grushevsky Pond, on dead stems of Ambrosia
artemisiifolia (Asteraceae), 12 May 2015, T.S. Bulgakov
(MFLU 15-1971, holotype, GZU 2016022413, isotype).
Notes: We could not obtain a culture from single conidia. Therefore fungal DNA was extracted directly from the
conidiomata. Muriphaeosphaeria ambrosiae is a sister to
M. galatellae with moderate support (Fig. 52). Both these
species are characterized by globose to subglobose,
uniloculate conidiomata, and pycnidial wall comprising
cells of textura angularis. They were found on Asteraceae.
The asexual morph of M. galatellae is distinct from
M. ambrosiae in having cylindrical to subclavate, 1–3septate conidia, while M. ambrosiae has 1-septate, oval to
oblong conidia with three inconspicuous appendages at the
apex (Phukhamsakda et al. 2015).
Nodulosphaeria Rabenh.
Nodulosphaeria is a relatively poorly studied genus
lacking a modern taxonomic treatment and phylogenetic
analyses (Hyde et al. 2013; Phookamsak et al. 2014). Since
2014, mycologists have attempted to resolve its natural
placement in Phaeosphaeriaceae by recollecting samples
and investigated their phylogeny (Ariyawansa et al. 2015c;
Li et al. 2015; Liu et al. 2015a). Presently, there are 28
sequences from seven species available in GenBank, with 64
epithets in Index Fungorum (2016). The most recent treatment of Nodulosphaeria which includes molecular data is
that of Mapook et al. (2016) and this is followed here.
Nodulosphaeria has relatively large ascomata, and
phragmospores or scolecospores, with an enlarged cell in
the upper part of the ascospores. Nodulosphaeria differs
from Ophiobolus in having brown, setae-like periphyses in
the ostiole, and ascospores do not separate into part spores
(Phookamsak et al. 2014; Mapook et al. 2016). Many
Nodulosphaeria species were transferred to Ophiobolus by
Shoemaker (1976, 1984). Subsequent authors introduced
new species and designated reference specimens to resolve
the placement of Nodulosphaeria (Phookamsak et al. 2014;
Ariyawansa et al. 2015a; Li et al. 2015; Liu et al. 2015a;
Mapook et al. 2016). However, many species still lack
modern taxonomic treatments or molecular data to clarify
their placement.
Nodulosphaeria italica Phookamsak, Camporesi & K.D.
Hyde, sp. nov.
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Fig. 72 RAxML phylogenetic tree generated from combined LSU c
and ITS sequence data from species of Tubeufiaceae. The final ML
optimization likelihood is -16263.541876. Bootstrap support values
greater than 50 % are shown at the nodes and Bayesian posterior
probabilities greater than 0.95 are marked with an asterisk (*). The
tree is rooted with Hysteropatella elliptica (Patellariaceae). Type
species are in bold and marked (T) and reference specimens are
marked (R)
Index Fungorum number: IF552202; Facesoffungi
number: FoF02261, Fig. 62
Etymology: The specific epithet ‘‘italica’’ refers to the
country, where the holotype was collected.
Holotype: MFLU 16-1359
Saprobic on Cirsium sp. Sexual morph Ascomata 240–
330 lm high (excluding necks), 280–330 lm diam., dark
brown to black, scattered, gregarious, immersed to erumpent through the host cortex, globose to subglobose, uniloculate, setose, covered by dark brown, septate, vegetative
hyphae. Papilla 190–330 lm high, 120–140 lm diam.,
truncate to cylindrical, composed of several layers of thick,
brown to dark brown, pseudoparenchymatous cells, arranged in a textura angularis, ostiole central, pore-like opening, with brown, 2–3-septate, setae-like periphyses.
Peridium 16–34 lm wide, thin-walled, of equal thickness,
composed of two types of brown to dark brown, pseudoparenchymatous cells, inner layers comprising 2–3 layers,
of flattened, dark brown cells, arranged in textura prismatica to textura angularis, outer layers comprising 1–2
layers, of thickened, subhyaline to black cells, arranged in
textura angularis to textura globulosa. Hamathecium
composed of dense, 2–4 lm wide, cellular pseudoparaphyses, distinctly septate, anastomosing at the apex,
embedded in a hyaline gelatinous matrix. Asci (157–)170–
200(–206) 9 (11–)12–14(–15) lm (
x = 183.5 9 13.2 lm,
n = 30), 8-spored, bitunicate, fissitunicate, cylindrical,
short pedicellate, with obtuse to furcate pedicel, apically
rounded, with well-developed ocular chamber. Ascospores
(114–)120–150(–158) 9 (2–)3–4(–5) lm (
x = 141.5 9
3.8 lm, n = 30), tri- to tetra-seriate in parallel, not fasciculate, yellowish-brown to brown, filiform, with rounded
ends, tapered towards lower cell, enlarged at the 8th cell,
multi-septate (17–20 septa), constricted at the central septum, smooth-walled, with guttules. Asexual morph
Undetermined.
Material examined: ITALY, Province of Trento[TN],
Mezzana, Marilleva 900, on dead stem of thistle (Cirsium
sp.), 3 June 2014, E. Camporesi, IT 549 (MFLU 16-1359,
holotype; MFLU 16-1360, isotype); ibid. 9 June 2014, E.
Camporesi (MFLU 16-1361); 16 July 2012, E. Camporesi
(MFLU 15-0449).
Notes: Nodulosphaeria italica is similar to N. cirsii (P.
Karst.) L. Holm in having brown, scolecosporous ascospores with multi-septa. However, they can be distinguished
Fungal Diversity (2016) 80:1–270
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123
108
Fig. 72 continued
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109
Fig. 73 Aquaphila albicans (MFLU 16-1136, reference specimen). a Conidia arise directly from hyphal cells on natural substrate. b–
d Conidiophores with attached conidia. e–h Conidia. i Germinating spore. j, k Colonies on MEA from surface and reverse. Scale bars
a = 200 lm, b–i = 50 lm, j, k = 20 mm
by the size of asci and ascospores, as N. italica has smaller
asci and ascospores than N. cirsii. Based on phylogenetic
analyses, N. italica sits with N. senecionis in a robust clade
(92 % ML, 0.95 PP), while N. scabiosae is basal to both
species (Fig. 52).
Poaceicola W.J. Li et al.
The genus Poaceicola was introduced by Li et al. (2015)
to accommodate taxa from Poaceae. The genus includes
Po. arundinis Li et al. (type species), Po. bromi
Wijayawardene et al. and Po. elongata (Wehm.)
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b Fig. 74 Chlamydotubeufia helicospora (MFLU 16–1338, holotype).
a Conidia arise directly from hyphal cells on natural substrate.
b Conidiophores with attached conidia. c Conidiogenous cells.
d Germinating conidium. e–h Conidia. i–j Colonies on MEA from
above and below. Scale bars a = 500 lm, b, e–h = 50 lm,
c = 20 lm, d = 100 lm, i–j = 20 mm
Shoemaker & C.E. Babc.) Li et al. Poaceicola is characterized by immersed to semi-immersed ascomata, subepidermal, globose, papillate, usually reddish-brown and
fusiform ascospores (Li et al. 2015). Specimens in the
genera produce brown conidia and are coelomycetous.
Poaceicola elongata has a sexual morph which is phaeosphaeria-like (Ariyawansa et al. 2014b). Li et al. (2015)
introduced the asexual morph into Phaeosphaeriaceae
based on combined LSU and ITS sequence data. From the
analysis they found Poaceicola is closely related to
P. elongata (96 % ML), and therefore they synonymized
Phaeosphaeria elongata under Poaceicola elongata. In this
study, we reconstruct the phylogeny of genera of
Phaeosphaeriaceae and our strain (MFLUCC 14-1060)
clusters together with Po. arundinis (MFLUCC 15-0702)
with high support. We therefore describe the sexual morph
for Po. arundinis.
Poaceicola arundinis W.J. Li, Camporesi, D.J. Bhat &
K.D. Hyde, Mycosphere 6 (6): 681 (2015)
Facesoffungi number: FoF02383, Fig. 63
Saprobic on dead stem of Dactylis sp. Sexual morph
Ascomata 164–230 lm high 9 210–290 lm wide diam.
(
x = 200 9 235 lm, n = 10), superficial, solitary, scattered, or sometimes gregarious, globose, black to dark
brown, ostiole central. Ostioles 70–85 lm high 9 87–
110 lm diam. (
x = 80 9 100 lm, n = 5), papillate, dark
brown, smooth, without periphyses. Peridium 21–35 lm
wide., up to 45 lm wide at the apex, composed of 6–7 layers
of textura angularis, outer region heavily pigment, cells 4–
8 lm wide, inner layer composed of hyaline gelatinous cells,
thin, merging with pseudoparaphyses. Hamathecium comprising numerous, long, 2–3 lm (n = 40) wide, transversely
septate, branched, cellular pseudoparaphyses. Asci 65–
105 9 8–19 lm (
x = 87 9 13 lm, n = 20), 7–8-spored,
bitunicate, cylindric-clavate, with short bulbose pedicel,
apically rounded, with an ocular chamber up to 1–2 lm
wide 9 1–2 lm high. Ascospores 23–34 9 4.5–8 lm
(
x = 29 9 6 lm, n = 30), bi-seriate or overlapping, hyaline when immature, pale brown to yellowish at maturity,
fusiform, tapering towards the ends, (5–)8–9-distoseptate,
slightly contricted at third septa, third cell from apex
enlarged, smooth-walled, surrounded by thick, distinct
mucilaginous sheath. Asexual morph Coelomycetous.
Conidiomata 100–150 lm high, 100–200 lm diam., pycnidioid, dark brown, solitary or aggregated, semi-immersed,
111
unilocular, globose, papillate. Pycnidial wall comprised of
cells of textura angularis, gradually merging with the outer,
surrounding layers of brown, textura oblita. Ostiole central.
Conidiophores reduced to conidiogenous cells. Conidiogenous cells enteroblastic, phialidic, hyaline, smooth-walled, discrete. Conidia 30–40 9 6.5–10 lm, pale brown,
cylindrical, flexuous, up to 8-euseptate, slightly curved,
smooth-walled, with middle cells wider than end cells, guttulate, with an acute apex, truncate at the base (Li et al. 2015).
Culture characteristics: Colonies on PDA reaching
43 mm diam. after 14 days at 16 °C, surface white, with
moderate aerial mycelium, fluffy, fimbriate margins;
reverse white at the edges, dark green at the center, radiating light green outwardly, dense, circular, flattened,
margin rough, not pigmented.
Material examined: ITALY, Trento [TN], Forte
Strino—Vermiglio, on dead stem of Dactylis sp. (Poaceae), 12 September 2014, E. Camporesi, IT 541 (MFLU
16-1336); KUN HKAS 94614, living culture, MFLUCC
14-1060, KUMCC 16-0029.
Notes: Based on the combined gene analysis, Poaceicola arundinis (MFLUCC 14-1060) has a close relationship with P. arundinis (MFLUCC 15-0702), reported from
Arundo plinii (98 % ML, 100 % PP support). Poaceicola
arundinis (MFLUCC 14-1060) is similar to P. elongata,
but relatively smaller than the epitype (Ariyawansa et al.
2014b). Li et al. (2015) introduced P. arundinis from the
asexual morph and the phylogeny support them to be the
same strain as our sexual morph. We therefore introduce
the sexual morph to accommodate P. arundinis.
Pseudophaeosphaeria Jayasiri, Camporesi & K.D. Hyde,
gen. nov.
Index Fungorum number: IF552207; Facesoffungi
number: FoF02345
Etymology: Named because of its morphological similarity to the genus Phaeosphaeria.
Saprobic on Rubus idaeus L. Sexual morph Ascomata
scattered, immersed to slightly erumpent through host tissue, visible as small irregular black spots on the host surface, uniloculate, globose to subglobose, glabrous, dark
brown, ostiole central, with a minute papilla. Peridium
wide, thin-walled, of equal thickness, composed of 3–5
layers of flattened cells, outer layer dark brown, inner layer
of pseudoparenchymatous cells, arranged in textura angularis. Hamathecium composed of numerous, wide, filiform,
distinctly septate, frequently anastomosing, narrow, cellular pseudoparaphyses, embedded in mucilaginous matrix
between and above asci. Asci 8-spored, bitunicate, fissitunicate, broadly cylindrical, subsessile with minute knoblike pedicel, apically rounded, with a well-developed
ocular chamber. Ascospores overlapping 1–3-seriate, hyaline, phragmosporous, narrowly fusiform, mostly 3-septate,
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112
indistinctly constricted at the septa, slightly curved,
smooth-walled. Asexual morph Undetermined.
Type species: Pseudophaeosphaeria rubi Jayasiri,
Camporesi & K.D. Hyde
Notes: Phylogenetic analyses of combined LSU, SSU
and ITS sequence data indicate that Pseudophaeosphaeria
is a distinct genus in Phaeosphaeriaceae, which forms a
clade sister to Paraphoma, Setomelanomma and
Xenoseptoria clades. Pseudophaeosphaeria, differs from
Setomelanomma in having irregular ascomata and narrowly fusiform ascospores, while Setomelanomma has
regular ascomata and ellipsoidal to broadly fusiform
ascospores. Paraphoma and Xenoseptoria are asexual
genera of this family. Pseudophaeosphaeria is morphologically most closely related to Phaeosphaeria sensu
stricto, which has immersed to slightly erumpent, irregular, black ascomata, broadly cylindrical asci with small
subsessile knob-like pedicels, and narrowly fusiform,
3-septate ascospores. However, this relationship is not
supported by sequence data, as Pseudophaeosphaeria is
phylogenetic distinct from Phaeosphaeria sensu stricto
(Fig. 52). Pseudophaeosphaeria rubi stands phylogenetically apart other genera with relatively high support
(Fig. 52).
Pseudophaeosphaeria rubi Jayasiri, Camporesi & K.D.
Hyde, sp. nov.
Index Fungorum Number: IF552174; Facesoffungi
number: FoF02221, Fig. 64
Etymology: The specific epithet rubi is based on the host
genus from which the taxon was collected.
Holotype: MFLU 15-1400
Saprobic on Rubus idaeus L. Sexual morph Ascomata
105–145 lm high, 125–135 lm diam., scattered, immersed
to slightly erumpent through host tissue, visible as small
irregular black spots on the host surface, uniloculate, globose to subglobose, glabrous, dark brown, ostiole central,
with a minute papilla. Peridium 12–18 lm wide, thinwalled, of equal thickness, composed of 3–5 layers of
flattened cells, outer layer dark brown, inner layer pseudoparenchymatous cells, arranged in textura angularis.
Hamathecium composed of numerous, 1.8–2.4 lm wide,
filiform, distinctly septate, frequently anastomosing, narrow cellular pseudoparaphyses, embedded in mucilaginous
matrix between and above asci. Asci 59–75 9 7–9 lm
(
x = 66.5 9 8.2 lm, n = 20), 8-spored, bitunicate, fissitunicate, broadly cylindrical, subsessile with minute knoblike pedicel, apically rounded, with a well-developed
ocular
chamber. Ascospores 17–21 9 2.3–3.9 lm
(
x = 18.6 9 2.8 lm, n = 30), overlapping 1–3-seriate,
hyaline, phragmosporous, narrowly fusiform, mostly
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Fig. 75 Helicoma guttulatum (holotype). a Conidiophores with c
attached apical conidium on natural substrate. b, c Conidiophores
with conidia at the apex. d Conidiogenous cell with conidiophores.
e Germinating conidium. f–h Conidia. i–j Colonies on MEA from
above and below. Scale bars a = 200 lm, b, c, e–h = 20 lm,
d = 10 lm, i–j = 20 mm
3-septate, slightly constricted at the septa, slightly curved,
smooth-walled. Asexual morph Undetermined.
Culture characteristics: Ascospores germinating on
MEA within 36 h. Colonies on MEA, reaching 2.2 cm
diam. in 1 week at 28 °C. Mycelium superficial, felty,
gummy, pale grey to black. No asexual morph formed in
culture.
Material examined: ITALY, Forlı̀-Cesena, Passo la
Calla—Santa Sofia, on dead branch of Rubus idaeus
(Rosaceae), 28 January 2014, Erio Camporesi IT 1680
(MFLU 15-1400, holotype), (isotype in KUN), ex-type
culture MFLUCC 14-0259, KUMCC.
Wojnowicia Sacc.
The genus Wojnowicia was introduced by Saccardo
(1892) to accommodate W. hirta. The sexual morph of
Wojnowicia has not been reported. Presently, there are 14
species epithets in Wojnowicia (Index Fungorum 2016)
which are coelomycetous fungi. However, W. graminis
(McAlpine) Sacc. & D. Sacc. and W. tenella Pat. have been
synonymized under W. hirta (Sutton 1980) and W. buxi
Bertault & Malençon under W. ephedrae Hollós (Farr and
Bills 1995). Wojnowicia bryophila Racov., W. exilis
(Corda) Sacc. & Traverso, W. viburni Wijayaw. et al. and
W. lophostoma (Höhn.) Sacc. were excluded from Wojnowicia (Sutton 1980; Farr and Bills 1995; Wijayawardene et al. 2013; Crous et al. 2015a). Li et al. (2016)
introduced and illustrated W. spartii based on morphological characters and phylogenetic analysis. In the present
study, the sexual morph of Wojnowicia is described and
eight species are accepted in the genus.
Wojnowicia italica Qing Tian, Camporesi & K.D. Hyde,
sp. nov.
Index Fungorum number: IF552183; Facesoffungi
number: FoF02254, Fig. 65
Etymology: In reference to the occurrence of this species
in Italy.
Holotype: MFLU 14-0732.
Colonies growing on a dead branch of Spartium junceum L. Sexual morph Ascomata 230–260 9 130–
165 lm (
x = 240 9 150 lm, n = 7), immersed or semiimmersed and erumpent in the host, solitary, scattered,
sometimes gregarious, subglobose to globose, black, up to
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114
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Fig. 76 Neoacanthostigma septoconstrictum (MFLU 16-1134).
a Conidia arise directly from hyphal cells on natural substrate.
b Conidiophores with conidia. c–e Conidia. f Germinating spore. g,
h Colonies on MEA from above and below. Scale bars a = 200 lm,
b–e = 100 lm, f = 50 lm, g–h = 30 mm
200 lm high. Ostiole inconspicuous. Peridium thick at the
apex 30–60.5 lm (
x = 42 lm, n = 7), thinner at the base
and sides 17–30 lm (
x = 24 lm, n = 7), one-layered,
composed of brown to black, thin-walled, cells of textura
angularis. Asci 60–93 9 15–20 lm (
x = 77 9 16 lm,
n = 10), 8-spored, bitunicate, clavate, short-pedicellate,
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115
Fig. 77 Tubeufia hyalospora (holotype). a Conidia arise directly
from hyphal cells on natural substrate. b, c Conidiophores with
conidia. d–f Conidia. g Germinating spore. h, i Colonies on MEA
from surface and reverse. Scale bars a = 100 lm, b, c = 50 lm, d–
g = 20 lm, h, i = 30 mm
apically rounded, with an inconspicuous ocular chamber.
Hamathecium consisting of 2–3 lm broad long, colourless,
branched, pseudoparaphyses, with transverse sept. Ascospores 18–27 9 5–8 lm (
x = 22 9 8 lm, n = 10),
overlapping uni-seriate or bi-seriate, light brown to brown,
ellipsoid to fusiform, slighty curved, muriform, with 4–7
transverse septa, and one longitudinal septum in some
central cells, constricted at the septa, end cells conical,
smooth-walled, lacking a mucilaginous sheath. Asexual
morphs Coelomycetous. Conidiomata pycnidial, dark
brown, scattered, immersed to semi-immersed, globose to
subglobose, glabrous, ostiolate. Ostiole centrally located,
papillate. Wall of conidiomata composed of 4–5-cell layers. Conidiogenous cells phialidic, hyaline, integrated,
flask-shaped, thick-walled. Conidia dark brown, fusiform
or cylindrical, 7–12-septate.
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Culture characteristics: Ascospore germinating on PDA
within 12 h. Colonies on PDA reaching 10 mm diam. in
7 days at 16 °C. Mycelium superficial, initially white, later
becoming light brown or grey, hairy, with entire edge.
Material examined: ITALY, Forlı̀-Cesena [FC] Province, Teodorano-Meldola, on dead stem of Spartium
junceum (Fabaceae), 15 December 2012, Erio Camporesi
(MFLU 14-0732, holotype), (HKAS 94522, isotype); extype living cultures, MFLUCC 13-0447, KUMCC 16-0017.
Notes: The sexual morph of Wojnowicia had not been
previously reported. However, we found the sexual morph
of W. italica. Its placement in Wojnowicia is supported by
Fig. 78 Tubeufia roseohelicospora (holotype). a Conidia arise
directly from hyphal cells on natural substrate. b, c Conidiophores
with conidia. d–f Conidia. g Germinating spore. h, i Colonies on
MEA from above and below. Scale bars a = 200 lm, b–f = 20 lm,
g = 50 lm, h, i = 20 mm
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117
Fig. 79 Phylogram generated from maximum likelihood analysis
based on combined LSU and SSU sequence data. Maximum
likelihood bootstrap values greater than 50 % are shown above the
branches. The ex-type strains are in bold and the new isolate is in
blue. The tree is rooted with Gloniopsis praelonga
phylogenetic analyses of combined ITS, LSU and SSU
sequence data.
Comoclathris italica Tibpromma, Camporesi & K.D.
Hyde, sp. nov.
Index Fungorum number IF552245; Facesoffungi number: FoF02387, Fig. 67
Etymology: refers to the name of the country where the
holotype was collected.
Holotype: MFLU 15-1491.
Saprobic on dead stem of Thalictrum sp. Sexual morph
Ascomata 195–215 lm high 9 225–250 lm diam.
(
x = 203 9 230 lm, n = 5), globose to subglobose, solitary, dark brown to black, immersed, black, smooth, ostiolar canal filled with sparse periphyses. Peridium 6–27 lm
wide, comprising 3–4 layers of brown to reddish-brown
Pleosporaceae Nitschke
The family Pleosporaceae was introduced by Nitschke
(1869) and Pleospora is the type genus of this family.
Pleosporaceae is the largest family in the Pleosporales and
is representative of the order (Zhang et al. 2012b; Hyde
et al. 2013; Wijayawardene et al. 2014b). A backbone tree
for Pleosporaceae was provided by Ariyawansa et al.
(2015c) and this is updated in Fig. 66. Taxa of this family
are pathogenic or saprobic on wood and dead herbaceous
stems or leaves (Ariyawansa et al. 2015c).
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b Fig. 80 Pleurotrema thailandica (holotype). a Appearance of
ascomata on host surface. b Close up of clypeus of ascomata. c, d
Vertical sections through ascomata. e Peridium. f–i Asci. j Apical
thickening. k Pseudoparaphyses. l–o Ascospores with sheath. p Germinating ascospore. Scale bars b = 1 mm, c, d, f–i = 100 lm, e, l–
p = 20 lm, j = 10 lm
cells of textura angularis. Hamathecium comprising 1.8–
3.7 lm wide, dense, septate, hyaline, pseudoparaphyses.
Asci 75–115 9 22–29 lm (
x = 94 9 25 lm, n = 10), 8spored, bitunicate, fissitunicate, cylindric-clavate, short
pedicellate, thick-walled at the apex, with a minute ocular
chamber. Ascospores 20–29 9 9–13 lm (
x = 24 9
11 lm, n = 20), overlapping 1–2-seriate, hyaline when
young, becoming golden yellow at maturity, muriform, 5–7
transverse septate with 2–3 longitudinal septa in all cells
and rarely in end cells, slightly constricted in the middle,
conical and narrowly rounded at the ends, surrounded by
large mucilaginous sheath. Asexual morph Undetermined.
Culture characteristics: Colonies on MEA 60 mm diam.
after 4 weeks at 16 °C, circular colony with entire edge,
smooth, raised on surface, white-grey.
Material examined: ITALY, near Mezzana, province of
Trento [TN], dead stem of Thalictrum sp. Tourn. ex L.
(Ranunculaceae), 16 July 2014, Erio Camporesi, IT1998B
(MFLU 15-1491, holotype); ex-type living culture,
MFLUCC 15-0073); ibid. (HKAS 94599 bis, paratypes).
Notes: Comoclathris lanata is the type species of Comoclathris which was introduced by Clements (1909). The
genus is characterized by ascomata with circular lid-like
openings and applanate, reddish-brown to dark reddishbrown, muriform ascospores, with a single longitudinal
septum (Shoemaker and Babcock 1992). The phylogenetic
classification in this study shows that Comoclathris italica
clusters with C. spartii (Fig. 66), although it is differentiated from C. spartii based on molecular phylogeny and
morphological support. Our new taxon is characterized by
ascomata with ostioles and golden yellow ascospores with
conical and narrowly rounded ends, while C. spartii has
yellow to pale brown ascospores with broadly fusiform,
obtuse ends, constricted at primary septum (Crous et al.
2014).
Roussoellaceae J.K. Liu et al.
Roussoellaceae was introduced by Liu et al. (2014) and
is typified with Roussoella. This family is characterized by
immersed, clypeate, gregarious ascostromata, cylindrical,
bitunicate asci, and brown, 2-celled ascospores, and is
phylogenetically close with Biatriosporaceae and Pleomassariaceae (Liu et al. 2014; Wijayawardene et al.
2014a). Appendispora, Cytoplea, Elongatopedicellata,
Neoroussoella, Roussoella and Roussoellopsis are placed
in this family (Ariyawansa et al. 2015a), and members
mostly occur on bamboo and palms in terrestrial habitats.
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In this study we introduce a new species of Neoroussoella
with support from molecular data (Fig. 68).
Neoroussoella lenispora J.F. Zhang, J.K. Liu, K.D. Hyde
& Z.Y. Liu, sp. nov.
Index Fungorum number: IF552324; Facesoffungi
number: FoF02443, Fig. 69
Etymology: Name reflects the ascospores being smoothwalled, from the Latin lenis referring to smooth and spora.
Holotype: GZAAS 16-0011.
Saprobic on decaying branch. Sexual morph Ascostromata semi-immersed to erumpent, clypeate, globose
to subglobose, solitary to gregarious, black, coriaceous,
centrally ostiolate, visible as black, dome-shaped spots.
Peridium up to 22–45 lm, composed of an outer stratum of
dark brown, thick-walled cells fusing with host tissue, and
an inner stratum of pale brown, thin-walled cells arranged
in a textura prismatica. Hamathecium comprising 1.4–
2.8 lm wide, anastomosing pseudoparaphyses, branched at
apex, and embedded in a gelatinous matrix. Asci (74–)89–
113(–127) 9 (5.9–)6.5–7(–9) lm (
x = 101 9 6.8 lm,
n = 20), 8-spored, bitunicate, cylindrical, long pedicellate,
with a furcate pedicel, apically rounded with a minute
ocular chamber. Ascospores (10–)12–14.5(–16) 9 (3.5–)
4.5–5 lm (
x = 13 9 4.5 lm, n = 30), uni- to bi-seriate or
overlapping, ellipsoidal to fusiform, 2-celled, constricted at
the septum, pale brown to brown, guttulate when young,
smooth-walled, without any mucilaginous sheath and
appendages. Asexual morph Undetermined.
Culture characteristics: Ascospore geminating on WA
within 12 h and colonies on PDA reaching 24 mm diam. in
10 days at 25 °C; circular, irregular, aerial hypha dense,
raised, white to pale brown in forward and rufous in
reverse. Mycelium superficial to immersed in/on media,
with branched, smooth-walled hyphae.
Material examined: CHINA, Guizhou, Libo County,
Maolan National Natural Preserve, on decaying branch of
unidentified host, 8 July 2015, J.F. Zhang, MLC-13
(GZAAS 16-0011, holotype), ex-type living culture,
GZCC16-0020.
Notes: The new taxon is typical of species in the
family Roussoellaceae, and the sequence data also shows
that it is most phylogenetically close to Neoroussoella
bambusae (Fig. 68). However, it differs in the morphology of ascostromata. In N. lenispora ascostroma are
globose or subglobose, while in N. bambusae, they are
flattened at the base. The asci and ascospores also differ
(asci: 101 9 6.8 lm vs. 72.2 9 5.6 lm; ascospores:
13 9 4.5 lm vs. 9 9 3 lm). The ascospores of N. bambusae are ornamented with longitudinal ribs and surrounded by a mucilaginous sheath, whereas those of
N. lenispora lack a sheath and are smooth-walled when
mature.
123
120
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Fungal Diversity (2016) 80:1–270
b Fig. 81 Phylogram generated from maximum likelihood analysis
based on combined LSU and SSU sequence data of selected taxa.
Maximum likelihood bootstrap support values greater than 60 % are
given above the nodes. The type strains of each genera are in bold and
the new isolates are in blue bold. The tree is rooted with
Arthoniomycetes taxa
Torulaceae Corda
= Dendryphiaceae Corda, Icon. fung. (Prague) 4: 32
(1840)
Corda introduced the family Torulaceae (Sturm 1829).
Crous et al. (2015a) revisited this family, provided
molecular data for four species of Torula; accepted two
genera (Dendryphion and Torula) and placed Torulaceae in
the order Pleosporales. Su et al. (2016a) introduced a new
genus Neotorula in this family and provided molecular data
for three species of Dendryphion including two new species. Presently, there are three genera accepted in the
family Torulaceae (Su et al. 2016a). Torulaceae is known
only by asexual taxa characterized by erect, micro- or
macronematous conidiophores, with or without apical
branches, and doliiform to ellipsoid or clavate, brown,
smooth to verruculose, mono- to polyblastic, often cupulate, conidiogenous cells and subcylindrical, phragmosporous, acrogenous, brown, dry, smooth to verrucose
conidia, characteristically produced in branched chains
(Crous et al. 2015a; Su et al. 2016a). A phylogenetic tree
for the family is presented in this paper (Fig. 70).
Neotorula Ariyawansa et al.
Neotorula was established by Su et al. (2016a) to
accommodate a Torula-like species collected from freshwater habitats in China, based on both morphology and
phylogeny. Neotorula shares similarities with Torula in
having clavate to subcylindrical, brown conidia, with
rounded ends that are borne in branched chains. The young
conidia of N. aquatica are pale green, and are pale yellow
in N. submersa. The conidiogenesis in Neotorula is polytretic, whereas Torula has polyblastic cupulate and darkcoloured conidiogenous cells. Neotorula has distinct
conidiophores which comprise a few cells, while comprise
a single or few cells in Torula. Our new species have
polytretic, macronematous conidiophores. In this study, we
introduce a new species in Neotorula based on morphological characters. The phylogenetic analysis also supports
our taxon as a new species (Fig. 70).
Neotorula submersa Z.L. Luo, H.Y. Su & K.D. Hyde, sp.
nov.
Index Fungorum number: IF552138; Facesoffungi
number: FoF02217, Fig. 71
Etymology: With reference to the submerged habitat.
Holotype: HKAS 92660.
121
Saprobic on submerged decaying wood. Sexual morph
Undetermined. Asexual morph Colonies on the substrate
superficial, effuse, gregarious, hairy, dark brown. Mycelium immersed, composed of septate, branched, thin-walled, smooth, pale brown hyphae. Conidiophores
mononematous, macronematous, branched or unbranched,
septate, erect, straight or flexuous, smooth, pale brown,
cylindrical, 2–5-septate, 18–48 lm (
x = 33 lm, SD = 15,
n = 10) long, 4–5 lm (
x = 4.5 lm, SD = 0.5, n = 10)
wide. Conidiogenous cells mono- to polytretic, integrated
or discrete, terminal, pale brown, doliiform or lageniform.
Conidia acrogenous, in short chains, dry, clavate to subcylindrical, rounded at the apex, 2–4-septate, dark bands at
the septa, verruculose, pale yellow when young, tawny to
brown when mature, 16–22 lm (
x = 19 lm, SD = 3,
n = 25) long, 5–6 lm (
x = 5.5 lm, SD = 0.5, n = 25)
wide. Conidial secession schizolytic.
Material examined: CHINA, Yunnan Province, saprobic
on decaying wood submerged in Dulong River, May 2015,
X.C. Tao, HD 1–10–7 (HKAS 92660, holotype), ex-type
culture, KUMCC 15-0280, MFLUCC.
Notes: Neotorula submersa was collected from northwestern Yunnan Province. It differs from N. aquatica in
having thinner conidiophores (4–5 lm vs. 6–7 lm) and
longer conidia (16–22 lm vs. 11–16 lm). The conidia of
N. aquatica are pale green when young and brown when
mature, in N. submersa they are pale yellow when young
and tawny to brown when mature. Phylogenies also reveal
a close relationship between these two taxa (Fig. 70).
Tubeufiales Boonmee & K.D. Hyde
For Tubeufiales, we follow Boonmee et al. (2014)
Tubeufiaceae M.E. Barr
The most recent treatments of the family and order are
Boonmee et al. (2014) and their findings are followed here.
In this study we provide and updated backbone tree for
Tubeufiaceae (Fig. 72) and introduce Chlamydotubeufia
helicospora, Helicoma guttulatum, Neoacanthostigma
septoconstrictum, Tubeufia hyalospora and T. roseoelicospora. A reference specimen is also designated for
Aquaphila albicans.
Aquaphila Goh et al.
Aquaphila is a genus known only from freshwater
habitats and is typified by A. albicans which is connected
with the sexual morph Tubeufia asiana Sivichai & K.M.
Tsui (Tsui et al. 2007). Boonmee et al. (2014) synonymized
T. asiana under Aquaphila and named Aquaphila asiana
based on phylogenetic evidence.
Aquaphila albicans Goh, K.D. Hyde & W.H. Ho, Mycol.
Res. 102(5): 588 (1998)
123
122
Fig. 82 Alloarthopyrenia italica (holotype). a Herbarium material.
b, c Ascostromata on host surface. d Section of ascostroma. e Ostiole.
f Section of peridium. g Pseudoparaphyses. h–i Ocular chamber
(i ocular chamber staining in Melzer’s reagent). j Immature asci.
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Fungal Diversity (2016) 80:1–270
k–l Mature asci. m Ascus staining in Melzer’s reagent. n–r
Ascospores. Scale bar b = 200 lm, c = 100 lm, d = 20 lm,
e–m = 50 lm, h–i = 5 lm, n–r = 10 lm
Fungal Diversity (2016) 80:1–270
Index Fungorum number: IF443558; Facesoffungi
number: FoF02356, Fig. 73
Reference specimen: MFLU 16-1136
Saprobic on decaying wood in flowing freshwater
stream. Colonies on natural substrate effuse, translucent,
chalky white or pale yellowish when dry. Mycelium partly
immersed in woody substrate and partly superficial, consisting of septate, branched, hyaline, smooth, thin-walled
hyphae. Sexual morph Undetermined. Asexual morph
hyphomycetous; phragmosporous. Conidiophores arising
singly as lateral branches from procumbent hyphae, up to
33 lm long, 4–6 lm wide, simple or branched, thin-walled
and smooth, indistinctly septate, flexuous, hyaline. Conidiogenous cells monoblastic or polyblastic, sympodial with
cylindrical denticles, hyaline, with numerous integrated
tiny pegs. Conidia 70–86 9 8–10 lm, holoblastic,
acrogenously borne on denticles, solitary, obclavate, predominantly fusoid to sickle-shaped, sometimes sigmoid,
slightly curved and acute at both ends, basal cell obconical,
up to 14-euseptate, slightly constricted at the septa, hyaline
to pale yellowish, densely guttulate, smooth-walled.
Culture characteristics: Conidia germinating on water
agar (WA) within 12 h and germ tubes produced at both
ends. Colonies on malt extract agar (MEA), reaching
10 mm in 4 weeks at 28 °C, brown to dark brown in MEA
media. Mycelium superficial and partially immersed,
branched, septate, hyaline to pale brown, smooth.
Material examined: THAILAND, Prachuap Khiri Khan,
Bang Sapan, Ron Thai, N11°140 47.02800 E99°190 59.68200 ,
elev. ca. 171 msl., on decaying wood in flowing freshwater
stream, 30 July 2015, K.D. Hyde TF04 (MFLU 16-1136,
reference specimen designated here), MFLU 16-1138,
BBH 41052,; ex-type living culture, MFLUCC 16-0010,
TBRC.
Sequence data: LSU = KX454166 and KX454168,
ITS = KX454165 and KX454167.
Notes: Two new collections of Aquaphila albicans
(MFLU 16-1136 and MFLU 16-1138) were found on
submerged wood in southern Thailand. Their morphology
is identical to the type species A. albicans (HKU (M) 2856)
collected in Queensland, Australia (Goh et al. 1998).
Additionally, phylogenetic analysis placed the two new
specimens in the same clade with other A. albicans strains
and its sexual morph A. asiana (strain BCC3463) with high
support (Fig. 72). We therefore designate MFLU 16-1136
as a reference specimen (sensu Ariyawansa et al.
2014a, b, c) for Aquaphila albicans. We do not designate
an epitype as this was not collected from Australia.
Chlamydotubeufia Boonmee & K.D. Hyde
Chlamydotubeufia is a genus recognized by black pigmented with multi-septate dictyochlamydosporous conidia
and a typical sexual morph (Boonmee et al. 2011).
123
Chlamydotubeufia a small genus with only four species
presently recorded and typified by Ch. huaikangplaensis
Boonmee & K.D. Hyde (Index Fungorum 2016).
Chlamydotubeufia helicospora Boonmee, Y.Z. Lu & K.D.
Hyde, sp. nov.
Index Fungorum number: IF552217; Facesoffungi
number: FoF02357, Fig. 74
Holotype: MFLU 16-1338.
Etymology: ‘helicospora’ referring to asexual
helicospores.
Saprobic on woody substrate. Mycelium partly immersed,
partly superficial, white, septate, sparsely branched hyphae,
with masses of crowded conidia. Sexual morph Undetermined. Asexual morph Conidiophores 15–25 lm
long 9 4–6 lm wide (
x = 18 9 5 lm, n = 10), pale
brown, macronematous, erect, short, smooth-walled. Conidiogenous cells 11–33 lm long 9 4–5 lm wide
(
x = 18 9 4.5 lm, n = 10), monoblastic, integrated, each
with single conidium. Conidia 63–119 lm diam. and conidial filament 5–7 lm wide (
x = 80 9 6 lm, n = 20),
405–546 lm long, loosely coiled 1–2.5 times, rounded at
apical end, up to 68-septate, slightly constricted at septa,
hyaline, smooth-walled.
Culture characteristics: Conidia germinating on water
agar (WA) within 8 h and germ tubes produced from
conidia. Colonies on malt extract agar (MEA) reaching
4 mm in 1 week at 28 °C, brown to dark brown. Mycelium
superficial and partially immersed, branched, septate,
hyaline to pale brown, smooth.
Material examined: THAILAND, Uttaradit, Laplae, Mae
Phun, Ban Ton Klua, on decaying wood in flowing freshwater stream, 24 October 2015, Saranyaphat Boonmee,
UTD15–1 (MFLU 16-1338, holotype, BBH 41053, isotype); ex-type living culture, MFLUCC 16-0213, TBRC.
Notes: According to phylogenetic analysis, Chlamydotubeufia helicospora clusters in the Chlamydotubeufia
clade with high support. The molecular analyses (Fig. 72)
confirms C. helicospora as a new taxon that is basal to
C. khunkornensis, while C. chlamydospora and
C. huaikangplaensis constitutes a different sublineage with
very high statistical support. Hence, we assign our collection as a new species based on morphology and phylogeny
(Fig. 72). Morphologically, C. helicospora is characterized
by slender hyphae, moderately long conidiophores and
helicosporous conidia. The conidia are different from other
species in Chlamydotubeufia (Boonmee et al. 2011).
Helicoma Corda
The genus Helicoma was established by Corda with the
type species H. muelleri Corda. The genus is distinguished
by its relatively short, erect, thick, dark brown, smooth
conidiophores, holoblastic conidiogenous cells and
123
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Fungal Diversity (2016) 80:1–270
Neotrematosphaeria biappendiculata KTC 1124
Coelodictyosporium muriforme MFLUCC 13 0351
Lophiopoacea paramacrostoma MFLUCC 11 0463
ravennica MFLUCC 14 0005
Lophiohelichrysum helichrysi MFLUCC 15 0701
Guttulispora crataegi MFLUCC 13 0442
Platystomum crataegi MFLUCC 14 0925e
Lophiostoma compressum IFRD 2014
Lophiostoma macrostomum KT508
Biappendiculispora japonica MAFF 239452
79
Capulatispora sagittiformis JCM 15100
93
Platystomum scabridisporum BCC 22835
Trematosphaeria terricola SC 12
Paucispora quadrispora MAFF 239455
85
96
Dimorphiopsis brachystegiae CPC 22679
Vaginatispora aquatic MFLUCC 11 0083
98 Angustimassarina populi MFLUCC 13 0034
75 Massarina corticola CBS 154.93
100 Angustimassarina quercicola MFLUCC 14 0506
Angustimassarina acerina MFLUCC 14 0505
Exosporium stylobatum CBS 160.30
Camarographium carpini CBS 128781
99 Preussia lignicola CBS 264.69
94
Sporormia lignicola CBS 363.69
Preussia minima AFTOL ID1256
90 Preussia funiculate 2577
100
Preussia funiculate CBS 659.74
Preussia terricola AFTOL ID 282
74 Westerdykella dispersa CBS 50875
Westerdykella ornate CBS 379 55
98
97 Eremodothis angulate CBS 610.74
Westerdykella cylindrical CBS 454.72
Sporormia fimetaria Gr.81.194
100 Decaisnella formosa BCC 25617
97
Decaisnella formosa BCC 25616
100
Lignosphaeria thailandica MFLUCC 11 0376
Lignosphaeria fusispora MFLUCC 11-0377
100
Berkleasmium nigroapicale BCC 8220
Bahusandhika indica GUFCC 18001
100
Misturatosphaeria kenyensis GKM L100Na
Pseudoaurantiascoma kenyense GKM 1195
95
Misturatosphaeria minima ANM 933
Misturatosphaeria minima GKM 169N
Curreya grandicipis CBS 114272
Misturatosphaeria claviformis GKM 1210
Macrodiplodiopsis desmazieri MFLUCC 12 0088
100 Floricola striata JK 5678I
Floricola striata JK 5603K
Misturatosphaeria tennesseensis ANM 911
Pseudomisturatosphaeria cruciformis SMH 5151
Floricola viticola IT 2178
Misturatosphaeria aurantonotata GKM 1238
Misturatosphaeria radicans ATCC 42522
Ramusculicola thailandica MFLUCC 13 0284
Neomassarina thailandica MFLUCC 10 0552
1 Sigarispora
6
98
100
GU479796_Pontoporeia_biturbinata
93
Halotthia posidoniae BBH 22481
Mauritiana rhizophorae BCC 28866
Sulcosporium thailandica MFLUCC 12-0004
Neolophiostoma pigmentatum MFLUCC 10 0129
Brunneoclavispora bambusae MFLUCC 11- 0177
Roussoellopsis tosaensis KT 1659
Roussoella pustulans KT 1709
Roussoella hysterioides HH 26988
95 Arthopyrenia salicis CBS 368.94
Arthopyrenia salicis CBMAI1330
74
Roussoella angustior MFLUCC 15-0186
Roussoella magnatum MFLUCC 15-0185
97
Neoroussoella bambusae MFLUCC 11-0124
Sporidesmium_australiense
100 Torula herbarum CBS 111855
98
Torula herbarum CBS 379.58
98
Torula herbarum CBS 220.69
100 Dendryphion europaeum CPC 22943
Dendryphion europaeum CPC 23231
Parameliola acaciae MFLU 15–0378
100
Parameliola dimocarpi MFLU15–0045
100 Pseudocoleodictyospora tectonae MFLUCC 12 0387
Pseudocoleodictyospora_tectonae MFLUCC 12 0385
72
Pseudocoleodictyospora thailandica MFLUCC 12 0565
100
Pseudocoleodictyospora sukhothaiensis MFLUCC 12 0554
Subglobosporium
tectonae MFLUCC 12 0393
100
Subglobosporium tectonae MFLUCC 12 0390
84 Occultibambusa bambusae MFLUCC 13-0855
87 Occultibambusa bambusae MFLUCC 11-0394
98
Occultibambusa chiangraiensis MFLUCC 16–0380
Occultibambusa aquatica MFLUCC11–0006
Occultibambusa pustula MFLUCC 11-0502
Versicolorisporium triseptatum JCM 14775
Occultibambusa fusispora MFLUCC 11-0127
100
Seriascoma didymospora_MFLUCC 11 0179
Seriascoma didymospora MFLUCC 11 0194
Neooccultibambusa chiangraiensis MFLUCC 12 0559
94_Biatriospora mackinnonii CBS 674.75
100
Nigrograna mackinnonii E5202H
Biatriospora marina CY_1228
100
Paradictyoarthriinium diffractum MFLUCC 12-0557
Paradictyoarthriinium tectonicola MFLUCC 13-465
Stagonospora pseudocaricis HKUCC 10833
Neottiosporina paspali CBS 331.37
Corynespora leucadendri CBS 135133
Corynespora olivacea CBS 114450
Massarina eburnea CBS 473.64
Pseudodidymosphaeria phlei MFLUCC14-1061
Deniquelata barringtoniae MFLUCC 110422
94
49
Kalmusia brevispora KT 1466
85
Montagnula aloes CPC 19671
100 Pseudocamarosporium pini MFLUCC 14-1091
Didymosphaeria rubi-ulmifolii MFLUCC 14-0023
96 Bambusicola splendida MFLUCC 11-0439
Bambusicola massarinia MFLUCC 11-0389
100
Latorua caligans CBS 576.65
Latorua grootfonteinensis CBS 369.72
Katumotoa bambusicola KT 1517a
86
Lentithecium fluviatile CBS 122367
Pseudomonodictys tectonae MFLUCC 12 0552
Trematosphaeria pertusa CBS 122371
79
Falciformispora lignatilis BCC 21118
Macrodiplodiopsis desmazieri CPC 24971
99 Macrodiplodiopsis desmazieri CBS 125026
Macrodiplodiopsis desmazieri CPC 24648
100 Moroshperia ramunculicola BCC 18404
85
Moroshperia ramunculicola BCC 18405
Helicascus_nypae BCC 36752
Longiostiolum tectonae MFLUCC 12 0562
Thyridaria rubronotata CBS 385.39
Massariosphaeria phaeospora CBS 611.86
100
Julella avicenniaea BCC 18422
Julella avicenniae BCC 20173
100 Cucurbitaria berberidis CBS 363.93
82
Cucurbitaria berberidis CBS 394.84
Pyrenochaeta nobilis CBS 407.76
89 Camarosporium quaternatum DQ377884
Camarosporium aloes CPC 21572
Phaeosphaeria oryzae CBS 110110
94
79
Ophiosphaerella herpotricha AFTOL-ID 1569
Paraphoma radicina CBS 111.79
100 Neophaeosphaeria filamentosa CBS 102203
100 Neophaeosphaeria filamentosa CBS 102202
Neophaeosphaeria agaves CPC 21264
83 Dothidotthia symphoricarpi CPC 12929
Dothidotthia aspera CPC 12933
100 Coniothyrium palmarum CBS 758.73
Coniothyrium palmarum CBS 400.71
100 Alternariaster helianthi CBS 134020
Alternariaster helianthi CBS 327.69
Leptosphaeria maculans AFTOL-ID 277
75
Pleospora herbarum CBS 191.86
100
Alternaria_alternata AFTOL-ID 1610
Pyrenophora phaeocomes AFTOL-ID 283
72 Phoma exigua CBS 431.74
97 Leptosphaerulina australis CBS 317.83
Didymella exigua CBS 183.55
100 Ascocylindrica marina MD6011
Ascocylindrica mari MD6012
97
Corynespora smithii CABI 5649b
Corynespora cassiicola CBS 100822
72
Beverwykella pulmonaria CBS 283.53
100
Melanomma pulvis-pyrius CBS 371.75
Byssosphaeria jamaicana SMH 1403
77
Lindgomyces ingoldianus ATCC 200398
73
Lindgomyces rotundatus KH 114
91
Massariosphaeria typhicola MAFF 239218
Lolia aquatic MF644
97 Anguillospora longissimi CCMF 10304
Anguillospora_longissima CS869-1D
Amniculicola lignicola Ying01
Anguillospora longissima F00980
Amniculicola parva CBS 123092
Spirosphaera cupreorufescens A20
100
Amniculicola immersa CBS 123083
100
Repetophragma ontariense HKUCC 10830
96
Massariosphaeria grandispora CBS 613 86
70
Pseudomassariosphaeria bromicola MFLUCC 15–0031
Murispora rubicunda IFRD 2017
Hysterium angustatum CBS 236.34
100
0.05
123
Fungal Diversity (2016) 80:1–270
b Fig. 83 RAxML maximum likelihood phylogenetic tree based on a
LSU and SSU sequenced data from species of order Pleosporales.
Maximum likelihood bootstrap support values greater than 50 % are
shown in near the nodes. Some branches were shortened to fit the
page—these are indicated by two diagonal lines with the number of
times a branch was shortened indicated next to the lines. The new
isolates are in red. The tree is rooted with Hysterium angustatum
helicoid, hyaline, thick-walled, brown to dark brown
conidia forming from terminal, denticulate conidiophores.
Helicoma guttulatum Y.Z. Lu, Boonmee & K.D. Hyde, sp.
nov.
Index Fungorum number: IF552218; Facesoffungi
number: FoF02358, Fig. 75
Holotype: MFLU 16-1339
Etymology: ‘guttulatum’ referring to conidia containing
numerous guttules.
Saprobic on woody substrates. Mycelium composed of
partly immersed and partly superficial, pale brown, septate
hyphae. Sexual morph Undetermined. Asexual morph
Conidiophores 74–182(197) lm long, 4–6 lm wide
(
x = 120 9 5 lm, n = 20), macronematous, crowded,
erect, subhyaline to yellowish, brown towards the base,
septate, unbranched, smooth-walled. Conidiogenous cells
monoblastic to polyblastic, subhyaline to pale brown,
smooth-walled. Conidia 18–23 lm diam., and conidial
filament 6–8 lm wide (
x = 20 9 7 lm, n = 20), tightly
coiled 1–1 times, hyaline to pale brown, tapering toward
flat end, 8–9-septate, rounded at the apex, conico-truncate
at the base, smooth-walled.
Culture characteristics: Conidia germinating on water
agar (WA) within 12 h. Colonies on malt extract agar
(MEA) reaching 5 mm in 1 week at 28 °C, pale brown to
brown. Mycelium superficial and partially immersed,
branched, septate, hyaline to pale brown, smooth.
Material examined: THAILAND, Chiang Rai, Muang,
Ban Nang Lae Nai, on decaying wood in flowing freshwater stream, 28 November 2015, Yong-Zhong Lu and
Saranyaphat Boonmee, TUB02–1 (MFLU 16-1339, holotype, BBH 41054, isotype); ex-type living culture,
MFLUCC 16-0022, TBRC.
Notes: Helicoma guttulatum was collected from northern Thailand. Phylogenetic results recognize H. guttulatum
as belonging to the genus Helicoma as well as its recognition as a new species. Helicoma guttulatum segregates
from H. dennisii and H. inthanonense with full support in
the phylogenetic analysis (Fig. 72). Helicoma guttulatum
forms an asexual morph on the natural substrate, which is
morphologically similar to H. muelleri (Boonmee et al.
2014). However, its conidiophores (74–182 lm) are longer
and narrower (4–6 lm) than H. muelleri (80–154 lm) and
(7–10 lm). The conidial diameter (18–23 lm) is also
125
greater than H. moelleri (16–19 lm). Therefore, H. guttulatum is introduced here as a novel species.
Neoacanthostigma Boonmee et al.
The genus Neoacanthostigma was introduced to
accommodate three sexual taxa, characterized by dark
pigmented ascomata, surrounded by reddish-brown to dark
brown setae, bitunicate asci, fasciculate cylindrical and
hyaline ascospores, and having a helicosporous asexual
state (Boonmee et al. 2014). Here we collected Acanthostigma septoconstrictum Promp. & A.N. Mill. and
transfer it to Neoacanthostigma.
Neoacanthostigma septoconstrictum (Promp. & A.N.
Mill.) S. Boonmee & K.D. Hyde, Fungal Diversity 68(1):
279 (2014)
: Acanthostigma septoconstrictum Promp. & A.N.
Mill., Mycologia 102(3): 579 (2010)
Index Fungorum number: IF550683; Facesoffungi
number: FoF02360, Fig. 76
Reference specimen: MFLU 16-1134
Saprobic on decaying wood in flowing freshwater
stream. Mycelium partly superficial, partly immersed,
composed of brown, septate, sparsely branched hyphae,
with masses of crowded conidia. Sexual morph Undetermined. Asexual morph hyphomycetous; helicosporous.
Conidiophores brown, macronematous, erect, short,
smooth-walled. Conidiogenous cells monoblastic, integrated, each with single conidium. Conidia 115–155 lm
diam., filament 9–11 lm wide (
x = 139 9 10 lm,
n = 20), 775–920 lm long, loosely coiled 1–2 times,
rounded at apical end, up to 71-septate, not constricted at
septa, pale brown to brown, smooth-walled.
Culture characteristics: Conidia germinating on water
agar (WA) within 12 h and germ tubes produced from
conidia. Colonies growing on malt extract agar (MEA),
reaching 25 mm in 4 weeks at 28 °C, slightly convex, with
an undulate edge, pale yellow to pale brown, brown in
MEA media. Mycelium superficial and partially immersed,
branched, septate, hyaline to pale brown, smooth.
Material examined: THAILAND, Prachuap Khiri Khan,
Bang Sapan, Ron Thai, on decaying wood in flowing
freshwater stream, 30 July 2015, K.D. Hyde, KH02 (MFLU
16-1134, BBH 41051); living culture, MFLUCC 15–1248,
TBRC.
Notes: Neoacanthostigma was introduced by Boonmee
et al. (2014) with N. fusiforme as the type species. An
asexual morph has not been previously reported in this
genus. In this study, N. septoconstrictum was found in its
asexual form (Fig. 72) on decaying wood in a flowing
freshwater stream and is characterized by pale brown to
brown, large filament, multi-septate conidia. Phylogenetic
analysis placed our collection in a cluster with the sexual
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Fig. 84 Neomassarina thailandica (holotype). a Appearance of
ascostromata on the host surface. b Section through the ascostroma.
c Section through peridium. d Pseudoparaphyses stained in Melzer’s
reagent. e–h Asci. i–k Ascospores. l Ascospores becoming pale
brown at maturity. m, n Culture characteristics (m = from above,
n = from below). Scale bars b = 100 lm, c = 20 lm, d–h,
l = 10 lm, i–k = 5 lm
morph N. septoconstrictum (strain ANM 536.1) with high
support (Fig. 72). Therefore, we describe this fungus under
the sexual name N. septoconstrictum.
Tubeufia hyalospora Y.Z. Lu, Boonmee & K.D. Hyde, sp.
nov.
Index Fungorum number: IF552220; Facesoffungi
number: FoF02361, Fig. 77
Holotype: MFLU 16-1135
Etymology: ‘hyalospora’ referring to hyaline helicospores of asexual morph.
Saprobic on decaying woody in flowing freshwater
stream. Mycelium partly immersed, partly superficial,
composed of pale brown, septate, sparsely branched
Tubeufia Penz. & Sacc.
The genus Tubeufia is the type genus of the family
Tubeufiaceae. It is characterized by white, cream-pink to
brownish, vertically oblong to ovoid ascomata and cylindrical, fusiform to vermiform, multi-septate ascospores
(Barr 1980).
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127
Fig. 85 Phylogenetic tree of Aspergillus koreanus and related species
based on maximum parsimony analysis of a combined dataset from
RNA polymerase II (RPB2), ITS and 28S rDNA sequence data.
Sequences data from Talaromyces bacillisporus was used as the
outgroup taxon. Numbers at the nodes indicate the bootstrap values
([50 %) from 1000 replicates. The bar indicates the number of
substitutions per position. New taxa are in blue
hyphae, with masses of crowded conidia. Sexual morph
Undetermined. Asexual morph hyphomycetous; helicosporous. Conidiophores pale brown, macronematous,
erect, short, smooth-walled. Conidiogenous cells
monoblastic, integrated, each with single conidium. Conidia 16–33 lm diam., conidial filament 3–5 lm wide
(
x = 22 9 4 lm, n = 20), 110–225 lm long, coiled 2–
3 times, rounded at apical end, multi-septate, slightly
constricted at septa, hyaline, smooth-walled.
Culture characteristics: Conidia germinating on water
agar (WA) within 12 h and germ tubes produced from
conidia. Colonies growing on malt extract agar (MEA),
reaching 26 mm in 3 weeks at 28 °C, with an undulate
edge, pale brown to brown, dark brown, white at margin.
Mycelium superficial and partially immersed, branched,
septate, hyaline to pale brown, smooth.
Material examined: THAILAND, Prachuap Khiri Khan,
Bang Sapan, Ron Thai, on decaying wood in flowing
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Fig. 86 Phylogenetic tree of Aspergillus koreanus and related species
within the sect. Cremei based on maximum parsimony analysis of ITS
sequence data. Sequence data for Aspergillus tamarii was used as the
outgroup taxon. Numbers at the nodes indicate the bootstrap values
([50 %) from 1000 replicates. The bar indicates the number of
substitutions per position. New taxa are in blue
freshwater stream, 30 July 2015, K.D. Hyde, TF03 (MFLU
16-1135, holotype); ex-type living culture, MFLUCC
15-1250, TBRC.
Notes: Tubeufia hyalospora grouped within the Tubeufia
clade, but did not cluster with any known species (Fig. 72).
Tubeufia hyalospora forms an asexual morph on the natural
substrate, which is morphologically similar to T. tectonae
(Doilom et al. 2016). Its conidia (diam. 16–33 lm) are
narrower than those of T. tectonae (32–55 lm) and are
tightly coiled, but in T. tectonae most are loosely coiled or
uncoiled.
Holotype: MFLU 16-1133.
Saprobic on decaying woody in flowing freshwater
stream. Colonies on natural substrate, superficial with
partly immersed mycelium and bright, septate, sparsely
branched hyphae, with masses of crowded conidia. Sexual
morph Undetermined. Asexual morph Hyphomycetes,
helicosporous. Conidiophores macronematous, erect, short,
pale brown, smooth-walled. Conidiogenous cells
monoblastic, integrated, each with single conidium. Conidia 36–48 lm diam., filaments 5–7 lm wide
(
x = 41 9 6 lm, n = 20), 210–283 lm long, coiled 2–
3 times, tight to loose, rounded at apical end, 29–35septate, slightly constricted at septa, light pink on substrate,
hyaline when seen under light microscope, smooth-walled.
Culture characteristics: Conidia germinating on water
agar (WA) within 12 h and germ tubes produced from
conidia. Colonies growing on malt extract agar (MEA)
reaching 10 mm in 2 weeks at 28 °C, with an undulate
Tubeufia roseohelicospora Y.Z. Lu, Boonmee & K.D.
Hyde, sp. nov.
Index Fungorum number: IF02362; Facesoffungi number: FoF02362, Fig. 78
Etymology: ‘roseohelicospora’ referring to light pink
helicospores of asexual morph.
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129
Fig. 87 Phylogenetic tree of Aspergillus koreanus and related species
within the sect. Cremei based on maximum parsimony analysis of
combined dataset for beta tubulin (BenA), calmodulin (CaM), ITS
rDNA, 28S rDNA and RPB2 sequence data. Sequence data for
Aspergillus tamarii is used as the outgroup taxon. Numbers at the
nodes indicate the bootstrap values ([50 %) from 1000 replicates.
The bar indicates the number of substitutions per position. New taxa
are in blue
edge, pale brown to brown, dark brown. Mycelium superficial and partially immersed, branched, septate, hyaline to
pale brown, smooth-walled.
Material examined: THAILAND, Prachuap Khiri Khan,
Bang Sapan, Ron Thai, on decaying wood in flowing
freshwater stream, 30 July 2015, K.D. Hyde KH01 (MFLU
16-1133, holotype; BBH 41050, isotype); ex-type living
culture, MFLUCC 15-1247, TBRC.
Notes: Tubeufia roseoelicospora clusters in Tubeufia
sister to T. tectonae based on the phylogenetic analysis of
combined LSU and ITS sequence data (Fig. 72). The
conidiophores of T. roseoelicospora are tightly coiled
while those of T. tectonae are mostly loosely coiled or
uncoiled; its conidia filament is 5–7 lm wide, while that of
T. tectonae is 2–5 lm wide. On the natural substrate
colonies of T. roseohelicospora are light pink, while
T. tectonae colonies are hyaline to white. Therefore,
T. hyalospora is introduced here as a novel species based
on its morphological and phylogenetic differences from
other Tubeufia species.
Dothideomycetes family, incertae sedis
Pleurotremataceae Walt. Watson
Currently Pleurotremataceae comprises the genus Pleurotrema and Dyfrolomyces species may need transferring to
this genus (Maharachchikumbura et al. 2016). Whether the
freshwater species of Saccardoella belong to this family will
require further studies at the molecular level.
Pleurotrema Müll. Arg.
Watson (1929) introduced the family Pleurotremataceae based on Pleurotrema polysemum (Nyl.) Mull. Arg.
Eriksson and Hawksworth (1993) suspected Pleurotremataceae to be synonym of Pyrenulaceae. Pleurotrema was synonymized under Lithothelium as a member
of Pyrenulaceae by Aptroot (1991). Later, Harris re-
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130
Fig. 88 Aspergillus koreanus (holotype). a, d Colonies on malt
extract agar (MEA). b, e Colonies on yeast extract sucrose agar
(YES). e, f Colonies on Czapek yeast autolysate agar (CYA) (a–
c from above, d–f from below). g–i Developing bi-seriate conidial
heads on stalks. j Phialides finely differentiated on metulae covering
the entire surface of vesicle. k Conidiophore and phialides (green
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Fungal Diversity (2016) 80:1–270
arrow). l Detail of a metula bearing 3 phialides (red arrow). m Initial
young conidia (yellow arrow) developed at the tips of phialides and
scars remaining after conidial detachment (white arrow). n Mature
conidia with slightly roughened scale-like surface. Scale bars g–j, k,
l = 20 lm, m, n = 5 lm
Fungal Diversity (2016) 80:1–270
examined the isotype of P. polysemum, and showed its
similarity to Melomastia and Saccardoella (Barr 1994).
Based on its cylindrical asci, with a non amyloid apical
ring and hyaline, distoseptate ascospores Barr (1994)
transferred Pleurotremataceae to Xylariales. Five genera,
Phomatospora, Melomastia, Pleurotrema, Saccardoella,
Daruvedia, were included in Pleurotremaceae on account
of their non-fissitunicate asci (Barr 1994). Hawksworth
et al. (1995) disagreed with this justification and retained
Pleurotrema in the order Pyrenulales. Saccardoella and
Melomastia were placed in Ascomycota genera incertae
sedis (Kirk et al. 2001; Lumbsch and Huhndorf 2010), but
there is no phylogenetic evidence to support this. Pleurotremataceae was accepted as monotypic via Pleurotrema
and placed under Chaetosphaeriales without giving a reason by Maharachchikumbura et al. (2015). However,
Maharachchikumbura et al. (2016) excluded this family
from class Sordariomycetes and this is an earlier name for
Dyfrolomycetaceae (Dothideomycetes).
Pleurotrema thailandica Dayarathne, Jones E.B.G. & K.D.
Hyde, sp. nov.
Index Fungorum number: IF552244; Facesoffungi
number: FoF02445, Fig. 80
Etymology: Name reflects the country Thailand, from
where the holotype was collected.
Holotype: MFLU 16-1173.
Saprobic on pneumatophores of Avicennia marina in
mangrove vegetation. Sexual morph Ascomata 430–
350 lm diam. 9 242–280 lm high (
x = 428 9 264 lm,
n = 10), solitary, semi-immersed, clypeate, globose to
subglobose, dark brown to black, ostiolate, ostiolar canal
filled with dark cells, apapillate. Peridium 26–35 lm wide,
comprising an outer layer of dark brown cells of textura
angularis, becoming lighter inwardly. Hamathecium comprising numerous, 2.1–3.6 lm wide, septate pseudoparaphyses embedded in a gelatinous matrix. Asci 146–
158 9 7–9 lm (
x = 152 9 8.2 lm, n = 10), 8-spored,
bitunicate, cylindrical, short-pedicellate, with a thickened
apex. Ascospores 24–32 9 6–8 lm (
x = 28 9 6.5 lm,
n = 20), slightly overlapping uni-seriate, hyaline, ellipsoidal, mostly 3–5-septate, slightly constricted at septa,
with several small guttules, surrounded by a gelatinous
sheath, 1.2–2.4 lm thick. Asexual morph Undetermined.
Culture characteristics: Colonies on PDA, reaching
3 cm in 14 days at 25–28 °C, white at first, becoming
yellowish when mature, undulate and reverse yellowishwhite.
Material examined: THAILAND, Phetchaburi Province,
Hat Chao Samran, 47°725060 E, 40°250380 N, 0 m asl., on
pneumatophores of Avicennia marina, 28 July 2015,
Monika Dayarathne, CHAM 006 (MFLU 16-1173, holotype); ex-type living culture, MFLUCC 15-0945, ICMP.
131
Notes: Pleurotrema thailandica is similar to other species of Pleurotrema, but can be differentiated based on
ascospore characteristics, in particular ascospore septation.
Ascospores of P. thailandica are 3–5-septate, while those
of P. tiomanensis, P. mangrovei, P. marinospora and
P. rhizophorae are 20–24, 7–9, 3 and 4–6 septate,
respectively. According to our maximum likelihood analysis based on combined LSU and SSU sequence data,
D. thailandica forms a distinct lineage, sister to P. rhizophorae with 99 % bootstrap support (Fig. 79).
Trypetheliaceae Eschweiler
Trypetheliaceae is mainly a lichen-forming family distributed mostly in tropical to subtropical areas (Harris
1984; Aptroot et al. 2008; Nelsen et al. 2014). The family
was introduced by Eschweiler (1824) and is typified by
T. eluteriae Sprengel. It is characterized by a corticolous
thallus, perithecioid ascomata often organized in pseudostromata, richly branched and anastomosing, net-like
pseudoparaphyses, bitunicate asci, ascospores often with
angular wall thickenings and diamond-shaped lumina, and
a trentepohlioid photobiont in lichenized taxa (Harris 1984;
Aptroot 1998, 2008; Hyde et al. 2013; Nelsen et al.
2009, 2014).
Based on phylogenetic analyses, the family forms a
monophyletic clade in Trypetheliales, sister to Polycoccaceae (Ertz et al. 2015). In this paper we introduce a new
genus, Alloarthopyrenia, in Trypetheliaceae to accommodate a unique, new, non-lichenized lineage resembling
Arthopyrenia (Fig. 81).
Alloarthopyrenia Phukhamsakda, Lücking & K.D. Hyde,
gen. nov.
Index Fungorum number: IF552236, Facesoffungi
number: FoF02379
Etymology: The generic name Alloarthopyrenia refers to
its morphological resemblance with Arthopyrenia.
Saprobic on living tree branches. Thallus absent but area
around the ascomata in part whitish. Sexual morph Ascostromata covered with a blackened pseudoclypeus, semiimmersed, with only ostioles visible, coriaceous, solitary,
scattered or gregarious, depressed globose to obpyriform,
wall rough, black to dark brown, ostiolate. Ostiole centrally
located, filled with periphyses. Peridium dark brown to
light brown, cells of textura angularis and textura epidermoidea, easy to break, at base indistinguishable from host
tissue. Hamathecium composed of branched, anastomosing, trabeculate pseudoparaphyses, embedded in a gelatinous matrix. Asci 8-spored, bitunicate, obviod to
suboblong, short pedicellate, thick-walled, faintly bluish in
IKI ? , apically rounded, with ocular chamber, lightly
IKI ? bluish. Ascospores partially overlapping or bi-seriate, hyaline, oviod or ellipsoid, septate, constricted at the
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Fungal Diversity (2016) 80:1–270
Unknown rock isolate1 TRN508
Unknown rock isolate3 TRN515
Unknown rock isolate2 TRN210
Cladophialophora australiensis CBS 112793
Cladophialophora minourae CBS 556.83
95/1.00 99/1.00
Cladophialophora carrionii CBS 160.54
Phialophora verrucosa AFTOL ID670
90/1.00
90/1.00 Capronia semiimmersa AFTOL ID658
Cladophialphora devriii CBS 147.84
Fonsecaea_monophora CBS 102243
69/-Cladophialophora emmonsii CBS 640.96
Herpotrichiellaceae
Exophiala xenobiotica CBS 115831
Atrokylindriopsis setulosa HMAS245592
86/-Exophiala salmonis AFTOL ID671
Minimelanolocus melanicus MFLUCC15-0415
73/0.99
85/1.00
Minimelanolocus asiaticus MFLUCC15-0237
78/1.00
Minimelanolocus submersus KUMCC15-0206
86/1.00
Minimelanolocus aquaticus MFLUCC15-0414
Minimelanolocus obscurus MFLUCC15-0416
Capronia fungicola CBS 614.96
Capronia mansonii CBS101.67
100/1.00 Capronia munkii AFTOL ID656
94/1.00
Ceramothyrium thailandicum MFLUCC10-0008
Ceramothyrium menglunense MFLUCC14-1120
78/1.00
Chaetothyrium brischoficola MFLUCC 10-0012
100/1.00
Ceramothyrium longivolcaniforme MFLU13-0632b
99/1.00
Ceramothyrium ficus MFLUCC15-0229
75/1.00
Ceramothyrium ficus MFLUCC15-0228
Chaetothyriaceae
Ceramothyrium podocarpi CPC19826
68/-Ceramothyrium carniolicum AFTOL ID1063
100/1.00
Ceramothyrium carniolicum CBS 175.95
100/1.00 Phaeosaccardinula multiseptata IFRDCC2639
100/1.00
Phaeosaccardinula ficus MFLUCC10-0009
Phaeosaccardinula dendrocalami IFRDCC2663
Aphanophora eugeniae CBS 124105
62/-Exophiala eucalyptorum CBS121638
62/-Camptophora hylomeconis CBS 113311
Vonarxia vagans CBS 123533
100/1.00 Vonarxia vagans CPC 15152
Brycekendrickomyces acacia CBS 124104
Phaeococcomyces catenatus CBS 650.76
100/1.00
Phaeococcomyces catenatus CPC 13707
Neophaeococcomyces aloes CPC 21873
100/1.00
100/1.00 Bradymyces alpinus CCFEE 5493
Bradymyces oncorhynchi CCF 4369
--/-0.99
Strelitziana malaysiana CPC 24874
100/1.00
92/1.00
Strelitziana syzygii CPC 26591
96/1.00 100/1.00
Strelitziana australiensis CBS 124778
Trichomeriaceae
Strelitziana eucalypti CBS 128214
Ceramothyrium melastoma CPC 19837
65/1.00
Trichomerium deniqulatum MFLUCC10-0884
100/1.00
Trichomerium foliicola MFLUCC10-0078
71/-100/1.00 Trichomerium gleosporum MFLUCC10-0087
63/0.95
Knufia
epidermidis CBS 120353
100/1.00
Knufia cryptophialidica DAOM 216555
100/1.00
Phaeococcomyces chersonesos Ch49
Knufia perforans CBS 885.95
100/1.00 Metulocladosporiella musae CBS 113863
Metulocladosporiella musae CBS 161.74
100/1.00
Metulocladosporiella musicola CBS 113873
100/1.00 Metulocladosporiella musicola CBS 113865
Cyphellophora reptans CBS 113.85
Cyphellophora sessilis CBS 243.85
73/0.98
Cyphellophora oxyspora CBS 698.73
99/1.00
69/0.98
Cyphellophora europaea CBS 101466
Cyphellophora ambigua CBS 235.93
99/1.00
77/0.99
Cyphellophora eucalypti CBS 124764
Cyphellophora guyanensis MUCL 43737
Cyphellophoraceae
Cyphellophora pluriseptata CBS 286.85
--/0.99
Cyphellophora olivacea CBS 122.74
81/0.98
Cyphellophora suttonii CBS 449.91
95/1.00
Cyphellophora laciniata CBS 190.61
Cyphellophora
vermispora CBS 228.86
97/1.00
Cyphellophora fusarioides MUCL44033
83/-Cyphellophora pauciseptata CBS 284.85
Epibryon sp.2 M175
Epibryon
hepaticola M224
92/1.00
100/1.00 Epibryon hepaticola M10
68/0.94
Epibryon sp.1 M274
100/1.00 95/1.00
Epibryaceae
Epibryon turfosorum M292
Epibryon interlamellare M1
100/1.00
Epibryon interlamellare M32
83/1.00
79/1.00
Epibryon interlamellare M223
Epibryon diaphanum M122
Epibryon plagiochilae M187
99/1.00
98/0.99
Catapyrenium daedaleum AFTOL ID2273
0.1
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Fungal Diversity (2016) 80:1–270
b Fig. 89 Phylogenetic tree generated from maximum parsimony
analysis (RAxML) of a combined dataset of LSU and ITS sequence
data. Bootstrap support values for maximum likelihood (ML, left),
equal to or greater than 60 %, are indicated above or below branches.
Bayesian posterior probabilities (PP, right) equal to or greater than
0.95 are indicated on the branches. New isolates are in blue, and other
ex-type strains are in bold. The tree is rooted with Catapyrenium
daedaleum
Fig. 90 Ceramothyrium menglunense (holotype). a, Herbarium
material. b Superficial ascoma on substrate. c, d Squash of ascomata
wall. e Section through ascoma. f Setae. g Minutely papillate ostiole.
133
septa, wall rough, indentations present when mature, surrounded by a mucilaginous sheath. Asexual morph
Undetermined.
Type species: Alloarthopyrenia italica Phukhamsakda,
Camporesi, Ariyawansa & K.D. Hyde.
Notes: Based on phylogenetic analysis, Alloarthopyrenia is introduced as a monospecific genus to accommodate
h Pseudoparaphyses. i–j Asci. k–n Ascospores. Scale bars c,
e = 100 lm, i, j = 50 lm, d, f, g, h, k–n = 10 lm
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Fungal Diversity (2016) 80:1–270
135
b Fig. 91 Minimelanolocus submersus (holotype). a, b Appearance of
the fungus on wood. c–g Conidiophores and conidia. h–m Conidia.
n Germinating conidium. o Surface view of culture on PDA.
p Reverse view of culture on PDA. Scale bars c = 20 lm, d–
e = 25 lm, f = 40 lm, g = 25 lm, h–i = 10 lm, j–m = 20 lm,
n = 25 lm
a species forming a distinct lineage in Trypetheliaceae
(Fig. 81). The genus Arthopyrenia sensu lato is polyphyletic with tropical, lichenized species clustering in
Trypetheliaceae and a single, non-lichenized taxon,
Arthopyrenia salicis, which falls in Pleosporales: there,
A. salicis clusters with Roussoella and Roussoellopsis, and
Liu et al. (2014) introduced Roussoellaceae to accommodate these taxa. Arthopyreniaceae was kept as a separate
family in Pleosporales, based on its morphology and
anatomy based on the type species, A. cerasi (Schrad.) A.
Massal. which has not yet been sequenced. The new genus,
Alloarthopyrenia is similar to other non-lichenized, temperate species formerly placed in Arthopyrenia sensu lato,
including A. cerasi, in having a carbonaceous pseudoclypeus, branched and anastomosing pseudoparaphyses,
bitunicate asci with shortpedicels, and hyaline ascospores
(Coppins 1988; Hyde et al. 2013). Upreti and Pant (1993)
illustrated species of Arthopyrenia from India. Alloarthopyrenia italica differs from Arthopyrenia cerasi in
having depressed-globose to obpyriform ascostromata,
wider hamathecium filaments (1.4 vs. 0.5–0.7 lm), obovoid to suboblong asci, and 1-septate, ovoid to ellipsoid
ascospores; in contrast, A. cerasi has K ? greenish, larger
and hemisphaerical ascostromata (300–500 lm), less
branched and narrower pseudoparaphyses, cylindrical-clavate asci, and 3-septate, oblong ascospores (Coppins 1988;
Nelsen et al. 2009; Hyde et al. 2013). It is therefore highly
unlikely that the two species are congeneric and that
A. cerasi also falls within Trypetheliaceae.
Although the family Trypetheliaceae mainly comprises
lichenized fungi, Nelsen et al. (2014) found some weakly
or non-lichenized taxa in the family, forming basal lineages
in the tree. This particularly applies to a non-lichenized,
temperate species of Julella, and hence, the placement of a
further, non-lichenized lineage basally in the family is not
surprising. The internal anatomy of Alloarthopyrenia is
otherwise typical of Trypetheliaceae, with the hamathecium forming a network of anastomosing hyphae embedded in a gelatinous matrix, bitunicate asci, transversely
septate, and hyaline ascospores (Harris 1984; Prado et al.
2006; Nelsen et al. 2009). In the present study we introduce
a new genus for this species in Trypetheliaceae, based on
data from analysis of combined LSU and SSU sequence
data.
Alloarthopyrenia italica Phukhamsakda,
Ariyawansa & K.D. Hyde, sp. nov.
Camporesi,
Index Fungorum number: IF552237; Facesoffungi
number: FoF02380, Fig. 82
Etymology: Name reflects the country ‘‘Italy’’, where the
holotype was collected.
Holotype: MFLU 15-0399.
Saprobic on living tree branches of Fraxinus ornus L.
Thallus absent, algae not visible. Sexual morph Ascostromata 110–185 lm high 9 100–270 lm diam. (
x = 139 9
173 lm, n = 5), covered with a blackened pseudoclypeus,
semi-immersed, with only ostioles visible, coriaceous, solitary, scattered or gregarious, depressed globose to obpyriform,
wall rough, black to dark brown, flattened, ostiolate. Ostiole
centrally located, oblong, filled with periphyses. Peridium 12–
23(–21 at apex) lm wide, composed of 5–6 layers of dark
brown to light brown, cells of textura angularis and textura
epidermoidea, easy to break, at base indistinguishable from
host tissue. Hamathecium of numerous, dense, 1–1.8 lm wide
(
x = 1.4 lm, n = 60), narrow, transversely septate, branched, anastomosing, trabeculate pseudoparaphyses, embedded in a gelatinous matrix. Asci 45–99 9 9–20 lm
(
x = 66 9 15 lm, n = 30), 8-spored, bitunicate, fisitunicate,
obviod to suboblong, short pedicellate, thick-walled, lightly
faintly bluish in IKI ? , apically rounded, with ocular
chamber up to 1–2 lm high. Ascospores 15–20 9 4–9 lm
(
x = 18 9 7 lm, n = 50), partially overlapping or bi-seriate,
hyaline, oviod or ellipsoid, slightly narrow at the apex when
young, rounded at the apex when mature, 1-septate, strongly
constricted at the septum, lower end tapering, wall rough, not
uniform, smooth, indentations present when mature, surrounded by 2.5–4 lm wide, mucilaginous sheath. Asexual
morph Undetermined.
Material examined: ITALY, Forlı̀-Cesena Province,
Monte Mirabello—Predappio, on dead branch of Fraxinus
ornus L. (Oleaceae), 15 September 2014, E. Camporesi, IT
122 (MFLU 15-0399, holotype, isotype in HKAS 94615).
Notes: Alloarthopyrenia italica was collected on living
branches of Fraxinus ornus (Oleaceae) in Italy; it is nonlichenized and probably saprobic. Due to the taxon having
similarities with Arthopyrenia, we compared its morphology with Arthopyrenia species reported from Fraxinus.
Five species have been recorded from Fraxinus in Europe,
including A. cerasi (Schrader) A. Massal (type species of
Arthopyrenia, see above), A. cinereopruinosa (Schaerer) A.
Massal., A. carneobrunneola B.J. Coppins, A. fraxini A.
Massal., and A. ranunculospora B.J. Coppins & P. James
(Coppins 1988; Index Fungorum 2016). Arthopyrenia
carneobrunneola and A. ranunculospora differ in being
lichenized (Coppins 1988). Arthopyrenia cinereopruinosa
has been reported from Italy, but it is weakly lichenized
and has larger ascostromata [(200–)300–440 lm] with
3-septate ascospores (Coppins 1988; Aptroot et al. 2008).
Arthopyrenia fraxini differs in having larger ascostromata
(250–500 lm), longer, clavate to cylindrical asci, and
123
136
Fungal Diversity (2016) 80:1–270
Fig. 92 Phylogram generated
from maximum likelihood and
Bayesian analyses based on ITS
sequence data from species of
Trichomeriaceae. The numbers
above the nodes are RAxML
bootstrap values expressed from
1000 repetitions with values
above 50 % shown. The ex-type
strains are in bold and the new
isolates is in blue.The tree is
rooted with Phaeococcomyces
catenatus
Trichomerium sp. MFUCC 13-0780
73
Trichomerium gleosporum MFLUCC 10-0087
Trichomerium foliicola MFLUCC 10-0078
100
73
Trichomerium foliicola MFLUCC 10-0073
Trichomerium sp. MFUCC 13-0789
Trichomerium foliicola MFLUCC 10-0054
90
Trichomerium foliicola MFLUCC 10-0058
99
Trichomerium siamensis MFUCC 12-0105
Trichomerium deniqulatum MFLUCC 10-0884
74
Chaetothyriales sp. CR07-3/2
95
81 Chaetothyriales sp CR08-2/1
100
100
93
Chaetothyriales sp CR08 -2/2
Trichomeriaceae
Chaetothyriales sp. AL4
Chaetothyriales sp. M-Mo2
71 Chaetothyriales sp. CN-Cre-Bo3-2
93 Chaetothyriales sp. CN-Cre-Bo1-4
58
99
94
Chaetothyriales sp. CN-Phe1-1
Chaetothyriales sp. M-Camp4
67
Chaetothyriales sp. M-Cre2
Trichomerium bamboosicola MFUCC 13-0097
100
Chaetothyriales sp CN Cre Bo1 2
Chaetothyriales sp CR07 2 1
99
100
Chaetothyriales sp CR07 2 4
Chaetothyriales sp. CR07 3 1
Phaeococcomyces catenatus CBS 650.76
Herpotrichiellaceae
10
thick-walled, larger ascospores (20–30 9 4–9 lm), with
blunt ends and lacking indentations (Coppins 1988; Upreti
and Pant 1993). Our species is also similar to A. grisea
(Schleich. ex Schaer.) Körb. but the latter has a yellow
thallus, clavate to cylindrical uni-seriate asci, and 3-septate
ascospores (Upreti and Pant 1993). As most Arthopyrenia
species lack sequence data, their placement is uncertain.
Pleosporales genera incetae sedis
The following tree (Fig. 83) represents taxa which do
not cluster in the main families of Dothideomycetes.
Neomassarina Phookamsak, Jayasiri & K.D. Hyde, gen.
nov.
Index Fungorum number: IF552225; Facesoffungi
number: FoF02259
Etymology: The generic epithet ‘‘Neomassarina’’ refers
to the resemblance to Massarina
123
Saprobic on Agave angustifolia. Sexual morph Ascostromata black, solitary, scattered to clustered,
immersed, erumpent through host surface, slightly raised,
globose to subglobose, uni- to bi-loculate, glabrous, ostiole
central, with a minute papilla. Peridium thin-walled, of
unequal thickness, composed of 3–5 layers of dark pseudoparenchymatous cells, arranged in a textura angularis.
Hamathecium composed of dense, 0.5–1 lm wide, cellular
pseudoparaphyses, anastomosing among the asci, embedded in a hyaline gelatinous matrix. Asci 8-spored, bitunicate, fissitunicate, cylindrical to cylindric-clavate, short
pedicellate. Ascospores overlapping uni- to bi-seriate,
hyaline, pale brown at maturity, fusiform, 1-septate, constricted at the septum, smooth-walled with guttules, surrounded by a distinct mucilaginous sheath. Asexual morph
Undetermined.
Type species: Neomassarina thailandica Phookamsak,
Jayasiri & K.D. Hyde
Fungal Diversity (2016) 80:1–270
Fig. 93 Trichomerium bambusae (holotype). a Appearance of
ascomata on leaf. b, c Section through ascoma. d Peridium at the
base of ascoma. e Mycelium. f Ascomatal setae. g Apical ring.
137
h Ascus. i Ascus in Melzer’s reagent. k, l Ascospores. j Ascospore in
Melzer’s reagent. Scale bars b = 100 lm, c, f = 50 lm, e, h,
i = 20 lm d, g, k–l = 10 lm, f–i = 10 lm
123
138
Notes: Neomassarina is introduced as a monotypic
genus to accommodate the massaina-like species, but not
congeneric with Massarina. Neomassarina is similar to
Massarina in having globose to subglobose ascostromata,
with a thin-walled peridium and typically hyaline, fusiform
ascospores, with broadly cellular pseudoparaphyses.
However, the phylogeny indicates that they are distinct.
Based on multi-gene phylogenetic analyses, Neomassarina
form a single clade basal to Floricolaceae (Fig. 83).
Therefore, the new genus is introduced in this study.
Neomassarina thailandica Phookamsak, Jayasiri & K.D.
Hyde, sp. nov.
Index Fungorum number: IF552226; Facesoffungi
number: FoF02260, Fig. 84
Etymology: The specific epithet ‘‘thailandica’’ refers to
the country, from which the holotype was collected.
Holotype: MFLU11-0144.
Saprobic on dead bract-like leaves from flower stalk of
Agave angustifolia Haw. Sexual morph Ascostromata
130–180 lm high, 100–200 lm diam., black, solitary,
scattered to clustered, immersed, erumpent through host
surface, slightly raised, with dark area around ostioles,
globose to subglobose, uni- to bi-loculate, glabrous, ostiole
central, with minute papilla. Peridium 7–16 lm wide, thinwalled, of unequal thickness, slightly thick at the apex,
composed of 3–5 layers of flattened, dark brown to black,
pseudoparenchymatous cells, arranged in a textura angularis. Hamathecium composed of dense, 0.5–1 lm wide,
cellular pseudoparaphyses, distinctly septate, anastomosing
among the asci, embedded in a hyaline gelatinous matrix.
Asci (70–)75–90(–93) 9 7–8(–8.5) lm (
x = 81.8 9
7.6 lm, n = 20), 8-spored, bitunicate, fissitunicate, cylindrical to cylindric-clavate, short pedicellate, apically
rounded, with an obtuse, ocular chamber. Ascospores
(17.5–)18–20 9 3–4(–5) lm (
x = 18.6 9 4 lm, n = 25),
overlapping uni- to bi-seriate, hyaline, pale brown at
maturity, fusiform, 1-septate, constricted at the septum,
Fig. 94 RAxML maximum likelihood phylogenetic tree based on
LSU sequence data for species of Rhytismataceae. Maximum
likelihood bootstrap values greater than 70 % are shown above the
123
Fungal Diversity (2016) 80:1–270
smooth-walled, guttulate, surrounded by a distinct
mucilaginous sheath. Asexual morph Undetemined.
Culture characteristics: Colonies on PDA reaching
20–25 mm diam. after 3 weeks at 25–30 °C; colony from
above, white to cream; from below, white to cream at the
margin, pale yellowish at the centre; medium dense,
irregular, flattened, with undulate edge, surface slightly
rough, cottony; not producing pigmentation in agar.
Material examined: THAILAND, Chiang Mai, Muang
District, Medicinal Plant Garden in Doi Suthep-Pui, on
dead bract-like leaves from flower stalks of Agave angustifolia (Asparagaceae), 23 November 2009, R. Phookamsak, RP0015 (MFLU 11-0144, holotype), ex-type living
cultures, MFLUCC 10-0552, BCC.
Eurotiomycetes
Eurotiales G.W. Martin ex Benny & Kimbr.
Section Cremei
Aspergillus P. Micheli ex Haller
The genus Aspergillus (Trichocomaceae, Eurotiales)
was established by Micheli (1729). The species belonging
to this genus are characterized by their conidiophore patterns and the production of conidial heads (Samson et al.
2014). Species of Aspergillus are ubiquitous and are particularly abundant in the soil and in contaminated foods;
about 339 species (Samson et al. 2014) are divided into
four subgenera with 20 sections (Houbraken et al. 2014).
About 60 species are known to be medically relevant
pathogens, while many species are common contaminants
as mycotoxins producers on various foods, others, such as
A. oryzae, are important in commercial microbial fermentations as Koji mould. Currently, aspergillum is known as a
name of an asexual spore-forming structure which is
common to all species of Aspergillus (Dyer and O’Gorman
2011); about one-third of species are known to have a
sexual morph. Among the sections of Aspergillus, section
Cremei (known as A. cremeus group) was first described by
Raper and Fennell (1965) for five species. Colonies of
nodes. The new isolate is in blue, and other ex-type strains are in bold.
The tree is rooted with Cudonia lutea
Fungal Diversity (2016) 80:1–270
139
Fig. 95 Terriera thailandica (holotype). a, b Hysterothecia on host surface. c Hand section of hysterothecia. d Arrangement of asci. e Peridium.
f–h Asci with ascospores. i, j Ascospores. Scale bars c = 50 lm, d = 50 lm, e, f = 20 lm, g–h = 30 lm, i–j = 20 lm
123
140
Fungal Diversity (2016) 80:1–270
Fig. 96 Phylogenetic tree
inferred from most likelihood
(ML) analysis and Bayesian
inference (BI) using combined
ITS and LSU data (ln
L = 2297.425977). Only the
topology generated from the
ML analysis is shown. Bayesian
Posterior Probability [0.95 are
indicated with thick branch.
New taxa are in blue
species belonging to this section are characterized by
shades of yellowish-brown to brown or grey-green, with biseriate conidial heads and long conidiophores (Samson
et al. 2014). Their conidia are pale grey-green to yellowbrown (Peterson 1995). Among species assigned to this
section, A. wentii is known as a source of enzymes (Raper
and Fennell 1965; Lowe 1992) and A. inflatus is reported to
123
produce sterigmatocystin—a precursor to the even more
potent compounds, the aflatoxins (Rank et al. 2011). During the investigation of the fungi from rhizosphere soil of
pine trees in forests of Gunsan, Korea in August 2015, an
interesting species of Aspergillus was discovered. It proved
to be sufficiently different from previously described species (Stolk and Malla 1971; Samson et al. 2014) to warrant
Fungal Diversity (2016) 80:1–270
141
Fig. 97 Helvella tinta (holotype). a Typical mature specimens. b Receptacle surface of pileus. c, d Asci and paraphyses. e–g Asci with
ascospores. Scale bars a = 1 cm, b = 50 lm, c–g = 20 lm
123
142
description as a new species, forming a distinct lineage.
Previously only one species, A. cibarius from traditional
Meju, has been reported as a new species in Korea (Hong
et al. 2012).
Aspergillus koreanus Hyang B. Lee, T.T. Duong & T.T.T.
Nguyen, sp. nov.
MycoBank number: MB816938; Facesoffungi number:
FoF02476, Fig. 88
Etymology: koreanus, referring to the country from
which the species was first isolated (Korea)
Holotype: EML-GSNP1-1
Colonies on MEA reaching 41–44 mm diam. at 25 °C in
7 days, initially white to white-cotton, turning greyishbrown in the center with age and the abundant sporulation;
reverse white to pale yellowish. Conidiophores hyaline,
smooth, septate, varying greatly in length, 4.5–6 lm diam.
Vesicles subglobose to globose, 7.5–12 lm diam. Conidial
heads bi-seriate, consisting of many metulae bearing several phialides. Phialides ampulliform, 6–9 9 1.5–2.5 lm,
extending from metulae. Conidia grey-green, slightly
roughened, borne in short or long chains, subglobose to
globose, 2.5–3.5 lm diam. Cleistothecia not observed.
Notes: Aspergillus koreanus is distinct from A. inflatus,
growing rapidly when cultivated on MEA producing subglobose to globose vesicles successively and bearing
metulae which develop simultaneously. The metulae consist of several (commonly 3) phialides. The conidial colour
is consistently grey-green with a thin band in the middle to
connect conidia into a chain, while that of A. inflatus is
brownish, with mostly two roughly parallel very thin
bands. A part from phylogenetic characterization; phylogenetic comparisons from single and multi-gene sequence
analyses clearly indicate that A. koreanus strains are related
to A. inflatus and represent a new taxon (Figs. 85, 86, 87).
Material examined: REPUBLIC OF KOREA, Jeonnam
Province, Gunsan City, Sinsido Island (35.82°N,
126.45°E), from a rhizosphere soil of pine tree in forest, 7
August 2015; EML-GSNP1-1 holotype, (ex-type) at Culture Collection of National Institute of Biological Resources (NIBR), Incheon, and preserved as glycerol stock at
-80 °C in the Chonnam National University Fungal Collection (CNUFC); living culture (ex-type) deposited at Jena
Microbial Resource Collection (University of Jena and
Leibniz Institute for Natural Product Research and Infection Biology, Jena, Germany) (JMRC:SF:012334).
The isolate was observed to grow over a wide range of
temperatures, with varying growth rates The average
growth rates on MEA, PDA, YES, OA, CYA, and CREA
were 6, 5.5, 4.5, 4, 3, and 0.2 mm per day, respectively at
25 °C. Optimal growth was observed at 25 °C, slow
growth was observed at below 20 °C and no growth at
35 °C (Fig. 88).
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Chaetothyriales M.E. Barr
Chaetothyriales comprises Chaetothyriaceae, Cyphellophoraceae and Herpotrichiellaceae based on molecular
analysis (Réblová et al. 2013), and includes the sooty
mould families Chaetothyriaceae, Coccodiniaceae and
Trichomeriaceae (Chomnunti et al. 2012b, 2014).
Chaetothyriaceae Hansf. ex M.E. Barr
For the family Chaetothyriaceae, we follow Chomnunti
et al. (2014).
Ceramothyrium menglunense Mapook, J.F. Li & K.D.
Hyde, sp. nov.
Index Fungorum number: IF552275, Facesoffungi
number: FoF02477, Fig. 90
Etymology: Named for the town, Menglun, where the
holotype was collected.
Holotype: MFLU 16-1874
Saprobic on a dead leaves of Syzygium sp. Sexual
morph Ascomata (205–)245–255(–370) lm high 9 140–
165 lm diam. (
x = 270 9 150 lm, n = 5), superficial,
solitary or scattered, coriaceous, globose to subglobose,
dark brown to black with scattered brown setae, Ostiole
central, minutely papillate. Peridium (10–)20–60 lm wide,
2-layered, outer layer comprising dark brown to black cells
of textura globulosa, inner layer comprising light brown to
hyaline cells of textura angularis. Hamathecium comprising 2–3 lm wide, cylindrical to filiform, septate, branched,
pseudoparaphyses.
Asci
50–75 9 24–30(–37)
lm
(
x = 65 9 28 lm, n = 10), 8-spored, bitunicate, clavate to
pyriform, short pedicellate, with an ocular chamber. Ascospores 25–35 9 10–12 lm (
x = 28 9 11 lm, n = 10),
overlapping, hyaline, ellipsoid to obovoid, muriform, with
4–7 transverse septa, and 1–5 vertical septa when mature,
constricted at the septa, smooth-walled, without mucilaginous sheath. Asexual morph Undetermined.
Material examined: CHINA, Yunnan Province,
Xishuangbanna, Menglun, on dead leaves, 13 February
2015, J.F. Li (MFLU 16-1874, holotype), ex-type culture
MFLUCC 14-1120, (isotype in HKAS, under the code of
HKAS 95077).
Notes: Maximum likelihood (RAxML) and Bayesian
analyses of combined LSU and ITS sequences data showed
that our collection is closely related to Ceramothyrium
thailandicum. However, our collection is morphologically
distinct in having larger ascomata with brown setae, shorter
and narrow asci and wider ascospores with vertical septa.
Thus, based on morphology and phylogeny support,
C. menglunense is a new species (Figs. 89, 90).
Herpotrichiellaceae Munk
The family Herpotrichiellaceae was established by
Munk and encompasses loculoascomycetes with small,
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143
Fig. 98 Helvella floriforma (holotype). a Typical mature specimens. b Receptacle surface of pileus. c Stipitipellis. d Asci and paraphyses. e, f
Asci. g Ascospores. Scalebars a = 1 cm, b–c = 50 lm, d–g = 20 lm
superficial, inconspicuous, setose ascomata, bitunicate asci
with a thickened endotunica and greenish-grey to brown,
septate ascospores (Munk 1953). Barr (1976) included the
family Herpotrichiellaceae in the order Chaetothyriales.
Phylogenetic analysis also supported the family Herpotrichiellaceae as belonging to the order Chaetothyriales
(Liu et al. 2015b).
Minimelanolocus R.F. Castañeda & Heredia
Castañeda-Ruiz et al. (2001) introduced the genus
Minimelanolocus with M. navicularis (R.F. Castañeda)
R.F. Castaneda as the type species. They transferred ten
species from the genera Pseudospiropes, Helminthosporium, and Belemnospora into Minimelanolocus, and
described one new species, M. curvisporus. Several species
of Minimelanolocus have since been described from a wide
range of hosts worldwide based on morphology (Zhang
et al. 2010; Ma et al. 2011a, b; Hernández-Restrepo et al.
2013; Xia et al. 2014; Liu et al. 2015b). Liu et al. (2015b)
listed 28 species of Minimelanolocus and provided the
morphological characters of these taxa. A phylogenetic tree
of Chaetothyriales, based on combined LSU and ITS
sequence data, is provided in this paper (Fig. 89).
Minimelanolocus submersus Z.L. Luo, H.Y. Su & K.D.
Hyde, sp. nov.
Index Fungorum number: IF552154, Facesoffungi
number: FoF02219, Fig. 91
123
144
Fungal Diversity (2016) 80:1–270
Fig. 99 Helvella oblongispora. a–c Typical mature specimens (a HKAS 87693, b HKAS 87726, c HKAS 87694). d Receptacle surface of
pileus. e, f Stipitipellis. g, h Asci and paraphyses. i–k Asci l. Ascospores. Scale bars a–c = 1 cm, d–f = 70 lm, g–l = 20 lm
Etymology: With reference to the submerged habitat.
Holotype: HKAS 92593.
Saprobic on submerged decaying wood. Sexual morph
Undetermined. Asexual morph Colonies on the substrate
superficial, effuse, hairy, scattered, brown. Mycelium
mostly immersed, composed of septate, dark brown,
smooth hyphae. Conidiophores mononematous, macronematous, unbranched, erect, straight or slightly flexuous,
123
smooth, cylindrical, septate, dark brown, gradually paler
towards the apex, 110–180 lm (
x = 145.5 lm,
SD = 35.5 lm, n = 10) long, 5–6 lm (
x = 5.5 lm,
SD = 0.5 lm, n = 10) wide. Conidiogenous cells
holoblastic, integrated, sympodially proliferating, terminal,
pale brown or subhyaline. Conidia acrogenous, clavate to
fusiform, solitary, immature conidia 1-septate, mostly 2–5septate at maturity, dry, subhyaline to pale brown, 19–
Fungal Diversity (2016) 80:1–270
33 lm (
x = 26 lm, SD = 7 lm, n = 20) long, 3.5–
4.5 lm (
x = 4 lm, SD = 0.5 lm, n = 20) wide. Conidial
secession schizolytic.
Culture characteristics: Colonies on PDA 25 mm diam.
after 3 weeks at room temperature, pale brown at the
margins, dark brown at the center; reverse steel grey,
medium dense, circular, fairly tight.
Material examined: CHINA, Yunnan Province, Dali,
Erhai Lake, saprobic on submerged decaying wood, June
2015, X.Y. Liu, S-323 (HKAS 92593, holotype), ex-type
culture KUMCC 15-0206, MFLUCC.
Notes: In the phylogenetic analysis, Minimelanolocus
submersus clustered with species of Minimelanolocus; and
formed a sister group to M. obscurus (Matsush.) R.F.
Castañeda & Heredia. M. submersus closely resembles
M. aquaticus H.Y. Su etal. and M. asiaticus H.Y. Su et al.
but differs from M. aquaticus in having 2–5-septate, narrower conidia (3.5–4.5 lm vs. 4–8 lm), and differs from
M. asiaticus in having longer and wider conidiophores
(110–181 9 5–6 lm vs. 112–162 9 3.5–4.5 lm) (Liu
et al. 2015b).
Trichomeriaceae Chomnunti & K.D. Hyde
The family was introduced and placed in Chaetothyriales by Chomnunti et al. (2012a), with the generic type
Trichomerium. Trichomeriaceae comprises several species
of sooty moulds. The family presently comprises three
genera, Bradymyces, Knufia, and Trichomerium (Hubka
et al. 2014). Trichomeriaceae includes strains that were
isolated from the surface of rocks as well as fungi associated with ants (Hubka et al. 2014). In this study, we
introduce a new species isolated from a plant, based on
morphology and phylogenetic analysis. We provide a tree
for Trichomeriaceae below (Fig. 92).
Trichomerium Lév.
Trichomerium species differ from Capnodiaceae species
in having loose mycelium beneath the ascomata, the
appearance of ascomatal setae, and fusoid ascospores, with
three transverse septa or sometimes with longitudinal septa.
Phylogenetic analyses from previous studies indicate that
Trichomeriaceae species belong to Eurotiomycetes, and
Capnodiaceae belongs to Dothideomycetes.
Trichomerium bambusae Hongsanan & K.D. Hyde, sp.
nov.
Index Fungorum number: IF552365; Facesoffungi
number: FoF02446, Fig. 93
Etymology: Named after the host bamboo.
Holotype: MFLU 16-2286
Epiphytes, saprobic on the surface of culm of bamboo.
Superficial hyphae 5–6 lm wide, branched, septate, constricted at the septa, brown. Sexual morph Ascomata 140–
145
180 high 9 105–130 wide lm (
x = 154 9 118 lm,
n = 7), superficial, solitary to gregarious, mainly globose
to subglobose, with long ostiole at the center, held to the
leaf surface by basal mycelium, brown to greyish, with
apical setae. Setae 50–80 9 5–8 lm (
x = 65 9 7.5 lm,
n = 10), surrounding the ascomata, usually at the upper
part of ascomata, dark brown to black, slightly narrower
and paler at the apex, straight, dark brown to black.
Peridium 13–17 lm (
x = 14 lm, n = 10), comprising two
layers, outer layer composed of pigmented, thick-walled
cells of textura angularis, inner layer composed of pale,
flattened cells of textura angularis. Hamathecium not
observed. Asci 53–62 9 19–24 lm (
x = 57 9 22 lm,
n = 10), 8-spored, bitunicate, broadly cylindrical or
oblong, short pedicellate, ocular chamber when immature.
Ascospores 15–18 9 5–7 lm (
x = 17 9 6 lm, n = 15),
2–3-seriate, oblong, hyaline, 3-septate, usually not constricted at the septa, but sometimes slightly constricted at
the lowest septum, with thin mucilaginous sheath at the
middle, smooth-walled, end cells narrow and smaller than
central cell. Asexual morph Undetermined.
Material examined: THAILAND, Chiang Rai, Bandoo,
on dead of culm of bamboo (Poaceae), December 2012,
SC Karunarathna HSA59 (MFLU 16-2286, holotype; isotype in KIB); ex-type living culture, MFLUCC 13-0097.
Notes: Trichomerium bambusae differs from other species within Trichomerium in having a long central ostiole,
and 2–3-septate ascospores, with very thin mucilaginous
sheath. The new species is most similar to T. gloeosporum
Chomnunti & K.D. Hyde based on the shape of ascomata,
with raised ostioles, however, T. bambusae has a longer
ostioles than T. gloeosporum. The ascospores of
T. gloeosporum have a thick mucilaginous sheath, while its
mucilaginous sheath is poorly developed in T. bambusae.
In the phylogenetic analysis T. bambusae clusters with
strains of Chaetothyriales isolated from carton fungi (fungi
associated on ant carton nests) and other Trichomerium
species. Ant fungi are a polyphyletic group which also
present in Capnodiales (Voglmayr et al. 2010).
Leotiomycetes
We follow Wang et al. (2006) for the classification of
this order.
Rhytismatales M.E. Barr ex Minter
Rhytismatales is an order of endophytic, parasitic or
saprotrophic fungi in the class Leotiomycetes (Ascomycota), the inoperculate discomycetes. Four families are
currently recognized in the order.
Rhytismataceae Chevall.
Rhytismataceae is largest family (Johnston 2001) and
includes 44 genera (Lumbsch and Huhndorf 2010),
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146
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Peziza vesiculosa DQ491509
Peziza vesiculosa JF908568
Peziza vesiculosa AF491625
Peziza vesiculosa AF491623
Peziza bovina JF908555
1/100
1/100 Peziza fimeti JF908537
Peziza sp. UDB015563
0.89/63
Peziza domiciliana JF908561
Peziza granularis JF908558
0.98/75
1/100 Peziza ammophila JX845423
Peziza ammophila JX845421
1/100 Peziza pseudoammophila JF908559
Peziza ammophila AF491622
Peziza ammophila AF491621
0.73/Peziza ammophila UDB019674
1/97
1/100 Peziza ammophila UDB019673
1/91 Peziza varia AF491553
Peziza varia AF491551
0.87/54
Peziza varia AF491550
Peziza varia AY789392
1/100 Peziza varia AF491549
1/82
Peziza varia AF491548
1/100 Peziza
varia AF491558
Peziza varia AF491557
1/
100 Peziza varia AF491556
1/81
Peziza fimeti JQ654490
Peziza varia AF491555
1/100
Peziza varia JF908557
1/99 Peziza varia AF491570
Peziza varia AF491569
Peziza varia AF491568
Peziza varia UDB018603
0.99/93
0.99/88
1/100
0.95/-
1/98
1/100
0.96/75
1/71
Peziza sp. JX415342
Peziza echinospora JF908533
Peziza echinospora AF491575
Peziza echinospora AF491573
Peziza sp. AF491572
1/100 Peziza sp. AF491571
1/100
0.99/77
Peziza varia JF908560
Peziza sp. AF491587
Peziza sp. AF491586
0.88/53
Peziza arvernensis JF908569
1/96
Peziza arvernensis AF491577
Peziza arvernensis AF491580
Peziza fimeti JQ654489
1/100 Peziza nivalis JX845425
Peziza nivalis AF491620
Peziza
nivalis
AF491619
1/100
Peziza fruticosa AMB 17136 Calambrone
1/69
Peziza fruticosa AMB 17135 Rosolina
1/55 0.86/Peziza ninguis JF908536
0.99
Peziza domiciliana AF491614
/100
Peziza perdicina JF908529
0.87/54
Peziza ampliata AF491590
Peziza ampliata AF491592
1/Peziza fimeti JQ654491
1/100 Peziza fimeti JQ654487
0.75/73
Peziza fimeti AF491605
Peziza fimeti JQ654495
Peziza fimeti JQ654493
Peziza fimeti AF491600
Peziza udicola JF908549
Peziza lohjaoensis AF491576
1/99
Peziza proteana JF908566
1/91
Peziza proteana DQ491497
Peziza petersii JF908550
1/100
0.76/54
Peziza petersii JF908527
Peziza petersii AF133179
Peziza fruticosa
1/100
Peziza michelii JN836748
Peziza michelii DQ200839
Peziza subcitrina AF491627
0.1 expected changes per site
Fig. 100 Phylogeny of Peziza based on a Bayesian and maximum
likelihood analysis of ITS dataset. Bayesian posterior probability
(BPP) values (in bold) C0.7 and maximum likelihood bootstrap
(MLB) values C50 % are shown on the branches. Thickened
branches indicate BPP C 0.95 and MLB support C70 %. Peziza
subcitrina was chosen as outgroup taxon
Lophodermium is the largest genus with more than 100
species currently accepted (Lantz et al. 2011).
Etymology: thailandica is based on country where species was found.
Holotype: MFLU 16-0945.
Saprobic on decaying woody branches in terrestrial
habitats, associated with bleached, pale brown areas. Sexual morph Ascomata hysterothecial, 1100–1900(–900)
long 9 222–250 high, 210–268 lm diam., in surface view,
matt, elliptical, ends rounded to subacute, margins diffuse,
the central part of the ascomata strongly raising the surface
of the substrate at maturity, opening by a longitudinal split
that extends almost the whole length of the ascoma.
Peridium 17–44 lm, carbonaceous, brittle, of heavily
pigmented, basal cells of textura angularis and globulosa,
covering stroma consisting of an outer layer of host cuticle.
Hamathecium comprising 1–2 lm wide, hyaline, aseptate,
excipulum moderately developed, borne in a gel matrix,
closely adhering to the covering stroma and the extension,
arising from the marginal paraphyses. Asci 80–105 9 3.4–
Terriera B. Erikss.
The genus Terriera is a member of Rhytismataceae
(Kirk et al. 2008). The type species is T. cladophila (Lév.
in Moug. & Nestl.) B. Erikss (syn. Lophodermium cladophilum (Lév. In Moug. & Nestl.) Rehm) (Eriksson 1970).
Johnston (1988, 1989) enumerated some important features
for delimitation of this genus, such as oblong to sublinear
ascomata, the lack of lip cells, covering stroma forming a
platform in vertical section, and the triangular space in
section between the covering stroma and basal stroma filled
with vertically oriented cells.
Terriera thailandica Jayasiri & K.D. Hyde, sp. nov.
Index Fungorum number: IF552171; Facesoffungi
number: FoF02222, Fig. 95
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147
Peziza exogelatinosa AY500501
0.85/ -
1.0/100
Peziza quelepidotia AF107809
Peziza natrophila AY500486
0.93/52
Peziza sp. PM 120 97 AY500488
0.95/54
Peziza subcitrina AY500520
Peziza howsei AY500493
0.98/ -
1.0/100
0.82/ -
Peziza emileia KJ728718
Peziza emileia KJ728719
0.99/54
Peziza lobulata AY500495
Peziza ampelina AY500492
1.0/100
Peziza vesiculosa DQ470898
Peziza vesiculosa AY500489
Peziza varia AY500498
1.0/100
Peziza varia AY500499
0.98/57
Peziza varia AY500519
0.96/53 1.0/96
Peziza varia AY500500
0.99/70
0.82/58
Peziza echinospora AY500496
1.0/85
Peziza arvernensis LN714689
Peziza arvernensis AY500497
1.0/100
Peziza ampliata AY500510
Peziza fruticosa
0.99/100 Peziza polaripapulata AY500514
Peziza polaripapulata AY500515
0.05 expected changes per site
Peziza obtuspiculata AY500490
Fig. 101 Phylogeny of Peziza based on a Bayesian and maximum
likelihood analysis of RPB2 dataset. Bayesian posterior probability
(BPP) values (in bold) C0.7 and maximum likelihood bootstrap
(MLB) values C50 % are shown on the branches. Thickened
branches indicate BPP C 0.95 and MLB support C70 %. Newly
sequence collections are in bold. Peziza polaripapulata and P.
obtuspiculata are chosen as the outgroup taxa
6.6 lm (
x = 90 9 5 lm, n = 15), 8-spored, crowded to
somewhat parallel, cylindrical, but cylindrical-clavate
when immature, apex obtuse to truncate, thin-walled,
without a circumapical thickening, long stalked J-, discharging spores through a small apical pore. Ascospores
38–60 9 1–1.5 lm (
x = 45 9 1.2 lm, n = 25), arranged
in a fascicle, hyaline, filiform, tapering slightly towards
both ends, aseptate, pluriguttulate, cell wall not clear,
without a gelatinous sheath or appendages. Asexual
morph Undetermined.
Culture characters: Colonies on MEA 25 mm diam. after
7 days at 25 °C, raised, with lobate margin; colony two
layered, outer layer off white and bound to media, inner layer
white and finely floccose to woolly aerial mycelia. Reverse
white with black colour patches, prominent in middle.
Material examined: THAILAND. Chiang Rai, Doi Pui:
dead branch of undetermined tree, 15 June 2014, Subashini
C. Jayasiri. (MFLU 16-0945, holotype), (isotype in KUN),
ex-type culture, MFLUCC 14-0818, KUNCC.
Notes: Terriera thailandica is introduced here based on
both morphology and phylogeny. Terriera was segregated
from Lophodermium based mainly on its swollen paraphyses at the apex to form an epithecium and the lack of lip
cells (Eriksson 1970). Terriera thailandica shares these
characters with other species of the genus. Terriera thailandica clusters with T. camelliicola in a clade with high
ML support and distinct from T. thailandica and T. minor
(Fig. 94). Terriera thailandica shares similar morphological characters with T. camelliicola, but the latter species
has hysterothecia which are more or less curved, lacks an
123
148
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b Fig. 102 Peziza fruticosa (holotype). a, b Ascospores in Lactic blue.
c–e Ascospores in Melzer’s reagent. f Hymenium in water. g Ascus
and paraphyses in Melzer’s reagent. h Asci and paraphyses in water.
i Apothecia in situ. Scale bars a = 8 lm, b = 4 lm, c = 3 lm, d,
e = 2.5 lm, f = 30 lm, g, h = 15 lm, i = 2 mm (Photo G.
Medardi)
epithecium, and has observable conidiomata near the hysterothecia (Zhang et al. 2015).
Pezizomycetes
Pezizales J. Schrot.
The order Pezizales has long attracted the attention of
mycologists from different parts of the world. Currently,
there are 16 families, namely, Ascobolaceae, Ascodesmidaceae, Caloscyphaceae, Carbomycetaceae, Chorioactidaceae, Discinaceae, Glaziellaceae, Helvellaceae,
Karstenellaceae, Morchellaceae, Pezizaceae, Pyronemataceae, Rhizinaceae, Sarcoscyphaceae, Sarcosomataceae
and Tuberaceae in the order (Hibbett et al. 2007; Kirk et al.
2008). This order is distinguished by asci generally open by
rupturing to form a terminal or eccentric lid or operculum
(Hansen and Pfister 2006).
Helvellaceae Fr.
This family was introduced by Fries (1822) and is
widely distributed in temperate to arctic-alpine areas (e.g.
Abbott and Currah 1997; Dissing 1966; Ying and Zang
149
1994; Zhuang 2004), with a few taxa known from the
tropics (Dissing 1979). To date, the family includes four
genera, viz. Balsamia., Barssia, Helvella and Wynnella,
that comprise approximately 100 species (Kirk et al. 2008;
Nguyen et al. 2013; Zhao et al. 2015b, 2016a, b; Wang
et al. 2016).
Helvella L.
Helvella is the type genus of Helvellaceae, is characterized by its epigeous, stipitate, cupulate, saddle-shaped to
irregularly-lobed ascomata, with whitish, cream, greyish,
brown to black hymenium, glabrous, pubescent to villose
receptacle surface, and terete or externally sulcate, solid,
hollow or lacunose stipe (Dissing 1966; Abbott and Currah
1997).
This genus has received much attention in China, Europe and North America, in the last decade (Zhuang 2004;
Zhuang and Yang 2008; Ariyawansa et al. 2015a; Hwang
et al. 2015; Zhao et al. 2015b, 2016a, b; Wang et al. 2016).
To date, 20 species have been described from China (Wang
et al. 2016). In this study we introduced two new species
based on molecular support (Fig. 96) and a new record for
Helvella oblongispora.
Helvella tinta Q. Zhao, B. Feng & K.D. Hyde, sp. nov.
Index Fungorum number: IF552355, Facesoffungi
number: FoF02447, Fig. 97
Etymology: Named because of its tints hymenium and
receptacle surface.
Fig. 103 Diagrammatic
representation of the
microscopic characters of
Peziza fruticosa (holotype).
a Vertical section of one
apothecium. a1 Hymenium. a2
Subhymenium. a3 Upper
medullar excipulum. a4 Median
medullar excipulum. a5 Lower
medullar excipulum. a6 Ectal
excipulum. b Ascospores.
c Asci and ascospores.
d Paraphyses. Scale bars
a = 100 lm, b = 10 lm, c,
d = 25 lm (drawing by Del.
G. Medardi)
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Fig. 104 Coronophora myricoides (holotype). a Herbarium material. b, c Appearance of ascomata on wood. d, e Vertical sections of ascomata.
f Peridium. g–i Asci. j–l Ascospores. Scale bars c = 200 lm, d = 50 lm, e, g, h = 20 lm, f, i–l = 10 lm
Holotype: HKAS 82560.
Symbiotic in coniferous forests of Picea euphratica
Oliv. Sexual morph Pileus cupulate to irregularly
cupulate, 2–3 cm high, 1.5–3.5 cm broad, margin slightly
flattened; hymenium glabrous, even, brown to dark brown,
usually mottled with patches of paler pigmentation, greyish to
dark brown when fresh, blackish-brown when dried; receptacle surface pubescent, concolourous with the hymenium,
ribs absent or extending onto basal quarter only. Stipe
2.5–4 cm long, 0.7–1.5 cm broad, glabrous, flaring and
123
merging with apothecium, white to smoky grey, becoming
cream and tough when dried, with 6–8 ribs, with roundededge, few anastomosis between ribs, basal mycelium white.
Medullary excipulum 370–500 lm broad, of textura intricata, hyaline, composed of 4–6 lm broad hyphae, J-. Ectal
excipulum 70–180 lm broad, of textura angularis, outermost
cells catenuliform in long fascicled tufts, 14–38 9 8–18 lm,
brown, evenly blue in cotton blue, J?. Asci
270–350 9 14–20 lm, pleurorhynchous base, 8-spored, uniseriate, subcylindrical to clavate. Paraphyses filiform,
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151
Fig. 105 One of 255 most
parsimonious trees obtained
from analyses of combined ITS,
EF-1a, b-tubulin and CAL
sequence data for all ex-types
from species in Diaporthe.
Isolate numbers of new species
and new host records are in
blue. Maximum parsimony
bootstrap values ([70 %) and
Bayesian inference values
([0.8) are given on the nodes.
The tree is rooted to Diaporthe
corylina
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152
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Fig. 105 continued
3–5 lm broad, slightly exceeding the asci, light brown, with a
slightly yellow refractive content in Melzer’s reagent, blue in
cotton blue, apex slightly enlarged, 4–5.5 lm broad.
123
Ascospores [30/2/2, in H2O] (15–)16–19(–20) 9 10–13(–14)
lm [Q = 1.36–1.77, Q = 1.59 ± 0.09)], ellipsoid, smoothwalled. All tissues J-. Asexual morph Undetermined.
Fungal Diversity (2016) 80:1–270
153
Fig. 106 Diaporthe aseana (MFLU 13-0256, holotype). a Specimen on dead leaf. b Conidiomata on the host surface. c Longitudinal section of
a conidioma. d, e Longitudinal section of a conidioma wall. f, g Conidiogenous cells with developing conidia. h–k Conidia. l Germinating
conidium. m, n Colonies on PDA, m from above n from below. Scale bars c = 100 lm, d = 50 lm, e–l = 10 lm
123
154
Habitat and distribution: Scattered or gregarious on the
moss, under Picea euphratica Oliv., forests. Known only
from southwestern China.
Material examined: CHINA, Sichuan Province, Hongyuan County, alt. 3300 m, 12 August 2013, Bang Feng
1454 (HKAS 82560, holotype).
Notes: Helvella tinta is well characterized by its cupulate to irregularly cupulate pileus with a consistently
mottled, greyish to dark brown hymenium, and a lacunose
stipe possessing ribs extending onto 1/4 of the receptacle
surface. Paraphyses light brown, 3–5 lm. The brown pigments of the receptacle surface are visible in cotton blue or
Melzer’s reagent.
Morphologically, H. tinta is similar to H. maculata N.S.
Weber, a species originally described from North America,
but the latter has a saddle-shaped pileus with a drab, buffy
brown to snuff brown hymenium, margin curved to
receptacle surface, a white to cream stipe and large
ascospores (Weber 1975). Helevlla floriforma also shares
some features with H. tinta, but it has a peculiar apothecial
shape and colour.
Phylogenetically, H. tinta clusters with H. griseoalba
N.S. Weber (66 % bootstrap support). However,
H. griseoalba differs from H. tinta in having grey to cinnamon hymenium and receptacle surface and hyaline
paraphyses (Weber 1972; Landeros et al. 2012).
Helvella floriforma Q. Zhao & K.D. Hyde, sp. nov.
Index Fungorum number: IF552356, Facesoffungi
number: FoF02448, Fig. 98
Etymology: the epithet refers to the flower-shaped
apothecia.
Holotype: HKAS 90224.
Symbiotic in the coniferous forests of Platycladus orientalis (L.) Franco. Sexual morph Pileus slightly
infundibuliform to central strongly depressed but not hollow when mature, 2–3 cm high, 2–6 cm broad, margin
rolled to receptacle surface, wavy; hymenium glabrous,
drab, buffy brown to snuff brown when fresh, yellowishbrown when dried; receptacle surface pubescent, cream,
pale, apricot greyish when fresh, becoming yellowish when
dried, branching and anastomosing ribs extending to marginal region. Stipe 2–4 cm long, 1.5–3 cm broad, cream,
becoming yellowish when dried, glabrous, sulcate, branched and anastomosed ribs extending to the receptacle
surface margin. Medullary excipulum 290–450 lm broad,
of textura intricata, hyaline, composed of 3–5 lm broad
hyphae, with a red refractive content in Melzer’s reagent,
blue in cotton blue, J-. Ectal excipulum 100–130 lm
broad, of textura angularis, outermost cells catenuliform in
long fascicled tufts, hyaline, evenly blue in cotton blue,
cylindrical end cells 13–25 9 7–13 lm, J?. Stipitipellis
50–70 lm, hyaline, composed of textura angularis,
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Fig. 107 Diaporthe eres (MFLUCC 12-0351). a Specimen on dead c
cone of Picea excelsa. b Conidiomata on the host surface. c Longitudinal section of a conidiomata. d–g Longitudinal section of
peridium. h–m Conidiogenous cells with developing conidia. n–
s Conidiogenous cells with developing conidia stained with lactophenol cotton blue. t Conidia. u Conidia stained with lactophenol cotton
blue. v Germinating conidium. w–x Colonies on PDA, w from top,
x from reverse. Scale bars c = 100 lm; d–g = 50 lm; h–u = 5 lm;
v = 10 lm
terminal cells 13–27 9 7–10 lm, clavate, with a yellow
refractive content in Melzer’s reagent, blue in cotton blue,
J-. Asci 260–310 9 16–22.5 lm, pleurorhynchous base,
8-spored, uni-seriate, subcylindrical to clavate. Paraphyses
filiform, 3–4 lm broad, slightly exceeding the asci, with a
slightly yellow refractive content in Melzer’s reagent, blue
in cotton blue, apex 4–5 lm broad, J-. Ascospores [40/2/1,
in H2O] (14)14.5–20 9 10.5–16 lm [Q = 1.16–1.59,
Q = 1.34 ± 0.11)], subglobose to subellipsoid, smoothwalled. Asexual morph Undetermined.
Habitat and known distribution: Scattered or gregarious
on the ground, under Platycladus orientalis. Currently
known only in southwestern China.
Material examined: CHINA, Yunnan Province,
Gucheng County, alt. 2500 m, 19 August 2013, Qi Zhao
2000 (HKAS 90224, holotype).
Notes: Helvella floriforma is characterized by its slightly
infundibuliform to strongly depressed pileus with a wavy,
curved to receptacle surface margin, a glabrous, drab, buffy
brown to snuff brown hymenium, and a pubescent, cream,
pale, apricot greyish, ribs branching and anastomosing
extending to marginal region receptacle surface. Stipe
cream, glabrous, sulcate, rounded-edge, branched and
anastomosed ribs extending to the receptacle surface margin. Stipe inner and ectal excipulum J?.
In our phylogenetic analysis, H. floriforma is sister to
H. robusta S.P. Abbott with relatively high statistical
support (91 %) (Fig. 96). However, the latter species is
irregularly cupulate with a large central depression to
irregularly bi-lobed pileus, free margin, a subpubescent to
pubescent, ribbed receptacle surface and a subpubescent to
pubescent, lacunose stipe with sharp ribs; medullary
excipulum and stipe inner and stipitipellis J?; ectal
excipulum J- (Landeros et al. 2012).
Helvella tinta and H. maculata also have mottled
hymenium surface and lacunose stipe. However, H. tinta
has a cupulate to irregularly cupulate pileus, with a greyish
to dark brown hymenium, a mottled receptacle surface only
near the stipe possessing ribs, all tissues J-. Helvella
maculata has a saddle-shaped pileus, margin rolled or
straight toward the hymenium and a white to cream stipe,
all tissues J- (Weber 1975; Abbott and Currah 1997;
Landeros et al. 2012).
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123
156
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b Fig. 108 Diaporthe foeniculina (MFLUCC 12-0668). a Specimen
on dead branch of Laburnum sp. b Conidiomata on the host surface. c,
d Longitudinal section of a conidiomata. e, f Longitudinal section of
peridium. g–l developing conidia. m–p Developing conidia stained
with lactophenol cotton blue. q Alpha conidia. r Alpha conidia
stained with lactophenol cotton blue. s Beta conidia. t Beta conidia
stained with lactophenol cotton blue. u Germinating conidium. v,
w Colonies on PDA, v from top, w from reverse. Scale bars c,
d = 100 lm; e, f = 50 lm; g–t = 5 lm; u = 10 lm
Helvella oblongispora Harmaja, Karstenia 18(2): 57
(1978)
Facesoffungi number: FoF02449, Fig. 99
Symbiotic in the coniferous forests of Picea euphratica
Oliv. Sexual morph Pileus cupulate to irregularly cupulate, 0.5–2.5 cm high, 0.5–2.5 cm broad, margin slightly
splitting; hymenium glabrous, even to slightly undulate at
center, greyish-brown to brown when fresh, blackishbrown when dried; receptacle surface finely pubescent,
greyish to brown at the margin, cream to white below,
white near stipe, ribs absent or extending onto basal quarter
only. Stipe 0.5–1.5 cm long, 0.3–0.5 cm broad, flaring and
merging with apothecium, white, becoming cream and
tough when dried, glabrous to finely pubescent, ribs
prominent, rounded, few anastomosis between ribs, basal
mycelium white. Medullary excipulum 220–315 lm broad,
of textura intricata, hyaline, composed of 3–4 lm broad
hyphae. Ectal excipulum 60–115 lm broad, of textura
angularis, hyaline, evenly blue in cotton blue, outermost
cells 17–45 9 7–27 lm, clavate to subglobose. Stipitipellis
100–160 lm, hyaline, composed of textura angularis, terminal
cells
14–59 9 8–21 lm,
clavate.
Asci
260–330 9 15–21 lm, aporhynchous base, 8-spored, uniseriate, subcylindrical to clavate. Paraphyses filiform,
slightly exceeding the asci, with a yellow refractive content
in Melzer’s reagent, 3–5 lm broad. Ascospores [80/4/4, in
H2O] (17–)18–20(–21) 9 11–13 lm [Q = 1.36–1.77,
Q = 1.59 ± 0.09)], subellipsoid, smooth-walled under the
light microscope. All tissues J-. Asexual morph
Undetermined.
Known distribution: Scattered or gregarious on moss
under Picea euphratica, P. retroflexa Mast. and P. purpurea Mast. Picea spp. forests. Known in Europe (Harmaja
1978) and Asia (Liu et al. 1985).
Material examined: CHINA, Sichuan Province, Jiuzhaigou County, on forest ground with Picea euphratica
Oliv., alt. 3200 m, 19 Jun 2014, Qi Zhao 2046 (HKAS
87726), Jiuzhaigou County, alt. 3300 m, 22 June 2014,
Qi Zhao 2064 (HKAS 87744), same location, 23 June
2014, Qi Zhao T24352, T24387, T24470, T24477,
respectively (HKAS 87693; HKAS 87694; HKAS 87698;
HKAS 87700), Xinjiang Autonomous Region,Zhaosu
County, alt. 2330 m, 9 July 2015, Qi Zhao 2380 (HKAS
90254).
157
Notes: Helvella oblongispora is characterized by deeply
cupulate pileus, slightly serrate margin, brown, even to
slightly undulate hymenium, white to pale brownish
receptacle surface, and prominent, rounded ribs, white stipe
without few crossed veins or pockets; aporhynchous asci,
paraphyses apex 3–5 lm broad and subellipsoid,
18–20 9 11–13 lm ascospores. The species was re-examined by Landeros et al. (2012).
Helvella oblongispora is very similar to H. leucomelaena in gross morphology and anatomical structure.
However, H. leucomelaena has dark grey to blackish
apothecia, deeply cupuliform, generally subsessile or with
a very short stipe and less or more visible ribs and
20–23 9 11–13 lm ascospores (Dissing 1966; Abbott and
Currah 1997).
Pezizaceae Dumort.
Pezizaceae, known as the cup-fungi, is recognized by
the fleshy, soft, brittle, cupulate ascomata, with amyloid
asci. This family formed a strongly supported monophyletic with Ascobolaceae as its sister group (Hansen and
Pfister 2006). To date, GBIF lists 54 genera, approximately
1214 species (http://www.gbif.org/species/8399, 27 July
2016) and, of these, 31 genera, 230 species were accepted
(Kirk et al. 2008).
Peziza Dill. ex Fr.
Peziza is the type genus of the family Pezizaceae,
estimated to comprise 104 species (Kirk et al. 2008).
Molecular phylogenetic studies have shown that the
genus Peziza is not monophyletic, composed of at least
14 distinct lineages (Hansen and Pfister 2006) (Figs. 100,
101).
Peziza fruticosa Lantieri, Medardi & Vizzini, sp. nov.
Index Fungorum number: IF552367; Facesoffungi
number: FoF02450, Figs. 102 and 103
Etymology: from Latin ‘‘fruticosa’’, clustered, for its
way of fruiting.
Holotype: AMB 17135.
Saprobic on the sand of the high dunes. Sexual morph
Apothecium cup shaped, sessile, irregularly rounded or
compressed-lobed because of mutual contact; on average
15 mm diam. Hymenium smooth, often wavy, dull to dark
brown, with reddish reflections, darker on the bottom.
Receptacle surface smooth, concolorous with hymenium;
margin whole (rarely cracked), even to lobed, wavy,
curved. Flesh fragile, waxy, up to 1.5 mm thick, pale
brown. Medullar excipulum up to 950 lm thick, 3-layered,
upper layer of textura globulosa-angularis, cells 30–80
(100) lm diam.; intermediate layer of textura intricate;
hyphae 4–6 lm diam., with presence of a few rounded
cells, 10–15 lm diam.; lower layer identical to the upper
123
158
Fig. 109 Diaporthe garethjonesii (MFLU 13-0261, holotype).
a Diseased leaves. b Conidiomata on the host surface. c Longitudinal
section of conidiomata. d Conidiomata on host. e–g Conidiogenous
cells with developing conidia. h–m Conidiogenous cells with
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Fungal Diversity (2016) 80:1–270
developing conidia stained with lactophenol cotton blue. n Alpha
conidia. o Beta conidia. p Germinating conidium. q, r Colonies on
PDA, q from above, r from below. Scale bars c, = 200 lm, e–
p = 10 lm
Fungal Diversity (2016) 80:1–270
Fig. 110 Diaporthe siamensis (MFLUCC 12-0300, NTCL056-1).
a Herbarium specimen. b Appearance of conidiomata on the host
surface. c Vertical section of conidioma. d Peridium. e Ostiole. f,
g Conidiogenous cells with developing conidia. h Alpha conidia.
159
i Beta conidium. j Germinating alpha conidium. k–l Culture on PDA,
k top, l reverse. Scale bars b = 500 lm, c, d = 100 lm, e = 50 lm,
f–j = 10 lm
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160
Fungal Diversity (2016) 80:1–270
Fig. 111 Maximum parsimony
(MP) majority rule consensus
tree of Cytospora isolates based
on a combined dataset of ITS,
LSU, RPB2 and ACT sequence
data. Values above the branches
indicate maximum parsimony
and maximum likelihood
bootstrap C70 %, (MPBS/
MLBS). Values at the third
positions, respectively, above or
below the branches represent
posterior probabilities (BI
PP C 0.90) from Bayesian
inference analysis. The tree is
rooted with Phomopsis vaccinii
(ATCC 18451). The strain
numbers are mentioned after the
species names. The species
obtained in this study are in blue
bold and ex-type strains are in
black bold
one. Ectal excipulum up to 250 lm thick, textura globulosa
or globulosa-angularis, cells 15–20 lm diam. Subhymenium up to 100 lm thick, textura globulosa-angularis
(cells 5–10 lm diam.), mixed with intricate hyphae. Asci
190–225 9 12–15 lm, cylindrical, amyloid, 8-spored,
amyloidity visible only on the top, lower part manifestly
destrinoid. Paraphyses clavate to irregularly swollen,
several bent in the upper part, a few with oil drops,
septate, simple or forked, with some moniliform cells,
2.5–4 lm in the lower part, up to 10 lm at the apex.
Ascospores 15–17 9 7.5–9.5 lm, elliptical, distinctly and
very delicately warted, ornamentation as very fine warts,
punctiform or slightly elongated (0.4–0.6 lm), hyaline,
123
without oil drops, uniseriate in the ascus. All tissues
hyaline to brownish, except the ectal excipulum that is
pale reddish-brown. Asexual morph Undetermined.
Habitat and known distribution: on the sand in dune
zones and on high dunes, near degraded remnants of Ammophila arenaria and Eryngium maritimum; so far known
only from Italy and Spain. Autumn-spring.
Material examined: ITALY, Calambrone, Pisa, Leg.
G. Medardi, 26 April 1996, AMB 17136, Rosolina Mare,
Rovigo, Leg. A. Lantieri and G. Medardi, 7 December 2014,
AMB 17135 (holotype). SPAIN, Playa de Xagò, Asturias,
Leg. C. Lopez-Alvarez, 15 February 2006, K(M) 140524 (sub
ammophila).
Fungal Diversity (2016) 80:1–270
161
Fig. 112 Maximum parsimony
(MP) majority rule consensus
tree of Cytospora isolates based
on ITS sequence data. Values
above the branches indicate
maximum parsimony and
maximum likelihood support
C 70 %, (MPBS/MLBS).
Values at the third positions,
respectively, above or below the
branches represent posterior
probabilities (BI PP C 0.90)
from Bayesian inference
analysis. The tree is rooted to
Phomopsis vaccinii. New strains
are in blue bold and ex-type
strains are in black bold
Notes: In the year 1996, on the dunes near Calambrone
(Pisa, Italy), we collected one portion of a strange fungal
structure, made up by a sort of hemisphaerical accumulation of small, brown apothecia, quite overlapping; due to
the particularly rainy period and the advanced age, the
specimen was not in good conditions to be photographed,
but we preserved it however (AMB 17136) because of its
interesting microscopic characters. In 2006 one Spanish
colleague collected one identical formation on sand in
Asturias, which revealed the same microscopic features,
still referable to any known species; we placed this collection in herbarium under a provisional name
[K(M) 140524], waiting for other more detailed studies. In
December 2014 near Rosolina Mare (Rovigo, Italy) we
newly found this fungus (AMB 17135). The microscopic
study showed the same characteristics of the two previous
collections, and the following molecular analyses confirmed it as a new species.
According to current information, Peziza fruticosa
seems to be a rare species, however its easily identifiable
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Fig. 113 Cytospora cotini (holotype). a Appearance of fruiting
bodies in wood. b Fruiting bodies on substrate. c Close up of fruiting
body. d Cross section of the conidioma. e Peridium. f Conidiophores
with conidia. g Conidia. h Germinating spore. i Culture characters on
MEA. Scale bars a = 2 mm, b = 1 mm, c = 400 lm, d = 300 lm,
e = 20 lm, f, g = 10 lm, h = 40 lm
by well-defined macro- and microscopic features, such as
the clustered-overlapping apothecia, amyloid asci only at the
apex, and the manifest destrinoidity. Phylogenetically,
P. fruticosa is closely related to P. domiciliana, P. perdicina
P. ampliata and P. ninguis (Fig. 100). However, P. domiciliana Cooke differs by growing gregarious or occasionally
cespitose on cellars, damp walls, plaster, mortar, mushroomcaves, and smaller ascospores, 14.5–15.5 9 7–8 lm, with
two oil drops (Seaver 1917; Le Gal 1941; Maas Geesteranus 1967; Rifai 1968; Hansen et al. 2002). Peziza
perdicina (Velen.) Svrček has ascospores 15–16 9
6–7 lm, and occurs on dung (Svrček 1976; Donadini 1981;
Häffner 1985; Cacialli et al. 1997; Doveri 2004; Medardi
2006). Peziza ampliata Pers. has ascospores 18–20.5 9
9.5–11.5 lm, smooth and grows on hardwood trunks, rich
soil mixed with wood-chips, chalk (Svrček 1970; Dennis
1981; Hansen et al. 2002; Medardi 2006).
Concerning P. ninguis Donadini & Trimbach, we had
the possibility to examine the collection ref. JF908536,
consisting of one specimen kept in MCVE (ref. 11883),
and we noticed it fully matches the characters of P. varia
(Hedw. : Fr.) Alb. & Schwein; the name under which its
sequence was filed in GenBank is therefore incorrect.
P. ninguis is one of the synonyms of P. nivalis (R. Heim &
L. Rémy) M.M. Moser, that has divergent microscopic
characters: ascospores remarkably larger and differently
ornamented, 18–22.5 9 9–10 lm vs. 14–16 (17.5) 9 9–11
(12) lm for P. varia (Medardi et al. 2012), with a very
feeble and inconstant punctuation instead of the typical
superficial crimp, and growth in nival environment only in
solstitial period, often immersed in water from melting
snow fields. The fungi of MCVE 11883 were collected on
04/05/1996 in the municipality of Trasaghis (Udine, Italy),
one town located only a few hundred meters above sea
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Fungal Diversity (2016) 80:1–270
level and therefore not in the expected typical glacial
environment, and moreover in an inappropriate period. The
sample was previously identified as ‘‘P. ninguis var. fortoulii’’, but also in this case the characters are not coincident, because the ascospores of P. fortoulii Donadini &
Neville are larger [16–18 (19) 9 10.5–12.5 lm] and
smooth (our personal observations).
Among the morphologically closest species, P. proteana
(Boud.) Seaver f. sparassoides (Boud.) Korf shares similar
habits with P. fruticosa, such as several apothecia often
coalesced into a large, irregular compound cauliflower-like
mass, but differs in smaller ascospores 10–13 9 5–7 lm
with a strongly warty ornamentation, and by growing on
163
burnt soil (Seaver 1917, 1928; Durand 1919; Korf
1956, 1973; Dennis 1981; Donadini 1981; Geesink 1984;
Hohmeyer 1986; Van Vooren 2003; Medardi 2006; Barseghyan and Wasser 2007).
The sand-associated P. ammophila Durieu & Lév. shows semi-hypogean, funnel-shaped not coalesced apothecia, at first
sphaerical and then opened and splitting into a stellate shape,
long stalked, and ascospores 16–18 9 9.5–11 lm, elliptical
and smooth (Donadini 1981; Hansen et al. 2002).
Sordariomycetes
For Sordariomycetes for follow Maharachchikumbura
et al. (2016).
Fig. 114 Phylogram generated
from maximum parsimony
analysis based on combined
ITS, GADPH, CHS, ACT and
TUB2 sequence data from
species of dematium species
complex. Maximum parsimony
bootstrap support values greater
than 50 % and Bayesian
posterior probabilities greater
than 0.70 are shown above the
branches. The new isolate is in
blue. The tree is rooted with
Colletotrichum nigrum
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Fungal Diversity (2016) 80:1–270
Fig. 115 Colletotrichum insertae (holotype). a Appearance of
conidiomata on host. b Black acervuli with setae. c Setae. d Blunt
rounded tip of setae. e Base of the setae. f–h Conidiogenous cells. i–
l Conidia. Scale bars a, b = 200 lm, c = 50 lm, d = 5 lm,
e = 150 lm, f = 20 lm g–l = 5 lm
Coronophorales Nannf.
Member of the order Coronophorales are wood inhabiting fungi and comprise the families Bertiaceae,
Chaetosphaerellaceae, Coronophoraceae, Nitschkiaceae
and Scortechiniaceae (Maharachchikumbura et al.
2015, 2016). The taxa in the order are characterized by
mostly superficial ascomata, sometimes with an extensive
hyphal subiculum or well-developed basal stroma, that
often becomes cupulate or collapsed, and in some cases the
ostiolar opening is either indistinct or lacking (Mugambi
and Huhndorf 2010; Maharachchikumbura et al. 2016).
characterized by immersed, erumpent or superficial ascomata, lack of ostiole, thin-walled asci with long stipe and
large ascospore numbers. Maharachchikumbura et al.
(2015, 2016) showed the family Coronophoraceae is
monophyletic, however it was only represented by the type
species C. gregaria.
Coronophoraceae Höhn.
The order and family were treated by Maharachchikumbura et al. (2015, 2016) and is followed here.
Coronophora Fuckel
The genus Coronophora was introduced by Fuckel (1864)
and Mugambi and Huhndorf (2010) reported a single collection of Coronophora gregaria Fuckel from USA. It is
123
Coronophora myricoides H.X. Wu & K.D. Hyde, sp. nov.
Index Fungorum number: IF552180; Facesoffungi
number: FoF02451, Fig. 104
Etymology: Referring to the resemblance of the ascomata (under the stereomicroscope) to the berries of Myrica
rubra (waxberry).
Holotype: IFRD 9201.
Saprobic on the surface of dead wood. Asexual morph
Undetermined. Sexual morph Ascomata 373–395 lm
high 9 282–340 lm diam. (
x = 383 9 308 lm, n = 10),
perithecial, superficial or erumpent through bark of host,
clustered in small to large groups, subglobose, surface
tuberculate, dark brown to black, lacking ostioles.
Fungal Diversity (2016) 80:1–270
165
Fig. 116 Phylogram generated from maximum likelihood analysis
based on combined LSU, ITS sequence data from species of
Glomerellales. Maximum likelihood bootstrap support values greater
than 50 % and Bayesian posterior probabilities greater than 0.90 are
shown in above and below and branches with maximum parsimony
bootstrap support values greater than 50 % are in bold. The new
isolates is in red and other ex-type strains are in bold. The tree is
rooted with Microascus longirostris
Peridium 27–135 lm (27–56 lm at the apex, 93–135 lm
at the base), composed of five layers of dark brown-walled
cells of textura globosa, inner layer of hyaline pseudoparenchymatic cells, slightly melanized on the outer surface. Paraphyses not seen. Asci 102–147 9 11–27 lm
(
x = 119 9 22 lm, n = 15), numerous, polysporous,
unitunicate, cylindrical to clavate, pedicel 11–38 9 4–
7 lm. Ascospores 6–8 9 1–2 lm (
x = 7 9 1.7 lm,
n = 30), numerous, hyaline, 1-celled, cylindrical or ellipsoidal, narrowly rounded at both ends, smooth-walled.
Material examined: CHINA, Yunnan Province, Jing
Dong, on dead wood of unidentified plant, 13 September
2013, Wu Hai Xia (IFRD 9201, holotype).
Notes: The genus Coronophora is typified by Coronophora gregaria and there are 28 epithets in Index Fungorum (2016). Kirk et al. (2008) estimated that there are five
species in Coronophora from wood. There are two sequences for Coronophora gregaria in GenBank. In this paper, we
introduce a new species of Coronophora; C. myricoides
which differs from C. gregaria in the shape of the ascomata
and also the shape of ascospores. The ascospores of Coronophora myricoides (6–8 9 1–2 lm) are smaller the type
species (8–12 9 2–3 lm). We directly extracted DNA
using ascomata and sequencing. We add the molecular
data to better understand the generic and familial
circumscriptions.
123
166
Fig. 117 Blastophorum aquaticum (DLU084, holotype). a–c Conidiophores and conidia. d Conidiogenous cell. e–k Conidia. l Germinating conidium. m Surface view of culture on MEA. n Reverse view
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Fungal Diversity (2016) 80:1–270
of culture on MEA. Scale bars a = 60 lm, b = 50 lm, e,
l = 20 lm, c = 15 lm, d = 10 lm, f–k = 5 lm
Fungal Diversity (2016) 80:1–270
Diaporthales Nannf.
Members of Diaporthales are pathogens, parasites, and
endophytes of plants, human-animal pathogens, saprobes and
soil inhabitants (Rossman et al. 2007). The order Diaporthales
is characterized by perithecia with an elongate beak, often
forming within stromatic tissues (Rossman et al. 2007). Asci
generally deliquesce at the base when mature and have a
characteristic refractive apical annulus. The order Diaporthales comprises 12 families: Cryphonectriaceae, Diaporthaceae, Gnomoniaceae, Harknessiaceae, Macrohilaceae, Melanconidaceae, Pseudoplagiostomataceae, Pseudovalsaceae, Schizoparmaceae, Stilbosporaceae, Sydowiellaceae and Valsaceae (Maharachchikumbura et al. 2016).
Diaporthaceae Höhn. ex Wehm.
Von Höhnel (1917) established the family Diaporthaceae and accommodated it in the order Diaporthales. The family has undergone various taxonomic
revisions during the past century (Wehmeyer 1975; Barr
1978; Castlebury et al. 2002; Dai et al. 2014).
Maharachchikumbura et al. (2015) accepted eleven genera
in Diaporthaceae: Allantoporthe, Apioporthella, Clypeoporthella, Diaporthe, Diaporthella, Diaporthopsis, Leucodiaporthe, Mazzantia, Mazzantiella, Ophiodiaporthe and
Pustulomyces.
Diaporthe Nitschke
Diaporthe has a worldwide distribution as endophytes,
pathogens and saprobes (Udayanga et al. 2011). The
asexual morph was previously known as Phomopsis. These
genera were linked in Wijayawardene et al. (2014a) and
Maharachchikumbura et al. (2015). Rossman et al. (2015)
proposed to conserve Diaporthe over Phomopsis, to resolve
nomenclatural problems. An update of the phylogeny of
Diaporthales species with new species and records is
presented herein (Fig. 105).
Diaporthe aseana Dissanayake, Tangthirasunun & K.D.
Hyde, sp. nov.
Index Fungorum number: IF551402; Facesoffungi
Number: FoF00925, Fig. 106
Etymology: In reference to the ASEAN, the ‘Association
of Southeast Asian nations’ where the fungus was collected
in Thailand, one of the member countries.
Holotype: MFLU 13-0256.
Saprobic on dead leaves. Sexual morph Undetermined.
Asexual morph Conidiomata 140–200 lm diam. 9 220–
300 lm high (
x = 185 9 260 lm, n = 10), associated
with necrotic leaf tissue; pycnidial, globose, unilocular, black,
erumpent, ostiolate, walls comprising 5–6 layers of dark
brown cells of textura angularis. Conidiophores
8–15 9 2–3 lm, hyaline, smooth, densely aggregated,
cylindrical, straight to sinuous. Conidiogenous cells
167
6–10 9 2–3 lm, phialidic, cylindrical, terminal and lateral,
slightly tapering towards the apex. Paraphyses not
observed. Alpha conidia 6–9 9 2–3 lm, aseptate, hyaline,
smooth, guttulate, fusoid to ellipsoid, tapering towards both
ends, straight, apex subobtuse. Beta conidia not observed.
Culture characteristics: Colonies covering 9 cm Petri
dish after 2 weeks in the dark at 28 °C. On PDA surface
with fluffy white aerial mycelium with patches of saffron,
in reverse patches of luteous to olivaceous-grey.
Material examined: THAILAND, Phayao, Jam Pa
Thong Waterfall, on dead leaf, 12 March 2012, N.
Tangthirasunun (MFLU 13-0256, holotype); ex-type living
cultures MFLUCC 12-0299a, KUMCC 15-0112.
Notes: Diaporthe aseana did not produce a sexual
morph in culture or on the host. Multi-gene phylogenies
reveal a close relationship of D. aseana to D. hongkongensis, D. lithocarpus, D. eucalyptorum, but support our
new taxa as not being conspecific. Our phylogenetic
analyses position D. aseana basal to the above species
(Fig. 105). Based on a Blast search of NCBI’s GenBank
nucleotide database, the closest matches for the ITS
sequence of D. aseana are an undescribed endophytic Diaporthe sp. InaCC F-238 from Cinchona calisaya,
Indonesia (AB899784; Identities = 544/549 (99 %),
Gaps = 2/549 (0 %), and an endophytic Diaporthe sp.
M23-2 from Abies beshanzuensis from Zhejiang Province,
China (HM595506; Identities = 538/549 (98 %)).
Diaporthe eres Nitschke, Pyrenomycetes Germanici 2: 245
(1870)
Facesoffungi number: FoF02182, Fig. 107
Saprobic on dead cone of Picea excelsa found on land.
Sexual morph Undetermined. Asexual morph Conidiomata
225–400 lm
diam. 9 235–280 lm
high
(
x = 330 9 255 lm, n = 10), pycnidial, sub-globose to
globose, unilocular, black, immersed to sub-immersed,
ostiolate, peridium consisting of 3–5 layers of dark brown
cells of textura angularis. Conidiophores 5–9 9 1–1.5 lm,
hyaline, smooth, cylindrical, straight to sinuous. Conidiogenous cells 0.5–1 lm diam, phialidic, cylindrical, terminal
and lateral, with slight taper towards apex. Paraphyses not
observed. Alpha conidia 5–6 9 2–3 lm aseptate, hyaline,
smooth, guttulate, fusoid to ellipsoid, tapering towards both
ends, straight, apex subobtuse. Beta conidia not observed.
Culture characteristics: Colonies white and pale brown
on surface, reverse pale brown to black. Aerial mycelium
white and brown, feathery, with concentric zonation,
margin fimbriate, with visible conidiomata at maturity.
Material examined: ITALY, on dead cone of Picea
excelsa (Pinaceae), 12 April 2012, Erio Camporesi (MFLU
13-0334), living cultures MFLUCC 12-0351.
Note: Diaporthe eres is an important plant pathogen
(Gomes et al. 2013; Udayanga et al. 2014a, b; Dissanayake
123
168
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Fungal Diversity (2016) 80:1–270
Fungal Diversity (2016) 80:1–270
169
et al. 2015) and has been reported on various woody hosts
(from 121 host genera from 61 host families; https://nt.arsgrin.gov/fungaldatabases). This is the first record of D. eres
on Picea excelsa. In our phylogenetic analysis of combined
ITS, EF-1a, b-tubulin and CAL sequence data of Diaporthe species (Fig. 105), strain MFLUCC 12-0351
grouped together with ex-type strains of D. eres (ex-epitype culture AR5193).
Diaporthe foeniculina Niessl, in von Thümen, Contr. Ad.
Fl. Myc. Lusit. 2: 30. 1880.
Facesoffungi number: FoF02183, Fig. 108
Saprobic on dead branch of Laburnum. Sexual morph
Undetermined. Asexual morph Conidiomata 215–390 lm
diam. 9 160–235 lm high (
x = 360 9 185 lm, n = 10),
pycnidial, eustromatic, unilocular, semi-immersed, dark
brown, scattered or aggregated, ostiolate, peridium consisting of brown thick-walled cells of textura angularis,
conidial mass globose to conical and exuding in cirrhi,
yellow to reddish brown. Conidiophores 20–28 9 2–3
(
x = 24 9 2 lm, n = 10) lm hyaline, subcylindrical or
cylindrical, filiform, tapering towards the apex. Alpha
conidia 6–9 9 2–3 lm (
x = 7 9 3 lm, n = 10), hyaline,
oblong to ellipsoidal, apex bluntly rounded, base obtuse to
Fig. 119 Ophiocordyceps hemisphaerica. a Overview of the stromata and the host (FURB 45181). b Fertile head (FLOR 59525,
holotype). c Cross section showing the complete immersed
perithecia. d Ascoma entirely isolated from the fertile head. e Conidia.
f Asci. g Apical cap. h–i Asci with part-spores. Scale bars a = 2 cm,
b–c = 1 mm, d = 1000 lm, e = 10 lm, h–i = 10 lm
b Fig. 118 Phylogram generated from Bayesian analysis based on ITS
sequence data of Ophiocordyceps. Metacordyceps taii is used as
outgroup taxon. Maximum likelihood bootstrap values greater than
50 % and Bayesian posterior probabilities over 0.90 are indicated
above and below the nodes (BBP/BP), respectively. The new species
are indicated in blue, holotypes in bold
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Fungal Diversity (2016) 80:1–270
sub-truncate, bi- to multi-guttulate. Beta conidia 26–
34 9 1–2 lm (
x = 28 9 1 lm, n = 10), hyaline, smooth,
slightly curved.
Culture characteristics: Colony is entirely white both on
surface and reverse. Aerial mycelium cottony and feathery,
colonies reaching 60 mm diam. after 5 days in 28 °C.
Fig. 120 Ophiocordyceps lacrimoidis. a Holotype, overview of the
stromata and the host attached by mycelium on a twig. b Fertile
region. c Fertile head in a cross section showing the complete
immersed perithecia. d Vertical section showing the superficial
perithecia. e Asci. f Layer of conidiogenous cells with a conidium
attached. g Conidia. Scale bars a = 5 mm, b–d = 1 mm, e = 20 lm,
f = 20 lm, g = 25 lm
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Material examined: ITALY, on dead branch of Laburnum (Fabaceae), 21 August 2012, Erio Camporesi (MFLU
13-0320), living cultures MFLUCC 12-0668.
Note: Diaporthe foeniculina has been reported on various woody hosts (from 20 host genera from 16 host families; https://nt.ars-grin.gov/fungaldatabases). This is the
first record of D. foeniculina on Laburnum sp. In our
phylogenetic analysis of combined ITS, EF-1a, b-tubulin
and CAL sequence data of Diaporthe species (Fig. 105),
strain MFLUCC 12-0668 grouped together with ex-type
strains of D. foeniculina with very high support (ex-epitype
culture CBS 111553).
Diaporthe garethjonesii Dissanayake, Tangthirasunun &
K.D. Hyde, sp. nov.
Index Fungorum number: IF551403; Facesoffungi
Number: FoF00926, Fig. 109
Etymology: In reference to the significant contribution of
E.B. Professor Gareth Jones made to Thai mycology.
Holotype: MFLU 13-0261.
Saprobic on dead leaves. Sexual morph Undetermined.
Asexual morph Conidiomata 85–125 lm diam. 9 80–
100 lm high (
x = 115 9 85 lm, n = 10), associated with
necrotic leaf tissue; pycnidial, globose, unilocular, black,
immersed to sub-immersed, ostiolate, walls consisting of
5–6 layers of dark brown cells of textura angularis.
Conidiophores 5–12 9 1–1.5 lm, hyaline, smooth, cylindrical, straight to sinuous. Conidiogenous cells 0.5–1 lm
diam., phialidic, cylindrical, terminal and lateral, with
slight taper towards apex. Paraphyses not observed. Alpha
conidia 5–6 9 2–3 lm, aseptate, hyaline, smooth, guttulate, fusoid to ellipsoid, tapering towards both ends,
straight, apex subobtuse. Beta conidia 40–50 9 3–4 lm
hyaline, smooth, less common than alpha conidia, straight,
curved or hamate.
Culture characteristics: Colonies with sparse aerial
mycelium covering the dish after 2 weeks in the dark at
28 °C. On PDA buff, honey to isabelline, reverse smokegrey.
Material examined: THAILAND, Kanjanaburi, on dead
leaf, 5 May 2012, Jayarama Bhat (MFLU 13-0261, holotype), ex-type living cultures MFLUCC 12-0542a,
KUMCC15-0117.
Notes: Diaporthe garethjonesii forms a well-supported
clade in our phylogenetic analysis with high bootstrap and
Bayesian values (Fig. 105). The BLAST comparison of the
ITS sequence of D. garethjonesii showed a 99 % match to
a fungal endophyte from Hong Kong (DQ485955) (Identities = 543/548 (99 %), Gaps = 4/548 (0 %)), a 98 %
match to the ITS sequence of USA isolate NY8658c
(HQ108026) (Identities 537/550 (98 %), Gaps 4/550
(0 %)). The distinct clade that D. garethjonesii forms in the
phylogenetic analysis, represents a separate species.
171
Diaporthe siamensis Udayanga et al., Cryptogamie
Mycologie 33: 298 (2012)
Facesoffungi number: FoF02398, Fig. 110
Saprobic on dead leaves of woody plants. Sexual morph
Undetermined. Asexual morph Conidiomata pycnidial,
185–200 lm high, 150–200 lm diam. (
x = 190 9 170 lm,
n = 10), ampulliform, scattered, immersed, ostiolate, with
elongate black neck. Pycnidial wall, comprising 2–3 layers,
with heavily pigmented outer layer, thick-walled, comprising
blackish to dark brown cells of textura angularis, with lighter
cells towards the inside, with inner layer composed of 1–2
layers, hyaline, thin-walled cells of textura angularis. Conidiophores reduced to conidiogenous cells. Conidiogenous
cells enteroblastic with percurrent annellations, integrated,
solitary, hyaline, smooth-walled and formed from the inner
layer of pycnidium wall. Paraphyses extending above conidiophores and hyaline, 20–35 lm long. Alpha conidia 3–
5 9 2–3 lm (
x = 3.5 9 2.4 lm, n = 30), hyaline, smoothwalled, mono- or bi-guttulate. Beta conidia15–20 9 1.5–
2 lm (
x = 18.5 9 1.8 lm, n = 30), aseptate, hyaline,
hamate or curved, apex acutely rounded.
Culture characteristics: Colonies on PDA 30 mm diam.
after 4 weeks at 25 °C, cream to white mycelium, cottony and
lobate at the margins, white at the center; reverse yellowish.
Material examined: THAILAND, Chiang Rai Province,
Thasud, Muang District, Mae Fah Luang University, on
dead leaves of Castanopsis sp. (Fagaceae), 21 March
2012, N. Tangthirasunun (MFLU 13-0257, NTCL056-1),
living culture, MFLUCC 12-0300.
Notes: Diaporthe siamensis was introduced by
Udayanga et al. (2012). This species was described on
leaves of Dasymaschalon sp. (Annonaceae) from Chiang
Rai, Thailand. Since then this species has not been reported
from any locality worldwide. Here we have collected
D. siamensis for the first time on Castanopsis sp. (Fagaceae) from a close locality in Thailand. The morphological characters are similar to the type specimen. Based
on the phylogenetic analysis of combined ITS, EF1-a, CAL
sequence data, shows high support to its ex-type strain
MFLUCC 10-0573a, clusters with high support with D. siamensis MFLUCC 12-0300 (Fig. 105). According to
Udayanga et al. (2012) this species was found in different
habitats of Dasymaschalon sp. whereas, in this study we
isolated it as a saprobic fungus. Identification of D. siamensis, within the same locality on two different plant
groups, reveals the potential of this species to occur on
multiple hosts in different habitats.
Valsaceae Tul. & C. Tul.
The family Valsaceae was introduced by Tulasne and
Tulasne (1861) and placed in Diaporthales by Barr (1978).
Most of Valsaceae species are plant pathogens causing
canker and dieback disease, with damage to several
123
Kobayasi and Shimizu (1982)
6–9 9 1
–
5–6 diam.
620–700 9 200–250
2.5–3.5 9 0.7–1.2
Beneath a branch
O. discoideicapitata
(Japan)
650–700 9 200–250
4–5 9 1
On twigs
(Brazil)
O. lacrimoidis
economic crops worldwide (Adams et al. 2005; Fan et al.
2014, 2015a, b; Ariyawansa et al. 2015a, Li et al. 2016,
Maharachchikumbura et al. 2015, 2016). Currently this
family comprises 13 genera: Amphicytostroma, Chadefaudiomyces, Cryptascoma, Cytospora, Ditopellina, Durispora, Harpostroma, Hypospilina, Kapooria, Leptosillia,
Maculatipalma, Pachytrype and Paravalsa (Maharachchikumbura et al. 2015, 2016). Valsaceae was
restricted to Cytospora as asexual morph and four sexual
morph genera viz. Valsa, Leucostoma, Valsella, and
Valseutypella (Fries 1838; Saccardo 1884; Gvritishvili
1982; Spielman 1985; Adams et al. 2002, 2005; Castlebury
et al. 2002; Bulgakov 2010; Yang et al. 2015; Li et al.
2016; Maharachchikumbura et al. 2015, 2016). Therefore,
Adams et al. (2005) synonymized all sexual genera under
Valsa as a subgenus or species without additional infrageneric rank, according to the International Code of
Nomenclature for Algae, Fungi, and Plants (ICN) in 2011
for placement on the list of protected fungi (Adams et al.
2005; Fotouhifar et al. 2010, Fan et al. 2015a; Wingfield
et al. 2012; Crous et al. 2015a; McNeill et al. 2012;
Rossman et al. 2015; Ariyawansa et al. 2015a, b, c; Li et al.
2016; Maharachchikumbura et al. 2015, 2016). Cytospora
species have been reported as the asexual morph of most
taxa in Valsaceae, and they are characterized by single or
labyrinthine locules, filamentous conidiophores and allantoid, hyaline conidia. In moist conditions, conidia emerge
from the fruiting bodies as yellow masses, which later
become orange to red (Spielman 1983, 1985; Adams et al.
2005, 2006; Li et al. 2016; Maharachchikumbura et al.
2015, 2016). There are 579 epithets for Cytospora in Index
Fungorum (2016). Ex-type sequence data are available for
only a few species and it is, thus, difficult to identify isolates to the species level (Liu et al. 2015a; Ariyawansa
et al. 2015a, b, c; Li et al. 2016; Maharachchikumbura
et al. 2015, 2016). Further studies and recommended on
Cytospora to clarify cryptic species with updated gene
trees (Adams et al. 2002; Fotouhifar et al. 2010; Hyde et al.
2010, 2014; Fan et al. 2015a, b; Liu et al. 2015a; Ariyawansa et al. 2015a, b, c; Yang et al. 2015; Li et al. 2016).
–
4–5 9 3–5
8–14 9 2
350–450 9 5
–
590–700 9 2
750–800 9 270–280
(Japan)
O. dipterigena
On twigs
3–6 9 0.3–0.5
–
–
–
–
5–10 9 1
On twigs
(Sri Lanka)
O. dipterigena
This study
Kobayasi and Shimizu (1978)
4.5–9 9 1.5–2.5
–
10 9 1.5
–
–
500–640 9 5–6
780–860 9 220–290
12–20 9 0.8–1
On twigs
(Brazil)
Ophiocordyceps hemisphaerica
–
Berkeley and Broome (1875)
This study
6.2–8.3 9 2.5–3.5
–
11–14 9 2
–
550–700 9 5
850–920 9 230–300
9–13 9 0.5
Under the leaf
Cordyceps muscicola
123
(Brazil)
Conidia
(lm)
Part-spores
(lm)
Ascospores
(lm)
Asci
(lm)
Ascomata
(lm)
Stromata
(mm)
Host behavior
Species
Table 1 Synopsis of Ophiocordyceps species discussed in the paper
Möller (1901)
Fungal Diversity (2016) 80:1–270
Reference
172
Cytospora Ehrenb.
Cytospora was introduced by Ehrenberg (1818), and it
contains species that are one of the most important
pathogens, causing canker disease on branches leading to
large areas of dieback on a wide range of plants (Adams
et al. 2005, 2006). The genus Cytospora has sexual morphs
in Valsa, Leucostoma, Valsella, and Valseutypella (Adams
et al. 2005). All sexual genera were synonymized with
Valsa either as a subgenus or species. Cytospora (1818) is
an older name than Valsa (1849) and the Cytospora morph
is more common in nature. Therefore, Valsa species are
treated as synonyms of Cytospora. Cytospora species
Fungal Diversity (2016) 80:1–270
173
Fig. 121 Maximum Parsimony
tree based on an analysis of
combined ITS and b-tub
sequence data. Bootstrap values
greater than 50 % are indicated
at the nodes. Type strains are
shown in bold and the new
taxon is in violet
Fig. 122 Purpureocillium sodanum (holotype). a–c Colonies on MEA, CMA and CYA. d, e Subglobose conidia. f Acremonium-like
conidiophores. g Paecilomyces-like conidiophores. h Cylindrical conidia. Scale bars d, g, h = 10 lm, e, f = 5 lm
123
174
Fungal Diversity (2016) 80:1–270
Fig. 123 Phylogram generated from maximum likelihood analysis
based on combined LSU, ITS and RPB2 sequence data for taxa of
Hypocreales. Maximum likelihood bootstrap support values greater
than 50 % are shown near the nodes. The new isolates are in blue and
ex-type sequences in bold. The tree is rooted with Gelasinospora
tetrasperma and Neurospora crassa
identification has generally been established according to
host affiliation, while morphological descriptions generally
lacked detail. However, a single species of Cytospora often
occurs on taxonomically unrelated host plants (Adams
et al. 2005; Wang et al. 2011, Fan et al. 2015a, b; Ariyawansa et al. 2015a, b, c; Liu et al. 2015a;
123
Fungal Diversity (2016) 80:1–270
175
Fig. 124 Alfaria spartii (holotype). a, b Ascomata on host substrate. c Cross section of ascoma. d Ostiole. e Peridium. f, g Asci. h–
k Ascospores. l Colonies on MEA. Scale bars a, b = 500 lm, c = 100 lm, d = 50 lm, e, f, g = 20 lm, h–k = 10 lm
123
176
Maharachchikumbura et al. 2015, 2016). There are 579
epithets for Cytospora in Index Fungorum (2016) with an
estimated 110 species in Kirk et al. (2008). In this study
with introduce two new species with support from molecular data (Figs. 111, 112).
Cytospora cotini Norphanphoun, Bulgakov & K.D. Hyde,
sp. nov.
Index Fungorum number: IF552231; Facesoffungi
Number: FoF02365, Fig. 113
Etymology: from generic name of host plant, Cotinus
coggygria.
Holotype: MFLU 14-0783.
Necrotrophic on dying branches of Cotinus coggygria
Scop. Sexual morph Undetermined. Asexual morph
Conidiomata 800–1000 lm diam. pycnidial, solitary,
immersed in host tissue, multi-locule, dark brown, ostiolate. Ostiole 250–350 lm diam. at the same level as the
disc surface. Peridium comprising a few to several layers
of cells of textura angularis, with inner most layer thin,
hyaline, outer layer brown to dark brown. Conidiophores
reduced to conidiogenous cells. Conidiogenous cells
enteroblastic, phialidic, formed from the innermost layer of
pycnidial wall, hyaline, smooth. Conidia (4.9–)5.6–
6.5 9 0.8–1.4(–1.7) lm (
x = 5.9 9 1.2 lm, n = 30),
unicellular, allantoid to subcylindrical, hyaline, smoothwalled.
Culture characteristics: Colonies on MEA, reaching
1.7 cm diam. after 7 days at 25 °C, producing dense
mycelium, lobate circular, with white rough margin, after
5 days, flat or effuse on the surface, without aerial
mycelium.
Material examined: RUSSIA, Rostov region, Shakhty
city, near Grushevsky pond, shelterbelt artificial forest, on
dead branches of Cotinus coggygria Scop. (Anacardiaceae), 18 May 2014, Timur S. Bulgakov (MFLU
14-0783, holotype; KUM, isotype); ex-type living cultures, MFLUCC 14-1050, KUMCC.
Notes: Cytospora cotini is a weak pathogen on Cotinus
coggygria Scop., and is often associated with Pseudocamarosporium cotinae Norphanphoun et al. The present
study using morphology and phylogenetic analyses, places
Cytospora cotini in Valsaceae. The new species has
immersed, multi-locular conidiomata, with a single ostiole
and shares common walls with the host tissue. Phylogenetic analyses, using ITS sequence data (Fig. 112), indicate
that C. cotini can be distinguished from other species
within the genus Cytospora. The analyses based on combined ITS, LSU, RPB2 and ACT sequence data also
demonstrate that C. cotini separates from other sequenced
species, and is close to Cytospora tanaitica Norphanphoun
et al. (Figs. 111, 112). However, C. tanaitica differs in
123
Fungal Diversity (2016) 80:1–270
having a single locule with smaller conidia than our
species.
Glomerellales Chadef. ex Réblová et al.
Order ‘‘Glomerellales’’ was proposed by Chadefaud
(1960), without a Latin diagnosis which made it invalid.
Glomerellales was validly published by Réblová et al.
(2011) and comprises of four families Australiascaceae,
Glomerellaceae, Plectosphaerellaceae and Reticulascaceae (Maharachchikumbura et al. 2016).
Glomerellaceae Locq. ex Seifert & W. Gams
The family Glomerellaceae was invalidly published by
Locquin (1984) and was validated in Zhang et al. (2006).
This family was accepted as one of the three families of
Glomerellales in Réblová et al. (2011). Glomerellaceae is a
monotypic family, mainly comprised pathogens, and
characterized by the Colletotrichum asexual morph and the
Glomerella sexual morph, which was synonymized under
Colletotrichum (Hyde et al. 2014; Maharachchikumbura
et al. 2015, 2016).
Colletotrichum Corda
This genus was introduced by Corda (1831), for C. lineola Corda (Damm et al. 2009). Colletotrichum comprises
mainly pathogens, as well as endophytes and saprobes
(Cannon et al. 2012; Hyde et al. 2014). Kirk et al.
(2001, 2008) and Réblová et al. (2011) placed Colletotrichum in the family Glomerellaceae and
Maharachchikumbura et al. (2015) further confirmed the
placement of this genus. In the latter study the use of the
name Colletotrichum over its sexual name Glomerella was
followed.
Colletotrichum insertae Jayawardena, Bulgakov & K.D.
Hyde, sp. nov.
Index Fungorum number: IF552260; Facesoffungi
number: FoF02399, Fig. 115
Etymology: Based on the host species.
Hoolotype: MFLU 15-1895.
Saprobic on dying twigs and leaf stalks of Parthenocissus inserta (A. Kern.) Fritsch. Sexual morph Undetermined. Asexual morph Conidiomata 250–620 lm
(
x = 420 lm, n = 10) diam., black, acervulus, oval, solitory, gregarious. Setae straight or ±bent, abundant, dark
brown, becoming paler towards the apex, opaque, smoothwalled, septate, 1–5-septate, 55–150 lm long, base cylindrical, 6.7–8.9 lm diam., apex rounded, Conidiophores
simple, to 20 lm long, hyaline to pale brown, smoothwalled. Conidiogenous cells 11–24 9 2.8–4.9 lm
(
x = 8.5 9 2.5 lm, n = 20), hyaline, smooth-walled,
cyllindrical to slighty inflated, opening 1–2 lm wide, collarette or periclinal thickening not observed. Conidia 17.3–
Fungal Diversity (2016) 80:1–270
177
Fig. 125 Maximum likelihood
tree based on an analysis of
combined ITS and b-tub
sequence data. Bootstrap values
greater than 50 % are indicated
at the nodes. New taxa are in
violet, and other ex-type strains
are in bold
23.4 9 2.6–5.2 lm (
x = 19.3 9 3.8 lm, n = 40), L/W
ratio 5, hyaline, smooth or verruculose, aseptate, curved,
gradually tapering towards the round to slightly acute apex
and truncate base, guttulate. Appressoria not observed.
Material examined: RUSSIA, Rostov Region, Rostovon-Don City, Botanical garden of Southern Federal
University, High Park, dying twigs and leafstalks, on
Parthenocissus inserta (L.) Planch. (Vitaceae), 15 April
2015, T.S. Bulgakov (T191), (MFLU 15-1895, holotype).
Notes: No culture is available for this species; hence
direct DNA extraction from conidiomata was conducted.
The Colletotrichum dematium species complex is mainly
characterized by curved conidia (Damm et al. 2009). Multigene analyses reveal a close association between C. insertae and C. dematium with high support (75 % BT/ 1.0 PP,
Fig. 114). This species differs from C. dematium in having
abundant setae per acervulus, which becomes hyaline
towards the rounded apex and 1–5 septa, as well as smaller
conidia (C. dematium conida L/W = 6). This species differs from C. lineola in having shorter setae with 1–5 septa
which can be observed along the seta, conidiogenous cells
not having a distinct collarette, as well as having smaller
conidia (C. lineola conidia L/W = 6.6) (Fig. 115).
Reticulascaceae Réblová & W. Gams
The family Reticulascaceae was introduced to accommodate two holomorphic genera Reticulascus and
Porosphaerellopsis Samuels & E. Müll., supported by
analysis of combined ITS, LSU, SSU and RPB2 sequence
data. Even though the characters and ontogeny of these
genera differ, the centrum and interthecial tissues are quite
similar (Réblová et al. 2011). Presently, three genera are
accepted in Reticulascaceae (Maharachchikumbura et al.
2015, 2016). In this study, we place the genus
Blastophorum in family Reticulascaceae based on phylogenetic analysis (Fig. 116) and morphological characters.
Blastophorum Matsush
Matsushima (1971) introduced the genus Blastophorum
with B. truncatum as the type species. Index fungorum
(2016) lists three additional species in this genus, namely
B. fusarioides K. Matsush. & Matsush., B. pini Minter &
Hol.-Jech., and B. uniseptatum Matsush. There is no
sequence data in GenBank, and fresh collections are needed to provide a natural classification of the genus.
Blastophorum aquaticum Z.L. Luo, Bhat & K.D. Hyde,
sp. nov.
Index Fungorum number: IF552241; Facesoffungi
number: FoF02218, Fig. 117
Etymology: In reference to the aquatic habitat of this
fungus.
Holotype: DLU 084.
Saprobic on submerged decaying wood. Sexual morph
Undetermined. Asexual morph Colonies on the substrate
superficial, effuse, velvety, brownish-grey to greyish-brown.
Conidiophores macronematous, mononematous, unbranched,
percurrently proliferating, smooth, septate, dark brown, 190–
250 lm (
x = 220 lm, SD = 26.5 lm, n = 10) long, 6–
8 lm (
x = 7 lm, SD = 1 lm, n = 10) wide. Conidiogenous
cells terminal, integrated, initially enteroblastic, subsequently
polyblastic and sympodial, subhyaline at the base and hyaline
above, with inconspicuous flattened denticles at conidiogenous loci. Conidia 1–3-septate, elongated, cuneiform,
rounded at the tip, narrowed towards the flattened base,
smooth, slimy, 19–24 lm (
x = 21.5 lm, SD = 2.5 lm,
n = 20) long, 6.5–7.5 lm (
x = 7 lm, SD = 0.5 lm,
n = 20) wide. Conidial secession schizolytic.
123
178
Fungal Diversity (2016) 80:1–270
Fig. 126 Emericellopsis persica (holotype). a, b Colonies on CYA and MEA. c, d Immature and mature ascomata. e Ascospores. f Asci.
g Acremonium-like conidiophores bearing hyaline conidia. h–j Chlamydospores. Scale bars c–j = 10 lm
Culture characteristics: Colonies on MEA 70 mm diam.
after 6 weeks at room temperature, dark brown at the
margins, cream to white at the center; reverse yellowish to
dark brown and orangish at the center, medium dense,
circular, umbonate.
123
Material examined: CHINA, Yunnan Province, Dali,
Cangshan Mountain, Heilong stream, saprobic on submerged decaying wood, March 2014, X.Y Liu, S-084
(DLU084, holotype), ex-type living culture, MFLUCC
15-0264.
Fungal Diversity (2016) 80:1–270
179
Fig. 127 Phylogram generated from Bayesian analysis based on LSU
sequence data from species of Meliolaceae. Maximum parsimony/likelihood bootstrap support values greater than 50 % and
Bayesian posterior probabilities greater than 0.9 are shown in the first
and second set, respectively. Ex-type strains are in bold and the new
sequence are in blue bold
Notes: Blastophorum resembles Kylindria in having
terminal, integrated, initially enteroblastic, subsequently
polyblastic and sympodial conidiogenous cells, but
Blastophorum differs in having cuneiform and larger
conidia. The phylogeny analysis (Fig. 116) also showed
Blastophorum separates from Kylindria. Blastophorum
aquaticum differs from B. pini in having elongated,
cuneiform and wider conidia (6.5–7.5 lm vs. 1.5–2.5 lm)
(Minter and Holuboá-Jechová 1981). It differs from
B. truncatum in having longer conidiophores (190–250 lm
vs. 18.3–132.6 lm), elongated, cuneiform, 1–3-septate and
larger conidia (19–24 9 6.5–7.5 lm vs. 8.7–17 9 2.6–
4.3 lm) (Chen and Tezan 2008).
Nectriaceae, Niessliaceae, Ophiocordycipitaceae, Stachybotriaceae and Tilachlidiaceae. Maharachchikumbura
et al. (2016)
Hypocreales Lindau
Hypocreales is an order within the class Sordariomycetes, represented by Bionectriaceae, Clavicipitaceae,
Cordycipitaceae,
Flammocladiaceae, Hypocreaceae,
Ophiocordycipitaceae G.H. Sung et al.
Ophiocordycipitaceae was proposed in Sung et al.
(2007), based mainly on phylogenetic analyses. According
to most recent studies on the family, it contains six genera,
Drechmeria, Harposporium, Ophiocordyceps, Polycephalomyces, Purpureocillium and Tolypocladium
(Quandt et al. 2014). Ophiocordyceps was first described to
accommodate species possessing ascospores that usually
do not break into part-spores at maturity, asci with thin
apical caps (Petch 1931a, 1932), and often producing a
dark pigmented, tough to pliant stromata, commonly with
perithecial apices (Sung et al. 2007). This genus is associated with several asexual morphologies, Sorosporella,
Hirsutella, Hymenostilbe, Stilbella, Syngliocladium, and
123
180
Fig. 128 Meliola citri-maximae (MFLU14-0288, holotype). a Host
leaves. b Colony on surface of leaf. c Ascoma on host substrate.
d Hyphae with hyphopodia and phialides. e Hyphae with hyphopodia.
f Hyphae with phialides. g Outer layer of peridium. h Inner layer of
123
Fungal Diversity (2016) 80:1–270
peridium. i Hamathecium. j Hyphal setae. k–q Asci from young state
to mature state. r–t Ascospore from young state to mature state. Scale
bars c, d = 200 lm, j = 100 lm, e–i, k–t = 20 lm
Fungal Diversity (2016) 80:1–270
181
Fig. 129 Meliola pseudosasae (MFU 16-2136, reference specimen). a Herbarium specimen. b Appearance of ascomata on host leaf. c Ascoma.
d Setae. e Vertical section of ascoma. f Superficial hyphae. g, h Ascospores. Scale bars d, e = 50 lm, f = 10 lm, g, h = 20 lm
Paraisaria (Quandt et al. 2014). Hymenostilbe was proposed by Petch (1931b) and contains species that produce
conidia from multiple denticles on conidiogenous cells
forming a palisade-like layer along the surface of synnemata (Mains 1950). There is evidence to restrict the use
of Hymenostilbe to the ‘‘O. sphecocephala clade’’ within
the genus Ophiocordyceps (Sung et al. 2007). The phylogenetic tree, based on ITS, composed by species of this
clade, including two new species, is presented in Fig. 118.
Ophiocordyceps hemisphaerica Mafalda-Freire, Reck &
Drechsler-Santos, sp. nov.
Index Fungorum number: IF552122; Faces of fungi
number: FoF02193, Fig. 119
Etymology: Referring to the hemisphaeric form of the
fertile head of the stroma.
Holotype: FLOR 59525
Specimens found on twigs of living plants, parasitic in
true flies of Muscidae. Sexual morph capitated. Stromata
12–20 9 0.8–1 mm, rarely branched, stipitate, usually 2,
up to 4, arising from the host thorax between the wings,
tough, with morphology unaffected when dried. Stipe 11–
19 mm long, 0.8–1 mm wide, cylindrical, with a fertile
apex, flexible when fresh, hard and tough when dried,
123
182
Fungal Diversity (2016) 80:1–270
Fig. 130 Consensus tree resulting from a Bayesian analysis of the
ITS sequence alignment of Arthrinium species. RAxML bootstrap
support values (MLB above 50 %) and Bayesian posterior probabilities (PP above 50 %) are given at the nodes (MLB/PP). The newly
introduced sequences are in blue bold. The scale bar represents the
expected number of changes per site. The tree is rooted to Xylaria
hypoxylon
smooth, brown to greyish-brown, epidermal layer compact,
medullar region white to cream, not compacted. Fertile
head 1–1.2 mm long, 2–4 mm diam., hemisphaerical,
truncate, applanate at the base, pale to dark yellowish,
ostioles inconspicuous on the upper surface, smooth-like.
Ascomata obpyriform, slightly curved, 780–860 9 220–
290 lm (
x = 820 9 255 lm, n = 40), completely
immersed, yellow, thick-walled, perithecia easily detachable from the fertile region. Asci narrow cylindrical, 500–
640 9 5–6 lm (
x = 570 9 5.5 lm, n = 40), apex thickened, 8-spored, hyaline. Apical cap 5–6 lm (
x = 5.5 lm,
n = 40) diam. Ascospores filiform, smooth, almost as long
as asci, hyaline, more than 52 septa, easily breaking into
part-spores. Part-spores cylindrical to unusually fusoid, 7–
10 9 1–1.5 lm (
x = 8.5 9 1.25 lm, n = 40), hyaline,
smooth. Asexual morph Hymenostilbe-like. Synnemata
cylindrical, 6–12 9 0.5–1 mm, simple or branched, 1 to 3,
smooth, brown to greyish-brown, arising from the thorax
and abdomen of the host, unusually arising from aborted
stromata. Conidiogenous cells phialidic, clavate, surface
slightly rugose, hyaline, compactly arranged. Conidia
obovoid,
6.2–8.3 9 2.5–3.5 lm
(
x = 7.25 9 3 lm,
n = 40), hyaline, slightly verrucous, with persistent
appendix.
123
Fungal Diversity (2016) 80:1–270
183
Fig. 131 Arthrinium paraphaeospermum (holotype). a Colony on MEA. b Sporodochium on MEA. c–f Conidiogenous cells giving rise to
conidia. h Conidia in side view. g Conidia and sterile cells. i Conidia in surface view. Scale bars a, b = 100 lm, c–i = 10 lm
Material examined: BRAZIL. Santa Catarina, Ituporanga, Rio do Norte, 27°240 0000 S, 49°360 0000 W, on dead
flies of Muscidae, 19 October 2014, Bittencourt FB 246
(FURB 45181); Florianópolis, 27°350 3300 S, 48°280 4200 W,
2010, Mafalda-Freire FMF 33 (FLOR 59524); Joaçaba,
Parque Natural Municipal Rio do Peixe, 278100 2200 S,
518300 2300 W, 10 March 2010, Mafalda-Freire FMF 104
(FLOR 59525, holotype); 24 January 2014, Mafalda-Freire
FMF 331 (FLOR 59526); Mafalda-Freire FMF 332 (FLOR
59527); Mafalda-Freire FMF 333 (FLOR 59528); 27
September 2014, Mafalda-Freire FMF 295 (FLOR 59529);
Mafalda-Freire FMF 305 (FLOR 59530); 27 September
2014, Mafalda-Freire FMF 296 (FLOR 59531); MafaldaFreire FMF 297 (FLOR 59532); Mafalda-Freire FMF 298
(FLOR 59533); Mafalda-Freire FMF 309 (FLOR 59534);
Mafalda-Freire FMF 310 (FLOR 59535); Mafalda-Freire
FMF 311 (FLOR 59536); Mafalda-Freire FMF 312 (FLOR
59537); Mafalda-Freire FMF 313 (FLOR 59538); MafaldaFreire FMF 314 (FLOR 59539); Mafalda-Freire FMF 315
(FLOR 59540); Mafalda-Freire FMF 316 (FLOR 59541);
Mafalda-Freire FMF 317 (FLOR 59542); Mafalda-Freire
FMF 318 (FLOR 59543); Mafalda-Freire FMF 319 (FLOR
59544); Mafalda-Freire FMF 320 (FLOR 59545); Mafalda-
Freire FMF 321 (FLOR 59546); Mafalda-Freire FMF 322
(FLOR 59547); Mafalda-Freire FMF 323 (FLOR 59548);
Mafalda-Freire FMF 324 (FLOR 59549); 28 September
2014, Mafalda-Freire FMF 328 (FLOR 59553); Águas
Mornas, Sı́tio Portal, 27°410 4700 S, 48°490 2900 W, 26 October
2013 Mafalda-Freire FMF 141 (FLOR 59550); Blumenau,
Parque das Nascentes, 26°570 4100 S, 49°040 1100 W, 22
February 2012, Drechsler-Santos DS 785 (FLOR 59551).
Additional material examined (as Ophiocordyceps
dipterigena): JAPAN, Ibaraki: Gozenyama, 29 July 2001,
F. Ihara 1021 (personal collection of F. Ihara); 26 July
2002, F. Ihara 02024 (personal collection of F. Ihara); F.
Ihara 2025 (personal collection of F. Ihara); 7 August 2002,
F. Ihara 04063 (personal collection of F. Ihara); 23 July
2007, F. Ihara 7044 (personal collection of F. Ihara); F.
Ihara 07045 (personal collection of F. Ihara); Kyushu, July
1950, (MICH 274571); USA, North Carolina, Cranberry,
August 1887, R. Thaxter, Reliquiae Farlowianae 612 (FH
5086); R. Thaxter, Reliquiae Farlowianae 612 (FH 5077);
R. Thaxter, Reliquiae Farlowianae 612 (MICH 274593).
Notes: Ophiocordyceps hemisphaerica is characterized
by the tough stipitate stromata, with hemisphaerical and
pale to dark yellowish fertile-head, with applanate base.
123
184
Fungal Diversity (2016) 80:1–270
Fig. 132 Phylogenetic tree of Neotruncatella endophytica EMLAS5-1 and EML-AS5-2 and related species based on maximum
likelihood analysis of ITS rDNA sequences. Sequence of Pestalotiopsis malayana was used as outgroup. Numbers at the nodes
indicate the bootstrap values ([50 %) from 1000 replications. The
bar indicates the number of substitutions per position. New taxa are in
blue
The tough stromata are resistant and the morphology is
unaffected even when dried. The primordium stromata has
conidiogenous cells and conidia during the development.
Additionally, the species parasitizes true big flies of Muscidae, which are not covered by mycelium. All parasitized
flies were found above branches of living trees and
always in advanced stage of decomposition. ITS based
phylogenies group all O. hemisphaerica isolates in a
strongly supported monophyletic clade and sharing a sister relationship to O. dipterigena species (Fig. 118).
Ophiocordyceps hemisphaerica is morphologically similar
to O. dipterigena (Berk. & Broome) G.H. Sung et al.
However, it can be differentiated by its larger stromata
(12–20 9 0.8–1 mm vs. 5–10 9 1 mm) and hemisphaerical fertile region, which are not observed in O. dipterigena
(Petch 1932; Kobayasi and Shimizu 1978, Luangsa-ard et al.
2008). Additionally, O. dipterigena was described originally
by Berkeley and Broome (1874) from material collected in
Sri Lanka (Ceylon). Cordyceps muscicola Möller, a heterotypic synonymy of O. dipterigena, was described from
123
Fungal Diversity (2016) 80:1–270
185
Fig. 133 Neotruncatella endophytica (holotype). a, d Colonies in
potato dextrose agar (PDA). b, e Colonies in oatmeal agar (OA). c, f
Colonies in malt extract agar (MEA) (a–c obverse view, d–f reverse
view). g, h Conidioma and peridium. i–k Conidia attached to
conidiogenous cells. l 3-septate conidia with apical and basal
appendages. Scale bars g = 50 lm, h–l = 20 lm
material collected in Blumenau, Santa Catarina, Brazil. In a
taxonomical perspective O. dipterigena corresponds to
a species-complex, O. hemisphaerica differs from
C. muscicola by its slightly smaller perithecia and asci
(780–860 9 220–290 lm and 500–640 9 5–6 lm vs.
850–920 9 230–300 lm and 550–700 9 5 lm, respec-
123
186
tively). Furthermore, C. muscicola presents six stromata,
smaller (9–13 mm) than those observed for O. hemisphaerica (12–20 mm); the host of C. muscicola was described as
covered by mycelium found adhered to the abaxial region of
the leaves (Möller 1901).
Ophiocordyceps lacrimoidis Mafalda-Freire, Reck &
Drechsler-Santos, sp. nov.
Index Fungorum number: IF552123, Facesoffungi
number: FoF02194, Fig. 120
Etymology: Referring to the lacrimoid form of the
conidia.
Holotype: FLOR 59552.
Specimen found attached by mycelium on a twig, parasitic in true flies of Muscidae (host very damaged). Sexual
morph capitated. Stromata 4–5 9 1 mm, simple, stipitate,
2 stromata arising from the thorax. Stipe 3–4 mm long,
1 mm wide, cylindrical, with a fertile apex, fleshy, robust,
smooth, epidermal layer brown, compact, medullar region
white to cream, soft, not compacted. Fertile head 1.2 mm
long, 1.8–2.2 mm diam., discoid, pale to dark yellowish,
ostioles inconspicuous on the upper surface. Ascomata
obpyriform, slightly curved, 650–700 9 200–250 lm
(
x = 675 9 225 lm, n = 40), completely immersed, yellow, thick-walled, perithecia easily detachable from the
fertile region. Asci narrow cylindrical, 350–450 9 5 lm
(
x = 400 9 5 lm, n = 40), apex thickened, 8-spored,
hyaline. Apical cap 3–4 9 5–6 lm (
x = 3.5 9 5.5 lm,
n = 40). Ascospores filiform, smooth, almost as long as
asci, hyaline, more than 56 septa, easily breaking into partspores.
Part-spores
cylindrical,
8–14 9 2 lm
(
x = 11 9 2 lm, n = 40), hyaline, smooth. Asexual
morph Hymenostilbe-like. Synnemata cylindrical,
3 9 0.3 mm, simple, unique, smooth, orange brown, arising from the abdomen of the host. Conidiogenous cells
phialidic, clavate, surface roughened, hyaline, pseudoparenchymatous, compactly arranged. Conidia lacrimoid,
4–5 9 3–5 lm (
x = 4.5 9 4 lm, n = 40), hyaline.
Material examined: BRAZIL. Amazonas, Manaus,
Reserva Florestal Adolfo Ducke, 3°000 1600 S 59°550 0700 W,
on a dead fly attached on a twig, 2012, Mafalda-Freire
FMF148 (FLOR 59552, holotype).
Additional material examined (as Ophiocordyceps discoideicapitata): JAPAN, Yamagata, 8 July 1982 (TNS-F.
181545, NMNS); Iwate, Morioka, 18 July 2007, F. Ihara
07031 (personal collection of F. Ihara); F. Ihara 07032
(personal collection of F. Ihara); 11 August 2007, F. Ihara
07116 (personal collection of F. Ihara); F. Ihara 07117
(personal collection of F. Ihara); F. Ihara 07118 (personal
collection of F. Ihara).
Notes: Ophiocordyceps lacrimoidis is characterized by a
robust short stipe, discoid fertile region, cylindrical synnemata and lacrimoid conidia. Additionally, the species
parasitizes true flies of Muscidae, which are not covered by
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Fungal Diversity (2016) 80:1–270
mycelium. According to phylogenetic analysis (Fig. 118),
O. lacrimoidis is in a distinct lineage in the ‘‘O. sphecocephala clade’’, as sister of O. australis (Speg.) G.H. Sung
et al. Morphologically, O. lacrimoidis is closely related to
O. discoideicapitata (Kobayasi & Shimizu) G.H. Sung
et al., but it can be differentiated by its darker fertile region
with inconspicuous ostioles. Ophiocordyceps lacrimoidis
slightly smaller perithecia than those of the O. discoideicapitata specimen studied by us (TNS-F. 181545:
700–820 9 270–280 lm) and similar to those described by
Kobayasi and Shimizu (1982, 620–700 9 250–300 lm).
The presence of synnemata and lacrimoid conidia was not
observed for O. discoideicapitata. Furthermore, O. discoideicapitata was originally described from Japan, while
the new species is described from Brazilian Amazon.
Ophiocordyceps dipterigena and O. hemisphaerica also
parasize flies, however the size of the stromata, and the
morphology of fertile region and part-spores are very different (Table 1).
Purpureocillium Luangsa-ard et al.
The genus Purpureocillium was introduced by Luangsaard et al. (2011) with P. lilacinum (Thom) Luangsa-ard
et al. as the type species, and it was reported as important
pathogens in humans. Luangsa-ard et al. (2011) also
showed that it belongs to Ophiocordycipitaceae,
Hypocreales. Besides the type species, Purpureocillium
comprises three species viz. P. atypicola (Yasuda) Spatafora et al., P. lavendulum Perdomo et al. and P. takamizusanense (Kobayasi) S. Ban, Azuma & Hirok. Sato (Index
Fungorum 2016).
Purpureocillium sodanum Papizadeh, Soudi, Wijayaw.,
Shahz.-Faz. & K.D. Hyde, sp. nov.
Faces of Fungi number: IF552178; Facesoffungi Number: FoF02247, Fig. 122
Etymology: Named after isolation of this species from
salt crystals.
Holotype: IBRC-H 2020
Saprobic on salt crystals. Sexual morph Undetermined.
Asexual morph Hyphae hyaline, smooth-walled, 1–2 lm
wide. Conidiophores growing from the superficial mycelium very long, consisting of a stipe, bearing 6–8 lm long
monoverticillate branches with terminal whorls of 2–3
phialides; stipe erect, septate, hyaline to pale purple-brown,
rough-walled, profusely verrucose and up to 5 lm wide.
Conidiophores growing from the aerial mycelium 25 to
40 nm long, irregularly branched, with 3–4 phialides per
branch, septate, hyaline and smooth-walled. Phialides
5–13 9 2–3 mm, with a cylindrical basal portion tapering
into a distinct neck up to 3 lm long. Conidia were in long
dry chains, unicellular, subglobose with apiculate base or
limoniform, 3.5–5.5 9 3–4.5 lm, smooth-walled, mostly
Fungal Diversity (2016) 80:1–270
187
Fig. 134 Consensus tree resulting from a maximum likelihood
analysis of combined LSU and ITS sequence alignment of taxa in
Xylariales. RAxML bootstrap support values (MLB above 50 %) and
Bayesian posterior probabilities (PP above 50 %) are given at the
nodes (MLB/PP). The newly introduced sequences are in blue bold.
The scale bar represents the expected number of changes per site. The
tree was rooted to Vialaea mangiferae MFLUCC 12-0808
subhyaline, pinkish-purple in mass. Acremonium-like
conidiophores were rarely observed, and when present, up
to 50 lm long, simple, usually reduced to a cylindrical
phialides, 9–22 9 1.5–2 mm. Chlamydospores and sexual
morph were not observed.
Culture characteristics: Colonies at 25 °C on MEA
attained 28–30 mm diam. after 7 days incubation, consisting of a dense basal felt of numerous conidiophores
giving a powdery texture, pale vinaceous (9B2), with
sparse aerial mycelium, and producing a very faint yellow
diffusible pigment. Colony features on CYA, CMA and
PDA were similar to those observed on MEA, except that
on PDA and CYA they were radially folded toward the
periphery. Such a folding pattern was observed on MEA
exclusively at 25 °C. Colonies on CYA were floccose and
pale vinaceous (9A2). The optimum growth temperature
was 30 °C (on MEA, 33–35 mm diam. after 7 days incubation) and the minimum was 20 °C. Colonies at 35 °C on
MEA attained 23–25 mm diam. after 7 days incubation.
Growth was insignificant at 37 °C and absent at 40 °C.
Material examined: IRAN, Markazi Province, Mighan
(Miqan) saline Lake, salt crystals, 2013, M. Papizadeh
(IBRC-H 2020, holotype), ex-type culture IBRC-M 30175.
Notes: In our phylogenetic analyses, the new strain
clusters as a distinct clade from other species (Fig. 121).
This clade is sister to the type species, but is separated
from high bootstrap value (86 %). Hence, we introduce
our strain as a new species based on DNA sequence
analyses. P. sodanum is distinguished from other Purpureocillium species based on its growth at 30 up to
37 °C.
Hypocreales incertae sedis
Alfaria Crous et al.
The monotypic genus Alfaria was introduced and typified by A. cyperi-esculenti Crous et al. The genus is placed
in Hypocreales incertae sedis based on molecular data
(Fig. 123).
Alfaria spartii Senan., Camporesi & K.D. Hyde, sp. nov.
Index Fungorum number: IF552303; Facesoffungi
number: FoF02455, Fig. 124
Etymology: In reference to the host genus Spartium.
Holotype: MFLU 16-1969.
Saprobic on Spartium junceum L. Sexual morph Ascomata 220–250 lm high 9 190–275 lm diam. (
x = 216 9
123
188
Fig. 135 Cainia globosa (MFLU 14-0738, holotype). a Appearance
of ascomata on host substrate as minute clypeate ostiolar dots.
b Section through ascoma. The blackened clypeal area is narrow.
c Peridium. d Papilla. e Pseudoparaphyses. f Ascus in Melzer’s
reagent showing lightly bluing apical apparatus. g–j Immature and
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Fungal Diversity (2016) 80:1–270
mature unitunicate ascospores in asci. k–o Ascospores. p Mucilaginous sheath. q Conidiomata on MEA. r Cross section of conidiomata.
s, t Conidiogenous cells with conidia. u Conidia. Scale bars a, q,
r = 200 lm, b = 100 lm, c, d, g–j, s–u = 50 lm, e, f, k–p = 20 lm
Fungal Diversity (2016) 80:1–270
189
99/1
100/1
Seimatosporium pseudorosae MFLUCC 14-0468
Seimatosporium pseudorosarum MFLUCC 14-0466
Seimatosporium lichenicola MFLUCC 14-0623
Discostroma fuscellum NBRC 32680
69 Seimatosporium vaccinii ICMP 7003
Seimatosporium sp. MFLUCC 14-0454
Seimatosporium sorbi MFLUCC 14-0469
63 AB593735 Seimatosporium glandigenum
94/1 Discostroma fuscellum Emma
Discostroma fuscellum NBRC 32625
94/1 Seimatosporium pseudocornii MFLUCC 14-0529
Seimatosporium cornicola MFLUCC 14-0448
Seimatosporium sp. IT1323
86/1 Seimatosporium cornii MFLUCC 14-0467
94/1 Seimatosporium cornii IT426
Discostroma cornii MFLUCC 14-1208
Discostroma tostum NBRC 32626
88/1 Seimatosporium parasiticum NBRC 32682
Seimatosporium physocarpi MFLUCC 14-0625
Seimatosporium vitis MFLUCC 14-0051
71/0.98
Discostroma botan DQ368629
64
70
Seimatosporium botan HMUC 316PD
80/1
Seimatosporium
botan NBRC 104200T
60
Seimatosporium discosioides NBRC 104201
Seimatosporium
pistaciae
CBS 138865
96/1
Seimatosporium rosae MFLUCC 14-0621
Seimatosporium hypericinum NBRC 32647
85/1 Seimatosporium eucalypti CPC 156
Seimatosporium eucalypti CPC 159
Seimatosporium eucalypti CPC 157
56 98/1 Seimatosporium falcatum CPC 13578
Seimatosporium falcatum CPC 13580
Seimatosporium eucalypti CPC 158
Seimatosporium obtusum CPC 12935
87/1 Discostroma stoneae NBRC 32690
74/0.98
Seimatosporium kriegerianum NBRC 32679
Seimatosporium elegans NBRC 32674
Seimatosporium biseptatum CPC 13584
AB593736 Seimatosporium hakeae
60 Sarcostroma restionis CBS 118154
66 Sarcostroma restionis CBS 118153
Sarcostroma bisetulatum CBS 122695
56
Seimatosporium grevilleae ICMP 10981
64 Seimatosporium leptospermi ICMP 11845
Seimatosporium mariae NBRC 32681
Seimatosporium walkeri CPC 17644
Seimatosporium quercina MFLUCC 14-1198
51
98/1
Seimatosporium foliicola NBRC 32676
Seimatosporium ficeae MFLUCC 15-0519
Strickeria kochii C138
Strickeria kochii C143
88/1
Sarcostroma lomatiae CBS 118144
Seimatosporium rhombusii MFLUCC 15-0543
100/1 Discosia neofraxinea MFLU 15-0375
Discosia fraxinea NTIT469
63
Discosia brasiliensis NTCL097-2
51
74/1 Discosia brasiliensis NTCL095
Discosia brasiliensis NTCL094-2
Discosia sp. MAFF 236709
92 Discosia artocreas NBRC 8975
55
65 Discosia pini MAFF 410149
Discosia aff. brasiliensis NBRC 104198
Discosia sp. MAFF 242783
Discosia pseudoartocreas CPC 21117
Discosia sp. MFLU:14-0299A
Discosia pseudoartocreas CBS 136438
Discosia aff. artocreas MAFF 242776
92/1
Adisciso yakushimense MAFF 242774
76/0.96
Adisciso yakushimen NBRC 104194T
Adisciso
tricellulare NBRC 32705
64 100/1
Adisciso tricellulare MAFF 237478
51
Discosia sp. MAFF 242778
Discosia sp. MFLU 14-0299
Discosia sp. MAFF 242785
Discosia sp. MFLU 14-0298
Discosia sp. MAFF 242777
100/1 Immersidiscosia eucalypti NBRC 104195
Immersidiscosia eucalypti NBRC 104196
72
Ciliochorella mangiferae MFLUCC 12-0310
82/0.98
Ciliochorella castaneae HHUF28800
Ciliochorella phanericola MFLUCC 14-0984
Pseudopestalotiopsis theae MFLUCC 12-0055
Seimatosporium
Discosia
Immersidiscosia
Ciliochorella
10
Fig. 136 One of the 16 equally most parsimonious trees obtained
from combined analyses set of LSU, ITS and b-tubulin sequence data
(CI = 0.627, RI = 0.821, RC = 0.514, HI = 0.373). MP values
([50 %) resulting from 1000 bootstrap replicates and Bayesian
posterior probabilities above 0.95 are given at the nodes. The tree is
rooted to Pseudopestalotiopsis theae (MFLUCC 12-0055). Ex-type
strains are in bold and newly introduced species is in blue
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190
Fungal Diversity (2016) 80:1–270
Fig. 137 Ciliochorella phanericola on Phanera purpurea. a, b Symptoms on leaf. c Surface of fruiting bodies. d Cross section of the
conidiostroma. e Peridium. f, g Mature conidia. h, i Colonies on MEA (I from below). Scale bars d = 300 lm, e = 30 lm and f, g = 20 lm
250 lm, n = 10), scattered, solitary, immersed, black, globose to subglobose, coriaceous, ostiolate. Ostiole lined
with hyaline, filiform periphyses. Hamathecium comprises
few, septate, hyphae-like, hyaline paraphyses. Peridium
comprising thick-walled, brown cells of textura angularis.
Asci 140–180 9 30–40 lm (
x = 155 9 37 lm, n = 20),
8-spored, unitunicate, oblong to fusiform, base sessile,
apex oblong, with J-apical ring. Ascospores 30–35 9 10–
12 lm (
x = 32 9 11 lm, n = 20), overlapping bi-seriate,
hyaline, oval to ellipsoid, unicellular, thick-walled, guttulate, guttules concentrated at the ends. Asexual morph
Undetermined.
123
Culture characteristics: Colonies on MEA reaching
2 cm after 14 days at 18 °C, circular, umbonate, filiform,
white, dense colonies, tightly attached to the media.
Material examined: ITALY, Santa Sofia, Province of
Forlı̀-Cesena [FC], Camposonaldo, on dead branch of
Spartium junceum (Fabaceae), 13 September 2013, Erio
Camporesi, IT 1452 (MFLU 16-1969, holotype; MFLU
16-1978, paratype), ex-type living culture, MFLUCC
13-0799.
Notes: Alfaria spartii is the second species of genus
Alfaria. In combined analyses of LSU and ITS sequence
data, A. spartii clustered with A. cyperi-esculenti with high
Fungal Diversity (2016) 80:1–270
support (Fig. 123). Alfaria spartii differs from A. cyperiesculenti in having large, oval ascospores with distinctly
periphysate ostioles.
Bionectriaceae Samuels & Rossman
Bionectriaceae Samuels & Rossman was introduced by
Rossman et al. (1999) to accommodate Bionectria and
other allied genera. Members of this family are characterized by uniloculate hyaline to brightly coloured, soft
perithecia with or without a well-developed stroma and
unitunicate asci (Maharachchikumbura et al. 2016).
Maharachchikumbura et al. (2016) accepted 39 genera and
confirmed the familial placement in Hypocreales, Sordariomycetes in their phylogenetic analyses.
Emericellopsis Beyma
The genus Emericellopsis was introduced by van Beyma
(1939-1940) with Emericellopsis terricola Beyma as the
type species. Grum-Grzhimaylo et al. (2013) placed this
genus in Bionectriaceae based on sequence analyses, but
Maharachchikumbura et al. (2016) treated it as Hypocreales, genera incertae sedis. Based on habitats, species of
Emericellopsis have been detected as saprobes. Moreover,
Grum-Grzhimaylo et al. (2013) showed that Emericellopsis
species can be classified into three categories based on their
origins; marine, soda lakes, and terrestrial. Our new collection, Emericellopsis persica, falls into the terrestrial
clade of this genus although detected in saline soils.
Emericellopsis persica Papizadeh, Wijayaw, Soudi & K.D.
Hyde, sp. nov.
Index Fungorum number: IF552358; Facesoffungi
Number: FoF 02517, Fig. 126
Etymology: Epithet taken from its isolation from Iran
(formerly known as Persia).
Holotype: IBRC-H 2022.
Isolated from soil from lake. Sexual morph Ascomata
dark brown, superficial on the substrate, globose, 280–
320 lm diam., non-ostiolate. Peridium 19–25 lm thick,
multi-layered, pseudoparenchymatous, composed of 3–5
layers of compressed cells. Asci saccate, 15–18 lm diam.,
with thin deliquescent wall, soon dissolving, unitunicate,
scattered irregularly in the ascocarp. Ascospores ellipsoid,
pale brown, with uneven surfaces, 8–11 9 4–5 lm
(
x = 9.5 9 4.5), surrounded by 3 longitudinal, hyaline,
sharp appendages, width up to 1 lm. Asexual morph
hyphomycetous, acremonium-like. Conidiophores mostly
simple orthotropic. Conidiogenous cells 30–40 lm long,
phialidic, tapering from 2–2.5 lm at the base to 0.5–
0.7 lm at the apex. Conidia narrowly ellipsoid, smoothsurfaced, 4.5–6.5 9 2 lm (
x = 5.5 9 2), hyaline, adhering in slimy heads. Chlamydospores mostly intercalary,
10–22 9 7–17 lm, hyaline.
191
Culture characteristics: Colonies on CYA agar fastgrowing, reaching 21–23 mm diam. in 7 days at 25 °C.
On MEA (pH 6.5) growing slower, reaching 17–18 mm
diam. in 7 days. Colonies orange-salmon-pink, later
darkening in centre due to the formation of ascomata with
tufted aerial mycelium. Reverse colourless. Exudate
absent. Decumbent vegetative hyphae thin-walled, hyaline, 0.5–1.5 lm wide. Mycelium consisting of hyaline,
smooth-walled, septate hyphae, 1–2 lm wide, often
fasciculate.
Material examined: IRAN, Lake Urmia, soil, 2011, M.
Papizadeh, M.R. Soudi (IBRC-H 2022, holotype), ex-type
culture IBRC-M 30046.
Sequence: ITS sequence GenBank KX668543. The
aligned dataset used in the analysis has been submitted
with the TreeBase under the submission ID 19688 for the
combined set of ITS and b-tub.
Notes: In our phylogenetic analyses, Emericellopsis
persica clusters as a distinct clade from other species.
Emericellopsis donezkii and E. humicola are the closest
species, but separated with high bootstrap support (100 %).
Hence, we introduce our strain as a new species based on
DNA sequence analyses (Figs. 125, 126).
Meliolales Gäum. ex D. Hawksw. & O.E. Erikss.
Hongsanan et al. (2015b) provided a monograph and a
backbone tree for Meliolales. Meliolales is in Sordariomycetes, which also comprises the family Armatellaceae.
A new tree incorporating Meliola citri-maximae and
M. pseudosasae (Fig. 127) is provided below.
Meliolaceae G.W. Martin ex Hansf.
The family Meliolaceae was introduced by Martin
(1941) without a Latin diagnosis and validated by Hansford
(1946). It is typified by Meliola with M. nidulans (Schwein.) Cooke as the type species. At various times, this
family has been placed in the orders Dothideales, Erysiphales, Meliolales, Myriangiales and Hypocreales (Martin
1941; Luttrell 1951, 1989; Roger 1953; Ainsworth et al.
1971; Müller and von Arx 1973; Yarwood 1973; Barr
1976; Eriksson 1981; Hawksworth et al. 1983). Kirk et al.
(2001) introduced a new subclass, Meliolomycetidae
(Sordariomycetes) for members of Meliolaceae, but without a description or diagnosis. The placement of Meliolomycetidae in Sordariomycetes was confirmed by
Justavino et al. (2015) based on their phylogenetic analyses
and validated in Maharachchikumbura et al. (2015).
Ainsworth et al. (1971) and Eriksson and Hawksworth
(1993) listed 50 genera in the family, but this was reduced
to 25 genera (Hawksworth et al. 1995) and later 22 genera
(Kirk et al. 2008). Lumbsch and Huhndorf (2010) placed
Meliolaceae in the class Sordariomycetes with 26 genera.
Hongsanan et al. (2015b) accepted only seven genera
123
192
Fungal Diversity (2016) 80:1–270
Fig. 138 Maximum parsimony (MP) majority rule consensus tree for
the analyzed Neopestalotiopsis isolates based on a combined dataset
of ITS, LSU, TUB and TEF sequence data. Values above or below the
branches indicate maximum parsimony bootstrap equal or greater
than 60 %. The tree is rooted with Pestalotiopsis trachicarpicola
(OP068). The strain numbers are mentioned after the species names.
The species obtained in this study are shown in blue bold and ex-type
strains are in black bold
Amazonia, Appendiculella, Asteridiella, Cryptomeliola,
Endomeliola, Irenopsis and Meliola in this family.
studies have provided sequence data from Meliola members (Gregory and John 1999, Justavino et al. 2015, Pinho
et al. 2012, 2014, Hongsanan et al. 2015b), and phylogenetic placement of Meliola was clarified. However, there is
still little sequence data for the entire subclass available in
GenBank.
Meliola Fr.
The genus Meliola is typified by M. nidulans (Schwein.)
Cooke. Meliola is the largest genus of Meliolaceae. Several
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Fungal Diversity (2016) 80:1–270
193
Fig. 139 Neopestalotiopsis cocoës (holotype). a Leaf bright disease
on coconut. b, c Conidiomata on host substrate. d Cross section of
conidioma. e, f Conidiogenous cells with developing conidia.
g Immature conidia. h–l Mature conidia. m Colony on PDA. Scale
bars a = 2 mm, b = 500 lm, c = 250 lm, d = 100 lm, e,
f = 20 lm, g–l = 15 lm
Meliola citri-maximae X.Y. Zeng, K.D. Hyde & T.C.
Wen, sp. nov.
Index Fungorum number IF552271; Facesoffungi number: FoF02253, Fig. 128
Etymology: Referring to the host Citrus maxima (Burm.
f.) Merr.
Holotype: MFLU 14-0288.
Epiphytes on the surface of living leaves. Colonies
hypophyllous, scattered to dense, sometimes confluent.
Hyphae superficial, brown, substraight, radiating
outwardly, branched, septate, darker at septa, reticulate,
with hyphal setae. Hyphal setae up to 320 lm long, dark
brown, denticulate at the apex. Hyphopodia 20–22 9 8–
10 lm (
x = 21 9 9 lm, n = 20), 2-celled, brown,
spathulate, form one or two near the septa, alternate to
unilateral, sometimes opposite antrorse. Sexual morph
Perithecia up to 250 lm, superficial, subdense, globose to
subglobose, thick-walled, with ostiole, without perithecial
setae. Peridium two strata, comprising hyaline inner cells
and dark brown outer wall of textura angularis.
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194
Fig. 140 Neopestalotiopsis musae (holotype). a Leaf bright disease
on banana. b, c Conidiomata on host substrate. d Cross section of
conidioma. e Peridium. f Apex. g, h Conidiogenous cells with
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developing conidia. i–j Mature conidia. k Germinating spore.
l Colony on PDA. Scale bars d, e = 50 lm, f–h = 20 lm, i–
j = 15 lm, k = 25 lm
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Fig. 141 Pestalotiopsis sequoiae (MFLU 16-1489, holotype). a Herbarium specimen. b, c, Appearance of black coniodiomata on the
host. d, e Vertical sections of conidiomata. f–h Conidiophores,
195
conidiogenous cells and developing conidia. i–m Conidia. Scale bars
d = 100 lm, e = 50 lm, f–h = 10 lm, i–m = 5 lm
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196
Hamathecium with evanescent paraphyses. Asci unitunicate, 2–3-spored, ovoid to ellipsoid when young, with short
pedicel, asci wall attenuated or broken when mature,
evanescent.
Ascospores
42–48 9 16–21 lm
(
x = 44 9 19 lm, n = 20), 2–3-seriate, cylindrical, hyaline at young state, becoming dark brown when mature, 3–
4-septate, constricted and darker at the septa, middle cell a
little larger, rounded at both ends, smooth-walled. Asexual
morph Phialides 15–22(–25) 9 7–9 lm (
x = 20 9 8 lm,
n = 10), ampulliform, form one at top of hyphal cell,
alternate or unilateral, sometimes mixed with capitate
hyphopodia. Conidia undetermined.
Material examined: THAILAND, Chiang Rai, Horticulture Research Centre, on living leaves of Citrus maxima
(Rutaceae), 12 March. 2014, Xiang-Yu Zeng (MFLU
14-0288, holotype)
Notes: Species of Meliolaceae found on Citrus include
Amazonia butleri, Meliola amyridis, M. camelliae and
M. citricola. Morphologically, the new collection is typical
to the genus Meliola in having hyphal setae, but differs
from M. camelliae and M. citricola as it has 2–3-spored
asci instead of 8-spored asci in M. camelliae and shorter
setae, larger perithecia and larger ascospores than M. citricola. It is most similar to M. amyridis in having
hypophyllous and dense colonies, alternate hyphopodia,
and ascospore size, but differs in having shorter setae,
larger perithecia and hyphopodia. Phylogenetically, our
new collection is related to M. brachyodonta and M. trichostroma. The sequence data of M. brachyodonta and
M. trichostroma, which are unpublished, were provided by
Pinho et al. (2013). However, these two species were found
on Croton curranii (Euphorbiaceae) and Psidium guajava
(Myrtaceae) respectively, which are on the different host
family from this new collection.
Although LSU phylogenies did not fully resolve the
affinities of M. citri-maximae with other known Meliola
species, we suggest it is a new taxon. Further analyses
should target a more variable region such as ITS or btubulin, as the LSU gene is too conserved to differentiate
species (Jeewon et al. 2002, 2003).
Meliola pseudosasae I. Hino, Bull. Faculty of Agriculture,
Yamaguchi University 9: 882 (1958)
Facesoffungi number: FoF02458, Fig. 129
Epiphytes on the upper surface of leaves of Sasa borealis (Hack.) Makino & Shibata. Superficial hyphae branched, septate, darker at the septa, brown, with hyphopodia,
hyphal setae present. Hyphopodia 10–11 lm diam.
(
x = 10.5 lm, n = 20), head cell capitate, 22–25 lm long
(
x = 23 lm, n = 20), alternate or opposite on hyphae,
near to hyphal septum, 2-celled, upper cell cylindrical and
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Fig. 142 Consensus tree resulting from a Bayesian analysis of the c
LSU and ITS sequence of taxa in Sordariomycetes. RAxML bootstrap
support values (MLB above 50) and Bayesian posterior probabilities
(PP above 90 %) are given at the nodes (MLB/PP). The newly
introduced sequences are in blue bold. The scale bar represents the
expected number of changes per site. The tree was rooted to Sordaria
fimicola (HKUCC3414)
curly, brown. Hyphal setae 170–210 lm high 9 9–12 wide
lm (
x = 180 9 10 lm, n = 10), arising from hyphae,
comprising 2–3 arms, with 2–4-branches in each arm, acute
and hyaline at the apex, septate, brown to dark brown,
smooth-walled. Sexual morph Ascomata 97–135 lm
high 9 105–160 lm diam. (
x = 120 9 140 lm, n = 10),
superficial on the surface of hosts, solitary to gregarious on
superficial hyphae, globose to subglobose, thick-walled,
ascomatal setae and appendages absent, surface of ascomata verrucose. Peridium 17–20 lm comprising dark
brown cells of textura angularis when viewed in squash
mounts, with two strata, outer stratum thick-walled, dark
brown cells of irregular textura angularis, and inner stratum of flattened, hyaline cells. Hamathecium not observed.
Asci not observed. Ascospores 45–52 lm high 9 18–23
wide lm (
x = 50 9 22 lm, n = 10), 2-seriate, oblong to
broadly cylindrical, hyaline to dark brown, 4-septate,
constricted and darker at the septa, smooth-walled, ends
rounded. Asexual morph Undetermined.
Material examined: CHINA, Xishuangbanna, on living
leaves of Sasa borealis (Pinaceae), 22 November 2015, R
Phookamsak Xb002 (MFLU 16-2136, in KIB, reference
specimen designated here).
Notes: Our fresh specimen is similar to Meliola pseudosasae both in host (Sasa borealis) and morphology. It
has hyphal setae with 2–3 arms and with 2–4-branches at
the apex, superficial hyphae with hyphopodia and 4-septate, brown ascospores as in the protologue. We were
unable to isolate our specimen in culture, thus, DNA was
extracted from ascomata and ascospores directly. Phylogenetic analysis (Fig. 127) indicate that our collection
clusters with members of Meliolaceae within the subclass
Meliolomycetidae, Sordariomycetes. Meliola pseudosasae
is related to M. clerodendricola Henn. strain, which was
published as a reference specimen by Hongsanan et al.
(2015b). Asteridiella, Appendiculella and Endomeliola
species clustered with Meliola species. Thus, further
molecular data are needed to clarify if they are polyphyletic. We designate our collection as a reference specimen (sensu Ariyawansa et al. 2014b).
Xylariales Nannf.
For the order Xylariales, we follow Daranagama et al.
(2016).
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198
Apiosporaceae K.D. Hyde et al.
The monophyletic family Apiosporaceae was introduced by Hyde et al. (1998), and is typified by the genus
Apiospora. Apiospora typified by A. montagnei Sacc. is
characterized by apiospores and a basauxic, Arthriniumlike conidiogenesis (Samuels et al. 1981; Hyde et al. 1998;
Bahl 2006, Senanayake et al. 2015).
Arthrinium Kunze
Crous and Groenewald (2013) provided a monograph on
this genus and Index Fungorum (2016) listed 60 species.
The older name Arthrinium was conserved as it is a more
commonly encountered name and more frequently used in
literature (Crous and Groenewald 2013). We provide an
updated tree here (Fig. 130).
Arthrinium paraphaeospermum Senan., & K.D. Hyde, sp.
nov.
Index Fungorum number: IF552137, Facesoffungi
number: FoF02459, Fig. 131
Etymology: In reference to the large conidia and conidiogenous cells and morphological similarity to the
A. phaeospermum.
Holotype: MFLU 16-1974
Saprobic on bamboo. Sexual morph Undetermined.
Asexual morph Mycelium consisting of smooth, hyaline,
branched, septate, 3–5 lm diam. hyphae. Conidiophores
reduced to conidiogenous cells. Conidiogenous cells 25–
30 9 4–6 lm (
x = 27 9 4.7 lm), basauxic, aggregated
in clusters on hyphae, hyaline, smooth, elongated, conical
to ampulliform. Conidia 10–19 lm diam., (
x = 13 lm)
brown, smooth, somewhat granular, globose to ellipsoid
in surface view, with a median scar, lenticular in side
view, with pale equatorial slit. Sterile cells 20–30 9 9–
13 lm (
x = 24 9 11 lm), forming on solitary loci on
hyphae, brown, finely roughened, ellipsoid to clavate.
Culture characteristics: Colonies on MEA flat, circular,
smooth, spreading, with fluffy aerial mycelium, mycelia
not tightly attached to the surface, dirty white, with patches
of iron-grey to black due to sporulation.
Material examined: THAILAND, Chiang Mai Prov.,
Doi Inthanon, Hwy 1009 at 25 km marker, N18°32.540 ,
E98°33.510 , alt. 1076 m, on dead clumps of Bambusa sp. L.
(Poaceae), 4 November 2012, I. C. Senanayake, CHUNI
33, (MFLU 16-1974, holotype), ex-type living culture,
MFLUCC 13-0644.
Notes: Arthrinium paraphaeospermum is phylogenetically closely related to A. rasikravindrii and A. hydei
(Fig. 130). However A. paraphaeospermum is phylogenetically distinct with high support (97/0.9, Fig. 130) and
morphologically differs in having ellipsoid to clavate,
brown, sterile tissues, elongated, conical conidiogenous
cells and conidia with a median scar.
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Bartaliniaceae Wijayaw. et al.
The family Bartaliniaceae was introduced by Senanayake et al. (2015) based on analysis of ITS and 28S
rDNA sequence data, and includes Bartalinia, Broomella,
Dyrithiopsis, Hyalotiella, Truncatella and Zetiasplozna.
The polyphyletic genera were placed in this family based
on similarities in morphology and their phylogeny (Senanayake et al. 2015; Maharachchikumbura et al. 2016).
Members of Bartaliniaceae including Truncatella species
produce secondary metabolites with potential use in
biotechnological applications including control of plant
diseases (Flores-Bustamante et al. 2010; Zhao et al.
2015c).
Truncatella was established by Steyaert (1949). The
genus was previously placed in Amphisphaeriaceae, Xylariales, a family including 41 genera, although some
genera are uncertain position (Lumbsch and Huhndorf
2010). The primary reason for introducing a separate genus
is to accommodate a species which is distinct in having
3-septate conidia (Steyaert 1949; Maharachchikumbura
et al. 2012). Molecular data has been used to evaluate
pestalotiod fungi, including species of Truncatella (Jeewon
et al. 2002; Lee et al. 2006; Maharachchikumbura et al.
2012). During a study on the Amphisphaeriales from an
Abies firma leaf in Korea, a new genus that differed morphologically and phylogenetically (Fig. 132) from related
species was isolated and is described as new.
Neotruncatella Hyang B. Lee & T.T.T. Nguyen, gen. nov.
MycoBank number: MB 817422; Facesoffungi number:
FoF02460
Etymology: The genus name ‘‘Neotruncatella’’ refers to
the similarity with Truncatella
Endophyte of Abies firma. Sexual morph Undetermined. Asexual morph Conidiomata globose to subglobose, black. Conidiogenous cells hyaline, ampulliform to
subcylindrical, formed from the inner cells of the peridial
wall. Conidia fusiform with rounded ends, yellow to yellowish-brown at maturity, mostly 3-septate; apical appendages short.
Type species: Neotruncatella endophytica Hyang B.
Lee, P.M. Kirk, K.D. Hyde, S.S.N. Maharachch., & T.T.T.
Nguyen
Neotruncatella endophytica Hyang B. Lee, P.M. Kirk,
K.D. Hyde, S.S.N. Maharachch., & T.T.T. Nguyen, sp.
nov.
MycoBank number: MB 816958; Facesoffungi number:
FoF02478, Fig. 133
Etymology: endophytica, referring to the endophytic
habitat
Holotype: EML-AS5-1
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199
Fig. 143 Anthostomella ravennica (holotype). a, b Appearance of
ascomata on host. c Cross section of ascoma. d Peridium. e Paraphyses. f–h Asci (f note the apical ring bluing in Melzer’s reagent).
i–k Ascospores without visible germ slits. Scale bars a = 200 lm, b,
c = 100 lm, d = 50 lm, f–k = 10 lm
Endophyte of plants. Sexual morph Undetermined.
Asexual morph Conidiomata globose to subglobose,
black, 75–190 9 75–215 lm diam. Conidiogenous cells
hyaline, phialidic, ampulliform to subcylindrical, formed
from the inner cells of the peridial wall. Conidia fusiform
with rounded ends, yellow to yellowish-brown at maturity,
mostly 3-septate, 25–34 9 3–5 lm; apical cell slightly
yellow, thin-walled, bearing a single apical appendage, 3–5
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b Fig. 144 Anthostomella thailandica (holotype). a–c Appearance of
ascomata on host surface. d Ascoma in cross section. e Peridium. f, g
Mature asci. h Apical ring bluing in Melzer’s reagent. i–n Immaturemature ascospores. Scale bars a = 1000 lm, b–c = 500 lm,
d = 100 lm, e–g = 20 lm, h = 5 lm, i–n = 10 lm
(av. 4) lm long; apical appendages short, 4.5–6.5 lm
long; two median cells yellow to yellowish-brown; second
cell from the base 7.5–11.5 lm long, third cell
6.5–10.5 lm long; basal cell conic or with a truncate base,
slightly yellowish-white, 5–7 lm long, basal appendages
present or absent, if appendages present, very shortly at
base, 1–3.5 lm long.
Culture characteristics: Colonies on PDA reaching
82–85 mm diam. at 25 °C after 14 days, initially greenishglaucous, but finally whitish with a white margin; reverse
yellowish with irregular margin; on OA powdery white.
The isolate was observed to grow over a wide range of
temperatures with average growth rates of EML-AS5-1 on
PDA, MEA, and OA of 6.8 mm, 5.8 mm, and 6.5 mm per
24 h, respectively. Optimal growth was observed around
23 °C–27 °C, slow growth was observed at below 15 °C,
and no growth at 35 °C (Fig. 132). Sporulation is excellent
on oatmeal agar and malt extract agar.
Notes: Neotruncatella is morphologically distinct from
Truncatella and Bartalinia by having narrower conidia
with only a shorter apical appendage. In the phylogenetic
trees, the ITS and 28S rDNA sequences (KX216520 and
KX216518, respectively) of the strain formed a separate
branch distinct with them, showing it represents a new
species. In addition, ITS based phylogenies resulted in a
clearly defined monophyletic subclade for our novel
genus basal to Bartalinia (Fig. 132). Morphological
comparison also reveals that Neotruncatella is characterized by a centric basal appendages as compared to
Bartalinia.
Material examined: REPUBLIC OF KOREA, Jeonnam
Province, garden of the Chonnam National University
located in Gwangju (35°100 N 126°550 E), from a Abies
firma leaf, 3 August 2015 (EML-AS5-1, holotype); EMLAS5-1 (ex-type) at Culture Collection of National Institute
of Biological Resources (NIBR), Incheon, Chonnam
National University Fungal Collection (CNUFC-EMLAS5-1) and Jena Microbial Resource Collection (University of Jena and Leibniz Institute for Natural Product
Research and Infection Biology, Jena, Germany)
(JMRC:SF:012333).
Cainiaceae J.C. Krug
The family Cainiaceae was introduced to accommodate
species with a characteristic apical ring comprising a series
of rings (Krug 1978). This family comprises Amphibambusa, Atrotorquata, Arecophila, Cainia, Ceriophora,
201
Reticulosphaeria, Seynesia and Ommatomyces (Kohlmeyer
and Volkmann-Kohlmeyer 1993; Hyde 1996; Senanayake
et al. 2015, Maharachchikumbura et al. 2016). Rhabdospora-like species have been reported as the asexual
morph of this family (Müller and Corbaz 1956).
Cainia Arx & E. Müll.
Cainia is typified by C. graminis (Niessl) Arx & E.
Müll., and comprises eight species (Index Fungorum
2016). Members of Cainia occur on grasses and are characterized by immersed ascomata, cylindrical asci with a
characteristic J? apical ring with series of rings, brown,
two-celled, ascospores with longitudinal striations, and
ascospores surrounded by a mucilaginous sheath (Krug
1978; Senanayake et al. 2015). An updated tree for Cainia
is provided (Fig. 134).
Cainia globosa Senan., Camporesi & K.D. Hyde, sp. nov.
Index Fungorum number: IF552136, Facesoffungi
number: FoF02461, Fig. 135
Etymology: In reference to the globose ascospores.
Holotype: MFLU 14-0738.
Saprobic on standing culms of Anthoxanthum odoratum
L. Sexual morph Ascomata 250–360 lm high 9 210–
315 lm diam. (
x = 326 9 264 lm, n = 10), solitary to
scattered, immersed beneath clypeus, dark brown to
black, globose to subglobose, uniloculate, coriaceous,
with a central ostiole. Peridium 20–25 lm wide, composed of inner, thick-walled, compressed, hyaline cells of
textura angularis and outer, dark brown to black cells of
textura angularis. Hamathecium comprising 3–4 lm
wide, septate, guttulate, sometimes branched, hyaline
paraphyses. Asci 175–190 9 40–50 lm (
x = 188 9
46 lm, n = 20), 8-spored, unitunicate, cylindric-clavate,
straight, short pedicellate, with a layered, apical ring,
bluing lightly in Melzer’s reagent. Ascospores 30–
35 9 14–17 lm (
x = 33 9 16 lm, n = 20), uni-seriate,
ellipsoid to oval, rounded at both ends, hyaline when
immature, becoming pale brown to dark brown when
mature, medianly 1-septate, not constricted at the septum,
guttulate when immature, striate, surrounded by a
mucilaginous sheath. Asexual morph Conidiomata pycnidia, superficial, solitary, formed at the colony margins.
Conidiophores reduced to conidiogenesis cells. Conidiogenous cells 4–7 9 1.5–3 lm, (
x = 5.5 9 2.4 lm),
holoblastic, cylindrical, hyaline, unbranched, septate.
Conidia 7–10 9 3.5–5 lm (
x = 8 9 4.4 lm, n = 20),
globose to oval, 0–1-septate, brown, striate.
Culture characteristics: Colonies on MEA reaching
2 cm after 14 days at 18 °C, circular, dense, white, velvety,
radiating towards the slightly undulate edge, not
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202
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Fig. 145 Phylogram generated from Bayesian analysis based on
combined LSU and ITS sequence data from species of Sordariomycetes. Maximum parsimony/likelihood bootstrap support values
greater than 50 % and Bayesian posterior probabilities greater than 0.90
are shown in above and below. The new isolates are in red and other extype strains are in bold. The tree is rooted with Sordaria fimicola
pigmented, after 4 weeks, small superficial, pycnidia
formed mainly at margin of the colonies.
Material examined: ITALY, Province of Forlı̀-Cesena,
Santa Sofia, near San Paolo in Alps, on dead culms of
Anthoxanthum odoratum (Poaceae), 2 May 2013, E.
Camporesi, IT 1217 (MFLU 14-0738, holotype), ex-type
living culture, MFLUCC 13-0663.
Notes: Cainia globosa is morphologically distinct from
other species in having ascospores with small guttules in
each cell, a very lightly bluing bi-lobed apical ring and
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Fig. 146 Sporidesmium pyriformatum (MFLU 15-1155, holotype).
a Colonies. b–e Fruiting body. f Conidiophores. g, h Conidiogenous
cell. i–n Conidia. o Germinated conidia on PDA medium. p, q
203
Culture, p from above, q from below. Scale bars a = 100 lm, b–f,
o = 30 lm, g, h = 20 lm, i–n = 10 lm
123
204
Fig. 147 Sporidesmium aquaticivaginatum (MFLU 15-1159, holotype). a Colonies on wood. b, c Conidiophores with conidia.
d Conidium developing on the apex of conidiophore. e Conidiophores.
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f–j Conidia. k Germinated conidium on PDA medium. l, m Culture,
l from above, m from below. Scale bars a = 100 lm, b–k = 30 lm
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Fig. 148 Sporidesmium olivaceoconidium (MFLU 15-1175). a Substrate. b Colonies on wood. c, d Conidiophores. e Conidiogenous cell.
f Fruiting body. g–k Conidia. l Germinated conidium on PDA
205
medium. m, n Culture, m from above, n from below. Scale bars
b = 50 lm, c–e, h–k = 15 lm, f = 20 lm, g = 30 lm, l = 40 lm
123
206
Fig. 149 Distoseptispora multiseptata (MFLU 15-1144, holotype).
a Colonies on wood. b–e Conidiophores and conidia. f–j Conidia.
k Conidiophore. l Germinated conidium on PDA medium. m, n
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Culture on MEA medium, m from above, n from below. Scale bars
a = 200 lm, b–g = 50 lm, h, i = 40 lm, j, k = 20 lm,
l = 100 lm
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1-septate, striate conidia. Cainia globosa clustered together
with C. graminis and C. anthoxanthis. However C. globosa
separates from these species with high support (85/
0.9).The asexual morph of Cainia observed from a pure
culture of C. desmazieresii was reported by Müller and
Corbaz (1956) as Rhabdospora-like. However, Krug
(1978) was unable to obtain the asexual morph from
C. desmazieresii in culture (Kang et al. 1999). Here we
observed a coelomycetous asexual morph from the culture
(Fig. 135).
Pestalotiopsidaceae Maharachch. & K.D. Hyde
Pestalotiopsidaceae was introduced by Maharachchikumbura and Hyde (2015), which possesses pestalotiopsis-like
asexual morphs (Senanayake et al. 2015, Maharachchikumbura
et al. 2016). Six genera belong to Pestalotiopsidaceae; Ciliochorella, Monochaetia, Neopestalotiopsis, Pestalotiopsis,
Pseudopestalotiopsis and Seiridium (Senanayake et al. 2015,
Maharachchikumbura et al. 2016). Most taxa in Pestalotiopsidaceae are phytopathogens that cause a variety of diseases in
plants and some are often isolated as saprobes or endophytes and
are widely distributed throughout tropical and temperate regions
(Guba 1961; Barr 1975; Nag Raj 1993; Maharachchikumbura
et al. 2014a, b, 2016).
Ciliochorella Syd.
Ciliochorella was introduced by Sydow & Mitter (1935)
with C. mangiferae Syd. as the type species. Seven species
of Ciliochorella (C. bambusarum Shanor, C. buxifoliae
Allegr., Eliades & Aramb., C. castaneae Munjal, C. eucalypti T.S. Viswan, C. indica Kalani, C. mangiferae Syd.,
C. splendida Nag Raj & R.F. Castañeda) are recorded in
Index Fungorum (2016). Only two strains of Ciliochorella
have sequence data (LSU) in GenBank (Tangthirasunun
et al. 2015). Tangthirasunun et al. (2015) provided
sequence data for the reference specimen of C. mangiferae
found on dead leaves in Thailand.
Ciliochorella phanericola C. Norphanphoun, T.C. Wen &
K.D. Hyde, sp. nov.
Index Fungorum number: IF552264; Facesoffungi
Number: FoF02462, Fig. 137
Etymology: In reference to the host Phanera, and Latin
cola meaning loving.
Holotype: MFLU 14-0776.
Pathogen or saprobic on leaf. Sexual morph Undetermined. Asexual morph Coelomycetous. Conidiomata semiimmersed, circular areas, carbonaceous, black, mostly aggregated, sometimes solitary, in cross section 1000–1200 lm
diam., 170–200 lm high. Peridium comprising a few to several
layers of cell of textura angularis, with inner most layer thin,
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pale-brown, outer layer dark brown to black. Conidiophores
reduced to conidiogenous cells. Conidiogenous cells enteroblastic phialidic, formed from the inner most layer of wall,
hyaline to pale brown, smooth. Conidia (12.5–)13–15 9 2.8–
3.5(–4) lm (
x = 15 9 3.7 lm, n = 30), allantoid to subcylindrical, 2-septate, hyaline to pale brown, smooth-walled;
apical cell with 2 tubular apical appendages, 15–23 lm
(
x = 20 lm), unequal; basal appendage 9–11.5 lm
(
x = 11 lm). Sexual morph Undetermined.
Culture characteristics: Colonies on MEA, reaching
2.5 cm diam. after 7 days at 25 °C, producing dense
mycelium, circular, margin rough, white, without aerial
mycelium.
Material examined: THAILAND, Chiang Rai, Mae Fah
Luang University, on dead leaves of Phanera purpurea (L.)
Benth. (Leguminosae), August 2014, Ausana Mapook
(MFLU 14-0776, holotype; KUN, isotype); ex-type-living
cultures, MFLUCC 14-0984, KUMCC.
Notes: Phylogenetic analyses based on maximum parsimony and Bayesian analyses of LSU, ITS and TUB
sequence data, provide good evidence that our strain
belongs in Ciliochorella. Phylogenetic analysis confirms
C. phanericola is a new species with high support
(Fig. 136). However, C. castaneae differs mainly by 0–2
septate, branched conidiophores occasionally reduced to
conidiogenous cells (Allegrucci et al. 2011). Ciliochorella
phanericola, has conidia of a similar size
(12–17 9 2–4 lm) as C. mangiferae, the latter has fusiform, 3-septate conidia, while those of C. phanericola are
allantoid to subcylindrical, 2-septate with longer basal and
apical appendages.
Neopestalotiopsis Maharachch. et al.
Neopestalotiopsis species are important plant pathogens,
causing post-harvest diseases, fruit rots, and leaf blights in
plants worldwide. Pestalotiopsis was divided into three
genera by Maharachchikumbura et al. (2014b), viz
Neopestalotiopsis, Pestalotiopsis, and Pseudopestalotiopsis with Neopestalotiopsis being distinguished from Pseudopestalotiopsis and Pestalotiopsis by its versicolorous
median cells. Conidiophores in Neopestalotiopsis are
indistinct and often reduced to conidiogenous cells (Xu
et al. 1999; Das et al. 2014, b; Maharachchikumbura et al.
2013, 2014b; Hyde et al. 2014; Jayawardena et al. 2015).
Neopestalotiopsis cocoës Norphanphoun, T.C. Wen &
K.D. Hyde, sp. nov.
Index Fungorum number: IF552232; Facesoffungi
Number: FoF02364, Fig. 139
Etymology: The specific epithet cocoes refers to the
host.
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208
Fungal Diversity (2016) 80:1–270
Fig. 150 Distoseptispora tectonae (holotype). a Colonies on dead branch. b, c Conidia attached to conidiophores on host. d–f Conidia. g,
h Conidia attached to conidiogenous cells and conidiophores. Scale bars d–h = 20 lm
Holotype: MFLU 15-0220.
Pathogenic on Cocos nucifera L. Sexual morph
Undetermined. Asexual morph Conidiomata 170–
123
260 9 140–190 lm (
x = 250 9 180 lm, n = 10), acervuli, with basal stroma and lateral wall of 1–2 layers, thin,
wall cells pale brown, textura angularis. Conidiophores
Ellis (1958), McKenzie (1995)
Dead culms of Pennisetum
purpureum on Freycinetia sp.
Paler, rounded apex
(38–)50–75(–85) 9 11–15
Up to 175 9 6–7
Di. leonensis
7–12-distoseptate
Malaysia
Shoemaker and White (1985),
Réblová (1999)
Decayed wood of Fagus sylvatica
Hyaline, hemisphaerical apex
(80)350–500 9 15–18
28–46 9 8–10
Di. adscendens
80-distoseptate
Hungary
Su et al. (2016a)
On decaying wood
submerged in stream
Paler towards the apex
125–250 9 13–15
21–33 9 5.5–6.5
D. fluminicola
17–34-distoseptate
China
This study
T. grandis
Slightly paler, rounded
to annellidic apex
(83–)148–225(–360)
9 (10–)11–12(–13)
Up to 110 9 5–11
Di. tectonigena
20–46-distoseptate
Thailand
This study
Tectona grandis
Slightly paler, rounded apex
20–28-distoseptate
(90–)130–140(–170)
9 (11–)13–14(–16)
Up to 40 9 4–6
Di. tectonae
Apex of conidia
Conidia (lm)
Conidiophores (lm)
Conidia septation
Thailand
209
Species
Table 2 Synopsis of Distoseptispora tectonae, D. tectonicola, D. adscendens and D. leonensis
Country
Host
Reference
Fungal Diversity (2016) 80:1–270
reduced to conidiogenous cells. Conidiogenous cells
enteroblastic, discrete, simple, short, filiform. Conidia 19–
22.5 9 7.5–9.5 lm (
x = 21 9 9.3 lm, n = 30), fusiform
to ellipsoid, straight to slightly curved, 4-septate; basal cell
conic, hyaline, thin and smooth-walled, 3.4–4.6 lm long
(
x = 4.5 lm); three median cells 13.4–14.5 lm long
(
x = 14 lm), brown to pale brown, versicolored, verruculose; second cell from base pale brown to olivaceous,
3.8–5.5 lm (
x = 5.2 lm); third cell darker brown to olivaceous, 4.2–5.5 lm (
x = 5.5 lm); fourth cell darker
brown, 3.6–4.9 lm (
x = 4–3 lm); apical cell 2.1–3.2 lm
long (
x = 2.3 lm), hyaline, cylindric to subcylindric;
apical appendages 14.9–21 lm long (
x = 19.6 lm), 2–3
(mostly 3), basal appendage present (1 or rarely absent),
filiform 3.2–8.1 lm (
x = 6 lm).
Culture characteristics: Colonies on MEA, reaching
4 cm diam. after 7 days at 25 °C, producing dense mycelium, circular, rough margin white.
Material examined: THAILAND, Chiang Rai, Mushroom Research Foundation, on leaf of Cocos nucifera L.
(Arecaceae), 4 February 2015, K.D. Hyde (MFLU
15-0220, holotype; KUN, isotype); ex-type-living cultures,
MFLUCC 15-0152, KUMCC.
Notes: Neopestalotiopsis cocoës was associated with
leaf blight disease of Cocos nucifera. In the phylogenetic
analyses Neopestalotiopsis egyptiaca Ismail et al., is sister
to our strain (Fig. 138). However, this species differs from
N. cocoës in having larger conidia (22.5–28 lm). The
conidia of N. cocoës are also wider (7.5–9.5 lm) than those
of N. egyptiaca (6–7.5 lm) (Jayawardena et al. 2016)
(Fig. 139).
Neopestalotiopsis musae Norphanphoun, T.C. Wen &
K.D. Hyde, sp. nov.
Index Fungorum number: IF552233, Facesoffungi
Number: FoF02363, Fig. 140
Etymology: Named after the host genus on which the
fungus occurs.
Holotype: MFLU 16-1279.
Pathogenic on Musa sp. Sexual morph Undetermined.
Asexual morph Conidiomata 175–180 9 110–160 lm
(
x = 175 9 140 lm), acervuli, with basal stroma and
lateral wall 1–3 cells thick; the wall cells hyaline, textura
angularis. Conidiophores reduced to conidiogenous cells.
Conidiogenous cells enteroblastic, discrete, simple, short,
filiform. Conidia 18.6–25 9 4.1–5 lm (
x = 20.5 9
4.5 lm), fusiform to ellipsoid, straight to slightly curved,
4-septate; basal cell conic, hyaline, thin and smoothwalled, 3.2–5.2 lm long (
x = 3.5 lm); three median cells
12.1–16 lm long (
x = 15 lm), hyaline, versicolored,
verruculose; second cell from base pale brown to olivaceous, 4.1–6.1 lm (
x = 5 lm); third cell pale brown to
olivaceous, 3.2–6.1 lm (
x = 5.5 lm); fourth cell pale
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210
brown to olivaceous, 3.3–6 lm (
x = 4.5 lm); apical cell
3.6–5.2 lm long (
x = 3.7 lm), hyaline, cylindric to subcylindric;
apical
appendages
16.3–25 lm
long
(
x = 16 lm), 2–3 (mostly 3), basal appendage filiform,
4.6–10.3 lm long (
x = 5 lm).
Culture characteristics: Colonies on PDA, reaching
5 cm diam. after 10 days at 25 °C, producing dense
mycelium, circular, rough margin white, after 2 weeks, flat
or effuse on the surface, without aerial mycelium.
Material examined: THAILAND, Chiang Rai, Pha
Chang, on leaf of Musa sp. (Musaceae), 4 February 2015,
Chada Norphanphoun (MFLU 16-1279, holotype; KUN,
isotype); ex-type-living cultures, MFLUCC 15-0776,
KUMCC.
Notes: Molecular analysis provides good evidence that
our strain belongs to Neopestalotiopsis and is closest to N.
honoluluana Maharachch. et al. and N. zimbabwana
Maharachch. et al. (Fig. 138). However, N. honoluluana
and N. zimbabwana has longer and wider conidia
(
x = 28 9 8.3 lm, = 25.3 9 7.7 lm, respectively) with
N. honoluluana having three dark median cells, unlike N.
musae which has smaller conidia (
x = 20.5 9 4.5 lm),
and pale-brown median cell of conidia (Maharachchikumbura et al. 2014b).
Pestalotiopsis sequoiae W.J. Li, Camporesi & K.D. Hyde,
sp. nov.
Index Fungorum number: IF552215; Facesoffungi
number: FoF02352, Fig. 141
Etymology: Named after the host genus Sequoia.
Holotype: 16-1489.
Saprobic on dead stem of Sequoia sempervirens (Lamb.
ex D. Don) Endl. (Cupressaceae). Sexual morph Undetermined. Asexual morph Coelomycetous. Conidiomata
160–305 lm diam. 9 162–326 lm high, black, solitary to
aggregated or confluent, subperidermal in origin, erumpent
at maturity, pycnidioid, globose to subglobose, unilocular,
glabrous, straw-coloured with dark center; wall 10–20 lm
wide, composed of thin-walled, hyaline cells of textura
angularis to textura globulosa. Conidiophores reduced to
conidiogenous cells. Conidiogenous cells 4–24 lm
long 9 2–4 lm wide, hyaline, enteroblastic, lageniform to
subcylindrical, arising from the cells lining the inner wall
of the pycnidium. Conidia 21–30 9 7.5–10 lm ( =
25 9 8.5 lm, n = 30 lm), fusiform, 4-septate, straight or
slightly curved, bearing tubular, unbranched, flexuous
appendage at both ends, basal cell 2.9–5.7 lm
(
x = 4.3 lm) long, hyaline, obconic with a truncate base, 3
median cells, together 14.7–20 lm long (second cell from
base 4.1–7 (
x = 5.6 lm, third cell 5.4–6.9 lm
(
x = 6.2 lm), fourth cell 4.6–6.7 lm (
x = 5.6 lm),
cylindrical, pale brown to brown and concolourous, septa
and periclinal wall darker, apical cell 2.9–4.8 lm
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(
x = 4 lm) long, hyaline, conical, with mostly 4 apical
appendages, 3–17 lm (
x = 11 lm) long, with independent
loci of origin along upper half of the apical cell, often
forming aggregated crest, basal appendage 4–11 lm
(
x = 6 lm) long, single, unbranched.
Culture characteristics: Colony on PDA reaching
40–50 mm diam. in 7 days, with circular margin, white and
fluffy, dense, aerial mycelium on the surface, reverse
similar in colour.
Material examined: ITALY. Province of Forlı̀-Cesena
[FC]), Castrocaro Terme, on dead stem of Sequoia sempervirens (Cupressaceae), 20 October 2012, Erio Camporesi, IT-832 (MFLU 16-1489, holotype); ex-type living
culture, MFLUCC 13-0399, ICMP; IT-832B (HKAS
95026, isotype); living culture, KUMCC 15-0642.
Notes: Phylogenetically, based on megablast search of
NCBIs GenBank nucleotide database, the closest hit using
ITS sequence is Pestalotiopsis hollandica (GenBank
KM199328; Identities = 548/548(100 %); Gaps = 0/
548(0 %)), Followed by uncultured fungus clone (GenBank = KP889337; Identities = 549/550(99 %); Gaps =
0/550(0 %)), and P. monochaeta (GenBank = KM199327;
Identities = 547/547(100 %); Gaps = 0/547(0 %)). Closest hits using the TEF sequence yielded highest similarity to
P. hollandica (GenBank = KM199481; Identities = 237/
238(99 %); Gaps = 1/238(0 %)), Pestalotiopsis sp. (GenBank = KP781881; Identities = 228/230(99 %); Gaps =
2/230(0 %)), P. verruculosa (GenBank = JX399061;
Identities = 233/238(98 %); Gaps = 1/238(0 %)).
Morphologically, the conidia of Pestalotiopsis sequoiae is
smaller than those of P. hollandica [(25–)25.5–33(–34) 9
8.5–10(–10.5) lm], P. monochaeta [(25–)27–40(–42) 9
7–11(–11.5) lm] and P. verruculosa [(28–35 9 9–11 lm ( =
30:6 9 10:3 lm)]. Based on morphology and phylogeny
(Fig. 138), Pestalotiopsis sequoiae is introduced as novel
species in Pestalotiopsis.
Xylariaceae Tul. & C. Tul.
The family Xylariaceae currently comprises 87 genera
(Maharachchikumbura et al. 2016). Xylariaceous members
are well-known for their secondary metabolite production
(Stadler 2011). Xylariaceae has been traditionally classified into two subfamilies (Chesters and Greenhalgh 1964;
Ju and Rogers 1996; Bitzer et al. 2008). In this study we
introduce new species to genus Anthostomella.
Anthostomella ravennica Daranagama, Camporesi & K.
D. Hyde, sp. nov.
Index Fungorum number: IF552286; Facesoffungi
number: FoF02418, Fig. 143
Etymology: ‘‘ravennica’’ refers to the province from
where it was first collected
Holotype: MFLU 16-0972
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Fig. 151 Distoseptispora tectonigena (holotype). a, b Colonies on
dead twig. c Conidiophores. d, e Conidia attached to conidiophores.
f–i Conidia. j, k Percurrently proliferating apex of conidia.
211
l Germinated conidium. m Colony on PDA after 7 months. Scale
bars a = 200 lm, b = 100 lm, d–h = 20 lm
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212
123
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b Fig. 152 Maximum likelihood phylogenetic tree generated by
RAxML (GTR?G model) based on LSU sequence data. ML/MP
values ([50 %) resulting from 1000 bootstrap replicates are given at
the nodes and branches with Bayesian posterior probabilities greater
than 0.90 are in bold. The original isolate or specimen numbers are
noted after the species names. The tree is rooted to Colletotrichum
asianum. Newly generated sequence is in red and other ex-type
strains are in bold
Saprobic on dead stem of Ammophila arenaria (L.)
Link. Sexual morph Ascomata 170–190 9 180–200 lm
(
x = 175 lm 9 192 lm, n = 10), immersed, visible as
black, irregular to dome-shaped areas, coriaceous, solitary,
scattered, in cross section globose, with inconspicuous
ostiole. Ostiole black, papillate. Peridium 30–50 lm wide
(
x = 35 lm, n = 10), with two cell layers, outwardly
comprising thick-walled, compressed, dark brown cells of
textura angularis and inwardly comprising a few layers
thin-walled, hyaline cells of textura angularis. Paraphyses
3.3–3.8 lm wide at base (
x = 3.5 lm, n = 30), shorter
than the asci, numerous, filamentous, septate. Asci 116–
127 9 11–13.6 lm (
x = 124 9 12.8 lm, n = 20), 8spored, unitunicate, cylindrical, short pedicellate, with
conspicuous, J?, discoid, apical ring, 0.7–1.5 9 0.2–
0.8 lm (
x = 1.2 9 0.5 lm, n = 20). Ascospores 14–
17 9 7–8 lm (
x = 16.2 9 7.6 lm, n = 20), uni-seriate,
inequilaterally ellipsoidal, with one convex surface, pointed ends, light brown-dark brown, smooth-walled, without
germ slit. Asexual morph Undetermined.
Culture characteristics: Colonies on OA 9 cm diam.
after 4 weeks at 27 °C, white at the margins, pale yellowish at the center; reverse yellowish to cream, colony
azonate, cottony appearance, dense.
Material examined: ITALY, Ravenna Province, Lido di
Dante, on dead stems of Ammophila arenaria (L.) Link, 28
January 2015, E. Camporesi, IT 2358 (MFLU 16-0972,
holotype), ibid. HKAS, isotype; ex-type living culture,
MFLUCC 15-0012, KUMCC.
Notes: Anthostomella ravennica is morphologically
reminiscent of A. zongluensis K.D. Hyde. However, the
new species can be distinguished as the latter has larger
ascomata with a central ostiolar canal, a thin peridium,
while A. ravennica has irregular or sometimes domeshaped ascomata, with an inconspicuous ostiole. Ascospores in A. zongluensis have conspicuous mucilaginous
sheaths and asci have a distinct, J?, wedged-shaped apical
ring, while ascospores in A. ravennica lack mucilaginous
sheaths and asci have a J?, discoid apical ring which is
inconspicuous. Due to these morphological differences we
designate our collection as a new species. Molecular data
also support A. ravennica to be phylogenetically distinct
(Figs. 142, 143)
213
Anthostomella thailandica Daranagama & K.D. Hyde, sp.
nov.
Index Fungorum number: IF552287; Facesoffungi
number: FoF02419, Fig. 144
Etymology: ‘‘thailandica’’ refers to the country it was
first collected.
Holotype: MFLU 16-0971
Saprobic on leaves of grasses. Sexual morph Ascomata
150–200 9 140–155 lm (
x = 167 9 148 lm, n = 10),
immersed, visible as black, irregular areas, coriaceous,
solitary, sometimes clustered in groups of two, scattered, in
cross section subglobose. Ostiole inconspicuous, without
conspicuous papilla. Peridium 14–22 lm wide (
x=
17 lm, n = 10), comprising two cell layers, outwardly
comprising thin-walled, loosely arranged, light brown cells
of textura globosa and inwardly comprising a few layers of
thin-walled, hyaline cells of textura angularis. Paraphyses
4.2–6.5 lm wide at base (
x = 4.9 lm, n = 30), as long as
asci, few, filamentous, septate. Asci 87–118 9 14.1–
17.8 lm (
x = 92.6 9 15.2 lm, n = 20), 8-spored, unitunicate, cylindrical-clavate, apedicellate, with conspicuous,
J ?,
discoid
apical
ring,
3.1–3.8 9 1.1–1.9 lm
(
x = 3.5 9 1.5 lm, n = 20). Ascospores 12–17 9 5.5–
9 lm (
x = 15.4 9 7.2 lm, n = 20), uni-seriate, equilateral ellipsoidal, with broad ends, olivaceous, smooth-walled, with one large, central guttule, without germ slit.
Asexual morph Undetermined.
Culture characteristics: Colonies on OA 9 cm diam.
after 4 weeks at 27 °C, white at the margins, pale yellowish at the center; reverse yellowish to cream, colony
azonate, cottony appearance, dense.
Material examined: THAILAND, Chiang Rai Province,
Mae Fah Luang University garden, on dead leaves of a
grass, 12 August 2015, D.A. Daranagama, KM24 (MFLU
16-0971, holotype), ibid. HKAS, isotype; ex-type living
culture, MFLUCC 15-0017, KUMCC.
Notes: Our collection is reminiscent of both
A. zongluensis K.D. Hyde and A. consanguinea (Ces.)
Sacc. However, A. zongluensis has ascospores with more
parallel sides, minutely verruculose walls and ascomata
with periphysate ostiolar canals (Lu and Hyde 2000).
Ascospores of A. thailandica are similar in size to those
of A. consanguinea (Ces.) Sacc. However, ascospores of
A. consanguinea have verruculose walls and a short,
straight germ slit, while those in A. thailandica have a
smooth wall and lack germ slits. In the phylogenetic
analysis A. thailandica clusters in a separate clade with
A. formosa Kirschst., A. conorum (Fuckel) Sacc., A. rubicola Speg. ex Sacc. & Trotter and A. obesa Daranagama, E. Camporesi & K.D. Hyde.
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Fig. 153 Paracapsulospora metroxyli (holotype). a Appearance of
ascomata on host substrate. b Close up of ascomata. c Peridium.
d Section of ascoma. e–j Asci. k–q Ascospores. r Germinated
ascospore. s Culture characters on MEA. Scale bars a = 1 mm,
b = 200 lm, c = 10 lm, d = 100 lm, e = 50 lm, f–j = 20 lm, k–
r = 5 lm
Sordariomycetes incertae sedis
Sporidesmiaceae Fr.
The family Sporidesmiaceae was introduced by Fries (1849),
but the taxon has not been widely used in modern classification.
The family is typified by Sporidesmium Link with S. atrum Link
as the type species. Su et al. (2016a) treated Ellisembia as a
synonym of Sporidesmium based on molecular data and placed
in the presently monotypic Sporidesmiaceae. Based on molecular data, Sporidesmiaceae shows a sister relationship to Papulosaceae and Trichosphaeriaceae (Fig. 145).
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215
Fig. 154 Phylogenetic tree of the Clavulinopsis and Ramariopsis
obtained by analyses from rDNA sequences. Taxa with two accession
numbers were analyzed by concatenated ITS and partial LSU rDNA;
only the partial LSU rDNA was used to analyze the other individuals.
Support values (from top) are maximum likelihood (ML) and
Bayesian analyses. Sequences obtained in this study are in boldface.
Only support values of at least 50 % are shown. The tree was rooted
with Clavaria acuta
Sporidesmium Link
The genus Sporidesmium was typified by S. atrum. Ellis
(1958) considered S. ehrenbergii be the same as the type.
Sporidesmium is characterized by unbranched or sparingly
branched conidiophores; monoblastic or percurrently
proliferating conidiogenous cells with broad scar;
euseptate, rostrate conidia, sometimes with a mucilaginous apex (Shenoy et al. 2006; Seifertk et al. 2011).
Sporidesmium is morphologically similar to the distoseptate genus Ellisembia. However, Su et al. (2016a)
considered the euseptate/distoseptate difference between
Sporidesmium and Ellisembia to have little taxonomic
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Fig. 155 Clavulinopsis aurantiaca (holotype). a Basidiomata. b, c Hyphae with clamps. d Basidia, one clamped at the base. e Basidia with
sterigmata and basidiospores. f Basidiospores with apiculus. Scale bars a = 1 cm, b = 20 lm
significant and that did not appear to be supported by
any molecular data. Sporidesmium is shown to be polyphyletic based on phylogenetic studies (Shenoy et al.
2006; Su et al. 2016a).
Sporidesmium pyriformatum J. Yang & K.D. Hyde, sp.
nov.
Index Fungorum number: IF552227; Facesoffungi
number: FoF02245, Fig. 146
Etymology: Referring to the pyriform conidia.
Holotype: MFLU 15-1155
Saprobic on submerged wood in a stream. Colonies on
substrate superficial, effuse, dark brown, hairy. Mycelium
mostly immersed, composed of branched, septate, hyaline
to pale brown hyphae. Sexual morph Undetermined.
Asexual morph Conidiophores macronematous, mononematous, solitary or caespitose, cylindrical, straight or
slightly flexuous, smooth, 2–5-septate, brown, paler
towards the apex, 65–190 9 3–5.5 lm (
x = 118.5 9
4 lm, n = 20). Conidiogenous cells holoblastic,
123
integrated, lageniform, terminal, brown to olivaceous,
9–24.5 9 4–7 lm, darkened and truncate at the apex,
with up to three lageniform percurrently proliferating.
Conidia acrogenous, solitary, pyriform or obclavate,
3–4-euseptate, the lower 3–4 cells olivaceous to brown,
with pale brown to subhyaline apical cell, darkened at
the septa, 18.5–32.5 9 8–15.5 lm (
x = 25.5 9 12 lm,
n = 20).
Culture characteristics: Conidia germinating on PDA
within 24 h. Germ tubes produced from the apex. Colonies
on MEA reaching 10–15 mm diam. after 2 weeks at 25 °C,
white in the middle, pale yellow at edge, with dense white
mycelium on surface, sparser to the edge; in reverse with a
yellow middle with yellowish irregular margin.
Material examined: THAILAND, Prachuap Khiri Khan
Province, Hua Hin, stream flowing outside Kaeng Krachan
National Park, on submerged wood, 25 December 2014,
Jaap van Strien Site4-26-3 (MFLU 15-1155, holotype); extype living culture, MFLUCC15-0620; ibid. (MFLU
15-1162, paratype).
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217
Fig. 156 Ramariopsis atlantica (holotype). a Basidiomata. b Hyphae with clamps. c Hyphal ends resembling gloeocystidia. d Basidiospores,
one of them germinating. Scale bars a = 1 cm, b = 20 lm
Note: Phylogenetic analysis of combined LSU and ITS
sequence data, supports the placement of our taxon within
Sporidesmium and reveals a close relationship (Fig. 145) to
S. parvum (S. Hughes) M.B. Ellis and S. fluminicola H.Y.
Su & K.D. Hyde. We therefore introduced this as a new
species, Sporidesmium pyriformatum. The conidial shape of
S. pyriformatum resembles those of S. penzigii Cooke &
Ellis, S. jasminicola M.B. Ellis, S. bambusinum N.D.
Sharma, S. fraxini-paxianae Jian Ma & X.G. Zhang,
S. submersum H.Y. Su & K.D. Hyde, S. parvum and S. fluminicola. However, the conidia of the new taxon are larger
than those of S. parvum (18–24 9 8–9 lm), but smaller
than those of the other taxa that listed above. Sporidesmium
pyriformatum is distinguished from the newly introduced
taxa S. submersum and S. fluminicola in having much longer
conidiophores and fewer septa (Su et al. 2016a).
Sporidesmium aquaticivaginatum J. Yang & K.D. Hyde,
sp. nov.
Index Fungorum number: IF552229; Facesoffungi
number: FoF02369, Fig. 147
Etymology: Referring to the aquatic habitat and the
mucilaginous sheath.
Holotype: MFLU 15-1159
Saprobic on decaying plant twigs. Sexual morph
Undetermined. Asexual morph Colonies effuse, olivaceous,
hairy. Mycelium partly immersed, partly superficial on the
substrate. Conidiophores macronematous, mononematous,
solitary or sometimes caespitose, cylindrical, straight or
slightly flexuous, smooth, dark brown, triangular or globose at
the base, 1–3-septate, 60–125 9 4–6 lm (
x = 82.5 9 5 lm,
n = 15). Conidiogenous cells holoblastic, monoblastic, integrated, terminal, determinate or with one percurrent proliferation, cylindrical, smooth, mid to dark brown. Conidia
acrogenous, solitary, dry, olivaceous to pale brown, paler at
the apex, obclavate, tapering gradually towards the apex,
smooth, straight or curved, 6–10-distoseptate, sometimes
with a mucilaginous sheath, 49.5–80.5 9 10.5–14 lm
(
x = 66.5 9 12 lm, n = 20), rounded and 2.3–3.5 lm at the
apex, with a thick, flat, colourless, sheath extending from the
tip to half way down, truncate and 3.5 9 5 lm at the base with
conspicuous darkened scars.
Culture characteristics: Colonies on PDA reaching
10–15 mm diam. after 4 weeks at 25–30 °C, colony from
above, dark greenish in the middle becoming greyish-green at
the umbonate margin; from below, greyish-green in the centre
and the third layer, dark greenish to black at the second layer
and the edge; medium raised to umbonate in the middle.
123
218
Material examined: THAILAND, Prachuap Khiri Khan
Province, Hua Hin, stream flowing outside Kaeng Krachan
National Park, on submerged wood, 25 December 2014,
Jaap van Strien, Site4-44-2 (MFLU 15-1159, holotype);
ex-type living culture, MFLUCC15-0624; ibid. (HKAS950
46, isotype).
Note: The combined LSU and ITS phylogenetic analyses indicate Sporidesmium aquaticivaginatum represents a
sister taxon to S. olivaceoconidium within the
Sporidesmiaceae (Fig. 145). In conidial shape and sheath,
Fig. 157 Phylogram resulting from the RAxML (Stamatakis 2014)
analysis of ITS sequence data. Bootstrap values greater than 50 % are
indicated above branches. The names in blue represent the species of
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Fig. 158 a, f Cortinarius fulvescens 04-935 (neotype, H). b, g c
C. fulvescentoideus 03-1675 (H). c, i C. pseudobulliardioides 11-452
(holotype, H). d, j C. tenuifulvescens 04-572 (holotype, H). e, h
C. nymphatus DBB21430 (UC). Scale bar 10 mm. (a–d Photographs
by K. Liimatainen, e by D. Bojantchev. f–j Drawings by T. Niskanen
and I. Kytövuori.)
S. aquaticivaginatum resembles S. bambusae M.B. Ellis,
S. minigelatinosa Matush., S. novozymium W.P. Wu. and
S. palmicola W.P. Wu. Conidia in these species are characterized by a globose to subclavate, mucilaginous sheath
Cortinarius presented in this paper and the specimens in boldface are
type specimens of the species. The tree is rooted with Obtusi species
Fungal Diversity (2016) 80:1–270
219
123
220
at the apex. The conidiophores of S. aquaticivaginatum
(60–125 9 4–6 lm) are obviously longer than those of
S. bambusae (30–70 9 4–5 lm), S. minigelatinosa
(40–100 9 4–5 lm) and S. palmicola (20–35 9 4–6 lm),
but shorter than S. novozymium (150–200 9 6–8 lm). The
conidia of S. hainanense (40–50 9 10–13 lm) and
S. minigelatinosa (45–52 9 8–12 lm) are smaller than the
conidia
of
S.
aquaticivaginatum
(49.5–80.5 9
10.5–14 lm). Moreover, among all these species, S. hainanense and S. palmicola are described as having euseptate
conidia while in others the conidia are euseptate and distoseptate. The olivaceous conidia with a basal scar of S. aquaticivaginatum is clearly distinguishable from the brown or dark
brown conidia of S. bambusae, S. minigelatinosa,
S. novozymium and S. palmicola.
Sporidesmium olivaceoconidium J. Yang & K.D. Hyde,
sp. nov.
Index Fungorum number: IF552230; Facesoffungi
number: FoF02368, Fig. 148
Etymology: Referring to the olivaceous conidia.
Holotype: MFLU 15-1175.
Saprobic on decaying plant twigs. Sexual morph
Undetermined. Asexual morph Colonies effuse, olivaceous, hairy. Mycelium partly immersed, partly superficial
on the substrate. Conidiohores macronematous, mononematous, solitary or sometimes caespitose, cylindrical,
straight or slightly flexuous, smooth, dark brown, slightly
swollen at the base, 1–3-septate, 25–55 9 3–4 lm
(
x = 40 9 3.5 lm,
n = 20).
Conidiogenous
cells
holoblastic, monoblastic, integrated, terminal, determinate,
cylindrical, lageniform, smooth, mid to dark brown, 11–
15 9 3–5 lm (
x = 13 9 4 lm, n = 20). Conidia acrogenous, solitary, dry, olivaceous to pale brown, paler towards
the tip, obclavate, smooth, straight or slightly curved, 6–
10-septate, sometimes with a rounded mucilaginous sheath
at the tip, 25–50 9 6–10 lm (
x = 40 9 8 lm, n = 20),
rounded at the apex, truncate at the base.
Culture characteristics: Conidia germinating on PDA
within 24 h and germ tube produced from the apex.
Colonies on MEA slow-growing, reaching 5–10 mm diam.
in 14 days, with dense white mycelium in the center,
becoming sparse at the edge, folded in the middle; in
reverse, brown in the middle and paler, smooth in the
margin.
Material examined: THAILAND, Chiang Rai Province,
stream flowing in Tham Luang Nang Non Cave, on submerged wood, 25 November 2014, Jing Yang YJ-14
(MFLU 15-1175, holotype), ex-type living culture,
MFLUCC 15-0380, GZCC 16-0008.
Notes: The placement of Sporidesmium olivaceoconidium is revealed by molecular data within Sporidesmiaceae as
sister taxon to S. aquaticivaginatum (Fig. 145). However,
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Fungal Diversity (2016) 80:1–270
Fig. 159 RPB2-based phylogram generated from maximum likeli- c
hood analysis (RAxML) depicting the placement of Rhodocybe
indica, R. luteobrunnea and R. griseoaurantia within the genus
Rhodocybe. Values at nodes indicate bootstrap support. BS values C50 % are indicated above or below the nodes, new species are
in blue bold. The tree is rooted with Tricholoma flavovirens
(KC816997) and Catathelasma imperiale (KC816994)
the phylogenetic tree contains a single long branch of the
taxa S. olivaceoconidium and S. aquaticivaginatum with all
other branches being short. It requires more collections for
the future study of these two taxa. Sporidesmium olivaceoconidium is morphologically similar to S. aquaticivaginatum. It differs from S. aquaticivaginatum by the shorter
conidiophores and smaller conidia.
Distoseptisporaceae K.D. Hyde & McKenzie
The family was introduced by Su et al. (2016a) and we
follow this here. We introduce a new species of Distoseptispora and provide an updated tree (Fig. 145).
Distoseptispora K.D. Hyde et al.
We follow Su et al. (2016a)
Distoseptispora multiseptata J. Yang & K.D. Hyde, sp.
nov.
Index Fungorum number: IF552206; Facesoffungi
number: FoF02244, Fig. 149
Etymology: Referring to the multi-septate conidia.
Holotype: MFLU 15-1144.
Saprobic on wood submerged in stream. Colonies
effuse, dark olive-green, hairy or velvety. Mycelium mostly
immersed, comprised of branched, septate, smooth, hyaline
to pale brown hyphae. Sexual morph Undetermined.
Asexual morph Conidiophores macronematous, mononematous, solitary, brown, 2–3-septate, straight or slightly
flexuous, erect, slightly tapering distally, truncate at the
apex, olive-green to mid brown, 23–65 9 4.5–8.5 lm
(
x = 41.5 9 6 lm,
n = 20).
Conidiogenous
cells
holoblastic, monoblastic, integrated, terminal, brown,
determinate, cylindrical. Conidia acrogenous, solitary,
obclavate, rostrate, multi-distoseptate, tapering towards the
apex, dark olivaceous green, 95–290 lm (
x = 160 lm,
n = 30) long, 11–20 lm (
x = 16 lm, n = 30) wide at the
broadest part, rounded and 5.5–9 lm (
x = 7.5 lm,
n = 30) wide at the apex, basal cell subcylindrical, truncate and 3–5.5 lm (
x = 4 lm, n = 30) wide at the base.
Conidial secession schizolytic.
Culture characteristics: Conidia germinating on PDA
within 24 h. Germ tubes produced from both ends. Colonies on
MEA reaching 10–15 mm diam. after 2 weeks at 25 °C,
greyish-green on the surface, with dense, fluffy mycelium folded in the middle, dark green in reverse, with smooth margin.
Material examined: THAILAND, Prachuap Khiri Khan
Province, Hua Hin, a stream flowing outside Kaeng
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221
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222
Krachan National Park, on decaying submerged wood, 25
December 2014, Jaap van Strien, Site4-4-2 (MFLU
15-1144, holotype), ex-type culture, MFLUCC 15-0609,
GZCC; ibid. (HKAS 95045, isotype).
Notes: Distoseptispora multiseptata is placed within the
recently established genus Distoseptispora and it is wellsupported by both morphology and molecular data. Distoseptispora multiseptata clusters with D. fluminicola,
D. tectonae and D. tectonigena. Distoseptispora multiseptata however, is distinguished by its dark olivaceous green
conidia, while the conidia are yellowish to dark reddish
brown in D. fluminicola, D. tectonae and D. tectonigena.
The new species also differs from D. fluminicola and
D. tectonae in having larger conidiophores and conidia, but
smaller than those of D. tectonigena (Su et al. 2016a).
Distoseptispora tectonigena is the only species in the genus
having percurrently proliferating conidiogenous cells at the
apex of conidiophores. Distoseptispora multiseptata
resembles D. aquatica in conidia shape and colour, but
differs by its larger conidia (Su et al. 2016a).
Distoseptispora tectonae Doilom & K.D. Hyde, sp. nov.
Index Fungorum number: IF552223; Facesoffungi
number: FoF01877, Fig. 150
Etymology: Name refers to the host genus Tectona on
which the fungus was collected.
Holotype: MFLU 15-3417.
Saprobic on dead twig and branch of Tectona grandis
L.f. Sexual morph Undetermined. Asexual morph Colonies on natural substrate, superficial, numerous, hairy, dark
brown, scattered, single or in groups. Conidiophores up to
40 lm long, 4–6 lm wide, macronematous, mononematous, simple, erect to slightly curved, unbranched, 2–4septate, slightly constricted at the septa, pale brown to dark
brown, cylindrical. Conidiogenous cells 7.5–9.5 9 3.5–
5 lm, monoblastic, integrated, terminal, cylindrical.
Conidia (90–)130–140(–170) 9 (11–)13–14(–16) lm
(
x = 128 9 14 lm, n = 20), 3.5–5.5 lm wide at the protruding truncate base (rostrum), 20–28-distoseptate, flexuous, cylindric-obclavate, elongate, straight or slightly
curved, rounded at the apex, obconically truncate at the
base, verruculose, dark reddish brown, slightly paler
towards the apex, thick-walled, smooth, secession
schizolytic.
Culture characteristics: Conidia germinating on PDA
within 24 h. Germ tubes produced at the apex and septa of
conidia. Colonies on MEA reaching 8–10 mm diam. after
7 days in the dark at 25 °C (
x = 8.9 mm, n = 5), undulate,
irregular shape, flat or effuse at the edge, velvety, brownish-grey (4E2) from above, olive brown (4F3) from below.
Material examined: THAILAND, Utaradit Province, on
dead twig of Tectona grandis (Lamiaceae), 29 December
2012, M. Doilom (MFLU 15-3417, holotype), ex-type
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living culture MFLUCC 12-0291, MKT 030, ICMP
21156; Phrae Province, Song District, on dead branch of
T. grandis, 30 October 2011, M. Doilom, (MFLU
15-3416), living culture MFLUCC 12-0288, MKT 025,
ICMP 21154.
Notes: Distoseptispora tectonae is introduced here as a
novel species based on its morphological and phylogenetic
differences from known Distoseptispora species. Phylogenetically, D. tectonae lies in a subclade together with
D. fluminicola McKenzie et al. 2016 and D. tectonigena sp.
nov. (this study), but in a distinct lineage with support
67 %MPBS, 55 % MLBS and 1.00 PP (Fig. 145). The
conidia of D. tectonae ((90–)130–140(–170) 9 (11–)
13–14(–16)) are shorter and slightly narrower than those of
D. fluminicola (125–250 9 13–15), D. adscendens
((80)350–500 9 15–18), and shorter and wider than
D. tectonigena ((83–)148–225(–360) 9 (10–)11–12(–13)),
but longer than those of D. leonensis ((38–)50–75
(–85) 9 11–15). The conidia of D. tectonae also have
more septa than those of D. leonensis, but fewer than those
of D. adscendens, D. fluminicola and D. tectonigena. In
addition, the conidiophores of D. tectonae are shorter than
those of D. adscendens, D. leonensis and D. tectonigena,
but longer than D. fluminicola (Table 2).
Distoseptispora tectonigena Doilom & K.D. Hyde, sp. nov.
Index Fungorum number: IF552224; Facesoffungi
number: FoF01878, Fig. 151
Etymology: Name refers to the host genus Tectona on
which the fungus was collected,
gena are used for a second species from the same host.
Holotype: MFLU 15-3418.
Saprobic on dead twig of Tectona grandis. Sexual
morph Undetermined. Asexual morph Colonies on natural substrate, superficial, numerous, hairy, dark brown,
scattered, single or in groups. Conidiophores up to 110 lm
long, 5–11 lm wide, macronematous, mononematous,
simple, erect, straight to slightly curved, septate, slightly
constricted at the septa, pale brown to dark brown, 2–6septate, cylindrical. Conidiogenous cells 4–10 9 2–5.5 lm,
monoblastic, integrated, terminal, lageniform or cylindrical,
brown, smooth, with lageniform or doliiform. Conidia (83–)
148–225(–360) (rostrum included) 9 (10–)11–12(–13) lm
thick (
x = 170 9 11 lm, n = 30), 2.5–6 lm wide at the
truncate base (rostrum), older conidia 20–46-distoseptate,
flexuous, cylindric-obclavate, elongated, straight or slightly
curved, verruculose, rounded at apex, sometimes percurrently proliferating 5–10 times at apex, obconically truncate
at base, dark reddish-brown and slightly paler towards the
apex, thick-walled, smooth, secession schizolytic.
Culture characteristics: Conidia germinating on PDA
within 24 h. Germ tubes produced at the apex and other
cells of conidia. Colonies on MEA reaching 6–13 mm
Fungal Diversity (2016) 80:1–270
223
Fig. 160 Rhodocybe indica (CAL 1323, holotype). a Basidiospores. b Basidium. c Pseudocystidia. d Stipitipellis. e Pileipellis with
pileocystidia. f Basidiocarp in the field. Scale bars a–e = 10 lm f = 10 mm (a–e Photo by K.P. Deepna Latha, f Photos by K.N. Anil Raj)
diam. after 7 days in the dark at 25 °C (
x = 9.9 mm,
n = 5), edge entire to undulate, flat or effuse, velvety,
aerial, medium sparse, grey (7F1) from both above and
below.
Material examined: THAILAND, Chiang Rai Province,
Mae Lao District, on dead twig of Tectona grandis
(Lamiaceae), 12 March 2012, M. Doilom (MFLU 15-3418,
holotype), ex-type living cultures MFLUCC 12-0292,
MKT 033, ICMP 21157.
Notes: Although phylogenies reveal a close affinity
between D. fluminicola, D. tectonae and D. tectonigena
(Fig. 145). Distoseptispora tectonigena differs from D. fluminicola and D. tectonae by its conidiophores and conidial
dimensions, conidial septation and apex of conidia as well as
by phylogenetic analyses. Conidiophores of D. tectonigena
are longer than those of D. tectonae and D. fluminicola. Its
conidia are longer and narrower than D. tectonae, shorter and
narrower than D. fluminicola. The conidia have more septa
than those of D. tectonae and D. fluminicola (Table 2). The
conidial apex of D. tectonigena is sometimes rounded like
that of D. tectonae, but sometimes D. tectonigena produces
an annellidic apex (Table 2).
Sordariomycetidae, genera incertae sedis
Paracapsulospora Konta. & K. D. Hyde, gen. nov.
Saprobic on dead Metroxylon sagu (Arecaceae). Sexual
morph Ascomata solitary, immersed, uniloculate, globose to
subglobose, surrounded by a dark brown layer of peridium
cells. Peridium composed of outer layers of black, thickwalled cells of textura angularis inner layers hyaline and
thin-walled. Asci, 8-spored, unitunicate, cylindrical, long
pedicellate. Ascospores uni-seriate, oblong to cylindrical,
unicellular, hyaline, with two guttules, thin-walled, smoothwalled, have appendages with a distinct mucilaginous
sheath. Asexual morph Undetermined.
Note: Paracapsulospora is introduced to accommodate
a single species P. metroxyli which was collected from
Fig. 161 Rhodocybe luteobrunnea (CAL 1322, holotype). Basidiocarps in the field. Scale bar 10 mm (Photo by K.N. Anil Raj)
Metroxylon in Thailand. The genus is morphologically
similar to Capsulospora and Xylochrysis in having cylindrical asci with uni-seriate and oblong to cylindrical,
aseptate ascospores. Paracapsulospora is most similar to
Capsulospora in having immersed ascomata, with uni-seriate, hyaline, oblong to cylindrical, aseptate ascospores.
Paracapsulospora differs from Xylochrysis in forming
immersed ascomata, with hyaline ascospores, while Xylochrysis forms ascostromata erumpent through host tissue
by long neck and forms paraphyses, with pale brown
ascospores However, these two genera can be distinguished
as Paracapsulospora has appendages at both ends of
ascospores and does not produce paraphyses at the centrum, whereas Capsulospora forms paraphyses buts lacks
appendages.
Phylogenetic
analyses
show
that
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Fig. 162 Rhodocybe luteobrunnea (CAL 1322, holotype). a Basidiospores. b Basidium. c Pseudocystidia. d Pileipellis with encrustations.
e Stipitipellis. Scale bars a–e = 10 lm (Photos by K.N. Anil Raj)
Paracapsulospora is close to the orders Magnaporthales
and Amplistromatales but cannot be confidently assigned to
these order due to weak support (Fig. 152). Based on the
limited sequences data in this group, we could not clarify
the natural placement of our genus. Thus, we treat the new
genus in the Sordariomycetes genera incertae sedis.
Type species: Paracapsulospora metroxyli Konta. & K.
D. Hyde
Paracapsulospora metroxyli Konta & K. D. Hyde. sp. nov.
Index Fungorum number: IF552257; Facesoffungi
number: FoF02367, Fig. 153
Etymology: The specific epithet ‘‘metroxyli’’ refers to
host of which the taxon was collected.
Holotype: MFLU 15-0704
Saprobic on dead Metroxylon sagu (Arecaceae). Sexual
morph Ascomata 87–95 lm high 9 158–167 lm wide,
solitary, immersed, uniloculate, globose to subglobose,
surrounded by a dark brown layer of peridium cells.
123
Peridium 9–17 lm wide, composed of outer layers of
black, thick-walled cells of textura angularis, inner layers
hyaline and thin-walled. Asci 93–100 9 3–4 lm
(
x = 97 9 4 lm, n = 10), 8-spored, unitunicate, cylindrical, long pedicellate. Ascospores 9–11 9 1.7–2.6 lm
(
x = 12.2 9 2.2 lm, n = 10), uni-seriate, oblong to
cylindrical, unicellular, hyaline, with two guttules, thinwalled, smooth-walled, with a distinct mucilaginous
sheath. Asexual morph Undetermined.
Culture characteristics: Ascospores germinating after 24 h
on MEA. Colony on MEA reaching 1.5–2.3 cm diam. after
13 weeks at 25 °C. Center of the colony grey, slightly wrinkled, margin uneven; reddish-brown pigment diffusing into
the agar.
Material examined: THAILAND, Krabi Province, on
dead Metroxylon sagu Rottb. (Arecaceae), 8 December
2014, S. Konta (MFLU 15-0704, holotype, HKAS
95031, isotype), ex-type living culture, MFLUCC
15-0250.
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225
Basidiomycota
Agaricales Underw.
Clavariaceae Chevall.
Clavariaceae is a family of the order Agaricales and is
characterised by erect, simple or branched, white, pale or
brightly coloured basidiomata. The hyphal system is monoor dimitic, with inflated hyphae or not, clamped or not. The
cystidia are rare and the basidiospores are variable in
shape, usually globose to ellipsoid, with smooth to ornamented walls, hyaline to pale yellow, and non-amyloid
(Donk 1964). The family has seven genera and about 120
species (Kirk et al. 2008), widely distributed in the world
(Corner 1950, 1970). They are reported as saprobes, ectomycorrhizal, and possibly biotrophs (Birkebak et al. 2013).
Clavulinopsis Overeem
A genus among the clavarioid fungi that comprises 33
species distributed worldwide (Kirk et al. 2008). The species are usually terrestrial, rarely lignicolous, solitary or
gregarious, and characterized by white, yellow, orange or
red, simple or branched basidiomata; hyaline or slightly
yellow, globose or ellipsoid basidiospores, usually smooth,
but echinulate in some species, and tramal hyphae and
basidia with clamp connections (Corner 1950, 1970;
Petersen 1978). However, the circumscriptions of
Clavulinopsis sensu Corner (1950, 1970) and Petersen
(1978) were not supported by Birkebak et al. (2013). The
phylogenetic tree is presented in Fig. 154.
Clavulinopsis aurantiaca Araujo-Neta, Silva & Gibertoni,
sp. nov.
MycoBank MB 816939; Facesoffungi number:
FoF02463, Fig. 155
Etymology: aurantiaca (Latin) = orange-coloured,
referring to the colour of the basidiomata.
Holotype: URM 84216.
Basidiomata simple, 1.5–3 cm, solitary or in small
groups, slender, with sharp tips, acute, robust, orange (48),
with whitish base (78 W) when fresh.
Basidiospores
globose,
rarely
subglobose,
5–6 9 5–6 lm, Q = 1.01, hyaline, guttulate or not, thinwalled, with short apiculus (0.2–0.5 lm), IKI-. Hyphal
system monomitic, tramal hyphae parallel, hyaline, abundantly clamped, 2–3 lm wide, inflated up to 10 lm, also
clamped, subhymenium tightly interwoven. Cystidia of any
kind absent. Basidia elongate-clavate, 35–37 9 5–9 lm,
hyaline, guttulate, with 4 (rarely 5) stout sterigmata,
5–6 lm long, clamped at the base.
Material examined: BRAZIL, Pernambuco: Tamandaré,
Reserva Biologica de Saltinho, August 2011, L.S. AraujoNeta (URM 84216, holotypus; isotype in O); BRAZIL,
Pernambuco, Tamandaré, Reserva Biológica de Saltinho,
February 2011, L.S. Araujo-Neta (URM 84212), July 2011,
Fig. 163 Rhodocybe griseoaurantia (CAL 1324, holotype). A Basidiome in the field. Scale bar 10 mm (Photo by K.N. Anil Raj)
L.S. Araujo-Neta (URM 84212), March 2012, L.S. AraujoNeta (URM 84272), June 2013, L.S. Araujo-Neta & V.R.
Coimbra 44NA (URM 85691), L.S. Araujo-Neta & V.R.
Coimbra 45NA (URM 85692), L.S. Araujo-Neta & V.R.
Coimbra 46NA (URM 85693).
Notes: This species was found as several solitary basidiomata scattered on soil and is characterized by the slender,
flexible, and orange basidiomata when fresh, brittle when
dry, and hyaline, mostly globose basidiospores.
The specimens of C. aurantiaca were placed in a separated, well supported clade (100/1.0) more closely related,
but with little support (63/0.58), to Ramariopsis aurantioolivacea R.H. Petersen, C. fusiformis (Sowerby) Corner,
C. helvola (Pers.) Corner and C. laeticolor (Berk. & M.A.
Curtis) R.H. Petersen (Fig. 154). These species have simple, orange basidiomata, but R. aurantio-olivacea has
narrower,
subglobose
to
ovate
basidiospores
(5.4–6.5 9 3.6–5 lm) and so far is restricted to New
Zealand (Petersen 1988). Clavulinopsis fusiformis,
C. helvola and C. laeticolor have been reported in the
Neotropics (Petersen 1968; Corner 1950, 1970; Furtado
et al. 2016), but C. helvola differs from C. aurantiaca by
the angular basidiospores. Clavulinopsis fusiformis
(Sowerby) Corner has, on average, larger, globose, subglobose to broadly ovate basidiospores [4.8–7.5(–
9.2) 9 4.5–7.2(–9.2) lm], while C. laeticolor (Berk. &
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Fig. 164 Rhodocybe griseoaurantia (CAL 1324, holotype). a Basidiospores. b Basidia. c Cheilocystidia. d Pileipellis. e Stipitipellis. Scale bar
a–e = 10 lm (Photos by K.N. Anil Raj)
M.A. Curtis) R.H. Petersen has broadly ellipsoid to subglobose basidiospores (4.3–7.5 9 3.5–6 lm).
Other species of Clavulinopsis with yellow to orange,
simple basidiomata have been reported from the Neotropics. Clavulinopsis aurantiocinnabarina (Schwein.) Corner,
however, has hyaline to pale yellow, subglobose basidiospores (5–7.1 9 4–7.1 lm), while C. calocera (G.W.
Martin) Corner has longer, cylindrical to suballantoid
basidiospores (8.5–10 9 4–5 lm). Clavulinopsis amoena
(Zoll. & Moritzi) Corner presents pale yellow basidiomata
and subglobose basidiospores (4–7 9 4–6.5 lm) (Petersen
1968; Corner 1950, 1970).
Ramariopsis (Donk) Corner
A genus among the clavarioid fungi that comprises 44
species distributed worldwide. The species have variously
coloured, branched or rarely simple basidiomata; hyaline,
ellipsoid or subglobose, finely verruculose or echinulate
basidiospores, and tramal hyphae and basidia with clamp
connections (Corner 1950, 1970). The species are terrestrial, rarely lignicolous, solitary or gregarious (Corner
1950, 1970; Petersen 1978). Garcı́a-Sandoval et al. (2005)
considered Ramariopsis sensu Corner (1950, 1970) as a
monophyletic group, limiting their species to those with
123
branched basidiomata, echinulate basidiospores and cyanophilous spore ornamentation derived from the tunica.
According to Birkebak et al. (2013), the circumscriptions
of Ramariopsis sensu Corner (1950, 1970) and Petersen
(1978) were not supported. A phylogenetic tree is presented in Fig. 154.
Ramariopsis atlantica Araujo-Neta, Silva & Gibertoni, sp.
nov.
MycoBank
MB816940;
Facesoffungi
number:
FoF02464, Fig. 156
Etymology: referring to the Atlantic Rain Forest.
Holotype: URM 84210.
Basidiomata branched, 5–3.2 cm, with dichotomous,
erect and cylindrical branches, solitary to gregarious, white
(2B) when fresh, beige to straw when dry (52B, 50S).
Basidiospores broadly ellipsoid, 4 9 3 lm, Q = 1.33,
hyaline to pale yellow, guttulate or not, echinulate, spines
0.2–0.5 lm, IKI-. Hyphal system monomitic, tramal
hyphae hyaline to yellow in mass, 2–3 lm, conspicuously
clamped, some inflated up to 10 lm, hardly clamped;
subhymenium loose. Cystidia of any kind absent, but
hyphal ends resembling gloeocystidia occasionally present.
Hymenium collapsed, basidia not seen.
Fungal Diversity (2016) 80:1–270
Material examined: BRAZIL, Pernambuco, Tamandaré,
Reserva Biológica de Saltinho, July 2011, L.S. AraujoNeta (URM 84210 holotype, isotype in O). BRAZIL,
Pernambuco, Tamandaré, Reserva Biologica de Saltinho,
July 2011, L.S. Araujo-Neta (URM 84213). A culture was
obtained from URM 84213 and deposited in the URM
Culture Collection (URM 6985).
Notes: Ramariopsis atlantica was collected as several
basidiomata scattered on soil and is characterized by
branched basidiomata, with dichotomous, cylindrical
branches, inflated hyphae and echinulate, ellipsoid to subglobose basidiospores. Ramariopsis kunzei (Fr.) Corner has
similar basidiomata (branched, whitish), but smaller
basidiospores (3–4.5 9 2.5–3 lm, minutely echinulate,
verruculose or asperulate).
Ramariopsis atlantica was placed in a separated, well
supported subclade (100/1.0), closely related with good
support (81/0.97) to R. aff. kunzei and R. tenuiramosa
Corner (Fig. 154). The new species can be distinguished
from R. tenuiramosa by the larger basidiospores
(3.5–4.5 9 3–3.5 lm in R. tenuiramosa) (Corner
1950, 1970). Additionally, R. atlantica was distantly related to all specimens identified as R. kunzei from the USA
and the UK.
Fig. 165 Phylogenetic tree generated with RAxML (GRT GAMMA)
based on ITS sequence data aligned with MAFFT. Bootstrap values
higher than 50 % are shown. The sequence of the holotype strain is in
bold. The tree is rooted with Nidula niveotomentosa SWFC3000.
Species names are followed by strain numbers. Ex-type strains are
highlighted in bold and new isolates are in blue
227
Cortinariaceae R. Heim ex Pouzar
The limits of the family Cortinariaceae remain
unknown. Currently, this family harbors about 12 genera,
2104 species (Kirk et al. 2008). The majority of the species
in this family are in the genus Cortinarius. Many genera
formerly placed in the Cortinariaceae, e.g., Phaeocollybia,
Hebeloma, and Galerina, and some others have been
moved to other families in Agaricales. On the other hand,
the sequestrate genera, Thaxterogaster, Quadrispora,
Protoglossum and Hymenogaster as well as Cuphocybe,
Rapacea and species of Rozites, once thought to be genera
within the Cortinariaceae, are currently included in the
genus Cortinarius (Peintner et al. 2001, 2002). The
basidiocarps range from agaricoid to sequestrate, and many
have poorly to well-developed veils. The basidiospores are
typically ornamented and cinnamon brown in deposit.
Cortinarius (Pers.) Gray
Cortinarius is the largest genus of Agaricales with a
cosmopolitan distribution and over 2000 described species
(Kirk et al. 2008). The species are important ectomycorrhizal fungi and are associated with trees and shrubs,
belonging to the families Fagaceae, Salicaceae, Caesalpiniaceae, Cistaceae, Dipterocarpaceae, Myrtaceae,
Rhamnaceae, Rosacea and Pinaceae, as well as some
herbaceous plants in the Cyperaceae and Polygonaceae.
Revealing the true diversity of species using only morphological and ecological characteristics has proven to be a
difficult if not an impossible task. The use of DNA
sequence data has made it possible to elucidate phylogenetic relationships within the genus, to show patterns of
speciation, and to help define new, convergent and cryptic
species (Fig. 157).
Cortinarius sect. Fulvescentes Melot and Cortinarius
sect. Laeti Melot were initially included in subgenus Telamonia (Brandrud et al. 1998), but recent phylogenetic
analyses suggest that they form a distinct lineage in genus
Cortinarius (Peintner et al. 2004; Harrower et al. 2011;
Garnica et al. 2016). These two sections were traditionally
distinguished from one another by the colour of the universal veil; the species of the section Fulvescentes have
pinkish to vinaceous universal veil and species of the
section Laeti yellow to ochraceous veil, but sometimes the
veil is very indistinct and difficult to observe. The phylogenetic analysis of Harrower et al. (2011) and Garnica et al.
(2016), however, suggested that the division based on the
veil colour is at least partly unnatural, and it seems a likely
hypothesis based on our analysis too. Additional studies are
needed to reveal the natural divisions within the clade
Fulvescentes/Laeti. Characteristics of the species in these
two sections are mat (dull and flat without a shine),
hygrophanous pileus, silky-shiny fibrillose stipe, uniformly
yellowish-brown context in the stipe, and indistinct smell.
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228
Fungal Diversity (2016) 80:1–270
Fig. 166 Cyathus pyristriatus (holotype). a Basidiomata. b Basidiomata in top view showing peridioles. c Peridioles and plications on
the inner surface of peridium by longitudinal section. d, e
Basidiomata in side view showing fluffy hairs. f Basidiospores.
g Purse hyphae showing clamp-connections. Scale bars a = 3 mm;
b–d = 1 lm; f, g = 10 lm
Below we propose a neotype for C. fulvescens Fr. and
describe four new species belonging to this clade.
hyphae 4–13 lm wide, smooth to encrusted. Hypoderm
distinct, elements up to 30 9 65 lm. Trama hyphae
4–13 lm wide, encrusted. ITS sequence (holotype) distinct
from the other known members of the clade Fulvescentes/
Laeti, and differs from them in the ITS region by more than
14 substitutions and indel positions.
Ecology and distribution: In mesic to dry coniferous
forests with Picea and/or Pinus and Tsuga. Known from
eastern North America and Europe.
Material examined: CANADA, Québec, Montebello,
mixed forest, under Tsuga canadensis, 27 September 2010,
leg. K. Liimatainen & T. Niskanen 10-153, 10-169 (H).
FINLAND, Varsinais-Suomi, Lohja, dry Pinus sylvestris
heath forest on sandy soil with lime dust effect, 19
September 2004, leg. I. Kytövuori & T. Niskanen 04-866/
H6030077 (H). Vihti, Sipilänmäki, Lintumäki, mesic
Picea-dominated forest, 7 October 2001, leg. H. Tuovila &
I. Kytövuori (H). Uusimaa, Kirkkonummi, submesic Picea
abies dominated forest with some Betula and Populus
tremula, in places herb-rich in others more Pinus dominated, 23 September 2004, leg. T. Niskanen 04-935,
H6031281 (neotype, H; isoneotype, K). Uusimaa, Sipoo,
Paippinen, dryish to mesic, mossy coniferous forest (Picea
abies, Pinus sylvestris) with some Betula, 14 September
Cortinarius fulvescens Fr., Epicr. syst. mycol. (Upsaliae):
311 (1838)
Facesoffungi number: FoF02465, Fig. 158
Neotype: T. Niskanen 04-935 (H), designated here.
Pileus 25–50 mm diam., conical to hemisphaerical, later
low convex with an umbo, narrowly pellucid-striate, surface mat, red brown to vinaceous red brown, hygrophanous. Lamellae adnexed, subdistant, moderately broad,
moderately thick, at first light medium brown, becoming
cinnamon brown, edges remaining pale for some time, then
concolorous. Stipe 50–120 mm long, apex 3–9 mm thick,
equal, whitish silky-fibrillose. Universal veil pale pink,
very sparse, forming incomplete girdles on the stipe. Basal
mycelium white. Context in pileus dark red brown, in stipe
medium yellow brown. Odor indistinct. Basidia 4-spored,
8–9 9 27–34 lm, clavate. Basidiospores 7.9–9.5 9
4.5–5.2 lm, Q = 1.64–1.90, narrowly amygdaloid to narrowly amygdaloid-ellipsoid, moderately, often somewhat
sharply verrucose, slightly to moderately dextrinoid.
Lamella trama hyphae moderately to fairly strongly
encrusted in MLZ. Pileipellis duplex (in MLZ): Epicutis
123
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229
Fig. 167 Phylogeny of Polyporus and related genera generated by
maximum likelihood based on combined ITS ? LSU sequence data.
Branches are labeled with maximum likelihood bootstrap proportions
(the former values) higher than 50 %, maximum parsimony bootstrap
proportions (the middle values) higher than 50 % and Bayesian
posterior probabilities (the latter values) more than 0.95
2004, leg. K. Liimatainen & T. Niskanen 04-792,
H6029527 (H). Etelä-Häme, Orivesi, Päilahti, Ojanperä,
mesic Picea abies forest with some Pinus sylvestris and
deciduous bushes, 26 September 1995, leg. I. Kytövuori
95-1547 (H).
Notes: Cortinarius fulvescens is a medium-sized species
and a typical member of traditional section Fulvescentes
with mat, red-brown to vinaceous red-brown pileus, whitish silky-fibrillose stipe, pale pink universal veil, and uniformly yellow brown context in the stipe. It can be
123
230
Fig. 168 Basidiome of Polyporus mangshanensis (holotype). Scale
bar 1 cm
distinguished from most other species with pinkish to
vinaceous veil by somewhat robuster appearance and the
narrowly amygdaloid, somewhat more strongly verrusoce
spores. The species was described by Fries (1838) and the
short description fits well to our species, except the pale
pinkish universal veil is not mentioned. This characteristics, however, is easy to miss since the veil is very sparse
and sometimes invisible as e.g. in the basidiomata in the
middle in Fig. 158. Fries also has two unpublished plates of
C. fulvescens (S0313 and S0314, available at the Krypto-S
database in the Swedish Museum of Natural History, http://
andor.nrm.se/fmi/xsl/kryptos/kbo/publFindspecies.xsl?-view
&-db=kbo_svampregister&-token.languagecode=en-GB),
painted after the description of the species. They both
represent typical species of traditional section Fulvescentes: the stature is slender, the colour of the pileus is
brown to reddish-brown, flesh is brownish-yellow and
they do not have any strong veil bands in the stipe.
However, the two plates may represent two different species. The basidiomata presented in S0313 illustrate a species common to coniferous heath forests of
Scandinavia. It also fits with the current interpretation
of the species (Niskanen and Kytövuori 2012). We
therefore propose specimen 04-935 as a neotype of the
species.
Cortinarius fulvescentoideus Kytöv., Niskanen & Liimat.,
sp. nov.
Index Fungorum number: IF552368; Facesoffungi
number: FoF02466, Fig. 158
Etymology: The name refers to affinity to C. fulvescens.
Holotype: K. Liimatainen & T. Niskanen 03-1634 (H).
Pileus 10–25 mm diam., conical to somewhat hemisphaerical, umbonate, later low convex to plane with an
umbo or not, up to 1/2 pellucid-striate, surface mat, warm
red-brown, hygrophanous. Lamellae adnexed, subdistant,
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Fungal Diversity (2016) 80:1–270
moderately broad to broad, moderately thick, at first light
medium brown, becoming rich brown, edges remaining
pale for some time, then concolorous. Stipe 40–100 mm
long, apex 3–5 mm thick, equal, whitish silky-fibrillose.
Universal veil pale pink, very sparse, forming incomplete
girdles on the stipe. Basal mycelium white. Context in
pileus dark red-brown, in stipe medium yellow-brown.
Odour indistinct. Basidia 4-spored, 8–9 9 30–35 lm,
clavate. Basidiospores 8.2–9.5 9 (4.5–)5–5.4 lm, Q =
1.64–1.76, ellipsoid to narrowly ellipsoid, finely to moderately verrucose, slightly to somewhat moderately dextrinoid. Lamella trama hyphae moderately to strongly
encrusted in MLZ. Pileipellis duplex (in MLZ): Epicutis
hyphae 5–12 lm wide, smooth to encrusted. Hypoderm
distinct, elements up to 28 9 60 lm. Trama hyphae
5–13 lm wide, encrusted. ITS sequence (holotype) differs
from the sister species C. fulvescens and C. tenuifulvescens
more than 7 substitutions and indel positions.
Ecology and distribution: In mesic to moist coniferous
forests with Picea and Tsuga, often in Sphagnum. Widely
distributed and found from Japan, western North America
and Europe.
Material examined: FINLAND, Etelä-Häme, Orivesi,
Päilahti, mesic to moist Picea abies dominated forest with
some Pinus sylvestris and hardwood bushes, 26 September
1995, leg. I. Kytövuori 95-1550 (H). SLOVAKIA, Liptovská kotlina basin, Važec, Važecké lúky, Picea abies
forest, 29 September 2003, leg. K. Liimatainen & T.
Niskanen 03-1634, H7018165 (holotype, H; isotype, K),
30 September 2003 leg. K. Liimatainen & T. Niskanen
03-1675, H7018157 (H).
Additional specimens: CANADA, British Columbia,
cloneSWUBC478, root tip, GenBank no. DQ481834.
JAPAN, Yamanashi, Mt. Fuji, Tsuga diversifolia, June
2011, spYM454 root tip, GenBank no. AB848431. U.S.A.,
Alaska, Bonanza Creek LTER, Site TKN0109, Picea
mariana, 2004, clone3199N24,
Notes: Cortinarius fulvescentoideus looks like a slender
C. fulvescens and forms a well-supported clade with
C. fulvescens and C. tenuifulvescens but strains from two
species are partitioned in two highly supported monophyletic subclades that support their delineation (Fig. 157).
It is morphologically very similar to C. tenuifulvescens and
the two species can easily be misidentified. Our data also
indicates that they may have somewhat different ecologies;
the collections of C. fulvescentoideus are made from mesic
to moist coniferous forests, whereas C. tenuifulvescens
prefers sandy, well-drained soils, at least in northern Europe. From the other members of the section Fulvescentes
C. fulvescentoideus can most easily be distinguished by the
spores. The spores of C. fulvescens are narrowly amygdaloid whereas the spores of C. bulliardioides, C. pseudobulliardioides and C. subfloccopus are broader, [5.5 lm
Fungal Diversity (2016) 80:1–270
Fig. 169 Polyporus mangshanensis (holotype). a Basidiospores.
b Basidia and basidioles. c Hyphae from context. c1 Generative
hyphae, c2 Skeleto-binding hyphae. d Hyphae from cuticle. d1
Contextual cuticle hyphae. d2 Stipe cuticle hyphae. e Hyphae from
231
stipe. e1 Generative hyphae. e2 Skeleto-binding hyphae. f Hyphae
from trama. f1 Generative hyphae. f2 Skeleto-binding hyphae. Scale
bars a–f = 10 lm
123
232
wide. The spores of C. badiovinaceus are small, ovoidsubglobose.
Cortinarius nymphatus Kytöv., Niskanen, Liimat. &
Bojantchev, sp. nov.
Index Fungorum number: IF551705; Facesoffungi
number: FoF02467, Fig. 158
Etymology: Name based on a Greek word nymphe,
meaning fairy, since the species is small and elegant
Holotype: I. Kytövuori 95-1549 (H)
Pileus 10–30 mm diam., at first conical to hemisphaerical, later low convex to almost plane with an umbo, surface mat, brown to dark red-brown, paler towards the
margin, hygrophanous. Lamellae adnexed, subdistant,
moderately broad, moderately thick, at first light brown,
becoming medium brown, edges remaining pale for some
time, then concolorous. Stipe 40–60 mm long, apex
2–4 mm thick, equal, whitish silky-fibrillose. Universal
veil yellow to ochraceous, forming incomplete girdles on
the stipe. Basal mycelium white. Context in pileus dark redbrown, in stipe medium yellow-brown. Odour indistinct.
Basidia 4-spored, 6.5–8 9 25–30 lm, clavate. Basidiospores 6.8–8.2 9 4.3–4.8 lm, Q = 1.56–1.76, amygdaloid, finely to moderately, sharply verrucose, slightly to
moderately dextrinoid. Lamella trama hyphae moderately
encrusted in MLZ. Pileipellis duplex (in MLZ): Epicutis
hyphae 4.5–10 lm wide, encrusted. Hypoderm distinct,
elements up to 30 9 60 lm. Trama hyphae 5–13 lm wide,
somewhat encrusted. ITS sequence (holotype) distinct from
the other known members of the clade Fulvescentes/Laeti,
and differs from them in the ITS region by more than 7
substitutions and indel positions.
Ecology and distribution: In dry to mesic coniferous
forests (Pinus, Picea). Known from Europe, Canada, British Columbia.
Material examined: CANADA, British Columbia, near
Squamish, temperate coniferous rainforest (Picea sitchensis, Pseudotsuga menziesii), 19 October 2009, leg. D. Bojantchev DBB21430 (UC). FINLAND, Etelä-Häme,
Orivesi, Päilahti, mesic to moist Picea abies forest with
some Pinus sylvestris and hardwood bushes, 26 September
1995, leg. I. Kytövuori 95-1549 (holotype, H; isotype, K).
Kainuu, Suomussalmi, W of Hossa, dry Pinus sylvestris
heath forest on sandy soil, 16 September 2002, leg.
I. Kytövuori, K. Liimatainen & T. Niskanen 02-565 (H).
Notes: Cortinarius nymphatus looks like a small Fulvescentes species, but with a yellow-ochraceous veil. Also
based on our phylogenetic analysis it seems to belong to
section Laeti s. str. with the yellow-veiled type species,
C. laetus, of section Laeti. With the combination of small
fruiting bodies and small spores it can be distinguished
from the other yellow veiled species of the clade Fulvescentes/Laeti.
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Cortinarius pseudobulliardioides Kytöv., Niskanen, Liimat. & Ammirati, sp. nov.
Index Fungorum number: IF551704; Facesoffungi
number: FoF02468, Fig. 158
Etymology: The name refers to affinity and similar
appearance to C. bulliardioides.
Holotype: J.F. Ammirati & K. Liimatainen 11-452 (H).
Pileus 25–45 mm diam., at first conical, later low conical to almost plane with an umbo surface mat, dark reddish-brown, somewhat paler towards the margin,
hygrophanous. Lamellae adnexed-emarginate, subdistant,
moderately broad, moderately thick, at first light medium
brown, becoming medium brown, edges remaining pale for
some time, then concolor. Stipe 40–90 mm long, apex
4–7 mm thick, equal, whitish silky-fibrillose. Universal
veil pink, forming broad, incomplete girdles on the stipe.
Basal mycelium white. Context in pileus dark red brown, in
stipe medium to light yellow brown. Odor indistinct. Basidia 4-spored, 9–11 9 35–40 lm, clavate. Basidiospores
8.4–10 9 5.5–6.6 lm, Q = 1.38–1.54, ellipsoid, finely
verrucose, moderately dextrinoid. Lamella trama hyphae
moderately to strongly encrusted in MLZ. Pileipellis
duplex (in MLZ): Epicutis hyphae 6–14 lm wide, encrusted. Hypoderm distinct, elements up to 25 9 40 lm.
Trama hyphae 5–12 lm wide, encrusted. ITS sequence
(holotype) distinct from the other known members of the
clade Fulvescentes/Laeti, and differs from them in the ITS
region by more than 7 substitutions and indel positions.
Ecology and distribution: In coniferous forests (Abies,
Picea, Pinus, Tsuga). Widely distributed and known from
western North America and Europe.
Material examined: FINLAND, Koillismaa, Kuusamo,
Oulanka National Park, forests close to Kiutaköngäs, dry
Pinus sylvestris heath forest with some Picea and Betula on
sandy and calcareous soil, 18 September 2002, leg.
I. Kytövuori, K. Liimatainen & T. Niskanen 02-689 (H).
U.S.A, Washington, Kittitas County, Table Mountain,
coniferous forest (Abies, Picea, Pinus, Larix), 9 October
2011, leg. J.F. Ammirati & K. Liimatainen 11-452 (holotype, H; isotype, K).
Additional specimens: CANADA, British Columbia,
Tsuga heterophylla, cloneSLUBC12.
Notes: Cortinarius pseudobulliardioides is a rather large
species in clade Fulvescentes/Laeti, in appearance similar
to sister species C. subfloccopus and somewhat more distantly related C. bulliardioides. Cortinarius subfloccopus,
however, usually has vinaceous red universal veil and
C. bulliardioides has broader spores, [6 lm wide.
Cortinarius tenuifulvescens Kytöv., Niskanen & Liimat.,
sp. nov.
Index Fungorum number: IF551703; Facesoffungi
number: FoF02469, Fig. 158
Fungal Diversity (2016) 80:1–270
233
Fig. 170 Phylogram generated
from maximum likelihood
method based on ITS-rDNA
sequences from MEGA6 under
Kimura 2-parameter model
(Kimura 1980). The tree with
the highest log likelihood
(-1342.1525) is shown. Onethousand bootstrap replicates
were analyzed to obtain the
nodal support values. The novel
species having GenBank
Accession Number KX234820
(ITS-rDNA) is shown in blue.
The R. brevipes, R. chloroides,
R. emetica are considered as the
out groups
Etymology: The species looks like a slender
C. fulvescens.
Holotypus: K. Liimatainen & T. Niskanen 04-572,
H6031278 (H)
Pileus 10–30 mm diam., conical to somewhat hemisphaerical, umbonate, later low convex to plane with an umbo
or not, up to 1/3 pellucid-striate, surface mat, warm redbrown, hygrophanous. Lamellae adnexed, subdistant, moderately broad, moderately thick, at first light brown, becoming
medium brown, edges remaining pale for some time, then
concolorous. Stipe 40–110 mm long, apex 2–4 mm thick,
equal, whitish silky-fibrillose. Universal veil pale pink, very
sparse, forming incomplete girdles on the stipe. Basal mycelium white. Context in pileus dark red-brown, in stipe medium
yellow-brown. Odour indistinct. Basidia 4-spored, 8–9 9
27–35 lm, clavate. Basidiospores 7.5–9.3 9 4.8–5.4 lm,
Q = 1.48–1.70, ellipsoid to narrowly ellipsoid, finely to fairly
finely, densely, evenly verrucose, slightly to somewhat
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Fig. 171 Russula indoalba (holotype). a–c Fresh basidiomata in the
field and base camp. d Transverse section through lamellae showing
pleuromacrocystidia. e–g Radial section through pileipellis showing
elements in chains. h SEM of basidiospores. Scale bars a = 100 mm,
d, e = 50 lm, f, g = 10 lm, h = 2 lm
moderately dextrinoid. Lamella trama hyphae moderately to
strongly encrusted in MLZ. Pileipellis duplex (in MLZ):
Epicutis hyphae 4.5–13 lm wide, smooth to encrusted. Hypoderm distinct, elements up to 30 9 70 lm. Trama hyphae
5–13 lm wide, encrusted. ITS sequence (holotype) distinct
from the other known members of the clade Fulvescentes/
Laeti, and differs from them in the ITS region by more than 7
substitutions and indel positions.
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235
Fig. 172 Russula indoalba
(holotype). a Fresh and/or
dissected basidiomata. b Radial
section through pileipellis.
c Basidiospores.
d Pleurocystidia.
e Cheilocystidia. f Basidia.
Scale bars a = 10 mm; b–
f = 10 lm
Ecology and distribution: In dry to mesic coniferous
forests, in northern Europe often on sandy soil, one collection from spruce-hardwood swamp. Widely distributed
and found from western and eastern North America and
Europe.
Material examined: CANADA, Newfoundland, Avalon
Peninsula, Butter Pot Provincial Park, mesic to damp Picea
dominated forest with some Abies, Larix and Betula, 25
September 2007, leg. K. Liimatainen & T. Niskanen
07-292, H7000956 (H). FINLAND, Perä-Pohjanmaa,
Rovaniemi, Pisavaara nature reserve area, 31 August 2004,
leg. K. Liimatainen & T. Niskanen 04-572, H6031278
(holotype, H; isotype, K). SWEDEN, Jämtland, Berg, dry
Pinus sylvestris heath forest with some Picea and Betula on
sandy soil, leg. I. Kytövuori, K. Liimatainen & T. Niskanen
03-907, H7018168 (H). Ångermanland, Strömsund,
Bodum, at a small sand pit, Picea-hardwood swamp, leg.
I. Kytövuori 97-500b (H).
Additional specimens: CANADA, British Columbia, Mt.
Washington, Trail to Rossiter Lake, in mossy detritus, 13
Sep 2001, leg. O. Ceska OC43, F17115 (UBC)
Notes: See discussion under C. fulvescentoideus.
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Fig. 173 Phylogram generated from maximum likelihood method
based on ITS-rDNA sequences: The evolutionary history was inferred
by using the maximum Likelihood method based on the Kimura
2-parameter model (Kimura 1980). The tree with the highest log
likelihood (-2215.6038) is shown. One-thousand bootstrap replicates
were analyzed to obtain the nodal support values. The novel species
having GenBank Accession Number KX234819 (ITS-r-DNA) is
shown in blue. The R. emetica and R. nana were considered as the out
group taxa. Evolutionary analyses were conducted in MEGA6
(Tamura et al. 2013)
Entolomataceae Kotl. & Pouzar
Entolomataceae was introduced by Kotlaba and Pouzar
(1972) and typified by Entoloma, is highly variable in
terms of sporocarp morphology and micromorphology
(Noordeloos 2004). To date, this family harbors four genera, 1071 species (Kirk et al. 2008).
namely Rhodocybe, Clitopilopsis, Rhodophana and Clitocella. The amended Rhodocybe now incorporates only
clampless species (Kluting et al. 2014). In the course of our
studies on the Entolomataceae of Kerala State, India, we
came across three species of Rhodocybe, which were found
to be new to science. They are described here based on
both morphology and molecular phylogeny. Maximum
likelihood analysis placed the three new species in Rhodocybe clade with significant (76 %) ML bootstrap support
(Fig. 159)
Rhodocybe Maire
The genus Rhodocybe is characterised by pink, vinaceous white or grey spore print and inamyloid, cyanophilic
basidiospores that are angular in polar view and with an
undulate-pustulate ornamentation (Baroni 1981; Singer
1986). Based on a molecular phylogenetic study, Kluting
et al. (2014) split Rhodocybe sensu lato into four genera,
123
Rhodocybe indica K.N.A. Raj & Manim., sp. nov.
MycoBank MB 816841; Facesoffungi number:
FoF02179, Fig. 160
Fungal Diversity (2016) 80:1–270
237
Fig. 174 Russula pseudoamoenicolor (holotype). a, b Fresh basidiomata. c–e Transverse section through lamellae showing pleuromacrocystidia. f Radial section through pileipellis. g–j Elements of
pileipellis. k Basidiospores. l SEM micrograph of basidiospores.
Scale bars a = 100 mm; c, d = 50 lm; e, g–k = 10 lm;
f = 100 lm; l = 2 lm
Etymology: The specific epithet refers to India, the
country where this species was first observed.
Holotype: CAL 1323.
Basidiocarp small, mycenoid. Pileus 15 mm diam.,
broadly convex with a small umbo; surface brown (6E7/
OAC599) on and around the umbo, brownish-yellow
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238
Fungal Diversity (2016) 80:1–270
Fig. 175 Russula
pseudoamoenicolor (holotype).
a Fresh and/or dissected
basidiomata. b Radial section
through pileipellis.
c Basidiospores.
d Pleurocystidia. e Basidia.
f Cheilocystidia. Scale bars
a = 10 mm; b–f = 10 lm
(5C7/OAC790) towards the margin, weakly hygrophanous
and becoming slightly paler, faintly pellucid-striate
towards the margin, glabrous, somewhat tacky; margin
slightly incurved and somewhat wavy. Lamellae narrowly
adnate to adnate, subventricose, close, greyish-orange
(6B3/OAC695), up to 3 mm wide, with lamellulae of 3–7
lengths; edge crenulate, concolourous with the sides. Stipe
30 9 2 mm, central, equal, slightly flexuous, cartilaginous, solid; surface brownish-yellow (5C7/OAC790),
glabrous to the naked eye, finely pruinose all over under a
123
lens; base with white mycelial cords. Odour and taste not
distinctive.
Basidiospores 6.5–8 9 5.5–7 (7.27 ± 0.49 9 6.1 ±
0.38) lm, (Q = 1–1.3, Qm = 1.19), subglobose or lacrymoid, undulate-pustulate, with or without a suprahilar
depression in profile view, 6 angled in polar view, hyaline,
thin-walled. Basidia 18–33 9 7–8 lm, narrowly clavate to
clavate, pale yellow, thin-walled, 4-spored; sterigmata up
to 5 lm long. Lamella-edge heterogeneous. Cheilocystidia and pleurocystidia present as pseudocystidia.
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239
Pseudocystidia 24–40 9 4–9 lm, scattered, fusiform,
lanceolate or ventricose-rostrate, with glittering, yellow,
granular contents, thin-walled. Lamellar trama subregular;
hyphae 4–7 lm wide, hyaline or pale yellow, thin-walled.
Subhymenium inconspicuous. Pileus trama subregular;
hyphae 5–14 lm wide, pale yellow, thin-walled. Pileipellis
an undifferentiated cutis, made up of closely septate and
compactly arranged hyphae with widely scattered pileocystidia; hyphae 5–13 lm wide, slightly gelatinised, with a
pale brownish-yellow wall pigment and fine hyaline
encrustations, thin- to slightly thick-walled. Pileocystidia
21–42 9 3–11 lm, scattered, versiform: nettle hairshaped, narrowly fusiform or flexuous, hyaline or pale
yellow, thin-walled. Stipitipellis a cutis, rarely disrupted by
flaring out hyphae; hyphae 3–9 lm wide, with a pale
brownish-yellow wall pigment, thin-walled. Caulocystidia
absent. Clamp connections not observed on any hyphae.
Habitat: on a decaying twig, solitary.
Specimens examined: INDIA, Kerala, Kollam District,
Thenmala, Thenmala Forest, 17 August 2013, K.
P. Deepna Latha DKP130 (CAL 1323, holotype).
Notes: Rhodocybe indica is well-characterized by its
small basidiocarps with a brownish-yellow, umbonate
pileus; subglobose or lacrymoid basidiospores; smaller,
fusiform, lanceolate or ventricose-rostrate pseudocystidia;
a pileipellis with pileocystidia and a stipitipellis lacking
encrusted hyphae. Characters such as the centrally stipitate
basidiocarps and the presence of pseudocystidia indicate
Fig. 176 Phylogram generated from maximum likelihood method
based on ITS-rDNA sequences: the evolutionary history was inferred
by using the maximum likelihood method based on the Kimura
2-parameter model (Kimura 1980). The tree with the highest log
likelihood (-1884.3936) is shown. One-thousand bootstrap replicates
were analysed to obtain the nodal support values. The novel species
having GenBank Accession Number KX254611 (ITS-r-DNA) is
shown in blue. The Lactifluus vellereus was considered as the out
group. Evolutionary analyses were conducted in MEGA6 (Tamura
et al. 2013)
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240
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Fig. 177 Lactarius dirkii (holotype). a Fresh basidiomata. b White
latex exuding from cut lamellae. c Transverse section through
lamellae edge showing cheiloleptocystidia. d, e Radial section
through pileipellis showing pileopseudocystidia (arrows). f Radial
section through pileipellis. g Terminal elements of pileipellis. h, i
SEM micrograph of basidiospores. Scale bars a = 100 mm; c,
f = 50 lm; d, e, g = 10 lm; h = 20 lm; i = 2 lm
the section Rhodocybe (Baroni 1981). Rhodocybe pruinosistipitata T.J. Baroni et al., a species reported from
Pakaraima Mountains of Guyana (Henkel et al. 2010), is
somewhat comparable to the present species because of a
similar looking pileus, a pruinose stipe, adnate lamellae,
somewhat similar sized basidiospores, abundant pseudocystidia and a similar habitat. However, R. pruinosistipitata
has a longer stipe, pip-shaped basidiospores, larger,
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241
Fig. 178 Lactarius dirkii
(holotype). a Fresh or dissected
basidiomata. b Radial section
through pileipellis showing
terminal and subterminal cells.
c Basidiospores.
d Cheiloleptocystidia.
e Pseudocystidia. f Basidia.
Scale bars a = 10 mm; b–
f = 10 lm
broadly ventricose to lageniform pseudocystidia, a
pileipellis lacking pileocystidia and a stipitipellis with
encrusted hyphae (Pegler 1977, 1986). Rhodocybe retroflexa (Berk. & Broome) Pegler reported from Sri Lanka
(Pegler 1977, 1986) has somewhat similar sized basidiocarps with similarly coloured pileus, almost similarly
attached lamellae, almost similar sized basidiospores of
similar shape, pseudocystidia with granular contents, and
an undifferentiated cutis-type pileipellis. However, that
species has a depressed pileus, a pseudoparenchymatous
subhymenium, and a pileipellis devoid of encrusted hyphae
and pileocystidia.
Comparison of the RPB2 (687 bp), ITS (412 bp) and
nLSU (884 bp) sequence data derived from Rhodocybe
indica with the nucleotide sequences of taxa available in
GenBank suggests that the sequences of the present species
are different. In a megablast search of the GenBank
nucleotide database using the RPB2 and ITS sequences, no
close hits with zero e-values were obtained. While using
the nLSU sequence, the closest hit was Rhodocybe fallax
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242
(GenBank AF261283; Identities = 853/884 (96 %)). Rhodocybe fallax (Quél.) Singer), belonging to the section
Decurrentes (Baroni 1981), and differs from the present
species in almost all macro- and micromorphological
characters.
In the resulting phylogenetic tree after the ML analysis
(Fig. 159), Rhodocybe indica, R. collybioides and R. caelata formed a distinct clade with significant support (73 %
BS). Within this clade, R. indica is related to R. collybioides with weak support (67 % BS).
Rhodocybe luteobrunnea K.N.A. Raj & Manim., sp. nov.
MycoBank MB 816842; Facesoffungi number:
FoF02180, Figs. 161 and 162
Etymology: The specific epithet refers to the yellowishbrown pileus.
Holotype: CAL 1322.
Basidiocarps small, mycenoid. Pileus 5–14 mm diam.,
initially somewhat conico-convex, becoming convex or
broadly convex with a small umbo; surface yellowishbrown (5E8/OAC734) on umbo and striations, brownishyellow (5D5, 5D6/OAC743) elsewhere, rather hygrophanous and becoming paler, finely pellucid-striate, glabrous; margin slightly incurved when young, becoming
deflexed to almost straight with age, finely wavy. Lamellae
sinuate or emarginate with a small decurrent tooth, close,
greyish-orange (5B3/OAC574), up to 3 mm wide, with
lamellulae of 3 lengths; edge entire to the naked eye, finely
torn under a lens, concolourous with the sides. Stipe
7–26 9 1–2 mm, central or slightly eccentric, terete,
equal, straight or somewhat flexuous, cartilaginous, solid;
surface brownish-orange (5C3/OAC730), glabrous to the
naked eye, finely appressed fibrillose all over under a lens,
finely pruinose towards the apex; base with white mycelial
cords. Odour and taste not distinctive.
Basidiospores 5–7 9 3–4.5 (5.67 ± 0.54 9 3.97 ±
0.49) lm, Q = 1.2–1.7, Qm = 1.44, lacrymoid or pipshaped, with or without a suprahilar depression in profile
view, up to 8, weak angular facets in polar view, finely
undulate-pustulate all over, thin-walled. Basidia
16–21 9 5.5–6.5 lm, clavate, pale yellow, thin-walled,
4-spored; sterigmata up to 4 lm long. Lamella-edge
heterogeneous. Cheilocystidia and pleurocystidia present
as pseudocystidia. Pseudocystidia 16–30 9 4–6 lm, narrowly fusiform, narrowly utriform or cylindrical with an
acute apex, with golden yellow or reddish-yellow contents,
thin-walled. Lamellar trama subregular; hyphae 4–9 lm
wide, hyaline or pale yellow, thin-walled. Subhymenium
inconspicuous. Pileus trama subregular; hyphae 4–7 lm
wide, with a pale yellow wall pigment and fine brown
spiral encrustations, thin- to slightly thick-walled.
Pileipellis an undifferentiated cutis; hyphae 4–7 lm wide,
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with a pale brownish-yellow wall pigment and fine brown
encrustations, thin- to slightly thick-walled. Stipitipellis a
cutis often disrupted by bunches of flaring out hyphae;
hyphae 3–7 lm wide, with a pale brownish-yellow wall
pigment and fine brown encrustations, thin- to slightly
thick-walled. Caulocystidia 10–14 9 2–4 lm, versiform:
lageniform, cylindrical with a short rostrate apex or flexuous, with a pale brownish-yellow wall pigment, thinwalled. Clamp connections not observed on any hyphae.
Habitat: in small groups, on forest floor, among
decaying litter.
Material examined: INDIA, Kerala State, Thrissur District, Peechi, Peechi Forest, 26 July 2010, K. N. Anil Raj
AR180 (CAL 1322, holotype).
Notes: Rhodocybe luteobrunnea is distinguished by its
yellowish-brown basidiocarps with a finely pellucid-striate
pileus; sinuate or emarginate lamellae; lacrymoid or pipshaped basidiospores; cylindrical to fusiform pseudocystidia and encrusted pileipellis. A combination of characters
such as the centrally stipitate basidiocarps and the presence
of pseudocystidia with coloured contents (in KOH) lead the
present species to the section Rhodocybe. Rhodocybe perplexa T.J. Baroni & Watling from Malaysia (Baroni and
Watling 1999) resembles R. luteobrunnea in having a
pileus of similar size and shape, somewhat similar sized
basidiospores, presence of pseudocystidia, pileipellis with
encrusted hyphae and clampless hyphae (Baroni and
Watling 1999). However, R. perplexa has differently
coloured basidiocarps, adnate lamellae, shorter stipe, subglobose to ellipsoid basidiospores and larger pseudocystidia. Rhodocybe naucoria Singer, an Argentinean species
has similar shaped pileus, similar-sized and somewhat
similar-shaped basidiospores and presence of pseudocystidia (Baroni 1981). However, that species differs from the
present one in having smaller sized and differentlycoloured basidiocarps, differently attached lamellae, larger
pseudocystidia and presence of birefringent crystals in the
hymenial hyphae.
Comparison of RPB2 (648 bp), ITS (675 bp) and nLSU
(830 bp) sequence data derived from the present Rhodocybe species with the nucleotide sequences of taxa available in GenBank suggests that the sequences are different.
In a BLASTn search using the ITS sequence derived from
the present species, no close hits with a zero e-value were
obtained. Rhodocybe aureicystidiata is the closest hit while
conducting a BLASTn search using both nLSU (GenBank
AY380407; Identities = 840/852 (99 %)) and RPB2
(GenBank AY337412; Identities = 604/648 (93 %))
sequences.
Rhodocybe aureicystidiata Lennox ex T.J. Baroni, a
species belonging to the section Rhodocybe (Baroni 1981),
shows some similarity to the present species in having
rather similar-coloured basidiocarps, encrusted hyphae in
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243
Fig. 179 Phylogenetic tree of
Mortierella fluviae EMLYR25716-1 and EMLYR25716-2 and related species
before based on maximum
likelihood analysis of ITS
rDNA sequences. Sequence of
Umbelopsis isabellina was used
as outgroup. Numbers at the
nodes indicate the bootstrap
values ([50 %) from 1000
replications. The bar indicates
the number of substitutions per
position. New taxa are in blue
Fig. 180 Phylogenetic tree of
Mortierella fluviae EMLYR25716-1 and EMLYR25716-2 and related species
based on maximum likelihood
analysis of 28S rDNA
sequences. Sequence of
Umbelopsis isabellina was used
as outgroup. Numbers at the
nodes indicate the bootstrap
values ([50 %) from 1000
replications. The bar indicates
the number of substitutions per
position. New taxa are in blue
the pileipellis and presence of pseudocystidia. However,
that species has larger basidiocarps, a depressed pileus
which becomes dark red on bruising, inrolled margin, larger and amygdaliform basidiospores, larger and differently
shaped pseudocystidia and a different geographical location (Baroni 1981).
In the phylogram (Fig. 159) generated from the ML
analysis, Rhodocybe luteobrunnea and R. aureicystidiata,
R. pruinosostipitata, R. mellea and Rhodocybe sp. formed
a distinct clade with significant support (99 % BS).
Within this, R. luteobrunnea and R. aureicystidiata
formed another subclade also with significant support
(90 % BS).
Rhodocybe griseoaurantia K.N.A. Raj & Manim., sp. nov.
MycoBank MB 816843; Facesoffungi number:
FoF02181, Figs. 163 and 164
Etymology: The specific epithet refers to the greyishorange colour of the pileus.
Holotype: CAL 1324.
Basidiocarp small. Pileus 14 mm diam., convex with a
slightly raised centre; surface greyish-orange (6B6/
OAC694) at the centre and pale orange (6A3/OAC695)
toward the margin, hygrophanous and soon becoming
orange-white (6A2/OAC696) after collection, not striate,
finely appressed squamulose on and around the centre and
minutely pubescent towards the margins; margin slightly
123
244
incurved, rather wavy. Lamellae adnate to short decurrent,
close, pale orange (6A3/ OAC695), up to 3.5 mm wide,
occasionally furcate, with lamellulae of 1–4 lengths; edge
entire or finely torn, concolourous with the sides. Stipe
31 9 4 mm, central, terete, slightly tapering towards the
apex, solid; surface orange-white (6A2/OAC696), finely
appressed-fibrillose all over, finely pruinose towards the
apex; base slightly swollen, with whitish mycelial cords.
Odour and taste not distinctive.
Basidiospores 5–7 9 3.5–4.5 (5.85 ± 0.67 9 3.82 ±
0.33) lm, Q = 1.25–2, Qm = 1.53; with 6–7 facets in
polar view, ellipsoid in profile with or without a suprahilar
depression, weakly undulate-pustulate all over, hyaline,
thin-walled. Basidia 19–25 9 5–7 lm, clavate, hyaline,
thin-walled, 4-spored; sterigmata up to 3 lm long.
Lamella-edge often fertile or occasionally heterogeneous.
Cheilocystidia 14–29 9 3–9 lm, infrequent, scattered,
often filiform or cylindrical, sometimes flexuous or narrowly fusiform. Pleurocystidia none. Lamellar trama subregular; hyphae 4–5.5 lm wide, hyaline, thin-walled.
Subhymenium inconspicuous. Pileus trama subregular;
hyphae 3.5–8 lm wide, pale yellow, thin-walled.
Pileipellis a cutis often disrupted by scattered or rarely
clustered ascending hyphae; hyphae 2.5–6 lm wide, with a
pale yellow wall pigment and occasionally with faint,
hyaline encrustations, thin- to slightly thick-walled. Stipitipellis a cutis occasionally disrupted by scattered or clustered flaring-out hyphae; hyphae 3.5–6 lm wide, hyaline or
pale yellow, thin-walled. Caulocystidia absent. Clamp
connections not observed on any hyphae.
Habitat: on soil, among moss, solitary.
Material examined: INDIA, Kerala State, Wayanad District, Muthanga, Muthanga Wildlife Sanctuary, 6 September
2011, K. N. Anil Raj AR865 (CAL 1324, holotype).
Notes: Rhodocybe griseoaurantia is well characterized
by its small basidiocarps with a greyish-orange pileus;
adnate to short decurrent lamellae; ellipsoid basidiospores
and filiform or cylindrical cheilocystidia are the diagnostic
features of R. griseoaurantia. Owing to the centrally stipitate basidiocarp, presence of cheilocystidia and the
absence of both pseudocystidia and clamp connections,
R. griseoaurantia can be placed in the section Rufrobrunnea (Baroni 1981). Rhodocybe alutacea Singer, a North
American species (Baroni 1981; Baroni and Horak
1994), is similar to the present species in having similar
coloured basidiocarps, a similar looking pileus, narrow
lamellae with similar type of attachment, a solid stipe,
almost similar-sized basidiospores of similar shape,
infrequent, scattered cheilocystidia, pileipellis with
encrusted hyphae and non-encrusted hyphae of stipitipellis. However, R. alutacea has larger basidiocarps, an
umbilicate pileus, basidiospores with more facets (7–9)
in profile view, larger, septate cheilocystidia, presence of
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Fig. 181 Mortierella fluviae (holotype). a, d Colony in potato c
dextrose agar. b, e Colony in oatmeal agar. c, f Colony in water agar
(a–c above view, d–f reverse view) (g–r light microscope; s–u SEM).
g–j Young and mature sporangia on sporangiophores and apophysis
(i, s) (white arrow). k–m Single and branched sporangiophores. n–
r Different shapes of columellae (r, t) (purple arrow) with collarette
(green arrow) and sporangial septum (yellow arrow). u Spores. Scale
bars g–j, n, p–r = 20 lm, k–m, o = 50 lm, s = 30 lm, t = 10 lm,
u = 5 lm
caulocystidia and a different habitat and geographical
location.
Comparison of the RPB2 (681 bp), ITS (733 bp) and
nLSU (908 bp) sequence data derived from R. griseoaurantia with the nucleotide sequences of taxa available in
GenBank suggests that the present species is different. In
the BLASTn search using the RPB2 sequence, the closest
hit was Rhodocybe truncata (GenBank EF421019; Identities = 616/659 (93 %)). For ITS sequence, the closest hit
was an unidentified Rhodocybe species (Rhodocybe sp. 1
GMB-2014) (GenBank KP012803; Identities = 719/737
(98 %)) from Australia, followed by R. truncata (GenBank
EF421110; Identities = 556/644 (86 %)). In the BLASTn
search with the nLSU sequence, the closest hits was Rhodocybe gemina (GenBank DQ071715; Identities = 894/
908 (98 %)). Rhodocybe truncata (Schaeff.) Singer, a
European species, differs from the present one in almost all
macro- and microscopic characters, although it has
cheilocystidia and rather similar-sized (5–6.5 9 4–5 lm)
basidiospores (Baroni 1981). Rhodocybe gemina (Paulet)
Kuyper & Noordel., another European species belonging to
the section Rufrobrunnea, differs from the present species
in having larger and differently-coloured basidiocarps,
crowded lamellae, a perfumed odour, unpleasant taste,
slightly larger and differently shaped basidiospores
(5–7 9 4–5.5 lm), and a cutis with transition to a trichoderm type of pileipellis (Noordeloos 1988).
In the phylogram (Fig. 159), R. griseoaurantia, R. truncata and R. gemina formed a strongly support clade. This
clade received significant support (98 % BS) and within it
R. griseoaurantia was found to be phylogenetically distinct
from R. truncata and R. gemina.
Agaricaceae Chevall.
We follow Li et al. (2016)
Cyathus Haller
The genus Cyathus belongs in the Agaricaceae, despite
the unusual form of its basidiomata (also known as peridia), which resemble a bird’s nest and grow on wood or
dung. These peridia are typically vase-, trumpet- or urnshaped with dimensions of 4–8 mm wide and 7–18 mm
high. The type species, Cyathus striatus (Huds.) Willd. was
described from Europe, but the genus has a cosmopolitan
distribution and comprises ca. 45 species (Das et al. 2016).
Fungal Diversity (2016) 80:1–270
245
123
246
Fungal Diversity (2016) 80:1–270
Fig. 182 Phylogenetic tree of
Cunninghamella constructed
using the ITS rDNA sequences.
Absidia anomala and
Halteromyces radiatus were
used as outgroup taxa.
Sequences are labeled with their
database accession numbers.
Support values are from
Bayesian inference. Sequences
obtained in this study are clones
annotated in blue
Besides the studies of Zhao et al. (Zhao et al. 2006, 2007),
the tropical species are less known, and we here describe a
new taxon from Thailand that was recently identified as a
123
producer of novel terpene alkaloid antibiotics. A phylogenetic tree in support of the new species is provided in
Fig. 165.
Fungal Diversity (2016) 80:1–270
Fig. 183 Phylogenetic tree of Cunninghamellaceae constructed
using the large subunit (LSU) rDNA sequence data. Mortierella
parvispora species was used as outgroup. Sequences are labeled with
247
their database accession numbers. Support values are from Bayesian
inference. The sequences obtained in this study are clones annotated
in blue
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248
123
Fungal Diversity (2016) 80:1–270
Fungal Diversity (2016) 80:1–270
b Fig. 184 Cunninghamella gigacellularis (holotype). a Colony sur-
face. b Sporophore with branches in pair. c Sporophore with branches
in two whorls of 4 and 3. d Sporophore with single and in pairs
branches. e, f Sporophore repeatedly branched. g Sporophore
branched. h Giant cels. i Sporangiola
Cyathus pyristriatus B. Thongbai, C. Richt. & M. Stadler,
sp. nov.
MycoBank number: MB817167; Facesoffungi number:
FoF02385, Fig. 166
Etymology: Named for its production of pyristriatins A
and B.
Holotype: MFLU15-1416.
Saprobic on rotten wood in forest with Fagaceae. Basidiomata clavate to broadly obconic, without stipe,
5.5–7 mm high and 4–6 mm diam., wide at the top, external
peridium covered with shaggy or fluffy yellowish-brown or
buff hairs with age, surface of inner peridium grey, darkening
with age, distinctly plicate. Peridioles 3–3.5 mm in diam.
wide, greyish-brown to dark grey covered with minute,
greyish to greyish-brown hairs. Basidiospores 14–17 9
8–10 lm (av. = 15.18 9 8.12, Q = 1.5–1.91, Qm = 1.70,
n = 40), ellipsoid to broadly ellipsoid, some ovoid, rarely
subglobose, hyaline, smooth, thin-walled, 1.5–3 lm thick.
Basidia not observed. Clamp connections present in all tissues including the mycelial culture.
Material examined: THAILAND, Chiang Mai Province,
Mae-Taeng District, near the Mushroom Research Centre
(http://www.mushroomresearchcentre.com/), N19°07.2000
E98°44.0440 , 12 August 2014, Thongbai, Richter & Stadler
M68 (MFLU 15-1416, holotype), ex-type culture
MFLUCC14-0770.
Notes: Cyathus pyristriatus closely resembles C. striatus
(Huds.) Willd. 1787 in the account of shaggy or fluffy yellowish-brown or buff hairs of peridium, distinctly plicate
inner peridium, while peridioles are greyish-brown to dark
grey. Based on morphological comparison between
C. striatus and C. pyristriatus the latter has smaller basidioma and also smaller ellipsoid basidiospores. However,
peridioles are covered with a distinctive minutely greyish to
greyish-brown hairs, while peridioles of C. striatus are
smooth. Remarkably, C. striatus was reported producing
pale to dark pigments from basidiomata, which vary in size
(Kuo 2014; Wood and Stevens 2015).
An initial BLAST search of the ITS nucleotide sequence
from NCBI database (http://www.ncbi.nlm.nih.gov/) gave
the closest hit to a Cyathus striatus strain (EU784194) with
a maximum similarity of 94.5 % (Brock et al. 2009). The
BLAST search of the LSU nucleotide sequence of
C. pyristriatus resulted in a high degree similarity of
98.8 % (DQ071742) with C. striatus (Garnica et al. 2007).
Figure 165 illustrates the phylogenetic relationships of
249
C. pyristriatus, C. striatus, and other Cyathus species. The
ITS based phylogenies clearly depict that our new taxon
belongs to the genus Cyathus. A close affinity between
C. pyristriatus and C. stercoreus collected from China, is
also noted with high support. Cyathus stercoreus, however,
differs in having a smooth surface of inner peridium and
larger globose to oval basidiospores (Zhao et al. 2008).
The ex-type culture of the new species also produces
novel terpene alkaloid antibiotics named pyristriatins, which
have never been found in Cyathus, despite the fact that this
genus has been evaluated intensively for secondary
metabolites for several decades (Richter et al. 2016).
Polyporales Gäum.
Polyporaceae Fr. ex Corda
We follow Zhou et al. (2016).
Polyporus P. Micheli ex Adans.
Polyporus, the type genus of Polyporaceae, is characterized by annual stipitate basidiocarps, a dimitic hyphal
system with generative hyphae and skeleto-binding
hyphae, hyaline and thin-walled cylindrical basidiospores,
and are the cause of white rot (Gilbertson and Ryvarden
1987). Phylogenetic analysis reports Polyporus as polyphyletic (Ko and Jung 2002; Krüger et al. 2006; Sotome
et al. 2008, 2011). Recently, based on a multi-locus phylogeny, species of Polyporus, Favolus and Neofavolus were
divided into six highly supported clades, i.e. Favolus clade,
Melanopus clade, Neofavolus clade, Polyporellus clade,
Polyporus clade and Squamosus clade (Zhou et al. 2016).
A phylogenetic tree in support of the new species is provided in Fig. 167.
Polyporus mangshanensis B.K. Cui, J.L. Zhou & Y.C.
Dai, sp. nov.
Index Fungorum number: IF552159; Facesoffungi
number: FoF02471, Figs. 168 and 169
Etymology: mangshanensis, referring to the locality of
the type specimen.
Holotype: BJFC 018267.
Basidiocarps annual, eccentrically to almost laterally
stipitate, solitary, corky when dry; pileus reniform, projecting up to 5.8 cm long, 10.5 cm wide and 2.5 cm thick;
pileal surface beige, saffron yellow to yellowish-orange
when dry, azonate, with slightly radial stripes, smooth and
glabrous; margin sharp and involute upon drying; pore
surface brown beige to olive brown when dry; pores
angular, 3–5 per mm, occasionally elongated to 1 mm long
and 0.5 mm wide; dissepiments thin, entire to lacerate;
context buff when dry, up to 1.2 mm thick; tubes concolorous with the pore surface, slightly decurrent on the stipe,
up to 1.5 mm long; stipe cylindrical, context of stipe buff
when dry, bearing dark brown cuticle, becoming tan
123
250
towards the tuber layer, glabrous, curvy and wrinkled upon
drying, up to 2 cm long and 7 mm diam. Hyphal system
dimitic; generative hyphae bearing both clamp connections
and simple septa; skeleto-binding hyphae IKI–, CB–; tissues
unchanged in KOH. Contextual generative hyphae frequent,
colourless, thin-walled, bearing clamp connections, frequently branched, 2.5–8 lm diam., usually inflating at the
branching areas or clamping areas, up to 14 lm in diam.;
contextual skeleto-binding hyphae dominant, colourless,
thick-walled with a narrow to wide lumen, frequently branched, flexuous, interwoven, 2.5–6.5 lm diam., occasionally
inflated up to 11 lm diam.; contextual cuticle hyphae simple
septate, thin-walled, frequently branched, interwoven,
1.5–4.5 lm diam. Tramal generative hyphae frequent,
colourless, thin-walled, bearing clamp connections and
simple septa, infrequently branched, 2.5–4 lm diam.; tramal
skeleto-binding hyphae dominant, colourless, thick-walled
with a narrow to wide lumen, frequently branched, flexuous,
interwoven, 1.3–4 lm diam. Stipe generative hyphae frequent, colourless, thin-walled, bearing clamp connections,
occasionally branched, 1.5–6 lm diam.; stipe skeleto-binding hyphae dominant, colourless, thick-walled, subsolid or
with a narrow to wide lumen, moderately branched, flexuous,
interwoven, 2.5–6 lm diam.; stipe cuticle generative hyphae
dominant, with buff inclusion inside, thick-walled with a
wide lumen, bearing clamp connections; stipe cuticle skeleto-binding hyphae frequent, with light brown to orange
brown inclusion inside, thick-walled with a narrow lumen,
occasionally branched, 2.4–4.5 lm diam. Cystidia and cystidioles absent; basidia clavate, with a basal clamp and four
sterigmata, 16.5–24 9 7–10 lm; basidioles in shape similar
to basidia, but smaller and without sterigmata. Basidiospores
cylindrical, few, oblong, colourless, thin-walled, smooth,
with guttules, IKI–, CB–, (6.5–)7.5–10.5(–11) 9 3.5–4.5
(–5) lm, L = 8.7 lm, W = 4.07 lm, Q = 1.84–2.53,
Qm = 2.14 (n = 60/1).
Type of rot: White rot.
Specimens examined: CHINA, Hunan Province, Yizhang County, Mangshan Nature Reserve, on fallen
angiosperm branch, 17 August 2014, Dai 15151 (BJFC
018267, holotype).
Notes: Recently, taxonomic and phylogenetic studies of
wood-rotting fungi in subtropical China have been carried
out, and many new species have been described based on
both morphological characters and molecular data (Chen
and Cui 2014, 2016; Chen et al. 2015c, 2016; Han et al.
2016; Li and Cui 2013; Li et al. 2014; Song et al.
2014, 2016; Zhao et al. 2013, 2015a). In the current study,
Polyporus mangshanensis is an additional new species
described from subtropical China on the basis of morphological characters and phylogenetic analysis. Morphologically, P. mangshanensis is characterized by its beige to
yellowish-orange pileal surface with slightly radial stripes,
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Fungal Diversity (2016) 80:1–270
brown beige to olive brown pore surface, dark brown stipe,
generative hyphae bearing both clamp connections and
simple septa, and oblong to cylindrical basidiospores
(7.5–10.5 9 3.5–4.5 lm).
Phylogenetically, P. mangshanensis clustered with P. leprieurii Mont. and P. guianensis Mont.; morphologically,
they produce similar pileal surface, pore surface and dark
stipe, however, P. mangshanensis differs from the latter two
species by producing both clamped and simple-septate
generative hyphae, contextual generative hyphae and skeleto-binding hyphae usually inflated over 10 lm in diam.
Moreover, pores and basidiospores of P. mangshanensis are
smaller than P. guianensis (pores 1–2 per mm, basidiospores
8–12 9 2.5–4 lm), while larger than P. leprieurii (pores
5–8 per mm, basidiospores 4.5–7 9 2–2.5 lm; Núñez and
Ryvarden 1995).
Polyporus subvarius C.J. Yu & Y.C. Dai has a similar
pileal and pore surface to P. mangshanensis, but the former
has larger pores (1–2 per mm) and basidiospores
(9.2–12.6 9 3.9–4.9 lm, L = 10.7 lm, W = 4.48 lm),
only clamped generative hyphae, and grows on living trees
of Salix (Dai et al. 2007).
Polyporus admirabilis Peck resembles P. mangshanensis in having a laterally dark stipe, a tan pileal surface and
similar pore size, but it differs in its subulate cystidioles,
only clamped generative hyphae, and slender basidiospores
(7.8–9 9 3–3.5 lm,
L = 8.29 lm,
W = 3.12 lm,
Qm = 2.66; Dai 1999).
Both Polyporus badius (Pers.) A.B. De and P. submelanopus
H.J. Xue & L.W. Zhou have simple-septate generative hyphae,
dark stipe and cylindrical basidiospores, which are similar to
P. mangshanensis. Polyporus badius has darker pileal surface
and smaller pores (5–6 per mm), only simple-septate generative
hyphae (Dai 1999); while P. submelanopus has larger pores
(2–3 per mm) and basidia (24–33 9 5–8 lm), and a terrestrial
habit (Xue and Zhou 2012).
Russulales Kreisel ex P.M. Kirk et al.
Russulaceae Lotsy
The family Russulaceae is one of the dominant and morphologically diverse (Miller et al. 2006) ectomycorrhizal
mushroom families in the Himalayas. Apart from its three
corticoid genera, i.e. Boidinia, Gloeopeniophorella and
Pseudoxenasma (Larsson and Larsson 2003; Miller et al.
2006), this family has four predominantly agaricoid genera:
Lactifluus, Lactarius, Multifurca and Russula (Buyck et al.
2008, 2010), some of which may also contain secotioidhypogeous or pleurotoid species.
Russula Pers.
The monophyletic agaricoid genus Russula (Buyck et al.
2008, 2010) has an enormous diversity in Indian Himalaya
Fungal Diversity (2016) 80:1–270
(Rawla 2001; Das and Sharma 2005; Das et al.
2006, 2010, 2013, 2014) showing its wide range of distribution from tropical to subalpine areas and associations with
broadleaf to coniferous trees. Apart from morphology-based
six subgenera (Sarnari 1998; R. subg. Compactae, R. subg.
Heterophyllidia, R. subg. Ingratula, R. subg. Amoenula,
R. subg. Incrustatula and R. subg. Russula), three additional
subgenera have been recently described (Hongsanan et al.
2015a; R. subg. Archaea Buyck & V. Hofstetter, R. subg.
Brevipes Buyck & V. Hofstetter and R. subg. Malodora
Buyck & V. Hofstetter). Here, two novel species (belonging
to R. subg. Heterophyllidia and R. subg. Amoenula respectively), collected from the northwestern part of Indian
Himalaya are introduced together with their morphology and
phylogenetic placement (Figs. 170, 173).
Russula indoalba A. Ghosh, Buyck, A. Baghela, K. Das &
R.P. Bhatt, sp. nov.
Index Fungorum number: IF552159; Facesoffungi
number: FoF02471, Figs. 171 and 172
Etymology: referring to white basidiomata growing in
India
Holotype: CAL 1328.
Basidiomata 70–100 mm. in height. Pileus 30–95 mm. in
diam., broadly convex when young, gradually planoconvex
to applanate with depressed center, finally uplifted at maturity; margin decurved to plane, entire, tuberculately striate;
surface dry, viscid when moist, cracked-areolate when
mature, white (1A1–2A1) with greyish-yellow (4B4–4B6) to
pale yellow (3A3) spots at the center; lemon-yellow or pale
yellow to pastel yellow (1A3–1A4) with KOH; cap context
white (1A1–2A1), lemon-yellow or pale yellow to pastel
yellow (1A3–1A4) with KOH. Lamellae adnexed to almost
free with age, equal, subdistant to close (7–10/cm), forked
near the stipe apex, white (1A1–1A2), with entire, concolourous gill edges. Stipe 20–65 9 10–23 mm., subclavate, central, dry, smooth, white (1A1–2A1), lemonyellow or pale yellow to pastel yellow (1A3–1A4) with
KOH; context stuffed, becoming hollow with age, white
(1A1–2A1), greyish-orange (5B4–5B5) with FeSO4,
unchanging with guaiacol and NH4OH. Taste mild. Odour
indistinctive. Spore print yellowish-cream. Basidiospores
5.5–7.42–9.5 9 5.5–6.4–8 lm (n = 50, Q = 1–1.16–
1.36), subglobose to broadly ellipsoid, rarely ellipsoid;
ornamentation amyloid, composed of short (0.3–0.5 lm)
and long (0.7–1 lm) conical to cylindric warts, mostly
connected to form incomplete reticulum, few isolated, apiculi up to 2 lm high; suprahilar spot not amyloid. Basidia
45–60 9 9–13 lm, 4-spored, cylindrical to subclavate,
sterigmata
up
to
5 lm
long.
Pleurocystidia
47–90 9 8–16 lm, subclavate to clavate with capitate,
moniliform, appendiculate and rounded apex, emergent up to
38 lm beyond the basidiole tips. Cheilocystidia
251
55–68 9 6–8 lm, cylindrical to subclavate with appendiculate apex. Subhymenium layer up to 26 lm thick, pseudoparenchymatous. Lamellar trama consists predominantly of
sphaerocytes. Gill edges fertile. Pileipellis up to 120 lm
thick, distinctly divided into suprapellis and supellis; subprapellis composed of erect to suberect chains of 3–6 rows of
cells; subterminal cells 8–20 9 6–18 lm, mostly rounded
(inflated), ellipsoid; terminal cells mostly, cylindrical to
subulate; subpellis composed of. horizontal, interwoven
hyphae. Pileocystidia rare, one-celled, slender, thin-walled,
up to 7 lm wide, dispersed as terminal cells in the
suprapellis.
Habitat and distribution: Under Quercus sp. in mixed
forests dominated by Quercus, Rhododendron, Abies and
Cupressus.
Material examined: INDIA, Uttarakhand, Rudraprayag
district, Baniyakund,
alt.
2630 m,
N30°28.9140
0
E79°10.854 , 1 September 2014, A. Ghosh, AG 15-541
(GUH); ibid., 14 July 2015, A. Ghosh, AG 15-628 (CAL
1328, holotype); ibid., A. Ghosh, 16 July 2015, AG 15-661
(GUH); ibid., 1 August 2015, A. Ghosh, AG 15-797
(GUH).
Notes: Russula indoalba which is characterized by white
pileus with greyish-yellow to pale yellow spots at center,
tuberculately striate and wavy to interrupted pilear margin,
equal gills, yellowish-cream spore print and mild taste is a
typical member of Russula subg. Heterophyllidia
sect. Virescentinae.
In the field Russula kanadii A.K. Dutta & K. Acharya
(also reported from India) appears to be quite similar to the
present taxon. However, R kanadii has a white spore print
(1A1),
smaller
basidiospores
[(4.5)–5.5–5.7–6.5
(–7) 9 (4.5)–5.3–5.5(–6) lm] and grows at very low altitudes (56 m) being ectomycorrhizal with dipterocarps, viz.
Shorea robusta (Dutta et al. 2015). Russula alboareolata
Hongo, described from Japan in association with Castanopsis (Fagaceae), but also reported from dipterocarps in
Taiwan (Watling and Lee 1998) and Thailand is also very
similar to R. indoalba but has smaller basidiospores
6.5–8.5 9 5.5–7 lm (Hongo 1979). Our phylogeny
(Fig. 170) shows that R. alboareolata is genetically closer
to R. kanadii than to our new species or to the other
Virescentinae (Fig. 170). Furthermore, the phylogeny also
suggests, although without support, that R. indoalba is
basal to the core group of Virescentinae (R. virescens and
allies) suggesting a migration from India via Asia
(Fig. 170, sequence UDB014226) to both Europe and
North America. All species in this core-group differ from
other Virescentinae such as those belonging to the
R. crustosa-R. mustelina lineage and also R. parvovirescens Buyck et al. (Buyck et al. 2006) in the absence
of dermatocystidia in the lower subpellis (Buyck, http://
www2.muse.it/russulales-news/id_virescentinae.asp).
123
252
Russula pseudoamoenicolor A. Ghosh, Buyck, K. Das, A.
Baghela & R.P. Bhatt, sp. nov.
MycoBank No.: MB 817101; Facesoffungi number:
FoF02522, Figs. 174 and 175
Etymology: referring to the lookalike of Russula
amoenicolor, an European species.
Holotype: CAL 1330.
Basidiomata up to 100 mm in height. Pileus
50–100 mm in diam., globose, plano-convex to applanate
with broadly depressed center, becoming uplifted when
mature; margin decurved, entire, tuberculately striate, torn
when mature, surface dry, viscid when moist, subvelvety,
purplish-red-violet-red (14B5–14B8) or light lilac, light
violet to pastel violet (16A4–16A5) with purplish-white
(14A2) to violet-white (16A2) towards margin, rarely dark
violet (15A6–15A8) towards the depression; cuticle peeling 3/4th of the radius; cap context white (1A1–2A1),
unchanging when bruised. Lamellae adnexed to subdecurrent, close to rather crowded, white (1A1–2A1), forked
near the stipe; edges marginate near the cap margin,
lamellulae absent. Stipe 45–70 9 10–14 mm, equal,
slightly tapered towards the base, dry, smooth, brittle,
central, reddish-white to pink-rose or pink-red
(12A3–12A5), context stuffed, white (1A1–2A1), pinkrose (12A3–12A5) with guaiacol. Taste mild. Spore print
not obtained.
Basidiospores 6–7.30–9.5 9 5–6.33–8 lm (n = 25,
Q = 1.03–1.16–1.33), subglobose to broadly ellipsoid,
rarely ellipsoid, ornamentation amyloid, composed mostly
of ridges and warts (up to 1 lm high) aligned or connected
to form an incomplete reticulum, with few isolated warts,
apiculi up to 2 lm high. Basidia 35–55 9 9–13 lm,
cylindrical, subclavate to clavate, 4-spored, sterigmata up
to 6 lm high. Subhymenium layer up to 30 lm thick,
pseudoparenchymatous. Hymenophoral trama mainly consisting of sphaerocytes measuring 17–40 9 16–34 lm.
Pleurocystidia 90–117 9 10–21 lm, ventricose, subfusiform to fusiform with blunt apex, thick walled (1 lm
thick), emergent up to 60 lm; content blank or insignificant. Gill edges fertile, with basidia and cystidia. Cheilocystidia 30–85 9 7–10 lm, same as pleurocystidia.
Pileipellis up to 90 lm thick, composed of clustered erect
to suberect elements composed of chains of 4–7 cells;
terminal cells ellipsoid, conical, subfusoid to occasionally
subulate (11–65 9 4–10 lm wide); subterminal cells
mostly cylindrical to rectangular, few ellipsoid or rounded
(inflated), measuring up to 14 lm wide.
Habitat and distribution: grows in close association with
Quercus sp. with undergrowth of Rhododendron sp. in
moist deciduous and mixed (broadleaf and coniferous)
forest.
Material examined: INDIA, Uttarakhand, Pauri Garhwal, along the road side of khirsu, alt. 1835 m.,
123
Fungal Diversity (2016) 80:1–270
N30°10.1550 E78°52.1340 , 24 July 2015, A. Ghosh, AG
15-739 (CAL 1330, holotype)
Notes: The combination of characters in Russula pseudoamoenicolor comprising a purplish-red to violet-red or
lilac subvelvety pileus with darker center, reddish-violet to
pink-rose stipe, occasional occurrence of typically subulate
terminal cells of pileipellis, absence of dermatocystidia and
inamyloid suprahilar spot place it in R. subg. Amoenula
Sarnari. In the field, the European species, R. amoenicolor
Romagn. appears to be quite similar to the present taxon
but the former (Sarnari 1998) has purple to green or variegated pileus, smaller basidiospores (6.7–8.4 9
5.6–7.4 lm) and mostly subulate terminal cells in
pileipellis (terminal cells mainly ellipsoid, conical, subfusoid or occasionally subulate in R. pseudoamoenicolor) and
its ITS nucleotide sequence are dissimilar (93 % identity
with R. pseudoamoenicolor for 100 % query coverage
using BLAST) from the present species. From a phylogenetic standpoint, R. pseudoamoenicolor is closely related to
R. violeipes and cluster together in a subclade with high
bootstrap support (Fig. 173).
The European species, R. violeipes Quél. was recently
reported from South Korea (Park et al. 2013), yet these
collections differs significantly from European material
(GenBank acc. no. AY061726). These Korean collections
(GenBank accession nos. KF361797 and KF361783) are
here, however, recovered as identical to our material from
the Indian Himalaya.
Interestingly, two Australian species, R. variispora T.
Lebel and R. rostraticystidia T. Lebel, previously of
undetermined position within R. subg. Heterophyllidia
(Lebel and Tonkin 2007) are here for the first time
recovered with significant support (93 % BS) as very closely related to or likely members of R. subsect. Amoeninae.
Russula variispora and R. rostraticystidia both are
sequestrate species and hence, have an altogether different
morphology from other known Amoeninae, or even from
all other known species in R. subg. Heterophyllidia as
secotioid-gasteroid taxa have never been reported from
other subsections in this subgenus (Kong et al. 2015). The
rostrate pleurocystidia of R. rostraticystidia and the similarly shaped terminal cells in the less developed pileipellis
of these Australian secotioid species (Lebel and Tonkin
2007) support their placement in Amoeninae.
Lactarius Pers.
The well-known milkcaps are split into two genera:
Lactarius and Lactifluus (Buyck et al. 2008, 2010). The
genus Lactarius comprises three subgenera, L. subg.
Piperites (Fr. ex J. Kickx f.) Kauffman, L. subg. Russularia
(Fr. ex Burl.) Kauffman and L. subg. Plinthogali (Burl.)
Hesler & A.H. Sm. and is one of the common ectomycorrhizal associates in Indian Himalaya (Rawla 2002; Das
Fungal Diversity (2016) 80:1–270
and Sharma 2005; Das and Verbeken 2011, 2012; Das and
Chakraborty 2014; Das et al. 2015). One undescribed
species of L. subg. Plinthogali collected from northwest
part of Indian Himalaya is introduced here with morphological details and phylogenetic evidence. A phylogenetic
tree is presented in Fig. 176.
Lactarius dirkii Uniyal, K. Das, A. Baghela & R.P. Bhatt,
sp. nov.
MycoBank: MB 817126; Facesoffungi number
FoF02521, Figs. 177 and 178
Etymology: commemorating Dr. Dirk Stubbe for his
contribution to Lactarius subg. Plinthogali.
Holotype: CAL 1332.
Pileus 20–86 mm. in diam.; convex when young,
becoming planoconvex with shallowly depressed center
with maturity; margin decurved, entire to irregularly
undulating, often interrupted; surface dry, slightly wrinkled
towards margin when mature; white to orange-white (6A2)
with a pinkish-white (7A2) or paler tinge, spotted with
reddish-grey (7B2) scrobicules, changing to light yellow
(2A5) with 10 % KOH. Lamellae subdecurrent, close to
rather crowded (13–14/10 mm), some forked near the stipe
apex, white to orange-white (5A2) to pale orange (5A3),
becoming pinkish to pale red (7A2) to pastel red (7A3/4)
when bruised or cut, lamellulae numerous. Stipe
30–58 9 5–8 mm., cylindrical, tapering toward base,
central, dry, concolourous with pileus, white at base with
minute hairs; Context whitish to yellowish-white (4A2),
turning light orange on cutting, hollow in stipe; Latex
abundant, white, unchanging when isolated, but drying
pinkish on cut lamellae. Taste bitter to acrid. Odour
insignificant. Spore print light yellow (4A5). Basidiospores
6.5–7.7–8.5 9 6–6.9–7.5 lm, (n = 70, Q = 1.06–1.11–
1.21), subglobose to broadly ellipsoid, rarely broadly
ellipsoid, winged; ornamentations amyloid, composed
mostly of broad wings and few small ridges but never
forming a reticulum, up to 2 lm high, edges smooth, isolated warts also present between ridges, plage amyloid.
Basidia 44–62 9 11–14 lm, subclavate, 4-spored, sterigmata 4–7 lm long. Hymenophoral trama composed of
lactifers and rosettes of sphaerocytes. Pleurocystidia
absent. Pseudocystidia abundant, emergent, cylindrical,
sometimes tortuous, branched, up to 5.5 lm wide. Lamellar edge sterile. Cheiloleptocystidia 24–35 9 4–5.5 lm,
mostly cylindrical, rarely subfusiform, thin walled. Subhymenium up to 20 lm thick, pseudoparenchymatous.
Lactifers in hymenophoral trama up to 7 lm wide.
Pileipellis trichopalisade, 55–121 lm thick, suprapellis
formed of cylindrical to subfusiform cells and septate
hyphal elements 10–57 9 3.5–6.5 lm; subpellis composed
of cells with irregular shapes, cells up to 11 lm wide;
Lactifers in pilear trama up to 8 lm wide, sometimes
253
becoming gradually thin toward suprapellis and projecting
in form of pileopseudocystidia, wide up to 4.5 lm. Stipitipellis trichoderm, up to 100 lm thick, hyphae up to 5 lm
wide.
Habitat & distribution: Under Quercus sp., Rhododendron arboreum in temperate mixed forest dominated by
Abies, Cupressus, Quercus and Rhododendron.
Material examined: INDIA, Uttarakhand, Rudraprayag
district, Baniyakund, 2630 m, N30°28.9140 E79°10.8540 ,
29 August 2015, P. Uniyal, PU 15-1004 (CAL 1332,
holotype); ibid., Baniyakund, 2630 m, N30°28.9140
E79°10.8540 , 9th Aug 2014, P. Uniyal, PU 15-360 (GUH);
ibid., 25 August 2014, P. Uniyal, PU 15-451 (GUH); ibid.,
1 August 2015, P. Uniyal, PU 15-803 (GUH).
Notes: The combination of morphological features such
as whitish dry pileus, concolourous (to pileus) stipe, light
yellow spore print, winged basidiospores, trichopalisade
pattern of pileipellis and absence of hymenial macrocystidia undoubtedly place Lactarius dirkii under L. subg.
Plinthogali (Burl.) Hesler & A.H. Sm. (Basso 1999; Das
and Sharma 2004; Stubbe et al. 2008). Among this subgenus, the new species is fairly easy to recognize in the
field by its white to orange-white basidiomata with pinkish
tinge and darker scrobicules on cap surface, yellow reaction with KOH, close to rather crowded lamellae turning
pinkish to orange on bruising, hollow stipe and abundant
white latex that remains unchanged when isolated but
becoming pinkish on cut lamellae. Micromorphologically,
occurrence of subglobose to broadly ellipsoid basidiospores with up to 2 lm high wing like ornamentations
and warts that never form a reticulum, abundant emergent
pseudocystidia and sterile lamellar edges containing
cylindrical cheiloleptocystidia are also worth mentioning.
In the field, L. dirkii can be mistaken for L. oomsisiensis
Verbeken & Halling (probably morphologically closest
species, which was reported from Papua New Guinea and
Thailand and labeled with GenBank accession numbers
EF560680 and EF560679, respectively in Fig. 176). Both
have whitish basidiomata but the latter can be distinguished
macromorphologically by distant orange-brown lamellae,
white latex drying cream on gills and mild taste (Le et al.
2007) and micromorphologically, by comparatively narrow
pileipellis (40–80 lm thick) with brown intracellular pigmentation in upper layer and absence of pileopseudocystidia in suprapellis. Lactarius dirkii is also closely related
to another Asian taxon, L. friabilis H.T. Le & Stubbe (reported from Thailand as well and labelled with GenBank
accession numbers EF560663 and EF560664 but, the latter
differs from L. dirkii in having distant (to subdistant)
lamellae,
larger
spores
(7.8–7.9–8.4–9.1 9 7.1–
7.5–7.8–8.7 lm) with incompletely reticulated ornamentation of warts and ridges with mostly crenulate edges (Le
et al. 2007). Phylogenetically, both L. oomsisiensis (95 %
123
254
identity with L. dirkii for 95 % query coverage using
BLAST) and L. friabilis (94 % identity with L. dirkii for
98 % query coverage using BLAST) are also distinct.
In L. subg. Plinthogali, molecular phylogeny already
confirmed the existence of eight European species (Stubbe
and Verbeken 2012) of which, seven species: L. acris
(Bolton: Fr.) Gray (GenBank acc. nos. JQ446084 and
JQ446083), L. azonites (Bull.) Fr. (GenBank acc. nos.
JQ446094 and JQ446095), L. fuliginosus (Fr.: Fr.) Fr.
(GenBank acc. nos. JQ446111 and JQ446110), L. picinus
Fr. (GenBank acc. nos. JQ446129 and JQ446130),
L. pterosporous Romagn. (GenBank acc. nos. JQ446138
and JQ446136), L. romagnesii Bon (GenBank acc. nos.
JQ446143 and EF560662) and L. ruginosus Romagn.
(GenBank acc. nos. JQ446106 and EF560660) are found to
be closely related (93–95 % identity with L. dirkii for
95–98 % query coverage using BLAST) to L. dirkii
(showing significant support with the Asian taxa of L. subg.
Plinthogali in Fig. 176). However, mostly these species are
dark coloured in contrast to the white basidiome of
L. dirkii, making it easily distinguishable from them in the
field. Moreover, L. acris shows a completely different
pattern of pileipellis i.e., ixooedotrichoderm to ixotrichopalisade with capitate terminal elements (HeilmannClausen et al. 1998); L. fuliginosus and L. picinus (also
reported from India) have dark coloured cap, scarce latex,
trichoepithelium pattern of pileipellis and lower (1 lm)
spore ornamentations (Heilmann-Clausen et al. 1998) and
strictly conifer association in later species (Stubbe and
Verbeken 2012). Among other close European taxa, L. romagnesii and L. ruginosus have fuscous colour of pileus,
irregularly crenate to grooved margin, distant gills and
rather sparse latex and spores with high ornamentations
that are reticulate in L. romagnesii but zebroid in L. ruginosus (Stubbe and Verbeken 2012), while L. azonites can
be separated from the present taxon by distant gills, larger
spores (7.3–9.3 9 6.8–8.3 lm), lower ornamentations (up
to 1 lm) and hyphoepithelium to trichoepithelium type of
pileipellis (Heilmann-Clausen et al. 1998). Lactarius
pterosporus shares similar pileipellis structure (a trichopalisade of 80–120 lm thick), but differs in having
dark greyish-buff coloured cap, sparse latex and higher (up
to 2.5 lm) spore ornamentations (Heilmann-Clausen et al.
1998).
Molecular data also indicates genetic closeness (95 %
identity with L. dirkii for 95 % query coverage using
BLAST) of L. dirkii with L. fumosibrunneus A.H. Sm. &
Hesler (reported from North American continent and
labelled with GenBank acc. nos. JQ797634 and JQ797633
in Fig. 176), which is different from the former in
brownish, rugose cap, negative reaction of KOH on surface
and reddening of tissue by latex. Micromorphologically,
hymenoepithelium nature of pileipellis and presence of
123
Fungal Diversity (2016) 80:1–270
abundant cheilocystidia in L. fumosibrunneus are also quite
distinct (Bandala and Montoya 2010).
Few Indian species of L. subg. Plinthogali namely,
L. crenulatus K. Das & Verbeken, L. montoyae K. Das &
J.R. Sharma, L. croceigalus K. Das & Verbeken and
L. vesterholtii K. Das & D. Chakr. partly resemble
micromorphologically (nature of pileipellis, absence of
macrocystidia, pinkish discoloration of flesh and high spore
ornamentations). But, L. crenulatus has distinctly smaller
papillate dark-coloured pileus and slender stipe (Das and
Verbeken 2012). Lactarius montoyae (also reported from
Thailand) shows subdistant to distant lamellae, brown
coloured basidiomata, white unchanging latex which does
not stain tissues (Le et al. 2007; Das and Sharma 2005).
Lactarius croceigalus differs from L. dirkii on the
basis of larger, papillate and dark colored pileus, distant lamellae and larger spores (9–9.2–10.1 9
7.8–8.7–9.5 lm) as mentioned by Das and Verbeken
(2012). Finally, Lactarius vesterholtii, which was
recently discovered from Himalayan India under L. subg.
Plinthogali, can be separated morphologically from the
present species by crowded lamellae, typically orange
discolouration of latex on exposed gills, brown to greyish-brown pileus, lack of pileopseudocystidia and showing palisade to lampropalisade nature of stipitipellis (Das
and Chakraborty 2014).
Mortierellomycotina Kerst. Hoffm. et al.
Mortierellales Caval.-Sm.
Mortierellaceae A. Fisch.
We follow Li et al. (2016).
Mortierella Coem.
The genus Mortierella (Mortierellaceae, Mortierellales)
was described by Coemans (1863) with type species
M. polycephala Coem. To date, nearly 100 species of
Mortierella have been described (Wagner et al. 2013). The
species belonging to this genus are characterized by the
production of a mainly coenocytic but becoming irregularly septate mycelium. Sporangiophores are simple or
variously branched terminating with sporangia and occasionally with a swelling at the base. Sporangia are globose,
multi-, few- or uni-spored. Species of Mortierella typically
exhibit rapid growth at temperatures ranging from 15 °C to
25 °C. They are frequently isolated from soil and dead or
dying plant tissue or from animal dung (Gams 1977; Benny
2006, 2008). Many of them show potential as producers of
polyunsaturated fatty acids (Shinmen et al. 1989; Ogawa
et al. 2012). In addition, several species of Mortierella
have been used as pesticide degrading agent, suggesting
that they might have potential for the bioremediation of
sites contaminated with organochlorine pesticides (Kataoka
et al. 2010).
Fungal Diversity (2016) 80:1–270
Based on morphological characters, Gams (1977) divided Mortierella into nine sections: Actinomortierella,
Alpina, Haplosporangium, Hygrophila, Mortierella, Schmuckeri, Simplex, Spinosa and Stylospora. Recently,
molecular data have been used to evaluate the genus
Mortierella (Petkovits et al. 2011; Wagner et al. 2013;
Ariyawansa et al. 2015a, b, c).
While examining the diversity of fungi of the order
Mortierellales isolated from freshwater sample of Yeongsan River in Gwangju, Korea, a new species was isolated
and is described here based on morphological characteristics and phylogenetic analyses.
Mortierella fluviae Hyang B. Lee, K. Voigt & T.T.T.
Nguyen, sp. nov.
MycoBank number: MB 817071; Facesoffungi number:
FoF02474, Fig. 181
Etymology: referring to the freshwater which from the
species was first isolated
Holotype: EML-YR25716-1.
Colonies reaching 45–48 mm diam. at 20 °C after
7 days incubation on PDA, cotton in the center with a
white margin, the reverse white and regularly zonate: on
OA, aerial hyphae dispersed on agar surface. Sporangiophores 115–350 lm long, 3.5–7.5 lm wide at the tip,
8.5–17 lm wide at the base, arising from the aerial stolons
with
1–4
branches
(av.
2–3).
Sporangia
19–51 9 23–52 lm, globose, multi-spored, with deliquescent wall, sometimes with a bell-shaped apophysis.
Columellae 3.5–8 9 5–10 lm, hemisphaerical or subglobose, with small projection on the apex. Collarette
appearing after sporangium maturation. Spores globose to
ellipsoidal or pyriform, 6.5–11.5 9 5.5–8.5 lm. Chlamydospores absent in aerial mycelia. Zygospores not
observed.
The isolate was observed to grow over a wide range of
temperatures with varying growth rates on PDA, OA
(oatmeal agar), and WA (water agar). The average growth
rates on PDA, OA, and WA were 6.5 mm, 6 mm, and
5 mm per 24 h, respectively. Optimal growth was observed
around 15–25 °C, maximum around 30 °C, and no growth
at 35 °C (Fig. 181).
Notes: Mortierella fluviae is similar to M. gamsii but
differs by shorter sporangiospores and having a bell-shaped
apophysis. Spores are variable in shape. In the phylogenetic trees the ITS and 28S rDNA sequences of the strain
formed a branch separate from other species of Mortierella,
showing it represents a new species (Figs. 179, 180,
respectively).
Material examined: REPUBLIC OF KOREA, Jeonnam Province, Yeongsan River located in Gwangju
(35°100 N 126°550 E), from a freshwater sample, 15
February 2016; holotype, ex-type living culture EML-
255
YR25716-1 at Culture Collection of Nakdonggang
National Institute of Biological Resources (NNIBR),
Sangju, Gyeongbuk Province, and preserved as glycerol
stock at -80 °C in the Chonnam National University
Fungal Collection (CNUFC) under deposition number
(CNUFC-EML-YR25716-1) and living culture (ex-type)
deposited at Jena Microbial Resource Collection
(University of Jena and Leibniz Institute for Natural
Product Research and Infection Biology, Jena, Germany)
(JMRC:SF:012332).
Mucoromycotina Benny
Mucorales Fr.
Cunninghamellaceae Naumov ex R.K. Benj.
Cunninghamella Matr.
Cunninghamella includes saprobes species commonly
isolated from soil, decaying fruit or rotten wood, and
some species have been responsible for disseminating
infections in humans, especially immunocompromised
patients (Yu et al. 2015). Species of Cunninghamella have
traditionally been distinguished based on their morphology, especially in their sporangial stages (Baijal and
Mehrotra 1980; Zheng and Chen 2001; Yu et al. 2015).
Since 1991, other characteristics, such as maximum
growth temperature, the color and texture of colonies,
mating compatibility, zygospore formation as well as
molecular tools have been used to delimitate the species
(Zhou and Huang 1991; Zheng and Chen 1994, 1998; Liu
et al. 2001; Gherbawy and Voigt 2010). Zheng and Chen
(2001) monographed the genus based on morphological
characteristics, maximum growth temperature and mating
experiments, as well as the entire length of the ITS
region, describing twelve species and three varieties:
C. bertholletiae Stadel, C. blakesleeana Lendn., C. binarie R.Y. Zheng, C. clavata R.Y. Zheng & G.Q. Chen,
C. echinulata var. antarctica (Caretta & Piont.) R.Y.
Zheng & G.Q. Chen, C. echinulata var. echinulata
(Thaxt.) Thaxt. ex Blakeslee, C. echinulata var. nodosa
R.Y. Zheng, C. echinulata var. verticillata (F.S. Paine)
R.Y. Zheng & G.Q. Chen, C. elegans Lendn., C. homothallica Komin. & Tubaki, C. intermedia K.B. Deshp.
& Mantri, C. multiverticillata R.Y. Zheng & G.Q. Chen,
C. phaeospora Boedijn and C. septata R.Y. Zheng. Liu
et al. (2011) and Yu et al. (2015) amplified the ITS rDNA
region and TEF-1a of all the species described by Zheng
and Chen (2001) and found their results to be consistent
with morphological studies performed by previous
authors.
Cunninghamella gigacellularis A.L. Santiago, C.L. Lima
& C.A. de Souza, sp. nov.
Index Fungorum number: IF552131; Facesoffungi
number: FoF02204, Fig. 184
123
256
Etymology: gigacellularis. A reference to the giant cells
produced in the mycelium
Holotype: URM 7400.
Colonies white, floccose, reaching 9 cm diam. and
touching the plate lid after 4 days on BDA at 25 °C.
Reverse light yellow. Rhizoids frequent, long or short,
simple or weakly branched. Stolons present, coenocytic,
some with septa near the site of origin of the sporophore.
Sporophores erect, straight or recumbent, smooth-walled,
arising from stolons or from aerial hyphae; main axes of
the sporophores usually equal in width throughout, (5–)
6.5–10(–12.5) lm diam., predominantly terminated with a
vesicle, but with some dividing in branches at the apices;
branches monopodial, verticillate and pseudoverticillate;
short and long branches (up to 500 9 7.5 lm) in the same
sporophore are common, repeatedly branching 1–6 times,
terminating with a vesicle. Some branches arise from a
bulbous base. Septa in sporophores may be present at its
base, near to the branches of the sporophore or below the
vesicles. Vesicles of the main sporophores light grey,
smooth-walled, globose and subglobose, some lightly
depressed, but rarely obovoid or piriform, (14–)20–35 lm
diam. Lateral vesicles light grey, smooth-walled, globose
and subglobose (10–)12.5–20(–25) lm diam. Sporangioles
hyaline, subsmooth to very shortly echinulate, mostly
globose 5–7.5 lm diam., some subglobose and ellipsoid
(12.5–)7.5–9 9 6–8(–10) lm, rarely with a pointed end.
The larger sporangioles are light grey to pale brown and
more echinulate than the smaller ones. Giant cells subglobose, some hypha-like and irregularly swollen, simple
or branched. Zygosporangia not observed.
Material examined: BRAZIL, Itambé: Instituto Agronômico de Pernambuco (7°970 6000 S, 35°470 8800 W), in soil
samples. Soil, 11.X.2013, leg. C. Lima (URM 7400,
holotype) and deposited in the Jena Microbial Resource
Collection (University of Jena and Leibniz Institute for
Natural Product Research and Infection Biology, Jena,
Germany) (JMRC:SF:012269).
Media and temperature tests: On BDA. At 15 °C—very
limited growth (2.6 cm in diam. in 120 h); total lack of
reproductive structures. At 20 °C—slow growth (5 cm in
120 h); good sporulation. At 25 °C—good growth (8 cm in
120 h); good sporulation. At 30 and 35 °C—excellent
growth (9 cm in 96 h); excellent sporulation. At 40 °C—
6.8 cm in 96 h; rare sporophores formed, very poor
sporulation. At 42 °C—total lack of growth and sporulation. The growth of C. gigacellularis on MEA was slightly
slower than on PDA at all tested temperatures, except at
25 °C. Abundant sterile mycelium was observed on PDA
and MEA cultures at 30, 35 and 40 °C.
Notes: Cunninghamella gigacellularis is distinguished
from other species of the genus as it produces giant cells
that have not so far been reported for this genus. The new
123
Fungal Diversity (2016) 80:1–270
species also simultaneously presents a white colony and
produces monopodially, verticillately and pseudoverticillately branched sporophores, as well as producing long and
short branches in the same repeatedly branched sporophore. At first, C. gigacellularis could be confused with
C. bertholletiae Stadel because of the branching pattern of
the sporophores. However, only the former produces giant
cells. Colonies of C. gigacellularis are persistently white,
in contrast to the drab to mouse grey colonies of
C. bertholletiae (Zheng and Chen 2001), and sporophores
of C. gigacellularis seem to be more repeatedly branched
than those of C. bertholletiae. Additionally, the sporangiospores of C. bertholletiae are 4.5–9(–11) or 5–14.5
(–16) 9 3.5–9(–11) lm (Zheng and Chen 2001), larger
than those of C. gigacellularis. Our molecular analysis
(ITS and LSU rDNA regions, Figs. 182, 183, respectively)
placed C. gigacellularis in a separate clade in ITS and LSU
trees, confirming that it is genetically different from other
species of the genus.
Acknowledgements K.D. Hyde would like to thank the Thailand
Research Fund grant no RSA5980068 entitled Biodiversity, phylogeny and role of fungal endophytes on above parts of Rhizophora apiculata and Nypa fruticans and the Chinese Academy
of Sciences, Project Number 2013T2S0030, for the award of
Visiting Professorship for Senior International Scientists at Kunming Institute of Botany. Financial support by the German Academic Exchange Service (DAAD) and the Thai Royal Golden
Ph.D. Jubilee-Industry program (RGJ) for a joint TRF-DAAD PPP
(2012–2014) academic exchange grant to K.D. Hyde and M.
Stadler, and the RGJ for a personal grant to B. Thongbai (No.
Ph.D/0138/2553 in 4.S.MF/53/A.3) is gratefully acknowledged.
Chayanard Phukhamsakda (PHD/0020/2557) acknowledges the The
Royal Golden Jubilee Ph.D. Program under the Thailand Research
Fund. Mingkwan Doilom acknowledges the Royal Golden Jubilee
Ph.D. Program (PHD./0072/2553 in 4.S.M.F./53/A.2) under the
Thailand Research Fund. Ausana Mapook is grateful to Research
and Researchers for Industries (RRI) PHD57I0012. Rungtiwa
Phookamsak sincerely appreciates The Royal Golden Jubilee Ph.
D. Program (PHD/0090/2551 in 4. S. MF/51/A.1) under the
Thailand Research Fund for financial support. Qi Zhao thanks the
National Natural Science Foundation of China (No. 31360015) and
the CAS/SAFEA International Partnership Program for Creative
Research Teams, and the Knowledge Innovation Program of the
Chinese Academy of Sciences (No. KSCX2-EW-Z-9 and
KIB2016002). KNAR acknowledges support from the University
Grants Commission (UGC), India, in the form of a Rajiv Gandhi
National Fellowship (Grant No. F. 14-2(SC)/2009 (SA-III)) and the
permissions given to him for collecting agaric specimens from the
forests of Kerala by the Principal Chief Conservator of Forests,
Government of Kerala (WL12-4042/2009 dated 05-08-2009). This
Project was funded by the National Plan for Science, Technology
and Innovation (MAARIFAH), King Abdulaziz City for Science
and Technology, Kingdom of Saudi Arabia, Award Number (12–
BIO2840–02). B.K. Cui thanked for the finance by the Fundamental Research Funds for the Central Universities (No.
2016ZCQ04) and the National Natural Science Foundation of
China (Project No. 31422001). We would like to thank Dr.
Marcela E. S. Cáceres for translating the German description of
Clavulinopsis, the Conselho Nacional de Desenvolvimento Cientı́fico (CNPq) for the master scholarship of LSAN, the Pós-
Fungal Diversity (2016) 80:1–270
Graduação em Biologia de Fungos (UFPE, Brazil), CNPq (Protax
562106/2010-3, Sisbiota 563342/2010-2, Universal 472792/2011-3)
and FACEPE (APQ-0788-2.03/12) for financing this research. H.B.
Lee was supported by the Graduate Program for the Undiscovered
Taxa of Korea, and by the Project on Survey and Discovery of
Indigenous Fungal Species of Korea, funded by NIBR and NNIBR
of the Ministry of Environment (MOE), and in part by a fund
from National Institute of Animal Science under Rural Development Administration, Republic of Korea. Aniket Ghosh, Priyanka
Uniyal and R.P. Bhatt are grateful to the Head, Department of
Botany & Microbiology, HNB Garhwal University, Srinagar
Garhwal for providing all kinds of facilities during the present
study. Kanad Das and Abhishek Baghela are thankful to the
Director, Botanical Survey of India, Kolkata and Director, MACS’
Agharkar Research Institute, Pune respectively for providing
facilities. UGC provided fellowship to Aniket Ghosh and Priyanka
Unial. Field assistance rendered by Mr. Tahir Mehmood and Mr.
Upendra Singh (HNBGU) are also duly acknowledged. Tuula
Niskanen, Kare Liimatainen, Ilkka Kytövuori, Joe Ammirati,
Bálint Dima, and Dimitar Bojantchev would like to acknowledge
Heino Vänskä for the help with nomenclature. We are grateful to
the curators of H and S. This work was partially supported by the
Ministry of Environment, Finland (YM38/5512/2009) and Oskar
Öflunds Stiftelse. The authors thanks Dr. Kerstin Voigt for the
inestimable help in critical reviewing the lower fungi entries, the
Coordenação de Aperfeiçoamento de Pessoal de Nı́vel Superior
(CAPES) and Fundação de Amparo à Ciência e Tecnologia do
Estado de Pernambuco (FACEPE) for the postgraduate scholarships to Diogo X. Lima and Carlos A. F. de Souza, respectively.
We also thank Conselho Nacional de Desenvolvimento Cientı́fico e
Tecnológico (CNPq) and FACEPE for financial support through
the projects: ‘Mucoromycotina in upland forests from the semi-arid
of Pernambuco’ (CNPq—458391/2014-0), and ‘Diversity of
Mucoromycotina in different ecosystems of the Pernambuco’s
Atlantic Rainforest’ (FACEPE—APQ 0842-2.12/14). Z.L Luo and
H.Y Su would like to thank the National Natural Science Foundation of China (Project ID: 31460015) for financial support on
Study of the distribution pattern and driving factors of aquatic
fungal diversity in the region of Three Parallel Rivers.
C. Phukhamsakda would like to thank Dr. Matthew P. Nelsen for
his valuable suggestions. Saranyaphat Boonmee thanks to the
Thailand Research Fund, project number TRG5880152 and Mae
Fah Luang University for a Grant Number 2559A30702006’’. C.G.
Lin and Y. Wang thank for the finance by the National Natural
Science Foundation of China (No. NSFC 31560489) and Fundamental Research on Science and Technology, Ministry of Science
and Technology of China (2014FY120100). Haixia Wu would like
to thank Dr. Shaun Pennycook for his kindly nomenclatural review
and thanked for the finance by the National Natural Science
Foundation of China (Project No. 31300019). S.C. Karunarathna,
P.E. Mortimer and J.C. Xu would like to thank the World
Agroforestry Centre, East and Central Asia Office; Key Laboratory
for Plant Diversity and Biogeography of East Asia, Kunming
Institute of Botany, Chinese Academy of Science; the Chinese;
Ministry of Science and Technology, under the 12th 5-year
National Key Technology Support Program (NKTSP)2013
BAB07B06 integration and comprehensive demonstration of key
technologies on Green Phosphate-mountaion Construction and the
CGIAR Research Program 6: Forest, Trees and Agroforestry for
partial funding. The National Research Council of Thailand
(NRCT), projects—Taxonomy, phylogeny and cultivation of
Lentinus species in northern Thailand (NRCT/55201020007) is
also thanked. K. Tanaka and A. Hashimoto would like to thank
the Japan Society for the Promotion of Science (JSPS; 26291084,
16K07474, 16J07243).
257
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