Fungal Diversity
DOI 10.1007/s13225-017-0378-0
Fungal diversity notes 491–602: taxonomic and phylogenetic
contributions to fungal taxa
Saowaluck Tibpromma1,2,3,4,5 • Kevin D. Hyde1,2,3,4,5 • Rajesh Jeewon29 •
Sajeewa S. N. Maharachchikumbura25 • Jian-Kui Liu18 • D. Jayarama Bhat9,10 •
E. B. Gareth Jones11 • Eric H. C. McKenzie12 • Erio Camporesi6,7,8 •
Timur S. Bulgakov27 • Mingkwan Doilom2 • André Luiz Cabral Monteiro de Azevedo Santiago15
Kanad Das34 • Patinjareveettil Manimohan33 • Tatiana B. Gibertoni42 •
Young Woon Lim30 • Anusha Hasini Ekanayaka2 • Benjarong Thongbai2 •
Hyang Burm Lee17 • Jun-Bo Yang55 • Paul M. Kirk60 • Phongeun Sysouphanthong53 •
Sanjay K. Singh22 • Saranyaphat Boonmee2 • Wei Dong20 • K. N. Anil Raj33 •
K. P. Deepna Latha33 • Rungtiwa Phookamsak1,2,4 • Chayanard Phukhamsakda1,2,3,4 •
Sirinapa Konta2,3,5 • Subashini C. Jayasiri2,3,5 • Chada Norphanphoun2,3,5 •
Danushka S. Tennakoon2,3,5 • Junfu Li2,3,5 • Monika C. Dayarathne2,3,5 •
Rekhani H. Perera2,3,5 • Yuanpin Xiao2,3,5 • Dhanushka N. Wanasinghe1,2,3,4,5 •
Indunil C. Senanayake1,2,3,4,5 • Ishani D. Goonasekara1,2,3,4,5 • N. I. de Silva1,2,4,13 •
Ausana Mapook2,3 • Ruvishika S. Jayawardena2,16 • Asha J. Dissanayake2,16 •
Ishara S. Manawasinghe2,16 • K. W. Thilini Chethana2,16 • Zong-Long Luo2,19 •
Kalani Kanchana Hapuarachchi2,3,28 • Abhishek Baghela22 • Adriene Mayra Soares42 •
Alfredo Vizzini23,40 • Angelina Meiras-Ottoni42 • Armin Mešić46 • Arun Kumar Dutta31 •
Carlos Alberto Fragoso de Souza15 • Christian Richter58 • Chuan-Gen Lin2,3,5,59 •
Debasis Chakrabarty48 • Dinushani A. Daranagama2,3,5 • Diogo Xavier Lima15 •
& Samantha C. Karunarathna
samanthakarunarathna@gmail.com
1
Key Laboratory for Plant Diversity and Biogeography of East
Asia, Kunming Institute of Botany, Chinese Academy of
Science, Kunming 650201, Yunnan, People’s Republic of
China
2
Center of Excellence in Fungal Research, Mae Fah Luang
University, Chiang Rai 57100, Thailand
3
School of Science, Mae Fah Luang University,
Chiang Rai 57100, Thailand
4
World Agroforestry Centre, East and Central Asia,
Kunming 650201, Yunnan, People’s Republic of China
5
Mushroom Research Foundation, 128 M.3 Ban Pa Deng T. Pa
Pae, A. Mae Taeng, Chiang Mai 50150, Thailand
6
A.M.B. Gruppo Micologico Forlivese ‘‘Antonio Cicognani’’,
Via Roma 18, Forlı̀, Italy
7
A.M.B. Circolo Micologico ‘‘Giovanni Carini’’,
C.P. 314, Brescia, Italy
8
Società per gli Studi Naturalistici della Romagna,
C.P. 144, Bagnacavallo, RA, Italy
9
Formerly Department of Botany, Goa University, Taleigão,
Goa, India
10
No. 128/1-J, Azad Housing Society, Curca, Goa Velha,
India
•
11
33 B St. Edwards Road Southsea Hants,
Hampshire PO5 3DH, UK
12
Landcare Research Manaaki Whenua,
Private Bag 92170, Auckland, New Zealand
13
Department of Biology, Faculty of Science, Chiang Mai
University, Chiang Mai 50200, Thailand
14
Division of Applied Science, College of International
Education, The Hong Kong Baptist University,
Hong Kong SAR, China
15
PostGraduate Program in Biology of Fungi, Department of
Mycology, Federal University of Pernambuco, Av. Nelson
Chaves, s/n, Recife, PE 50670-420, Brazil
16
Institute of Plant and Environment Protection, Beijing
Academy of Agriculture and Forestry Sciences,
Beijing 100097, People’s Republic of China
17
Environmental Microbiology Lab, Division of Food
Technology, Biotechnology & Agrochemistry, College of
Agriculture and Life Sciences, Chonnam National University,
Yongbong-Dong 300, Buk-Gu, Gwangju 61186, Korea
18
Guizhou Key Laboratory of Agricultural Biotechnology,
Guizhou Academy of Agricultural Sciences,
Guiyang 550006, Guizhou, People’s Republic of China
19
College of Agriculture and Biological Sciences, Dali
University, Dali 671003, Yunnan, People’s Republic of
China
123
Fungal Diversity
Dyutiparna Chakraborty34 • Enrico Ercole40 • Fang Wu26 • Giampaolo Simonini38 •
Gianrico Vasquez37 • Gladstone Alves da Silva15 • Helio Longoni Plautz Jr.44 •
Hiran A. Ariyawansa61 • Hyun Lee30 • Ivana Kušan46 • Jie Song26 •
Jingzu Sun2,3,21 • Joydeep Karmakar36 • Kaifeng Hu54 • Kamal C. Semwal47 •
Kasun M. Thambugala2,3,5,18 • Kerstin Voigt59 • Krishnendu Acharya31 •
Kunhiraman C. Rajeshkumar22 • Leif Ryvarden45 • Margita Jadan46 •
Md. Iqbal Hosen51 • Michal Mikšı́k41 • Milan C. Samarakoon2,4,5,13 •
Nalin N. Wijayawardene2 • Nam Kyu Kim30 • Neven Matočec46 • Paras Nath Singh22 •
Qing Tian2,3,5 • R. P. Bhatt49 • Rafael José Vilela de Oliveira15 • Rodham E. Tulloss50 •
S. Aamir22 • Saithong Kaewchai24 • Sayali D. Marathe22 • Sehroon Khan1,4 •
Sinang Hongsanan2 • Sinchan Adhikari35 • Tahir Mehmood49 • Tapas Kumar Bandyopadhyay36
Tatyana Yu. Svetasheva39 • Thi Thuong Thuong Nguyen17 • Vladimı́r Antonı́n32 •
Wen-Jing Li1,2,3,5 • Yong Wang59 • Yuvraj Indoliya48 • Zdenko Tkalčec46 •
Abdallah M. Elgorban57 • Ali H. Bahkali57 • Alvin M. C. Tang14 • Hong-Yan Su19 •
Huang Zhang20 • Itthayakorn Promputtha13 • Jennifer Luangsa-ard56 •
Jianchu Xu1,4 • Jiye Yan16 • Kang Ji-Chuan28 • Marc Stadler58 • Peter E. Mortimer1,4 •
Putarak Chomnunti2,3 • Qi Zhao52 • Alan J. L. Phillips43 • Sureeporn Nontachaiyapoom3 •
Ting-Chi Wen28 • Samantha C. Karunarathna1,4
•
Received: 6 March 2017 / Accepted: 20 March 2017
Ó School of Science 2017
Abstract This is a continuity of a series of taxonomic and
phylogenetic papers on the fungi where materials were
collected from many countries, examined and described. In
addition to extensive morphological descriptions and
appropriate asexual and sexual connections, DNA
20
21
22
Yunnan Institute of Food Safety, Kunming University
of Science & Technology, Kunming 650500, Yunnan,
People’s Republic of China
State Key Laboratory of Mycology, Institute of
Microbiology, Chinese Academy of Sciences (IMCAS),
No. 3 Park 1, West Beichen Road, Chaoyang District,
Beijing 100101, People’s Republic of China
National Fungal Culture Collection of India, Biodiversity and
Palaeobiology Group, MACS Agharkar Research Institute,
G.G. Agarkar Road, Pune 411 004, India
sequence data are also analysed from concatenated datasets
to infer phylogenetic relationships and substantiate systematic positions of taxa within appropriate ranks. Wherever new species or combinations are proposed, we apply an
integrative approach using morphological and molecular
29
Department of Health Sciences, Faculty of Science,
University of Mauritius, Reduit 80837, Mauritius
30
School of Biological Sciences, Seoul National University,
Seoul 08826, Korea
31
Molecular and Applied Mycology and Plant Pathology
Laboratory, Department of Botany, University of Calcutta,
Kolkata, West Bengal 700019, India
32
Moravian Museum, Department of Botany, Zelný trh 6,
659 37 Brno, Czech Republic
23
Institute for Sustainable Plant Protection (IPSP)-CNR,
Viale P.A. Mattioli 25, 10125 Turin, Italy
33
Department of Botany, University of Calicut, Malappuram,
Kerala 673 635, India
24
Faculty of Agriculture, Princess of Naradhiwas University,
No. 99 Khok Khian, Amphur Muang, Narathiwat Province,
Thailand
34
Botanical Survey of India, Cryptogamic Unit,
P.O. Botanic Garden, Howrah 711103, India
35
25
Department of Crop Sciences, College of Agricultural and
Marine Sciences, Sultan Qaboos University, P.O. Box 34,
123 Al-Khod, Oman
Department of Botany, University of Kalyani,
Kalyani, Nadia, West Bengal 741235, India
36
Department of Molecular Biology & Biotechnology,
University of Kalyani, Kalyani, Nadia, West Bengal 741235,
India
37
Laboratorio di Isto-Cito-Patologia s.n.c., Via del Bosco, 96,
95125 Catania, Italy
38
Via Bell’Aria, 8, 42121 Reggio nell’Emilia (RE), Italy
39
Biology and Technologies of Living Systems Department,
Tula State Lev Tolstoy Pedagogical University, 125, Lenin
av., Tula, Russia 300026
26
Institute of Microbiology, Beijing Forestry University,
PO Box 61, Beijing 100083, China
27
Russian Research Institute of Floriculture and Subtropical
Crops, 2/28 Yana Fabritsiusa Street, Sochi, Krasnodar region,
Russia 354002
28
Engineering Research Center of Southwest
Bio-Pharmaceutical Resources, Ministry of Education,
Guizhou University, Guiyang 550025, Guizhou,
People’s Republic of China
123
Fungal Diversity
data as well as ecological features wherever applicable.
Notes on 112 fungal taxa are compiled in this paper
including Biatriosporaceae and Roussoellaceae, Didysimulans gen. nov., 81 new species, 18 new host records and
new country records, five reference specimens, two new
combinations, and three sexual and asexual morph reports.
The new species are Amanita cornelii, A. emodotrygon,
Angustimassarina alni, A. arezzoensis, A. italica,
A. lonicerae, A. premilcurensis, Ascochyta italica, A. rosae,
Austroboletus appendiculatus, Barriopsis thailandica,
Berkleasmium ariense, Calophoma petasitis, Camarosporium laburnicola, C. moricola, C. grisea, C. ossea,
C. paraincrustata, Colletotrichum sambucicola, Coprinopsis cerkezii, Cytospora gelida, Dacrymyces chiangraiensis,
Didysimulans italica, D. mezzanensis, Entodesmium italica,
Entoloma magnum, Evlachovaea indica, Exophiala italica,
Favolus gracilisporus, Femsjonia monospora, Fomitopsis
flabellata, F. roseoalba, Gongronella brasiliensis, Helvella
crispoides, Hermatomyces chiangmaiensis, H. chromolaenae, Hysterium centramurum, Inflatispora caryotae, Inocybe brunneosquamulosa, I. luteobrunnea, I. rubrobrunnea,
Keissleriella cirsii, Lepiota cylindrocystidia, L. flavocarpa,
L. maerimensis, Lophiotrema guttulata, Marasmius luculentus, Morenoina calamicola, Moelleriella thanathonensis,
Mucor stercorarius, Myrmecridium fluviae, Myrothecium
septentrionale, Neosetophoma garethjonesii, Nigrograna
cangshanensis, Nodulosphaeria guttulatum, N. multiseptata, N. sambuci, Panus subfasciatus, Paraleptosphaeria
padi, Paraphaeosphaeria viciae, Parathyridaria robiniae,
Penicillium
punicae,
Phaeosphaeria
calamicola,
Phaeosphaeriopsis yuccae, Pleurophoma italica, Polyporus
brevibasidiosus, P. koreanus, P. orientivarius, P. parvovarius, P. subdictyopus, P. ulleungus, Pseudoasteromassaria spadicea, Rosellinia mearnsii, Rubroboletus
demonensis, Russula yanheensis, Sigarispora muriformis,
Sillia italica, Stagonosporopsis ailanthicola, Strobilomyces
longistipitatus, Subplenodomus galicola and Wolfiporia
pseudococos. The new combinations are Melanomma populina and Rubroboletus eastwoodiae. The reference specimens
are
Cookeina
tricholoma,
Gnomoniopsis
sanguisorbae, Helvella costifera, Polythrincium trifolii and
Russula virescens. The new host records and country
records are Ascochyta medicaginicola, Boletellus emodensis, Cyptotrama asprata, Cytospora ceratosperma, Favolaschia auriscalpium, F. manipularis, Hysterobrevium
mori, Lentinus sajor-caju, L. squarrosulus, L. velutinus,
Leucocoprinus cretaceus, Lophiotrema vagabundum,
Nothophoma quercina, Platystomum rosae, Pseudodidymosphaeria phlei, Tremella fuciformis, Truncatella spartii
and Vaginatispora appendiculata and three sexual and
40
50
P. O. Box 57, Roosevelt, NJ 08555-0057, USA
51
State Key Laboratory of Applied Microbiology Southern
China, Guangdong Provincial Key Laboratory of Microbial
Culture Collection and Application, Guangdong Institute of
Microbiology, Guangzhou 510070, People’s Republic of
China
52
Key Lab of Southwestern Crop Gene Resources and
Germplasm Innovation Ministry of Agriculture, Yunnan
Provincial Key Laboratory of Agricultural Biotechnology,
Institute of Biotechnology and Germplasm Resources,
Yunnan Academy of Agricultural Sciences,
Kunming 650223, Yunnan, People’s Republic of China
53
Ecology Division, Biotechnology and Ecology Institute,
Ministry of Science and Technology,
P.O.Box: 2279, Vientiane Capital, Lao People’s Democratic
Republic
54
State Key Laboratory of Phytochemistry and Plant Resources
in West China, Kunming, People’s Republic of China
55
Germplasm Bank of Wild Species in Southwest China,
Kunming Institute of Botany, Chinese Academy of Science,
Kunming 650201, Yunnan, People’s Republic of China
56
Microbe Interaction Laboratory (BMIT), BIOTEC, National
Science and Technology Development Agency (NSTDA),
113 Thailand Science Park, Thanon Phahonyothin, Tombon
Khlong Nueng, Amphoe, Khlong Luang,
Pathum Thani 12120, Thailand
Department of Life Sciences and Systems Biology,
University of Turin, Viale P.A. Mattioli, 25,
10125 Turin (TO), Italy
41
Lečkova 1521/15, 14900 Prague 4, Czech Republic
42
Universidade Federal de Pernambuco (UFPE), Centro de
Biociências (CB), Departamento de Micologia, Avenida da
Engenharia, S/N - Cidade Universitária, Recife,
PE 50740-600, Brazil
43
University of Lisbon, Faculty of Sciences, Biosystems and
Integrative Sciences Institute (BioISI), Campo Grande,
1749-016 Lisbon, Portugal
44
Instituto de Ciências Biológicas, Laboratório de
Microbiologia, Av. Augusto Corrêa, Belém, Pará 66075-110,
Brazil
45
Department of Botany, University of Oslo, Blindern,
P. O. Box 1045, 0316 Oslo, Norway
46
Rud̄er Bošković Institute, Bijenička cesta 54, 10000 Zagreb,
Croatia
47
Department of Biology, College of Sciences, Eritrea Institute
of Technology, P. Box 12676, Mai Nafhi, Asmara, Eritrea
48
Genetics and Molecular Biology Division, CSIR-National
Botanical Research Institute,
Lucknow 226001, Uttar Pradesh, India
49
Department of Botany & Microbiology, H.N.B. Garhwal
University, Srinagar, Garhwal 246174, Uttarakhand, India
123
Fungal Diversity
asexual morphs are Aposphaeria corallinolutea, Dothiora
buxi and Hypocrella calendulina.
Keywords Agaricomycetes Ascomycota
Basidiomycota Dacrymycetes Dothideomycetes
Eurotiomycetes New combination Mucoromycotina
New genus New records New species Pezizomycetes
Phylogeny Sordariomycetes Taxonomy
Table of contents
Ascomycota
Dothideomycetes
Asterinales M.E. Barr ex D. Hawksw. & O.E. Erikss.
Asterinaceae Hansf.
Asterinales genera incertae sedis
491. Morenoina calamicola Konta & K.D. Hyde, in Fungal Diversity 83: 10 (2017), new species
Botryosphaeriales C.L. Schoch, Crous & Shoemaker
Botryosphaeriaceae Theiss. & Syd
492. Barriopsis thailandica Dissanayake, Senan., & K.D.
Hyde, in Fungal Diversity 83: 13 (2017), new species
Hysteriales Lindau
Hysteriaceae Chevall.
495. Hysterium centramurum Senan., in Fungal Diversity
83: 22 (2017), new species
496. Hysterobrevium mori (Schwein.) E.W.A. Boehm &
C.L. Schoch, Stud. Mycol 64: 62 (2009), new host record
Pleosporales Luttr. ex M.E. Barr
Amorosiaceae Thambugala, K. D. Hyde
497. Angustimassarina alni Jayasiri & K.D. Hyde, in
Fungal Diversity 83: 26 (2017), new species
498. Angustimassarina arezzoensis Tibpromma, Camporesi & K.D. Hyde, in Fungal Diversity 83: XX (2017),
new species
499. Angustimassarina premilcurensis Tibpromma,
Camporesi & K.D. Hyde, in Fungal Diversity 83: 29
(2017), new species
500. Angustimassarina italica Tibpromma, Camporesi &
K.D. Hyde, in Fungal Diversity 83: 33 (2017), new species
501. Angustimassarina lonicerae Tibpromma, Camporesi
& K.D. Hyde, in Fungal Diversity 83: 28 (2017), new
species
502. Biatriosporaceae K.D. Hyde
Capnodiales Woron
Mycosphaerellaceae Lindau
493. Polythrincium trifolii Kunze, in Kunze & Schmidt,
Mykologische Hefte (Leipzig) 1: 14 (1817), reference
specimen
Dothideales Lindau
Dothioraceae Theiss. & Syd.
494. Dothiora buxi Jayasiri, Camporesi & K.D. Hyde, in
Fungal Diversity 81: 30 (2016), asexual morph report
57
Department of Botany and Microbiology, College of Science,
King Saud University, P.O. Box: 2455, Riyadh 1145, Saudi
Arabia
58
Department of Microbial Drugs, Helmholtz Centre for
Infection Research and German Centre for Infection Research
(DZIF), Partner Site Hannover/Braunschweig,
Inhoffenstrasse 7, 38124 Braunschweig, Germany
59
Department of Plant Pathology, College of Agriculture,
Guizhou University, Guiyang 550025, Guizhou, People’s
Republic of China
60
Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3DS,
UK
61
Department of Plant Pathology and Microbiology, National
Taiwan University, No. 1, Sec. 4, Roosevelt Road,
Taipei 106, Taiwan, People’s Republic of China
123
Lophiostomataceae Sacc.
503. Berkleasmium ariense Rajeshkumar & Marathe, in
Fungal Diversity 83: 35 (2017), new species
504. Platystomum rosae Wanas., Thambug., Camporesi &
K.D. Hyde, Fungal Diversity 74: 234 (2015), new host
record
505. Sigarispora muriformis Tibpromma, Camporesi &
K.D. Hyde, in Fungal Diversity 83: 37 (2017), new species
506. Vaginatispora appendiculata Wanas., E.B.G. Jones &
K.D. Hyde, Studies in Fungi 1 (1): 60 (2016), new host
record
Lophiotremataceae K. Hiray. & Kaz. Tanaka.
507. Lophiotrema guttulata Boonmee, Tibpromma & K.D.
Hyde, in Fungal Diversity 83: 40 (2017), new species
508. Lophiotrema vagabundum (Sacc.) Sacc., Michelia 1
(4): 338 (1878), new host record
509. Hermatomyces chiangmaiensis J.F. Li, Bhat & K.D.
Hyde, in Fungal Diversity 83: 45 (2017), new species
510. Hermatomyces chromolaenae J.F. Li, Mapook &
K.D. Hyde, in Fungal Diversity 83: 47 (2017), new
species
Melanommataceae G. Winter.
511. Melanomma populina (Died.) Phukhamsakda &
K.D. Hyde, in Fungal Diversity 83: 49 (2017), new
combination
Fungal Diversity
512. Aposphaeria corallinolutea Gruyter, Aveskamp &
Verkley, Stud. Mycol 75: 28 (2012), sexual morph report
Nigrogranaceae Jaklitsch & Voglmayr
513. Nigrograna cangshanensis Z.L. Luo, H.Y. Su & K.D.
Hyde, in Fungal Diversity 83: 52 (2017), new species
514. Roussoellaceae J.K. Liu, Phook., D.Q. Dai & K.D.
Hyde Q. Tian & K.D. Hyde
Thyridariaceae Q. Tian & K.D. Hyde
515. Parathyridaria robiniae Mapook, Camporesi & K.D.
Hyde, in Fungal Diversity 83: 52 (2017), new species
Sub order Massarineae
Didymosphaeriaceae Munk.
516. Paraphaeosphaeria viciae de Silva, Camporesi &
K.D. Hyde, in Fungal Diversity 83: 57 (2017), new species
Latoruaceae Crous
517. Pseudoasteromassaria spadicea W. Dong, H. Zhang
& K.D. Hyde, in Fungal Diversity 83: 59 (2017), new
species
Lentitheciaceae Y. Zhang ter, C.L. Schoch, J. Fourn.,
Crous & K.D. Hyde.
518. Keissleriella cirsii R.H. Perera, Bulgakov, Wanasinghe & K.D. Hyde, in Fungal Diversity 83: 62 (2017), new
species
519. Pleurophoma italica Tibpromma, Camporesi & K.D.
Hyde, in Fungal Diversity 83: 62 (2017), new species
Massarinaceae Munk
520. Pseudodidymosphaeria phlei Phukhams., Camporesi
& K.D. Hyde, Fungal Diversity 78: 57 (2016), new host
record
Massarineae genera incertae sedis
521. Inflatispora caryotae Wanasinghe & K.D. Hyde, in
Fungal Diversity 83: 68 (2017), new species
Suborder Pleosporineae
Didymellaceae Gruyter, Aveskamp & Verkley
522. Ascochyta italica Tibpromma, Camporesi & K.D.
Hyde, in Fungal Diversity 83: 70 (2017), new species
523. Ascochyta medicaginicola Q. Chen & L. Cai, Stud.
Mycol. 82: 187 (2015), new host record
524. Ascochyta rosae Tibpromma, Camporesi & K.D.
Hyde, in Fungal Diversity 83: 74 (2017), new species
525. Calophoma petasitis Tibpromma, Camporesi &
K.D. Hyde, in Fungal Diversity 83: 74 (2017), new
species
526. Didysimulans Tibpromma, Camporesi & K.D. Hyde,
in Fungal Diversity 83: 76 (2017), new genus
527. Didysimulans italica Tibpromma, Camporesi & K.D.
Hyde, in Fungal Diversity 83: 78 (2017), new species
528. Didysimulans mezzanensis Tibpromma, Camporesi &
K.D. Hyde, in Fungal Diversity 83: 78 (2017), new
species
529. Nothophoma quercina (Syd. & P. Syd.) Q. Chen & L.
Cai, Stud. Mycol. 82: 213 (2015), new host record
530. Stagonosporopsis ailanthicola Manawasinghe, Camporesi & K.D. Hyde, in Fungal Diversity 83: 81 (2017),
new species
Leptosphaeriaceae M.E. Barr
531. Paraleptosphaeria padi Phukhamsakda, Bulgakov, &
K.D. Hyde, in Fungal Diversity 83: 82 (2017), new
species
532. Subplenodomus galicola Phukhamsakda, Tibpromma, Camporesi & K.D. Hyde, in Fungal Diversity 83:
86 (2017), new species
Phaeosphaeriaceae M.E. Barr.
533. Entodesmium italica Tibpromma, Camporesi & K.D.
Hyde, in Fungal Diversity 83: 88 (2017), new species
534. Neosetophoma garethjonesii Tibpromma, EBG Jones
& K.D. Hyde, in Fungal Diversity 83: 88 (2017), new
species
535. Nodulosphaeria guttulatum Tibpromma, Camporesi
& K.D. Hyde, in Fungal Diversity 83: 92 (2017), new
species
536. Nodulosphaeria multiseptata Tibpromma, Camporesi
& K.D. Hyde, in Fungal Diversity 83: 92 (2017), new
species
537. Nodulosphaeria sambuci Tibpromma, Camporesi &
K.D. Hyde, in Fungal Diversity 83: 93 (2017), new
species
538. Phaeosphaeria calamicola Konta & K. D. Hyde, in
Fungal Diversity 83: 96 (2017), new species
539. Phaeosphaeriopsis yuccae Dayarathne, Bulgakov,
E.B.G. Jones & K.D. Hyde, in Fungal Diversity 83: 96
(2017), new species
Pleosporineae genera incertae sedis
540. Camarosporium laburnicola R.H. Perera, Bulgakov,
& K.D. Hyde, in Fungal Diversity 83: 99 (2017), new
species
541. Camarosporium moricola Chethana, Bulgakov & K.
D. Hyde, in Fungal Diversity 83: 101 (2017), new species
123
Fungal Diversity
Eurotiomycetes
Eurotiales G.W. Martin ex Benny & Kimbr.
Trichocomaceae E. Fisch.
542. Penicillium punicae Hyang B. Lee, P.M. Kirk &
T.T.T. Nguyen, in Fungal Diversity 83: 104 (2017), new
species
543. Exophiala italica Tibpromma, Camporesi & K.D.
Hyde, in Fungal Diversity 83: 106 (2017), new species
Mucoromycotina Benny
Mucorales Fr.
Cunninghamellaceae Naumov ex R.K. Benj.
544. Gongronella brasiliensis C.A. de Souza, D.X. Lima
& A.L. Santiago, in Fungal Diversity 83: 110 (2017), new
species
Rhizopodaceae K. Schum.
545. Mucor stercorarius Hyang B. Lee, P.M. Kirk, K.
Voigt & T.T.T. Nguyen, in Fungal Diversity 83: 111
(2017), new species
Sordariomycetes
Diaporthales Nannf.
Gnomoniaceae G. Winter
546. Gnomoniopsis sanguisorbae (Rehm) D.M. Walker,
Mycologia 102 (6): 1494 (2010), reference specimen
Sydowiellaceae Lar.N. Vassiljeva
547. Sillia italica de Silva, Camporesi & K.D. Hyde, in
Fungal Diversity 83: 117 (2017), new species
Valsaceae Tul. & C. Tul
548. Cytospora gelida Norphanphoun, Bulgakov, T.C.
Wen & K.D. Hyde, in Fungal Diversity 83: 120 (2017),
new species
549. Cytospora ceratosperma (Tode) G.C. Adams &
Rossman, in Rossman, Adams, Cannon, Castlebury &
Crous, IMA Fungus 6: 147 (2015), new record
Stachybotryaceae L. Lombard & Crous
552. Myrothecium septentrionale J.F. Li, Phookamsak &
K.D. Hyde, in Fungal Diversity 83: 129 (2017), new species
Glomerellales Chadef. ex Réblová et al.
Glomerellaceae Locq. ex Seifert & W. Gams
553. Colletotrichum sambucicola Jayawardena, Camporesi & K.D. Hyde, in Fungal Diversity 83: 131 (2017),
new species
Myrmecridiales Crous
554. Myrmecridium fluviae Hyang B. Lee & T.T.T.
Nguyen, in Fungal Diversity 83: 136 (2017), new species
Xylariales Nannf.
Bartaliniaceae Wijayawardene et al.
555. Truncatella spartii Senan., Camporesi & K.D. Hyde,
Fungal Diversity 73: 91 (2015), new host record
Xylariaceae Tul. & C. Tul.
556. Rosellinia mearnsii Tennakoon, Phookamsak & K.D.
Hyde, in Fungal Diversity 83: 139 (2017), new species
Ascomycota, families incertae sedis
Evlachovaea B.A. Borisov & Tarasov
557. Evlachovaea indica P.N. Singh, A. Baghela, S.K. Singh
& S. Amir, in Fungal Diversity 83: 139 (2017), new species
Pezizomycetes
Pezizales J. Schröt.
Helvellaceae Fr.
558. Helvella costifera Nannf., Schriften der NaturforschendeGesellschaft zu Leipzig 1: 114 (1953), reference specimen
559. Helvella crispoides Q. Zhao & K.D. Hyde, in Fungal
Diversity 83: 147 (2017), new species
Sarcoscyphaceae Le Gal ex Eckblad
560. Cookeina tricholoma (Mont.) Kuntze, Revis. gen. pl.
(Leipzig) 2: 849 (1891), reference specimen
Basidiomycota
Hypocreales (Lindau) Earle ex Rogerson
Agaricomycetes Doweld
Clavicipitaceae Earle.
550. Hypocrella calendulina Hywel-Jones & Mongkolsamrit, Mycol. Res. 113 (6–7): 687 (2009), asexual morph
reported
551. Moelleriella thanathonensis Y.P. Xiao, T.C. Wen, &
K.D. Hyde, in Fungal Diversity 83: 125 (2017), new
species
123
Agaricales Underw.
Agaricaceae Chevall.
561. Amanita cornelii Mehmood, K. Das, Iqbal Hosen,
Tulloss, & R.P. Bhatt, in Fungal Diversity 83: 152 (2017),
new species
Fungal Diversity
562. Amanita emodotrygon Mehmood, Tulloss, K. Das,
Iqbal Hosen & R.P. Bhatt, in Fungal Diversity 83: 157
(2017), new species
563. Lepiota cylindrocystidia Sysouphanthong K.D Hyde
& Vellinga, in Fungal Diversity 83: 160 (2017), new
species
564. Lepiota flavocarpa Sysouphanthong, K.D. Hyde &
Vellinga, in Fungal Diversity 83: 162 (2017), new species
565. Lepiota maerimensis Sysouphanthong K.D Hyde, &
Vellinga, in Fungal Diversity 83: 163 (2017), new species
566. Leucocoprinus cretaceus (Bull.) Locq., Bull. mens.
Soc. linn. Soc. Bot. Lyon 14: 93 (1945), new record
Entolomataceae Kotl. & Pouzar
567. Entoloma magnum K.N.A. Raj & Manim, in Fungal
Diversity 83: 168 (2017), new species
Marasmiaceae Roze ex Kühner.
568. Inocybe brunneosquamulosa K.P.D. Latha & Manim,
in Fungal Diversity 83: 172 (2017), new species
569. Inocybe luteobrunnea K.P.D. Latha & Manim, in
Fungal Diversity 83: 176 (2017), new species
570. Inocybe rubrobrunnea K.P.D. Latha & Manim, in
Fungal Diversity 83: 178 (2017), new species
571. Marasmius luculentus A.K. Dutta, K. Acharya &
Antonı́n, in Fungal Diversity 83: 181 (2017), new species
Mycenaceae Overeem
572. Favolaschia auriscalpium (Mont.) Henn., Botanische
Jahrbücher für Systematik Pflanzengeschichte und
Pflanzengeographie 22: 93 (1895), new record
573. Favolaschia manipularis (Berk.) Teng, Zhong Guo
De Zhen Jun [Fungi of China]: 760 (1963), new record
Physalacriaceae Corner
574. Cyptotrama asprata (Berk.) Redhead & Ginns,
Canadian Journal of Botany 58 (6): 731 (1980), new record
Boletales E.-J. Gilbert
Boletaceae Chevall.
575. Austroboletus appendiculatus Semwal, D. Chakr., K.
Das, Indoliya, D. Chakrabarty, S. Adhikari & Karunarathna, in Fungal Diversity 83: 189 (2017), new species
576. Boletellus emodensis (Berk.) Singer, Annales Mycologici 40: 19 (1942), new record
577. Rubroboletus demonensis Vasquez, Simonini, Svetasheva, Mikšı́k & Vizzini, in Fungal Diversity 83: 197
(2017), new species
578. Rubroboletus eastwoodiae (Murrill) Vasquez, Simonini, Svetasheva, Mikšı́k & Vizzini, in Fungal Diversity 83:
206 (2017), new combination
579. Strobilomyces longistipitatus D. Chakr., K. Das & S.
Adhikari, in Fungal Diversity 83: 207 (2017), new species
Cantharellales Gäum.
Clavulinaceae Donk
580. Clavulina grisea Meiras-Ottoni & Gibertoni, in
Fungal Diversity 83: 211 (2017), new species
581. Clavulina ossea Meiras-Ottoni & Gibertoni, in Fungal
Diversity 83: 211 (2017), new species
582. Clavulina paraincrustata Meiras-Ottoni & Gibertoni,
in Fungal Diversity 83: 212 (2017), new species
Polyporales Gäum.
Fomitopsidaceae Jülich
583. Fomitopsis flabellata Soares & Gibertoni, in Fungal
Diversity 83: 215 (2017), new species
584. Fomitopsis roseoalba Soares, Ryvarden & Gibertoni,
in Fungal Diversity 83: 215 (2017), new species
Polyporaceae Fr. ex Corda
585. Favolus gracilisporus H. Lee, N.K. Kim & Y.W.
Lim, in Fungal Diversity 83: 219 (2017), new species
586. Lentinus sajor-caju (Fr.) Fr., Epicrisis Systematis
Mycologici: 393 (1838), new record
587. Lentinus squarrosulus Mont., Annales des Sciences
Naturelles Botanique 18: 21 (1842), new record
588. Lentinus velutinus Fr., Linnaea 5: 510 (1830), new
record
589. Panus subfasciatus Thongbai, Karunarathna & K.D.
Hyde, in Fungal Diversity 83: 228 (2017), new species
590. Polyporus brevibasidiosus H. Lee, N.K. Kim & Y.W.
Lim, in Fungal Diversity 83: 230 (2017), new species
591. Polyporus koreanus H. Lee, N.K. Kim & Y.W. Lim,
in Fungal Diversity 83: 230 (2017), new species
592. Polyporus orientivarius H. Lee, N.K. Kim & Y.W.
Lim, in Fungal Diversity 83: 232 (2017), new species
593. Polyporus parvovarius H. Lee, N.K. Kim & Y.W.
Lim, in Fungal Diversity 83: 234 (2017), new species
594. Polyporus subdictyopus H. Lee, N.K. Kim & Y.W.
Lim, in Fungal Diversity 83: 235 (2017), new species
595. Polyporus ulleungus H. Lee, N.K. Kim & Y.W. Lim,
in Fungal Diversity 83: 236 (2017), new species
596. Wolfiporia pseudococos F. Wu, J. Song & Y.C. Dai,
in Fungal Diversity 83: 237 (2017), new species
Psathyrellaceae Locq.
597. Coprinopsis cerkezii Tkalčec, Mešić, I. Kušan &
Matočec, in Fungal Diversity 83: 239 (2017), new species
Russulales Pers.
Russulaceae Lotsy
598. Russula yanheensis T.C. Wen, K. Hapuarachchi &
K.D. Hyde, in Fungal Diversity 83: 244 (2017), new
species
123
Fungal Diversity
599. Russula virescens (Schaeff.) Fr., Anteckningar öfver
de i Sverige växande ätliga svampar: 50 (1836), reference
specimen
Dacrymycetales Henn.
Dacrymycetaceae J. Schröt.
600. Dacrymyces chiangraiensis
C. Karunarathna, Q. Zhao & K. D.
Diversity 83: 248 (2017), new species
601.
Femsjonia
monospora
C. Karunarathna, Q. Zhao & K. D.
Diversity 83: 251 (2017), new species
Ekanayaka, S.
Hyde, in Fungal
Ekanayaka,
S.
Hyde, in Fungal
Tremellales Fr.
Tremellaceae Fr.
602. Tremella fuciformis Berk., Hooker’s Journal of Botany and Kew Garden Miscellany 8: 277 (1856), new record
Materials and methods
Sampling, isolation and identification
Specimens in this study were collected from Brazil, China,
Croatia, Germany, India, Italy, Korea, Russia, Sri Lanka,
Thailand and the UK. Soil samples to isolate Gongronella
brasiliensis were collected from Taquaritinga do Norte,
state of Pernambuco, Brazil, following methods outlined in
Benny (2008). Gongronella brasiliensis was cultured in
triplicate, in MEA and PDA, and incubated at 10, 15, 20,
25 and 30 °C, over 7 days. Colony characteristics were
recorded, while alternative morphs were induced in culture
using sterilized pieces of plant material (Phookamsak et al.
2015). Plant material was examined with a Carl Zeiss
GmbH (AxioCam ERC 5 S) stereo microscope. Fruiting
bodies of ascomycetes were rehydrated in water, lactic acid
or 5% KOH. Cut sections were observed with a compound
microscope; micro-morphological characteristics (e.g.
ascomata sections, peridium structures, asci and ascospores
in ascomycetes and basidia, cystidia, basidiospores in
basidiomycetes) were examined, described and photographed. Measurements (e.g. from basidiospores and
ascospores) were taken from at least 20 representatives and
the mean and the standard deviation for length and width,
together with the range of spore quotient (Q, the length/
width ratio) and its mean value (Qm) are given. Herbarium
specimens and ex-type living cultures were deposited in
various collections and are listed under each taxon
description. Facesoffungi numbers (FoF) and Index Fungorum (IF) numbers were obtained as explained in Jayasiri
et al. (2015) and Index Fungorum (2017). New species
were established as per recommendations outlined by
Jeewon and Hyde (2016).
123
DNA extraction, PCR amplification and sequencing
For most ascomycetous fungal samples, total genomic
DNA was extracted from fresh fungal mycelium grown on
appropriate media at 16 °C or room temperature (as outlined by Jeewon et al. 2002), while for basidiomycetes, dry
fruiting bodies were used. DNA was extracted following
the instructions of the Biospin Fungus Genomic DNA
Extraction Kit (BioFluxÒ, Hangzhou, P.R. China) or other
fungal DNA extraction kits according to the manufacturer’s instructions. When fungi failed to grow in culture,
DNA was extracted directly from ascomycete fruiting
bodies using aseptic techniques. For Gongronella
brasiliensis, fungal biomass was obtained in MEA cultures
in test tubes kept at 28 °C for up to six days and genomic
DNA extraction of G. brasiliensis was carried out with
previously macerated material according to Góes-Neto
et al. (2005).
DNA amplification for all samples was performed by
Polymerase Chain Reaction (PCR) using universally standard primers. The primers used and PCR conditions are
shown in the Table 1. For Gongronella brasiliensis specimens, thermal profiling and amplification reactions of ITS
regions were conducted as described by Oliveira et al.
(2014) and the final amplicons were purified with the
PureLink PCR Purification Kit (Invitrogen), sequenced
directly or cloned with a Clone JETTM PCR Cloning Kit
(Fermentas; Carlsbad, USA), following the manufacturer’s
instructions. The quality of the PCR products was checked
by electrophoresis on 1% agarose gels. Purification and
sequencing of PCR products (with same primers used in
PCR) was done by commercial sequencing providers
depending on the regions where the studies were carried
out.
Sequence alignment and phylogenetic analyses
A careful verification of sequence data generated in this
study was carried out with appropriate reference sequences
following BLAST searches in the nucleotide database of
GenBank (http://blast.ncbi.nlm.nih.gov/). It was ensured
that no erroneous sequences were used in the further
analyses and then data were submitted to GenBank. Following sequence verification and Blast search, DNA
sequences from appropriate taxonomic ranks were downloaded based on recent publications to construct datasets
for phylogenetic analyses. BioEdit sequence alignment
editor (Hall 1999), 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 sequence alignment. Under most circumstances,
concatenated DNA datasets were analyzed to generate
phylogenetic trees, but in cases of limited availability of
Fungal Diversity
Table 1 Details of genes/loci with PCR primers and PCR profiles
Gene/loci
PCR primers (forward/reverse)
PCR condition
References
Actin
ACT512F/ACT783R
95 °C: 40 s, 58 °C: 30 s,b
Carbone and Kohn (1999)
72 °C: 1 min (40 cycles)e
Calmodulin
cmdD3/cmdD4
91 °C: 1 min, 45 °C: 1 min,b
72 °C: 1 min (35 cycles)
CHS-1
CHS79F/CHS345R
Wang and Zhuang (2004)
e
95 °C: 30 s, 59 °C: 30 s,c
Carbone and Kohn (1999)
72 °C: 45 s (35 cycles)f
GAPDH
GDF/GDR
95 °C: 30 s, 54 °C: 30 s,c
Templeton et al. (1992)
72 °C: 45 s (35 cycles)f
ITS
ITS5/ITS4
94 °C: 30 s, 52 °C: 50 s,a
White et al. (1990)
d
White et al. (1990)
ITS1/ITS4
72 °C: 1 min (35 cycles)
LSU
LROR/LR5
NL1/NL4
94 °C: 30 s, 55 °C: 50 s,a
72 °C: 1 min (35 cycles)d
Vilgalys and Hester (1990)
O’Donnell (1993)
RPB1
RPB1-Af/RPB1-Cr
95 °C: 30 s, 4 °C: 30 s,b
Stiller and Hall (1997)
72 °C: 1 min (30 cycles)d
RPB2
SSU
fRPB2-5f/fRPB2-7cR
95 °C: 1 min, 52 °C: 2 min,b
Liu et al. (1999)
bRPB-6F/bRPB-7.1R
72 °C: 1.30 s (35 cycles)d
Matheny (2005)
NS1/NS4
94 °C: 30 s, 55 °C: 50 s,a
White et al. (1990)
72 °C: 1 min (35 cycles)d
TEF1
EF1-983F/EF1-2218R
94 °C: 30 s, 52 °C: 50 s,a
72 °C: 1 min (35 cycles)
b-tubulin
Rehner (2001)
d
Btub2Fd/Btub4Rd
95 °C: 30 s, 53 °C: 30 s,c
Woudenberg et al. (2009)
Bt2a/Bt2b
72 °C: 45 s (35 cycles)f
Glass and Donaldson (1995)
BT1/BT2
a
Initiation step of 94 °C: 3 min
b
Initiation step of 95 °C: 5 min
c
Initiation step of 95 °C: 3 min
d
Final elongation step of 72 °C: 10 min and final hold at 4 °C
e
Final elongation step of 72 °C: 5 min and final hold at 4 °C
f
Final elongation step of 72 °C: 1 min and final hold at 4 °C
DNA sequences from respective gene regions, phylogenies
were inferred from single or a combination of two gene
datasets and either a consensus of these gene phylogenies
or one of the most parsimonious phylogenies was used to
infer phylogenetic relationships across taxa sampled.
Selection of outgroup(s) for rooting purposes was based on
knowledge of potential common ancestors to the in-group
as well as taxon sampling from previously published
studies. Phylogenetic analyses were performed by neighbor
joining (NJ), maximum parsimony (MP), maximum likelihood (RAxML) and Bayesian inference (BI) analyses.
The maximum parsimony analyses were performed using
PAUP v. 4.0b10 (Swofford 2003), bootstrap analysis with
1000 replicates. All multiple, equally parsimonious trees
were saved and descriptive tree statistics for parsimony
consistency index (CI), retention index (RI), rescaled
consistency index (RC) and homoplasy index (HI) were
Carbone and Kohn (1999)
calculated. Other details are as described by Jeewon et al.
(2003b, 2004, 2013), Hu et al. (2007) and Promputtha et al.
(2007). The robustness of the best parsimonious tree was
estimated by a bootstrap (BT) value with 1000 replicates,
each with 100 replicates of random stepwise addition of
taxa (Liu et al. 2011, 2012). Maximum likelihood (ML)
analysis was performed using RAxMLGUI v. 1.3 (Silvestro
and Michalak 2010) with 1000 rapid bootstrap replicates.
The substitution model comprised a generalized time
reversible (GTR) for nucleotides with a discrete gamma
distribution (Silvestro and Michalak 2012). Bayesian
analyses were performed by MrBayes v. 3.0b4 (Ronquist
and Huelsenbeck 2003) with the best-fit model of sequence
evolution estimated with MrModeltest 2.2 as described in
Nylander et al. (2008). Markov Chain Monte Carlo sampling (BMCMC) was used to determine the posterior
probabilities (PP) (Rannala and Yang 1996; Zhaxybayeva
123
Fungal Diversity
and Gogarten 2002) in MrBayes v. 3.0b4 (Ronquist and
Huelsenbeck 2003). Six simultaneous Markov chains were
run at least 1000,000 generations or depending on individal
settings for the fungal group and trees were sampled every
100th/1,000th generation (Cai et al. 2006). Tracer v. 1.6
(Rambaut and Drummond 2003) was used to check the
effective sampling sizes (ESS) and burn-in value. The run
was stopped automatically as soon as the average standard
deviation of split frequencies fell below 0.01 (Maharachchikumbura et al. 2015). Phylograms were visualized in Treeview (Page 2001) or FigTree 1.4.2 (Rambaut
2014) with bootstrap values above or below the nodes and
reorganized in Adobe Illustrator CS v. 6., Microsoft Powerpoint (v.2007), portable document format (PDF) and
Adobe Photoshop version CS3 (Adobe Systems, USA). All
the sequences generated in this study were deposited in
GenBank and accession numbers have been provided
where appropriate.
Dothideomycetes
Dothideomycetes is the largest class of Ascomycota. In
this study, we follow the classifications in the recent studies
of Hyde et al. (2013) and Wijayawardene et al. (2014a).
Asterinales M.E. Barr ex D. Hawksw. & O.E. Erikss.
Hyde et al. (2013) accepted Asterinaceae, Aulographaceae, and Parmulariaceae in Asterinales and a
revision of genera in Asterinales was provided by Hongsanan et al. (2014). In recent publications members of
Asterinales clustered in two clades (Ertz et al. 2016; Hyde
et al. 2016). Hyde et al. (2016) treated the Asterinales
clade (not clade containing Parmulariaceae) as Asterinales sensu stricto, because most of the Asterinales
strains, from both Asterina and Lembosia clustered in this
clade and also because the large clade that contains Asterinales as circumscribed by Ertz et al. (2016) does not
have any phylogenetic support. Hyde et al. (2016) also
synonymized the younger Asterotexiales under Asterinales. In this paper, we provide an updated tree for Asterinales (Fig. 1).
Asterinaceae Hansf.
The family Asterinaceae has characteristic superficial,
web-like, black colonies on the upper and lower surfaces of
leaves (Hyde et al. 2013). Hofmann et al. (2010) placed
Asterinaceae in Dothideomycetes based on multi-gene
sequences, and this was substantiated by Hongsanan et al.
(2014).
Morenoina Theiss
= Aulographella Höhn.
Morenoina was introduced by Theissen (1913) with
Morenoina antarctica (Speg.) Theiss. as the type species.
123
Morenoina resembles Aulographum, differing only in the
morphology of the scutellum, which comprises inordinately arranged cells and a hypostroma of subcuticular
hyphae beneath the thyriothecium (Ellis 1980).
Morenoina calamicola Konta & K. D. Hyde, sp. nov.
Index Fungorum number: IF552661; Facesoffungi
number: FoF02757, Fig. 2
Holotype: MFLU 15-0013
Saprobic on Calamus sp. Sexual morph Thyriothecia
100–325 lm, occurring on host surface, solitary, aggregated, or gregarious, easily removed from the host surface,
superficial, ellipsoid, oblong, curved, flat, with longitudinal, slit-like opening, linear fissure, which are branched at
the margin, from the center to the outer rim, lacking free
hyphae and appressoria at the margin, in transverse Sect.
24–33 lm high 9 86–121 lm diam. (
x = 27 9 103 lm,
n = 5). Upper wall comprising linear cells, with irregular,
filiform hyphae, radiating from the center to the outer rim.
Asci 13–20 9 10–17 lm (
x = 17 9 14 lm, n = 10), 8spored, bitunicate, globose to subglobose or clavate, or
saccate to globose, apedicellate, with a distinct, thickened
apical region. Ascospores 9–11 9 3–4 lm (
x = 10 9 3.9
lm, n = 10), irregularly arranged, oblong or fusiform,
wider at apex, with slightly acute ends, 1–septate, with two
large guttules in each cell, hyaline, smooth-walled. Asexual morph Undetermined.
Culture characteristics: Ascospores germinating on
MEA within 24 h and germ tube produced from each cell.
Colonies on MEA reaching 3–4 cm diam., after 5 days,
grey to olivaceous, dense, with fairly fluffy surface,
hyphae, septate, branched and smooth-walled.
Material examined: THAILAND, Chiang Mai Province, on
dead stem of Calamus sp. (Arecaceae), 11 August 2014,
Sirinapa Konta, P04a (MFLU 15-0013, holotype; ex-type
living culture, MFLUCC 14-1162).
GenBank Numbers LSU: KY511424; ITS: KY511430;
SSU: KY511427.
Notes: Hongsanan et al. (2014) transferred Morenoina to
Aulographaceae based on morphology and phylogeny. In
the current phylogenetic analyses, Morenoina calamicola
clustered with Melaspilea lekae (Melaspileaceae) (94% in
ML, 1.00 in BYPP). The morphology of Morenoina
calamicola is distinct and Melaspilea lekae is lichenicolous
(Kalb et al. 2012). Morenoina calamicola is similar to
Aulographaceae (Asterinales) in lacking appressoria and
flattened, elongate thyriothecia, opening by a slit. In the
phylogenetic analyses (Fig. 1) Morenoina calamicola is
well-separated from Aulographaceae.
Morenoina calamicola is most similar to M. palmicola J.
Fröhl., K.D. Hyde & Joanne E. Taylor in characteristics of
asci, ascospores and host genus (Calamus australis), but
differs in the size and shape of ascospores.
Fungal Diversity
Fig. 1 Phylogram generated from maximum likelihood and Bayesian
analyses based on LSU sequence data from selected species of
Asterinales. The RAxML bootstrap value greater than 50% and the
posterior probabilities value C0.95 are shown above the nodes.
Lichenized taxa are in green and lichenicolous fungi in red. The new
isolate is in blue bold and other ex-type strains are in black bold. The
tree is rooted with Saccharomyces cerevisiae
Botryosphaeriales C.L. Schoch et al.
Botryosphaeriales is a diverse order with a worldwide
distribution, comprising species that vary from endophytes
to pathogens (Slippers and Wingfield 2007; Phillips et al.
2013; Chethana et al. 2016; Daranagama et al. 2016; Dissanayake et al. 2016; Giambra et al. 2016; Konta et al.
2016a, b; Linaldeddu et al. 2016a, b, c; Manawasinghe
et al. 2016; Tennakoon et al. 2016; Zhang et al. 2017).
Since it was introduced by Schoch et al. (2006) this order
has undergone significant taxonomic changes with the
addition of several new families. Currently, nine families
are recognized, namely, Aplosporellaceae, Botryosphaeriaceae, Endomelanconiopsisaceae, Melanopsaceae, Phyllostictaceae, Planistromellaceae, Pseudofusicoccumaceae,
Saccharataceae and Septorioideaceae (Schoch et al. 2006;
Minnis et al. 2012; Wikee et al. 2013; Slippers et al. 2013;
Wyka and Broders 2016; Dissanayake et al. 2016; Yang
et al. 2017), although several authors considered this to be
too many (Chethana et al. 2016; Daranagama et al. 2016;
Dissanayake et al. 2016; Giambra et al. 2016; Konta et al.
2016a, b; Linaldeddu et al. 2016a, b, c; Manawasinghe
et al. 2016; Tennakoon et al. 2016; Liu et al. 2016; Zhang
et al. 2017).
Botryosphaeriaceae Theiss. & Syd.
The family Botryosphaeriaceae was described in the
1820s, originally for species of Sphaeria (Fr.) (Crous et al.
2006; Schoch et al. 2006). von Arx and Müller (1954). Barr
123
Fungal Diversity
Fig. 2 Morenoina calamicola (MFLU 15-0013, holotype). a Appearance of thyriothecia on host substrate. b Close up of thyriothecium. c, e Cell walls of thyriothecium with radial arrangement.
d Section of thyriothecium. f–i Asci. j–k Ascospores. l Germinated
ascospore. m Culture on MEA media. Scale bars a = 500 lm,
b = 100 lm, c, d = 20 lm, e–i = 10 lm, j, k = 5 lm, l = 20 lm
(1987) included only nine genera, which are mostly different from those of von Arx and Müller (1954). Hawksworth et al. (1995) listed five genera. Lumbsch and
Huhndorf (2010) included eleven genera, while Hyde et al.
(2011) and Wijayawardene et al. (2012) listed 16 genera.
Liu et al. (2012) included 17 genera in the family based on
molecular data and examination of generic types. Phillips
et al. (2013) provided comprehensive descriptions, notes
and phylogenies for the 17 genera and 110 species known
from cultures. Twenty-three genera and 187 species of
123
Fungal Diversity
Botryosphaeriaceae were listed by Dissanayake et al.
(2016).
Barriopsis A.J.L. Phillips, A. Alves & Crous
Barriopsis was introduced when Phillips et al. (2008)
transferred Physalospora fusca to Barriopsis fusca. Stevens
(1926) originally placed this species in Physalospora, but
was obviously hesitant to do so on account of its brown
ascospores. Petrak and Deighton (1952) then transferred it
to Phaeobotryosphaeria as P. fusca. Although von Arx and
Müller (1954) considered Phaeobotryosphaeria a synonym
of Botryosphaeria, Phillips et al. (2008) showed that it is
morphologically and phylogenetically distinct from other
genera in Botryosphaeriaceae. However, the fungus considered by Stevens (1926) and Petrak and Deighton (1952)
does not have apiculi on its ascospores, and thus does not
fall within the concept of Phaeobotryosphaeria, which has
small, hyaline apiculi on the ascospores. For this reason
Phillips et al. (2008) introduced the genus Barriopsis.
Dissanayake et al. (2016) list four species.
Barriopsis thailandica Dissanayake, Senan. & K.D. Hyde,
sp. nov.
Index Fungorum number: IF551363; Facesoffungi
number: 00913, Figs. 4, 5
Etymology: Name refers to the country Thailand, where
the fungus was collected. Holotype: MFLU 15-1414
Saprobic on decaying bark of Tectona grandis L.f.
Sexual morph Ascostromata 170–220 lm high 9 150–170
lm diam. (
x = 194 9 160 lm, n = 10), scattered, initially
immersed, becoming erumpent through bark when mature,
solitary or gregarious, uniloculate, globose to subglobose,
or flask-shaped when cut horizontally, coriaceous, ostiolate, papillate. Papilla 70–80 9 60–65 lm (
x = 77 9 64
lm, n = 10), short, periphysate. Periphyses 15–20 lm
(
x = 20 lm, n = 10), brown, curved towards the outside,
long, flat, leaf-like. Peridium 20–45 lm wide (
x = 30 lm,
n = 10) comprising outer, 5–10 layers of brown, thickwalled, large cells of textura angularis to textura globulosa
and inner, 4–7 layers of hyaline, thin-walled, small cells of
textura angularis to textura globulosa. Hamathecium
comprising 2–4 lm wide (
x = 3.5 lm, n = 10), hyphalike, numerous, septate, pseudoparaphyses, slightly constricted at septa. Asci 50–110 9 19–21 lm (
x = 70 9 21
lm, n = 20), (2) 4 (8)-spored, bitunicate, fissitunicate,
cylindric-clavate or clavate, with long or short pedicel,
apically wider than base, apex rounded with an ocular
chamber. Ascospores 15–20 9 9–11 lm (
x = 18 9 10 lm,
n = 30), mostly biseriate, rarely overlapping uniseriate,
ellipsoidal or rhomboidal, inequilateral, wider in the center,
initially hyaline, becoming yellow, pale brown to reddishbrown when mature, aseptate, ends blunt when mature,
thick-walled, smooth-walled or verruculose. Asexual
morph Conidiomata stromatic, uniloculate, dark brown to
black. Hamathecium 20–35 lm long (
x = 28 lm, n = 20),
hyaline, cylindrical, mostly aseptate, sometimes branched,
ends slightly swollen and rounded, arising amongst the
conidiogenous cells. Conidiophores reduced to conidiogenous cells. Conidiogenous cells 4–7 9 4–6 lm (
x = 6.5
9 5 lm, n = 20), hyaline, smooth, cylindrical, slightly
swollen at the base, enteroblastic, proliferating percurrently
to form one or two closely spaced annellations. Conidia
15–19 9 7.5–8.5 lm (
x = 17 9 7.8 lm, n = 20), ellipsoidal at apex, rounded at base, widest at the center, thickwalled, initially hyaline, aseptate, becoming 1-septate, dark
brown, striated after released from the conidiomata.
Culture characteristics: Colonies on MEA reaching 3–4
cm diam., after 5 days in the dark at 18 °C, flattened,
velvety, lobate, fimbriate, initially white at the edge,
becoming olive green in the center, after 7 days becoming
greenish-ash, woolly with erect mycelia.
Material examined: THAILAND, Uttaradit Province,
on decaying bark of Tectona grandis (Lamiaceae), 12
October 2014, I.C. Senanayake (MFLU 15-1414, holotype), ex-type living culture MFLUCC 14-1190, KUMCC
16-0185.
GenBank Numbers ITS:KY115675; TEF1:KY115676.
Notes: Barriopsis is one of the less well-known genera
in the family Botryosphaeriaceae. To date only four species; Barriopsis archontophoenicis, B. fusca, B. iraniana
and B. tectonae have been reported for this genus (Phillips
et al. 2008; Abdollahzadeh et al. 2009; Doilom et al. 2014,
Konta et al. 2016a, b). Barriopsis tectonae has both sexual
and asexual morphs (Doilom et al. 2014; Konta et al.
2016a, b). We describe both a sexual and asexual morph
for Barriopsis thailandica (Figs. 4, 5). The asexual morph
formed in culture grown on MEA (Fig. 5). In the phylogenetic analysis of combined ITS and TEF1 sequence data,
this new taxon is related to B. iraniana (Fig. 3), but phylogenetically distinct with very good support (100% maximum parsimony and 1.0 in Bayesian analysis) to justify its
establishment as a new species. The small conidial
dimensions of B. thailandica (17 9 8 lm, L/W ratio = 2.1)
clearly distinguish it from B. iraniana (27 9 16 lm, L/W
ratio = 1.6). The asexual morph has not been reported for
B. iraniana, whereas that of B. thailandica is reported
herein.
Capnodiales Woron.
The order Capnodiales introduced by Woronichin
(1925) was accepted in Dothideomycetes based on phylogenetic studies (Schoch et al. 2006, 2009; Crous et al.
2009; Chomnunti et al. 2014). The Capnodiales are mainly
defined by their shared ecological niches (Hughes 1976;
Crous et al. 2009; Chomnunti et al. 2011; Hyde et al.
2013).
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Fungal Diversity
Fig. 3 Phylogram generated
from maximum parsimony
analysis of all available type/
authentic sequences of
Barriopsis based on combined
ITS and TEF1 sequence data.
Parsimony bootstrap support
values for MP C 75% and
Bayesian posterior probabilities
C0.9 are indicated on the nodes.
Isolate numbers of all ex-types
and reference strains are in bold.
Novel species are indicated in
blue. The tree is rooted with
Sphaeropsis visci
Mycosphaerellaceae Lindau
Mycosphaerellaceae is a large family in the order
Capnodiales (class Dothideomycetes). Species in Mycosphaerellaceae have cosmopolitan distributions, occurring on various hosts worldwide (Crous et al.
2009, 2013a, b; Schoch et al. 2009; Farr and Rossman
2012; Hyde et al. 2013; Quaedvlieg et al. 2013). Some
genera of Mycosphaerellaceae e.g. Cercospora, Pallidocercospora, Pseudocercospora and Septoria contain
important and well known pathogens, causing serious diseases of economic crops (Crous et al. 2007, 2009, 2013a, b;
Cheewangkoon et al. 2008; Schoch et al. 2009; Hyde et al.
2013; Quaedvlieg et al. 2013). Several sexual genera in
Mycosphaerellaceae are morphologically similar, but can
be distinguished by their asexual morphs (Crous et al.
2007, 2009, 2013a, b). Many phylogenetic studies of the
asexual genera in Mycosphaerellaceae have been carried
out (Crous and Braun 2003; Crous et al.
2007, 2009, 2013a, b; Quaedvlieg et al. 2013). However,
phylogenetic affinities of sexual genera such as Achorodothis, Polythrincium (=Cymadothea), Euryachora,
123
Gillotia, Melanodothis, Polyascosporella and Wernerella
are largely unresolved partially due to limited availability
of collections and DNA sequence data (Hyde et al. 2013).
The phylogeny of Polythrincium trifolii is shown in Fig. 6.
Polythrincium Kunze
Polythrincium was introduced in Kunze and Schmidt
(1817) and is typified by P. trifolii Kunze. Polythrincium
trifolii is an obligate pathogen causing sooty blotch of
clover in temperate regions (Simon et al. 2005a, b, 2009;
Rossman et al. 2015). The sexual morph of Polythrincium
has been reported as Cymadothea trifolii (Wolf 1935) and
it is characterized by small, black, pustulate ascostromata
on the lower surface of clover leaflets, bitunicate, clavate
asci, lack of pseudoparaphyses and hyaline to subhyaline,
oblong to clavate and 1-septate ascospores (Wolf 1935;
Simon et al. 2009).
Polythrincium is relatively poorly studied with only
seven epithets in Index Fungorum (2017), but rDNA based
phylogenies indicate that P. trifolii belongs to the family
Mycosphaerellaceae (Simon et al. 2009).
Fungal Diversity
Fig. 4 Barriopsis thailandica (MFLU 15-1414, holotype). a, b
Appearance of ascostroma on substrate. c Cross section of
ascostroma. d Peridium. e Papilla with setae-like periphyses.
f Pseudoparaphyses. g–k Asci. l–q Ascospores. Scale bars a,
b = 500 lm, c = 50 lm, d, e, g–k = 20 lm, f = 10 lm
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Fungal Diversity
Fig. 5 Barriopsis thailandica (MFLUCC 14-1190, ex-type living culture on MEA). a Conidiomata in culture. b–e Immature conidia attached
to the conidiogenous cells. f, g Mature conidia. Scale bars a = 200 lm, b–g = 10 lm
Polythrincium trifolii Kunze, in Kunze & Schmidt, Mykologische Hefte (Leipzig) 1: 14 (1817) Synonym: Cymadothea
trifolii(Pers.)F.A.Wolf,Mycologia27(1):71(1935)
Facesoffungi number: FoF 2765, Fig. 7
Reference specimen: HKAS 96227
Obligate biotroph causing black blotch or sooty blotch on
leaves of Trifolium repens L. Colonies on host substrate
brown to dark brown, scattered to gregarious, punctiform,
effuse to dense, eventually covering most of the leaf surface.
Mycelium immersed, brown to dark brown, smooth-walled.
Sexual morph reported as Cymadothea trifolii (Wolf 1935).
Asexual morph hyphomycetous forming on lower surface
of leaves. Sporodochia 40–120 lm high, 35–80 lm diam.,
with cushion-like pseudoparenchymatous cells at the base,
composed of compact, fasciculate, foot-like cells at the base,
with tightly aggregated, parallel, cylindrical to subcylindrical conidiophores. Conidiophores (25–)30–60(–62) 9 (4.5–
)5–7 lm (
x = 45.1 9 6.2 lm, n = 50), macronematous,
mononematous, caespitose, erect to flexuous, unbranched,
aseptate, torsive, brown to dark brown, smooth-walled, largely ampulliform at the base, undulated at the upper part.
Conidiogenous cells polyblastic, sympodial, integrated,
terminal, cylindrical, undulate, with large, cicatrized scars.
Conidia (21–)24–27(–29) 9 (13–)15–17(–19) lm (
x = 25.3
9 16.6 lm, n = 50), acropleurogenous, solitary, simple,
cuneiform to pyriform, brown, 1-septate, rough-walled,
verruculose, upper cell larger than lower cell.
123
Material examined: CHINA, Yunnan Province, Kunming Institute of Botany, on lower surface of leaves of
white clover Trifolium repens L., (Fabaceae), 19 September 2016, R. Phookamsak, KIB001 (HKAS 96227, reference specimen designed here).
GenBank Numbers HKAS96227A LSU:KY554951;
ITS:KY554953. HKAS96227B LSU:KY554952.
Notes: Polythrincium trifolii differs from P. shiraianum
Henn. and P. trifolii var. platense Speg. in having larger
conidia (P. shiraianum = 15–30 9 7–8 lm and P. trifolii
var. platense = 16–18 9 12–14 lm) (Hennings 1905;
Spegazzini 1910). Polythrincium lathyrinum Syd. has
conidia of a similar size (P. trifolii: (21–)24–27(–29) 9
(13–)15–17(–19) lm; P. lathyrinum: 22–32 9 15–18 lm),
but occurs on Lathyrus maritimus Willd. (Farr and Rossman 2012) (Sydow 1928). while P. trifolii has been
reported from Trifolium, Astragalus sinicus L. (from
China) and Stemphylium sarcinaeforme (Cavara) Wiltshire.
Phylogenetic analyses based on combined LSU and ITS
sequence data show that P. trifolii belongs to Mycosphaerellaceae and concurs with Simon et al. (2009).
This is the first report on T. repens from China.
Dothideales Lindau
Thambugala et al. (2014b) revised the order Dothideales
and synonymized Dothioraceae under Dothideaceae, and
accepted only two families in the order.
Fungal Diversity
Fig. 6 The best scoring of the RAxML tree based on analysis of
combined dataset of ITS and LSU sequence data. Bootstrap support
values for maximum parsimony (right) and maximum likelihood (left)
greater than 50% are given above the nodes. Bayesian posterior
probabilities (BYPP) greater than 0.95 are shown as bold branches.
The tree is rooted to Dissoconium aciculare. The new strain is
indicated by bold blue. Ex-type strains are shown in bold
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Fungal Diversity
Fig. 7 Polythrincium trifolii (HKAS 96227, reference specimen).
a–c Appearance of fungal colonies on lower surface of white clover
leaves. d, e Section through sporodochia arising from leaf tissue. f, g
Conidiophores. h–j Conidiogenous cells with developing conidia. k–
q Conidia. Scale bars d–f = 50 lm, g = 20 lm, h–j = 10 lm, k–
q = 5 lm
Dothioraceae Theiss. & Syd.
Dothideaceae was established by Chevallier (1826) as
‘‘Dothideae’’, and later Fuckel (1870) introduced Dothidea as the type genus and D. gibberulosa (Fr.) Fr. as
the type species. The characteristic features of Dothideaceae are immersed to erumpent or superficial, 1 or
multi-loculate ascostromata; 8-or polysporous, bitunicate
asci and hyaline or brown, transversely septate, sometimes muriform (Dothiora Fr.) ascospores (Thambugala
et al. 2014b). Dothideaceae was revised by Thambugala
et al. (2014b) who included ten sexual genera and five
asexual genera. The phylogeny of Dothiora buxi along
with other members of the Dothideaceae is shown in
Fig. 8.
123
Dothiora Fr.
Dothiora was introduced by Fries (1849) with
D. pyrenophora (Fr.) Fr. as the type species. Thambugala
Fungal Diversity
Fig. 8 RAxML maximum
likelihood phylogenetic tree
based on analysis of LSU and
ITS sequence data from species
of Dothideaceae. Maximum
likelihood bootstrap values
greater than 50% are shown
above the nodes. The new
isolates are in blue and other extype strains are in bold. The tree
is rooted with Elsinoë phaseoli
et al. (2014b) based a study of Dothiora on both morphology and phylogeny.
Dothiora buxi Jayasiri, Camporesi & K.D. Hyde, in Fungal
Diversity 81: 30 (2016)
Facesoffungi number: FoF00078, Fig. 9
Saprobic and/or parasitic on dying leaves and twigs
(hemibiotrophic) of Buxus sempervirens L. Sexual morph
in Hyde et al. 2016. Asexual morph Conidiomata 182–263
lm high 9 383–447 lm diam. (
x = 232 9 422 lm,
n = 5), pycnidial, globose to subglobose, scattered,
solitary, visible as brown to black, pustulate on the lower
leaf surface, semi-immersed to erumpent, exposed by
breaking leaf epidermis. Peridium 21–43 lm wide, comprising 1–3 layers of thickened, pale brown, cells arranged
in textura prismatica. Conidiogenous cells 11–17 9 6–11
lm diam. (
x = 14 9 9 lm, n = 10), phialidic, subglobose,
hyaline. Conidia 10–16 9 6–19 lm diam. (
x = 13 9 7 lm,
n = 10), ellipsoid to obovoid, rounded at top, narrow at
base, guttulate, smooth, 1-celled, hyaline.
Culture characteristics: Colonies on MEA, reaching
2–2.5 cm diam. after 10 days at 16 °C, flat, margin with
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Fungal Diversity
Fig. 9 Dothiora buxi (MFLU 15-2356, asexual morph). a Conidiomata on the upper leaf surface. b Conidiomata on the lower leaf
surface. c–e Close up of conidiomata. f Section of conidiomata.
123
g Cells of peridium. h–o Conidiogenous cells. p–u Conidia. v Germinated conidium. Scale bars a, b = 5 mm, c = 1000 lm, d = 500
lm, e = 200 lm, f = 100 lm, g = 20 lm, h–u = 5 lm, v = 10 lm
Fungal Diversity
entire edge, white, producing conidiomata and numerous
conidia.
Material examined: RUSSIA, Rostov Region, Shakhty
City, 20th anniversary of Red Army microdistrict, street
plantation, street shrubs, dying leaves and twigs (hemibiotrophic) of Buxus sempervirens, 1 May 2015, T. Bulgakov, T421 (MFLU 15-2356, asexual morph); living
culture MFLUCC 15-0823.
GenBank Numbers LSU:KY511425; SSU:KY511428
(Fig. 10).
Notes: Fries (1849) introduced the genus Dothiora, with
D. pyrenophora (Fr.) Fr. as the type species. In the current
study, the phylogenetic analyses indicate that our strain is
closely related to Dothiora buxi Jayasiri, Camporesi &
K.D. Hyde (MLFU 15-3404) with 92% bootstrap support.
The morphology could not be compared as the latter was
reported in its sexual morph with no information on its
asexual morph (Hyde et al. 2016). However, we found the
asexual morph on the same host, Buxus sempervirens. In
culture, conidiomata were produced from germinated
ascospores after one week’s incubation. Conidiomata,
conidiogenous cells, and conidia found in culture are
similar to those of D. buxi found on the host. Thus, we
introduce the new record of asexual morph of D. buxi based
on the morphological and phylogenetic evidence.
Fig. 10 Phylogram generated from maximum parsimony analysis
based on LSU, SSU and TEF1 sequence data from taxa of
Hysteriaceae. Maximum likelihood bootstrap support values greater
than 70% are shown above the nodes. The tree is rooted with
Delitschia winteri. 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 isolate is in
blue
Hysteriales Lindau
The order Hysteriales, consisting of a single family, was
introduced by Lindau (1896). DNA sequence data showed
that Hysteriales should be accommodated in Dothideomycetes (Boehm et al. 2009a, b; Shearer et al. 2009;
Suetrong et al. 2009). This order is characterized by carbonaceous navicular ascoma with a longitudinal dehiscent
slit and ascospores that are typically dark and vary in
septation (Boehm et al. 2009a).
Hysteriaceae Chevall.
Chevallier (1826) introduced the family Hysteriaceae as
‘Hysterineae’. This family has been treated with different
genera by authors (Zogg 1962; von Arx and Muller 1975;
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Fungal Diversity
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. 2016).
Hysterium Tode
Hysterium was introduced by Persoon (1797) with
H. pulicare Pers. as type species. The genus is characterized by pigmented, versicolorous or concolorous, asymmetric phragmoascospores, three- or more transverselyseptate, borne in hysterothecia (Boehm et al. 2009a).
Hysterium centramurum Senan. sp. nov.
Index Fungorum number: IF552708; Facesoffungi
number: FoF00164, Figs. 11, 12
Etymology: The species epithet from the combination of
two Latin words ‘‘centrum’’ and ‘‘murum’’, meaning centre
and septum or wall, in reference to the ascospore having a
median septum.
Holotype: MFLU 14-0096
Saprobic on decaying wood. Sexual morph Ascomata
0.9–1.8 mm long, 400–450 lm wide, 370–380 lm high
(
x = 1250 9 400 9 375 lm, n = 20), a hysterothecium,
scattered, superficial, base immersed in substrate, elongate
and depressed conchate, globose, surface black, shiny,
longitudinally striate, apex compressed, opening by longitudinal slit. Periphyses aseptate, hyaline with swollen ends.
Peridium 30–45 lm wide (
x = 34 lm, n = 10), carbonaceous, brittle, of heavily pigmented, small, prosenchymatous cells. Hamathecium comprising 0.75–0.85 lm wide
(
x = 0.8 lm), aseptate, branched, trabeculate pseudoparaphyses, borne in a gel matrix which is pinkish and granular
above the asci. Asci 170–195 9 30–55 lm (
x = 179 9 36
lm, n = 20), 8-spored, bitunicate, oblong to clavate, with a
short pedicel, apically thickened, with a distinct ocular
chamber. Ascospores 110–130 9 20–25 lm (
x = 125 9 23
lm, n = 20), crowded to biseriate, fusiform when young,
oblong at maturity, with a single median septum, hyaline to
light yellow, wall greatly thickened towards the apex,
cytoplasm clearly subdivided into numerous compartments
in Melzer’s and Cotton blue reagent, muriform when
mature, endoplasts appearing cup-shaped when young,
smooth-walled. Sheath present, but not very obvious.
Asexual morph coelomycetous. Conidiomata 100–110 lm
(
x = 107 lm), superficial, solitary or aggregated, black to
brown, appearing as a mycelium mass, prosenchymatous,
without a prominent wall. Conidiophores 8–10 lm long 9
1–2 lm wide (
x = 9 9 1 lm), cylindrical, attached to the
mycelium, hyaline. Conidiogenous cells 2–2.5 9 1–1.5 lm
(
x = 2.4 9 1.2 lm), holoblastic, terminal or intercalary on
conidiophores, cylindrical, short, smooth, hyaline. Conidia
123
2–2.5 lm diam. (
x = 2.3 lm, n = 20) oval to globose,
hyaline, brown when mature.
Culture characteristics: Colonies on MEA, greenishwhite, circular, with smooth margin, slow growing,
attaining 2 cm diam. within 60 days at 20 °C, tightly
arranged, short, aerial mycelium. Orange-brownish exudates released into the media. The mycelial mats produce
erumpent, globose, light brownish, viscous droplets when
incubated at 20 °C for 3 weeks and later becoming lighter
in colour.
Material examined: THAILAND, Chiang Mai Province,
Doi Suthep-Pui, Sangasabhasri Lane to Huai Kok Ma
Village (N1848.620 , E9854.600 , elev. 1145 m), on decaying
wood, 17 December 2012, I.C. Senanayake CHUNI 70
(MFLU 14-0096, holotype, HKAS 82633, isotype), extype living culture MFLUCC 12-0808.
GenBank Numbers ITS:KM272258; LSU:KM272256;
SSU:KM272257; TEF1:KM277819.
Notes: Hysterium centramurum is morphologically
similar to Ostreichnion curtisii (Duby) M.E. Barr and
O. thailandicum Tennakoon, Phookamsak & KD Hyde.
However, phylogenetically, our strain is not closely related
to the type species of Ostreichnion, O. sassafras. Combined gene sequences analysis of LSU, SSU and TEF1
(Fig. 10) places H. centramurum in the genus Hysterium.
Morphologically H. centramurum differs from other Hysterium species in having hyaline to light yellow ascospores,
surrounded by a sheath, with a median septum. Additionally, the coelomycetous asexual morph for this species was
obtained in culture. The major secondary metabolic exudate produced in cultures of H. centramurum was identified
by mass spectroscopy and 1D and 2D NMR spectroscopy
data analysis as physcion (1,8-dihydroxy-6-methoxy-3methyl-9,10-dioxoanthracene-2-carboxylic acid), which is
useful in the textile industry as a dye and in the pharmaceutical industry as an antibiotic and anti-cancer agent
(Velmurugan et al. 2010).
Hysterobrevium mori (Schwein.) E.W.A. Boehm & C.L.
Schoch, Stud. Mycol 64: 62 (2009)
Facesoffungi number: FoF 2766, Fig. 13
Saprobic dead branch of Cornus sanguinea L. Sexual
morph Ascomata 200–220 lm high 9 240–290 lm diam.
(
x = 209 9 262 lm, n = 5), superficial on wood, elongate,
narrowly elliptical to fusiform, straight or irregularly
curved, solitary or scattered on the host surface, dull black,
carbonaceous, with a conspicuous longitudinal cleft and
often with inconspicuous parallel striations, the cleft
gradually widening to expose the hymenium, not easier to
remove. Peridium 45–65 lm wide, thick, composed of very
dark, near opaque, thick-walled brown to black cells, basal
region similar, with a layer of hyaline thin-walled cells
Fungal Diversity
Fig. 11 Hysterium centramurum (MFLU 14-0096, holotype). a–c
Ascomata on substrate. d Cross section of ascoma. e Granulated ends
of pseudoparaphyses near to slit. f Pseudoparaphyses. g–i Asci.
j Ascospores releasing from the broken asci. k–l Immature
ascospores. m–o Ascospores at maturity. p Cup-shaped endoplasts.
q–s Fully mature ascospores. t Sheath. Scale bars a, b. 500 lm,
c = 250 lm, d = 100 lm, e = 50 lm, f = 10 lm, g–j = 100 lm, k–
t = 50 lm
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Fig. 12 Hysterium centramurum (MFLUCC 12-0808, asexual morph of ex-type culture). a, b Colonies on MEA. c Mycelium. d Conidiomata on
MEA. e, f Conidiomata. g, h Conidia attached to conidiogenous cell. i Conidia. Scale bars b = 1 mm, c, g, h = 5 lm, e, f = 50 lm, i = 10 lm
forming the cavity floor, dull cells arranged in a textura
angularis to textura globulosa. Hamathecium of 1.2–2.2
lm wide, long cylindrical, hyaline, smooth, septate, branched, cellular, pseudoparaphyses. Asci 50–70 9 10–20 lm
(
x = 61 9 16 lm, n = 15), 8-spored, bitunicate, cylindrical to clavate, with short pedicel, tapering to a rounded
or irregular base, thick-walled. Ascospores 15–21 9 5–10
lm (
x = 19 9 7 lm, n = 20), mostly ellipsoidal, muriform, usually with 3–4 transverse septa and 1–2 longitudinal septa, with slightly paler ends, conical and narrowly
rounded at the ends, hyaline, with mucilaginous sheath.
Asexual morph Undetermined.
123
Culture characteristics: Colonies on MEA at 16 °C
reaching 4 cm in two weeks, circular with undulate, yellow-grey mycelium, velvety and flat on the media.
Material examined: ITALY, Forlı̀-Cesena Province,
Castrocaro Terme, dead branch of Cornus sanguinea
(Cornaceae), 6 January 2012, Erio Camporesi, IT86
(MFLU 15-1476, HKAS 94526, new host record); living
culture, MFLUCC 14-0520).
GenBank Numbers LSU:KY496718; ITS:KY496739;
SSU:KY501109; RPB2:KY514403.
Notes: Phylogenetically this species resides in a distinct
subclade in Hysterobrevium and is H. mori (Fig. 10). Our
Fungal Diversity
Fig. 13 Hysterobrevium mori (MFLUCC 14-0520). a Appearance of
ascomata on host substrate. b Section of ascoma. c, d Section of
peridium. e Pseudoparaphyses. f–h Asci. i–k Ascospores.
l Germinated ascospore. Scale bars a = 200 lm, b = 100 lm,
c = 10 lm, d = 20 lm, e = 2 lm, f–h = 20 lm, i–l = 5 lm
new isolate from Italy matches H. mori in having ellipsoidal, hyaline, muriform ascospores, with 3–4 transverse
septa and 1–2 longitudinal septa that pass through one to
two cells, with a mucilaginous sheath. Thus, we confirm
(based on both morphological characteristics and phylogenetic data) that our taxon belongs to H. mori even though
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Angustimassarina Thambug., Kaz. Tanaka & K.D. Hyde
Thambugala et al. (2015a) introduced Angustimassarina
to accommodate three new ascomycetous species placed in
the family Amorosiaceae. The members of the genus are
considered as fungicolous or they may be parasitic on other
fungi and appear to grow within other ascomata or on other
ascomycetes. A synopsis of Angustimassarina species is
provided (Table 2).
with a minute ocular chamber. Ascospores 19–22 9 3–4
lm diam. (
x = 21 9 3.5 lm, n = 30), uni to biseriate,
partially overlapping, hyaline, fusiform to cylindrical or
ellipsoidal-fusiform, straight to curved, widest at the centre
and tapering towards the ends, 3-septate, constricted at the
primary septum, smooth-walled, filled with two different
sized guttules per cell and surrounded by a mucilaginous
sheath. Asexual morph Undetermined.
Material examined: GERMANY, loamy sand, acid,
fresh, ditch with alder, 40 m a.s.l., mesotroph, twig of Alnus glutinosa (Betulaceae), 1 March 2014, Rene K.
Schumacher 044, (MFLU 16-2981, holotype); ex-type
living culture, MFLUCC 15-0184, KUMCC 16-0071.
GenBank Numbers LSU:KY548097; SSU:KY548098;
ITS:KY548099.
Culture characteristics: Ascospores germinating on WA
within 24 h. Colonies growing on MEA, reaching 5 mm
diam. in 1 week at 18 °C. Mycelium superficial, slightly
effuse, ashy white, inner brown, sparsely hairy, with
undulate edge, lower surface black.
Notes: Angustimassarina alni fits with generic concept
of Angustimassarina (Thambugala et al. 2015a). Angustimassarina alni forms a sister clade with Exosporium stylobatum
(CBS
16030;
asexual
morph)
and
A. premilcurensis (MFLUCC 15-0074. We could not
obtain an asexual morph in culture. Angustimassarina alni
is similar to A. premilcurensis in having immersed to semiimmersed ascomata, cylindrical to clavate asci and hyaline,
fusiform to cylindrical ascospores with a mucilaginous
sheath, but differs in having a carbonaceous peridium and
different sized ascospores with guttules.
Angustimassarina alni Jayasiri & K.D. Hyde, sp. nov.
Index Fungorum number: IF552551; Facesoffungi
number: FoF 2689, Fig. 15
Etymology: The specific epithet alni is based on the host
genus.
Holotype: MFLU 16-2981
Saprobic on Alnus glutinosa (L.) Gaertn. Sexual morph
Ascomata 160–250 lm high 9 130–200 lm diam.
(
x = 208 9 188 lm, n = 5), scattered to gregarious,
immersed to semi-immersed, carbonaceous, dark brown to
black, globose to subglobose. Ostiolar canal inner layer
lined with black to dark brown cells, outer layer with dark
brown to black pseudoparenchymatous cells, with a pore
like opening or opening through cracks in the host surface.
Peridium 28–44 lm wide, equally thick, comprising dark
brown to black cells of textura angularis. Hamathecium
comprising 1–1.5 lm septate, unbranched, cellular pseudoparaphyses, embedded in gelatinous matrix, between and
above the asci. Asci 71–89 9 8–10 lm (
x = 82 9 9 lm,
n = 15), 8-spored, bitunicate, fissitunicate, cylindric-clavate, with furcate or knob-like pedicel, rounded at the apex
Angustimassarina arezzoensis Tibpromma, Camporesi &
K.D. Hyde, sp. nov.
Index Fungorum number: IF552690; Facesoffungi
number: FoF 2769, Fig. 16
Etymology: refers to the Arezzo (Italy) where the fungus
was collected.
Holotype: MFLU 14-0681
Saprobic on dead stem of Salvia sp. Sexual morph
Ascomata 169–234 lm high 9 166–245 lm diam.
(
x = 208 9 188 lm, n = 4), immersed to erumpent, visible as raised, dark spots on the host surface, uniloculate,
subglobose, solitary or in small groups, scattered on the
host surface, without papilla, short ostiole in the center,
thick-walled, carbonaceous, dark brown or usually black.
Peridium 22–41 lm wide, inner cells hyaline to pale
brown, outer layer cells brown to dark brown, composed of
flattened cells of textura angularis. Hamathecium 1.6–2.9
lm wide, comprising numerous, septate, cylindrical, frequently anastomosing, cellular, pseudoparaphyses. Asci
67–95 9 10–15 lm (
x = 82 9 13 lm, n = 15), 8-spored,
bitunicate, fissitunicate, broadly cylindrical to cylindric-
it was found on a different host (Cornus sanguinea). Previously, H. mori has been reported from many hosts, such
as Acer, Amelanchier, Aspidosperma, Castanea, Cercocarpus, Cotinus, Crataegus, Fraxinus, Gleditsia, Juniperus, Melia, Morus, Olea, Ostrya, Pinus, Pistacia, Prunus,
Pyrus, Quercus, Rhus, Rubus, Salix, Ulmus, various
Fabaceae, Vitis and Ziziphus (Zogg 1962).
Pleosporales Luttr. ex M.E. Barr
For an account of Pleosporales Luttr. ex M.E. Barr see
Zhang et al. (2012b) and Hyde et al. (2013).
Amorosiaceae Thambugala & K.D. Hyde
Amorosia was introduced by Mantle et al. (2006) with
A. littoralis Mantle & D. Hawksw. Previous studies, based
on phylogenetic analysis, have referred it to the
Sporormiaceae (Mantle et al. 2006), but Thambugala et al.
(2015a) introduced Amorosiaceae as a new family for this
species and described three new species of Angustimassarina in this family. We introduce six new species of
Angustimassarina based on morphology and phylogeny
(Fig. 14).
123
Fungal Diversity
Fig. 14 Phylogram generated from maximum parsimony analysis of
combined ITS, SSU, LSU and TEF1 sequence data for Lophiostomataceae and Amorosiaceae. Parsimony bootstrap support values for
MP C 70% and Bayesian posterior probabilities C0.8 are indicated at
the nodes. Isolate numbers of all ex-types and reference strains are in
bold. Novel species/new host records are indicated in blue. The tree is
rooted with Melanomma pulvis-pyrius
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Fungal Diversity
Table 2 Synopsis of ascomata, peridium, asci and ascospores of Angustimassarina species
Taxa
Ascomata (lm)
Peridium (lm)
Asci (lm)
Ascospores (lm)
References
Angustimassarina acerina
MFLUCC 14-0505
200–350 9 164–183
15–26
92–105 9 7.5–8.6
21–23 9 4.1–4.6
Thambugala
et al. (2015a)
Angustimassarina alni
MFLUCC 15-0184
160–250 9 130–200
28–44
71–89 9 8–10
19–22 9 3–4
This study
Angustimassarina
arezzoensis MFLUCC
13-0578
169–234 9 166–245
22–41
67–95 9 10–15
19–21 9 5–6
This study
Angustimassarina italica
MFLUCC 15-0082
127–159 9 97–131
23–40
78–103 9 10–12
15–22 9 3–6
This study
Angustimassarina lonicerae
MFLUCC 15-0087
193–203 9 170–220
10–18
55–81 9 9–13
19–25 9 4–7
This study
Angustimassarina populi
MFLUCC 13-0034
125–175 9 100–120
14–32
80–95 9 9.5–13
19–22 9 3.2–5.5
Thambugala
et al. (2015a)
Angustimassarina
premilcurensis MFLUCC
15-0074
231–238 9 290–311
20–30
64–93 9 11–15
19–23 9 4–7
This study
Angustimassarina
quercicola MFLUCC
14-0506
200–250 9 150–265
14–27
60–94 9 8.8–13
17–21 9 4–6
Thambugala
et al. (2015a)
clavate, with short bulbous, furcate pedicel, rounded at the
apex, with a poorly develop ocular chamber. Ascospores
19–21 9 5–6 lm (
x = 20 9 6 lm, n = 20), overlapping
1–2-seriate, hyaline, fusiform, usually 3-septate, narrowly
fusoid with rounded ends, constricted at the septum, deeply
constricted at the central septum, enlarged at the second
cell, guttulate, smooth-walled, surrounded by mucilaginous
sheath. Asexual morph Undetermined.
Culture characteristics: Colonies on MEA at 16 °C
reaching 1.5 cm in 1 week, irregular, whiteish-grey, with
undulate edge, lower surface black.
Material examined: ITALY, near Poppi Province, Arezzo, dead stem of Salvia sp. (Lamiaceae), 4 June 2013,
Erio Camporesi, IT1321 (MFLU 14-0681, holotype); extype living culture, MFLUCC 13-0578; (HKAS 94552 bis,
paratype).
GenBank Numbers LSU:KY496722; ITS:KY496743;
SSU:KY501113; TEF1:KY514392.
Notes: Angustimassarina arezzoensis clustered with
A. populi in phylogeny (Fig. 14), but they are morphologically different (Table 2).
Angustimassarina lonicerae Tibpromma, Camporesi &
K.D. Hyde, sp. nov.
Index Fungorum number: IF552691; Facesoffungi
number: FoF 2770, Fig. 17
Etymology: named for its occurrence on the host plant
(Lonicera sp.).
Holotype: MFLU 15-1485
Saprobic on dead branch of Lonicera sp. Sexual morph
Ascomata 193–203 lm high 9 170–220 lm diam.
123
(
x = 198 9 201 lm, n = 5), semi-immersed to erumpent
through host tissue, solitary, globose to subglobose, visible
on host surface as raised ostiolar dots, black. Ostioles
central, long, conspicuous at the surface, shiny, black.
Peridium 10–18 lm wide, thick at the sides, broad at the
apex and thinner at the base, comprising brown to dark
brown cells of textura angularis, fusing at the outside with
the host tissues. Hamathecium comprising 0.7–1.1 lm
wide, dense, cylindrical, septate, branched, pseudoparaphyses. Asci 55–81 9 9–13 lm (
x = 71 9 12 lm, n = 15),
8-spored, bitunicate, cylindrical, short pedicellate, apex
rounded, with a minute ocular chamber. Ascospores 19–25
9 4–7 lm (
x = 22 9 6 lm, n = 15), overlapping 1–2seriate, fusiform, 1–3-septate, guttulate, enlarged at the
second cell, constricted at the center, conical at the ends,
smooth-walled, hyaline, surrounded by a mucilaginous
sheath. Asexual morph Undetermined.
Culture characteristics: Colonies on MEA reaching 6
cm diam., after 7 days in the dark at 16 °C, irregular, grey,
with undulate edge, lower surface black.
Material examined: ITALY, Forlı̀-Cesena Province,
Fiumicello di Premilcuore, on dead branch of Lonicera sp.
(Caprifoliaceae), 19 November 2013, Erio Camporesi,
IT1524 (MFLU 15-1485, holotype); ex-type living culture,
MFLUCC 15-0087; (HKAS 94563 bis, paratype).
GenBank Numbers LSU:KY496724; ITS:KY496759.
Notes: Angustimassarina lonicerae is similar to A. italica however the peridium in A. lonicerae is 10–18 lm
wide, with 1-septate ascospores swollen above centre
septum, versus 23–40 lm wide in A. italica, and 1–3septate ascospores enlarged at the second cell. The
Fungal Diversity
Fig. 15 Angustimassarina alni (MFLU 16-2981, holotype). a, b Appearance of ascomata on host surface. c Ostiole. d Section through
ascomata. e Peridium. f Pseudoparaphyses. g, h Asci. i–k Ascospores. Scale bar c, e = 20 lm, d = 50 lm, f = 5 lm, g, h = 20 lm, i–k = 5 lm
phylogeny (Fig. 14) also supports this new species as distinct as they cluster in different subclades within
Amorosiaceae with good statistical support (86% in ML/
1.00 in BYPP)
Angustimassarina premilcurensis Tibpromma, Camporesi
& K.D. Hyde, sp. nov.
Index Fungorum number: IF552692; Facesoffungi
number: FoF 2771, Fig. 18
123
Fungal Diversity
Fig. 16 Angustimassarina
arezzoensis (MFLU 14-0681,
holotype). a Appearance of
ascomata on host surface.
b Cross section of ascoma.
c Peridium.
d Pseudoparaphyses. e–h Asci.
i–k Ascospores. l Germinated
ascospore. Scale bars a,
b = 100 lm, c = 20 lm, d = 2
lm, e–h = 20 lm, i–l = 5 lm
Etymology: refers to the Fiumicello di Premilcuore
(Italy) where the fungus was collected.
Holotype: MFLU 14-0703
Saprobic on dead branch of Carpinus betulus L. Sexual
morph Ascomata 231–238 lm high 9 290–311 lm diam.
123
(
x = 235 9 326 lm, n = 5), immersed, solitary or scattered on the host surface, globose to subglobose, black,
without papilla, ostiole in the center. Peridium 20–30 lm
wide, comprising 2–4 layers of irregular, brown-walled
cells arranged in a textura angularis. Hamathecium
Fungal Diversity
Fig. 17 Angustimassarina lonicerae (MFLU 15-1485, holotype). a,
b Appearance of ascomata with ostiole on host substrate. c Section of
ascoma. d Section of peridium. e Pseudoparaphyses. f, g Asci. h–
k Ascospores. l Germinated ascospore. Scale bars a = 100 lm, b,
c = 50 lm, d = 10 lm, e = 2 lm, f, g = 20 lm, h–l = 5 lm
comprising 0.5–0.9 lm, cylindrical, filamentous, septate,
pseudoparaphyses. Asci 64–93 9 11–15 lm (
x = 77 9 13
lm, n = 10), 8-spored, bitunicate, cylindrical to cylindricclavate, short pedicellate or sessile, rounded at the apex,
with ocular chamber. Ascospores 19–23 lm 9 4–7 lm
(
x = 21 9 6 lm, n = 15), overlapping bi-seriate, fusiform,
1-septate, constricted at the centre, straight or slightly
curved, swollen near the septum, surrounded by
123
Fungal Diversity
Fig. 18 Angustimassarina premilcurensis (MFLU 14-0703, holotype). a, b Appearance of ascomata with erumpent ostiole. c Section of
ascoma. d Section of peridium. e Pseudoparaphyses. f–h Ascus with
123
pedicel. i Ocular chamber. j–m Ascospores. n Germinated ascospore.
Scale bars a = 500 lm, b = 200 lm, d = 20 lm, e = 2 lm, f–
i = 10 lm, j–n = 5 lm
Fungal Diversity
mucilaginous sheath, hyaline, smooth-walled. Asexual
morph Undetermined.
Culture characteristics: Colonies on MEA reaching 4
cm diam., after 7 days in the dark at 16 °C, irregular,
white–grey, flat on the surface, lower surface black.
Material examined: ITALY, Province of Forlı̀-Cesena
[FC], Fiumicello di Premilcuore, on dead branch of
Carpinus betulus L. (Betulaceae), 10 February 2014, Erio
Camporesi, IT1716 (MFLU 14-0703, holotype); ex-type
living culture, MFLUCC 15-0074; (HKAS 94575 bis,
paratype).
GenBank Numbers LSU:KY496725; ITS:KY496745;
RPB2:KY514404.
Notes: Angustimassarina premilcurensis differs from
other species in Angustimassarina in possessing 1-septate
ascospores, constricted and swollen near the septum. In the
phylogenetic analyses, A. premilcurensis clustered with
Exosporium stylobatum. Exosporium is a hyphomycete
(Crous et al. 2011) with more than 120 epithets (Index
Fungorum 2017), but DNA sequence data are not available
for its type species (Thambugala et al. 2015a). Any conclusive association between A. premilcurensis and Exosporium stylobatum is premature until further DNA
sequence data becomes available.
Angustimassarina italica Tibpromma, Camporesi & K.D.
Hyde, sp. nov.
Index Fungorum number: IF552707; Facesoffungi
number: FoF 2785, Fig. 19
Etymology: refers to the country (Italy) where the fungus
was collected.
Holotype: MFLU 15-1494
Saprobic on dead branch of Ilex aquifolium L. Sexual
morph Ascomata 127–159 lm high 9 97–131 lm diam.
(
x = 143 9 114 lm, n = 5), semi-immersed to erumpent
through host tissue, solitary, uniloculate, globose to subglobose, black, without hairs, with long ostiole in the
center. Peridium 23–40 lm wide, comprising several layers
of hyaline to pale brown cells of textura angularis. Hamathecium 1.1–1.9 lm wide, comprising numerous, aseptate, unbranched, pseudoparaphyses. Asci 78–103 9 10–12
lm (
x = 89 9 11 lm, n = 15), 8-spored, bitunicate,
cylindrical with bulbous pedicel, rounded at the apex, with
a well-developed ocular chamber. Ascospores 15–22 9 3–6
lm (
x = 19 9 4 lm, n = 15), overlapping 1-seriate,
hyaline, 1-septate, constricted at centre, swollen above
centre septum, conical at the ends, guttulate, surrounding
by a large mucilaginous sheath, smooth-walled. Asexual
morph Undetermined.
Culture characteristics: Colonies on MEA reaching 5–6
cm diam., after 7 days in the dark at 16 °C, irregular,
white–grey to black, flat on the surface, lower surface
black.
Material examined: ITALY, Campigna—Santa Sofia, on
dead branch of Ilex aquifolium (Aquifoliaceae), 10 October
2014, Erio Camporesi, IT2159 (MFLU 15-1494, holotype); ex-type living culture MFLUCC 15-0082; (HKAS
94605 bis, paratype).
GenBank Numbers LSU:KY496736; ITS:KY496756;
SSU:KY501124; TEF1:KY514400.
Notes: Angustimassarina italica is phylogenetically
closely related to A. acerina and A. quercicola (Fig. 14).
However, it differs in having hyaline, 1-septate ascospores
while A. quercicola has brown, 1(–3)-septate ascospores,
which are finely verruculose at maturity and A. acerina has
brown, 1(–3)-septate ascospores (Thambugala et al.
2015a).
Biatrosporaceae K.D. Hyde
Jaklitsch and Voglmayr (2016) dismissed the family
Biatriosporaceae, whose type clustered amongst Nigrograna species, and transferred several species from Biatriospora to Nigrograna. They introduced the family
Nigrogranaceae and accepted a single genus Nigrograna
based on morphology and phylogenetic analyses and provided DNA sequence data. Jaklitsch and Voglmayr (2016)
doubted the validity of the strain of B. marina in GenBank.
However, based on the type species, the family Biatriosporaceae is morphologically distinct and the family
should be maintained. Nevertheless, we accept that it
seems odd that the strain of Biatriospora marina in GenBank clusters with Nigrograna species and thus B. marina
needs recollecting for sequence analysis to establish if
Nigrograna is related, which morphologically seems unlikely. A phylogenetic tree for Nigrogranaceae is presented
in this paper (Fig. 59), where the continued use of Biatriosporaceae, which is unlike Nigrogranaceae based on
its morphological distinctiveness, is recommended.
Lophiostomataceae Sacc.
The family Lophiostomataceae is an interesting and
diverse family in the order Pleosporales, Dothideomycetes.
Most members of this family are saprobic and occur mainly
on twigs, stems or bark of various woody plants and
herbaceous plants in both terrestrial and aquatic environments (Hirayama and Tanaka 2011; Hyde et al. 2013;
Ariyawansa et al. 2015c; Thambugala et al. 2015a).
Thambugala et al. (2015a) revised the family and accepted
16 genera, including eleven newly introduced genera. In
this study, we place two new species and two new host
records in Lophiostomataceae (Figs. 14, 20).
Berkleasmium Zobel.
Zobel (1854) established the genus Berkleasmium by recombining the species Sporidesmium concinnum Berk., and
renamed the taxon in honour of Corda, as Berkleasmium
123
Fungal Diversity
Fig. 19 Angustimassarina italica (MFLU 15-1494, holotype).
a Appearance of ascomata on host substrate. b Section of ascoma.
c Section of peridium. d Pseudoparaphyses. e, f Asci. g, h
123
Ascospores. i Ascospore in Indian ink to show sheath. j Germinating
ascospore. Scale bars b = 40 lm, c = 20 lm, d = 2 lm, e, f = 20
lm, g–j = 5 lm
Fungal Diversity
Fig. 20 Phylogenetic tree generated from neighbor joining analysis
of LSU sequence data from Zhang et al. (2009a) with Tubeufiaceae
(Tubeufia javanica and Tubeufia paludosa) as out group taxa.
Bootstrap support values for maximum likelihood greater than 50%
are given above the nodes. New sequences are in blue
cordeanum. Hughes (1958) later corrected this type species
name as Berkleasmium concinnum. Moore (1958) and Ellis
(1971) accepted this genus for sporodochial taxa that produce dark dictyosporous conidia on short simple unbranched conidiophores or directly on the hyphae. Fourty
species are currently listed as legitimate under Berkleasmium in MycoBank (2017). According to Seifertk et al.
(2011) the genus Berkleasmium comprises 32 valid species
recorded from Africa, Asia, Australasia and North America. Most of these species are recorded from dead substrates, such as dead wood, litter and bark of various plants
(Moore 1958; Rao and Rao 1963; Ellis 1971, 1976; Morris
1972; Chouhan and Panwar 1980; Matsushima 1981, 1983;
Holubová-Jechová 1987; Yip 1988; Bussaban et al. 2001;
Pinnoi et al. 2007; Qu et al. 2014; Hüseyin et al. 2014). The
new species B. ariense from India is similar to the type
species B. concinnum, however, the conidia are longer.
Three species of Berkleasmium were previously recorded
from India; B. abuense (current name: Monodictys
abuensis), B. osmaniae and B. papillatum (current name:
Acrodictys papillata). Phylogenetic affinities of the new
taxon with other Berkleasmium species, based on the LSU
and ITS gene sequences, is shown in Fig. 20.
Berkleasmium ariense Rajeshkumar & Marathe, sp. nov.
MycoBank number: MB 819049; Facesoffungi number:
FoF 2779; Fig. 21
Etymology: ariense, referring to the Agharkar Research
Institute, a reputed school of mycology in India.
Holotype: AMH 9828
Saprobic on natural substrates. Sexual morph Undetermined. Asexual morph Conidiomata sporodochial, raised,
superficial, scattered, dark brown to blackish, velvety, with
immersed mycelia. Stroma rudimentary. Setae and
hyphopodia absent. Conidiophores micronematous, short,
closely packed together into sporodochia, flexuous, unbranched, 2–5 lm. Conidiogenous cells monoblastic, integrated,
terminal, determined. Conidia 119–238 9 41–60 lm, solitary, dry or sticky, acrogenous, simple, clavate, ellipsoidal,
oblong or irregular, muriform, pale to dark brown or blackishbrown, hilum not protruding; conidial base 11–15 lm. Young
conidia hyaline or subhyaline, 56 9 17 lm.
123
Fungal Diversity
Fig. 21 Berkleasmium ariense
(AMH 9828, holotype). a–
c Habit. d, e Conidia. f–i SEM
of sporodochia and conidia.
Scale bars d, e = 50 lm, f–
h = 20 lm, i = 15 lm
Culture characteristics: Colonies on MEA slow to
medium growing, 30 to 45 mm diam. after 20 days, dark
brownish-black, mycelia immersed, centrally fluffy, margin irregular, reverse dark brown to blackish. No sporulation observed.
123
Material examined: INDIA, Maharashtra, Tamhini Village, on unidentified stem litter, 9 June 2015 (AMH 9828,
holotype); ex-type living culture NFCCI 4026; additional
culture NFCCI 4027.
Fungal Diversity
GenBank Numbers ITS:KY039163, KY039164;
LSU:KY039165, KY039166.
Notes: Our new isolate is morphologically similar and
phylogenetically related to Platystomum rosae (strain
MFLUCC 15-0633) (Fig. 14), but associated with a different host. Platystomum rosae (holotype) was recorded on
Rosa canina L (Rosaceae) and our collection was found on
Cornus sp. (Cornaceae).
Platystomum Trevis.
Platystomum was introduced by Trevisan (1877) and is
typified by Platystomum compressum. Platystomum compressum was reported as a synonym of Lophiostoma
compressum and Thambugala et al. (2015a) accepted
Platystomum in Lophiostomataceae as a distinct genus, and
accepted five species (Platystomum actinidiae, P. compressum, P. crataegi, P. rosae and P. salicicola), based on
new collections and combinations (Fig. 14).
Platystomum rosae Wanas., Thambug., Camporesi & K.D.
Hyde, Fungal Diversity 74: 234 (2015)
Facesoffungi number: FoF 2780, Fig. 22
Saprobic on species of Cornus sp., in terrestrial environments. Sexual morph Ascomata 300–500 lm high 9
250–450 lm diam. (
x = 424.7 9 305.3 lm, n = 5), scattered to gregarious, immersed, coriaceous, dark brown to
black, subglobose to conical, ostiolate. Ostioles 150–200
lm long, 40–60 lm diam. (
x = 175.4 9 49.1 lm, n = 10),
papillate, black, smooth, with short, setae. Peridium 20–25
lm wide at the base, 25–35 lm wide at the sides, broad at
the apex, two–layered, outer layer composed of small,
blackish to dark brown, thick-walled cells of textura prismatica, inner layer composed of smaller, compressed,
hyaline cells. Hamathecium of 1.5–3 lm wide, septate,
cellular pseudoparaphyses, embedded in gelatinous matrix
between and above the asci. Asci 120–170 9 15–20 lm
(
x = 144.9 9 17.1 lm, n = 15), 8-spored, bitunicate, fissitunicate, cylindrical to cylindric-clavate, pedicel, narrowly rounded at the apex, with an indistinct ocular
chamber. Ascospores 19–23 9 7–9 lm (
x = 21.2 9 8.4
lm, n = 40), overlapping 1–2-seriate, muriform, mostly
ellipsoidal, with 3–4 transverse septa with 1 vertical septum in each row, slightly curved, deeply constricted at the
centre septum, slightly constricted at the remaining septa,
initially hyaline, becoming brown at maturity, rounded at
the ends, without a mucilaginous sheath. Asexual morph
Undetermined.
Culture characteristics: on PDA, reaching 25–30 mm
diam. after 21 days at 16 °C, surface dirty green, spreading
with moderate aerial mycelium, and even, smooth margins;
reverse dark green.
Material examined: ITALY, Province of Forlı̀-Cesena,
near Fiumicello di Premilcuore, on Cornus sp.
(Cornaceae), 11 January 2015, Erio Camporesi, IT 2333
(MFLU 15-2569, new host record).
GenBank Numbers LSU:KY264746; ITS:KY264742;
SSU:KY264750.
Notes: Our new isolate is morphologically similar and
phylogenetically related to Platystomum rosae (strain
MFLUCC 15-0633), but associated with a different host.
Platystomum rosae (holotype) was recorded on Rosa canina L (Rosaceae), while our collection was found on
Cornus sp. (Cornaceae). The phylogenetic placement of
this species is shown in Fig. 14.
Sigarispora Thambug. & K.D. Hyde.
Thambugala et al. (2015a) introduced the genus in
Lophiostomataceae and it is typified by S. ravennica
(Tibpromma et al.) Thambugala & K.D. Hyde. Sigarispora
is characterized by immersed to semi-immersed ascomata,
a small crest-like ostiole, and brown, cigar-shaped or
ellipsoidal-fusiform, multi-septate or muriform ascospores.
Seven species are accepted in the genus.
Sigarispora muriformis Tibpromma, Camporesi & K.D.
Hyde, sp. nov.
Index Fungorum number: IF552696; Facesoffungi
number: FoF 2781, Fig. 23
Etymology: refers to the muriform ascospores.
Holotype: MFLU 14-0676
Saprobic on dead stem of Helichrysum italicum (Roth)
G. Don fil. Sexual morph Ascomata 425–660 lm high
(including ostiole) 9 335–560 lm diam. (
x = 506 9 460
lm, n = 10), semi-immersed on host tissue, solitary,
scattered, globose to subglobose, conspicuous at the surface, papillate, carbonaceous. Ostioles central, lacking
periphyses in ostiole canal, dark, dull. Peridium 25–47 lm
wide, coriaceous to carbonaceous, comprising an outer
stratum of thick-walled and black cells arranged in a textura angularis and fusing with host cells, and inner layer of
hyaline cells of textura angularis. Hamathecium of 1.9–3.2
lm wide, cylindrical, septate, branched, guttulate, cellular
pseudoparaphyses. Asci 84–130 9 12–18 lm (
x = 104 9
15 lm, n = 10), 8-spored, bitunicate, cylindric-clavate,
slightly curved, long pedicellate, with apex rounded, with a
minute ocular chamber. Ascospores 18–28 9 7–12 lm
(
x = 24 9 9 lm, n = 20), overlapping 2–3-seriate, muriform, ellipsoidal, hyaline then become reddish–brown with
age, with 5–8 transverse septa and 2–3 longitudinal septa,
guttulate, constricted at the centre septum, conical at the
ends, smooth-walled, lacking a mucilaginous sheath.
Asexual morph Undetermined.
Culture characteristics: Colonies on MEA reaching 8
cm diam., after 7 days in the dark at 16 °C, surface darkbrown, spreading with moderate aerial mycelium, and
even, margins smooth; reverse dark.
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Fig. 22 Platystomum rosae (MFLU 15-2569). a Appearance of ascomata on host substrate. b Section of ascoma. c Section of peridium.
d Pseudoparaphyses. e, f Asci. g–l Ascospores. Scale bars b = 100 lm, c = 20 lm, d = 10 lm, e, f = 20 lm, g–l = 10 lm
Material examined: ITALY, Forlı̀-Cesena Province,
near Premilcuore, on dead stem of Helichrysum italicum
(Asteraceae), 24 May 2013, Erio Camporesi, IT1302
(MFLU 14-0676, holotype); ex-type living culture,
MFLUCC 13-0744); Ibid. (HKAS 94549 bis, paratype).
GenBank Numbers LSU:KY496719; ITS:KY496740;
SSU:KY501110.
Notes: The new species clearly differs from other
Sigarispora species in having semi-immersed ascomata,
123
cylindric-clavate asci and ellipsoidal, muriform ascospores, with 5–8 transverse septa and 2–3 longitudinal
septa. This species separates from other Sigarispora
species in the phylogenetic analyses (Fig. 14). Sigarispora coronillae and S. muriformis have muriform
ascospores, but S. coronillae differs from S. muriformis
in having brown to dark brown ascospores with 4–6
transverse septa and 2–4 longitudinal septa (Thambugala
et al. 2015a).
Fungal Diversity
Fig. 23 Sigarispora muriformis (MFLUCC 13-0744, holotype).
a Appearance of papilla of erumpent ascomata on host substrate.
b Section of ascoma. c Section of peridium. d Pseudoparaphyses. e–
g Asci. h–k Ascospores. l Germinated ascospore. Scale bars a,
b = 200 lm, c, d = 5 lm, e–h = 10 lm, f, g = 20 lm, h–l = 10 lm
Vaginatispora K.D. Hyde
Vaginatispora was introduced by Hyde (1995) in Massarinaceae Munk to accommodate Vaginatispora aquatica
K.D. Hyde. Vaginatispora was long considered to be a
synonym of Massarina (Hyde et al. 1992; Read et al. 1997)
but recently Thambugala et al. (2015a) confirmed it as a
separate genus in Lophiostomataceae based on both morphological characteristics and phylogeny. Vaginatispora is
characterized by ‘depressed globose ascomata, immersed
beneath a blackened neck, with a slot-like ostiole,
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numerous and filamentous pseudoparaphyses, cylindrical to
clavate asci and narrowly ellipsoidal, hyaline, 2-celled
ascospore with a mucilaginous collar around its equator
and a spreading papilionaceous sheath’ (Hyde 1995; Zhang
et al. 2014). Currently the genus comprises Vaginatispora
appendiculata, V. aquatica, V, armatispora and V. fuckelii.
Vaginatispora appendiculata Wanas., E.B.G. Jones &
K.D. Hyde, Studies in Fungi 1 (1): 60 (2016)
Facesoffungi number: FoF 2782, Fig. 24
Saprobic on Calamus rotang L. in terrestrial habitats.
Sexual morph Ascomata 250–300 lm high, 360–400 lm
diam. (
x = 266 9 389 lm, n = 5), scattered to gregarious,
immersed, coriaceous to carbonaceous, black, subglobose,
ostiolate. Necks 70–90 lm long, 30–50 lm diam., long
slot-like, black, filled with dark brown cells. Peridium 9–12
lm wide at the base, 30–40 lm wide at the sides, comprising 3–4 layers, fused at the outside with the host tissue,
composed of thick-walled reddish-brown to dark angular or
relatively compressed pseudoparenchymatous cells. Hamathecium of 1–1.5 lm wide, filiform, septate, anastomosing, branched, pseudoparaphyses. Asci 115–130 9 14–
18 lm (
x = 123.6 9 15.19 lm, n = 20), 8-spored, bitunicate, fissitunicate, cylindrical to cylindric-obclavate,
short pedicellate, apically rounded. Ascospores 40–45 9 6–
8 lm (
x = 42.7 9 7.2 lm, n = 30), 2–3-seriate, fusiform,
narrowing towards the acute ends, 1-septate, slightly wide
above and below the septa, constricted at the septum,
hyaline, smooth-walled, guttulate, surrounded by a distinct
mucilaginous sheath. Asexual morph Undetermined.
Culture characteristics: on MEA reaching 2 cm diam.
after 2 weeks at 25 °C, dirty white at the beginning and
blackish-brown at maturity, convex on the surface, undulate, margins smooth; reverse blackish-green.
Material examined: THAILAND, Chiang Rai Province,
Chiang Sean, on dead Calamus rotang stem (Arecaceae),
21 September 2013, Dhanushka Wanasinghe, DHA17
(MFLU 16-2887, new host record); living culture,
MFLUCC 13-0835.
GenBank Numbers LSU:KY264745; SSU:KY264749
(Fig. 25).
Notes: As morphological characters examined largely
overlap with those of Vaginatispora appendiculata, we
report our collection as a new host record of V. appendiculata from Calamus rotang in Thailand. Combined multigene based phylogenies also show that both strains
(MFLUCC 13-0835 and MFLUCC 11-0083) of V. appendiculata cluster together, but with a low bootstrap support
(Fig. 14).
Lophiotremataceae K. Hiray. & Kaz. Tanaka.
Hirayama and Tanaka (2011) introduced the family
Lophiotremataceae with its type genus Lophiotrema Sacc.
123
Phylogenetic analyses based on LSU and SSU sequence
data supported this new familial placement and its distinctiveness from Lophiostomataceae. Lophiotremataceae
comprises Aquasubmersa, Hermatomyces and Lophiotrema
(Hyde et al. 2016; Doilom et al. 2017). The genus Hermatomyces was introduced by Spegazzini (1911), with
H. tucumanensis as the type species and includes 13 species (Index Fungorum 2017). Morphologically, the genus is
characterized by lenticular to cylindrical, muriform conidia, often with subhyaline to pale brown peripheral cells,
and dark brown central cells. Conidia are cylindrical and
comprise 1–4 columns with 2–11 cells and are irregularly
pigmented (Castañeda and Heredia 2000; Leão-Ferreira
et al. 2013; Doilom et al. 2017).
Lophiotrema Sacc.
Lophiotrema was proposed by Saccardo (1878) to
accommodate species having hyaline ascospores and typified by Lophiotrema nucula, which was excluded from
Lophiostoma based on ascospore characteristics. This
genus includes many lignicolous bitunicate fungi characterized by immersed-erumpent through woody substrates,
dark-pigmented ascomata with ostiolate papilla and fusiform, 1-septate and hyaline ascospores (Barr 1992; Tang
et al. 2003; Hirayama and Tanaka 2011). Currently, 161
epithets are listed in Index Fungorum, but only 97 species
were accepted in Lophiotrema based on current morphological data and phylogenetic evidence (Hyde et al. 2016;
Doilom et al. 2017).
Lophiotrema guttulata Boonmee, Tibpromma & K.D.
Hyde, sp.nov.
Index Fungorum number: IF552688; Facesoffungi
number: FoF02735; Figs. 26, 27
Etymology: ‘guttulata’ referring to the guttules in this
species.
Holotype: MFLU 10-0971
Saprobic on dried terrestrial branch. Sexual morph
Ascomata 248.5–310 lm high 9 271–340 lm diam.
(
x = 279 9 305.5 lm, n = 5), uniloculate, immersed,
scattered, subglobose, dark brown, ostiolate. Ostioles 98–
137 9 29–53 lm diam. (
x = 124 9 37 lm, n = 5),
plugged by periphyses, apically with a crest-like papilla,
dark, carbonaceous, fragile, with irregular opening.
Peridium 23–38.5 lm wide, outer layer narrow, composed
of dark cells of textura angularis, inner layer of light
pigmented to hyaline cells of textura subglobosa. Hamathecium comprising 1–2 lm wide, branched, septate,
cylindrical pseudoparaphyses. Asci (78-)90–125 9 8–11.5
lm (
x = 96.5 9 9 lm, n = 20), 8-spored, bitunicate, fissitunicate, cylindric-clavate, sessile to short pedicellate,
apically rounded with an ocular chamber. Ascospores 23–
29 9 4–6 lm (
x = 26 9 5 lm, n = 10), uniseriate,
Fungal Diversity
Fig. 24 Vaginatispora appendiculata (MFLU 16-2887). a Appearance of ascomata on host substrate. b Section of ascoma. c Close up
of ostiole d Section of peridium. e Pseudoparaphyses. f–h Asci. i–
m Ascospores (note the ascospore stained in Indian ink to show the
mucilaginous sheath in m). n Germinated ascospore. Scale bars
a = 500 lm, b = 100 lm, c = 50 lm, d = 20 lm, e = 10 lm, f–
h = 20 lm, i–n = 10 lm
ellipsoidal to broadly fusiform, 1-septate, constricted at the
septum, with 1–2 guttules, with distinct large guttules in
each cell, with terminal appendages, hyaline, smoothwalled. Asexual morph Undetermined.
Culture characteristics: Ascospore germinating on
MEA within 12 h. Colonies on MEA reaching 10 mm
diam. in 7 days at 28 °C, flat to slightly effuse, entire edge,
dark brown. Aerial mycelium radiating outwards, partially
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Fig. 25 Phylogenetic construction using RAxML-based analysis of a
combined LSU and SSU, dataset. Bootstrap support values for
maximum likelihood (ML, black) equal to or greater than 50% and
Bayesian posterior probabilities (BYPP, red) equal to or greater than
123
0.95 are shown above the nodes. The tree is rooted to Lophiostoma
macrostomum. The type strains are in black bold and the newly
generated sequences are indicated in blue bold
Fungal Diversity
Fig. 26 Lophiotrema guttulata (MFLU 10-0971, holotype). a Herbarium specimens. b–f Cross sections through ascomata. g Periphyses
of ostiole neck. h Peridium. i Pseudoparaphyses. j, k Asci.
l Ascospores. Scale bars a = 200 lm, b–f = 100 lm, h, j–k = 50
lm, g = 20 lm, i = 5 lm, l = 10 lm
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Fungal Diversity
Fig. 27 Lophiotrema guttulata (MFLUCC 10-0929, ex-type living
culture on WA). a, b Germinating ascospores. c Single ascospore
colonies on MEA. d, e Colonies on MEA from surface and reverse at
4 weeks. f, g Squash mount of immature and mature hyphae.
123
h Growth of asexual morph on plant tissues. i–k Squash mount of
vegetative hyphae. Scale bars a, b, f, g, i–k = 10 lm, c, e, h = 10
mm
Fungal Diversity
superficial, light brown to dark brown, with thick cells,
nonsporulating on culture.
Material examined: THAILAND, Chiang Mai Province,
Jomthong, Ban Luang, N18°310 E98°290 , on dried branch, 6
October 2010, R. Phookamsak, ITN-02: MFLU 10-0971,
holotype; ex-type living culture: MFLUCC 10-0929, BCC
52151.
GenBank Numbers LSU:KY498006; SSU:KY498007.
Notes: Lophiotrema guttulata differs from other species
of Lophiotrema in having a protruding ascomatal neck,
carbonaceous papilla and lacking a mucilaginous sheath in
the ascospores. Phylogenetic analyses of the combined
LSU and SSU dataset reveal that L. guttulata is nested
independently between L. lignicola and L. neohysterioides
and therefore provides evidence to support our new species
(Fig. 25).
Lophiotrema vagabundum (Sacc.) Sacc., Michelia 1 (4):
338 (1878)
Facesoffungi number: FoF 2783, Fig. 28
Saprobic on dead stem of Sambucus ebulus L. Sexual
morph Ascomata 489–614 lm high 9 622–801 lm diam.
(
x = 551 9 762 lm, n = 5), semi-immersed to erumpent
through host tissue, under to clypeus, solitary, uniloculate,
globose, conspicuous at the surface, black, subcarbonaceous, without hairs, ostiolate, smooth-walled. Peridium
25–41 lm wide, comprising 5–6 layers of hyaline to reddish-brown cells of textura angularis. Hamathecium 1.1–
1.9 lm wide, comprising numerous, aseptate, unbranched,
anastomosing pseudoparaphyses. Asci 96–129 9 8–10 lm
(
x = 118 9 9 lm, n = 10), 8-spored, bitunicate, fissitunicate, cylindrical, furcate, rounded at the ends, short
pedicellate, lacking an ocular chamber. Ascospores 22–30
9 4–7 lm (
x = 25 9 5 lm, n = 15), overlapping 1–2seriate, hyaline, 3-septate, constricted at the second septum, swollen at second cell, conical at the ends, guttulate,
surrounding by large mucilaginous sheath, smooth-walled.
Asexual morph Undetermined.
Culture characteristics: Colonies on MEA reaching 4–5
cm diam., after 7 days in the dark at 16 °C, flat to slightly
effuse, entire edge, light brown to dark brown, not sporulating in culture.
Material examined: ITALY, near San Benedetto in
Alpe, on dead stem of Sambucus ebulus (Adoxaceae), 24
October 2014, Erio Camporesi, IT2196 (MFLU 15-1497,
new host record); living culture MFLUCC 15-0081,
(HKAS 94608 bis).
GenBank Numbers LSU:KY496737; ITS:KY496757;
SSU:KY501125; TEF1:KY514401.
Notes: Lophiotrema vagabundum was introduced by
Saccardo (1878), based on a sexual morph and was
recorded from Vitis coignetiae, Epilobium angustifolium
and woody plants. Based on rDNA phylogenetic analyses,
the collection of L. vagabundum in this study (MFLUCC
15-0081) clustered with high statistical support to other
L. vagabundum strains analyzed (Fig. 25). Morphological
characters are also identical to other L. vagabundum strains
and therefore we report this collection as a new host record
on Sambucus ebulus (Adoxaceae), from Italy.
Hermatomyces Speg.
The genus Hermatomyces was introduced by Spegazzini
(1911), with H. tucumanensis as the type species. Currently
the genus contains six species (Index Fungorum 2017).
Morphologically, the genus is characterized by having
lenticular to cylindrical, muriform conidia, often with
subhyaline to pale brown peripheral cells, and dark brown
central cells. Conidia are cylindrical and comprise 1–4
columns with 2–11 cells and are irregularly pigmented
(Castañeda and Heredia 2000; Leão-Ferreira et al. 2013;
Doilom et al. 2017). Based on the phylogenetic study by
Doilom et al. (2017), Hermatomyces is accommodated in
Lophiotremataceae.
Hermatomyces chiangmaiensis J.F. Li, Bhat & K.D.
Hyde, sp. nov.
Index fungorum number: IF552306; Facesoffungi number: FoF02480, Fig. 29
Etymology: refers to the location where the fungus was
collected, Chiang Mai.
Holotype: MFLU 16-1930
Saprobic on dead hanging leaves of Pandanus sp. Sexual morph Undetermined. Asexual morph Colonies discrete on host, black, powdery. Mycelium immersed to
superficial in the substrate, composed of septate, branched,
smooth, light brown-coloured hyphae. Conidiophores 3.8–
5.3 lm long 9 4.5–5 lm diam. (
x = 4.8 9 4.7 lm,
n = 10), micronematous, mononematous, flexuous, hyaline, smooth, moderately thin-walled, comprising 1–2
subcylindrical to subglobose cells, arising from prostrate,
thick-walled hyphae. Conidiogenous cells 3.8–4.5 lm long
9 4.8–6.7 lm diam. (
x = 4.4 9 5.8 lm, n = 10), monotretic, integrated, terminal, ampulliform to doliiform, subhyaline to hyaline, smooth, thin-walled, dark-coloured at
conidiogenous loci. Conidia dimorphic, thick-walled,
smooth: lenticular conidia (10–)13.5–18 lm long 9 (7.3–)
8.6–10.4 lm diam. (
x = 13.5 9 9.6 lm, n = 20), multiseptate, with central cells dark brown to black, with
peripheral cells subhyaline to pale brown, slightly constricted at the septa, smooth, in side view composed of one
column of 4–6 cells, hyaline to light brown at the lower and
upper cells, often carrying remnant of conidiogenous cell at
base; cylindrical conidia: 10–16.4 lm high, 9 16–19.5 lm
wide in broadest part of lower cells, (
x = 13.2 9 17.5 lm,
n = 20), with 3–4 columns of 2–4 cells arising from a
common basal cell, each column, with rectangular to
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Fig. 28 Lophiotrema
vagabundum (MFLU 15-1497).
a Appearance of ascomata on
host substrate. b Section of
ascoma. c Section of peridium.
d Ostiole. e Pseudoparaphyses.
f, g Ascus. h–k Ascospores.
Scale bars b = 200 lm, c,
d = 10 lm, e = 2 lm, f,
g = 20 lm, h–k = 5 lm
globose cells, constricted at septa, pale to subhyaline,
granulate, smooth, terminating with a brown upper cell.
Cultural characteristics: Conidia germinating on PDA
within 10 h and germ tubes produced from the apex.
Colonies growing on PDA, reaching 5 cm in 14 days at
30 °C, mycelium partly superficial, partly immersed,
123
slightly effuse, cottony, with regular edge, olive-green to
grayish-green; sexual or asexual spores not formed within
60 days.
Material examined: THAILAND, Chiang Mai Province,
Mae Taeng, Musroom Research Centre, on a dead hanging
leaf of Pandanus sp., 25 March 2016, Junfu Li,
Fungal Diversity
Fig. 29 Hermatomyces chiangmaiensis (MFLU 16-2930, holotype).
a Colonies on dead leaf of Pandanus. b Conidia in mass. c Hyphae.
d Conidiophores with conidiogenous cells. e Hypha with
conidiogenous cell and conidia. f–m Conidia. Scale bars a = 100
lm, b = 10 lm, c–m = 5 lm
H-MRC40B (MFLU 16-2930, holotype). ex-type living
culture MFLUCC 16-2817, KUMCC (paratype in HKAS).
GenBank Number LSU:KY559394.
Notes: Hermatomyces chiangmaiensis is morphologically similar to H. krabiensis (Tibpromma et al. 2016b) in
conidial shape, but the former has smaller and ampulliform
to cylindrical conidia.
Index Fungorum number: IF552309; Facesoffungi
number: FoF02481, Fig. 30
Etymology: refers to the Sai Khu waterfall where the
fungus was collected.
Holotype: MFLU 16-1931
Saprobic on dead stem of Chromolaena odorata Linn.
Sexual morph Undetermined. Asexual morph Colonies
on natural substrate dry, blackish-brown, velvety, circular,
doughnut-shaped, dull, consisting of a sterile mycelial
outer zone and a round, glistening, abundantly sporulating
Hermatomyces chromolaenae J.F. Li, Mapook & K.D.
Hyde, sp. nov.
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Fungal Diversity
Fig. 30 Hermatomyces chromolaenae (MFLU 16-2931, holotype). a Colonies on dead stem. b Conidia in mass. c, d Hypha. e–h Conidiophores
with conidia. i–m Conidia. Scale bars a = 100 lm, b = 10 lm, c–e, h, n = 5 lm, i–m = 2 lm
centre, with conidia readily liberated when agitated.
Mycelium 1.3–2.1 lm wide, superficial, composed of a
network of branched, septate, brown, thick-walled, smooth
hyphae. Conidiophores 3–4.3 lm long 9 1.8–1.9 lm wide
123
(
x = 3.6 9 1.8 lm, n = 10), micronematous, flexuous,
pale brown, smooth, unbranched, arising from prostrate
hyphae at the centre of circular colony. Conidiogenous
cells holoblastic, monoblastic, integrated, terminal,
Fungal Diversity
cylindrical, hyaline to subhyaline. Conidia 9.2–10.4 lm
high 9 10.2–11.5 lm wide (
x = 9.7 9 10.7 lm, n = 20)
monomorphic, thick-walled, smooth, disk-shaped, with
central cells dark brown to black, with peripheral cells pale
brown. Two halves of the disk-shaped conidia symmetrically adpressed, forming a deep constriction at lower and
upper end in lateral view, each half with 6–7 cells, hyaline
to light brown at lower and upper cells, dark brown in
middle cells.
Cultural characteristics: Conidia germinating on PDA
within 10 h and germ tubes produced from the apex.
Colonies growing on PDA, reaching 5 cm in 14 days at
30 °C, mycelium partly superficial, partly immersed,
slightly effuse, cottony, with regular edge, olivaceousgreen to grayish-green; sexual or asexual spores not formed
within 60 days.
Material examined: THAILAND, Chiang Rai Province,
Doi Pui, on a dead (not land) stem of Chromolaena odorata
(Asteraceae), 5 October 2016, Junfu Li, DPK4 (MFLU
16-2931, holotype). ex-type living culture MFLUCC
16-2818, KUMCC (paratype in HKAS).
GenBank Number LSU:KY559393.
Notes: Morphologically, Hermatomyces chromolaenae
is similar to H. saikhuensis in having dark coloured conidia
and conidiophores arising from prostrate hyphae at the
centre of a circular colony. However, H. chromolaenae is
unique in having darker peripheral cells and smaller conidia and the phylogeny also segregates these two species. A
close phylogenetic association, albeit with moderate support, between H. chromolaenae and H. pandanicola is also
noted. Hermatomyces pandanicola, however, has dark
brown to black central cells in the conidia, with subhyaline
to pale brown peripheral cells, and in lateral view obovoid,
guttulate, hyaline to light brown at lower and upper ends
and dark brown at the central cells.
Melanommataceae G. Winter.
The family Melanommataceae, typified by Melanomma,
was established by Winter (1885) to accommodate Bertia,
Bombardia, Crotonocarpia, Rosellinia, Melanomma and
Melanopsamma. Melanommataceae is characterized by
globose or depressed perithecial ascomata, bitunicate and
fissitunicate asci, pigmented and phragmosporous ascospore (Tian et al. 2015). Tian et al. (2015) revised this
family by combining morphology and multi-locus phylogenetic analyses using LSU, SSU, TEF1 and RPB2 datasets
and 20 genera were accepted in the family.
Melanomma Nitschke ex Fuckel
Melanomma was validly published by Fuckel (1870) and
is typified by Melanomma pulvis-pyrius (Pers.) Fuckel.
Melanomma is characterized by the carbonaceous ascomata, and hyaline or brown, 2–3-septate ascospores (Tian
et al. 2015). Asexual morphs of Melanomma were reported
as coelomycetous and hyphomycetous (Hyde et al. 2011).
In this study, we transfer Aposphaeria populina Died. into
Melanomma.
Melanomma populina (Died.) Phukhamsakda & K.D.
Hyde, comb. nov.
Index Fungorum number: IF552757; Facesoffungi
number: FoF 2887
Basionym: Aposphaeria populina Died., Krypt.-Fl.
Brandenburg (Leipzig) 9: 206 (1912)
Notes: Aposphaeria is considered as an asexual genus in
the family Melanommataceae. Tian et al. (2015) mentioned
Aposphaeria and phoma-like asexual morphs have been
reported for Melanomma species (Sivanesan 1984). Additionally, this study reports the sexual state for Aposphaeria;
A. corallinolutea based on multi-gene phylogeny and the
identity of the asexual morph produced from culture.
Diedicke (1912) introduced A. populina as a phoma-like
species reported from Populus canadensis. De Gruyter
et al. (2013) designed an epitype from the same host, and
generated sequences for A. populina. Based on phylogenies, A. populina is related to Melanomma pulvis-pyrius as
they clustered together with high statistical support (Tian
et al. 2015; Hyde et al. 2016). Given that the sexual morph
morphology in A. corallinolutea and M. pulvis-pyrius are
different (Mugambi and Huhndorf 2009), and phylogeny
refer them to distinct clades, we synonymize Aposphaeria
populina under Melanomma populina.
Aposphaeria Sacc.
The genus Aposphaeria was introduced by Saccardo
(1880) and has a phoma-like structure. Aposphaeria is
characterized by pycnidial, superficial conidioma, hyaline,
short conidiophores, enteroblastic, phialidic, discrete,
conidiogenous cells, and hyaline, aseptate, and ellipsoidal
conidia (Sutton 1980). Tian et al. (2015) reconsidered the
family Melanommataceae and as a result of lack of
sequence data for A. pulviscula (Sacc.) Sacc (the type
species), Aposphaeria was retained in Melanommataceae.
Based on the phylogenetic analysis, A. populina is closely
related to Melanomma pulvis-pyrius, the type genus of
Melanommataceae (82%ML/82%MP/0.90BYPP, Fig. 31)
and is a synonym of Melanomma. It is however, important
that the type of Aposphaeria is recollected to establish if
the genus sensu stricto, belongs in Melanommataceae. In
this study, we describe the sexual morph of Aposphaeria
corallinolutea, bearing in mind that Aposphaeria sensu
stricto may not belong in Melanommataceae.
Aposphaeria corallinolutea Gruyter, Aveskamp & Verkley, Stud. Mycol 75: 28 (2012)
Facesoffungi number: FoF 2787, Fig. 32
123
Fungal Diversity
Fig. 31 Phylogenetic tree
generated from maximum
parsimony analysis based on
combined ITS and LSU
sequence data of the family
Melanommataceae. Bootstrap
support values for maximum
likelihood and maximum
parsimony greater than 50% and
Bayesian posterior probabilities
greater than 0.80 are indicated
above or below the nodes as
ML/MP/BYPP. Ex-type strains
and reference strains are in bold.
The new isolates are in red. The
tree is rooted with Pleomassaria
siparia (CBS 279.74)
Saprobic on the dead twigs of Prunus padus L. Sexual
morph Ascomata 194–418 lm high 9 183–397 lm wide
diam. (
x = 306 9 290 lm, n = 5), scattered or clustered in
small to large groups, initially immersed, becoming
erumpent to superficial, globose, black to dark brown,
subcarbonaceous, ostiole central. Ostioles 149 lm high 9
119 lm diam., papillate, dark brown, smooth, periphyses
filled ostioles cannel. Peridium 35–50 lm wide, up to 72
lm, thick, composed of 9–13 layers of textura angularis,
outer region haeavily pigment cells, black to dark-brown,
inner layer composed of hyaline gelatinous cells, thin,
merging with pseudoparaphyses. Hamathecium comprising
numerous, long, 1.3–2.2 lm (
x = 1.8 lm, n = 40) wide,
broad, transversely septate, branched, trabeculate pseudoparaphyses. Asci 66–154 9 6–14 lm (
x = 129 9 12 lm,
n = 20), (4)–8-spored, bitunicate, cylindrical, with clubshaped pedicel, apically rounded, with an ocular chamber
up to 2–4 lm wide. Ascospores 16–23 9 7–11 lm (
x = 19
9 9 lm, n = 50), overlapping uniseriate, hyaline, broadfusiform, conical at the apex, 1-transversely septate,
strongly constricted at the septa, wall rough, not uniform,
123
indentations present when mature. Asexual morph
Coelomycetous formed in cultures. Conidiomata 300 lm
high, 200 lm diam., pycnidia, dark brown, solitary,
aggregated, or clustered in groups, globose, papillate. Pycnidial wall comprising of cells of textura angularis,
ostiole absent. Conidiophores reduced to conidiogenous
cells. Conidiogenous cells 5–10–(17) 9 1.8–3 lm (
x=9
9 2.5 lm, n = 20), holoblastic, phialidic, hyaline, smoothwalled, discrete. Conidia 3–7 9 1.6–3 lm, 1-celled, hyaline, ellipsoid to oblong, slightly curved, smooth-walled.
Material examined: RUSSIA, Arkhangelsk Region,
Arkhangelsk City, Northern Maimaksa urban micro-district, wood waste landfill, on dead branches of Prunus
padus (Rosaceae), 11 May 2015, G.V. Okatov, T-583
(MFLU 15-2752, sexual morph); living culture MFLUCC
17-0001.
GenBank Numbers LSU:KY554197; SSU:KY554200;
ITS:KY554202; TEF1:KY554205; RPB2:KY554207.
Notes: Aposphaeria corallinolutea was introduced by
De Gruyter et al. (2013) based on Pleurophoma sp. cultures
in CBS culture collection. Aposphaeria corallinolutea
Fungal Diversity
Fig. 32 Aposphaeria corallinolutea (MFLU 15-2752, sexual
morph). a Host substrate. b, c Appearance of ascomata on host.
d Vertical section of ascoma. e Ostioles. f Section through peridium.
g Pseudoparaphyses. h–i Developmental stages of asci. j–n Developmental stages of ascospores. o Germinated ascospore. p Culture
characters. q Conidiomata produced in culture. r–v Conidiogenesis
developing. w Conidia. Scale bars b = 1000 lm, c, q = 500 lm,
d = 200 lm, e, g–i = 50 lm, f = 100 lm, j–m, r–w = 5 lm, n,
o = 10 lm
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Fungal Diversity
(MFLUCC 16-1059, sexual morph) has characters similar
to Melanommataceae such as nearly superficial and carbonaceous ascomata, papillate ostioles, thick-walled
peridium, trabeculate pseudoparaphyses, cylindrical uniseriate asci, and 1-septate ascospores. Our collection also
produced an asexual morph in culture, which is very similar to the characteristics of A. corallinolutea (CBS 131286,
CBS 131286). On the other hand, A. corallinolutea strains
are sister taxa to Melanomma but lack support. In the
phylogenetic tree, our strain Aposphaeria corallinolutea
(MFLUCC 16-1059) clustered together with A. corallinolutea (CBS 131286, CBS 131286) (Fig. 31) and we
describe the sexual morph of A. corallinolutea with the
support of asexual morph production from culture.
Nigrogranaceae Jaklitsch & Voglmayr
See notes under Biatriosporaceae.
Nigrograna De Gruyter et al.
Nigrograna was described by De Gruyter et al. (2013) as
a monotypic genus with Nigrograna mackinnonii as type
species. The sexual morph of Nigrograna is characterized
by depressed globose to globose, immersed or superficial,
black ascomata, ostioles periphysate, Ascospores asymmetric, fusoid to narrowly ellipsoid with the second cell
slightly wider than others (Jaklitsch and Voglmayr 2016).
The asexual morph of Nigrograna is characterized by
pycnidia similar to ascomata and usually co-occurring with
them, filiform conidiophores, simple to sparsely branched.
Conidia forming on pegs and phialides, oblong, 1-celled,
smooth (Jaklitsch and Voglmayr 2016). Five species are
accepted in Nigrograna (Jaklitsch and Voglmayr 2016)
(Fig. 33).
Nigrograna cangshanensis Z.L. Luo, H.Y. Su & K.D.
Hyde, sp. nov.
Index Fungorum number: IF 552681; Facesoffungi
number: FoF 2888, Fig. 34
Etymology: With reference to the collection site of this
fungus.
Holotype: HKAS 83978
Saprobic on decaying wood. Sexual morph Ascomata
120–135 9 135–155 lm, perithecioid, gregarious,
immersed, subglobose or ellipsoidal, black, with an ostiole,
neck long. Ostiole mostly central, brittle. Peridium 3-layered, composed of angular cells, outer layer, dark brown,
thick-walled cells, middle layer and inner layer, pale
brown, with thick-walled cells. Hamathecium composed of
1.5–2.5 lm (
x = 2 lm, n = 25) wide, numerous, tapering
distally, pseudoparaphyses, longer than asci, embedded in a
gelatinous matrix. Asci 50–70 lm long (
x = 60 lm,
n = 25), 6.5–8.5 lm wide (
x = 7.5 lm, n = 25), 8-spored,
bitunicate, fissitunicate, clavate to cylindric-clavate, short
123
pedicellate, apically rounded, sometimes with an ocular
chamber. Ascospores 11–14 lm long (
x = 12.5 lm,
n = 35), 3.5–4.5 lm wide (
x = 4 lm, n = 35), biseriate,
olive-gray to yellowish-brown, 3-septate, constricted at the
septa. Ascospores germinate from germ tubes from all four
cells. Asexual morph Undetermined.
Material examined: CHINA, Yunnan Province, saprobic
on decaying wood submerged in stream in Cangshan
Mountain, March 2014, Z.L. Luo, LQXM 25
(HKAS83978, holotype), ex-type living culture, MFLUCC
15-0253.
GenBank Numbers LSU:KY511064; SSU:KY511065;
ITS:KY511063; TEF1:KY511066.
Notes: All species of Nigrograna are morphologically
very similar and thus, Jaklitsch and Voglmayr (2016)
considered them to be cryptic species. Morphologically,
N. cangshanensis is very similar to other species in the
genus. However, N. cangshanensis differs from N. fuscidula, N. norvegica and N. obliqua in having a thicker
peridium. Phylogenetic analysis also showed that N. cangshanensis is distinct from other Nigrograna species
(Fig. 33).
Roussoellaceae J.K. Liu, Phook., D.Q. Dai & K.D. Hyde
The family Thyridariaceae was introduced by Hyde
et al. (2013) to accommodate the genus Thyridaria based
on morphology and phylogenetic analyses with Thyridaria
incrustans as the type species. Jaklitsch and Voglmayr
(2016) introduced a new genus, Parathyridaria, with two
species in the family, and provided a phylogenetic analysis
of the family based on the combined ITS, LSU, SSU, RPB2
and EF-1a sequence data. In this study, a new species,
Parathyridaria robiniae, is introduced with evidence from
molecular data analysis and morphological comparisons.
Jaklitsch and Voglmayr (2016) treated Roussoellaceae as a
synonym of Thyridariaceae. We obtained a similar topology in our phylogenetic analysis, however, the two families
have distinct morphological characters. The families are
also well resolved in the analyses. This is in agreement
with the results of Liu et al. (2014) where they provide
evidence that Roussoellaceae is a good family in
Pleosporales with high diversity (Crous et al. 2014;
Ariyawansa et al. 2015b; Dai et al. 2017). We follow Liu
et al. (2014) and treat Roussoellaceae as a well-resolved
family in Pleosporales. The phylogenetic tree, based on
combined LSU, ITS, EF-1a and RPB2 sequence data is
presented in Fig. 35.
Thyridariaceae Q. Tian & K.D. Hyde
See notes for Roussoellaceae above.
Parathyridaria robiniae Mapook, Camporesi & K.D.
Hyde, sp. nov.
Fungal Diversity
Fig. 33 RAxML tree based on a combined dataset of LSU, SSU, ITS,
RPB2 and TEF1 partial sequences. Bootstrap support values for
maximum likelihood (ML) higher than 70% and Bayesian posterior
probabilities (BYPP) greater than 0.95 are given above the branches.
The ex-type and reference strains are in bold; the new isolates are in
blue. The tree is rooted to Melanomma pulvis-pyrius
(Melanommataceae)
123
Fungal Diversity
Fig. 34 Nigrograna cangshanensis (HKAS 83978, holotype).
a Wood. b, c Appearance of ascomata on wood. d Section of ascoma.
e Section of the peridium. f Paraphyses. g–k Asci. l–p Ascospores. q–
123
s Germinating ascospore. t, u Culture grow on MEA. Scale bars
d = 100 lm, e = 50 lm, f = 20 lm, g–k = 10 lm, l–p = 5 lm, q–
s = 10 lm
Fungal Diversity
Fig. 35 Phylogram generated from RAxML based on combined
LSU, ITS, TEF1 and RPB2 sequenced data. Maximum likelihood
(ML) bootstrap support values greater than 60% and Bayesian
posterior probabilities (BYPP) greater than 0.95 are shown at the
nodes. The ex-type strains are in bold and the new isolate is in blue.
The tree is rooted with Occultibambusa bambusae
Index Fungorum number: IF552776; Facesoffungi
number: FoF 2794, Fig. 36
Etymology: Name reflects the host genus Robinia.
Holotype: MFLU 17-0015
Saprobic on a dead branch of Robinia pseudoacacia L.
Sexual morph Ascomata 450–470 lm high 9 255–270 lm
diam. (
x = 455 9 260 lm, n = 5), semi-immersed to
immersed, scattered, coriaceous, in groups, ovoid to
obpyriform, reddish-brown to brown, ostiole central.
Peridium 10–25 lm wide, comprising light brown to
hyaline cells of textura angularis. Hamathecium comprising 0.5–1.5 lm wide, cylindrical to filiform, septate,
123
Fungal Diversity
Fig. 36 Parathyridaria robiniae (MFLU 17-0015, holotype). a, b
Appearance of superficial ascomata on substrate. c Section through of
ascomata. d Ostiole e Peridium. f Pseudoparaphyses. g, h Asci at
immature and mature. i–k Ascospores. l Ascospore surrounded by
hyaline gelatinous sheath in Indian ink. Scale bars a, b = 200 lm,
c = 100 lm, d, g–h = 50 lm, e = 20 lm, i–l = 10 lm, f = 5 lm
branched, pseudoparaphyses. Asci 105–115 9 10–15 lm
(
x = 110 9 11.5 lm, n = 10), 8-spored, bitunicate,
cylindric-clavate, pedicellate, apically rounded with a
small ocular chamber. Ascospores 15–20 9 4–7 lm
(
x = 18.5 9 5 lm, n = 15), overlapping 1–2-seriate,
hyaline, broadly fusiform to inequilateral, 1-septate,
123
Fungal Diversity
constricted at the septa, widest at the centre and tapering
towards the narrow ends, smooth-walled, with mucilaginous sheath. Asexual morph Undetermined.
Material examined: ITALY, Forlı̀-Cesena Province,
Collina di Pondo—Santa Sofia, on dead branch of Robinia
pseudacacia (Fabaceae), 22 September 2014, E. Camporesi (MFLU 17-0015, holotype), ex-type living culture
MFLUCC 14-1119 (isotype in HKAS, under the code of
HKAS).
GenBank Numbers LSU:KY511141; ITS:KY511142;
TEF1:KY549682.
Notes: Phylogenetic analysis indicates that our collection belongs to the family Thyridariaceae and P. robiniae
groups together with members of Parathyridaria a genus
that was introduced by Jaklitsch and Voglmayr (2016). The
phylogeny also indicates that P. robiniae lies in an independent terminal taxon nested between P. percutanea and
P. ramulicola and therefore can be considered as a new
species (Fig. 35). Parathyridaria robiniae is also distinct in
having 1-septate ascospores surrounded by a hyaline
gelatinous sheath.
Massarineae
Massarineae is a suborder of Pleosporales, originally
established by Barr (1979) to accommodate Massarinaceae
and Arthopyreniaceae (Hawksworth et al. 1983, 1995; Barr
1987; Eriksson and Winka 1998; Kirk et al. 2008; Lumbsch
and Huhndorf 2010). The taxonomic framework, phylogenetic relationships, biology, and species diversity of
Massarineae are poorly understood (Tanaka et al. 2015;
Phukhamsakda et al. 2016).
Didymosphaeriaceae Munk.
The family Didymosphaeriaceae was introduced by
Munk (1953) and typified by Didymosphaeria, with
D. epidermidis (Fr.) Fuckel as the type species. Ariyawansa
et al. (2014a) synonymized Montagnulaceae under Didymosphaeriaceae based on priority of the oldest name.
Currently, 25 genera are accommodated in Didymosphaeriaceae (Ariyawansa et al. 2015b; Crous et al. 2015a;
Thambugala et al. 2014a; Wijayawardene et al. 2014a;
Tanaka et al. 2015; Wanasinghe et al. 2016). We introduce
introduce a new species of Paraphaeosphaeria with an
updated phylogenetic tree for the family.
Paraphaeosphaeria O.E. Erikss.
Paraphaeosphaeria was introduced by Erikss (1967)
and species of this genus are similar to Phaeosphaeria
Miyake 1909. Sutton (1980) mentioned that all known
species of Paraphaeosphaeria occur on monocotyledons
and taxa differ in ascomatal structure, ascospores septation,
size, host and in asexual characteristics. Taxonomic studies
have also reported that this genus is polyphyletic (Câmara
et al. 2002; Checa et al. 2002)
Paraphaeosphaeria viciae de Silva, Camporesi & K.D.
Hyde, sp. nov.
Index Fungorum number: IF552714; Facesoffungi
number: FoF 2776, Fig. 38
Etymology: Name reflects the host genus Vicia.
Holotype: MFLU 15-1231
Saprobic on Vicia sp. Sexual morph Undetermined.
Asexual morph Conidiomata 113–163 lm diam. semiimmersed, solitary to gregarious, scattered, globose to
subglobose, uniloculate, pale brown, ostiolate. Ostioles
circular, without papilla. Conidiomata wall 8–13 lm wide,
composed of thin-walled, hyaline to light brown cells of
textura angularis. Conidiophores indistinct. Conidiogenous cells 3–5 9 2–4 lm (
x = 4.1 9 3 lm), enteroblastic, phialidic, cylindrical to subcylindrical, ampulliform,
discrete, hyaline, smooth. Conidia 4–5 9 1–2 lm diam.
(
x = 4.8 9 2.1 lm), subglobose to ellipsoid or obovoid,
hyaline to pale, smooth and thick-walled, guttulate. Sexual
morph Undetermined.
Culture characteristics: Colonies on PDA 35 mm diam.
after 21 days at 25 °C, irregular, flat, smooth surface with
undulate edge, white colour colony and reverse pale brown.
Material examined: ITALY, Forlı̀-Cesena Province,
Monte Mirabello-Predappio, on dead branch of Vicia sp.
(Fabaceae), 18 May 2015, Erio Camporesi, IT2482
(MFLU 15-1231 holotype), ex-type living culture
KUMCC 16-0184.
GenBank Numbers LSU:KY397948; SSU:KY397947;
ITS:KY379969.
Notes: Paraphaeosphaeria viciae shares some similarities with P. spartii in having subglobose to ellipsoid or
obovoid, aseptate conidia with guttules. Paraphaeosphaeria viciae however, has 113–163 lm diam.,
semi-immersed, globose to subglobose conidiomata as
compared to 200–300 lm diam. in P. spartii (Liu et al.
2015; Wijayawardene et al. 2016). DNA sequence analyses
confirm the position of our new taxon in Didymosphaeriaceae (Fig. 37) and indicate the genus is polyphyletic.
Latoruaceae Crous
The family Latoruaceae was introduced by Crous et al.
(2015b) and typified by Latorua caligans (Bat. & H.P.
Upadhyay) Crous. Four genera Latorua, Matsushimamyces, Polyschema, Pseudoasteromassaria are accommodated in this family. Both sexual morph and asexual
morph of the genus Pseudoasteromassaria have been
reported and are characterized by fusiform, 1–3-septate,
brown ascospores and cylindrical, 6–7-septate, hyaline
conidia (Ariyawansa et al. 2015c) (Fig. 39).
123
Fungal Diversity
Fig. 37 RAxML tree based on a combined dataset of LSU, ITS, SSU
and TEF1 partial sequences from Didymosphaeriaceae. Bootstrap
support values for maximum likelihood higher than 60% and
Bayesian posterior probabilities higher than 0.90 are defined as
above the nodes respectively. The tree is rooted to Pleospora
herbarum and Pleospora tarda. The new species is in blue
Pseudoasteromassaria M. Matsum. & Kaz. Tanaka
Pseudoasteromassaria was introduced by M. Matsum.
& Kaz. Tanaka (2015) with Ps. fagi M. Matsum. & Kaz.
Tanaka as the type species. The morphological characteristics of this genus are similar to those of Asteromassaria
(Pleomassariaceae) (Ariyawansa et al. 2015c).
123
Fungal Diversity
Fig. 38 Paraphaeosphaeria viciae (MFLU 15-1231, holotype). a, b
Conidiomata on the host. c Section through conidiomata. d Conidia
with conidiogenous cells. e–h Conidia. i Germinating conidia. j Upper
view of culture. k Lower view of culture. Scale bars c = 80 lm,
d = 5 lm, e–h = 10 lm, i = 20 lm
Pseudoasteromassaria spadicea W. Dong, H. Zhang &
K.D. Hyde, sp. nov.
Index Fungorum number: IF 552730; Facesoffungi
number: FoF 2825, Fig. 40
Etymology: spadicea in reference to the colour of the
germinated conidia.
Holotype: MFLU 15-2683
Saprobic on submerged wood in freshwater habitats.
Asexual morph Conidiomata 290–320 lm high 9 460–
480 lm diam., pycnidial, immersed, solitary or scattered on
the host surface, ellipsoidal, unilocular, ostiolate in the
center, circular, dark-brown, with a short papilla. Peridium
23–28 lm wide, composing several layers of irregular cells
arranged in a textura angularis, brown to dark brown.
Conidiophores reduced. Conidiogenous cells phialidic, up
to 8 lm long, determinate, discrete, cylindrical or pyramidal, hyaline, smooth, forming from the inner wall cells.
Conidia 11–15 9 7–10 lm (
x = 13 9 9 lm, n = 20),
globose to subglobose or ellipsoidal or obovoid, without
septum, swollen with one large guttules in the cell, apex
and base of cell rugose or smooth, hyaline, becoming light
brown when germinated in culture, thick-walled. Sexual
morph Undetermined.
Culture characteristics: on PDA reaching 3 cm diam.
after 30 days at 25 °C, with circular colony, edge undulate,
surface rough with raised elevation, brown to dark brown.
Material examined: THAILAND, Prachuap Khiri Khan
Province, on submerged wood, 30 July 2015, Wei Dong,
34B (MFLU 15-2683, holotype); ex-type living culture
MFLUCC 15-0973).
GenBank Numbers LSU:KY522724; SSU:KY522725;
ITS:KY522726.
Notes: Phylogenetic analyses of combined genes indicate that Pseudoasteromassaria spadicea clustered with
other species of Pseudoasteromassaria and in particular it
shares a close affinity to P. fagi. Pseudoasteromassaria
spadicea differs from the asexual strain of P. fagi mainly
by its globose or obovoid, aseptate conidia and conidial
123
Fungal Diversity
Fig. 39 Phylogenetic tree generated from maximum likelihood
analysis based on combined LSU, SSU, TEF1 and RPB2 sequence
data of the suborder Massarineae. Bootstrap support values for
maximum likelihood greater than 70% are indicated above or below
123
the nodes as ML. Ex-type strains and reference strains are in black
bold. The new species are in blue. The tree is rooted with Pleospora
herbarum and Alternaria alternata
Fungal Diversity
Fig. 40 Pseudoasteromassaria spadicea (MFLU 15-2683, holotype). a, b Appearance of conidioma on host substrate. c, d Section of
conidioma. e Section of neck. f Section of peridium. g–i Conidiogenous cells and conidia. j Germinated conidium producing light
brown pigment. k–p Conidia. q Colony on PDA (from top). r Colony
on PDA (from reverse). Scale bars b = 200 lm, c–d = 100 lm,
e = 20 lm, f, j = 10 lm, g–i = 5 lm, k–p = 5 lm
123
Fungal Diversity
size (11–15 9 7–10 lm vs. (52–)61–71 9 10–12.5 lm)
(Ariyawansa et al. 2015c).
Lentitheciaceae Y. Zhang ter, C.L. Schoch, J. Fourn.,
Crous & K.D. Hyde
The family Lentitheciaceae was introduced by Zhang
et al. (2009a) with L. fluviatile (Aptroot & Van Ryck.) K.D.
Hyde (2009) as the type species. This family now includes
six genera (Wanasinghe et al. 2014).
Keissleriella Höhn.
Keissleriella was introduced by Höhnel (1919) to
accommodate Keissleriella aesculi Sacc. (:Pyrenochaeta
aesculi Höhn.) as the type. The genus is characterized by
ascomata with ostiolar necks filled with black setae, and 1
to multi-septate, hyaline ascospores (Barr 1990; Liu et al.
2015). Munk (1957) placed the genus in Lophiostomataceae, while von Arx and Muller (1975) placed it in
Pleosporaceae. Later Barr (1990) transferred it to Melanommataceae. Lumbsch and Huhndorf (2007) maintained Keissleriella in Massarinaceae, while Zhang et al.
(2009b) placed the genus in Lentitheciaceae, and this has
been followed by subsequent authors (Hyde et al. 2013;
Tanaka et al. 2015; Wijayawardene et al. 2014a; Zhang
et al. 2012b) (Fig. 41).
Keissleriella cirsii R.H. Perera, Bulgakov, Wanasinghe &
K.D. Hyde, sp. nov.
Index Fungorum number: IF552705; Facesoffungi
number: FoF 2767, Fig. 42
Etymology: Named after the host genus.
Holotype: MFLU 15-2900
Saprobic on Cirsium sp. Sexual morph Ascomata 245–
370 lm high, 180–368 lm diam. (
x = 285 9 255 lm,
n = 10), semi-immersed, appearing as black raised spots
on the host, solitary, globose to subglobose, wall black,
uniloculate, black, ostiolate. Ostioles with dark brown to
black setae, Peridium 30–67 lm wide, composed of 4–6
layers of pale brown to 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), 8spored, bitunicate, clavate, with a short furcate pedicel,
rounded at the apex. Ascospores 23.5–29 9 5–7 lm
(
x = 27 9 6 lm, n = 40), uniseriate to biseriate, fusiform,
3–5-septate, constricted at the septa, hyaline, inequilateral,
with a mucilaginous sheath. Asexual morph
Undetermined.
Culture characteristics: Colonies on PDA reaching
50–55 mm diam. in 21 days, white, spreading, with moderate aerial mycelium, slightly irregular, with smooth
margins.
123
Material examined: RUSSIA, Voronezh Oblast, Lysogorka, on stems of Cirsium vulgare (Savi) Ten., 25 July
2015, T. Bulgakov, T 755 (MFLU 15-2900, holotype); extype living cultures MFLUCC 16-0454, MFLUCC
16-0455.
GenBank Numbers LSU:KY497780; SSU:KY497782;
ITS:KY497783; TEF1:KY497786; RPB2:KY497787.
Notes: Keissleriella cirsii is typical of Keissleriella in its
ostiolar neck covered by short dark setae (Munk 1953;
Tanaka et al. 2015)., K. cirsii is a distinct species in the
genus, supported by molecular and morphological characteristics. Keissleriella cirsii clustered as sister to K. taminensis with 100% bootstrap support. Keissleriella cirsii
differs from K. taminensis in having larger ascomata and
ascospores with fewer longitudinal septa (3–4 vs. 4–5)
(Tanaka et al. 2015).
Pleurophoma Höhn.
The genus Pleurophoma, based on P. pleurospora (De
Gruyter et al. 2009), lacks any known sexual morph (De
Gruyter et al. 2010). We introduce a new species of
Pleurophoma italica, and describe both morphs.
Pleurophoma italica Tibpromma, Camporesi & K.D.
Hyde, sp. nov.
Index Fungorum number: IF552689; Facesoffungi
number: FoF 2768, Figs. 43, 44
Etymology: refers to the country where the holotype was
collected.
Holotype: MFLU 15-1254
Saprobic on cone of Pinus nigra J.F. Arnold lying on the
ground. Sexual morph Ascomata 154–167 lm high 9
141–155 lm diam. (
x = 157 9 148 lm, n = 5), semiimmersed or erumpent, solitary, or in small groups,
uniloculate, globose to subglobose, conspicuous at the
surface, black, without hairs, ostiolate. Peridium 6–12 lm
wide, comprising 4–5 layers of reddish-brown cells of
textura angularis. Hamathecium 2.2–3.4 lm wide, comprising dense, aseptate, unbranched pseudoparaphyses.
Asci 51–106 9 7–11 lm (
x = 72 9 9 lm, n = 15), 8spored, bitunicate, fissitunicate, cylindrical, with short,
club-shaped pedicel. Ascospores 13–16 9 3–5 lm (
x = 14
9 4 lm, n = 20), overlapping 1–2-seriate, hyaline, fusiform, 1-septate in the center, deeply constricted at the
septum, swollen at the first cell, conical at the ends, with
guttulate, lacking a mucilaginous sheath. Asexual morph
Coelomycetous, produced on sterilized bamboo pieces and
pine needles on water agar. Conidiomata 68–105 lm high
9 77–98 lm diam., pycnidial, superficial on substrate,
solitary, black, dull, uniloculate, globose to subglobose,
conspicuous at the surface. Conidiophores reduced to
conidiogenous cells arising from the innermost wall-layer
Fungal Diversity
Fig. 41 RAxML tree based on analysis of a combined LSU, SSU,
TEF1 and ITS sequence dataset. Bootstrap support values for
maximum likelihood, maximum parsimony higher than 60% and
Bayesian posterior probabilities greater than 0.95 are given above
each branch. The ex-type and reference strains are in bold; the new
species are in blue. The tree is rooted to Massarina eburnea and
Massarina cisti in Massarinaceae
cells of the conidiomata. Conidiogenous cells 5.3–8.6 9
3.2–5.1 lm (
x = 6.6 9 3.9 lm, n = 20), holoblastic,
phialidic, globose to oblong, hyaline, and formed from the
inner layer of the pycnidium wall. Conidia 0.9–3 9 0.7–1.8
lm (
x = 1.9 9 1.2 lm, n = 40), globose to oval, aseptate,
hyaline, smooth-walled, spermatia-like.
Culture characteristics: Colonies on MEA at 16 °C
reaching 2.5 cm in 1 week, circular with wavy margin,
white–grey, smooth surface and raised.
Material examined: ITALY, near Ortignano-Raggiolo,
on cone of Pinus nigra (Pinaceae) on ground, 29 July
2014, Erio Camporesi, IT2024 (MFLU 15-1254,
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Fungal Diversity
Fig. 42 Keissleriella cirsii (MFLU 15-2900, holotype). a Type
material. b Appearance of ascomata immersed in host substrate.
c Section through ascoma. d Section through ostiole with internal
setae. e Close up of the peridium. f. Pseudoparaphyses. g, h Asci.
i Ascus in Melzer’s reagent. j, k Ascospores. Scale bars b = 500 lm,
c = 100 lm, d = 50 e, f = 10 lm, g–k = 20 lm
holotype); ex-type living culture MFLUCC 15-0061
(HKAS 94600 bis, paratype).
GenBank Numbers LSU:KY496734; ITS:KY496754;
SSU:KY501122; TEF1:KY514398.
Notes: Pleurophoma italica is introduced as a new
species with both sexual and asexual morphs. The
phylogeny indicates a close relationship of P. italica to
P. pleurospora (type species), but constitutes a separate
branch (Fig. 41). Pleurophoma italica shares similar
asexual morphology with P. pleurospora. However, our
taxon also clustered with Keissleriella sparticola,
K. cladophila and K. genistae, but the morphological
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Fungal Diversity
Fig. 43 Pleurophoma italica
(MFLU 15-1254, holotype).
a Appearance of ascomata on
host substrate. b Section of
ascoma. c Section of peridium.
d Pseudoparaphyses. e, f Ascus.
g–j Ascospores. Scale bars
b = 50 lm, c = 10 lm, d = 2
lm, e, f = 5 lm, g–j = 2 lm
characteristics of sexual morphs are different. In the future,
if Keissleriella sparticola, K. cladophila and K. genistae
asexual morphs are discovered, and with more phylogenetic evidence, these three species could be synonymized
under Pleurophoma. Hence, based on our phylogenetic tree
(Fig. 41) we agree with Crous et al. (2015a) that the link
between Pleurophoma and Keissleriella needs to be
resolved.
Massarinaceae Munk
Details of this family can be seen in Hyde et al. (2013),
Tanaka et al. (2015) and Hyde et al. (2016).
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Fungal Diversity
Fig. 44 Pleurophoma italica
(MFLUCC 15-0061, ex-type
living culture on WA).
a Appearance of conidioma on
bamboo. b Conidiomata. c, d
Conidiogenous cells. e Conidia.
Scale bars a = 500 lm, b = 50
lm, c–e = 2 lm
Pseudodidymosphaeria Thambug. & K.D. Hyde
The genus Pseudodidymosphaeria Thambugala & K.D.
Hyde was introduced by Thambugala et al. (2015b) and
accommodated in the family Massarinaceae. Two species
are accepted in the genus, viz. P. spartii (Fabre) Thambugala et al. and P. phlei Phukhamsakda et al. (Li et al.
2016).
Pseudodidymosphaeria phlei Phukhams., Camporesi &
K.D. Hyde, Fungal Diversity 78: 57 (2016)
Facesoffungi number: FoF 2786, Fig. 45
Saprobic on species of Arundo donax L., in terrestrial
environments. Sexual morph Ascomata 300–500 lm high 9
250–450 lm diam. (
x = 184.6 9 146.2 lm, n = 5), solitary,
immersed, coriaceous, dark brown to black, globose to
subglobose. Peridium 7–12 lm wide at the base, 12–20 lm
wide at the sides, two-layered, and outer layer composed of
dark brown, thin-walled cells of textura angularis, inner
layer composed of smaller, compressed, hyaline cells. Hamathecium of 1.5–2.5 lm wide, septate, hyaline, cellular
pseudoparaphyses, embedded in gelatinous matrix between
and above the asci. Asci 100–130 9 12–20 lm (
x = 112.9 9
123
15.1 lm, n = 15), 8-spored, bitunicate, fissitunicate, cylindrical to cylindric-clavate, with a short pedicel, narrowly
rounded at the apex, with an indistinct ocular chamber. Ascospores 19–22 9 9–11 lm (
x = 20.2 9 10.2 lm, n = 40),
uniseriate to obliquely uniseriate, slightly overlapping,
hyaline when immature, becoming golden brown when
mature, ellipsoid, with broadly obtuse ends, 1-septate, upper
cell slightly broader than the lower one, constricted at the
septum, verrucose, surrounded by a thick mucilaginous
sheath. Asexual morph Undetermined.
Culture characteristics: on PDA growing slowly,
reaching 25–30 mm diam. after 21 days at 16 °C, surface
greenish-brown, spreading with moderate aerial mycelium,
and uneven, smooth margins; reverse buff.
Material examined: ITALY, Forlı̀-Cesena, Ravaldino in
Monte, on dead Arundo donax stem (Poaceae), 11
November 2014, Erio Camporesi, IT 2233 (MFLU
15-1360, new host record);
GenBank Numbers LSU:KY264748; ITS:KY264744;
SSU:KY264752.
Notes: Li et al. (2016) introduced Pseudodidymosphaeria phlei from Phleum pretense L. (Poaceae)
Fungal Diversity
Fig. 45 Pseudodidymosphaeria phlei (MFLU 15-1360). a Appearance of ascomata on the host substrate. b Section of ascoma.
c Section of peridium. d Pseudoparaphyses. e–g Asci. g–l Ascospores.
Scale bars a = 500 lm, b = 100 lm, c = 20 lm, d = 10 lm, e,
f = 20 lm, h–l = 10 lm
collected in Forlı̀-Cesena Province of Italy as the second
species of Pseudodidymosphaeria. Pseudodidymosphaeria
phlei is similar to type species, P. spartii, in having globose
to subglobose ascomata, cylindric-clavate asci and 1-septate ascospores. Our new isolate is also morphologically
similar to P. phlei (strain MFLUCC 14-1061) and
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Fungal Diversity
phylogenetically groups with 100% ML support. However,
it is associated with a different host. Pseudodidymosphaeria phlei (holotype) was recorded on Phleum pretense
and our collection was found on Arundo donax (Poaceae).
Massarineae genera incertae sedis
Inflatispora Y. Zhang ter, J. Fourn. & K.D. Hyde
Zhang et al. (2011) introduced Inflatispora as a lignicolous genus of Pleosporales from France to accommodate
I. pseudostromatica Y. Zhang ter, J. Fourn. & K.D. Hyde.
The genus is characterized by ‘widely porate ascomata,
bitunicate asci, 3-septate, hyaline, cylindrical ascospores,
with a swollen upper central cell and surrounded by a thin
sheath, and narrow cellular pseudoparaphyses that anastomose between and above the asci (Zhang et al. 2011). Inflatispora pseudostromatica resembles species of
Nodulosphaeria in having ascospores with an enlarged
supramedian cell, but it can be distinguished from Nodulosphaeria by the presence a sheath surrounding the whole
ascospore, while Nodulosphaeria has terminal ascospore
appendages. The genus Inflatispora seems more typical of
Massarineae, but the familial placement is still uncertain.
Thus, we maintain the genus in Massarineae genera incertae sedis. It is essential to collect more taxa to establish
their placement in Massarineae (Fig. 39).
Inflatispora caryotae Wanasinghe & K.D. Hyde, sp. nov.
Index Fungorum number: IF552767; Facesoffungi
number: FoF 2795, Fig. 46
Etymology: refers to the host genus (Caryota).
Holotype: MFLU 16-2886
Saprobic on Caryota sp. in terrestrial habitats. Sexual
morph Ascomata 350–400 lm high, 200–300 lm diam.
(
x = 403 9 260 lm, n = 5), scattered, immersed, black,
subglobose, papillate. Peridium 15–25 lm wide at the base,
10–15 lm wide at the sides, comprising 2–3 layers, fused
with the host tissue at the outside, composed of thickwalled, black-brown, angular or relatively compressed,
pseudoparenchymatous cells. Hamathecium of 1.5–3 lm
wide, filiform, septate, anastomosing and branched, pseudoparaphyses. Asci 110–140 9 15–30 lm (
x = 122.6 9
23.1 lm, n = 20), 8-spored, bitunicate, fissitunicate, clavate to cylindric-clavate, pedicellate, apically rounded,
with an ocular chamber. Ascospores 40–50 9 5–8 lm
(
x = 46.5 9 6.6 lm, n = 30), 2–3-seriate, hyaline, fusiform, narrow towards and with acute ends, 1-septate,
constricted at the septum, guttulate, smooth-walled, surrounded by a distinct mucilaginous sheath. Asexual morph
Undetermined.
Culture characteristics: on MEA reaching 2 cm diam.
after 2 weeks at 25 °C, dirty white at the beginning, with
pale brown centre, dirty white towards outer margin at
123
maturity, convex on the surface, undulate, margins smooth;
reverse coffee brown.
Material examined: THAILAND, Narathiwat Province,
on dead Caryota rachis, August 2013, Saithong Kaewchai,
DHA07 (MFLU 16-2886, holotype); ex-type living culture, MFLUCC 13-0825).
GenBank Numbers LSU:KY264747; ITS:KY264743;
SSU:KY264751.
Notes: We introduce the second species in Inflatispora,
I. caryotae from Thailand based on morphological characteristics coupled with molecular data (Fig. 39). This
species morphologically resembles I. pseudostromatica in
having fusiform, hyaline ascospores with acute ends and a
distinct mucilaginous sheath surrounding the ascospores.
However, I. pseudostromatica has semi-immersed ascomata and 3-septate ascospores, while I. caryotae has
immersed ascomata and 1-septate ascospores.
Suborder Pleosporineae
Didymellaceae Gruyter, Aveskamp & Verkley
The family Didymellaceae was introduced for the
‘‘Didymella clade’’ to accommodate the type species
Didymella exigua and includes phoma and phoma-like
genera (De Gruyter et al. 2009). Seventeen genera are
presently included in Didymellaceae based on morphology
and phylogeny, and these are Allophoma, Ascochyta, Boeremia, Calophoma, Dactuliochaeta, Didymella, Epicoccum, Heterophoma, Leptosphaerulina, Macroventuria,
Neodidymelliopsis, Neoascochyta, Nothophoma, Paraboeremia, Phoma, Phomatodes and Xenodidymella (Hyde
et al. 2013; Wijayawardene et al. 2014a; Chen et al. 2015).
We provide an updated tree for Didymellaceae.
Ascochyta Lib
Ascochyta was introduced by Lib. in (1830) with A. pisi
as the type species. A recent study by Aveskamp et al.
(2010), reported that Ascochyta is polyphyletic. We introduce two new species and one new host record of Ascochyta and provide a new phylogenetic tree for
Didymellaceae (Fig. 47).
Ascochyta italica Tibpromma, Camporesi & K.D. Hyde,
sp. nov.
Index Fungorum number: IF552693; Facesoffungi
number: FoF 2772, Fig. 48
Etymology: Named after the country where the fungus
occurs.
Holotype: MFLU 14-0724
Saprobic on dead stem of Onobrychis viciifolia Scop.
Sexual morph Ascomata 127–161 lm high 9 167–196 lm
diam. (
x = 142 9 187 lm, n = 5), immersed, solitary, or in
Fungal Diversity
Fig. 46 Inflatispora caryotae (MFLU 16-2886, holotype). a Appearance of ascomata on host substrate. b, c Section of ascoma. c Close up
of ostiole. d Section of peridium. e Pseudoparaphyses. f–h Asci. i–
k Ascospores. l Germinated ascospore. Scale bars a = 500 lm, b,
c = 100 lm, d = 20 lm, e = 10 lm, f–l = 20 lm
small groups, uniloculate, globose, conspicuous at the surface, dull black, carbonaceous, without hairs, with short
ostiolate. Peridium 16–37 lm wide, comprising 5–6 layers of
hyaline to yellow–brown cells of textura angularis.
Hamathecium 1.3–2.7 lm wide, comprising few dense,
septate, unbranched, pseudoparaphyses. Asci 42–66 9 9–16
lm (
x = 59 9 14 lm, n = 10), 8-spored, bitunicate, fissitunicate, cylindric-clavate, with short club-shaped pedicel,
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Fungal Diversity
Fig. 47 Phylogram generated from maximum likelihood analysis of a
LSU, ITS, b-tubulin and RPB2 combined dataset for Didymellaceae.
Leptosphaeria doliolum is used as the outgroup taxon. Bootstrap
support values for maximum likelihood greater than 60% and
Bayesian posterior probabilities greater than 0.80 are given at the
nodes. The ex-type strains are in bold. Novel species/new host records
sequences are in blue
thick-walled, rounded at the apex. Ascospores 14–20 9 7–10
lm (
x = 16 9 8 lm, n = 20), overlapping 1–2-seriate,
2-celled, hyaline, oval to ellipsoid, 1-septate in the centre,
deeply constricted at the septum, enlarge at the first cell,
rounded to conical at the ends, with small guttules, lacking a
mucilaginous sheath. Asexual morph Undetermined.
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Fungal Diversity
Fig. 47 continued
Culture characteristics: Colonies on MEA reaching 6
cm diam., after 7 days in the dark at 16 °C, circular with
entire edge, smooth, convex, white to white–gray with age.
Material examined: ITALY, Premilcuore Province, near
Fiumicello, on dead stem of Onobrychis viciifolia (Faboideae),
24 June 2014, Erio Camporesi, IT1954 (MFLU 14-0724,
holotype); ex-type living culture MFLUCC 15-0077.
GenBank Numbers LSU:KY496732; ITS:KY496752;
SSU:KY501120; RPB2:KY514410; TEF1:KY514397.
Notes: The sexual morph of Ascochyta species have
been referred to Didymella and Mycosphaerella (Corlett
1981; Peever et al. 2007). Our phylogeny indicates Ascochyta italica is close to A. viciae (asexual morph) which
was collected from the Netherlands on Vicia sepium. There
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Fungal Diversity
Fig. 48 Ascochyta italica (MFLU 14-0724, holotype). a Appearance
of ascomata on host substrate. b Section of ascoma. c Section of
peridium. d Pseudoparaphyses. e, f Ascus. g–i Ascospores.
j Germinating ascospore. Note: stained with cotton blue reagent in
f and i. Scale bars b, c = 20 lm, d = 2 lm, e–f = 10 lm, g–j = 5
lm
are five base pair differences in RPB2 nucleotide sequences
between A. viciae and A. italica.
Saprobic on dead branch of Scabiosa sp. Sexual morph
Ascomata 143–162 lm high 9 144–233 lm diam.
(
x = 152 9 196 lm, n = 5), semi-immersed to erumpent,
visible as raised, dark spots on the host surface, subglobose, solitary or small groups, scattered on the host surface,
without papilla, with short ostiole in the centre, thin-walled, carbonaceous, dark brown or usually black. Peridium
Ascochyta medicaginicola Q. Chen & L. Cai, Stud. Mycol.
82: 187 (2015)
Facesoffungi number: FoF 2773, Fig. 49
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Fungal Diversity
Fig. 49 Ascochyta medicaginicola (MFLU 15-1481). a, b Appearance of ascomata on host substrate. c Section of ascoma. d Section of
peridium. e Pseudoparaphyses. f–i Ascus. j–l Ascospores. Scale bars
a = 100 lm, b = 200 lm, c = 100 lm, d = 20 lm, e = 2 lm, f–
h = 10 lm, j–l = 5 lm
17–28 lm wide, thin-walled, inner cells hyaline to pale
brown, outer layer cells brown to dark brown, composed of
flattened cells of textura angularis. Hamathecium 2.5–3.8
lm wide, comprising numerous, septate, cylindrical, cellular, pseudoparaphyses. Asci 46–67 9 12–15 lm (
x = 55
9 14 lm, n = 15), 8-spored, bitunicate, fissitunicate,
cylindrical to cylindric-clavate, short pedicellate, rounded
at the apex with acute ends, lacking an ocular chamber.
Ascospores 17–22 9 7–9 lm (
x = 19 9 8 lm, n = 15),
overlapping 1–2-seriate in the ascus, fusiform, 2-celled,
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Fungal Diversity
narrowly fusoid with rounded ends, the cells above central
septum often broader than the lower ones, constricted at the
septum, deeply constricted at the septum, enlarged at the
first cell, with small guttules in each cell, hyaline, lacking a
mucilaginous sheath, smooth-walled. Asexual morph
Undetermined.
Material examined: ITALY, Province of Arezzo, near
Moggiona, on dead branch of Scabiosa sp. (Caprifoliaceae), 1 June 2013, Erio Camporesi, IT1314 (MFLU
15-1481, new host record); Ibid. (HKAS 94550 bis).
GenBank Numbers LSU:KY496720; ITS:KY496741;
SSU:KY501111.
Notes: Ascochyta medicaginicola isolates sampled for
LSU, ITS, b-tubulin and RPB2 based phylogenies clustered
in the same subclade with A. medicaginicola var. medicaginicola (Fig. 47). We report our collections as new host
records for A. medicaginicola. This is the first report of A.
medicaginicola on Scabiosa sp. (Caprifoliaceae) from Italy
and first report of a sexual morph for A. medicaginicola.
Ascochyta rosae Tibpromma, Camporesi & K.D. Hyde, sp.
nov.
Index Fungorum number: IF552694; Facesoffungi
number: FoF 2774, Figs. 50, 51
Etymology: Named after the host family (Rosaceae).
Holotype: MFLU 14-0723
Saprobic on dead branch of Rubus ulmifolia Sexual
morph Ascomata 398–451 lm high 9 364–398 lm diam.
(
x = 446 9 380 lm, n = 5), immersed, solitary, or in
small groups, uniloculate, globose, conspicuous at the
surface, dull black, without hairs, long ostiolate. Peridium
16–29 lm wide, comprising 4–5 layers of reddish-brown
cells of textura angularis. Hamathecium comprising 0.8–
2.3 lm wide, dense, septate, unbranched, pseudoparaphyses. Asci 48–74 9 7–9 lm (
x = 58 9 8 lm, n = 25),
8-spored, bitunicate, fissitunicate, cylindrical, with short,
club-shaped pedicel. Ascospores 13–17 9 3–5 lm (
x = 16
9 4.5 lm, n = 20), overlapping 1–2-seriate, fusiform,
hyaline, 2-celled, 1-septate in the centre, deeply constricted at the septum, enlarged at the second cell, conical
at the ends, with large guttules and with appendages at the
ends. Asexual morph Produced on sterilized bamboo
pieces with pine needles on water agar. Coelomycetous.
Conidiomata 68–105 9 77–98 lm, pycnidial, yellow
orange to black, globose, covered by dense vegetative
hyphae, uniloculate, solitary to scattered. Conidiophores
reduced to conidiogenous cells. Conidiogenous cells 5.3–
8.6 9 3.2–5.1 lm (
x = 6.6 9 3.9 lm, n = 20),
holoblastic, annelidic, globose to oblong, hyaline, and
formed from the inner layer of pycnidium wall. Conidia
3.2–4.4 9 1.2–1.8 lm (
x = 3.7 9 1.5 lm, n = 40),
cylindrical, rounded at both ends, sometimes slightly
curved, aseptate, hyaline, guttulate.
123
Culture characteristics: Colonies on MEA reaching 3–4
cm diam., after 7 days in the dark at 16 °C, circular with
entire edge, smooth, convex, flossy and velvety, white to
yellow-white with age. Asexual morph on WA at room
temperature (26 °C), convex, flossy and velvety, yellowish.
Material examined: ITALY, Galeata Province, near
Strada San Zeno, on dead branch of Rubus ulmifolia
(Rosaceae), 19 June 2014, Erio Camporesi, IT1946 (MFLU
14-0723, holotype); ex-type living culture MFLUCC
15-0063. (HKAS 94596bis, paratype).
GenBank Numbers LSU:KY496731; ITS:KY496751;
SSU:KY501119; RPB2:KY514409.
Notes: Ascochyta rosae is introduced with both sexual
and asexual morphs. The phylogeny indicates that A. rosae
clusters with A. herbicola with high statistical support
(100% in ML/1.00 in BYPP) (Fig. 47). Ascochyta rosae
and A. herbicola differ in the sizes of conidiomata and
conidia morphology. Conidia of A. rosae are smaller than
A. herbicola (3.2–4.4 9 1.2–1.8 vs. 7–10.5 9 2–2.5) and
the conidia are cylindrical to subcylindrical, without guttules (De Gruyter et al. 1998) while, A. rosae has cylindrical, sometimes slightly curved conidia with guttules.
RPB2 nucleotide sequence comparison reveals six base
pair differences with Ascochyta herbicola.
Calophoma Q. Chen & L. Cai
Calophoma was introduced by Chen & Cai (2015) with
C. clematidina (Thüm.) Q. Chen & L. Cai as the type
species. The asexual morph of this genus is characterized
by subglobose, subcylindrical, ellipsoidal, somewhat
obclavate-fusiform conidia with 0–1-septa and chlamydospores are produced in one species. The sexual morph of
this genus has not been reported (Chen et al. 2015).
Calophoma petasitis Tibpromma, Camporesi & K.D.
Hyde, sp. nov.
Index Fungorum number: IF552695; Facesoffungi
number: FoF 2775, Fig. 52
Etymology: Named after the host genus.
Holotype: MFLU 15-1489
Saprobic on dead stem of Petasites sp. Sexual morph
Ascomata 573–661 lm high 9 437–508 lm diam.
(
x = 624 9 494 lm, n = 5), superficial, flattened at the
base, solitary, black subglobose, solitary or aggregated,
ostiolate. Peridium 20–50 lm wide, comprising 1–2 layers,
the outer layer heavily encrusted with pigments and often
longitudinally striate on the surface pigmentation and the
inner layer comprising thick-walled, hyaline cells of textura angularis. Hamathecium comprising few, 0.6–1.5 lm
wide, septate, guttulate pseudoparaphyses. Asci 60–92 9
9–12 lm (
x = 81 9 10 lm, n = 10), 8-spored, bitunicate,
cylindrical, slightly curved, with short, furcate pedicel.
Ascospores 14–16 9 4–6 lm (
x = 15 9 6 lm, n = 20),
Fungal Diversity
Fig. 50 Ascochyta rosae
(MFLUCC 15-0063, holotype).
a, b Appearance of ascomata on
host substrate. c Section of
ascoma. d Ostiole. e Section of
peridium. f Ascus with
pseudoparaphyses. g–
j Ascospores. Scale bars
c = 100 lm, d = 10 lm,
e = 20 lm, f = 10 lm, g–j = 5
lm
overlapping 1–2-seriate, 2-celled, hyaline, 1-septate, guttulate, enlarged at the first cell, constricted at the septum,
conical at the ends, smooth-walled, lacking a mucilaginous
sheath. Asexual morph Undetermined.
Culture characteristics: Colonies on MEA reaching 7
cm diam. after 7 days at 16 °C, edge entire, brown to black
mycelium, raised, producing fruiting bodies on media.
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Fungal Diversity
Fig. 51 Ascochyta rosae (MFLU 14-0723, ex-type living culture on WA). a, b Appearance of cconidioma on culture. c Conidiomata. d–
f Conidiogenous cells. g Conidia. Scale bars a = 500 lm, b = 100 lm, c = 200 lm, d–f = 5 lm, g = 2 lm
Material examined: ITALY, Province of Forlı̀-Cesena,
near Campigna—Santa Sofia, dead stem of Petasites sp.
(Asteraceae), 9 June 2014, Erio Camporesi, IT1920
(MFLU 15-1489, holotype); ex-type living culture,
MFLUCC 15-0076); Ibid. (HKAS 94594 bis, paratype).
GenBank Numbers LSU:KY496729; ITS:KY496749;
SSU:KY501117; RPB2:KY514407.
Notes: Calophoma petasitis is introduced as a new
species with the first report of a sexual morph for this
genus. The sexual morph of C. petasitis is characterized by
subglobose ascomata and smooth-walled, guttulate, hyaline
ascospores lacking a mucilaginous sheath. Multigene
phylogenetic analyses support the position of the new
taxon in Calophoma and its establishment as a new species.
Calophoma petasitisi clearly segregates from other known
species (Fig. 47). We attempted to induce the asexual
morph of this species on WA media and found that they
produce ascomata on media.
Didysimulans Tibpromma, Camporesi & K.D. Hyde, gen.
nov.
Index Fungorum number: IF552770; Facesoffungi
number: FoF 2884
Etymology: refers to the didymella-like characteristics.
Type species: Didysimulans mezzanensis Tibpromma,
Camporesi & K.D. Hyde
123
Saprobic on dead stems. Sexual morph immersed under
the host tissue, solitary, black, globose to irregular, conspicuous at the surface, with or lacking ostioles. Peridium
composed of brown cells arranged in a textura angularis.
Hamathecium comprising dense, septate, guttulate pseudoparaphyses. Asci 8-spored, bitunicate, cylindrical,
slightly curved, short pedicellate, apically rounded. Ascospores 2-celled, hyaline, 1-septate, guttulate, enlarged at
the first cell, constricted at the septum, conical at the ends,
lacking a mucilaginous sheath. Asexual morph Chlamydospores occasionally produced, hyaline to yellow–brown
mycelium, in spiral chains.
Notes: Didysimulans is introduced as a new genus
based on morphology and phylogenetic support (LSU,
ITS, RPB2 and b-tubulin DNA sequence data). The
ascomata of Didysimulans are similar to those of
Didymellaceae in having a peridium of textura angularis
and cylindrical asci with 2-celled, hyaline, 1-septate
ascospores. Didysimulans produces chlamydospores with
hyaline to yellow–brown mycelium, with spiral chains.
Leptosphaerulina (muriform ascospores), Platychora (ascospores septate near the lower end), Monascostroma
(ascospores narrowly inequilateral), Macroventuria (ascomata with hairs or setae) and Didymella (ovoid to
ellipsoidal ascospores) (Hyde et al. 2013), are all easily
distinguishable from Didysimulans.
Fungal Diversity
Fig. 52 Calophoma petasitis (MFLU 15-1489, holotype). a Appearance of ascomata on host substrate. b Section of ascoma. c Section of
peridium. d Pseudoparaphyses. e–g Asci. h–j Ascospores.
k Germinating ascospore. Scale bars b = 50 lm, c = 5 lm, d = 2
lm, e–g = 10 lm, h–j = 2 lm
Didysimulans italica Tibpromma, Camporesi & K.D.
Hyde, sp. nov.
Index Fungorum number: IF552771; Facesoffungi
number: FoF 2885, Fig. 53
Etymology: refers to the country where the holotype was
collected.
Holotype: MFLU 14-0722
123
Fungal Diversity
Fig. 53 Didysimulans italica (MFLU 14-0722, holotype). a Appearance of ascomata on host substrate. b Section of ascoma. c Section of
peridium. d Pseudoparaphyses. e, f Ascus. g–i Ascospores.
j Germinating ascospore. Scale bars b = 50 lm, c = 20 lm, d = 2
lm, e, f = 10 lm, g–j = 2 lm
Saprobic on dead stem of Epilobium sp. Sexual morph
Ascomata 149–167 lm high 9 165–188 lm diam.
(
x = 159 9 177 lm, n = 5), immersed, solitary or in small
groups, uniloculate, globose, conspicuous at the surface,
dull, black, smooth, without hairs, ostiolate. Peridium 27–
58 lm wide, thick, comprising 5–6 layers of hyaline to
dark brown cells of textura angularis. Hamathecium
comprising 2.3–3.3 lm wide, dense, numerous, septate,
123
Fungal Diversity
unbranched, pseudoparaphyses. Asci 32–49 9 9–15 lm
(
x = 42 9 12 lm, n = 15), 8-spored, bitunicate, fissitunicate, cylindric, short pedicellate, thick-walled. Ascospores 9–14 9 3–5 lm (
x = 11 9 4 lm, n = 20),
overlapping 1–2-seriate, 2-celled, hyaline, oval or obclavate, 1-septate, slightly constricted at the septum, enlarged
at first cell, conical at the ends, lacking a mucilaginous
sheath. Asexual morph Undetermined.
Culture characteristics: Colonies on MEA reaching 4
cm diam., after 7 days in the dark at 16 °C, rhizoid colony
with undulate, smooth, raised on surface, white.
Material examined: ITALY, Galeata Province, Passo
delle Forche, on dead stem of Epilobium sp. (Onagraceae),
16 June 2014, Erio Camporesi, IT1936 (MFLU 14-0722,
holotype); ex-type living culture MFLUCC 15-0059
(HKAS 94595 bis, paratype).
GenBank Numbers LSU:KY496730; ITS:KY496750;
SSU:KY501118; RPB2:KY514408.
Notes: Didysimulans italica is introduced as a new species
with distinct ascospores which are conical at the ends.
Although D. italica is closely related to D. mezzanensis (type
species), they can be distinguished by morphology. Ascomata and ascospore characters are different (Figs. 53, 54).
Didysimulans mezzanensis Tibpromma, Camporesi &
K.D. Hyde, sp. nov.
Index Fungorum number: IF552772; Facesoffungi
number: FoF 2886, Figs. 54, 55
Fig. 54 Didysimulans mezzanensis (MFLU 15-1490, holotype). a Appearance of ascomata on host substrate. b Section of ascoma. c Section of
peridium. d Pseudoparaphyses. e–g Asci. h–j Ascospores. Scale bars b = 50 lm, c = 5 lm, d = 2 lm, e–g = 10 lm, h–j = 2 lm
123
Fungal Diversity
Fig. 55 Didysimulans
mezzanensis (MFLUCC
15-0067, ex-type living culture
on WA). a Growth of asexual
morph on plant tissues.
b Growth of asexual morph on
WA. c, e Squash mount of
vegetative hyphae. Scale bars a,
b = 200 lm, c = 20 lm, d,
e = 10 lm
Etymology: Named after the province which the fungus
was found.
Holotype: MFLU 15-1490
Saprobic on dead stem of Thalictrum sp. Sexual morph
Ascomata 83–95 lm high 9 110–139 lm diam. (
x = 89 9
124 lm, n = 5), immersed in host tissues, solitary, black,
irregular, conspicuous at the surface, ostiolate. Peridium 6–
29 lm wide, composed of two strata, an outer stratum of
thin-walled and brown cells arranged in a textura angularis
and inner layer of hyaline cells of textura angularis. Hamathecium comprising 0.8–1.2 lm wide, dense, septate,
guttulate pseudoparaphyses. Asci 45–60 9 10–13 lm
(
x = 53 9 12 lm, n = 15), 8-spored, bitunicate, cylindrical, slightly curved, short pedicellate, apically rounded.
Ascospores 13–16 9 4–8 lm (
x = 15 9 7 lm, n = 20),
overlapping 1–2-seriate, 2-celled, hyaline, 1-septate, guttulate, enlarged at the first cell, constricted at the septum,
conical at the ends, smooth-walled, lacking a mucilaginous
sheath. Asexual morph Colonies superficial to immersed;
123
mycelium composed of septate, branched, thick-walled,
smooth, guttulate, yellow green, coiled, smooth, hyphae,
producing thick-walled, small chlamydospores; sometimes,
hyphae packed into knot-like structures.
Culture characteristics: Ascospore germinating on
MEA within 12 h. Colonies on MEA reaching 5 mm diam.
in 7 days at 16 °C, rhizoidal with lobate edge, rough, flat
on surface, hyaline at beginning and becoming yellow
green, with septate, branched, guttulate, hyphae packed in
circular knots; not sporulating in culture.
Material examined: ITALY, near Mezzana Province,
Trento, dead stem of Thalictrum sp. (Ranunculaceae), 16
July 2014, Erio Camporesi, IT1998A (MFLU 15-1490,
holotype); ex-type living culture, MFLUCC 15-0067);
Ibid. (HKAS 94598 bis, paratype).
GenBank Numbers LSU:KY496733; ITS:KY496753;
SSU:KY501121; RPB2:KY514411.
Notes: Didysimulans mezzanensis is introduced as the
type species of Didysimulans. ITS nucleotide sequence
Fungal Diversity
comparison reveals eight base pair differences between
D. mezzanensis and D. italica.
Nothophoma Q. Chen & L. Cai
The genus Nothophoma was introduced with the type
species N. infossa (Ellis & Everh.) Q. Chen & L. Cai. (syn.
Phoma infossa) to accommodate five synonymized Phoma
species, including Nothophoma anigozanthi (Tassi) Q.
Chen & L. Cai (syn. Phoma anigozanthi), Nothophoma
arachidis-hypogaeae (V.G. Rao) Q. Chen & L. Cai (syn.
Phoma arachidis-hypogaeae), Nothophoma quercina
(Syd.) Q. Chen & L. Cai (syn. Phoma fungicola), and
Nothophoma gossypiicola (Gruyter) Q. Chen & L. Cai
(syn. Phoma gossypiicola) (Chen et al. 2015). This genus is
ubiquitous, species-rich and includes many important plant
pathogens, some of which are of quarantine concern
(Aveskamp et al. 2008, 2010; Chen et al. 2015).
Nothophoma quercina (Syd. & P. Syd.) Q. Chen & L. Cai,
Stud. Mycol. 82: 213 (2015)
Facesoffungi number: FoF 2688, Fig. 56
Basionym: Cicinobolus quercinus Syd., Ann. Mycol. 13:
42 (1915)
: Ampelomyces quercinus (Syd.) Rudakov, Mikol.
Fitopatol. 13: 109 (1979).
: Phoma fungicola Aveskamp et al., Stud. Mycol. 65:
26 (2010)
Saprobic on dead branch of Ulmus 9 hollandica Mill.
Sexual morph Undetermined. Asexual morph Conidiomata produced on surface of culture in PDA, 0.22–0.43
mm (
x = 0.28 mm, n = 10) diam., pycnidial, solitary,
scattered, globose to irregularly-shaped, black, ostiolate.
Pycnidia multi-layered, composed of pale brown, pseudoparenchymatous cells, with thick outer layer and thin inner
layer. Conidiogenous cells phialidic, hyaline, simple,
doliiform to ampulliform, variable in size. Conidia 2–5.5 9
1–4 lm (
x = 3.5 9 2.5 lm, n = 40), variable in size and
shape, subglobose to oval or obtuse, initially hyaline, but
brown at maturity, aseptate, smooth-walled. Conidial
exudates not recorded.
Culture characteristics: Colonies on PDA reaching 80
mm diam. after 7 days at 25 °C, with regular margin, dull
white aerial mycelium, surface floccose to downy, greenish-olivaceous to olivaceous near the centre and reverse
dark ochreous in the centre and white at the margin.
Material examined: ITALY, Forlı̀-Cesena Province, San
Lorenzo in Noceto—Forlı̀, on dead branch of Ulmus 9
hollandica (Ulmaceae), 31 March 2016, Erio Camporesi
(MFLU 16-2676, new host record), living culture,
MFLUCC 16-1392, KUMCC 16-0128.
GenBank Numbers LSU:KY053897; ITS:KY053896;
RPB2: KY053898; b-tubulin: KY053899.
Notes: Nothophoma quercina has been reported as a
saprobe from Quercus sp. in Ukraine (Aveskamp et al.
2010; Chen et al. 2015) and as a pathogen causing stem
canker and fruit blight on Pistacia vera L. in the USA
(Chen et al. 2013). This is the first record of N. quercina on
Ulmus species. When comparing our species with the type
specimen of N. quercina (CBS 633.92), they are similar in
morphology, but differ in their host. The conidiomata are
larger, while conidia are smaller compared to the type
specimen. Based on our phylogenetic analysis of combined
ITS, TEF1, b-tubulin and RPB2 sequence data of Nothophoma species (Fig. 47), our strain (MFLUCC 16-1392)
Fig. 56 Nothophoma quercina (MFLU 16-2676). a, b Conidiomata on the host tissue. c Pycnidia on PDA. d Section of pycnidial wall.
e Conidiogenous cells. f Conidia. Scale bars a = 1 mm, b = 0.2 mm, c = 2 mm, d–f = 20 lm
123
Fungal Diversity
clustered with the ex-type strain of Nothophoma quercina
with relatively high bootstrap and Bayesian probabilities
(99%MP/1.00BYPP).
Stagonosporopsis Died.
Stagonosporopsis produces dimorphic conidia, and most
of these conidia are aseptate, hyaline, ellipsoidal to subglobose, thin-walled and smooth-walled. The other type of
conidia are smaller, 1-celled or with up to 3 septa, produced both in vivo and in vitro in the same pycnidium
(Chen et al. 2015). Aveskamp et al. (2010) and Chen et al.
(2015) have detailed this genus.
Stagonosporopsis ailanthicola Manawasinghe, Camporesi
& K.D. Hyde, sp. nov.
Index Fungorum number: IF552548; Facesoffungi
number: FoF 2682, Fig. 57
Etymology: Name reflects the host genus Ailanthus.
Holotype: MFLU 16-2678
Saprobic on dead branch of Ailanthus altissima (Mill.)
Swingle. Sexual morph Undetermined. Asexual morph
Conidiomata 41–74 lm diam. (
x = 55 lm, n = 10), scattered, fusing when mature, globose to subglobose. Pycnidial wall multi-layered, comprising 2–3 layers, heavily
pigmented, thick-walled, comprising dark brown celled of
textura prismatica. Conidiophores reduced to conidiogenous cells. Conidiogenous cells enteroblastic, solitary,
hyaline, smooth-walled. Conidia 3.3–5.9 9 1.7–3.2 lm
(
x = 4.7 9 2.3 lm, n = 40), aseptate, hyaline, 1-celled,
ellipsoid, apex acutely rounded.
Culture characteristics: Colonies on PDA reach 60–65
mm diam., after 7 days at 25 °C. Floccose aerial mycelium
covering the whole colony.
Material examined: ITALY, Forlı̀-Cesena (FC) Province, Via Pietro Nenni—Forlı̀, on dead branch of Ailanthus altissima (Simaroubaceae), 2 May 2015, Erio
Camporesi IT2473, (MFLU 16-2678, holotype); ex-type
living culture, MFLUCC 16-1439.
GenBank Numbers ITS:KY100872; TEF1:KY100874;
b-tubulin:KY100878; RPB2:KY100876.
Notes: Stagonosporopsis species are characterized by
their unique dimorphic conidia, chlamydospores and
conidiomata (De Gruyter and Noordeloos 1992; Chen et al.
2015). Molecular data is important to identify and characterize these species. Combined phylogenetic analysis of
LSU, ITS, b-tubulin and RPB2 sequence data places
S. ailanthicola within the clade of Stagonosporopsis in the
Phoma species complex. Stagonosporopsis ailanthicola
differs from S. dorenboschii in having chlamydospores and
larger conidia. It forms a sister taxon with S. heliopsidis
with 100% bootstrap support and 1.00 Bayesian posterior
probability (Fig. 47). This is the first host record of
Stagonosporopsis species on Ailanthus.
123
Leptosphaeriaceae M.E. Barr
The family Leptosphaeriaceae was introduced by Barr
(1987) in the order Pleosporales and is typified by Leptosphaeria doliolum (Pers.) Ces. & De Not. Taxa commonly occur as saprobes or parasites (Hyde et al. 2013).
Members of this family are characterized by superficial or
erumpent ascomata, with single papillate ostiole lined with
periphyses, a relatively thick peridium wall, bitunicate
cylindrical asci and pale brown to brown, transversely
septate ascospores (Kodsueb et al. 2006; Ariyawansa et al.
2015a). The asexual morphs of the family Leptosphaeriaceae can be coelomycetous or hyphomycetous (Shenoy
et al. 2006; Zhang et al. 2012b; De Gruyter et al. 2013;
Ariyawansa et al. 2015a). Currently, seven genera are
accepted in the family: Alternariaster, Heterospora, Leptosphaeria, Neophaeosphaeria, Paraleptosphaeria, Plenodomus, and Subplenodomus (Ariyawansa et al. 2015a).
We introduce a new species in both Paraleptosphaeria and
Subplenodomus. Updated phylogenetic analyses followed
taxon sampling as outlined by Ariyawansa et al. (2015a)
(Fig. 58).
Paraleptosphaeria Gruyter, Aveskamp & Verkley
Paraleptosphaeria is typified by P. nitschkei (Rehm ex
G. Winter) Gruyter, Aveskamp and Verkley. De Gruyter
et al. (2013) introduced the genus to accommodate the
heterogeneous section of Leptosphaeria and its related
species in Leptosphaeriaceae. Paraleptosphaeria is characterized by having superficial or immersed, subglobose
ascomata, a thick-walled peridium comprising cells of
textura angularis, cylindric-clavate asci and transversly
septate ascospores. Ariyawansa et al. (2015a) included six
species in this genus.
Paraleptosphaeria padi Phukhamsakda, Bulgakov, Okatov
& K.D. Hyde, sp. nov.
Index Fungorum number: IF552709; Facesoffungi
number: FoF 2777, Fig. 59
Etymology: The species habitat regarding to the host
Prunus padus L.
Holotype: MFLU 15-2756
Saprobic on dead and dying branches of Prunus padus
L. Sexual morph Ascomata 288–345 lm high 9 330–396
lm diam. (
x = 316 9 363 lm, n = 5), solitary, scattered
or clustered in small groups, immersed under epidermal
layer, superficial on host surface, globose, black to dark
brown, coriaceous, ostiole central. Ostioles 119 9 83 lm,
papillate, shiny, dark brown, smooth, periphyses filling
ostiole channel. Peridium 9–53 lm wide, up to 72 lm at
base, outer region heavily pigmented, composed of 6–8
layers of textura angularis, black to dark-brown, 7–17(–20)
lm wide cells, inner layer composed of thick hyaline
gelatinous cells. Hamathecium comprising numerous, 2–
Fungal Diversity
Fig. 57 Stagonosporopsis
ailanthicola (MFLU 16-2678,
holotype). a Conidiomata.
b Chlamydospores.
c Developing chlamydospores.
d Conidia. e Conidiogenous
cells. f Pycnidial wall. g Surface
view of culture on PDA. Scale
bars b–e = 10 lm
3.1 lm (
x = 2 lm, n = 50) wide, long, transversely septate, branched, cellular pseudoparaphyses. Asci 74–127 9
8–14 lm (
x = 106 9 11 lm, n = 30), 8-spored, bitunicate,
broad-cylindrical, with club-shaped pedicel, apically
rounded, ocular chamber clearly visible when immature.
Ascospores 16–21 9 6–7 lm (
x = 18 9 6 lm, n = 50),
overlapping biseriate, pale brown, broad-fusiform, conical
at the apex, (2)–3 transversely septate, constricted at the
septa, enlarged at the second cell from apex, wall rough,
indentations present when mature. Asexual morph
Undetermined.
Material examined: RUSSIA, Arkhangelsk Region,
Arkhangelsk City, near northern Maimaksa urban
microdistrict, wood waste landfill, on dead and dying
branches of Prunus padus (Rosaceae), 11 May 2015, G.V.
Okatov, T-589 (MFLU 15-2756, holotype).
GenBank Numbers LSU:KY554198; SSU:KY554201;
ITS:KY554203; TEF1: KY554206.
123
Fungal Diversity
Fig. 58 Phylogenetic tree generated from maximum likelihood
analysis based on combined LSU, SSU, ITS, RPB2, and b-tubulin
sequence data from Leptosphaeriaceae. Bootstrap support values for
maximum likelihood greater than 50% and Bayesian posterior
probabilities greater than 0.80 are indicated above or below the
nodes as ML/BYPP are indicated above or below the nodes. Ex-type
strains and reference strains are in black bold. The new species are in
blue. The tree is rooted with Leptosphaeria doliolum (CBS 505.75)
Notes: Paraleptosphaeria padi is typical of Paraleptosphaeria based on the thick-walled peridium, immersed
under host epidermal layer, scleroplectenchymatous cell
layers, broad-cylindrical asci with a bulbous pedicel and
transversally septate ascospores (Ariyawansa et al.
2015a). We compared the new species with P. praetermissa that has been reported from Rosaceae. Even
though P. padi is similar to P. praetermissa in its
peridium, asci, and ascospore characteristics; the ascomata of P. praetermissa are erumpent to immersed,
whereas in P. padi they are superficial and smaller in
size (300–500 lm high 9 350–625 lm wide diam. vs.
288–345 lm high 9 330–396 lm wide diam.).
Paraleptosphaeria padi also shares similarities with
P. dryadis. However, P. dryadis has smaller ascomata
(180–275 lm high 9 150–250 lm wide and larger
ascospores (20–25 lm high 9 6–7 lm wide diam. The
phylogeny also supports the establishment of P. padi as a
new species with high support (Fig. 58).
123
Subplenodomus Gruyter, Aveskamp & Verkley
Subplenodomus was established by De Gruyter et al.
(2013), typified with S. violicola (P. Syd.) Gruyter, Aveskamp & Verkley, to accommodate some Phoma-like species that belong to Leptosphaeriaceae. Based on
morphological
characterization
and
multi-gene
Fungal Diversity
Fig. 59 Paraleptosphaeria
padi (MFLU 15-2752,
holotype). a Host substrate. b, c
Close up of ascomata on host.
d Vertical section of ascoma.
e Vertical section of ascoma
stained with cotton blue to show
the hyaline inner layer.
f Ostioles filled with periphyses.
g Section through peridium.
h Cellular pseudoparaphyses
(stained with cotton blue
showing septate). i–k
Developmental stages of asci. l–
p Developmental stages of
ascospores. q Ascospore stained
with cotton blue to show septa.
Scale bars c = 500 lm, d,
e = 200 lm, f = 100 lm, g–
k = 50 lm, l–q = 10 lm
phylogenetic analysis, Subplenodomus are accepted as
asexual morphs of Leptosphaeriaceae (De Gruyter et al.
2013; Hyde et al. 2013; Ariyawansa et al. 2015a). The
genus includes S. apiicola, S. drobnjacensis, S. valerianae
and S. violicola. Subplenodomus is characterized by thickwalled pycnidia, consisting of pseudoparenchymatous or
sometimes scleroplectenchymatous cell types, ostiole,
phialidic, ampulliform to doliiform conidiogenous cells
and hyaline, aseptate, and ellipsoid conidia (De Gruyter
et al. 2013). We reconstruct the phylogeny for
Leptosphaeriaceae using a concatenated dataset of RPB2
and b-tubulin. Our new collection of S. galicola (MFLU
15-1368) is close to S. violicola (Fig. 59).
Subplenodomus galicola Phukhamsakda, Tibpromma,
Camporesi, & K.D. Hyde, sp. nov.
Index Fungorum number: IF552711; Facesoffungi
number: FoF 2778, Fig. 60
Etymology: The species epithet refers to the host.
Holotype: MFLU 15-1368
123
Fungal Diversity
Fig. 60 Subplenodomus galicola (MFLU 15-1368, holotype) a Host
substrate. b Close up of ascomata on host. c Vertical section of
ascoma. d Ostioles filled with periphyses. e Section through peridium.
f Cellular pseudoparaphyses (stained with cotton blue showing septa).
g–i Developmental stages of asci. j–n Developmental stages of
ascospores. Scale bars b = 500 lm, c = 200 lm, d, e, g–i = 50 lm,
f, j–n = 20 lm
Saprobic on dead stem of Galium sp. Sexual morph
Ascomata 254–285 lm high 9 311–314 lm wide diam.
(
x = 269 9 313 lm, n = 5), solitary, scattered or sometimes gregarious in small groups, immersed under epidermis layer, superficial on host surface, depressed-globose,
ampulliform, base flat, dark brown to brown, coriaceous,
ostiole central. Ostioles 70–98 lm high 9 98–117 lm
diam. (
x = 87 9 104 lm, n = 5), papillate, shiny, brown
to reddish-brown, smooth, filled with periphyses. Peridium
32–60 lm wide, up to 75 lm at ostiole part, thick, outer
region of heavily pigment cells, outer layer of 8–10 layers
of black to dark-brown cells of textura angularis, inner
layer composed of thick-walled, hyaline, angular cells.
Hamathecium comprising numerous, long, 1.5–2.8 lm
(
x = 2 lm, n = 50) wide, broad, transversely septate,
branched, cellular pseudoparaphyses. Asci 66–120 9 12–
17 lm (
x = 98 9 14 lm, n = 20), 8-spored, bitunicate,
broad cylindrical, with club-shaped pedicel, apically
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Fungal Diversity
rounded, ocular chamber visible when immature. Ascospores 30–40 9 6–9 lm (
x = 37 9 7 lm, n = 50),
overlapping biseriate, pale brown, broad fusiform, tapering
at the ends, with (3)–4 transverse septa, slightly constricted
at the septa, enlarged at the second cell from apex, wall
rough, indentations present when mature. Asexual morph
Undetermined.
Material examined: ITALY, Forlı̀-Cesena [FC], Tontola
di Predappio, on dead stem of Galium sp., 4 April 2014,
Erio Camporesi, IT 1797 (MFLU 15-1368, holotype).
GenBank Numbers LSU:KY554199; ITS:KY554204.
Notes: Subplenodomus galicola is similar to other Subplenodomus species in its pseudoparenchymatous peridium
cell types (De Gruyter et al. 2013). Subplenodomus galicola also shares similarities with Plenodomus in its cylindric asci and yellowish ascospores with 3–5 transverse
septa. However, they are distinct, as the ascomata of
S. galicola are, superficial, whereas Plenodomus has
immersed to erumpent ascomata (Ariyawansa et al. 2015a).
Currently, the species of Subplenodomus are known from
herbaceous plants or woody dicotyledonous trees (De
Gruyter et al. 2013).
Phaeosphaeriaceae M.E. Barr.
The family Phaeosphaeriaceae, introduced by Barr
(1979), was originally characterized by immersed to
superficial, globose to subglobose ascomata, short papilla,
bitunicate asci and hyaline, yellowish or brown, fusiform to
ellipsoidal, filiform, or muriform, septate ascospores (Barr
1979; Shoemaker 1984; Shoemaker and Babcock 1989;
Zhang et al. 2012b; Hyde et al. 2013; Phookamsak et al.
2014). Most species in this family are widely distributed on
plants as necrotrophic pathogens or as saprobes (Shoemaker and Babcock 1989; Carson 2005; Stukenbrock et al.
2006; Cannon and Kirk 2007). Phaeosphaeriaceae shows
diverse morphological characters and comprise more than
35 sexual and asexual genera (Phookamsak et al. 2014;
Wijayawardene et al. 2014a; Hyde et al. 2016).
Entodesmium Riess
Entodesmium was introduced by Riess in 1854 with
E. rude Riess as the type species. Entodesmium has
immersed, globose ascomata, and filiform, multi-septate
ascospore, some breaking into part ascospores (Shoemaker
1984; Barr 1992; Zhang et al. 2012b; Phookamsak et al.
2014) (Fig. 61).
Entodesmium italica Tibpromma, Camporesi & K.D.
Hyde, sp. nov.
Index Fungorum number: IF552697; Facesoffungi
number: FoF 2788, Fig. 62
Etymology: refers to the country where the fungus was
collected.
Holotype: MFLU 14-0716
Saprobic on decaying plant stems of Lathyrus sp. Sexual morph Ascomata 256–289 lm high, 176–201 lm
diam. (
x = 266 9 190 lm, n = 5), immersed to semiimmersed, scattered beneath the host periderm or on
decorticated wood, visible as small black dots on the host
surface, ampulliform, solitary, with central, shiny ostiole.
Ostioles central, long, inconspicuous at the surface.
Peridium 25–32 lm wide, comprising 3–5 layers, of dark
brown cells of textura angularis. Hamathecium comprising
numerous, 0.9–3.7 lm wide, filamentous, unbranched,
cellular, guttulate pseudoparaphyses. Asci 72–98 9 9–10
lm (
x = 89 9 10 lm, n = 10), 8-spored, bitunicate, fissitunicate, cylindric-clavate, with long, club-shaped pedicel. Ascospores 24–30 9 4–5 lm (
x = 267 9 4 lm,
n = 15), overlapping biseriate, yellow-green, usually
4-septate, constricted or not at the septum, conical at the
ends, guttulate, smooth-walled, not separating at the septa,
lacking a mucilaginous sheath. Asexual morph
Undetermined.
Culture characteristics: Colonies on MEA reaching 3–4
cm diam., after 7 days in the dark at 18 °C, yellowish,
irregular, flattened to slightly raised, with rough surface,
fairly fluffy, not producing pigment in agar.
Material examined: ITALY, Province of Arezzo, Poppi,
on dead stem of Lathyrus sp. (Fabaceae), 14 May 2014,
Erio Camporesi, IT1875 (MFLU 14-0716, holotype); living culture, MFLUCC 14-0526; (HKAS 94588 bis,
paratype).
GenBank Numbers LSU:KY496727; ITS:KY496747;
RPB2:KY514406; TEF1:KY514395.
Notes: Molecular analysis indicates that the new species
belongs in the genus Entodesmium in the Phaeosphaeriaceae clade and clusters with E. rude (type) (Fig. 61).
Entodesmium rude differs in having longer filiform
ascospores that break into part spores (Shoemaker 1984;
Phookamsak et al. 2014).
Neosetophoma Gruyter, Aveskamp & Verkley
Neosetophoma is typified by N. samararum (Desm.) and
most species are pathogens causing leaf spots of various
hosts (Phookamsak et al. 2014). The genus is characterized
by globose to irregular conidiomata, with papillate ostioles,
and yellowish conidia that are attenuate at one end (De
Gruyter et al. 2010) and no sexual morph is known.
Neosetophoma garethjonesii Tibpromma, EBG Jones &
K.D. Hyde, sp. nov.
Index Fungorum number: IF552698; Facesoffungi
number: FoF 2789, Fig. 63
Etymology: Named in honour of Professor E.B. Gareth
Jones who turned 80 years old in January 2017 and in
recognition of his immense contribution to mycology.
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Fungal Diversity
b Fig. 61 RAxML tree based on analysis of a combined dataset of ITS,
LSU, SSU and RPB2 partial sequences. Bootstrap support values for
maximum likelihood (ML, black) higher than 50% and Bayesian
posterior probabilities (BYPP, black) greater than 0.90 are defined as
above the nodes. The tree is rooted to Didymella exigua. All type
strains are in bold. New species are given in blue
Holotype: MFLU 16-1921
Saprobic on dead stem. Sexual morph Ascomata 403–
444 lm high 9 332–363 lm diam. (
x = 424 9 348 lm,
n = 5), globose to subglobose, solitary, dark brown to
black, immersed, black, smooth, ostiolate canal filled
without sparse periphyses. Peridium 10–30 lm wide, thin,
comprising 3–4 layers of brown to reddish-brown cells of
textura angularis. Hamathecium of 1.5–2 lm wide, guttulate pseudoparaphyses. Asci 44–75 9 5–9 lm (
x = 61 9
7 lm, n = 10), 8-spored, bitunicate, fissitunicate, cylindrical, short pedicellate, thin-walled at the apex, with a
minute ocular chamber. Ascospores 16–26 9 3–5 lm
(
x = 21 9 4 lm, n = 20), overlapping 1–2-seriate, hyaline
when young, becoming yellow pale at maturity, 4–5-celled,
usually 3-septate, fusiform, narrowly fusoid with rounded
ends, constricted at the septum, enlarged at the second cell,
guttulate, lacking a mucilaginous sheath, smooth-walled.
Asexual morph Undetermined.
Culture characteristics: Colonies on MEA reaching 3–4
cm diam., after 5 days in the dark at 18 °C, brown to darkbrown irregular, flattened to slightly raised, with rough
surface, fairly fluffy.
Material examined: UK, Hampshire, the New Forest, on
dead stem of unknown plant, 30 March 2014, EBG Jones,
GJ004, (MFLU 16-1921, holotype); (HKAS 94525 bis,
paratype), ex-type living culture MFLUCC 14-0528.
GenBank Numbers LSU:KY496738; ITS:KY496758;
SSU:KY501126; TEF1:KY514402.
Notes: The phylogenetic analyses place N. garethjonesii
in Neosetophoma as a new species with good statistical
support (95% in ML/0.99 in BYPP) (Fig. 61). The sexual
morph of Neosetophoma had not been previously reported
(De Gruyter et al. 2010, Liu et al. 2015). Neosetophoma
garethjonesii is introduced as a new species and is the first
report of the sexual morph of Neosetophoma.
Nodulosphaeria Rabenh.
The genus Nodulosphaeria was introduced by Rabenhorst (1858) and typified with N. hirta Rabenh. The characteristics of this genus are ascomata lined with brown
setae, and three- to multi-septate ascospores, with a swollen cell and some with terminal appendages (Holm 1961;
Shoemaker 1984; Mapook et al. 2016). Clements and Shear
(1931) considered Nodulosphaeria a synonym of Leptosphaeria, but Holm (1957) showed Nodulosphaeria was
different to Leptosphaeria. Recent studies by Phookamsak
et al. (2014), Ariyawansa et al. (2015a) and Mapook et al.
(2016) included a reference strain of Nodulosphaeria
modesta, morphological characteristics and investigation of
the familial placement and generic affinities based on
combined DNA sequence data.
Nodulosphaeria guttulatum Tibpromma, Camporesi &
K.D. Hyde, sp. nov.
Index Fungorum number: IF552699; Facesoffungi
number: FoF 2790, Fig. 64
Etymology: ‘guttulatum’ referring to ascospore containing numerous guttules.
Holotype: MFLU 14-0712
Saprobic on dead stem of Scabiosa sp. Sexual morph
Ascomata 197–218 lm high 9 163–244 lm diam.
(
x = 206 9 199 lm, n = 5), immersed, solitary, or in
small groups, uniloculate, globose to subglobose, conical at
the base, black, hairy. Ostiole papillate, protruding, with
numerous, short, brown to dark brown setae. Peridium 15–
30 lm wide, comprising 6–7 layers of hyaline to reddishbrown cells of textura angularis. Hamathecium 2.1–3.9 lm
wide, comprising dense, numerous, aseptate, distinctly
guttulate pseudoparaphyses. Asci 40–89 9 10–15 lm
(
x = 71 9 13 lm, n = 20), 8-spored, bitunicate, fissitunicate, cylindric-clavate, with long, club-shaped pedicel,
thick-walled at the apex, with a minute ocular chamber.
Ascospores 29–38 9 5–6.5 lm (
x = 33 9 6 lm, n = 15),
overlapping 1–2-seriate, hyaline when young, becoming
golden yellow at maturity, fusiform, 4-septate, slightly
constricted at the septum, enlarged at the second cell from
apex, conical and narrowly rounded at the ends, with
numerous guttulate, with circular sheath at both ends.
Asexual morph Undetermined.
Culture characteristics: Colonies on MEA reaching 3–4
cm diam., after 7 days in the dark at 16 °C, white–gray to
dark, cottony, flat, dense, with sparse aerial mycelium on
the surface.
Material examined: ITALY, Predappio Province, Rocca
delle Caminate, on dead stem of Scabiosa sp. (Caprifoliaceae), 21 April 2014, Erio Camporesi, IT1826 (MFLU
14-0712, holotype); ex-type living culture MFLUCC
15-0069 (HKAS bis, paratype).
GenBank Numbers LSU:KY496726; ITS:KY496746;
SSU:KY501115; RPB2:KY514405; TEF1:KY514394.
Notes: Nodulosphaeria guttulatum differs from
N. aconita, N. modesta, N. scabiosae, and N. spectabilis
in having ascospores that are slightly constricted at the
septa, enlarged at the second cell from the apex, numerous guttules, and with a circular sheath at both ends.
Phylogenetic analyses indicate that N. guttulatum is closely related to N. multiseptata with high support (90% in
ML/0.98 in BYPP) (Fig. 61), but is morphologically
different.
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Fungal Diversity
Fig. 62 Entodesmium italica (MFLU 14-0716, holotype). a Fruiting
bodies on host surface. b Section through ascoma. c Section through
papilla. d Section through peridium. e Pseudoparaphyses. f–h Asci. i–
123
k Ascospores. l Germinated ascospore. Scale bars a = 100 lm, b,
c = 50 lm, d = 5 lm, e = 5 lm, f–h = 20 lm, i–l = 10 lm
Fungal Diversity
Fig. 63 Neosetophoma garethjonesii (MFLU 16-1921, holotype). a,
b Appearance of ascomata on host substrate. c Section of ascoma.
d Section of peridium. e Pseudoparaphyses. f–g Asci. h–k Ascospores.
l Germinating ascospore. Scale bars c = 50 lm, d = 10 lm, e = 5
lm, f, g = 10 lm, h–k = 10 lm, l = 5 lm
Nodulosphaeria multiseptata Tibpromma, Camporesi &
K.D. Hyde, sp. nov.
Index Fungorum number: IF552700; Facesoffungi
number: FoF 2791, Fig. 65
Etymology: refers to the ascospore morphology.
Holotype: MFLU 14-0720
Saprobic on dead stem of Sambucus ebulus L. Sexual
morph Ascomata 232–260 lm high 9 267–272 lm diam.
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Fungal Diversity
Fig. 64 Nodulosphaeria guttulatum (MFLU 14-0712, holotype).
a Appearance of ascomata on host substrate. b Section of ascoma.
c Section of peridium. d Hairs. e Ostiole. f Pseudoparaphyses. g–i
Ascus. j–l Ascospores. m Germinating ascospore. Scale bars b = 50
lm, c–e = 10 lm, f = 5 lm, g–l = 20 lm, j–m = 10 lm
(
x = 246 9 269 lm, n = 5), immersed to semi-immersed,
solitary, uniloculate, globose to subglobose, black, smoothwalled. Ostiole 60–74 9 83–102 lm (
x = 67 9 96 lm,
n = 5), papillate, protruding from the centre of ascomata,
with numerous, short, internal brown to dark brown setae.
Peridium 19–27 lm wide, comprising of dark brown cells
of textura angularis. Hamathecium 3.3–4.5 lm wide,
comprising numerous, aseptate, distinctly guttulate, pseudoparaphyses. Asci 74–127 9 17–21 lm (
x = 100 9 19
lm, n = 20), 8-spored, bitunicate, fissitunicate, cylindric-
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Fungal Diversity
Fig. 65 Nodulosphaeria multiseptata (MFLU 14-0720, holotype).
a Appearance of ascomata on host substrate. b Section of ascoma.
c Section of peridium. d, e Section of ostiole. f Pseudoparaphyses. g,
h Asci. i, j Ascospores. k Germinating ascospore. Scale bars a = 200
lm, b = 50 lm, c = 10 lm, d, e = 20 lm, f = 5 lm, g–k = 20 lm
clavate, with short, club-shaped pedicel, thick-walled at the
apex, with a minute ocular chamber. Ascospores 56–70 9
5–6 lm (
x = 65 9 6 lm, n = 20), overlapping 1–2-seriate,
hyaline when young, becoming golden yellow at maturity,
fusiform, 12–14-septate, slightly constricted at the septum,
enlarged at fifth cell from apex, conical and narrowly
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Fungal Diversity
rounded at the ends, guttulate, with mucilaginous appendages at the ends. Asexual morph Undetermined.
Culture characteristics: Colonies on MEA reaching 8
cm diam. after 7 days at 16 °C, whitish-gray, cottony,
dense, with sparse aerial mycelium on the surface.
Material examined: ITALY, Province of Arezzo, near
Ortignano-Raggiolo, on dead stem of Sambucus ebulus
(Adoxaceae), 25 May 2014, Erio Camporesi, IT1889
(MFLU 14-0720, holotype); ex-type living culture
MFLUCC 15-0078 (HKAS 94591, paratype).
GenBank Numbers LSU:KY496728; ITS:KY496748;
SSU:KY501116; TEF1:KY514396.
Notes: In the phylogenetic analysis N. multiseptata
clustered with N. guttulatum with high statistical support
(90% in ML, 0.98 in BYPP), but the species are different.
Nodulosphaeria guttulatum has 4-septate, fusiform ascospores, enlarged at the second cell from the apex, with
conical and narrowly rounded ends (Fig. 65).
Nodulosphaeria sambuci Tibpromma, Camporesi & K.D.
Hyde, sp. nov.
Index Fungorum number: IF552701; Facesoffungi
number: FoF 2792, Fig. 66
Etymology: named for its occurrence on the host plant.
Holotype: HKAS94536
Saprobic on dead stem of Sambucus ebulus L. Sexual
morph Ascomata 204–220 lm high 9 173–215 lm diam.
(
x = 213 9 198 lm, n = 5), superficial, solitary, or in
small groups, uniloculate, globose to subglobose, conspicuous at the surface, black, hairy. Ostioles 78–92 9
107–126 lm (
x = 86 9 117 lm, n = 5), papillate, protruding from the centre of the ascomata, with numerous,
short, brown to dark brown setae. Peridium 23–44 lm
wide, comprising of light-brown to dark brown cells of
textura angularis. Hamathecium 1.1–3.4 lm wide, comprising dense, numerous, aseptate, distinctly guttulate
pseudoparaphyses. Asci 53–80 9 7–10 lm (
x = 68 9 9
lm, n = 20), 8-spored, bitunicate, fissitunicate, cylindricclavate, with small furcate pedicel, apically rounded, with a
minute ocular chamber. Ascospores 32–35 9 3–5 lm
(
x = 34 9 4 lm, n = 15), overlapping 1–2-seriate, hyaline
when young, becoming golden yellow at maturity, fusiform, 8-septate, slightly constricted at the septum, enlarged
at four cell, conical and narrowly rounded at the ends,
guttulate, with small mucilaginous appendages at both
ends. Asexual morph Undetermined.
Culture characteristics: Colonies on MEA reaching 4
cm diam. after 7 days at 16 °C, gray to dark, cottony, flat,
dense, with sparse aerial mycelium on the surface
Material examined: ITALY, Province of Forlı̀-Cesena,
Monte Fumaiolo, on dead stem of Sambucus ebulus, 10
July 2012, Erio Camporesi, IT558 (HKAS 94536, holotype); ex-type living culture MFLUCC 15-0068.
123
GenBank Numbers LSU:KY496721; ITS:KY496742;
SSU:KY501112.
Notes: Nodulosphaeria sambuci shares similarities with
species having 8-septate ascospores (Nodulosphaeria
derasa, N. dolioloides and N. pellita), but those species
lack molecular data. It is phylogenetically closely related to
N. senecionis with high statistical support (100% in ML,
1.00 in BYPP). However, N. sambuci has fusiform, golden
yellow ascospores, that are enlarged at the fourth cell from
the apex, and conical and narrowly rounded at the ends,
while N. senecionis has long fusiform to almost cylindrical,
yellowish brown ascospores, enlarged at the third cell from
the apex and tapering towards the rounded ends (Ariyawansa et al. 2015c).
Phaeosphaeria I. Miyake
Phaeosphaeria was established by Miyake (1909) with
P. oryzae as the type species. The epitypification of
Phaeosphaeria oryzae I. Miyake and Phaeoseptoria
papayae Speg. are confirmed through sexual and asexual
morph connections and based on evidence from phylogenetic analysis (Quaedvlieg et al. 2013; Phookamsak et al.
2014).There are 203 epithets for Phaeosphaeria in Index
Fungorum (2017) with 19, 363 nucleotide sequences
available in GenBank.
Phaeosphaeria calamicola Konta & K. D. Hyde, sp. nov.
Index Fungorum number: IF552662; Facesoffungi
number: FoF02758, Fig. 67
Etymology: Referring to the host substrate (Calamus sp.).
Holotype: MFLU 15-0019
Saprobic on Calamus sp. Sexual morph Ascomata 76–
110 lm high 9 103–125 lm diam. (
x = 95 9 117 lm,
n = 5), solitary, scattered, semi-immersed, subglobose,
with ostiole at the centre. Peridium 9–14 lm wide, composed of outer layers of light brown, thick-walled, cells of
textura angularis, inner layers of hyaline and thin-walled
cells. Hamathecium 1–3 lm wide, comprising dense,
numerous, septate, pseudoparaphyses. Asci 41–63 9 8–11
lm (
x = 54 9 9 lm, n = 10), 8-spored, bitunicate, overlapping 2-seriate cylindric-clavate, short-pedicellate, apically rounded, lacking a distinct ocular chamber.
Ascospores 18–22 lm high 9 4–5 lm diam. (
x = 20 9 5
lm, n = 10), 2–3-seriate, initially hyaline, becoming pale
brown at maturity, fusiform, 3-septate, guttulate, rounded
at the ends, surrounded by a thick mucilaginous sheath.
Asexual morph Undetermined.
Culture characteristics: Ascospore germinating on
MEA within 24 h and germ tube produced from both ends
of the ascospore. Colonies on MEA reaching 3–4.5 cm
diam. after 4 weeks at 25 °C, white from the centre to
margins and slightly fluffy at the margins, dense, septate,
branched, smooth-walled.
Fungal Diversity
Fig. 66 Nodulosphaeria sambuci (HKAS94536, holotype). a Appearance of ascomata on host substrate. b Section of ascoma. c, d
Section of ostiole. e Section of peridium. f Pseudoparaphyses. g, h
Asci. i–k Ascospores. l Germinating ascospore. Scale bars a = 100
lm, b = 50 lm, c–e = 20 lm, f = 5 lm, g, h = 20 lm, i–l = 10 lm
Material examined: THAILAND, Chiang Rai Province,
Doi Chang, on dead stem of Calamus sp. (Arecaceae), 8
November 2014, Sirinapa Konta, DC02a (MFLU 15-0019,
holotype); ex-type living culture, MFLUCC 14-1168.
GenBank Numbers LSU:KY511423; ITS:KY511429;
SSU:KY511426.
Notes: Phaeosphaeria calamicola clustered with P. chiangraina in the Phaeosphaeria clade with high support (95%
ML, 0.97 BYPP). The phylogenetic analyses (Fig. 61)
indicate that P. calamicola belongs in Phaeosphaeria within
Phaeosphaeriaceae. Phaeosphaeria calamicola differs from
P. chiangraina, P. papayae and P. oryzae in its much smaller
ascomata, asci and ascospores.
Phaeosphaeriopsis M.P.S. Câmara, M.E. Palm & A.W.
Ramaley
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Fungal Diversity
Fig. 67 Phaeosphaeria calamicola (MFLU 15-0019, holotype).
a Appearance of ascomata on host substrate. b Close up of ascomata.
c Section of ascoma. d Peridium. e Pseudoparaphyses. f–j Asci. k–
123
o Ascospores. Note sheath in o in India Ink. Scale bars a = 500 lm,
b = 100 lm, c = 50 lm, d = 20 lm, e = 10 lm, f–j = 20 lm, k–
o = 10 lm
Fungal Diversity
Phaeosphaeriopsis, typified by P. glaucopunctata, was
introduced to accommodate non-congeneric species of
Paraphaeosphaeria by Câmara et al. (2003) and followed
by Phookamsak et al. (2014).
Phaeosphaeriopsis yuccae Dayarathne, Bulgakov, E.B.G.
Jones & K.D. Hyde, sp. nov.
Index Fungorum number: IF552710; Facesoffungi
number: FoF 2793, Fig. 68
Etymology: Named referring to the host genus.
Holotype: MFLU 15-2938
Saprobic on dying leaves of Yucca sp. Sexual morph
Undetermined. Asexual morph Conidiomata 97–161.5 lm
high 9 104–178 lm diam., pycnidial, solitary, semi-immersed to immersed, visible as slightly, raised, small black
spots on host surface, globose to subglobose or cup-shaped
when sectioned, with centrally placed ostiole. Conidiomata
wall 13–34 lm wide, composed of 3–4 layers of brown
cells, arranged in textura angularis, outer layers comprising 2–3 layers of brown to dark brown, thick-walled cells,
inner layer comprising 2–3 layers of hyaline, thin-walled
cells. Conidiophores simple, rarely branched, doliiform to
cylindrical or ampulliform, septate, hyaline, mostly
reduced to conidiogenous cells. Conidiogenous cells 3–5 9
2–4 lm, holoblastic, phialidic, single, discrete, sometimes
integrated, ampulliform or cylindric-clavate, hyaline, arising from basal stratum. Conidia 2–3.5 9 2–3 lm (
x = 2.9
9 2.8 lm, n = 30), initially hyaline, becoming brown to
dark brown, globose to subglobose, aseptate, smoothwalled.
Culture characters: Colonies on potato dextrose agar
(PDA) 24–30.5 mm diam. after 14 days at 25–30 °C, white
to cream at the margins, pale yellowish to yellowish-white,
white at the centre; reverse white to cream at the margins,
brown to orange-brown in the centre and pale yellowish to
yellowish at the centre; medium dense, irregular, flattened
to slightly raised, with rough surface, undulate edge with
entire to slightly radiating margins, floccose to fairly fluffy,
no pigments produced in agar.
Material examined: RUSSIA, Rostov Region, Rostovon-Don City, Botanical Garden of Southern Federal
University,
ornamental
nursery,
(47.23711210 E,
0
39.6443796 N), on Yucca sp. (Agavaceae), 22 July 2015,
Timur Bulgakov (MFLU 15-2938, holotype); ex-type living culture, MFLUCC 16-0558.
GenBank Numbers SSU:KY554480; LSU:KY554481;
ITS:KY554482.
Notes: Phaeosphaeriopsis yuccae is characterized by
having pycnidial conidiomata, doliiform to cylindrical or
ampulliform conidiophores and globose to subglobose,
dark brown conidia at maturity. Paraphaeosphaeria species differ in their asexual morph characteristics, including
conidiomatal structure, type of conidiogenesis, and
conidial morphology. Some asexual morphs are Coniothyrium-like, whereas others are more typical of Microsphaeropsis Höhn (Câmara et al. 2003). This new strain
is Coniothyrium-like and morphologically most similar to
asexual morph of Phaeosphaeriopsis dracaenicola in
conidiomata, conidiophores and conidial nature, but they
are phylogenetically distant in our phylogram (Fig. 62).
According to phylogenetic analyses P. phacidiomorpha has
a close affinity to P. yuccae, but the two strains are wellresolved with high statistical support (83% ML, 1.00
BYPP) (Fig. 61). Furthermore, the conidiomata of
P. phacidiomorpha is cylindrical, dark brown and 3 mm
long and typical of the Microsphaeropsis-like asexual
morph of Phaeosphaeriopsis (Farr et al. 2006; Câmara
et al. 2003).
Pleosporineae genera incertae sedis
Camarosporium Schulzer
Camarosporium was established by Schulzer (1870),
with Camarosporium quaternatum as the type species. The
genus Camarosporium has been referred to different families (Liu et al. 2015; Wijayawardene et al.
2014b, c, 2015, 2016; Ariyawansa et al. 2015b). Currently,
there are more than 500 names listed in the genus (Index
Fungorum 2017). Based on multi-locus phylogeny,
Wijayawardene et al. (2014c) placed Camarosporium
sensu stricto in Pleosporineae, Pleosporales. This genus
has been reported as the asexual morph of many important
fungal families and orders including Cucurbitariaceae,
Leptosphaeriaceae,
Phaeosphaeriaceae
and
Botryosphaeriales (Kirk et al. 2008; Wijayawardene et al.
2012; Hyde et al. 2013) (Fig. 69).
Camarosporium laburnicola R.H. Perera, Bulgakov, &
K.D. Hyde, sp. nov.
Index Fungorum number: IF552706; Facesoffungi
number: FoF 2784; Fig. 70
Holotype: MFLU 15-1522
Etymology: Named after the host genus on which the
fungus occurs.
Saprobic on branches of Laburnum anagyroides Medik.
Sexual morph Ascomata 280–370 lm high, 220–320 lm
diam. (
x = 331.4 9 274.2 lm, n = 5), erumpent through
the host epidermis, globose to subglobose or obovoid,
solitary or arranged in groups of 2–22, black, Ostioles
central, short, periphysate. Periphyses cellular, aseptate,
with a blunt apex, hyaline. Peridium 22–69 lm wide, thick,
comprising 8–14 layers, outer layer heavily pigmented,
thin- walled, comprising dark brown cells of textura
globulosa to textura angularis, inner layer composed of
hyaline to brown thin-walled cells of textura globulosa to
textura angularis. Hamathecium comprising numerous,
1.1–3.1 lm (n = 30) wide, filamentous, branched,
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Fungal Diversity
Fig. 68 Phaeosphaeriopsis yuccae (MFLU 15-2938, holotype) a, b
Appearance of conidiomata on Yucca sp. c Vertical section of
conidioma. d Peridium. e–g Developing stages of conidia on
conidiogenous cells. h–k Conidia. l Germinating conidium. m, n
Cultures on PDA (m upper, n lower). Scale bars b = 500 lm,
c = 100 lm, d = 20 lm, e = 10 lm. f, g, l = 5 lm, h–k = 2 lm
aseptate, pseudoparaphyses. Asci 125–150 9 9–11 lm
(
x = 134 9 9.8 lm, n = 30), 8-spored, bitunicate, fissitunicate, cylindrical, thick-walled at the apex, with a
minute ocular chamber. Ascospores 15–21 9 6–8 lm
(
x = 17.4 9 6.6 lm, n = 40), uniseriate to overlapping
uniseriate, muriform, ellipsoidal, 6–7 transversely septate,
with 5–6 vertical septa constricted at the septa, initially
hyaline, becoming yellowish-brown at maturity, conical
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Fungal Diversity
Fig. 69 Maximum likelihood tree based on combined LSU, SSU and
ITS sequenced data of Camarosporium sensu stricto. Maximum
likelihood and parsimony bootstrap support values greater than 50%
are indicated near the nodes, and branches with Bayesian posterior
probabilities greater than 0.95 are given in bold. The ex-types are in
bold and newly introduced species are in blue. The scale bar indicates
0.01 changes. The tree is rooted with Pyrenochaeta lycopersici (CBS
306.65)
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Fungal Diversity
Fig. 70 Camarosporium laburnicola (MFLU 15-1522, holotype). a,
b Appearance of ascomata on host substrate. c Vertical section
through ascomata. d Close up of ostiole. e Close up of the peridium.
f Pseudoparaphyses. g–i Immature and mature asci. j–n Immature and
mature ascospores. o Germinating ascospore. p, q Colonies on MEA.
Scale bars b = 2 cm, b = 200 lm, d–i = 50 lm, j–o = 10 lm
and narrowly rounded at the ends, without a mucilaginous
sheath. Asexual morph Undetermined.
Culture characteristics: Colonies on PDA reaching
40–45 mm diam. in 28 days, pinkish-grey, spreading with
moderate aerial mycelium, slightly irregular, margins
smooth, with underneath pinkish-grey.
Material examined: RUSSIA, Rostov Region, Rostovna-Donu City, Botanical garden of Southern Federal
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Fungal Diversity
University, Higher Park, dead branches of Laburnum
anagyroides Medik. (=Cytisus laburnum L.), 5 March
2014, T. Bulgakov, T-81 (MFLU 15-1522, holotype; ibid.
HKAS 89691, isotype); ex-type living culture MFLUCC
14-0565).
GenBank Numbers LSU:KY497779; SSU:KY497781;
ITS:KY497784; TEF1:KY497785.
Notes: Camarosporium uniseriatum clusters with two
coelomycetous species, C. clematidis and C. spartii with
high bootstrap support (Fig. 69). Camarosporium uniseriatum, C. elongatum and C. arezzoensis, are the sexual
morphs reported for the genus Camarosporium (Mirza
1968; Hyde et al. 2013; Tibpromma et al. 2016a; Thambugala et al. 2016). Camarosporium laburnicola differs
from C. elongatum and C. arezzoensis in having smaller
asci and ascospore with a different number of longitudinal
and transverse septa. Even though asci and ascospore sizes
overlap, C. laburnicola can be distinguished from C. uniseriatum which has 3–5 transverse septa and 1–2(–3) longitudinal septa; (Thambugala et al. 2016). However, there
are three coelomycetous Camarosporium species (i.e.
C. cytisi, C. laburni, C. laburnicum) reported from the
same host (Laburnum spp.) for which molecular data are
unavailable (Saccardo 1882, 1892).
Camarosporium moricola Chethana, Bulgakov & K.
D. Hyde, sp. nov.
Index Fungorum number: IF552502; Facesoffungi
number: FoF 2687, Fig. 71
Etymology: The specific epithet moricola is named after
the host genus Morus.
Holotype: MFLU 15-2075
Necrotrophic on dead and dying branches of Morus
alba L. Sexual morph Undetermined. Asexual morph
Conidiomata 150–340 lm (
x = 206 lm, n = 10) diam.,
pycnidial, solitary, scattered, immersed, unilocular, globose, black, with a papillate ostiolate. Pycnidial wall 25–
47 lm (-67 lm at apex), multi–layered, with outer layer
composed of 3–4 layers of thick, dark brown cells, with
inner 4–6 layers of hyaline cells of textura angularis.
Conidiophores reduced to conidiogenous cells. Conidiogenous cells 5–10 9 3–7 lm (
x = 7.6 9 5.3 lm,
n = 20), enteroblastic, phialidic, doliiform, hyaline,
smooth-walled, formed from the inner most layer of the
pycnidial wall, with an opening of 2.3 lm diam. Conidia
8–15 9 4–7.5 lm (
x = 11.1 9 5.4 lm, n = 30), oblong,
ellipsoid, straight or rarely slightly curved, initially hyaline, pale to dark brown at maturity, smooth-walled,
rounded at both ends, muriform, with 1–3 transverse septa
and 1–2 longitudinal septa.
Culture characteristics: Colonies on PDA, circular,
fimbriate, margin rough, with both surfaces greyish-white
at the margin and grey olivaceous towards the centre,
reaching 3 cm diam. after 7 days at 20 °C.
Material examined: RUSSIA, Rostov Region, Shakhty
City, Grushevka steppe slopes near Grushevsky pond, shrubbery, on dead and dying branches of Morus alba L. (Moraceae),
14 May 2015, T. S. Bulgakov (MFLU 15-2075, holotype), extype living culture, MFLUCC 16-1396, KUMCC 16-0132;
RUSSIA, Rostov Region, Krasnosulinsky District, Donskoye
forestry, artificial, forest edge, on dead and dying branches of
Morus alba L. (Moraceae), 18 June 2015, T. S. Bulgakov
(MFLU 15-2222), ex-type living culture, MFLUCC 16-1397,
KUMCC 16-0133; RUSSIA, Rostov region, Krasnosulinsky
District, Donskoye forestry, ravine forest edge, on dead and
dying branches of Morus alba L. (Moraceae), 18 June 2015, T.
S. Bulgakov (MFLU 15-2223, paratype), ex-type living culture, MFLUCC 16-1398, KUMCC 16-0134.
GenBank Numbers MFLUCC 16-1396 LSU:KY053890;
ITS:KY053887; SSU:KY053893. MFLUCC 161397
LSU:KY053891;
ITS:KY053888;
SSU:KY053894.
MFLUCC 16-1398 LSU:KY053892; ITS:KY053889;
SSU:KY053895.
Notes: Based on phylogenetic analyses and morphological comparison, our collection belongs to Camarosporium
sensu stricto, which is a distinct lineage in Pleosporinae,
Pleosporales. In our phylogenetic analyses of combined
LSU, SSU and ITS sequence data of Camarosporium sensu
stricto (Fig. 52), our collection of C. moricola clustered in a
well-defined monophyletic clade sister to C. robiniicola
(MFLUCC 13-0527) and C. aureum (MFLUCC 14-0620)
with relatively high bootstrap and Bayesian probabilities
(90% MP/97% ML/1.00 BYPP). Camarosporium moricolais distinct from C. robiniicola as the latter has obclavate
conidiophores and larger conidia (18–28 9 7–11 lm) with
4–6 transverse septa and 1 longitudinal septum (Wijayawardene et al. 2014b, c). Camarosporium moricola
differs from C. aureum in having smaller conidiomata
(400–550 9 350–550 lm), smaller conidia ((19–)–
19.5–22.5(–24) 9 8–9(–10) lm) with 4–5 transverse septa,
with 2–3 longitudinal septa (Liu et al. 2015). Camarosporium moricola also shows morphological similarities to its
sister taxa, C. clematidis and C. spartii. Our species is distinct from C. clematidis by having conidiomata (370–425 9
350–380 lm), enteroblastic, annellidic conidiogenous cells
and wider conidia 2–3 transverse and 1 longitudinal septa
(Wijayawardene et al. 2014c). They differ from C. spartii by
not having annelidic conidiogenous cells, larger conidia
(16–13 9 6–7 lm), 3–4 transverse septa and 2–3 longitudinal
septa (Wijayawardene et al. 2014b).
Eurotiomycetes O.E. Erikss. & Winka.
Eurotiomycetes was introduced by O.E. Erikss. &
Winka (1997). Eurotiomycetes is a monophyletic class
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Fungal Diversity
Fig. 71 Camarosporium moricola (MFLU 15-2075, holotype). a
Appearance of conidiomata on dead branch of Morus alba.
b Immersed conidiomata on the host surface. c Longitudinal section
of a conidioma. d Longitudinal section of a conidioma wall.
e Longitudinal section of a conidioma wall showing cell organization.
f Conidiogenous cells with developing conidia. g Conidia. h–
i Different septation patterns on conidia. j–k Upper view (j) and
the reverse view (k) of the colony on PDA. Scale bars a = 2 mm,
b = 0.5 mm, c = 50 lm, d, e = 20 lm, f, g = 10 lm, h, i = 5 lm
(Geiser et al. 2006) comprising three major clades,
Chaetothyriomycetidae, Eurotiomycetidae, and Mycocaliciomycetidae. Some are human pathogens, many of the
black yeasts, and some lichenized families (Kirk et al.
2008).
Eurotiales G.W. Martin ex Benny & Kimbr.
Eurotiales was introduced by G.W. Martin and validated
Benny & Kimbr in 1980. The order contains three families
(Elaphomycetaceae, Monascaceae and Trichocomaceae)
of well known and common fungi, particularly those with
123
Fungal Diversity
phialidic Aspergillus and Penicillium asexual stages (Kirk
et al. 2008).
Trichocomaceae E. Fisch.
Trichocomaceae was introduced by Fischer (1897).
Species of Trichocomaceae are economically important in
industry and medicine (Houbraken and Samson 2011).
Phylogenetic relationships are well known for only certain
genera of Trichocomaceae (Aspergillus, Penicillium and
Paecilomyces) (Peterson 2000a, b; Samson et al. 2004;
Peterson 2008; Samson et al. 2009), and the relationships
are still poorly studied (Houbraken and Samson 2011).
Section Exilicaulis
Penicillium Link.
Species of Penicillium (Trichocomaceae, Eurotiales) are
well known and abundant in soil, air, indoor environments and
in contaminated foods (Frisvad and Samson 2004; Samson et al.
2010). Their main function in nature is the decomposition of
organic materials and many are known for the postharvest
decay of fruits and as mycotoxin producers on various foods
(Frisvad and Samson 2004; Houbraken et al. 2016).
The genus Penicillium is divided into four subgenera—
Aspergilloides, Penicillium, Biverticillata and Furcatum
comprising 25 sections (Visagie et al. 2014). Section Exilicaulis was described by Pitt (1980), and is characterized by
monoverticillate conidiophores and typified by P. restrictum.
Recently, Houbraken and Samson (2011) revised the classification of the genus Penicillium based on phylogenetic data.
They redefined section Exilicaulis to include species with
biverticillate conidiophores, such as P. corylophilum,
P. melinii, and P. raciborskii. Recently, section Exilicaulis
was revised using three genes (BenA, CaM, and RPB2) to
comprise eight clades defined by the inclusion of the following
species: P. melinii, P. corylophilum, P. restrictum, P. citreonigrum, P. decumbens and P. parvum (Visagie et al. 2016).
As far as can be determined, only four species of
Penicillium. P. daejeonium, P. koreense, P. samsonianum
and P. jejuense, have been reported as new from Korea:
P. daejeonium from grape and schisanda fruit, P. koreense
from soil, P. samsonianum from stems and leaves of Viscum album var. coloratum, and P. jejuense from marine
environments of Jeju Island.
During an investigation of the fungi associated with
pomegranate fruit in Korea, a new species was isolated and
is described here based on morphological characteristics
and phylogenetic analyses (Fig. 72).
Penicillium punicae Hyang B. Lee, P.M. Kirk & T.T.T.
Nguyen, sp. nov.
MycoBank number: MB 818233; Facesoffungi number:
FoF 2797, Fig. 73
Etymology: punicae. Referring to pomegranate fruit
from which the species was first isolated.
Holotype: CNUFC-FP2-1
Colonies grow rapidly on MEA, reaching 35–37 mm
diam. at 25 °C in 7 days, green in the center with a white
margin; reverse white. Conidiophores mostly biverticillate,
with some triverticillate; stipes rough, 2–4.5 lm wide.
Metulae 2–4 per stipe. Vesicles subglobose to globose
when present. Phialides smooth ampulliform, 4–7 per
metula, 6–10.5 9 1.5–3 lm. Conidia slightly roughened,
globose, 2–3.5 lm 9 2–3.5 lm.
Material examined: REPUBLIC OF KOREA, Jeonnam
Province, Gwangju (35100 N 126550 E), from a fruit of Punica granatum (pomegranate) from a grocery store and
then stored in the fridge, 5 February 2016 (CNUFC-FP2-1,
preserved as glycerol stock at -80 °C in the Chonnam
National University Fungal Collection; isotype in Culture
Collection of National Institute of Biological Resources
(NIBR), Incheon); 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:12421).
GenBank Numbers CNUFC-FP2-1 RPB2:KX839675;
CaM:KX839671; b-tubulin:KX839673. CNUFC-FP2-2
RPB2:KX839670; CaM:KX839672; b-tubulin:KX839674.
Culture characteristics: The isolate was observed to
grow over a wide range of temperatures with varying
growth rates on MEA (malt extract agar), CYA (Czapek
yeast autolysate agar), and YES (yeast extract sucrose
agar). The average growth rates of CNUFC-FP2-1 on
MEA, CYA, and YES were 5, 3, and 2.5 mm per 24 h,
respectively. Optimal growth was observed around 25 °C,
slow growth was observed at below 20 °C, and restricted
growth at 37 °C.
Notes: Penicillium punicae is distinct from P. cravenianum and P. pagulum, growing rapidly when cultivated on
MEA and CYA; whereas P. cravenianum and P. pagulum
have restricted growth on MEA, and CYA, respectively. The
number of phialides per metula and metulae are fewer than in
P. cravenianum and P. pagulum. In the phylogenetic tree
based on multiple genes, the strain formed a separate branch
from other species of Penicillium sect. Exilicaulis and is
considered to represent a new species.
Exophiala J.W. Carmich.
Exophiala was introduced by Carmichael (1966) to
accommodate the type species E. salmonis from cerebral
lesions of Salmo clarkii. It is characterized by melanin or
melanin-like pigments in the cell walls, annelidic conidiogenesis and an abundance of budding cells (de Hoog and
Hermanides-Nijhof 1977; Haase et al. 1999). Exophiala
includes mostly black yeast asexual morphs belonging to
Chaetothyriales (Untereiner et al. 1995; de Hoog et al.
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Fungal Diversity
Fig. 72 Phylogenetic tree of Penicillium punicae CNUFC-FP2-1 and
CNUFC-FP2-2 and related species within the P. corylophilum clade
of the sect. Exilicaulis based on maximum likelihood analysis of
combined datasets for b-tubulin, CaM and RPB2. Sequence of
Penicillium restrictum CBS 367.48 was used as outgroup taxon.
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 and ex-type strains in bold
2011; Zeng et al. 2014), occasionally linked to peculiar
coelomycetes synanamorphs, viz. E. placitae Crous &
Summerell (Crous et al. 2007). The sexual morph of Exophiala has been reported as Capronia (Herpotrichiellaceae, Chaetothyriales) (Carmichael 1967; Hironaga et al.
1981; de Hoog et al. 2011). They are present in extreme
ecological niches causing animal and human diseases (Li
et al. 2008, 2009; Sudhadham et al. 2008; de Hoog et al.
2011; Najafzadeh et al. 2013; Wen et al. 2015), in soil, rock
surfaces, air, rhizosphere, and plant tissues (Addy et al.
2005; Julou et al. 2005; Bates et al. 2006; Neubert et al.
2006; Bukovská et al. 2010; de Hoog et al. 2011; Seyedmousavi et al. 2011; Ferrari et al. 2011; Isola et al. 2013)
(Fig. 74).
Etymology: refers to the country where the fungus was
collected.
Holotype: MFLU 15-2551
Saprobic on dead branch of Cytisus scoparius (L.) Link.
Sexual morph Ascomata 391–489 lm high 9 388–417
lm diam. (
x = 438 9 398 lm, n = 5), semi-immersed to
erumpent through host tissue, solitary, uniloculate, oval,
conspicuous at the surface, black, without papilla, ostiole
with reddish to brown setae in centre. Peridium 14–22 lm
wide, thin, comprising 6–7 layers of hyaline to pale
brown cell of textura angularis. Hamathecium of 1.6–2.4
lm wide, numerous, septate, branched, pseudoparaphyses.
Asci 62–100 9 8–12 lm (
x = 77 9 10 lm, n = 20), 8spored, bitunicate, fissitunicate, cylindric, with a clubshaped pedicel, thin-walled, round at the apex. Ascospores
13–22 9 3–7 lm (
x = 19 9 5 lm, n = 20), overlapping
1-seriate, fusiform, hyaline, 1-septate in the centre, apical
cell longer than basal cell, conical at the ends, guttulate,
smooth-walled, surrounded by large mucilaginous sheath.
Exophiala italica Tibpromma, Camporesi & K.D. Hyde,
sp. nov.
Index Fungorum number: IF552702; Facesoffungi
number: FoF 2798, Figs. 75, 76
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Fungal Diversity
Fig. 73 Penicillium punicae (CNUFC-FP2-1, holotype). a, d
Colonies in malt extract agar. b, e Colonies in yeast extract sucrose
agar. c, f Colonies in Czapek yeast autolysate agar. (a–c obverse
view, d–f: reverse view). (h–m, SEM). g Fruit rot inside of
pomegranate stored in fridge. h–l Verticillate conidiophore shapes
and phialides on metula. m Conidia. Scale bars h–j, l = 10 lm,
k = 5 lm, m = 2 lm
Asexual morph Mycelium consisting of smooth, branched, septate, pale brown, 1.2–3.5 lm wide hyphae,
forming hyphal strands. Conidiophores subcylindrical,
medium brown, smooth, consisting of a supporting cell
and a single conidiogenous cell, or reduced to a conidiogenous cell, straight to curved. Conidiogenous cells
123
Fungal Diversity
Fig. 74 RAxML tree based on a combined dataset of LSU, SSU, ITS
and TEF1 of Exophiala. Bootstrap support values for maximum
parsimony and maximum likelihood higher than 70% and Bayesian
posterior probabilities greater than 0.95 are defined above the nodes
respectively. The tree is rooted to Cyphellophora laciniata (CBS
190.61). Ex-type and ex-epitype strains are in bold. Newly generated
sequences are blue
10.6–26.5 9 1.2–2.7 lm (
x = 17.74 9 1.78 lm, n = 20),
phialidic, pale to medium brown, ellipsoid or subclavate,
smooth, with a somewhat flared collarette. Conidia 2.1–
5.2 9 1.7–3.4 lm (
x = 3.36 9 2.44 lm, n = 40),
1-celled, ellipsoid or oval, smooth-walled, guttulate,
hyaline, becoming pale with age, becoming aggregated in
chains.
Culture characteristics: Colonies reaching 9 cm diam.
after 1 week at 16 °C, colonies erumpent, spreading, with
sparse to dense aerial mycelium on water agar media with
sterile toothpicks and pine needles, thin to wide, smooth,
sporulation sparse.
Material examined: ITALY, near Passo la Calla, on
dead branch of Cytisus scoparius (Fabaceae), 25 September 2013, Erio Camporesi, IT1464 (MFLU 15-2551,
holotype); ex-type living culture MFLUCC 16-0245,
(HKAS 94561bis, paratype).
GenBank Numbers LSU:KY496723; ITS:KY496744;
SSU:KY501114; TEF1:KY514393.
Notes: We introduce a new species based on both
morphology and phylogeny. Exophiala italica was found in
Italy, where the sexual morph is characterized by semiimmersed to erumpent oval ascomata, ostiole, cylindric
asci, club-shaped pedicel and 2-celled hyaline ascospores,
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Fungal Diversity
Fig. 75 Exophiala italica (MFLU 15-2551, holotype). a Appearance
of ascomata on host substrate. b Section of ascoma. c Ostiole. d
Section of peridium. e Pseudoparaphyses. f, g Asci. h–l Ascospores.
m Germinating ascospore. Scale bars b = 100 lm, c = 50 lm,
d = 20 lm, e = 2 lm, f, g = 20 lm, h–m = 2 lm
constricted at the second septum. The species was mainly
identified based on phylogenetic analyses with high statistical support (96% in ML/1.00 in BYPP) and clustered
with E. hongkongensis. We compared the asexual morph of
E. italica and found that E. hongkongensis is different (see
Woo et al. 2013). E. italica is the first sexual morph report
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Fungal Diversity
Fig. 76 Exophiala italica (MFLUCC16-0245, ex-type living culture on WA). a Colony on WA and sterile pine needles. b Hyphae. c, e
Conidiophores. f, g Conidial. Scale bars a = 200 lm, b = 50 lm, c–e = 10 lm, f, g = 2 lm
123
Fungal Diversity
in Exophiala based on morphological characteristics and
molecular data (Fig. 74).
Mucoromycotina Benny
Mucorales Fr.
Mucorales has the largest number of species within the
Mucoromycota and is characterized by the formation of
coenocytic hyphae with septa either at the base of the
reproductive structures or irregularly distributed in old
colonies (Santiago et al. 2013). Species of this order are
mostly saprotrophic, and are commonly isolated from soil,
stored grains, plants, and animal excrement, especially that
of herbivores and rodents (Santiago et al. 2013; de Souza
et al. 2016). The genus Gongronella, proposed by Ribaldi
(1952), belongs to the phylum Mucoromycota, subphylum
Mucoromycotina, order Mucorales and family Cunninghamellaceae (Benny et al. 2016; Spatafora et al. 2016). It
includes species characterized by relatively slow growing
colonies that form poorly developed stolons and rhizoids,
as well as erect or circinately branched sporangiophores.
The sporangia are small with a deliquescing or breaking
wall, and the columellae are reduced in size, with the
presence of an apophysis meaning the sporangium never
has a pyriform shape. The sporangiospores are 1-celled,
hyaline, smooth and variously shaped, while the brown to
black zygospores are borne aerially, and have parallel
suspensors devoid of appendages (Hesseltine and Ellis
1964).
Cunninghamellaceae Naumov ex R.K. Benj.
Gongronella Ribaldi
Gongronella was established to accommodate a single
species: G. urceolifera Ribaldi (currently G. butleri
Lendn.), previously allocated within Mortierellaceae. The
presence of a distinct globose apophysis and the reduced
size of the columellae were the main characteristics in the
introduction of the genus (Upadhyay 1969). Peyronel and
Dal Vesco (1955) and Pici (1955) transferred Absidia
butleri Lendn. to Gongronella, based on the presence of a
characteristic apophysis similar to those observed in the
type species. Later, Hesseltine and Ellis (1961) proposed
the addition of another species, G. lacrispora Hesselt. &
J.J. Ellis, and recently, three new species have been
described (Adamčı́k et al. 2015; Ariyawansa et al. 2015c;
Li et al. 2016). Currently, the genus includes five species:
G. butleri, G. guangdongensis F. Liu, T.T. Liu & L. Cai,
G. koreana Hyang B. Lee & T.T.T. Nguyen, G. lacrispora
and G. orasabula Hyang B. Lee, K. Voigt, P.M. Kirk &
T.T.T. Nguyen. Species of Gongronella have traditionally
been distinguished based on their morphology and sporangial stages. In recent years, however, molecular analysis
has been used as an additional tool to delimitate species,
causing an increase in the number of new taxa (Adamčı́k
et al. 2015; Ariyawansa et al. 2015c; Li et al. 2016).
Gongronella brasiliensis C.A. de Souza, D.X. Lima &
A.L. Santiago, sp. nov.
Index Fungorum number: IF552552; Facesoffungi
number: FoF 2699, Fig. 78 Etymology: Brasiliensis,
referring to country where the species was first isolated.
Holotype: URM 7487
Colonies white, cottony, low and exhibiting slow growth
(6 cm diam. and 1–2 mm in height) after 7 days on MEA at
25 °C. Reverse cream to buff with irregular margins.
Rhizoids frequent, slightly branched, long or short, up to 45
lm length, hyaline. Stolons present, 2–6 lm diam., hyaline, generally coenocytic, although some septa can be
observed in old colonies. Odourless. Sporangiophores
erect, straight or slightly recumbent, smooth-walled, hyaline, 26.5–320 9 2.5–5 lm; solitary, arising from stolons,
or in whorls of two, often with a single branch, predominantly terminating in a sporangium, while those emerging
directly from aerial mycelia branch repeatedly. Some
branches end in a sterile sporangial, globose, 5–17 lm
diam. One or two septa may form below the apophysis.
Sporangia first yellowish then becoming light brownish,
globose, subglobose, 9.5–30 lm diam., smooth-walled,
some leaving collars. Columellae hyaline to light grey,
smooth-walled, globose, subglobose, (3–)4–8(–9) lm or
conical-cylindrical, 1.5–2.5 9 2–3 lm, some so small that
they are almost inconspicuous, up to 1 lm diam., some
with an evident collar. Apophysis hyaline, smooth-walled,
variable in shape, globose, (3–)4–5(–6) lm diam., vaseshaped, (3–)4 9 12(–14.5) lm and ellipsoidal, 5–10(–12)
9 3–7(–8.5) lm. Sporangiospores variable in shape, hyaline, smooth-walled, some containing oil droplets, reniform, 1.5–4 9 1.5–2.5 lm, ellipsoid to fusiform, 2–6.5 9
1.5–3 lm, ellipsoid with a flattened end, some almost
falciform, 2.5–7.5 9 1.5–4. Chlamydospores present in the
aerial mycelium, globose, subglobose and doliiform. Giant
cells globose, subglobose, ovoid, some hypha-like and
irregularly swollen, up to 48 lm diam. Zygosporangia not
observed.
Media and temperature tests: On MEA at 10 °C lack of
growth and sporulation. At 15 °C—colonies with slow
growth, reaching 3.6 cm in diam. after 192 h; poor
sporulation. At 20 °C—low (1–2 mm in 168 h) colonies
with slow growth (4.3 cm in 168 h); good sporulation.
Sporangiophores with mainly single branches, rarely multibranched; Chlamydospores present, globose to subglobose;
giant cells absent. At 25 °C—low (1–2 mm in 168 h)
colonies, better growth (6 cm in 168 h); excellent sporulation. At 30 °C—good growth 5.6 cm in 168 h; excellent
sporulation. The growth of G. brasiliensis on PDA was
slightly slower than on MEA at all tested temperatures.
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Fungal Diversity
Material examined: BRAZIL, Taquaritinga do Norte,
state of Pernambuco (7520 2800 S, 35590 2500 W), in soil samples. Soil, 5.XI.2015, leg. C.A.F de Souza (URM 7487,
holotype). Gene bank numbers ITS:KY114930;
KY114931.
Notes: According to the phylogenetic (ITS) and morphophysiological analyses, G. brasiliensis exhibits wellsupported genetic datasets and morphological characteristics that differentiate it from other species of the genus
(Fig. 78). Gongronella brasiliensis produces elliptical
apophysis, conical-cylindrical columellae and giant cells
not so far reported for this genus. G. brasiliensis may have
been previously confused with G. butleri because of the
branching pattern of the sporangiophores and for the size
and shape of the sporangia. However, only the former
produces elliptical apophysis, conical-cylindrical columellae and giant cells (Fig. 77). Additionally, colonies of
G. brasiliensis are persistently white, in contrast to the
colonies of G. butleri that are at first white, later becoming
olive-buff or olive-gray (Upadhyay 1969). In most species
of Gongronella, one septum is commonly observed below
the sporangia. The presence of two septa delimiting the
sporangia was only observed in G. orasabula and
G. brasiliensis. Additionally, the sporangiospores of
G. brasiliensis are variable in shape and larger than those
observed in G. butleri, G. guangdongensis, G. koreana and
G. orasabula (Figs. 79, 80).
Rhizopodaceae K. Schum.
Mucor Fresen.
The genus Mucor includes primarily saprotrophic species characterized by the production of fast-growing colonies, simple or branched sporangiophores without basal
rhizoids, non-apophysate sporangia, and seldomly produced zygospores which are borne from opposed suspensors, possess a thick pigmented and ornamented
zygosporangium (Schipper and Samson 1978). The species
within the genus can be easily isolated from soil, dung,
water, stored grains and other plant parts (Benny 2008; Li
et al. 2016). Mucor spp. are commonly reported in different
herbivore faeces (Thilagam et al. 2015; Li et al. 2016;
Nguyen et al. 2016). On these substrates, Zygomycetes
usually appear first, followed by ascomycete and finally
basidiomycete fungi. Among the Zygomycetes found in
dung, species of Mucor were dominant (Benny 2008). In
Korea, a small number of Mucorales have reported from
animal fecal (Li et al. 2016; Nguyen et al. 2016).
Traditional taxonomy of species of Mucor was based on
morphological characteristics such as the size and shape of
sporangia, as well as the mode of reproduction (sexual or
asexual). Recently, molecular data have been used to
evaluate mucoralean species (Hoffmann et al. 2013; Walther et al. 2013; Ariyawansa et al. 2015c).
123
Fig. 77 Phylogenetic tree of Gongronella constructed using the ITS
sequence data. Sequences are labeled with their database accession
numbers. Support values are from Bayesian inference and maximum
likelihood analyses. The new species are in blue
During a study on the Mucorales from animal faecal
samples in Korea, a species of Mucor that differed morphologically and phylogenetically from other species was
isolated and is thus described as new.
Mucor stercorarius Hyang B. Lee, P.M. Kirk, K. Voigt &
T.T.T. Nguyen, sp. nov.
MycoBank number: MB 818383; Facesoffungi number:
FoF 2799, Fig. 81
Etymology: stercorarius. Named for rat dung from which
the species was first collected.
Holotype: CNUFC-UK2-1
Colonies growing fast on SMA, white, grayish-white in
reverse, reaching 42–45 mm diam. at 20 °C after 2 days of
incubation. Sporangiophores 4.5–7 lm wide, erect, branched sympodially. Sporangia globose to subglobose, yellow, multi-spored, reaching 30–38.5 9 27.5–37 lm;
sporangial wall at maturity fully deliquescent, leaving a
small collar on the sporangiophore. Columellae of diverse
shape, globose to subglobose or pyriform, 15.5–22 9
Fungal Diversity
Fig. 78 Gongonella brasiliensis (URM 7487, holotype). a–c Sporangiophores with variously shaped apophysis and sporangia.
d Branched sporangiophore with apophysis and columella.
e Branched sporangiophore with two septa (arrows), sporangia and
a small columella with collarette. f Simple sporangiophore with
apophysis and columella. g Simple sporangiophore with apophysis
and inconspicuous columella with collarette (arrow). h Giant cells.
i Sporangiospores. Scale bars a, b, d = 25 lm, c, e–i = 20 lm
123
Fungal Diversity
Fig. 79 Phylogenetic tree of Mucor stercorarius CNUFC-UK2-1 and
CNUFC-UK2-2 based on maximum likelihood analysis of ITS
sequence data. Sequences of Syncephalastrum racemosum were used
as the outgroup taxon. Numbers at the nodes indicate the bootstrap
values ([50%) from 1000 replications. The bar indicates the number
of substitutions per position. The new species is in blue and ex-type
strains in bold
17–25 lm. Sporangiospores ellipsoidal, 6.5–10 9 3.5–6
lm. Zygospores 38–83 9 45–86 lm.
Material examined: REPUBLIC OF KOREA, Jeonnam
Province, garden of the Chonnam National University
located in Gwangju (35100 N 126550 E), from a rat fecal
sample, 24 February 2016 (CNUFC-UK2-1 preserved as
glycerol stock at -80 °C in the Chonnam National
University Fungal Collection (CNUFC); isotype in Culture
Collection of National Institute of Biological Resources
(NIBR), Incheon; 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:12422).
GenBank Numbers CNUFC-UK2-1 LSU:KX839685;
SSU:KX839684;
ITS:KX839689;
TEF1:KX839681;
Actin:KX839688.
CNUFC-UK2-2
LSU:KX839682;
SSU:KX839686; ITS:KX839680; TEF1:KX839687; Actin:
KX839683.
Culture characteristics: The isolate grew over a wide
range of temperatures with varying growth rates on SMA,
PDA (potato dextrose agar), and MEA (malt extract agar)
of 21.5, 17.5 and 23 mm per 24 h, respectively. Optimal
growth was 20–25 °C, slow growth at 5 °C, and no growth
at 37 °C.
Notes: Mucor stercorarius is similar in morphology and
closely related to M. genevensis but differs by forming
smaller sporangia. Columellae are diverse in shape and
smaller than those of M. genevensis; sometimes a, bellshaped apophysis was present. Moreover, the new species
grows and sporulates at 35 °C, whilst M. genevensis does
123
Fungal Diversity
Fig. 80 Phylogenetic tree of Mucor stercorarius CNUFC-UK2-1 and
CNUFC-UK2-2 and related species based on maximum likelihood
analysis of SSU and LSU, Actin TEF1 sequence data. Sequences of
Umbelopsis nana and U. isabellina were used as outgroup taxa.
Numbers at the nodes indicate the bootstrap values ([50%) from 1000
replications. The bar indicates the number of substitutions per
position. The new species is in blue and ex-type strains in bold
not. In the phylogenetic tree of ITS and multiple genes, the
strain forms a branch separate from other species of Mucor,
supporting its establishment as a new species.
fungicolous, and lichenized or lichenicolous taxa (Maharachchikumbura et al. 2016) (Figs. 79, 80).
Sordariomycetes O.E. Erikss. & Winka
Sordariomycetes is the second largest class of
Ascomycota (Hyde et al. 2013) including 6 subclasses, 32
orders, 105 families and 1331 genera (Maharachchikumbura et al. 2016). The Sordariomycetes have a cosmopolitan distribution comprising terrestrial and aquatic
taxa (Jones et al. 2015; Pratibha et al. 2014). Species are
characterized by perithecial ascomata and inoperculate
unitunicate asci (Zhang et al. 2006) or non-fissitunicate asci
(Kirk et al. 2008). The class includes many important plant
pathogens, endophytes, saprobes, epiphytes, coprophilous,
Diaporthales Nannf.
The Diaporthales was introduced by Nannfeldt (1932),
and includes a number of plant pathogenic fungi and many
asexually reproducing fungi (Castlebury et al. 2002;
Rossman et al. 2007).
Gnomoniaceae G. Winter
Gnomoniaceae was introduced by G. Winter (1886)
with Gnomonia Ces. & De Not. 1863 as the type genus.
Species in Gnomoniaceae can exist as saprobes as well as
cause serious tree diseases (Rossman et al. 2007). They are
characterized by ascomata that are generally immersed,
123
Fungal Diversity
Fig. 81 Mucor stercorarius (CNUFC-UK2-1, holotype). a Colony
on synthetic mucor agar. b Sporangium with sporangial wall net.
c Sporangium without wall net. d Spore bearing sporangium on
sporangiophore. e–j Columellae with clear collar present at the apex
of the sporangiophores. Scale bars b = 15 lm, c–j = 20 lm
solitary, without a stroma, or aggregated with a rudimentary stroma (Sogonov et al. (2008).The genera of Gnomoniaceae were listed by Sogonov et al. (2008) (Fig. 82).
immersed, solitary, or in small groups, uniloculate, globose, conspicuous at the surface, black, without hairs, long
ostiolate. Peridium 15–21 lm wide, comprising 3–4 layers
of pale brown cells of textura prismatica. Hamathecium
0.5–1.2 lm wide, numerous, comprising dense, septate,
unbranched, paraphyses. Asci 15–19 9 2–5 lm (
x = 17 9
4 lm, n = 25), 8-spored, bitunicate, fissitunicate, cylindric
to cylindric-clave, short pedicellate, thin-walled, round at
the apex. Ascospores 6–10 9 1–3 lm (
x = 9 9 2 lm,
n = 20), overlapping 1–2-seriate, hyaline, 1-septate, constricted at the septum, first cell longer than second cell,
round at the ends, guttulate, smooth-walled, lacking amucilaginous sheath. Asexual morph Undetermined.
Culture characteristics: Colonies on MEA, reaching 6
cm diam. after 7 days at 16 °C, producing dense mycelium,
circular, rough margin white–gray with raised on media
surface.
Material examined: ITALY, Premilcuore Province,
Castel dell’ Alpe, on dead stem of Sanguisorba minor
(Rosaceae), 22 August 2014, Erio Camporesi, IT2058
(MFLU 15-1255, HKAS 94604, reference specimen designated here); living culture MFLUCC 15-0062.
GenBank Numbers LSU:KY496735; ITS:KY496755;
SSU:KY501123; TEF1:KY514399.
Notes: Walker et al. (2010) chose a lectotype of
G. sanguisorbae but their illustration is not clear. However,
our phylogenetic tree shows that our strain groups with
Gnomoniopsis Stoneman
Gnomoniopsis Berl. was introduced based on the type
species G. chamaemori (Fr.) Berl. (Barr 1978; Monod
1983; Walker et al. 2010). Gnomoniopsis is characterized
by having small, black perithecia, solitary or in groups in a
minimal stroma, and 1-septate, oval to fusiform ascospores.
Sogonov et al. (2008) identified some Sirococcus species as
the asexual morph of Gnomoniopsis. Most species of
Gnomoniopsis occur on plants of the Fagaceae, Onagraceae and Rosaceae. Gnomoniopsis also contains economically important pathogens of rosaceous crop plants,
including blackberry, raspberry, and strawberry (Bolay
1971; Monod 1983; Maas 1998). An updated phylogeny of
Gnomoniopsis with new isolates based on a combined gene
analysis is presented (Fig. 83).
Gnomoniopsis sanguisorbae (Rehm) D.M. Walker,
Mycologia 102 (6): 1494 (2010)
Facesoffungi number: FoF 2800, Fig. 83
Reference specimen: MFLU 15-1255
Saprobic on dead stem of Sanguisorba minor Scop.
Sexual morph Ascomata 320–432 lm high 9 397–466 lm
diam. (
x = 388 9 442 lm, n = 5), immersed to semi-
123
Fungal Diversity
Fig. 82 Consensus tree resulting from a maximum likelihood
analysis of a combined TEF1 and ITS sequence alignment for taxa
of Gnomoniopsis. Species are indicated in coloured blocks. RAxML
bootstrap support values (ML above 50) and Bayesian posterior
probabilities (BYPP above 90%) are given at the nodes (ML/BYPP).
The scale bar represents the expected number of changes per site. The
tree is rooted to Gnomonia gnomon AFTOL ID 952. New sequences
are in blue bold
123
Fungal Diversity
Fig. 83 Gnomoniopsis
sanguisorbae (MFLU 15-1255,
reference specimen). a, b
Appearance of ascomata on host
substrate. c Section of ascoma.
d Ostiole. e Section of peridium.
f Pseudoparaphyses. g–i Ascus.
j–m Ascospores. n Germinating
ascospore. Note: stained with
cotton blue in i, j, l and m. Scale
bars c = 100 lm, d, e = 10
lm, f = 2 lm, g–i = 5 lm, j–
n = 2 lm
other strains of G. sanguisorbae. We also compared the
morphology of our strain with G. sanguisorbae and found
that it matches. We therefore designate G. sanguisorbae
(MFLU 15-1255) as a reference specimen (sensu Ariyawansa et al. 2014b).
123
Sydowiellaceae Lar.N. Vassiljeva
Sydowiellaceae was established in Diaporthales by
Lar.N. Vassiljeva (1987) based on S. fenestrans (Duby)
Petr. (Rossman et al. 2007, Senanayake et al. 2017). The
family accommodates 15 genera with 43 species, but
Fungal Diversity
several taxa do not share any clear morphological features
in common (Rossman et al. 2007; Liu et al. 2015;
Maharachchikumbura et al. 2015, 2016; Senanayake et al.
2017). Members of Sydowiellaceae are found on herbaceous, dicotyledonous plants and hardwood trees as
pathogens and saprobes (Rossman et al. 2007).
Sillia P. Karst.
Sillia was introduced by P. Karsten (1873), with the type
S. ferruginea (Pers.) P. Karst. There are six species listed in
Index Fungorum (2017) and Senanayake et al. (2017)
added a new species, S. karstenii. Sillia is characterized by
stromata with numerous ascomata with long cylindrical
necks and stromatic tissues, which turn to red in 10% KOH
(Iznova and Rukseniene 2012). A revised phylogeny
among members of the Diaporthales is presented in Fig. 84
Sillia italica de Silva, Camporesi & K.D. Hyde, sp. nov.
Index Fungorum number: IF552716; Facesoffungi
number: FoF 2801, Fig. 85
Etymology: Referring to the country where the fungus
was first collected.
Holotype: MFLU 16-0056
Saprobic on dead branch of Corylus sp. (Corylaceae).
Sexual morph Stroma erumpent from cracks, up to 1–2.5
mm in diam., black on the surface. Ascomata 480–745 lm
high 9 136–346 lm diam. (
x = 640 9 242 lm, n = 10),
semi-immersed in black stromatic tissues gregarious, globose to subglobose, ostiolate, soft and found in a valsoid
arrangement. In 10% KOH solution stroma becomes
brownish-red. Ostioles 330–570 lm long, cylindrical,
forming at the center of the ascomata, raised from the host
surface. Peridium 20–34 lm thick, with outer layer of
brown and thick-walled cells of textura prismatica; inner
layer of hyaline and thin-walled cells of textura angularis.
Paraphyses 4–7 lm wide, cylindrical, cellular, septate,
mixed with asci. Asci 147–234 9 10–15 lm (
x = 190 9 13
lm, n = 25), 8-spored, unitunicate, narrowly clavate,
straight or somewhat curved, with a short obtuse pedicel.
Ascospores 99–128 9 3.1–4.6 lm (
x = 109 9 3 lm,
n = 30), fasciculate, hyaline, filiform, elongate, narrow
and curved at the ends, guttulate, smooth-walled, with 3
transverse septa. Asexual morph Undetermined.
Material examined: ITALY, Forlı̀-Cesena Province,
Monte Mirabello–Predappio, on dead twig of Corylus sp.
(Corylaceae), 11 January 2016, Erio Camporesi, IT2754
(MFLU 16-0056 holotype, BBH isotype).
GenBank Numbers LSU:KY397949; ITS:KY523484.
Notes: Sillia italica strains analyzed herein are sister
taxa to S. ferruginea, S. karstenii and this relationship is
well-supported (Fig. 84). This close phylogenetic affinity is
also supported by morphological similarities. Both S. ferruginea and S. italica become reddish in 10% KOH
solution (Iznova and Rukseniene 2012). Sillia italica
however, has some distinct characters as compared to
S. ferruginea. Stromatic tissues of S. ferruginea comprise
black tissues on the surface and inner yellow stromatic
tissues mixed with black tissues (Iznova and Rukseniene
2012), while S. italica contains black stromatic tissues. The
ascomata arrangement of S. italica is valsoid, whereas in
S. ferruginea it is diatrypoid. Sillia italica has clavate asci
similar to S. ferruginea, but they are longer (147–234 lm
vs. 96–115 lm). Ascospores of S. italica are 99–128 lm
long with 3 septa, while in S. ferruginea they are 65–72 lm
long with 3–5 septa. Senanayake et al. (2017) introduced
Sillia karstenii which morphologically differs from S. italica in having large ascomata with very poorly developed
stromata. The phylogenetic analysis of Senanayake et al.
(2017) also showed that S. italica is phylogenetically distinct from other Sillia species (Fig. 86).
Cytospora Ehrenb.
Cytospora Ehrenb., Sylv. mycol. berol. (Berlin): 2
(1818)
Cytospora was introduced by Ehrenberg (1818) and is
an important pathogenic genus causing canker disease on
branches leading to dieback on a wide range of plants.
(Adams et al. 2005, 2006; Hyde et al. 2016; Norphanphoun
et al. 2017). See Norphanphoun et al. (2017) for a recent
account of the genus. An updated tree for the genus is
provided in Fig. 87.
Cytospora gelida Norphanphoun, Bulgakov, T.C. Wen &
K.D. Hyde, sp. nov.
Index Fungorum number: IF552712; Facesoffungi
number: FoF 2802, Fig. 88
Etymology: the Latin epithet ‘‘gelida’’ meaning colonies
on media look like ice.
Holotype: MFLU 15-3779, Fig. 87
Pathogen on twigs and branches of Cotinus coggygria
Scop. Sexual morph Undetermined. Asexual morph
Conidiomata 650–1000 lm diam. 9 350–450 lm high.
immersed in host tissue, solitary, scattered, erumpent,
unilocular, with ostiolate. Ostiole 300–400 lm diam. at the
same level as the disc surface. Peridium comprising several
layers of cell of textura angularis, with inner most layer
thick, brown, outer layer dark brown to black. Conidiophores branched, reduced to conidiogenous cells. Conidiogenous cells branched, blastic, enteroblastic, phialidic,
hyaline, smooth. Conidia (5.3–)5.7–8 9 1.4–1.8(–2) lm
(
x = 6.9 9 1.8 lm, n = 30) 1-celled, allantoid to subcylindrical, hyaline, smooth-walled.
Culture characteristics: Colonies on MEA, reaching 4
cm diam. after 7 days at 25 °C, producing dense mycelium,
circular, rough margin white mycelium, without aerial
mycelium: reverse of the colony whitish-yellow
123
Fungal Diversity
Fig. 84 Consensus tree resulting from a maximum likelihood
analysis of combined LSU and ITS sequence alignment of genera
of Diaporthales. Genera in Sydowiellaceae are indicated in blue and
ash blocks. Maximum parsimony bootstrap support (MPB above
50%), RAxML bootstrap support (MLB above 50%), and Bayesian
123
posterior probabilities (PP above 0.9) are given at the nodes (MPB/
MLB/PP). The scale bar represents the expected number of changes
per site. The tree is rooted to Gnomonia gnomon (CBS 829.79; CBS
199.53). Ex-type strains are in bold and new strains are in blue bold
Fungal Diversity
Fig. 85 Sillia italica (MFLU 16-0056, holotype). a Appearance of
ascomata erumpent through cracks in bark on branch of Corylus sp. b,
c Vertical sections of stromata and ascomata. d Peridium. e–h Asci.
i Pseudoparaphyses. i–n Ascospores. Scale bars b, c = 80 lm,
d = 30 lm, e–h = 80 lm, i = 20 lm, j–n = 80 lm
Material examined: RUSSIA, Rostov Region, Krasnosulinsky District, Donskoye forestry, lining-out nursery,
trees and shrubs, dying twigs and branches of Cotinus
coggygria (Anacardiaceae), 27 October 2015, T. Bulgakov; T-1117 (MFLU 15-3779, holotype), ex-type living
culture, MFLUCC 16-0634, KUMCC.
123
Fungal Diversity
Fig. 86 Phylogram generated from one of 1000 most parsimonious
trees based on combined ITS, LSU, RPB2 and Actin sequence data
from Cytospora isolates. The tree is rooted to Phomopsis vaccinii
(ATCC 18451). Maximum likelihood bootstrap values C60% are
given at the nodes. The species obtained in this study are in blue and
ex-type are in blue bold. Ex-type taxa are in bold
Notes: In the phylogenetic analyses (Fig. 87), C. gelida groups with C. cotini (strain MFLUCC 14-1050) and
C. ribis (strain 284-2). Cytospora gelida can be distinguished from C. cotini by its multi-loculate ([4) conidiostromata, while C. cotini has 2–3 locules in the
conidiostromata. The conidia of C. gelida are larger than
C. cotini (6.9 9 1.8 vs. 5.9 9 1.2 lm). Cytospora gelida
differs from C. cotini in the polymorphic nucleotides of
ITS, RPB2 and Actin sequence data. In ITS it differs
with two polymorphisms,
polymorphisms.
123
and
RPB2
in
17
Cytospora ceratosperma (Tode) G.C. Adams & Rossman,
in Rossman, Adams, Cannon, Castlebury & Crous, IMA
Fungus 6: 147 (2015)
Facesoffungi number: FoF 2803, Fig. 88
Pathogen on twigs and branches of Acer platanoides L.
Sexual morph: Ascomata 300–450 lm high 9 500–550
Fungal Diversity
Fig. 86 continued
lm diam. (
x = 300 9 500 lm, n = 5), immersed in host
tissue, scattered, erumpent through the surface of bark,
lenticular, extending to a large circular area, with 3–6ostioles per group. Locules dark brown, arranged circularly, flask-shaped to sphaerical with one ostiole. Ostiole
450–550 lm diam. numerous, dark brown to black, at the
same level as the disc, occasionally area below disc a
lighter entostroma. Peridium composing several layers of
irregular cells arranged in a textura angularis, with inner
most layer thick, dark brown, outer layer dark brown to
black. Hamathecium long cylindrical, cellular, anastomosed, paraphyses. Asci (58–)60–65 9 10–11(–12) lm
123
Fungal Diversity
Fig. 87 Cytospora gelida (MFLU 15-3779, 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 Ostiole.
f Peridium. g, h, i Conidiophores with conidiogenous cells. j Conidia.
123
k, l Culture characters on MEA. Scale bars a = 2 mm, b = 1 mm,
c = 500 lm, d = 200 lm, e = 100 lm, f = 50 lm, g = 20 lm, h,
i = 5 lm, j = 10 lm
Fungal Diversity
Fig. 88 Cytospora ceratosperma (MFLU 15-3757). a Appearance of
fruiting bodies in wood. b Fruiting bodies on substrate. c Close up of
fruiting body. d Cross section of the ascomata. e Peridium. f Ostiole.
g, h, i Asci. j Ascus strain in Melzer’s reagent. k Paraphyses. l Apical
ring. m, n, o, p Ascospores. q, r Culture characters on MEA. Scale
bars a = 2 mm, b = 1 mm, c, d = 500 lm, e = 50 lm, f = 100 lm,
g–j = 10 lm, k = 50 lm, l–p = 5 lm
123
Fungal Diversity
(
x = 63 9 11 lm, n = 15), 6–8-spored, unitunicate, clavate to elongate obovoid, with apical ring J-, short pedicel
late or sessile. Ascospores (11–)12.2–15 9 3.1–4(–4.2) lm
(
x = 14.5 9 3.7 lm, n = 20), biseriate, elongate-allantoid,
1-celled hyaline, lacking a mucilaginous sheath, smooth
walled. Asexual morph: Undetermined.
Culture characteristics: Colonies on MEA, reaching 5
cm diam. after 7 days at 25 °C, producing dense mycelium,
circular, rough margin white, without aerial mycelium.
Material examined: RUSSIA, Rostov Region, Krasnosulinsky District, Donskoye forestry, Kabanya Balka (Boar
gully), artificial forest, dying twigs and branches (necrotrophic) on Acer platanoides (Sapindaceae), 27 October
2015, T. Bulgakov; T-1095 (MFLU 15-3757, new record);
living culture, MFLUCC 16-0625, KUMCC.
Notes: Cytospora ceratosperma has been reported on
hosts worldwide (Ahmad 1969; Pantidou 1973; Simonyan
1981; Truszkowska and Chlebicki 1983; Cannon et al.
1985; Spielman 1985; Ahmad et al. 1997; Dudka et al.
2004; Mulenko et al. 2008; Fotouhifar et al. 2010; Rossman et al. 2015). The characters of the sexual morph in our
collection indicate it belongs to Cytospora in having a large
circular area, with 3–6-ostioles per group, with asci
(
x = 63 9 11 lm) and ascospores (
x = 14.5 9 3.7 lm). In
the phylogenetic analyses of our collection (strain
MFLUCC 16-0625), groups with C. rosarum (strain 218),
collected from Rosa canina (Fotouhifar et al. 2010)
(Fig. 86). The current study represents the first record of
maple canker caused by C. ceratosperma in Russia.
Cytospora ceratosperma has not been epitypified and
therefore we treat MFLUCC 16-0625 as a reference specimen (Ariyawansa et al. 2014b). Fresh collections of
C. ceratosperma from the type location and host are needed
for epitypification.
Hypocreales Lindau
Bionectriaceae, Clavicipitaceae, Cordycipitaceae,
Flammocladiaceae, Hypocreaceae, Nectriaceae, Niessliaceae, Ophiocordycipitaceae, Stachybotryaceae and Tilachlidiaceae are included in the order Hypocreales
(Maharachchikumbura et al. 2015). Hypocreales are highly
diverse and found in the tropics and subtropics (Põldmaa
2011). They are characterized by pigment producing,
brightly coloured perithecial ascomata, and typically ostiolate perithecial fruiting body (Rogerson 1970; Rehner and
Samuels 1995).
Clavicipitaceae Earle
The families Clavicipitaceae, Cordycipitaceae and
Ophiocordycipitaceae were recognized by Sung et al.
(2007) based on phylogenetic analyses. The family
Clavicipitaceae (Hypocreales) is a very heterogeneous
group comprising obligate saprotrophes, parasites and
123
symbionts with insects and, fungi or grasses, rushes or
sedges (Sung et al. 2007; Schardl et al. 2013; Kepler et al.
2013; Li et al. 2016). Maharachchikumbura et al. (2015)
listed 48 genera under this family.
Species of Hypocrella, Moelleriella and Aschersonia
belong to the family Clavicipitaceae and are pathogens of
scale insects and whiteflies and are common in tropical
regions (Chaverri et al. 2008). Most species of Hypocrella
produce an Aschersonia asexual morph that is characterized by pycnidia or acervuli on a stroma, often filled with
brightly coloured, slimy conidia (White et al. 2003; Chaverri et al. 2005). Moelleriella was described by Bresadola
(1896) and it produces filiform, multi-septate ascospores,
that disarticulate at the septa within the ascus and an
aschersonia-like asexual morph with fusoid conidia (Chaverri et al. 2008) (Fig. 89).
Hypocrella calendulina Hywel-Jones & Mongkols., Mycol
Res 113 (6–7): 687 (2009)
Facesoffungi number: FoF 2805, Fig. 90
Parasitic on scale insect nymph (Hemiptera), forming
yellow to brownish stromata on the underside of bamboo
leaves. Sexual morph See Mongkolsamrit et al. 2009.
Asexual morph Coelomycetous. Conidiomata 1.4–2.2 9
0. 45–0.55 mm (
x = 1.784 9 0.493 mm, n = 10), reddishbrown, solitary, irregular in shape, surface roughened,
flattened pulvinate, multi-loculate, locules 520–638 9
385–455 lm (
x = 579 9 420 lm, n = 20), brown,
immersed, globose or subglobose to obpyriform, ostiolate.
Ostioles subcylindrical, single, well-developed, centrally
located. Wall of conidiomata 19–29 lm wide, composed of
relatively thick-walled, brown cells of textura oblita in the
outer layers and thin-walled, hyaline cells of textura
angularis in the inner layers. Paraphyses 163–308 9 1.2–
1.6 lm (
x = 236 9 1.4 lm, n = 150), hyaline, filiform,
tapering at the apices, arising from the hymenium of the
conidioma. Conidiophores reduced to conidiogenous cells.
Conidiogenous cells 8–14 9 1–1.5 lm (
x = 11 9 1.2 lm,
n = 90), hyaline, phialidic, cylindrical to subcylindrical,
broad at base, discrete, smooth, arising from the inner
layers of conidioma. Conidia 11–13 9 2.1–2.9 lm (
x = 12
9 2.5 lm, n = 150), hyaline, fusiform, unicellular, straight
or slightly curved, guttulate, thick-and smooth-walled,
accumulating in a slimy yellowish mass above the
conidioma.
Culture characteristics: Cultures were obtained from
germinating ascospores and conidia. The ascospores and
conidia germinated within 48 h on PDA. Colonies on PDA
were irregular and approx. 2 cm diam. after 4 weeks at
20 °C. The colonies derived from germinating ascospores
or conidia formed a compact mycelium. The colonies have
rough surface, viscous texture, knobly protuberances and
curled margin. The conidial mass was pale yellow to
Fungal Diversity
Fig. 89 Phylogram of
Hypocrella, Samuelsia,
Moelleriella and Aschersonia
generated from maximum
likelihood analysis of RPB1,
RPB2 and TEF1 sequence data.
Shimizuomyces paradoxus
Kobayasi is used as outgroup
taxon. Maximum likelihood
bootstrap values greater than
50% and Bayesian posterior
probabilities over 0.5 are
indicated above the nodes. The
new species and epitype species
are indicated in blue
yellow appearing as abundant slimy masses scattered over
the surface of stromatic colonies.
Material examined: THAILAND, Chiang Rai Province,
on insect nymph (Hemiptera) on bamboo leaf, 10 February
2015, Jingzu Sun & Yuanpin Xiao, MRC 15021007
(MFLU 16-2918, asexual morph report) MFLU 16-2919,
MFLU 16-2919, MFLU 16-2920, MFLU 16-2921, living
culture, MFLUCC 17-0057, MFLUCC 17-0058, MFLUCC
17-0059, MFLUCC 17-0060. GenBank Numbers
RPB1:KY646196; RPB2:KY646197; TEF1:KY646198.
Notes: The sexual and asexual morphs of Hypocrella
calendulina were introduced by Mongkolsamrit et al.
(2009) and is a widespread species in Thailand, which can
be found throughout the year (Mongkolsamrit et al. 2009).
In this study, for the first time we provide, illustrations of
the asexual morph of this species, which includes wall of
conidiomata, conidiogenous cells and conidia.
Moelleriella thanathonensis Y.P. Xiao, T.C. Wen & K.D.
Hyde, sp nov.
Index Fungorum number: IF552794; Facesoffungi
number: FoF 2804, Fig. 91
Etymology: The species epithet refers to Headquarter of
Thanathon orchard, the location of the holotype collection.
Holotype: MFLU 16-2922
Parasitic on an unidentified insect, forming yellow to
yellowish conidiomata on the underside of leaves. Sexual
morph Undetermined. Asexual morph Coelomycetous,
Conidiomata 1.6–1.7 9 0.7–0.9 mm (
x = 1.65 9 0.8 mm,
n = 10), yellow to orange, solitary or gregarious, with
pycnidium-like depression in the stroma, lacking well-defined walls, irregular in shape, surface pruinose, flattened
pulvinate, base slightly constricted, surrounded by a white
hyphae, up to 3 mm wide, multi-locular, with locules, 309–
443 9 234–342 lm (
x = 376 9 288 lm, n = 20), globose
to subglobose, thick-walled., surface minutely tomentose,
covered with confluent conidial masses that are deep yellow to orange yellow. Wall of conidiomata 25–58 lm wide,
thin, composed of thick-walled, brown to hyaline cells.
Paraphyses 43–61 9 0.9–1.5 lm (
x = 52 9 1.2 lm,
n = 90), arising from the hymenium of the conidioma,
filiform, tapering at the apices. Conidiophores reduced to
conidiogenous cells. Conidiogenous cells 13–18 9 1–1.4
lm (
x = 15.6 9 1.2 lm, n = 60), hyaline, phialidic,
123
Fungal Diversity
Fig. 90 Hypocrella calendulina (MFLU 16-2918, asexual morph).
a Habitat. b Appearance of brownish conidiomata on the host. c, d
Vertical section of conidiomata. e, g, h Base of conidioma with
conidiophores, conidiogenous cells and developing conidia. f Paraphyses arising from the hymenium of the conidioma. i–n Immature to
123
mature conidia. o Germinating conidium. p, q Culture on PDA
medium notes q reverse. Scale bars b = 1000 lm, c = 200 lm,
d = 500 lm, e, g = 20 lm, f = 100 lm, h = 50 lm, i–o = 5 lm, p–
q = 1 cm
Fungal Diversity
Fig. 91 Moelleriella thanathonensis (MFLU 16-2922, holotype).
a Habitat. b Appearance of yellow to yellowish conidiomata. c, d
Vertical section of conidiomata. e–g Base of conidioma with
conidiogenous cells, conidiophores and developing conidia. h–n
Immature to mature conidia. o–r Culture on PDA medium, notes p r
reverse. Scale bars b, c = 1000 lm, d = 500 lm, e = 50 lm, f–
g = 20 lm, h–n = 5 lm, o–p = 5 cm, q–r = 1 cm
123
Fungal Diversity
cylindrical, undetermined, discrete, thick and smoothwalled. Conidia 11–13.5 9 1.6–2.1 lm (
x = 13 9 1.9 lm,
n = 150), hyaline, fusiform, small, 1-celled, often brightly
yellow in mass.
Culture characteristics: Cultures were obtained from
germinating conidia. The conidia germinated within 24 h
on PDA. The colonies were 2 cm diam. after 4 week at
25 °C. The colonies were circular, umbonate, with entire
margin and cream to pale yellow conidial masses formed in
two circular areas on the stromatic colonies.
Material examined: THAILAND, Chiang Rai Province,
Headquarter of Thanathon orchard, on an unidentified
insect, Yuanpin Xiao, 10 February 2015, MRC 15021007,
MRC 15021007 (MFLU 16-2922, holotype); (MFLU
16-2923, paratypes), ex-type living cultures, MFLUCC
17-0057, MFLUCC 17-0058, MFLUCC 17-0059,
MFLUCC 17-0060).
GenBank
Numbers
PBP1:KY646195;
RPB2:KY646199; TEF1:KY646200.
Notes: The sexual morph of this species was not found
in the field although several attempts were made to find it
throughout the year. Moelleriella thanathonensis is similar
to three widespread species in Thailand, namely Moelleriella raciborskii (Zimm.) P. Chaverri, M. Liu & K.T.
Hodge, M. libera (Syd. & P. Syd.) P. Chaverri & M. Liu
and Aschersonia placenta Berk. Aschersonia placenta is
the asexual morph of Moelleriella raciborskii (syn.
Hypocrella raciborskii Zimm.) (Liu et al. 2009). Moelleriella libera is the sexual morph of Aschersonia aleyrodis
Webber (Chaverri et al. 2008). Our new species M.
thanathonensis is characterized by multi-locular conidiomata, fusiform conidia and phialidic conidiogenous cells.
In this paper, we provide a synopsis of morphologically
similar asexual morph species to M. thanathonensis in
Table 3. Phylogenetic analysis of combined TEF1, RPB1
and RPB2 sequence data (Fig. 89) also provides evidence
that M. thanathonensis is a new species.
Stachybotryaceae L. Lombard & Crous
Stachybotryaceae was introduced with its type genus
Stachybotrys (Corda 1837) and is typified by S. chartarum
(Ehrenb.) S. Hughes (Wang et al. 2015). Presently, 33
genera are accepted in the family Stachybotryaceae
(Lombard et al. 2016; Lin et al. 2016) (Fig. 92).
Myrothecium Tode
Myrothecium was introduced by Tode (1790) with
M. inundatum Tode as the type species. Species of this
genus are saprobes in the soil and plant pathogens (Ellis
and Ellis 1985; Watanabe 1994; Chen et al. 2016). More
than 30 species of this genus have been recorded worldwide (Seifertk et al. 2011; Chen et al. 2016; Lombard et al.
2016).
Myrothecium septentrionale J.F. Li, Phookamsak & K.D.
Hyde, sp. nov.
Index Fungorum number: IF 552310; Facesoffungi
number: FoF02482, Fig. 93
Etymology: ‘‘septentrionale’’ means ‘‘northern’’ in Latin,
named after the location where the species collected,
northern Thailand.
Holotype: MFLU 16-2889
Saprobic on dead hanging leaves of Pandanus sp. Sexual
morph Undetermined. Asexual morph Colonies effuse on
host, grey. Mycelium superficial on the substrate, composed
of aseptate, unbranched, smooth, subhyaline hyphae. Conidiophores 8–11 lm long 9 1–1.6 lm diam. (
x = 9.8 9 1.2
lm, n = 20), macronematous, mononematous, solitary,
erect, scattered, straight to curved, subhyaline to hyaline,
smooth, thin-walled, aseptate, unbranched, arising from a
stromatic base. Conidiogenous cells 1–1.5 lm long 9 1–1.6
lm diam. (
x = 1.3 9 1.3 lm, n = 20), phialidic, terminal,
globose to ellipsoid, hyaline, smooth and thin-walled.
Conidia 8.6–12 lm long 9 1.2–2.1 lm diam. (
x = 10.5 9
1.6 lm, n = 20) solitary, acrogenous, hyaline, smoothwalled, 0–1-septate, ampulliform to doliiform, chiefly subcylindrical. Conidial secession rhexolytic.
Cultural characteristics: Conidia germinating on PDA
within 14 h and germ tubes produced from the apex.
Colonies growing on PDA, reaching 5 cm in 14 days at
30 °C, mycelium partly superficial, partly immersed,
slightly effuse, powdery, vertical, with regular edge, maroon to yellowish-red; sexual or asexual spores not formed
within 60 days.
Table 3 Synopsis of morphologically similar asexual morph species to Moelleriella thanathonensis
Species
Conidiomata (mm)
Conidiogenous
cells (lm)
Conidia (lm)
Paraphyses (lm)
References
Aschersonia placenta (asexual morph
of Moelleriella raciborskii)
Aschersonia aleyrodis (asexual morph
of Moelleriella libera)
1–3 9 0.2–0.7
7–22 9 1–1.5
9–16 9 1.5–2
40–70
Liu et al. (2006)
1–1.5
23–40 9 0.9–1.5
9–14 9 0.9–1.9
65–100
Saccardo (1899)
1.6–1.7 9 0.7–0.9
13–18 9 1–1.4
11–13.5 9 1.6–2.1
43–61 9 0.9–1.5
This study
Moelleriella thanathonensis
123
Fungal Diversity
Fig. 92 Phylogenetic construction using RAxML analysis of the
combined LSU and SSU, dataset. Bootstrap support values for
maximum likelihood (black) equal to or greater than 50% and
Bayesian posterior probabilities (red) equal to or greater than 0.95 are
shown above the nodes. The tree is rooted to Valsonectria pulchella
and Scopinella solani. The type strains are in black bold and the new
species is in blue bold
Material examined: THAILAND, Chiang Mai Province,
Mae Taeng, dead hanging leaves of Pandanus sp. (Pandanaceae), 25 March 2016, Junfu Li, H-MRC 41B (MFLU
16-2889, holotype), ex-type living culture MFLUCC
16-2819, KUMCC; (isotype in HKAS).
GenBank Numbers KY559395.
123
Fungal Diversity
Fig. 93 Myrothecium septentrionale (MFLU 16-2889, holotype). a Colonies on dead leaf of Pandanus sp. b–g Conidiophores with
conidiogenous cells. h–o Conidia. Scale bars a = 100 lm, b–o = 1 lm
Notes: Myrothecium septentrionale is similar to M. chiangmaiense D.Q. Dai & K.D. Hyde in morphology.
However, M. septentrionale and M. chiangmaiense are
phylogenetically distinct and the latter is basal to M cinctum and M. gramineum, while M. septentrionale is closely
related to M. roridum with high support (Fig. 92). Myrothecium septentrionale differs from M. roridum in having larger ampulliform conidia (8.6–12 9 1.2–2.1 lm vs.
5–6 9 1–1.2 lm), while M. roridum has cylindrical
conidia.
123
Glomerellaceae Locq. ex Seifert & W. Gams
Glomerellaceae is a monotypic family which mainly
comprises plant pathogens. The family was invalidly
published by Locquin (1984) and was validated in Zhang
et al. (2006). This family was accepted as one of the four
families of Glomerellales in Maharachchikumbura et al.
(2015). This family is characterized by the Colletotrichum
asexual morph and the Glomerella sexual morphs, which
are now named as Colletotrichum (Hyde et al. 2014;
Maharachchikumbura et al. 2015).
Fungal Diversity
Colletotrichum Corda.
This genus was introduced by Corda (1831), for C. lineola Corda (Damm et al. 2009). Colletotrichum species
are mainly plant pathogens, but also endophytes and saprobes (Yang et al. 2009; Hyde et al. 2009; Cannon et al.
2012; Jayawardena et al. 2016a, b). Kirk et al.
(2001, 2008), Réblová et al. (2011) referred Colletotrichum
to the family Glomerellaceae and Maharachchikumbura
et al. (2016) further confirmed the familial affinities of this
genus (Fig. 94).
Colletotrichum sambucicola Jayawardena, Camporesi &
K.D. Hyde, sp. nov.
Index Fungorum number: IF552553; Facesoffungi
number: FoF 2683, Fig. 95
Holotype: MFLU 16-2675
Saprobic on dead branch of Sambucus ebulus L. Sexual
morph Undetermined. Asexual morph Conidiomata 0.13–
0.4 mm (
x = 0.27 mm, n = 10) diam., black, acervulus,
oval, solitary, gregarious. Setae straight or ± bent, abundant, dark brown, becoming paler towards the apex, opaque, smooth walled, septate, 1–6-septate, 110–145 lm
long, base cylindrical, slightly inflated, 6.1–8.5 lm diam.,
apex acute to rounded, smooth. Conidiophores simple, to
20 lm long, hyaline to pale brown, smooth-walled. Conidiogenous cells 9–16 9 2–4 lm (
x = 11.9 9 2.9 lm,
n = 20), phialidic, hyaline, smooth-walled, cyllindrical to
slighty inflated, opening 1–2 lm wide, collarette or periclinal thickening observed. Conidia 12.2–29.7 9 3–5 lm
(
x = 17.7 9 3.8 lm, n = 15), L/W ratio 4.7, hyaline,
smooth or verruculose, aseptate, curved, both sides
Fig. 94 Phylogram generated
from maximum parsimony
analysis based on combined
ITS, GADPH, CHS, Actin and
TUB2 sequenced data from
species of the dematium
complex. Maximum parsimony
bootstrap support values greater
than 75% and Bayesian
posterior probabilities greater
than 0.80 are shown above the
branches. The ex-type strains
are in bold and the new species
is in blue. The tree is rooted
with Colletotrichum nigrum
123
Fungal Diversity
Fig. 95 Colletotrichum sambucicola (MFLU 16-2675, holotype).
a Appearance of conidiomata on host. b Base of setae. c Tip of setae.
d Setae with conidiophores. e, f Conidiogenous cells. g, h Conidia.
123
i Upper view of 7 days old culture. j reverse view of 7 days old
culture. Scale bars a = 0.2 mm, b = 20 lm, d = 10 lm scale bar of
b applies to c, e and scale bar of d applies to f, g, h
Fungal Diversity
gradually tapering towards the round to slightly acute apex
and truncate base, guttulate. Appressoria not observed.
Culture characteristics: Colonies on PDA flat with
entire margin, aerial mycelium sparse, short, pale olivaceous-grey, colony surface buff, some sectors dark greyolivaceous to dark-olivaceous-grey, iron-grey acervuli can
be observed mainly on the edge of the colony. Reverse
olivaceous green, concentric rings can be clearly observed,
reaching 40–55 mm in 7 days at 25 °C.
Material examined: ITALY, Province of Forlı̀-Cesena
[FC], Meldola, on dead branch of Sambucus ebulus
(Adoxaceae), 30 March 2016, Erio Camporesi IT 2902
(MFLU 16-2675, holotype); ex-type living culture,
MFLUCC 16-1388, KUMCC 16-0127.
GenBank Numbers ITS:KY098781, KY595193;
GAPDH: KY098780, KY595192; CHS:KY098779,
KY595190; Actin: KY098778, KY595190; TUB2:
KY098782, KY595194 respectively.
Notes: Colletotrichum dematium species complex is
mainly characterized by species with curved conidia
(Damm et al. 2009; Jayawardena et al. 2016a, b). Colletotrichum sambucicola falls within the C. dematium
species complex clade and forms a separate branch with
100% bootstrap support and 1.00 Bayesian posterior
probability and is a sister taxon to C. anthrisci (Fig. 95).
Among the species with angular conidia, conidia of C. anthrisci have the highest L/W ratio (7.8) and the apex is
strongly pointed. Colletotrichum sambucicola differs from
C. anthrisci in having conidiogenous cells with a distinct
collarette, as well as by having smaller conidia. A BLASTn
search of NCBI GenBank with the ITS sequence of
MFLUCC 16-1388 showed 99% similarity to quite a
number of Colletotrichum species with angular conidia.
The closet match in a BLASTn search in GenBank with the
GAPDH sequence of MFLUCC 16-1388 was GenBank
GU228237 (98% identity, three bp differences) and CHS
GU228335 (98% identity 4–7 bp differences). The newly
described taxon differs from its sister taxon C. anthrisci in
six bp in ITS, eight bp in GAPDH, four bp in CHS, 39 bp in
Actin and four bp in TUB2 (Fig. 96).
Xylariales Nannf.
Xylariales was introduced by Nannf. 1932 and this order
is the largest order of perithecial ascomycetes. Familial
relationships in this order have been investigated (e.g. Tang
et al. 2009) and currently 22 families are accepted
Maharachchikumbura et al. (2016).
Bartaliniaceae Wijayaw. et al.
Based on morphology and DNA sequence data, Senanayake et al. (2015) introduced the family Bartaliniaceae
to accomodate the genera Bartalinia, Broomella,
Dyrithiopsis, Hyalotiella, Truncatella and Zetiasplozna.
Later Wijayawardene et al. (2016) included the genera
Doliomyces Steyaert and Morinia Berl. & Bres. Doliomyces is known to be morphologically similar to the
other genera of the family but there is no sequence data yet
to confirm this placement (Wijayawardene et al. 2016).
Morinia Berl. & Bres. has uniquely distinct morphological
features but phylogenetically groups within Bartaliniaceae
(Wijayawardene et al. 2016). Bartaliniaceae is phylogenetically related to the families Sporocadaceae, Pestalotiopsidaceae and Robillardaceae (Jeewon et al.
2002, 2003a; Senanayake et al. 2015; Maharachchikumbura et al. 2016).
Truncatella Steyaert
The genus Truncatella, typified by T. truncata was
established by Steyaert (1949) to accommodate distinct
species having 3-septate, verruculose, pigmented conidia
(Steyaert 1949; Jeewon et al. 2002; Maharachchikumbura
et al. 2012, 2015). The genus was previously placed in
Amphisphaeriaceae, Xylariales (Jeewon et al. 2003a;
Lumbsch and Huhndorf 2010), and later included in Bartaliniaceae by Senanayake et al. (2015) based on both
morphological features and molecular evidence. Several
studies have contributed to the phylogenetic understanding
of Truncatella, such as its affinity to Bartalinia (Jeewon
et al. 2002; Lee et al. 2006; Maharachchikumbura et al.
2012, 2014a, b, 2016, 2017; Senanayake et al. 2015;
Maharachchikumbura et al. 2017) and Neotruncatella
(Hyde et al. 2016). The sexual morph of Truncatella
remains largely unresolved, however Senanayake et al.
(2015) illustrated both the sexual and asexual morph for
T. spartii.
Truncatella spartii Senan., Camporesi & K.D. Hyde,
Fungal Diversity 73: 91 (2015) Facesoffungi number: FoF
00685, Fig. 97
Saprobic on cone of Piceae sp., appearing as black,
irregular lesions on the host surface. Sexual morph See
Senanayake et al. 2015. Asexual morph Conidiomata
(240–)320–370(–433) lm diam., 160–200 lm, (
x = 340 9
180 lm, n = 8), pycnidial or acervular, semi-immersed to
superficial, becoming erumpent at maturity, globose with a
flattened base, solitary to aggregated, black. Conidiomata
wall consisting of outer light brown to hyaline cells of
textura angularis and inner hyaline cells of textura angularis to textura globulosa. Conidiophores 2.5–4 lm long,
hyaline, cylindrical, simple or branched. Conidiogenous
cells 8.5–14.2 lm long hyaline, simple, filiform, annelidic,
indeterminate, integrated. Conidia 17.5–24.8 9 5.3–7.2 lm
(
x = 21.5 9 6.5 lm, n = 30), broadly fusiform, 3-septate,
slightly constricted at the mid septum, verruculose, median
cells brown to yellowish-brown, 10.5–15 lm long, thickwalled, globules present, basal cell hyaline, conical
123
Fungal Diversity
Fig. 96 Phylogram generated
from maximum likelihood
analysis based on ITS sequence
data from species of
Bartaliniaceae. Maximum
likelihood bootstrap support
values higher than 70% and
Bayesian prosterior
probabilities greater than 0.96
are indicated above or below the
branches. The new isolate is
shown in blue bold and ex-type
strains in black bold. The tree is
rooted to Pseudopestalotiopsis
theae
tapering towards the base with an acute apex, apical cell
hyaline, bearing 3 appendages. Appendages 10–13.5 lm
long, hyaline, simple, filiform, similar in length or shorter
than conidia.
Culture characteristics: On PDA, slow growing,
attaining 15 mm in 7 days at 18 °C, circular, margin serrate, from above white, reverse greyish-white.
123
Material examined: ITALY, Province of Forlı̀-Cesena
[FC], near Camposonaldo—Santa Sofia, on dead cone
attached to Picea excelsa (Pinaceae), 6 May 2013, Camporesi Erio, IT1226 (HKAS96302, new host record), living culture KUMCC 15-0127.
GenBank Numbers ITS:KY 474383.
Fungal Diversity
Fig. 97 Truncatella spartii (KUMCC 15-0127). a Appearance of
conidiomata on cone of Picea sp. b Close-up of conidiomata. c, d
Cross section of conidioma. e Conidioma wall. f–i Developing
conidia from conidiogenous cells. j–m Conidia. n Germinating
spores. Scale bars c, d = 50 lm, e–i, n = 10 lm, j–m = 20 lm
Notes: This is the first record of Truncatella sparti on
Picea excelsa. This species was introduced by Senanayake et al. (2015) from the host Spartium with both
sexual and asexual morphs illustrated. Truncatella spartii
is closely related to T. hartigii (Tubeuf) Steyaert, T. helichrysi (Severini) B. Sutton and T. restionacearum S.
Lee & Crous, but differs in the apical cell
characteristics of its conidia (Lee et al. 2006; Senanayake et al. 2015).
Myrmecridiales Crous
Myrmecridium Arzanlou, W. Gams & Crous.
The genus Myrmecridium was described by Arzanlou
et al. (2007) with the type species M. schulzeri. To date, 10
123
Fungal Diversity
species of Myrmecridium have been described (Peintner
et al. 2016). Species belonging to this genus are characterized by the production of obovoid or fusiform conidia,
tapering towards a narrowly truncate base, hyaline mycelium with pale to unpigmented, and pimple-like denticles.
They are frequently isolated from soils and plant tissues
(Crous et al. 2011; Jiea et al. 2013; Peintner et al. 2016).
Some species possess potent antimicrobial activity (Zhang
et al. 2012a). Molecular data have been used to evaluate
species of Myrmecridium (Crous et al. 2015a; Réblová
et al. 2016; Peintner et al. 2016). In a previous study a new
species, Myrmecridium fluviae, was reported from a
freshwater sample from Yeongsan River in Gwangju,
Korea (Hyde et al. 2016).
While examining the diversity of fungi 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
(Fig. 98).
Myrmecridium fluviae Hyang B. Lee & T.T.T. Nguyen, sp.
nov.
MycoBank number: MB 818334; Facesoffungi number:
FoF 2806, Fig. 99
Fig. 98 Phylogenetic tree of
Myrmecridium fluviae CNUFCYR61-1 and CNUFC-YR61-2
and related species based on
maximum likelihood analysis of
a ITS, b LSU sequences.
Sequences of Veronaeopsis
simplex were used as outgroup.
Numbers at the nodes indicate
the maximum likelihood
bootstrap values ([50%) from
1000 replications. The bar
indicates the number of
substitutions per position. New
taxa are in blue and ex-type
strains in bold
123
Etymology: fluviae. Referring to fresh water from which
the species was first isolated.
Holotype: CNUFC-YR61-1
Sexual morph Undetermined. Asexual morph Colonies
on OA growing very slowly, grey olivaceous, reaching
15–17 mm diam. at 25 °C after 7 days incubation; reverse
olivaceous black with a white margin. Conidiophores
arising vertically from aerial hyphae, straight to irregular,
branched, septate, varying greatly in length, 3–4.5 lm
diam. Conidiogenous cells cylindrical, holoblastic, polyblastic, forming a rachis with scattered pimple-shaped
denticles. Conidia produced laterally or apically, obovoid
to fusoid, wall smooth to slightly rough, 7.5–12 9 3–4 lm.
On PDA growth is faster than on OA and MEA, greenisholivaceous, reverse olivaceous-buff; on MEA, surface
greyish-sepia.
Material examined: REPUBLIC OF KOREA, Jeonnam
Province, Yeongsan River located in Gwangju (35100 N
126550 E), from a freshwater sample, 15 February 2016
(CNUFC-YR61-1, preserved as glycerol stock at -80 °C
in the Chonnam National University Fungal Collection;
isotype in Culture Collection of Nakdonggang National
Institute of Biological Resources (NNIBR), Sangju,
Gyeongbuk Province); living culture (ex-type) deposited at
Fungal Diversity
Fig. 99 Myrmecridium fluviae
(CNUFC-YR61-1, 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–q light microscope, r–
v SEM). g–q Conidia and
conidiophores (i, j, s, t lateral
production of spore on
conidiophore and k–n, r apical
production of conidia on
conidiophore). v Rachis with
scattered, pimple-shaped
denticles (yellow arrow). Scale
bars g = 50 lm, h–q = 20 lm,
r–v = 3 lm
Jena Microbial Resource Collection (University of Jena
and Leibniz Institute for Natural Product Research and
Infection Biology, Jena, Germany) (JMRC:SF:12423).
GenBank Numbers CNUFC-YR61-1 LSU:KX839677;
ITS:KX839678.
CNUFC-YR61-2
LSU:KX839676;
ITS:KX839679.
123
Fungal Diversity
Culture characteristics: The isolate grew over a wide
range of temperatures with varying growth rates on PDA,
MEA and OA with average growth rates of 2.8, 2.5, and 2.2
mm per 24 h, respectively, for CNUFC-YR61-1. Optimal
growth around 25 °C to 27 °C, slow growth at 10 °C, and
no growth at 35 °C.
Notes: Myrmecridium fluviae is morphologically distinct
from other species, especially from M. phragmitis and
M. flexuosum by having larger conidia with smooth to
slightly rough-walls, sometimes forming a spiny layer. Our
phylogenies provide sufficient evidence to justify the new
species, albeit not strong, possibly due to inadequate taxon
sampling and sequence data from other genes.
Xylariaceae Tul. & C. Tul.
The family Xylariaceae was introduced by Tulasne and
Tulasne (1963) using the term ‘‘Xylariei’’ and the rank of
the name was not certain as it did not address the family
concept (Stadler et al. 2013). Even though the exact
number of currently accepted taxa varies, the recent outline
of Xylariaceae (Maharachchikumbura et al. 2015) accepted
87 genera. The segregation of Xylariaceae into the subfamilies Hypoxyloideae and Xylarioideae have been supported by molecular data (Tang et al. 2009; Stadler et al.
2013; Daranagama et al. 2015; Maharachchikumbura et al.
2015). The xylariaceae are mainly saprobes, while several
species can also be found as pathogens and endophytes.
Rosellinia De Not.
Rosellinia was introduced by De Notaris (1844) with
R. aquila (Fr.) Ces. & De Not. as the type species.
According to the world monograph by Petrini (2013) only
142 species are accepted, of which 37 species are described
as new species (Fig. 100).
Rosellinia mearnsii Tennakoon, Phookamsak & K.D.
Hyde, sp. nov.
Index Fungorum number: IF552779; Facesoffungi
number: FoF 2718, Fig. 101
Etymology: Name reflects the host Acacia mearnsii,
from which the holotype was collected.
Holotype: MFLU 16-1382
Saprobic on dead stem of Acacia mearnsii De Wild.
Sexual morph Stromata globose, with a conical pointed
top, 700–750 lm high 9 550–580 lm diam. (
x = 735 9
565 lm, n = 10), black, shiny, solitary, aggregated into
small groups, carbonaceous, flattened base, surrounded by
grey to pale brown, woolly to felty subiculum, confined to
the stromatal base, ostiolate. Ostioles black, distinctively
papillate. Ascomata, globose, uni-loculate, 550–625 lm
high 9 390–485 lm diam. (
x = 576 9 445 lm, n = 20).
Peridium black, thick-walled, with several compressed cell
layers, carbonaceous. Ascomata 600–650 9 550–580 lm
123
(
x = 635 9 565 lm, n = 10), nearly semiglobose, surface
convex, dark, smooth, with a central black papilla and
ostiole. Peridium black, thick walled, compressed and
carbonaceous. Paraphyses at base (3.6–)3.8–4.6(–4.9) lm
wide (
x = 4.2 lm), distinct septate, slightly longer than
asci, filamentous, unconstructed and hyaline. Asci (198.2–)
202–224(–232.5) 9 (9.6–)11.5–14.2(–15.5) lm (
x = 215
9 12.5lm, n = 20) 8-spored, bearing part cylindrical,
short pedicellate, with barrel-shaped, subapical, J ? ring,
15.6–15.9 lm high, 7.9–8.4 lm diam., apically rounded
with well-developed ocular chamber. Ascospores (44.7–)
46.2–53.4(–56.3) 9 (7.9–)8.2–8.8(–9.1) lm (
x = 49.3 9
8.6 lm, n = 20), overlapping, inequilaterally fusiform,
uni-seriate, initially hyaline to slight brown, becoming
brown colour, aseptate, straight to curved, smooth walled
with prominent guttules. Asexual morph Undetermined.
Material examined: CHINA, Yunnan Province,
Xishuangbanna, Nabanhe, dead branch of Acacia mearnsii
(Mimosaceae), 20 November 2015, D.S. Tennakoon, DXH
21 (MFLU 16-1382, holotype; HKAS 93712, isotype).
GenBank Numbers LSU:KY514060; ITS:KY514059;
RPB2:KY514061.
Notes: Rosellinia mearnsii resembles R. lamprostoma in
having asymmetrically ellipsoidal to slightly fusoid, light
brown ascospores (Petrini 2013). However, R. mearnsii
differs from R. lamprostoma in its globose ascomata and
prominent guttules in the ascospores, whereas stromata in
R. lamprostoma are conical, pear shaped and ascospores
with inconspicuous guttules (Petrini 2013). The size ranges
of ascomata also differ (550–625 lm high 9 390–485 lm
diam., in R. mearnsii and 700–1050 lm high 9 625–1250
lm diam., in R. lamprostoma). Rosellinia mearnsii is also
reminiscent to R. capetribulensis, which has fusiform,
brown ascospores and a barrel-shaped apical ring (Bahl
et al. 2005). However, in R. capetribulensis, the asci are
clavate as compared to the cylindrical asci in R. mearnsii.
Although the ascospores in R. mearnsii and R. capetribulensis different in length, the spores of R. capetribulensis
are 103–155 9 15–22.5 lm, whereas those in R. mearnsii
are 46.2–53.4 9 8.2–8.8 lm. Furthermore, Rosellinia
mearnsii resembles R. chiangmaiensis in having ellipsoidal
to slightly fusoid, brown ascospores, but R. chiangmaiensis
can easily be distinguished from R. mearnsii in having an
inverted hat-shaped apical ring and ascospores with a thin
mucilaginous sheath (Li et al. 2016).
Phylogenetic analyses of combined ITS, LSU and
RPB2 sequence data show that R. mearnsii clusters with
R. lamprostoma with strong support (100% ML, 100%
MP, 1.00 BYPP, Fig. 101). Besides the morphological
differences, R. mearnsii differs from R. lamprostoma in
having six ITS base pair and seven RPB2 base pair
differences.
Sordarimycetes genera, incertae sedis
Fungal Diversity
Fig. 100 Consensus tree resulting from a maximum likelihood
analysis of a combined ITS, RPB2 and LSU sequence alignment for
taxa in Rosellinia and other selected isolates of Xylariaceae. Type
species are indicated in black bold and the new species is in blue bold.
Maximum Parsimony bootstrap support values (MP above 50%),
RAxML bootstrap support values (ML above 50) and Bayesian
posterior probabilities (BYPP above 0.9) are given at the nodes (MP/
ML/BYPP). The scale bar represents the expected number of changes
per site. The tree is rooted to Lopadostoma turgidum CBS 133207
Evlachovaea B.A. Borisov & Tarasov
The genus Evlachovaea was established by Borisov and
Tarasov (1999) and since then this genus is a monotypic
with E. kintrischica Borisov and Tarasov as the type
species. Only one species (Evlachovaea kintrischica B.A.
Borisov & Tarasov 1999) has so far been reported from this
genus (Index Fungorum 2017) (Figs. 102, 103).
Ascomycota families incertae sedis
123
Fungal Diversity
Fig. 101 Rosellinia mearnsii (MFLU 16-1382, holotype). a Ascomata on host. b Close-up of ascomata. c section of ascoma.
d Pseudoparaphyses. e–h Asci. i Ascus stain with Indian ink. j–q Ascospores. Scale bars d–i = 50 lm, j–q = 10 lm
Evlachovaea indica P.N. Singh, A. Baghela, S.K. Singh &
S. Amir, sp. nov.
MycoBank number: MB 816184; Facesoffungi number:
FoF 2807, Figs. 104, 105
Etymology: specific epithet ‘indica’ refers to the country
of origin.
Holotype: AMH 9760
On live Mantispa sp. in terrestrial habitats. Asexual
morph Conidiophores mononematous, unbranched to
branched, sometimes apical region extends to form long
whip like structure producing solitary and short conidiogenous cells unilaterally or verticillately, smooth-walled,
hyaline, 25–80 9 1.5–2.5 lm (
x = 47 9 3.29 lm, n = 30).
123
Conidiogenous cells phialides in whorls of 2–8, or more,
occasionally solitary on conidiophores or arising from a
vegetative hypha, with slightly inflated and narrowed
towards base, smooth-walled, hyaline, tapering with long
slender neck, 8–40 9 1–2.39 lm (
x = 16.24 9 1.49 lm,
n = 30). Conidia produced in long basipetal chains,
forming a zip-like pattern, oval or subglobose, broadly
ellipsoidal to clavate, smooth-walled, hyaline, 1.75–9 9 1–
3 lm (
x = 6.40 9 2.07 lm, n = 30). Sexual morph
Undetermined.
Culture characteristics: on semi-synthetic agar mediumsucrose (HAM–S) reaching 5.5 cm diam. in 14 days at
25 °C. After two weeks of incubation with irregular
Fungal Diversity
Fig. 102 Phylogram generated
from maximum likelihood
method based on ITS
sequences: The tree with the
highest log likelihood (2225.7653) is shown. Onethousand bootstrap replicates
were analyzed to obtain nodal
support values. The novel
species is shown in blue. The
tree is rooted with Paecilomyces
carneus, P. lilacinus,
P. marquandii and Metarhizium
anisopliae. Phylogenetic
analyses were conducted in
MEGA6 (Tamura et al. 2013)
margin, smooth surface, cottony, puffy, umbonate to pulvinate, white (1A1-2) towards margin, pinkish-white at
centre (7A2), rosy vinaceous with age, raised, sometimes
reaching about 1 cm high, and touching upper lid of Petri
plates. Reverse light yellow to pale orange (4A4–5A3)
towards centre, yellowish-white (4A2) towards margin,
wrinkled; hyphae septate, unbranched to branched, pigmented, smooth and thin walled 2.5–4.35 lm wide.
Material examined: INDIA, Maharashtra, Pune District,
on live Mantispa sp. (Mantispidae) 30 August 2012, P.N.
Singh; (AMH 9760 holotype-dried colony on PDA); extype living culture, NFCCI 3970, National Fungal culture
Collection of India-WDCM 932);
GenBank Numbers ITS:KU983517; TEF1:KU983516.
Notes: Colony characters and growth rates of E. indica
vary when grown on HAM medium supplemented with
different carbon sources, viz., arabinose, galactose, glucose, glycerol, fructose, lactose, maltose and sucrose
(Fig. 106). Arabinose favored maximum radial growth of
colony (56–60 mm) on pre-defined incubation conditions
123
Fungal Diversity
Fig. 103 Phylogram generated from maximum likelihood method
based on TEF1 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 (-3415.2087) is shown. One-thousand bootstrap replicates
were analyzed to obtain nodal support values. The novel species is
shown in blue. The tree is rooted with Paecilomyces lilacinus,
P. marquandii and Metarhizium robertsii
of 25 °C, and for 14 days, followed by galactose (55–56
mm), lactose (54–56 mm), sucrose (52–55 mm). Colony
characters and growth pattern of E. indica resembles colony characters of E. kintrischica, but the growth rate was
faster in E. indica compared to E. kintrischica on most of
the carbon sources used in present study. In addition,
E. indica and E. kintrischica expressed very prominent
wrinkles on reverse sides of the colony on glucose, glycerol, and sucrose. Evlachovaea indica expressed less
prominent wrinkles on maltose compared to distinctly
prominent in E. kintrischica. Both the species did not
produce wrinkles on galactose and lactose. Besides,
E. indica produced no wrinkles on fructose, while it has
been reported prominent in E. kintrischica.
Evlachovaea indica is morphologically distinct in having smaller and slender conidiophores, and significantly
larger conidia from the type species, E. kintrichica, and
from Evlachovaea-like isolates group I and II (Humber
et al. 2013) in having longer conidia and conidiogenous
cells. However, length–width ratio of conidia (L/W
1.8–3.4) and conidiogenous cells (L/W 4.8–32.9) in
E. indica is significantly greater as compared to L/W ratio
of conidia (L/W 1.3–2.5) and conidiogenous cells (L/W
2–7) of E. kintrichica. Proposed species is also distinct
from type species in lacking synnemata and triangular or
pomegranate like conidia (Borisov and Tarasov 1999).
The internal transcribed spacer (ITS)-rDNA and translation elongation factor (TEF1) regions have been shown to
be taxonomically and phylogenetically informative for
various entomopathogenic fungi such as Beauveria,
Cordyceps, Isaria, Metarhizium and Paecilomyces (Liu
et al. 2002; Luangsa-ard et al. 2005). These two gene
sequences were also found to be useful in unambiguously
placing the proposed isolate in a unique place in the phylogenetic trees. Though, it shows some level of relatedness
with Evlachovaea-like isolates group I, is still distinct from
123
Fungal Diversity
Fig. 104 Evlachovaea indica (AMH 9760, holotype) on PDA.
a Fungal growth appeared near prothorax (arrow). b Colony
morphology (front view). c Colony morphology (reverse view).
d Basipetal chain of conidia on PDA showing zipper pattern.
e enlarged view of cylindrical and elongated conidia. f Conidiophores
producing phialides in whorls with intact conidia. g Enlarged view of
phialides and intact conidia. h Branched conidiophore bearing conidia
in basipetal chains. i Apical region of conidiophore extends to form
long whip like structure producing unilateral short phialides (arrow).
j Chlamydospores in chain. Scale bars d–j = 10 lm
Evlachovaea-like isolates group I, II, and Evlachovaea
kintrischica. The phylogenetic analysis clearly establishes
Evlachovaea indica as a novel species. Humber et al.
(2013), utilized above mentioned genes to place the evlachovaea-like Brazilian isolates to Cordycipitaceae. However, they could not determine the correct species
identifications, possibly due to the unavailability of TEF1
and other gene sequences of species of Isaria and other
allied taxa. This indicates that a comprehensive analysis of
more of genes, in addition to, ITS and TEF1 and their
phylogenetic analysis needs to be undertaken to resolve the
phylogenetic ambiguity of this group of fungi. Until the
123
Fungal Diversity
Fig. 105 Colony characteristics (front and reverse view) of Evlachovaea indica (AMH 9760, holotype) on different sugars. a, b Arabinose. c,
d Fructose. e, f Galactose. g, h Glucose. i, j Glycerol. k, l Lactose. m, n Maltose. o, p Sucrose
phylogenetic ambiguity is resolved, the present isolate is
placed as a novel species under the genus Evlachovaea
Borsov and Tarasov. To our knowledge, this fungus was
isolated for the first time from a four-winged live Mantispa
sp. collected from Western Ghats region in India. Moreover, this genus is being reported for the first time from
India.
Pezizales J. Schröt.
Helvellaceae Fr.
Helvella L. 1753
123
Species of Helvella L. are widely distributed in the
northern hemisphere (e.g. Hyde et al. 2016; Zhao et al.
2015, 2016a, b, c). Since Linnaeus (1753) proposed the
generic name Helvella, numerous studies have been conducted to address the taxonomic issue and infrageneric
division in Asia, Europe and North America (e.g. Dissing
1966; Weber 1972; Häffner et al. 1987; Abbott and Currah
1997; Hyde et al. 2016; Zhao et al. 2015, 2016a, b, c).
Helvella costifera Nannf., Schriften der Naturforschenden
Gesellschaft zu Leipzig 1: 114 (1953)
Fungal Diversity
Fig. 106 Phylogenetic tree
inferred from most likelihood
(ML) analysis using 28S data
(-ln L = -3407.754468).
Thickened branches represent
Bayesian posterior probabilities
(PP) above 95% are given at the
nodes; the new species are in
blue. The tree is rooted with
Wynnella subalpina
123
Fungal Diversity
Fig. 107 Helvella costifera (HKAS 87704, reference specimen). a,
b Typical mature specimens (a HKAS 69817. b HKAS 87759, HKAS
69817, HKAS 78900, HKAS78902). c Receptacle surface of pileus.
d Stipitipellis. e Asci and paraphyses. f–i Asci, j Ascospores. Scale
bars a, b = 1 cm. c, d = 50 lm. e–j = 20 lm
Facesoffungi number: FoF03127, Fig. 107
Reference specimen: HKAS 87704
Symbiotic in the coniferous, mixed or deciduous forests.
Sexual morph Pileus shallowly cupulate to cupulate,
margin expanding at maturity, 0.5–2 cm high, 0.5–4 cm
broad, hymenium even, pale grey to greyish-brown when
fresh, dark brown to blackish-brown when dried, receptacle surface above concolorous with hymenium, lower part
white to cream, pubescent, with branched, sharp or rarely
blunt ribs, ribs reaching from the half to the pileus edge.
Stipe 0.5–6 cm long, 0.2–2 cm diam., expanding and
merging with apothecium, below equal or base rarely
slightly enlarged, costate or lacunose, sharp-edged ribs,
white to light yellow brown, pubescent. Medullary excipulum 350–550 lm broad, of textura intricata, hyaline,
composed of 3–5 lm diam., hyaline, walls thickened, J?.
Ectal excipulum 80–110 lm broad, of textura angularis,
yellowish-brown, terminal cells 15–20 9 7–12 lm, blue in
cotton blue, J-. Stipitipellis 80–100 lm, of textura angularis, yellowish-brown, terminal cells 16–25 9 9–15 lm,
123
Fungal Diversity
blue in cotton blue, J-. Asci pleurorhynchous, 8-spored,
subcylindrical to clavate, with apex rounded, 230–300 9
12–16 lm, J- in Melzer’s reagent, blue in cotton blue.
Paraphyses filiform, 3–4 lm diam., slightly exceeding the
asci, apex enlarged, 6–7 lm diam., brown, J-. Ascospores
[60/2/2, in H2O] 15–19 9 10–13(–13.5) lm
[Q = 1.20–1.64, Q = 1.39 ± 0.08)], broadly ellipsoid to
ellipsoid, smooth-walled under the light microscope,
uniguttulate. Asexual morph Undetermined.
Habitat: Scattered or gregarious on soil in coniferous
forests.
Distribution: Europe (Dissing 1966; Häffner et al.
1987); North America (Abbott and Currah 1997).
Material examined: CHINA, Yunnan Province, Panlong
County, under Pinus spp., alt. 2000 m, 10 July, 2014, Qi
Zhao 1643 (HKAS 87704, reference specimen designed
here), Qi Zhao 1645 (HKAS 87706).
Notes: Helvella costifera has a greyish hymenium,
pubescent receptacle surface with blunt edged ribs and a
non-lacunose stipe, pleurorhynchous asci, broadly ellipsoid
ascospores 15–19 9 10–13(–13.5) lm, slightly enlarged
paraphyses (6–7 lm), and ectal excipulum J? in Melzer’s
reagent. Helvella costifera can be confused with H. acetabulum, which also have cupulate pileus and ribbed stipe
(Dissing 1966; Häffner et al. 1987; Abbott and Currah
1997). However, H. acetabulum is distinguished by its
brownish cup-shaped pileus and ribbed or lacunose stipe,
with ribs reaching to the half or to the receptacle surface
edge (Abbott and Currah 1997).
Helvella crispoides Q. Zhao & K.D. Hyde, sp. nov.
Index Fungorum number: IF 551935; Facesoffungi
number: FoF 01338, Fig. 108
Etymology: crispoides is proposed because of the eastern Asian mushroom’s similarity to Helvella crispa.
Holotype: HKAS 90535
Sexual morph Pileus irregularly lobed to saddleshaped, up to 4 cm high, 1–3 cm broad; margin inflexed to
receptacle surface when young, expanded at maturity; hymenium undulate-rugose, cream, becoming brownish-yellow when dried; receptacle surface pubescent, with blunt
ribs covering almost 1/3 of the pileus, grey, becoming
greyish when dried. Stipe 7–10 cm long, 1–2 cm thick,
deeply ribbed, with cross-veins and pockets, finely pubescent, slightly brittle, white, becoming cream when dried;
basal mycelium white. Medullary excipulum 280–450 lm
broad, of textura intricata, hyaline, composed of 2.5–4 lm
hyphae broad, J-. Ectal excipulum 80–150 lm broad,
outermost cells catenuliform in long fascicled tufts, hyaline, evenly blue in cotton blue, with 23–55 9 9–18 lm
cylindrical to subclavate, slightly thick-walled, J- end
cells. Stipitipellis 60–100 lm, hyaline, terminal cells 22–32
9 10–18 lm, clavate, J-. Asci 250–290 9 13–15 lm,
pleurorhynchous, 8-spored, subcylindrical to clavate. Paraphyses filiform, 4–5 lm broad, slightly exceeding the
asci, apex enlarged, 6–7.5 lm broad, slightly blue in cotton
blue, J-. Ascospores [100/4/4, in H2O] 15–17(–17.5) 9
9.5–11.5(–12) lm [Q = (1.4) 1.45–1.7(–1.78), Q = 1.56
± 0.09)], ellipsoid, smooth-walled under the light microscope, uniguttulate. Asexual morph Undetermined.
Habitat: Solitary or gregarious on the ground. In
northern Thailand in conifer forest dominated by Pinus
kesiya, and in southwestern China in conifer forest dominated by P. yunnanensis Franch.
Distribution: Only known from Eastern Asia.
Material examined: THAILAND, Chiang Mai Province,
Chiang Mai County, Mushroom Research Center, in
coniferous forest dominated by Pinus kesiyi, alt. 640 m, 31
August 2015, Qi Zhao 2661 (HKAS 90535, holotype).
CHINA, Yunnan Province, Yulong County, alt. 1840 m, 21
Aug 2010, Q. Zhao 8292 (HKAS 69968).
Notes: Gross morphological differences between
H. crispoides and H. pseudoreflexa Zhao et al. (2015) are
subtle, the broad medullary excipulum of H. crispoides is
usually 280–450 lm, while in the latter species it is generally 200–280 lm. The main differences between them are
probably their ecological preferences, H. crispoides is
associated with coniferous forests, while H. pseudoreflexa
is mainly distributed in deciduous forests (Zhao et al.
2015). Both H. crispoides and H. papuensis Weber exhibit
an irregular saddle-shaped pileus, which is deeply ribbed,
with cross-veins and stipe pockets. However, H. papuensis
is a tropical species originally described from Papua New
Guinea, in the southern hemisphere by Dissing (1979). It is
diagnosed by oxide yellow to brownish-yellow hymenium,
with a villose to tomentose sterile surface which is concolorous or slightly darker than the hymenium surface
(Dissing 1979). Helvella crispoides has a cream hymenium
and a grey, pubescent, receptacle surface.
Sarcoscyphaceae Le Gal ex Eckblad
Cookeina Kuntze
Cookeina is a genus of cup fungi in the family Sarcoscyphaceae, members can be found in tropical and
subtropical regions of the world. Species in the Cookeina
have a deep, cup-shaped to funnel-shaped fruiting bodies,
or apothecia, may be found on fallen branches of angiosperms, trunks, and sometimes on fruits (Weinstein et al.
2002) (Fig. 109).
Cookeina tricholoma (Mont.) Kuntze, Revis. gen. pl.
(Leipzig) 2: 849 (1891)
Facesoffungi number: FoF 3136, Figs. 110, 111
Apothecia deeply cup-shaped, 4–16 mm diam. and up to
43 mm high, more or less long stalked, funnel-shaped.
Hymenial surface smooth, pale yellow-orange or pale pink-
123
Fungal Diversity
Fig. 108 Helvella crispoides (HKAS 90535, holotype). a–c Typical mature specimens (a–b, HKAS 90535, c HKAS 69968). d Receptacle
surface of pileus. e–f Stipitipellis. g Asci and paraphyses. h–m Asci. n. Ascospores. Scale bars a–c = 1 cm, d–i = 40 lm, j = 17 lm
orange to orange-reddish. Outer surface covered with long,
whitish or very faintly orange brown hairs (1–2 lm diam.),
obviously denser at the margin, grouped in bundles,
123
concolorous to the hymenial surface, sometimes rough;
margin smooth, whole, often introflexed. Stalk cylindrical,
long, slightly expanded at the top, about 3 mm diam.,
Fungal Diversity
Fig. 109 Phylogenetic
relationships inferred from
maximum parsimony analysis
of ITS-rDNA sequences of 22
taxa. The percentages of
replicate trees in which the
associated taxa clustered
together in the bootstrap test
(1000 replicates) are shown next
to the branches. The tree is
drawn to scale, with branch
lengths in the same units as
those of the evolutionary
distances used to infer the
phylogenetic tree. The new Sri
Lankan record: Cookeina
tricholoma having GenBank
Accession Number KY649459
and Herbarium Number MFLU
12-2390 is shown in bold and
blue. The tre is rooted with
Microstoma floccosum.
Evolutionary analysis was
conducted in PAUP 4.0b 10
smooth, white-cream. Flesh whitish or very pale whitishorange, waxy. Ascospores variable in size, 33–36 9 13–16
lm, fusiform, few irregularly citriform, occasionally with
slightly mucronate extremities, with 10–20 longitudinal
thin striations, straight and more or less parallel, wall up to
2–3 lm thick, usually with a large cylindrical-ellipsoidal
oil drop, rarely 2 oil-drops, hyaline, uniseriate in the asci.
Asci 300–400 9 15–20 lm, 8-spored, cylindrical, wall 1–2
lm thick, with rapidly tapered base that ends in a thin and
usually curved appendage 15–35 lm long, not amyloid,
suboperculate. Paraphyses 2–3 lm diam., thread-like,
cylindrical, simple or pluri-forked. Subhymenium not
noticeable. Medullar excipulum 230–300 lm thick; textura
intricata, hyphae of 6–9 lm diam., interwoven, variously
anastomosed, septate, hyaline or slightly yellowish. Ectal
excipulum 300–350 lm thick; textura globulosa, cells
18–30 lm diam., rounded, rarely angular, globose or
slightly ellipsoidal, wall thin or somewhat thickened, grey-
123
Fungal Diversity
Fig. 110 Fruiting body of Cookeina tricholoma in the field (MFLU
12–2390, reference specimen)
brown or pale brown. Presence of catenulate marginal
cells, apex irregularly shaped or enlarged. Hairs up to 260
lm long, 40–50 lm large at the apex and 100–150 lm at
Fig. 111 Cookeina tricholoma
(MFLU 12-2390). a Asci with
ascospores. b Ascospores.
c Hairs. d Paraphyses
123
the base, arising from the medullar excipulum, formed by
more or less parallel fenced hyphae 10–15 lm wide,
cylindrical or slightly enlarged in some points, with rounded, capitulate tip, irregularly thickened in some points,
wall 2–4 lm thick, septate, yellow–brown, less dense near
the top and more numerous and shorter near the base.
Habit, habitat and distribution: as groups of more or less
close fruit-bodies, usually on dead or decaying wood,
May–August. Our collection was collected on a decaying
wood at Hanthana Mountains Range, Peradeniya.
Specimens examined: SRI LANKA, Kandy District,
Hanthana Mountains Range, 19 June 2012, Samantha C.
Karunarathna (MFLU 12-2390, reference specimen designated here).
GenBank Number ITS:KY649459.
Notes: The main distinguishing characteristics of C. tricholoma are pale yellow-orange apothecia and large
ascospores. However, colorful C. sulcipes (Berk.) Kunze,
are usually orange-red, cherry-red or scarlet, might be
confused with C. tricholoma; its ascospores, although with
the same kind of ornamentation, are smaller in size (25–30
9 11–14 lm). This is the first report of C. tricholoma with
the molecular phylogenetic confirmation from Sri Lanka
Fungal Diversity
(Fig. 111). We therefore designate it as a reference specimen (sensu Ariyawansa et al. 2014b).
Basidiomycota
Agaricomycetes Doweld
Agaricales Underw.
Agaricaceae Chevall.
The family Agaricaceae was introduced by the French
botanist François Fulgis Chevallier in 1826 (Chevallier
1826). Named after the type genus Agaricus, the family
was originally circumscribed by Carl Linnaeus in 1753 in
his work on Species Plantarum.
Amanita Pers.
The morphologically diverse and monophyletic genus
Amanita Pers. (Persoon 1797; Bas 1969; Zhang et al. 2004;
Wolfe et al. 2012b; Tulloss et al. 2016) is dominantly
mycorrhizal however, at least one species is saprobic
(Wolfe et al. 2012a) and another three dozen or so are nonmycorrhizal with precise trophic modes unclear. The genus
is divided into two subgenera: Amanita subg. Amanita
(with inamyloid spores) comprising three sections, viz
sect. Amanita, sect. Caesareae Singer, sect. Vaginatae
sensu Yang (1997) and A. subg. Lepidella (E.-J. Gilbert)
Veselý emend. Corner & Bas (with amyloid spores)
includes four sections namely, sect. Amidella (E. J. Gilbert)
Konrad & Maubl., sect. Lepidella sensu Bas (1969),
sect. Phalloideae (Fr.) Quél., and sect. Validae (Fr.) Quél.
More than 530 species are described worldwide with valid
accepted names and about 50 taxa are reported from India
(Yang 2000; Bhatt et al. 2003; Tulloss and Yang 2016).
Here, we describe two new species belonging to subg.
Amanita sect. Vaginatae. They were collected from the
northwestern Indian Himalayas and are presented here with
morphological and molecular data. The respective single
nrLSU gene tree is presented in Fig. 112.
Amanita cornelii Mehmood, K. Das, Iqbal Hosen, Tulloss,
& R.P. Bhatt, sp. nov.
MycoBank number: MB 817887; Facesoffungi number:
FoF 2946, Figs. 113, 114 Etymology: in honor of Cornelis
Bas for his major revision of the study of Amanita.
Holotype: CAL 1337
Pileus 40–60 mm wide, initially convex then planoconvex, umbonate, dark brown (8F4–6) over centre,
otherwise having brown to greyish-brown (6D3) rather
coarse and irregular radial streaks on pallid ground,
smooth, tacky, shiny. Pileus context 3–5 mm thick, thinning evenly toward margin, white, unchanging when
exposed or bruised. Margin short-striate up to 10 mm, nonappendiculate. Universal veil on pileus absent. Lamellae
free, subdistant (7–8 per 10 mm at margin), 3 mm broad,
white (1A1) in mass and in side view. Lamellulae scarce,
truncate, of at least 2 lengths, unevenly distributed. Stipe
100–120 9 8–12 mm, slightly tapering upward, with white
(2A1) ground, densely covered by light brown to gray
(6F3–5) fibrils arranged in ‘‘flame’’ or ‘‘zebroid’’ pattern,
with fibrils darkening to nearly black where handled. Stipe
context white, hollow, unchanging when bruised or
exposed. Partial veil absent. Universal veil at stipe base
saccate, 33–39 9 18–24 mm, white (1A1–2A1) thick,
membranous, persistent. Spore print white. Odor indistinct.
Taste not recorded. Basidiospores [50/2/1] (8–)9–11.5
(–13.5) 9 (7.5–)8–10.5(–13) lm (L = 9–10 lm; L0 = 9.31
lm; W = 8–10 lm; W0 = 8.92 lm; Q = (1–)1.02–1.10
(–1.25); Q = 1.05–1.11; Q0 = 1.06), hyaline, colourless,
thin walled, inamyloid, globose to subglobose, rarely
broadly ellipsoid, with uniguttulate contents, apiculus lateral to sublateral 1.5 9 1 lm. Basidia (39–)42–51(–57) 9
(11–)12–14(–15) lm, 2- to 4-spored, thin-walled, colourless; sterigmata up to 4 9 1.5 lm. Clamp connections not
observed at the base of Basidia. Subhymenium wstnear = 52–75 lm thick, wst-far = 60–88 lm, Basidia
arising from small inflated cells (up to 12 9 14 lm wide).
Lamellae edge sterile; inflated cell clavate or pyriform,
35–50 9 22–31 lm, colourless, thin walled; clamp connections not observed Hymenophoral trama bilateral,
divergent; wcs = 50–80 lm; well rehydrated, filamentous
undifferentiated hyphae (3–16) lm wide; vascular hyphae
11–15 9 3–4 lm wide. Pileipellis 130–195 lm thick, in
two layers, with gelatinized colorless suprapellis (40–55
lm thick), and ungelatinized subpellis (95–130 lm thick);
ungelatinized filamentous undifferentiated hyphae 2–6 lm
wide, subradially arranged densely arranged in subpellis,
with brown intracellular pigment vascular hyphae 7–10 9
2.8–3.5 lm. Pileus trama filamentous, undifferentiated
hyphae 2.5–16 lm wide, constricted at septa, thin-walled,
branching, forming loose open matrix; acrophysalides
common up to 120 9 23 lm, thin-walled, hyaline, vascular
hyphae rare up to 15 9 5 lm. Exterior surface of universal
veil (on stipe base) filamentous, undifferentiated hyphae
3–12 lm wide; inflated cells subclavate (up to 84 9 34
lm), narrowly ellipsoid to elongate (e.g. 120 9 18 lm).
Interior surface of universal veil (on stipe base) filamentous, undifferentiated hyphae 3–9 lm wide; inflated cells
subglobose to ovoid (up to 127 9 119 lm), infrequent.
Stipe context longitudinally acrophysalidic; filamentous
undifferentiated hyphae (2–9 lm wide); acrophysalides
dominating, 151–190 9 31–42 lm; clamp connections not
observed.
Habitat and distribution: Under Quercus sp., in temperate mixed forest dominated by Abies and Quercus.
Material examined: INDIA, Uttarakhand, Rudraprayag
District, Baniyakund, 2630 m, N3028.9140 E7910.8540 , 14
July 2015, T. Mehmood, TM 15–625 (CAL 1337, holotype); 18 July 2015, ibid. (TM 15–693, paratype).
123
Fungal Diversity
Fig. 112 RAxML tree and putative relationships of Amanita subg.
Amanita sect. Vaginatae inferred from analysis of LSU sequence data.
Bootstrap support values ([50%) obtained from maximum likelihood
analysis are shown above or below the branches at the nodes. The new
taxa are highlighted in bold and blue font in the tree. GenBank
numbers accession numbers are provided after each species name and
followed by country of origin. Amanita caesareae (Scop.) Pers.
served as the outgroup taxon. The tree is artificial in that it embodies
two somewhat distant clusters of taxa associated with the two new
taxa by morphology and/or BLAST searches. This results in a
compact single figure and, also, in low support for nodes near the
tree’s root
GenBank Number ITS: KX528072.
Notes: The combination of macro- and micromorphological features such as inamyloid spores, saccate volva,
absence of a bulb and absence of Basidia clamps place
Amanita cornelii in Amanita [subg. Amanita] sect. Vaginatae sensu Yang (Yang 1997). In the field the present
123
Fungal Diversity
Fig. 113 Amanita cornelii (CAL 1337, holotype). a–d Fresh basidiomata in the field. e Basidia and basidiole. f, g Lamellae edges cells.
h Elements of universal veil from stipe base. i Basidiospores. Scale bars a, b = 50 mm, e–i = 50 lm
species is distinct from other known species of Amanita
sect. Vaginatae by its pileus with coarse, irregular, radial,
brown streaks. In eastern Asia, Amanita cornelii might be
confused with Amanita brunneofuliginea Zhu L. Yang
(Yang 1997) (originally described from China); however,
the latter has a pileus that is blackish-brown to brownishblack over the centre. The latter also differs by lacking
coarse, irregular, brown stripes on the pale background on
123
Fungal Diversity
Fig. 114 Amanita cornelii
(CAL 1338, holotype).
a Basidiomata. b Lamellae edge
cells. c Basidiospores.
d Elements of universal veil at
stipe base. e Basidia and
elements of hymenium and
subhymenium. Scale bars
a = 10 mm, b–e = 10 lm
the pileus, bearing a white to dirty white patches of universal veil on the pileus, its having a saccate volva which is
dirty white with plentiful pale-leather to orange spots on its
outer surface and by its subglobose to broadly ellipsoid
basidiospores ‘‘(10–)10.5–13(–14) 9 (9–)9.5–12(–12.5)
lm, (Q = (1.04–)1.06–1.24(–1.26); Q0 = 1.13 ± 0.06)’’
(Yang 1997). Amanita lignitincta Zhu L. Yang (Yang
1997) (originally described from China), is somewhat
similar to the present taxon with a wood brown to leather
brown pileus; but it differs by its slightly depressed pileus,
a contrastingly darker ring–like zone at the inner end of its
marginal striations, by its larger basidiospores ‘‘(9.5–
123
)10–13(–13.5) 9 (8.5–)9–12(–13) lm’’ (Yang 1997) and
by lacking the unusual striping pattern of the present species. Our nrLSU-tree clearly indicates the genetic dissimilarities of Amanita cornelii from six taxa. BLAST search
indicates relative closeness (96% identity with 95% query
coverage) of A. cornelii to A. friabilis (GenBank Numbers
KU248120, KU248119); (96% identity to 96% query
coverage) of A. cornelii to A. basiana (GenBank Numbers
KP258987); (96% identity with 94% query coverage) of
A. cornelii to A. affin. fulva (GenBank Numbers
HQ539697); (96% identity with 95% query coverage) of
A. cornelii to A. olivaceogrisea (GenBank Numbers
Fungal Diversity
KU867877); (95% identity to 95% query coverage) of
A. cornelii to A. affin. ceciliae (GenBank Numbers
KU139438, KU139439); (97% identity with 95% query
coverage) of A. cornelii to A. populiphila (GenBank
Numbers KP221314, KP221304).
The European A. basiana Tulloss & M. Traverso (Tulloss and Traverso 2000) and A. friabilis (Karst.) Bas (Bas
1974) are easily segregated from the present species
because, in those two taxa, hyphae provide extensive and
persistent interconnection between the pileipellis and the
universal veil (Bas 1974; Tulloss and Traverso
2000, 2001). Moreover, A. friabilis is separated from
A. cornelii by its subglobose to elongate spores
‘‘Q = (1.06–)1.13–1.62(–1.89);
Q = 1.23–1.43;
Q0 = 1.34’’
(www.amanitaceae.org/?Amanita%20friabi
lis); and A. basiana also differs from the present taxon due
to its broadly ellipsoid to ellipsoid spores Q = (1.08–
)1.13–1.44(–2.34); Q = 1.21–1.33(–1.36); Q0 = 1.27
(www.amanitaceae.org/?Amanita%20basiana).
Volval differences also serve to segregate species that
could be mistaken for A. ceciliae (European) and A. olivaceogrisea (European) from the present species. It should
be noted that many species in eastern Asia, eastern North
America and other regions have been mistaken for A. ceciliae in the past. This group have friable volva’s that
quickly become gray with age often leaving warts and
small patches on the pileus, fragments on the lower stipe,
and a white cupulate structure on the stipe’s base (Bas
1984; Tulloss 1994, 2001). Amanita olivaceogrisea also
differs from Amanita cornelii by its subglobose to broadly
ellipsoid spores showing slightly higher Q values, i.e.
Q = (1.03–)1.04–1.14(–1.27); Q = 1.09–1.11; Q0 = 1.10
(www.amanitaceae.org/?Amanita%20olivaceogrisea).
Amanita affin. fulva (North America) possesses an
orange-brown pileus that becomes paler at the margin and
brown to dark brown in the center, lacks the irregular
striping pattern of the present species, and has a white
external volva surface that becomes orange-brown to rusty
in numerous spots. The matching GenBank Numbers
sequence is a nearly perfect match to sequences derived
from Newfoundland material presently treated under a
provisional name by Tulloss (www.amanitaceae.org/
?Amanita%20daimonioctantes).
Amanita populiphila Tulloss & Moses (Tulloss and
Moses 1995), described from North America differs from
the present taxon by an unstriped, white to pale yellowish
pileus that takes on rusty stains with time; cream-colored
Lamellae having pale orange or pinkish tint; occurrence of
plentiful lamellulae and a weakly structured, white volva
that tends to change color as the pileus does. Amanita
populiphila has only been recorded with species of Populus
[cottonwood and aspen] and has subglobose spores with
slightly higher Q values than that of the present taxon;
Q = (1–)1.04–1.20(–1.61); Q = (1.06–)1.08–1.15(–1.19);
Q0 = 1.11.
(www.amanitaceae.org/?Amanita%20populi
phila). Macro- and micromorphology coupled with the
LSU-based genetic distances, segregate Amanita cornelii
from previously known species of Amanita sect. Vaginatae.
Amanita emodotrygon Mehmood, Tulloss, K. Das, Iqbal
Hosen & R.P. Bhatt sp. nov.
Index Fungorum number: IF552463; Facesoffungi
number: FoF 2947, Figs. 115, 116
Etymology: emodos (Grk., India) ? trygon (Grk., dove),
to indicate possible genetic grouping with North American
A. trygonion Tulloss, Pastorino & Kudzma nom. prov.
(www.amanitaceae.org/?Amanita%20trygonion)
Holotype: CAL 1338
Pileus 96–132 mm wide, initially convex then campanulate, initially brownish-grey to greyish-brown (7E2–3)
and finally olive (1E4–4E7) smooth, dry, shiny. Pileus
context 8–10 mm thick, first thinning slowly then rapidly
toward margin, membranous at margin, off-white,
unchanging when bruised or exposed. Margin with distinct
regular veins or striations and corresponding grooves (up to
36 mm wide few veins forked more toward margin, nonappendiculate, whitish to greyish in between the grooves.
Universal veil on pileus absent Lamellae free, crowded
(11–13 per 10 mm at margin), 6 mm broad, initially white
(1A1) fading pale yellow with age. Lamellulae truncate, of
3–4 lengths, unevenly distributed. Stipe 150–183 9 16–19
mm, equal or slightly tapering upward initially off-white
then greenish-grey to greyish-green (1C2–3), with olive
brown tinge toward the base. Context off-white, hollow,
stuffed in button stage. Partial veil absent. Universal veil at
stipe base saccate, 33–41 9 19–29 mm, white to off-white
(1A1–2A1) with many olive-brown spots on exterior surface when age, bilimbate, membranous, thick at bottom
thin toward margin. Spore print cream. Odor indistinct.
Taste not recorded. Basidiospores [60/3/1] (8.5–)9.8–12
(–13.5) 9 (8–)9.5–11(–13) lm (L = 10–11.5 lm; L0 =
10.9 lm; W = 9.5–10.5 lm; W0 = 10.2 lm; Q = (1–)
1.05–1.12(–1.17); Q = 1.05–1.11; Q0 = 1.08), globose to
subglobose, rarely broadly ellipsoid, inamyloid, hyaline,
colourless, thin walled, with uniguttulate contents, apiculus
lateral to sublateral to 1.5 9 1.2 lm. Basidia (46–)52–65
(–70) 9 (14–)14.5–16(–17) lm, 2- to 4-spored, thin-walled, colorless; sterigmata 4–6 9 1.5–2 lm. Clamp connections not observed at the base of Basidia. Subhymenium
wst-near = 35–65 lm; wst-far = 50–75 lm, Basidia arising mostly from inflated cells to irregular cell (up to 10 9 8
lm wide). Lamellae edge sterile; inflated cells; pyriform or
clavate, 43–48 9 13–22 lm, colourless, thin-walled, infrequent; clamp connections not observed. Hymenophoral
trama bilateral, divergent; wcs = 70–90 lm filamentous
undifferentiated hyphae (4–9) lm wide; inflated cells
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Fungal Diversity
Fig. 115 Amanita emodotrygon (CAL 1338, holotype). a–c Fresh basidiomata in the field. d Basidia and basidiole. e Elements of universal veil
from stipe base. f Basidiospores. Scale bars b = 100 mm, d–f = 50 lm
19–27 9 38–45 lm; vascular hyphae not observed.
Pileipellis 125–190 lm thick, in two layers; slightly gelatinized colorless suprapellis (30–60 lm thick) of radially
arranged filamentous undifferentiated hyphae 1–3 lm
wide, thin-walled, colourless; nongelatinized subpellis
(90–130) lm thick); filamentous undifferentiated hyphae
2–7 lm wide, subradially and compactly arranged with
brown intracellular pigment; vascular hyphae not observed.
Pileus trama filamentous, hyphae 2–12 lm wide, constricted at septa, slightly thick-walled, hyaline, branching,
acrophysalides common up to 135 9 31 lm, slightly thickwalled, vascular hyphae up to 17 9 3.5 lm. Exterior surface of universal veil (on stipe base) filamentous, undifferentiated hyphae 4–12 lm wide, slightly thick walled,
septate and frequently branched, colourless, hyaline; globose to subglobose cells up to 102 9 108 lm; ellipsoid
123
cells up to 116 9 39 lm. Interior surface of universal veil
(on stipe base) filamentous, undifferentiated hyphae 4–8
lm wide; inflated cells subglobose cells to 45 9 41 lm,
clavate cells up to 78 9 23 lm, infrequent. Stipe context
longitudinally acrophysalidic; filamentous hyphae 4–9 lm
wide, acrophysalidic dominating, (151–190 9 31–42 lm);
Clamp connections not observed.
Habitat and distribution: Solitary to scattered, under
Pinus roxburghii in temperate coniferous forest of western
Himalaya.
Specimen examined: India, Uttararakhand, Rudraprayag,
Hariyali Devi Forest, 1651 m, N3015.9550 E7903.7190 , 27
July 2015, T. Mehmood, TM 15-659 (CAL 1338,
holotype).
GenBank Number ITS:KX539266.
Fungal Diversity
Fig. 116 Amanita
emodotrygon (CAL 1338,
holotype). a Basidiomata.
b Basidiospores. c Basidia and
element of subhymenium.
d Lamellae edges cells.
e Elements of universal veil at
stipe base. Scale bars a = 10
mm, b–e = 10 lm
Notes: The combination of macro- and micromorphological features such as longitudinally acrophysalidic stipe
context; bilateral, divergent lamella trama, inamyloid
spores, absence of a stipe bulb and absence of Basidia
clamps place Amanita emodotrygon in Amanita [subg.
Amanita] sect. Vaginatae Sensu Yang (Yang 1997).
In the field this species is distinct from other known
species of Amanita sect. Vaginatae by its greyish-brown to
olive campanulate pileus, which is deeply grooved towards
the margin, crowded white lamellae that becomes pale
yellowish with age and occurrence in a coniferous forest
under Pinus roxburghii. Micromorphologically, presence
of globose to subglobose spores, branched, septate and
thick-walled, filamentous hyphae in the universal veil at
the stipe base are also quite striking. Based on the key in
Tulloss (1994) and (www.amanitaceae.org/?stirps%
20Pachyvolvata), there is a group of North American and
European taxa that are morphologically similar to
A. emodotrygon in having a large, robust basidiome and
universal veil-strips Pachyvolvata of Tulloss. The relevant
section of the key is largely based on cap colour and spore
size and shape. Two of the taxa are described as having an
olivaceous tint to the cap. Two are described as having a
ring around the inner ends of marginal striations. Hence,
123
Fungal Diversity
gross morphology of this group is similar to that of
A. emodotrygon. One of the species has both the cited
pileus characters and is, hence, most similar to the present
taxon:
Amanita magnivolvata Aalto (1974) (originally reported
from Finland) is segregated from A. emodotrygon by its
gray ring on the inner edge of the marginal striations, white
Lamellae becoming gray with age and a white exterior
surface of the universal veil, developing small, rusty, yellow spots. It occurs in mixed forest including Alnus, Betula, and Picea in Scandinavia and has been reported from
Italy (Liguria). Its spores are subglobose to broadly ellipsoid and have comparatively higher Q values than occur in
the
present
taxon-Q = (1.03–)1.06–1.36(–1.52);
Q = 1.18.
(www.amanitaceae.org/?Amanita%
20magnivolvata)
Our nrLSU-tree clearly indicates the genetic dissimilarities of Amanita emodotrygon from three taxa. BLAST
search indicates relative closeness (98% identity with 98%
query coverage) of Amanita emodotrygon to Amanita trygonion (GenBank Numbers KU186810); (94% identity
with 94% query coverage) of Amanita emodotrygon with
Amanita penetratrix (GenBank Numbers KU186834,
KU186833, KU186832, KU186831); (93% identity to 93%
query coverage) of Amanita emodotrygon to Amanita
myrmeciae (GenBank Numbers KU852505 and
KU186806).
The relatively small number of taxa that have been
sampled for nrLSU in section Vaginatae and the great
difficulty in dealing with the genus by morphological
means alone create an unusual situation in the present case,
i.e. all of the sequences for which close BLAST comparisons exist in the case of A. emodotrygon are taxa described
provisionally. Hence, we can say, at the time of writing, the
present taxon is quite distinct from all published species of
sect. Vaginatae for which sequence data exists.
Amanita trygonion Tulloss, Pastorino & Kudzma, nom.
prov. is a small gray species of the sandy coastal plain in
southeastern Texas, USA, where it occurs in Pinus dominated mixed forest and has subglobose to broadly ellipsoid
spores with higher Q values than those of the present
taxon-Q = (1.05–)1.10–1.33(–1.40);
Q = 1.17–1.20;
Q0 = 1.19.
(www.amanitaceae.org/?Amanita%
20trygonion)
Amanita penetratrix Tulloss & Kudzma, nom. prov. is a
large species with a hard-fleshed, prominent umbo in the
center of a gray to gray-brown to white pileus. It is known
from a few sites in the northeastern USA. It occurs in
mixed forest including Pinus strobus, Betula, Fagus, and
Quercus and has spores that are dominantly subglobose to
broadly ellipsoid with higher Q values than those of the
present
taxon-Q = (1.04–)1.05–1.25(–1.34);
Q = 1.09–1.16; Q0 = 1.12. The primordium of this
123
mushroom develops deep in forest soil. The unusual umbo
allows penetration of up to 154 mm of soil dense with roots
and is reminiscent of the perforatorium of species of Termitomyces.
(www.amanitaceae.org/?Amanita%
20penetratrix)
Amanita myrmeciae Tulloss, Kudzma & Albertella,
nom. prov. is known only from association with eucalypts
in the Blue Mountains of New South Wales, Australia. This
species is unique among those discussed here by being a
significant food source for ants of the genus Myrmecia. The
species is known from one specimen with a light brown to
brown pileus 62 mm wide and has subglobose spores with
slightly higher Q values than those of the present taxon—
Q = 1.06–1.18; Q 1.10. (www.amanitaceae.org/?Ama
nita%20myrmeciae)
Lepiota (Pers.) Gray
Lepiota (Pers.) Gray (Agaricaceae) is a large group of
saprotrophic basidiomycetes in temperate and tropical
areas, comprising six sections and an estimated 500 species
(Singer 1986; Vellinga 2004; Kirk et al. 2008). A tree to
three new species from Thailand is provided below
(Fig. 117).
Lepiota cylindrocystidia Sysouphanthong & K.D. Hyde,
sp. nov.
Index Fungorum number: IF552903; Facesoffungi
number: FoF 2948, Figs. 118, 119
Etymology: The name of this species ‘‘cylindricystidia’’
comes from its cylindrical Cheilocystidia.
Holotype: MFLU 12-2035
Pileus 16 mm, broadly convex with inflexed margin,
covered with spine-like or pyramidal squamules over the
whole surface, slightly crowded at center and more distant
towards margin, squamules darker or yellowish-brown,
light brown to golden-brown (5D5–8, 5E5–6) on greyishyellow to greyish-orange (4B4–5, 5B4–5) fibrillose or
floccose background; at marginal zone, fibrillose, concolorous with background. Lamellae white, free, ventricose, 1.5–2 mm wide, crowded, with white eroded edge.
Stipe 38 9 2.5–3 mm, cylindrical or slightly wider at base
and apex, covered with crowded fibrillose squamules from
annular zone downward base, with squamules at basal
zone, concolorous with those on pileus, with white to
yellowish-white to pale yellow (4A2–3) fibrillose at
annular zone upward from apex. Annulus an annular zone
with fibrillose and fibrillose squamules as referred to on
stipe. Context in pileus white, 3 mm wide; white and
hollow in stipe. Taste and smell not observed. Spore print
white. Basidiospores [25, 1, 1] 4–5 9 3–3.2 lm, avl 9
avw = 4.62 9 3.1 lm, Q = 1.33–1.56, avQ = 1.49, in
side-view ellipsoid ovoid, in frontal view ellipsoid, hyaline, thin-walled, dextrinoid, congophilous, cyanophilous,
Fungal Diversity
Fig. 117 Bayesian tree based on ITS sequence alignment of Lepiota
species represent from each section. Support values from Bayesian
posterior probabilities values (BYPP) more than 0.95 are given above
tree branches, and bootstrap percentage more than 50% from
maximum likelihood analyses are indicated below the tree branches.
New sequences from Thailand are as bold. Agaricus bisporus and
Clarkeinda trachodes are outgroup taxa
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Fungal Diversity
not metachromatic. Basidia 20–26 9 5–6.5 lm, clavate,
4-spored, hyaline, thin-walled. Lamella edge sterile.
Cheilocystidia 22–34 9 7–17 lm, mostly cylindrical,
rarely narrowly clavate, hyaline, thin-walled, often branch
and septate. Pleurocystidia absent. Pileus covering an
epithelium made up of globose elements in lower layer
(10–20 lm), with subglobose to oblong elements and
often with subclavate elements in upper layer (8–32 9
5–20 lm, thick-walled, parietal and intracellular pigment
in upper elements and parietal in lower elements. Stipe
covering in squamules an epithelium similar to pileus
covering. Clamp-connections present in all tissues.
Habitat and distribution: Growing solitary, on humusrich soil with decaying leaves and wood in deciduous
forest; only known from Mae Taeng District, Chiang Mai
Province, Thailand.
Material examined: THAILAND, Chiang Mai Province,
Mae Taeng District, Forest of Pha Deng Village,
N19°070 13.700 , E98°430 52.900 , alt. 905 m, 30 June 2011, P.
Sysouphanthong (MFLU 12-2035, holotype).
GenBank Number ITS:JN224828.
Notes: All materials of Lepiota cylindrocystidia are
recognized as having a yellow basidiomata, with cylindrical cheilocystidia.The ITS sequence data from the holotype
clustered with species in sect. Echinatae and related to, but
distinct from species with ovoid basidiospores.
Lepiota flavocarpa Sysouphanthong & K.D. Hyde sp. nov.
MycoBank number: MB 519962; Facesoffungi number:
FoF 2949, Figs. 118, 120
Etymology: named because of its yellow basidiomata.
Holotype: MFLU 10-0581
Pileus 20–45 mm, convex at first, then umbonate with
low umbo, applanate to plano-concave with straight
margin, at center covered by crowded and large pyramidal
warts, with or without recurred tips, and distant towards
margin, dark yellow to yellowish-brown (4C8, 5D8), on
yellow to vivid yellow (3A6–8), fibrillose back ground,
when mature, trips of pyramids around margin fragile,
pyramids becoming large squamules, and later fibrillose
squamules; margin appendiculate with fibrillose or cortinate, concolorous with back ground or yellowish-white to
pale yellow (1A2–3). Lamellae free, crowded, ventricose,
4–8 mm wide, concolorous with pileus background,
consistent, turning brownish-red to dark red (10C6–8)
with KOH. Stipe 40–40 9 4–8 mm, cylindrical or slightly
tapering to apex, background fibrillose, concolorous with
pileus, completely covered with crowded lanate flocculate
or squamules from base upward to annular zone, then
absent toward apex, at annular zone with concolorous
fibrillose or cortinate as on pileus margin. Annulus an
123
Fig. 118 Habits of Lepiota species in the field. e L. flavocarpa. f–h
L. cynlindrocystidia
annular zone, cortinate or fibrillose, yellowish-white to
pale yellow (1A2–3). Context in pileus concolorous with
surface, up to 5 mm wide; in stipe concolorous with
surface, hollow. Taste and smell not observed. Spore print
white. Basidiospores [50, 2, 2] 4–4.5 9 3–3.2 lm, avl 9
avw = 4.24 9 3.1 lm, Q = 1.33–1.40, avQ = 1.36, in
side-view ellipsoid ovoid, in frontal view ellipsoid, dextrinoid, congophilous, cyanophilous, not metachromatic in
Fungal Diversity
Fig. 119 Lepiota cylindrocystidia (MFLU 12-2035, holotype). a Basidiospores. b Pileus and stipe covering. c Cheilocystidia
Cresyl Blue. Basidia 16–20 9 6–10 lm, narrowly clavate,
4-spored, hyaline to pale yellow-walled. Lamella edge
sterile. Cheilocystidia 20–28 9 7–13 lm, clavate, fusiform or utriform, hyaline to pale yellow and thin-walled.
Pleurocystidia absent. Pileus covering a sub hymeniderm
or epithecium made up of agglutinate chains of mostly
subglobose to oblong elements, 35–75 9 20–27 lm,
sometimes globose, 18–25 lm, yellow to pale brown
parietal and intracellular pigments. Stipe covering a sub
hymeniderm made up of subglobose to oblong, 30–60 9
18–25 lm. Clamp connections present in all tissues.
Habitat and distribution: growing on a small group, on
humus-rich soil mixed decayed leaves and branches; in
deciduous forest dominated by Dipterocarpus and Castanopsis spp.; only found in rainy season (June–August)
of Hui Kok Ma, Doi Suthep District, Chiang Mai,
Thailand.
Material examined: THAILAND, Chiang Mai Province,
Muang District, Hui Kok Ma od Doi Suthep, N18°480 6200 ,
E98°540 6000 , alt. 1145 m, 20 June 2010, P. Sysouphanthong, P63 (MFLU 10-0581, holotype); ibid., 18 June
2010, P. Sysouphanthong, P47 (MFLU 10-0565); ibid., 19
June 2010, P. Sysouphanthong, P50 (MFLU 10-0568);
ibid., 27 August 2010, P. Sysouphanthong, P110 (MFLU
10-0628).
GenBank Number ITS:KP348285.
Notes: All material of Lepiota flavocarpa is recognized
as having a yellow basidiomata, with dark yellow to
yellowish-brown pyramidal squamules on the pileus,
turning brownish-red to dark red in KOH for whole
basidiomata. The characters of ovoid Basidiospores and the
subhymenidermal pileus covering made up of globose to
oblong elements located this species in sect. Echinatae.
The ITS sequence data from the holotype clustered with
species in sect. Echinatae and related to, but distinct from
species with ovoid basidiospores.
Lepiota maerimensis Sysouphanthong & K.D Hyde, sp.
nov.
Index Fungorum number: IF552904; Facesoffungi
number: FoF 2950, Fig. 121
Etymology: The name of this species ‘‘cylindricystidia’’
comes from its cylindrical Cheilocystidia
Holotype: MFLU 12-2036
Pileus 8–17 mm, at first parabolic to broadly convex,
expanding umbonate with low umbo, inflexed margin; with
warts, or pyramidal squamules, larger at centre and smaller
towards margin, brown and turning dark brown (6F5–8,
6E4–6), on orange-white (5A2) background; margin
cortinate in young samples, squamulose, fibrillose.
Lamellae free, crowded, ventricose, 1.5–2 mm wide, white,
yellowish-white when mature, with white serrulate edge.
Stipe 28–32 9 1.8–2 mm, cylindrical, with bulb, 2.2–2.4
mm wide, completely floccose or with crowded squamules
from annular zone toward base, light brown (6D5–6), white
to yellowish-white (4A2) at apex, hollow. Annulus an
123
Fungal Diversity
Fig. 120 Lepiota flavocarpa
(MFLU 10-0581, holotype). a
Basidiospores. b Basidia.
c Cheilocystidia. d Pileus
covering
annular zone, floccose or squamulose, light brown
(6D5–6). Context white in pileus; white in stipe. Smell and
taste not observed. Spore print white. Basidiospores [25]
4–5 9 2.8–3 lm, avl 9 avw = 4.3 9 3 lm, Q = 1.1–1.6,
Qav = 1.4, oblong ovoid in side-view, oblong in frontal
view, thick-walled, hyaline, not dextrinoid, congophilous,
cyanophilous, not metachrometic. Basidia 10–12.5 9 4–5
lm, clavate, 4-spored. Lamella edge sterile. Cheilocystidia
10–22 9 4–9 lm, clavate, narrowly clavate, rarely utriform, hyaline, thin-walled. Pileus covering an epithelium
made up of several chains elements, terminal elements
10–25 9 7–17 lm, globose to ellipsoid, clavate, subclavate, with brown walls, slightly thick-walled; under
layer with hyaline hyphae, cylindrical, up to 4 lm wide.
Clamp connections present.
123
Habitat and distribution: growing solitary or in small
groups, saprotrophic on rich humus soil, sand or loamyclayey, nutrient-rich soil.
Material examined: THAILAND, Chiang Mai Province,
Mae Rim District, Mae Sa Valley, 12 July 2008, P.
Sysouphanthong (MFLU 12-2036, holotype).
GenBank Number ITS:KP348284.
Notes: Lepiota microspila, a species from Sri Lanka is
similar to L. maerimensis in its ochraceous tawny to rufous
squamules on the pileus and stipe covering. Lepiota
microspila differs from L. maerimensis in having a smaller
spore size (3.3–4.5 9 1.7–2.3 (3.8 9 2) lm, shorter Basidia
(6.5–8 9 3.3.5 lm), and thicker wall of elements in the
pileus covering (up to 1 lm). Unfortunately, Cheilocystidia
of Lepiota microspila were not recovered in the Sri Lankan
Fungal Diversity
Fig. 121 Lepiota maerimensis
(MFLU 12-2036, holotype).
a Basidiocarps. b Basidiospores.
c Basidia. d Cheilocystidia.
e Pileus and stipe covering
material (Pegler 1972). Some species of section Echinatae
from India are compared with this species, for example
Natarajan and Manjula (1982) described Lepiota subrufa
from South India and it differs from L. maerimensis in its
pale red background on the pileus and stipe, and reddishbrown pyramidal squamules, pastel red Lamellae and dull
red membranous annulus. Lepiota babruka is also similar
to L. maerimensis in having brown to dark brown pyramidal squamules on the pileus, but it differs in lacking a
squamulose stipe and yellowish-white to pale yellow
lamellae (Arun Kumar and Manimohan 2009) (Fig. 121).
Leucocoprinus Pat.
Leucocoprinus belongs to the family Agaricaceae. Its
best-known member is the yellow pot-plant mushroom
Leucocoprinus birnbaumii, found worldwide. The type
species of this genus is L. cepistipes (Sowerby) Pat.
Leucocoprinus cretaceus (Bull.) Locq., Bull. mens. Soc.
linn. Soc. Bot. Lyon 14: 93 (1945)
Facesoffungi number: FoF 3137, Figs. 122, 123, 124
Pileus 3–6 cm broad, hemispherical when young,
expanding to planoconvex to campanulate, margin
decurved, slightly sulcate-striate, when young with many,
dense, soft floccules, readily collapsing or wearing off to
leave farinose covering; light buff to white at center, white
elsewhere. Lamellae distantly free, rather broad, thin;
white, edge slightly fimbriate. Stipe 5–8 cm long, 4–6 mm
broad at apex, gradually enlarged downward to a broadly
clavate to somewhat fusiform 6–14 mm broad base;
sometimes coarsely farinose to slightly flocculose-farinose
below annulus, subfarinose above; white to ivory yellow
tinted. Annulus white, very soft, rather flaring, median to
superior. Basidiospores 8–11 9 5–8 lm, Q = 1.32–1.47,
metachromatic, dextrinoid, ellipsoid to amygdaliform,
thick-walled, with an apical germ pore covered with a
hyaline lens. Basidia 16–22 9 9–12 lm, pyriform to clavate with a bulbous base, 4-spored, surrounded by 4
pseudoparaphyses. Sterigmata 1–2 9 0.7–1.5 lm. Pseudoparaphyses 10–13 9 7–10 lm, sphaeropedunculate to
broadly clavate to pyriform, very broad point of attachment. Cheilocystidia 27–75 9 7–18 lm, thin-walled,
hyaline, subcylindrical to narrowly fusiform to slightly
narrowly lageniform, mucronate, rarely obtuse, moderately
pedicellate. Pileus covering a sparse layer of terminal cells
essentially absent on mature pilei, most prevalent near the
center of young pilei, 45–117 9 7–9 lm, cylindrical to
narrowly lageniform, with flexuous necks. Stipe covering
123
Fungal Diversity
Fig. 122 Phylogenetic relationships inferred from maximum parsimony analysis of ITS-rDNA sequences of 11 taxa. The percentages of
replicate trees in which the associated taxa clustered together in the
bootstrap test (1000 replicates) are shown next to the branches. The
tree is drawn to scale, with branch lengths in the same units as those
of the evolutionary distances used to infer the phylogenetic tree. The
new Sri Lankan record: Leucocoprinus cretaceus having GenBank
Accession Number KY649466 and Herbarium Number MFLU
12-1893 (ITS-rDNA) is shown in bold and blue. The topology is
rooted with Leucoagaricus gongylophorus. Evolutionary analysis was
conducted in PAUP 4.0b 10
structure like that of pileus, with terminal cells even more
sparse. Stipitipellis a cutis of narrowly cylindrical, 5–8 lm
broad elements. Clamp connections absent.
Habit, habitat and distribution: as a group, usually on
manure or wood chips, May–August. Our collection was
collected on cow manure at Hanthana Mountains Range,
Peradeniya.
Specimen examined: SRI LANKA, Kandy District,
Hanthana Mountains Range, 6 June 2012, Samantha C.
Karunarathna (MFLU 12-1893, new record).
GenBank Number ITS:KY649466.
Notes: This is widely distributed in North America and
this is the first report of L. cretaceus with molecular phylogenetic confirmation from Sri Lanka.
Entolomataceae Kotl. & Pouzar
Entolomataceae is a large family of pink-spored terrestrial gilled mushrooms introduced by Kotl. and Pouzar
in 1972 and typified by Entoloma. The family is highly
123
Fig. 123 Basidiocarps of Leucocoprinus cretaceus in the field
(MFLU 12-1893)
variable in terms of sporocarp morphology and micromorphology (Noordeloos 2004). This family contains four
genera and 1071 species (Kirk et al. 2008).
Entoloma Fr. ex P. Ku mm.
The genus Entoloma is characterized by pink or pinkishbrown spore print and basidiospores that are angular in all
views (Noordeloos 1992; Noordeloos and Gates 2012).
There is no agreement regarding the limits of the genus
Entoloma as some authors have set a very wide limit to the
genus (e.g. Donk 1949; Romagnesi 1978; Noordeloos
1981, 1992, 2004; Co-David et al. 2009; Noordeloos and
Gates 2012) and others have applied a very restricted
concept of the genus and have introduced several segregate
genera such as Alboleptonia, Claudopus, Inocephalus,
Leptonia, Nolanea, and Pouzarella (e.g. Largent 1974;
Mazzer 1976; Baroni et al. 2011). In the course of a continuing study on the Entolomataceae of Kerala State, India,
we came across a species of Entoloma, which was found to
be new to science. It is herein formally described as new
based on both morphology and molecular phylogeny.
Maximum likelihood analysis placed the new species
within a clade that represents the subgenus Entoloma with
significant (74%) ML bootstrap support (Fig. 125).
Fungal Diversity
Fig. 124 Leucocoprinus
cretaceous (MFLU 12-1893).
a Cheilocystidia.
b Basidiospores. c Basidia.
d Pileus covering
Entoloma magnum K.N.A. Raj & Manim., sp. nov.
MycoBank number: MB 818742; Facesoffungi number:
FoF02637; Figs. 126, 127
Etymology: The specific epithet refers to the large size
of the basidiocarp.
Holotype: CAL 1385
Basidiocarp large, tricholomatoid. Pileus 102 mm
diam., somewhat plano-convex with a low central umbo
surrounded by a shallow depression; surface dark brown
(7F4/OAC636) at the centre and light brown (7D4/
OAC659) towards the margin, not hygrophanous, tacky,
finely radially wrinkled, innately fibrillose, plicate-striate
towards the margin; margin somewhat incurved to almost
straight, finely crenate, fissile. Lamellae narrowly adnate to
adnate, moderately crowded, subventricose, orange white
(6A2/OAC634), up to 13 mm wide, with lamellulae of 3
lengths; edge crenate to somewhat wavy, concolorous with
the sides. Stipe 99 9 18 mm, central, terete, slightly
tapering towards both the apex and the base, robust, solid;
surface yellowish-brown (5D4/OAC743) towards the apex,
whitish towards the base, appressed-fibrillose all over; base
with scanty basal mycelium. Odour and taste not
distinctive.
Basidiospores 7–10 9 7–9 (8.6 ± 0.73 9 7.6 ± 0.66)
lm, Q = 1–1.2, Qm = 1.13, predominantly pentagonal or
occasionally 4- or 6-angled in profile, isodiametric to sub-
isodiametric, hyaline, thick-walled. Basidia 31–57 9 9–11
lm, clavate, hyaline, thin-walled, 4-spored; sterigmata up
to 7 lm long. Lamella-edge fertile. Cheilocystidia and
pleurocystidia absent. Lamellar trama subregular; hyphae
4–7 lm wide, pale brownish-yellow, thin-walled. Subhymenium inconspicuous. Pileus trama subregular; hyphae
7–13 lm wide, pale yellow, thin-walled. Pileipellis a
transition between an ixocutis and an ixotrichoderm;
hyphae 5–10 lm wide, with a pale brownish-yellow wall
pigment, thin-walled. Stipitipellis a cutis rarely disrupted
with flaring out hyphae; hyphae 5–12 lm wide, pale
brownish-yellow, thin-walled. Caulocystidia absent.
Oleiferous hyphae present in both lamellar and pileus
trama. Clamp connections observed on all hyphae.
Habitat: on soil, among litter, solitary.
Material examined: INDIA, Idukki District, Munnar,
Eravikulam National Park, 6 July 2011, K.N. Anil Raj
AR683 (CAL 1385, holotype).
GenBank Number ITS:KY002062.
Notes: Entoloma magnum is characterized by a large,
tricholomatoid basidiocarp; a brown-coloured, umbonate
pileus; adnexed to narrowly adnate lamellae; a fleshy-fibrous stipe; isodiametric to sub-isodiametric basidiospores;
absence of cheilocystidia; a pileipellis that is a transition
between an ixocutis and an ixotrichoderm and the presence
of clamp connections on all hyphae. Owing to these
123
Fungal Diversity
Fig. 125 ITS-based phylogram generated from maximum likelihood
analysis using RAxML-HPC2 v. 8.2.8 (Stamatakis 2014) on XSEDE
platform employed in CIPRES Science Gateway web server (Miller
et al. 2010) depicting the placement of Entoloma magnum
(KY002062) within Entoloma subg. Entoloma. Bootstrap support
values C50% are indicated at the nodes. The tree is rooted with
Rhodocybe luteobrunnea, R. tugrulii, Clitopilus prunulus and
C. reticulosporus
characters, E. magnum fits into the section Entoloma of the
subgenus Entoloma (Noordeloos 2004; Co-David et al.
2009; Noordeloos and Gates 2012).
Entoloma brihadum Manim., A.V. Joseph & Leelav., a
species previously described from Kerala (Manimohan
et al. 1995), somewhat resembles E. magnum in having
robust, tricholomatoid basidiocarps with a subumbonate
pileus, crowded lamellae, a fertile lamella-edge, and
absence of caulocystidia. However, E. brihadum has a
smaller (50 mm), pinkish-white, glabrous pileus with pellucid striations, free to adnexed lamellae, a pinkish-white,
and glabrous stipe, smaller and exclusively quadrate
basidiospores (6–8 9 6–7 lm) and a cutis-type pileipellis
composed of hyaline hyphae.
123
Fungal Diversity
Inocybaceae Jülich
The type genus of the Inocybaceae, Inocybe, had traditionally been placed within the Cortinariaceae family
(Kirk et al. 2008, Singer 1986). Despite this, Jülich placed
the genus in its own family, the Inocybaceae (Jülich 1982).
Members of this family have a widespread distribution in
tropical and temperate areas (Cannon & Kirk 2007).
Fig. 126 Entoloma magnum (CAL 1385, holotype). Scale bar 20
mm (photo by K. N. Anil Raj)
Entoloma myochroum Noordel. & E. Ludw., a species
from Germany (Noordeloos 2004), is close to E. magnum
in having almost similar-sized basidiocarps, a concave and
glabrous pileus, adnate lamellae with concolorous edges, a
solid stipe with a fibrillose surface, almost similar-sized
basidiospores (7–10 9 6.5–8.5 lm) with 4–6 angles, a
fertile lamella-edge and clamped hyphae. However, that
species has a mouse-grey-coloured pileus, a flexuous or
irregularly shaped stipe, a cutis-type pileipellis made up of
hyphae with intracellular pigment and lamellar and pileus
trama with short inflated elements.
In a BLASTn search using the ITS sequence (628 bp)
derived from E. magnum, the closest hit was E. ochreoprunuloides (GenBank Number KC710092; 92% identity).
Entoloma ochreoprunuloides Morgado & Noordel., is a
species from Germany (Noordeloos 2004; Morgado et al.
2013), resembles E. magnum in having a similar-shaped,
brownish pileus with radially wrinkled and innately fibrillose surface, isodiametric basidiospores, a fertile-lamella
edge, and clamped hyphae. However, E. ochreoprunuloides differs from E. magnum owing to its smallersized and differently-coloured basidiocarps, emarginate
lamellae, smaller (5.9–7.1 9 5.7–7.2 lm) basidiospores
and hyphae of the pileipellis with intracellular pigment.
In the phylogram (Fig. 126) generated from the ML
analysis, E. magnum nested within a clade, which represents the subgenus Entoloma. Within this clade, E. magnum, E. luteobasis, E. ochreoprunuloides, E. madidum and
collections of E. bloxamii formed a subclade. Within this
subclade, E. magnum was differentiated as an independent
lineage separated from other species with significant (74%)
bootstrap support.
Inocybe (Fr.) Fr.
The genus Inocybe (Inocybaceae) is a species-rich genus
of Agaricales and is well-known for their ectomycorrhizal
ecology and toxicity of most species (Matheny 2009).
Matheny (2009) proposed seven major clades or lineages
consisting of Inocybe and its allies within the family Inocybaceae. Of the seven clades or lineages (Inocybe s. str.,
Nothocybe, Pseudosperma, Mallocybe, Inosperma, Auritella and Mallocybella), Auritella and Mallocybella were
formally recognised as distinct genera, Auritella and
Tubariomyces (Matheny and Bougher 2006a, b; Alvarado
et al. 2010). Species of Inocybe are characterised by mostly
brownish or rarely whitish basidiomata occasionally with a
purplish or lilac hue, a fibrillose-rimose or squamulose
pileus, brownish lamellae, brown spore-print, a fibrillosepruinose stipe at times with a distinct marginate-bulbous
base, a characteristic odour, smooth, warty, nodulose or
spinulose basidiospores and metuloidal cystidia often with
crystalloid deposits at their apices. Several species are
devoid of metuloidal cystidia and they are characterised by
abundant, thin-walled cheilocystidia (Matheny 2005;
Larsson et al. 2009). During the course of our studies on
this genus in Kerala State, India, we discovered several
new species of Inocybe. Three of these species belonging to
the Pseudosperma clade (Fig. 128) are described here.
Inocybe brunneosquamulosa K.P.D. Latha & Manim., sp.
nov.
MycoBank number: MB 816735; Facesoffungi number:
FoF: 2176, Figs. 129, 130
Etymology: referring to the brown squamules on the
pileus surface.
Holotype: CAL 1308
Basidiocarps small. Pileus 7–11 mm diam., convex with
a small umbo when young, becoming broadly convex still
with a small umbo at maturity; surface initially dark brown
(6F8/OAC636) on the squamules and brownish-orange
(6C4/OAC653) elsewhere, becoming dark brown (6F7/
OAC637) on the squamules, greyish-orange (6B3/
OAC633) on the fibrils and brownish-grey (6C2/OAC662)
elsewhere at maturity, appressed- to recurved-squamulose
all over when young, becoming appressed- to recurved
squamulose on and around the umbo and appressed-fibrillose and rimose towards the margin; margin incurved,
becoming decurved, crenate or wavy. Lamellae sinuate,
123
Fungal Diversity
Fig. 127 Entoloma magnum (CAL 1385, holotype). a Basidiospores. b Basidium. c Stipitipellis. d Pileipellis. Scale bars a, b = 10 lm; c,
d = 100 lm (photos by K. N. Anil Raj)
close, initially orange grey (6B2/OAC634), becoming
brownish-orange (6C4/OAC655) or light brown (6D4/
OAC686) at maturity, up to 1.5 mm wide, with lamellulae
of 1 length; edges fimbriate, whitish. Stipe 17–19 9 1–2
mm, central, terete, equal, cartilaginous, solid; surface
brownish-orange (6C3/OAC633) all over, appressed-fibrillose in most parts, slightly-recurved fibrillose and finely
pruinose towards the apex; base not enlarged. Odour and
taste not distinctive. Basidiospores 8–10 9 5–6.5 (9 ± 0.6
9 5.9 ± 0.4) lm, Q = 1.3–1.9, Qm = 1.5, smooth, ellipsoid to subphaseoliform, slightly thick-walled, yellowishbrown. Basidia 21–30 9 11–13 lm, clavate, thin-walled,
hyaline, 4-spored; sterigmata up to 4.5 lm long. Pleurocystidia absent. Lamella-edge heterogeneous. Cheilocystidia 17–39 9 11–18 lm, versiform: clavate, utriform,
fusiform, cylindrical with an obtuse apex, occasionally
subglobose or rarely pedicellate or septate, hyaline with
faint hyaline encrustations, thin- to slightly thick-walled.
Lamellar trama subregular; hyphae 6–15 lm wide, thin-
123
walled, hyaline or pale yellow. Subhymenium pseudoparenchymatous. Pileus trama subregular, composed of
both narrow and inflated hyphae; hyphae 3–30 lm wide,
pale yellow, thin-walled. Pileipellis a cutis frequently
disrupted with trichodermal patches, often a perfect trichoderm at the centre; hyphae 8–16 lm wide, thin- to
slightly thick-walled, with a brown wall pigment and
dense, yellowish-brown or brown spiral encrustations;
terminal cells 25–52 9 7–11 lm, clavate or cylindrical
with an obtuse apex, thin- to slightly thick-walled. Stipitipellis a cutis often disrupted by loose hyphal projections
scattered over the entire surface of the stipe and with
bunches of caulocystidia confined to the extreme stipe
apex; hyphae 5–11 lm wide, thin- to slightly thick-walled,
with a pale yellowish-brown wall pigment and faint hyaline
encrustations; terminal cells, 21–49 9 5–7 lm, cylindrical
or flexuous-cylindric, slightly thick-walled, with a pale
yellowish-brown wall pigment and faint hyaline encrustations. Caulocystidia 22–50 9 11–14 lm, catenulate,
Fungal Diversity
Fig. 128 Phylogram generated from maximum likelihood (RAxML)
analysis based on RPB2 sequence data matrix for 34 Inocybe species.
Sequences of Inocybe species belonging to the Pseudosperma clade
used in this study have been selected from a previous analysis of
Kropp et al. (2013). Values at nodes indicate bootstrap support. BS
values C50% are shown. Inocybe luteobrunnea, I. brunneosquamulosa
and I. rubrobrunnea are in pink to highlight its phylogenetic position
in the tree. The tree is rooted with I. adaequata and I. calamistrata of
Inosperma clade
123
Fungal Diversity
Fig. 129 Inocybe brunneosquamulosa (CAL 1308, holotype). a, b Basidiocarps. Scale bars a, b = 5 mm (photos by K. P. Deepna Latha)
clavate, inflated clavate or obovoid, rarely septate, hyaline,
occasionally with faint, hyaline encrustations, thin- to
slightly thick-walled. Stipe trama hyphae with dense, yellowish-brown oleaginous contents. Clamp connections
seen on all hyphae.
Habitat: on the ground, scattered around Vateria indica
(Dipterocarpaceae) trees.
Specimen examined: INDIA, Kerala State, Ernakulam
District, Kochi, Thevakkal, Ponnakkudam Kavu sacred
grove, 25 August 2014, K.P.D Latha DKP264 (CAL 1308,
holotype).
GenBank Numbers ITS: KX073582; LSU: KX073586;
RPB2: KX073589.
Notes: Small basidiocarps with a dark brown, squamulose and fibrillose-rimose pileus; a fibrillose stipe with a
finely pruinose apex and an abruptly ending base; smooth,
ellipsoid to subphaseoliform basidiospores; a hymenium
devoid of pleurocystidia; versiform cheilocystidia with
occasional, faint, hyaline encrustations; a cutis-type
pileipellis which is disrupted with trichodermal patches and
a cutis-type stipitipellis disrupted by loose, hyphal projections and often with caulocystidia at the extreme stipe
apex are the salient features of I. brunneosquamulosa.
Inocybe fuscospinulosa, a species originally described from
Indonesia (Horak 1980b) and also reported from Sri Lanka
123
(Pegler 1986), seems to be somewhat similar to I. brunneosquamulosa in having a pileus of rather similar colour
and surface features, a fimbriate lamella-edge, a fibrillose
stipe, a hymenium devoid of pleurocystidia, a trichodermtype pileipellis and the presence of cheilocystidia. Inocybe
fuscospinulosa, however, is distinguished from the
I. brunneosquamulosa in having larger basidiocarps with a
densely squamulose pileus, crowded, adnexed, tobacco
brown lamellae, a reddish-brown tinted stipe with occasional scales, smaller (6.5–8 9 4–5 lm) and ovoid basidiospores, cylindric to subfusoid cheilocystidia devoid of
encrustations and a stipitipellis lacking caulocystidia. Inocybe brunneosquamulosa is also somewhat similar to
I. umbrinovirens E. Horak, a species so far known only
from Papua New Guinea (Horak 1980b), in having a
somewhat similar-coloured pileus with almost similar
surface features, a fibrillose stipe, smooth basidiospores, a
hymenium devoid of pleurocystidia, the presence of
cheilocystidia with encrusting pigment and a trichodermtype pileipellis. However, the characters such as the larger
basidiocarps with differently-shaped pileus, chocolate
brown, adnexed, crowded lamellae, a hollow stipe with a
greenish base, larger (10–12.5 9 7–8.5 lm) and ovoid
basidiospores, larger basidia, cheilocystidia that are terminal elements of lamellar trama, the absence of
Fungal Diversity
Fig. 130 Inocybe brunneosquamulosa (CAL 1308, holotype). a Basidiospores. b Basidium. c Cheilocystidia. d Stipitipellis. e Caulocystidia.
f Pileipellis. Scale bars a–e = 10 lm, f = 100 lm (photos by K. P. Deepna Latha)
caulocystidia and a strong odour make I. umbrinovirens
different from I. brunneosquamulosa.
Inocybe squamata J.E. Lange, a species widespread in
Europe and USA (Cripps 1997) and also reported from
Kerala (Pradeep and Vrinda 2010; Mohanan 2011) is
similar to I. brunneosquamulosa in having a pileus of
somewhat similar texture, rather similarly attached
lamellae, a fibrillose stipe, subphaseoliform basidiospores,
a hymenium devoid of pleurocystidia, the presence of
cheilo- and caulocystidia and similar type of pileipellis.
However, I. squamata has larger basidiocarps with a
differently-coloured pileus, thick, broad, yellow brown
lamellae, a longer, white stipe, larger basidiospores
(9–11.5 (13) 9 (5) 5.5–6.5 lm), occasional 2-spored
basidia and narrowly clavate cheilo- and caulocystidia
lacking encrustations.
123
Fungal Diversity
Comparison of the ITS (679 bp), LSU (935 bp) and
RPB2 (699 bp) sequences derived from I. brunneosquamulosa with the nucleotide sequences available in GenBank Numbers showed that I. brunneosquamulosa has
distinct sequences. Inocybe species MCA562 resulted as
the closest hit in megablast searches with ITS (GenBank
Numbers JQ408785; Identities = 630/682 (92%)), LSU
(GenBank Numbers JN975016; Identities = 921/936
(98%)) and RPB2 (GenBank Numbers JQ421077; Identities = 673/699 (96%)) sequences. Inocybe species
MCA562 is a collection from Japan, but its morphological
features are unavailable for comparison as it remains
unpublished.
The phylogenetic placement of I. brunneosquamulosa is
shown in the phylogram (Fig. 128) generated from the ML
analysis of RPB2 sequence data matrix. In the ML analysis,
I. brunneosquamulosa nested in the Pseudosperma clade
with maximum support (100% ML) where it paired with
Inocybe species MCA562 and had a strong support (96%
ML).
Inocybe luteobrunnea K.P.D. Latha & Manim., sp. nov.
MycoBank number: MB 816734; Facesoffungi number:
FoF 2177; Figs. 131, 132
Etymology: referring to the yellowish-brown pileus.
Holotype: CAL 1260
Basidiocarps small. Pileus 6–14 mm diam., narrowly
conical when very young, becoming conico-convex and
finally convex with a small umbo; surface brown (6F6/
OAC636) on the squamules and yellowish-brown (5D8/
OAC775) elsewhere when young, becoming dark brown
(6F7/OAC639) at the centre and on the squamules and
brownish-orange (5C4, 5C5/OAC806) elsewhere at maturity, with appressed- to slightly recurved, minute squamules on and around the umbo, appressed-fibrillose
towards the margin; margin incurved when young,
becoming decurved to somewhat straight with age, crenate
or somewhat wavy, finely fissile. Lamellae emarginate,
subventricose, rarely furcate, close, greyish-orange (5B3,
5B4/OAC793), up to 2 mm wide, with lamellulae of 3
lengths; edges fimbriate, rather whitish. Stipe 13–22 9
2–2.5 mm, central, equal, fistulose; surface initially orange
grey (6B2/OAC634), becoming greyish-orange (5B2/
OAC675) at maturity, appressed-fibrillose all over, finely
pruinose towards the apex; base somewhat bulbous, not
marginate-bulbous. Odour and taste not distinctive.
Basidiospores 7–8 (9) 9 5–6 (6.5) (7.9 ± 0.7 9 5.9 ±
0.5) lm, Q = 1.2–1.6, Qm = 1.4, smooth, ovoid to
amygdaliform, slightly thick-walled, pale yellowishbrown. Basidia 23–34 9 8–10 lm, clavate, thin-walled,
Fig. 131 Inocybe luteobrunnea (CAL 1260, holotype). a, b Basidiocarp in the field. Scale bars a, b = 5 mm (photos by K. P. Deepna Latha)
123
Fungal Diversity
Fig. 132 Inocybe luteobrunnea (CAL 1260, holotype). a Basidiospores. b Basidium. c Cheilocystidia. d Caulocystidia. e Pileipellis. Scale bars
a–d = 10 lm, e = 100 lm (photos by K. P. Deepna Latha)
hyaline, 4-spored or rarely 2-spored; sterigmata up to 5 lm
long. Mature basidia slightly projecting beyond the
hymenial surface. Pleurocystidia absent. Lamella-edge
sterile with abundant cheilocystidia. Cheilocystidia 17–52
9 9–15 lm, versiform: clavate, inflated clavate, cylindrical
or cylindrical with apical constriction, cylindro-flexuous,
cylindrical with irregular constriction, narrowly utriform,
utriform with a median constriction, ovoid, fusiform,
submoniliform or short-pedicellate, often septate, thin- to
slightly thick-walled, hyaline. Lamellar trama subregular,
composed of both narrow and inflated hyphae; hyphae
3–25 lm wide, thin- to slightly thick-walled, hyaline, at
times with faint, hyaline encrustations especially towards
the edge of the hymenium. Subhymenium poorly developed. Pileus trama subregular; hyphae 9–28 lm wide, pale
yellow, thin- to slightly thick-walled. Pileipellis a cutis
123
Fungal Diversity
often transitioning to a trichoderm towards the centre;
hyphae 4–11 lm wide, tangled, slightly thick-walled, with
a pale-yellow wall pigment and brown spiral encrustations.
Stipitipellis a cutis frequently disrupted with bunches of
caulocystidia towards the apex; hyphae 5–12 lm wide,
thin-to slightly thick-walled, with a pale-yellow wall pigment and hyaline encrustations, some hyphae with dense,
yellowish-brown, amorphous contents towards the base.
Caulocystidia 13–74 9 7–13 lm, versiform: clavate, narrowly clavate, moniliform, submoniliform, cylindricoflexuous, cylindrical with an obtuse apex, utriform with a
median constriction, cylindrical or ovoid with a rostrate
apex, often septate, thin- to slightly thick-walled, hyaline.
Clamp connections observed on all hyphae.
Habitat: scattered among bryophytes, on a mud wall.
Material examined: INDIA, Kerala State, Idukki District, Munnar, on the way to Mattupetti top hill station, 31
August 2013, K. P. Deepna Latha DKP167 (CAL 1260,
holotype); GenBank Numbers ITS: KX073580; LSU:
KX073584; RPB2: KX073588; INDIA, Kerala State,
Idukki District, Munnar, on the way to Mattupetti top hill
station, 9 November 2013, K. P. Deepna Latha DKP251
(CAL 1260, holotype; CAL 1261, paratype).
GenBank Numbers ITS: KX073581; LSU: KX073585.
Notes: Inocybe luteobrunnea is characterised by a yellowish-brown pileus with a fibrillose to minutely squamulose surface; a fibrillose stipe with a finely pruinose
apex and a bulbous base; smooth, ovoid to amygdaliform
basidiospores; a hymenium lacking pleurocystidia; abundant, versiform cheilocystidia; subregular lamellar trama
with faint hyaline encrustations; a pileipellis that is a cutis
with a transition to a trichoderm towards the centre and a
cutis-type stipitipellis frequently disrupted with bunches of
caulocystidia confined to the stipe apex. Inocybe
palaeotropica E. Turnbull & Watling, a widespread species
reported from Singapore, Malaysia, Sabah, Papua New
Guinea, Solomon Islands (Horak 1980b) and also recorded
from Kerala by Vrinda et al. (1997, as I. umbrina Massee),
shares a few characters with I. luteobrunnea such as a
similar-shaped
pileus,
similar-sized
basidiospores
(7.5–10.5 9 4.5–6 lm), a sterile lamella-edge and caulocystidia on the stipe apex. Inocybe palaeotropica, however,
has larger basidiocarps with dissimilar colour and surface
features, adnexed lamellae, a fibrillose stipe devoid of
apical pruinosity, ovoid to short ellipsoid basidiospores,
only 4-spored basidia, clavate to cylindro-clavate cheilocystidia, hyphae of lamellar trama lacking encrustations, a
cutis-type pileipellis and smaller, cylindro-clavate caulocystidia. Inocybe luteobrunnea is somewhat similar to
I. fuscospinulosa Corner & E. Horak, an Indonesian species
(Horak 1980b) and also reported from Sri Lanka (Pegler
1986), in having a somewhat similar-shaped pileus, a fibrillose stipe, almost similar-sized basidiospores (6.5–8 9
123
4–5 lm), cheilocystidia, a hymenium devoid of pleurocystidia and an almost similar pileipellis structure. However, I. fuscospinulosa differs from I. luteobrunnea owing
to its larger basidiocarps with conspicuous, erect, spiny
squamules on the pileus, crowded, adnexed, tobacco-brown
lamellae, a red-brown tinged stipe with occasional brown
squamules and without a distinct bulbous base, ovoid
basidiospores, consistently 4-spored basidia, larger, differently-shaped cheilocystidia and a stipitipellis devoid of
caulocystidia.
The distinctive status of (CAL 1260: 675 bp; CAL 1261:
674 bp), LSU (CAL 1260: 838 bp; CAL 1261: 746 bp) and
RPB2 (CAL 1260: 651 bp) sequences of I. luteobrunnea
was confirmed in the BLASTn searches. An unnamed
Australian species, Inocybe species AU95 was the closest
hit in a megablast search for ITS (GenBank Numbers
KP636851; Identities = 540/588 (92%)), LSU (GenBank
Number KP171053; Identities = 831/839 (99%)) and
RPB2 (GenBank Number KM555145; Identities = 638/
651 (98%)) sequences.
The phylogram generated from the Maximum Likelihood (ML) analysis (Fig. 129) depicts the relative placement of I. luteobrunnea. The ML analysis placed
I. luteobrunnea in the Pseudosperma clade with full support (100% ML) based on RPB2 sequence data matrix.
Within this clade, I. luteobrunnea clustered with an
unnamed Australian species, Inocybe species AU95
(KM555145) with maximum support (100% ML). The
macro-morphological and microscopic data of that species
are not available for comparison.
Inocybe rubrobrunnea K.P.D. Latha & Manim., sp. nov.
MycoBank number: MB 816736; Facesoffungi number:
FoF: 2175; Figs. 133, 134
Etymology: referring to the reddish-brown colour of the
pileus.
Holotype: CAL 1307
Basidiocarps small. Pileus 6–12 mm diam., somewhat
hemispherical or paraboloid when young, becoming convex or plano-convex with a small umbo; surface reddishbrown (8E5, 8E6/OAC609, OAC610) when young,
becoming dark brown (7F7/OAC621) at the centre and on
the squamules, orange grey (6B2) elsewhere, appressed- to
slightly recurved squamulose and appressed-fibrillose all
over when young, becoming appressed- to recurved squamulose on and around the centre, appressed-fibrillose and
rimulose towards the margin; margin initially incurved,
becoming decurved to somewhat straight with age, crenate.
Lamellae sinuate or emarginate, close, initially greyishorange (6B3/OAC654), becoming brownish-orange (6C4/
OAC695), up to 1.5 mm wide, with lamellulae of 3 lengths;
edges fimbriate, whitish. Stipe 11–20 9 1–2 mm, central,
terete, equal or slightly tapered towards the base,
Fungal Diversity
Fig. 133 Inocybe rubrobrunnea (CAL 1307, holotype). a, b Basidiocarp in the field. Scale bars a, b = 5 mm (photos by K. P. Deepna Latha)
cartilaginous, solid; surface brownish-orange (6C3/
OAC633), fading towards the apex, appressed-fibrillose all
over, slightly pruinose at the apex; base slightly enlarged.
Odour and taste not distinctive.
Basidiospores 7–9 9 5–6 (8.32 ± 0.63 9 5.57 ±
0.43) lm, Q = 1.2–1.7, Qm = 1.4, smooth, ellipsoid to
slightly phaseoliform, slightly thick-walled, pale yellowish-brown. Basidia 18–27 9 9–12 lm, clavate, thinwalled, hyaline, 4-spored; sterigmata up to 3 lm long.
Pleurocystidia absent. Lamella-edge heterogeneous.
Cheilocystidia 17–44 9 6–15 lm, abundant, versiform:
narrowly clavate, clavate, cylindrical, oblong, ellipsoid,
fusiform, obovoid, cylindrical with a median constriction
or with a subcapitate apex, occasionally septate, exuding
some amorphous material at the apex, slightly thickwalled (up to 1 lm thick), hyaline or rarely with pale
yellowish-brown, amorphous contents, occasionally with
faint, hyaline encrustations. Lamellar trama subregular,
composed of both narrow and inflated hyphae; hyphae
3–23 lm wide, thin- to slightly thick-walled, hyaline.
Subhymenium pseudoparenchymatous. Pileus trama subregular; hyphae 6–27 lm wide, hyaline or pale yellow,
thin-walled. Pileipellis a cutis often disrupted with bundles of tangled ascending hyphae towards the centre;
hyphae 5–10 lm wide, with a pale brownish-yellow wall
pigment and brown spiral encrustations. Stipitipellis a
cutis composed of hyaline or pale yellow hyphae (3–10
lm wide), devoid of encrustations, frequently disrupted
by bundles of loose, tangled hyphae towards the apex
(2–6 lm wide), with a pale-yellow wall pigment and
pale yellow spiral encrustations, thin- to slightly thickwalled, occasionally with yellowish-brown amorphous
contents. Caulocystidia absent. Clamp connections
observed on all hyphae.
Habitat: on soil, solitary or in small groups, near Hopea
ponga (Dipterocarpaceae) trees.
Specimen examined: INDIA, Kerala State, Wayanad
District, Muthanga, Muthanga Wildlife Sanctuary, 21
August 2013, K. P. Deepna Latha DKP142 (CAL 1307,
holotype).
GenBank Numbers ITS:KX073583; LSU:KX073587;
RPB2:KX073590.
Notes: Small basidiocarps with an appressed-fibrillose to
squamulose pileus; a stipe with a slightly enlarged base and
a fibrillose surface; smooth, ellipsoid to slightly phaseoliform basidiospores; a hymenium devoid of pleurocystidia; versiform cheilocystidia exuding some amorphous
material at the apex; a cutis-type pileipellis disrupted with
bundles of tangled ascending hyphae; and a cutis-type
stipitipellis often disrupted by bundles of loose, tangled
123
Fungal Diversity
Fig. 134 Inocybe rubrobrunnea (CAL 1307, holotype). a Basidiospores. b Basidium. c Cheilocystidia. d Stipitipellis. e Caulocystidia.
f Pileipellis. Scale bars a–c = 10 lm, d, e = 100 lm (photos by K. P. Deepna Latha)
hyphae towards the apex are the diagnostic features of
I. rubrobrunnea.
Following the key of Kobayashi (2002), I. rubrobrunnea
keys out close to I. quercina Hongo, a species known from
Japan, because of its reddish-brown pileus with almost
similar size, shape and surface features, a whitish lamellaedge, a solid stipe with a fibrillose surface, almost similarshaped basidiospores, the absence of pleurocystidia and the
123
presence of cheilocystidia. However, the characters such as
the adnexed lamellae that become red when cut, a longer
stipe, larger basidiospores (8.2–10.8 9 4.8–5.5 lm) and
basidia, smaller cheilocystidia devoid of encrustations, a
cutis-type pileipellis lacking encrustations, the presence of
caulocystidia and a strong odour of that species make it
different from I. rubrobrunnea. Inocybe fuscospinulosa, a
species originally described from Indonesia (Horak 1980b)
Fungal Diversity
and also reported from Sri Lanka (Pegler 1986), has a
pileus with dark brown squamules, a whitish lamella-edge,
basidiospores of similar size (7–9 9 4.2–5.5 lm) and shape
and the absence of pleuro- and caulocystidia. However,
that species differs in having larger basidiocarps with a
campanulate to applanate pileus, a white context discolouring purplish-red on exposure, cheilocystidia devoid
of encrustations and a trichoderm-type pileipellis.
Comparison of the ITS (676 bp), LSU (924 bp) and
RPB2 (702 bp) sequences of I. rubrobrunnea with the
nucleotide sequences of taxa available in GenBank
Numbers suggests that it has distinct sequences. In a
megablast search of the GenBank Numbers database
using ITS sequence of I. rubrobrunnea, the closest hit
was Inocybe species MCA562 (GenBank Numbers
JQ408785; Identities = 616/668 (92%)) followed by
Inocybe species AU43 (GenBank Numbers KJ729878;
Identities = 605/659 (92%)). An undescribed Inocybe
species, Inocybe species AU44 was the closest hit in
BLASTn search with LSU (GenBank Numbers
KJ729906; Identities = 896/925 (97%)) sequence. Inocybe species AU43 (GenBank Numbers KJ729935; Identities = 670/702 (95%)) resulted as the closest hit in
BLASTn search with RPB2 sequences. But, the details of
Inocybe species MCA562, an unnamed Inocybe collection
from Japan and Inocybe species AU43 and Inocybe species AU44, another unnamed collection from Australia,
are not available for comparison.
The RPB2-based ML phylogeny (Fig. 128) placed Inocybe rubrobrunnea in the Pseudosperma clade with full
support (100% ML). Within this clade, I. rubrobrunnea
clustered with Inocybe species AU43 with a significant
support (95% ML).
Marasmiaceae Roze ex Kühner.
The family Marasmiaceae is characterized by white
spores. The members of this family mostly have tough
stems and the capability of shrivelling up during a dry
period and later recovering. According to Kirk et al.
(2008), the family contains 54 genera and 1590 species.
Marasmius Fr.
Marasmius is a genus of mushroom-forming fungi in the
family Marasmiaceae. It contains about 500 species (Kirk
et al. 2008) of which a few, such as Marasmius oreades,
are edible. However, most members of this genus are small,
unimpressive, brown mushrooms. Their humble appearance contributes to them not being readily noticeable to the
layman, and therefore these mushrooms are seldom collected by mushroom hunters. Quite a few of the species are
known to grow in the characteristic fairy ring pattern
(Fig. 135).
Marasmius luculentus A.K. Dutta, K. Acharya & Antonı́n,
sp. nov.
MycoBank number: MB 816959; Facesoffungi number:
FoF 2192, Figs. 136, 137
Etymology: referring to the beautiful (luculentus)
appearance of the pileus.
Holotype: CUH AM120
Pileus 5–10 mm in diam., conic to hemispherical when
very young, becoming convex in age, with a small conic to
convex papilla that ranges from reddish-brown (8D6) to
violet brown (10E5–6, 11E6–7) when young, but later
turning brown (7D7, 7E8) to dark brown (7F7) at maturity,
very rarely forming a central dot, often with a depression
around the papilla; striate to plicate up to center, margin
often crenate; surface dry, glabrous, hygrophanous,
brownish-orange (7C4) to light brown (7D5–6) when very
young, later reddish-grey (7–8B2) and finally turning white
(1A1) to off-white at maturity, greyish-yellow (1–2B4,
4B4–5) on drying; context very thin, white to cream.
Lamellae adnexed or adnate to a collarium, distant to
subdistant (L = 12–14, l = 0), white (1A1), regular,
slightly intervenose, edge concolorous. Stipe 5–9(–12) 9
0.1–0.3 mm, central, glabrous, wiry, pliant, cylindric,
equal, simple and insititious on the substratum, dark
brownish-black to black overall, often accompanied by
black rhizomorphs, 30–42 mm long. Odour and taste
indistinct. Spore print white.
Basidiospores (9.3–)10–10.3–10.5(–11) 9 (3.9–)
4.3–4.8–5.4(–5.8) lm, Q = 1.7–2.2–2.6, ellipsoid to
ellipsoid-fusoid, smooth, hyaline, IKI-, thin-walled. Basidia not observed. Basidioles 24–25 9 5.5–7 lm, fusoid to
clavate, hyaline, thin-walled. Lamellae edge sterile. Pleurocystidia absent. Cheilocystidia composed of Siccus-type
broom cells; main body 14–18 9 4–6(–7.5) lm,
sphaeropedunculate to (broadly) clavate or irregular in
outline, hyaline, apically thick-walled; apical setulae
2–3.5(–5.5) lm long, cylindrical or irregular in outline,
obtuse, pale yellow to light brown in KOH, thick-walled.
Pileipellis a hymeniform layer, composed of Siccus-type
broom cells; main body (17–)19–23(–25) 9 (7–)8–11
(–14.5) lm, sphaeropedunculate to clavate, broadly clavate, often branched, pale yellow to light brownish in
KOH, thick-walled; apical setulae (1.5–)3–4(–5.5) lm
long, cylindrical, obtuse, thick-walled. Pileus trama composed of 4.5–7.5 lm broad, interwoven, cylindrical, hyaline, inamyloid, thin-walled hyphae. Lamellar trama
hyphae 4–6.5 lm broad, interwoven, cylindrical, hyaline,
inamyloid, thin-walled. Stipitipellis hyphae 5.5–6.5 lm
broad, parallel to subparallel, cylindrical, smooth, non-gelatinous, hyaline to pale yellow in KOH, non-dextrinoid to
weakly dextrinoid, thick-walled. Stipe trama hyphae
6.8–7.5 lm broad, parallel to subparallel, cylindrical,
123
Fungal Diversity
Fig. 135 Phylogram generated from maximum likelihood (RAxML)
analysis using a GTR ? I ? G model of nucleotide evolution based on
ITS for 55 Marasmius and two outgroup sequences (Crinipellis
brunneipurpurea and C. malesiana). Maximum likelihood bootstrap
support values greater than 50% and Bayesian posterior probabilities
greater than 0.5 are indicated above or below the nodes. Numbers to
the left of/are ML bootstrap percentages, and those to the right are
Bayesian posterior probabilities. Sequences used in this study mostly
have been sampled from a previous study (Tan et al. 2009; Wannathes
et al. 2009). The newly generated sequences are in blue, and the
newly described taxa is placed in bold font to highlight its
phylogenetic position in the tree. GenBank accession numbers for
all of the sequences are indicated in the tree. G—sect. Globulares;
L—sect. Leveilleani; MM—sect. Marasmius subsect. Marasmius;
MS—sect. Marasmius subsect. Sicciformes; N—sect. Neosessiles;
SA—sect. Sicci ser. Atrorubentes; SH—sect. Sicci ser. Haematocephali; SL—sect. Sicci ser. Leonini; and SS—sect. Sicci ser.
Spinulosi
Fig. 136 Marasmius luculentus (CUH AM120, holotype). Habit
(AKD 100/2015) (photo A.K. Dutta)
smooth, non-gelatinous, hyaline, inamyloid, thin- to thickwalled. Caulocystidia absent. Clamp connections present in
all tissues.
Material examined: INDIA, West Bengal, North-24parganas, Barasat, Berunanpukhuria, N22°440 22.7832’’,
E88°260 25.2384’’, 10.809 m alt., on dried leaves of Bambusa bambos (L.) Voss plant, A.K. Dutta & S. Paloi, 25
June 2015, AKD 100/2015 (CUH AM120, holotype).
INDIA, West Bengal, North-24-parganas, Barasat,
Berunanpukhuria, N 22 440 21.642’’, E 88 260 26.6676’’, ca
10.747 m alt., on dried leaves of B. bambos plant, A.K.
Dutta & S. Paloi, 29 June 2015, AKD 132/2015 (CUH
AM125, paratype).
GenBank Numbers CUH AM120 ITS: KX138604;
LSU:KX138606. CUH AM125 ITS:KX138605.
123
Fungal Diversity
Fig. 137 Marasmius
luculentus (CUH AM120,
holotype). a Basidiomata.
b Basidiospores. c Siccus-type
cheilocystidia. d Siccus-type
cells of the pileipellis. Scale
bars a = 5 mm, b–d = 5 lm.
(drawings A.K. Dutta)
Notes: Distinctive features of M. luculentus include a
small basidiomes with a convex, plicate, light brown to
reddish-grey pileus when young that later turns into white
to off-white, with a reddish-brown to brown or dark brown
central papilla, collariate and distant to subdistant, often
intervenose lamellae (12–14), a small central, insititious
stipe arising directly from the substratum, obtuse apical
setulae of the pileipellis broom cells and cheilocystidia of
the Siccus-type, and moderately-sized basidiospores (10.3
9 4.8 lm; mean Q = 2.2). These combinations of macroand micro-morphological features easily categorizes the
present specimen belonging to sect. Marasmius and subsect. Sicciformes Antonı́n (Wannathes et al. 2009).
Being a well representative of sect. Marasmius, the new
Indian species is morphologically similar to several other
species. Marasmius conicopapillatus Henn., described
from Cameroon for the first time (Hennings 1895) and later
subsequently reported to occur in other African countries
(Antonı́n 2007), Bolivia (Singer 1976), Uganda (Pegler
1977) and Java (Desjardin et al. 2000) differs by its much
smaller pileus (1–7 mm diam.) coloured pure white to offwhite when young that turns cream-buff at maturity, nonintervenose lamellae, and a considerably larger stipe
(12–30 mm long, Desjardin et al. 2000; 22–35 mm long,
Pegler 1977) coloured white towards apex. Marasmius
pallenticeps Singer, originally described from Argentina,
has a much smaller pileus (1–2 mm diam.), a copper brown
to reddish-brown coloured stipe that arise directly from the
rhizomorphs, and distinctly smaller basidiospores (7–8.5 9
2.7–4.5 lm; Singer 1976). Marasmius chrysochaetes Berk.
& M.A. Curtis differs by its much smaller pileus (1.2–2
mm), a longer stipe (22–25 mm long), and differently sized
basidiospores (9–10 9 3–3.5 lm, Dennis 1951; 8–9 9
3.5–4.2 lm, Singer 1976). Marasmius stypinoides Petch,
described from Hakgala, differs by its much smaller (up to
5 mm diam.) and minutely rugulose pileus, a longitudinally
striate and initially white stipe, and a serrate lamellae edge
(Petch 1948). Marasmius gracilichorda Corner has smaller
basidiomes with a minutely velutinous pileus surface
coloured brownish-orange to dark brown overall, yellowish-white lamellae with brownish edges (Tan et al. 2009).
Among other species that grows on bamboo leaves,
Marasmius kuthubutheenii Y.S. Tan, Desjardin, Vikineswary & Noorlidah differs by its much smaller (1–3 mm)
pileus with a minutely pruinose surface coloured brownishorange to light brown overall and a very smaller stipe (2–4
mm long; Tan et al. 2009). As revealed in Fig. 136,
M. luculentus appears to be phylogenetically close to
several other taxa like M. nigrobrunneus (Pat.) Sacc.,
M. ruforotula Singer, and M. subruforotula Singer.
Marasmius nigrobrunneus, originally described from
Vietnam and previously reported from India, differs by its
123
Fungal Diversity
smaller pileus (1–4 mm diam.) coloured greyish-black to
brown black with slightly paler striae, a long slender stipe
(up to 73 mm long), and somewhat differently sized
basidiospores (Singer 1976; Manimohan and Leelavathy
1989; Antonı́n and Buyck 2006; Antonı́n 2007). Marasmius ruforotula has a greyish-orange to brownish-orange
pileus with a black papilla, and yellowish-white lamellae,
and presence of numerous black rhizomorphs (Tan et al.
2009). Marasmius subruforotula has a minutely velutinous
pileus surface coloured greyish-brown to orange brown
often with white striae, discoloured and pale orange
coloured lamellae edge, slightly smaller basidiospores with
a mean of 9.5 9 4.5 lm (Q mean 2.1), and habitat on
dicotyledonous leaves or on wood (Pegler 1977; Antonı́n
2007; Wannathes et al. 2009).
The new species is macro-morphologically similar to
M. capillaris, M. limosus, M. rotalis, and M. rotula, but the
latter have cheilocystidia and pileipellis broom cells of the
Rotalis-type, and belong in subsect. Marasmius (Petch
1948; Gilliam 1976; Singer 1976; Pegler 1977, 1986;
Desjardin 1989; Bas et al. 1995; Corner 1996).
Mycenaceae Overeem
The Mycenaceae is a family of fungi in the order
Agaricales which the members are saprobic, have a cosmopolitan distribution, and are found in almost all ecological zones. According to Kirk et al. (2008), the family
contains 10 genera and 705 species. This is one of the
several families that were separated from the Tricholomataceae as a result of phylogenetic analyses.
Favolaschia (Pat.) Pat.
Favolaschia is a genus of mushrooms in the family
Mycenaceae. The genus has a widespread distribution, and
contains about 50 species worldwide (Kirk et al. 2008)
(Fig. 138).
Favolaschia auriscalpium (Mont.) Henn., Botanische
Jahrbücher für Systematik Pflanzengeschichte und
Pflanzengeographie 22: 93 (1895)
Facesoffungi number: FoF 3138, Figs. 139, 140
Pileus small, orange, becoming yellow and ultimately
dirty pallid when dry, orbicular in outline, convex, glabrous. Pores concolorous or yellowish, some irregularly
elongated, dried 0.2–0.4 mm wide, frequently hexagonal.
Pseudostipe concolorous with the pileus, laterally attached
to the pileus, subvelutinous, insititious, up to 1.8 9 0.3
mm. Basidiospores 8–11 9 5–7 lm, ellipsoid, sometimes
slightly constricted in the middle or reniform, smooth,
amyloid. Basidia 23–26 9 7–8 lm, 4–spored, rarely 2–
spored, mostly clavate or constriction in the middle.
Gloeocystidia few or none. Hyphae filamentous, hyaline,
truly gelatinized, thin walled in parts of the trama, non-
123
gelatinized and often broader and somewhat thick-walled
in other parts, predominantly in the pseudostipe, all hyphae
inamyloid, and clamp connections present. Gloeocystidia
21–35 9 9–22 lm, mostly clavate or ventricose to subvesiculose, rarely cylindrical, hyaline to yellowish. Dermatocystidia 19–29 9 6–14 lm, hyaline, thin-walled,
erect, subvesiculose or broadly ventricose, deep purplishpink in Cresyl blue mounts, obtuse or rarely somewhat
mucronate, crowded at the epicutis, otherwise few.
Habit, habitat and distribution: as a group on dead
wood, inflorescences, leaf sheaths or petioles of Palmae,
May–September. Our collection was collected on a
decaying wood near Hanthana Mountains Range,
Peradeniya.
Specimens examined: SRI LANKA, Kandy District,
Hanthana Mountains Range, 30 June 2012, Samantha C.
Karunarathna (MFLU 12-1891, new record).
GenBank Number ITS:KY649461.
Notes: This species was first described from Gulf Coast
of South America and this is the first report of F. auriscalpium with the molecular phylogenetic confirmation
from Sri Lanka. Favolaschia auriscalpium is morphologically closely related to F. calocera R. Heim. but F. calocera differs by having 2 spored, clavate, 38–40 9 7–10 lm
size basidia whereas F. auriscalpium has 4 spored, clavate
23–26 9 7–8 lm size basidia. Phylogenetic analysis
(Fig. 138) also supports to place the species in F.
auriscalpium.
Favolaschia manipularis (Berk.) Teng, Zhong Guo De
Zhen Jun [Fungi of China]: 760 (1963)
Facesoffungi number: FoF 3139, Figs. 141, 142
Pileus 0.3–2.5 cm in diam., conico-campanulate to
convex with conical umbo, translucently reticulate,
hygrophanous, white pruinose, whitish, when moist greyish
to hyaline, when dried up purely white to yellowish. Hymenophore tubular, adnate or adnate-emarginate with a
slightly decurrent tooth. 1–5 mm long, white, arranged as
radial rows, 5–7 in a row, with angular-round pores of
0.5–1.5 mm wide. Stipe 20–65 9 0.5–3 mm, hollow,
hyaline to white, completely white, pruinose cylindrical,
thickened in the base. Odour not distinct. Basidiospores
6–8 9 4–5 lm, Q = 1.2–1.6, smooth, white, ellipsoid to
broadly ellipsoid and amyloid. Basidia 16–18 9 6–8 lm,
narrowly clavate, clamped, with up to 6 lm long sterigmata. Cheilocystidia 51–75 9 7–13 lm, sterile lamellae
edge, lageniform, subcylindrical or subclavate, mostly
diverticulate or irregularly branched in the apex, simple
lageniform with more or less developed neck. Pleurocystidia not visible. Hyphae of the pileipellis 5–16 lm wide,
clamped, with diverticulate terminal cells. Pileocystidia
25–34 9 8–11 lm, abundant, irregularly shaped, lageniform to subclavate, with excrescences. Hyphae of the
Fungal Diversity
Fig. 138 Phylogenetic
relationships inferred from
maximum parsimony analysis
of ITS-rDNA sequences of 31
taxa. The percentages of
replicate trees in which the
associated taxa clustered
together in the bootstrap test
(1000 replicates) are shown next
to the branches. The tree is
drawn to scale, with branch
lengths in the same units as
those of the evolutionary
distances used to infer the
phylogenetic tree. The new Sri
Lankan records: Favolaschia
auriscalpium having GenBank
Accession Number KY649461
and Herbarium Number MFLU
12-1891; Favolaschia
manipularis having GenBank
Accession Number KY649462
and Herbarium Number MFLU
12-1172 (ITS-rDNA) are shown
in bold and blue. The topology
is rooted with Mycena crocata.
Evolutionary analysis was
conducted in PAUP 4.0b 10
cortical layer of the stipe 3–8 lm wide, clamped, with
diverticulate terminal cells. Caulocystidia 24–39 9 6–8
lm, abundant, irregularly shaped.
Habit, habitat and distribution: as a group, usually on
dead or decaying wood, May–August. Our collection was
collected on a decaying wood at Hanthana Mountains
Range, Peradeniya.
Specimens examined: SRI LANKA, Kandy District,
Hanthana Mountains Range, 3 June 2012, Samantha C.
Karunarathna (MFLU 12-1172, new record)
GenBank Number ITS:KY649462.
Notes: The mycelium and fruiting bodies of the mushroom are bioluminescent. This is found in Australasia,
Malaysia, and the Pacific islands and this is the first report
of F. manipularis with the molecular phylogenetic confirmation from Sri Lanka.
Physalacriaceae Corner
Physalacriaceae is a family of mushrooms in the order
Agaricales which was originally defined by the English
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Fungal Diversity
(2008), 15 species have been recorded worldwide
(Fig. 143).
Fig. 139 Basidiocarps of Favolaschia auriscalpium in the field
(MFLU 12-1891)
mycologist E.J.H. Corner in 1970 (Corner 1970). Species
in this family have a widespread distribution, ranging from
the Arctic, to the tropics and from marine sites and fresh
waters to semiarid forests.
Cyptotrama Singer
Cyptotrama is a genus of mushrooms in the family
Physalacriaceae. The type species of the genus is Cyptotrama macrobasidia Singer. According to Kirk et al.
Fig. 140 Favolaschia
auriscalpium (MFLU 12-1891).
a Gloeocystidia.
b Acanthocysts. c Epicular
elements (smooth
dermatocystidia). d Pores.
e Basidia. f Basidiospores
123
Cyptotrama asprata (Berk.) Redhead & Ginns, Canadian
Journal of Botany 58 (6): 731 (1980)
Facesoffungi number: FoF 3140, Figs. 144, 145
Pileus 1–5 cm in diam., convex, hemispherical to
applanate, whitish, cream to yellow, densely covered by
orange, golden yellow coloured squamules, becoming
whitish-yellow when mature. Lamellae adnate to slightly
decurrent, height up to 5 mm, distant, white to cream, with
lamellulae. Stipe 1–6 cm in length, 0.1–0.6 cm in diam.,
floccose, cylindrical or enlarged at base, fistulose, concolorous with pileus. Context thin, white. Odour indistinct.
Basidiospores (7–11 9 4.5–8.5 lm, Q = 1.2–1.9 lm,
Qm = 1.4), non-amyloid, non-dextrinoid, thin-walled,
smooth, ellipsoid-limoniform to broadly ellipsoid, colourless and hyaline. Basidia 4-spored, 35–60 9 6–8 lm, clavate, colourless or hyaline; sterigmata 4–6 lm long.
Basidioles abundant, 40–60 9 4–8 lm, fusiform with
subacute apices, thin walled, colourless. Cheilocystidia and
pleurocystidia present and similar in size and form, scattered, 50–90 9 8–17 lm, fusiform with acute to subacute
apices, thin-to slightly thick-walled, colourless or hyaline.
Pileocystidia absent. Squamules on pileus composed of
Fungal Diversity
Fig. 141 Basidiocarps of Favolaschia manipularis in the field (MFLU12-1172)
inflated chained, narrowly clavate, cylindrical to fusiform
cells, 18–65 9 3–16 lm, thick-walled, golden yellow to
brownish-yellow. Pileipellis under squamules an epithelium, composed of 1–4 layers, broadly ellipsoid, subglobose to obovoid cells, 24–40 9 12–28 lm, thin to
somewhat thick walled, nearly colourless. Squamules on
stipe composed of interwoven, filamentous to narrowly
cylindrical hyphae 2–5 lm wide, thin walled, colourless to
yellowish, hyaline, with few terminal cells, 25–40 9 5–10
lm, slightly thick walled. Stipitipellis composed of vertically arranged, colourless to yellowish filamentous hyphae,
3–10 lm in diam. Stipe trama composed of vertically
arranged, colourless or hyaline hyphae, 3–16 lm in diam.
Clamp connections abundant.
Habit, habitat and distribution: solitary on a dead wood,
widely distributed in tropical, subtropical or even temperate regions of many parts of the world, May–September.
Our collection was collected on a decaying wood near Doi
Suthep Pui national park, Chiang Mai.
Specimens examined: THAILAND, Chiang Mai Province, Doi Suthep Pui national park, 30 July 2012,
Samantha C. Karunarathna (MFLU 12-1892, new record).
GenBank Number ITS:KY649460.
Notes: Cyptotrama asprata was first described from Sri
Lanka. It is widely distributed in South and Southeast Asia
(Sri Lanka and Laos), East Asia (China, Japan and Korea),
New Zealand, Hawaii, and Central America (Martinique and
France) (Berkeley 1847; Redhead and Ginns 1980; Yang
1990; Moreau et al. 2015; Qin and Yang 2016). Cyptotrama
chrysopepla morphologically resembles to C. asprata
because of its’ gold yellow and scruffy appearance, but differs by having smaller squamules, hymeniderm pileipellis,
thick walled elements in squamules of stipe, the non-acute
subcapitate pleurocystidia and larger basidiospores (8.5–13
9 6.5–10 lm). This is the first report of C. asprata with the
molecular phylogenetic confirmation from Thailand.
Boletales E.-J. Gilbert
Boletaceae Chevall.
Boletaceae is a family of mushrooms, first described by
the French botanist François Fulgis Chevallier in 1826 as a
family distinct from Agaricaceae, primarily characterized
by developing their spores in small pores on the underside
of the mushroom, instead of gills, as are found in agarics.
As a whole, the typical members of the family are commonly known as boletes.
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Fungal Diversity
Fig. 142 Favolaschia
manipularis (MFLU 12-1172).
a Basidia. b Basidiospores.
c Cheilocystidia.
d Pileocystidia. e Caulocystidia
Austroboletus
The genus Austroboletus (Corner) Wolfe was established by Wolfe (1979) with the boletoid mushrooms
showing the combination of striking morphological features: white to pale cream coloured tube and pore surface
which gradually turn pink to pink-rose or sometimes
vinaceous on bruising; lacunose to lacerate or reticulate
stipe surface; spore print pink to vinaceous or purple brown
to rust cinnamon, basidiospores with warted to pitted or
reticulate ornamentations; ixotrichodermium or trichodermium nature of pileipellis (Wolfe 1979; Das and Dentinger
2015). According to recent phylogenetic analysis of the
members of Boletaceae, this genus is placed under subfamily Austroboletoideae (Wu et al. 2014). About 30
species are reported globally (Kirk et al. 2008) in this
genus. Here we describe a new Austroboletus species with
123
the support of morphology and phylogeny (Fig. 146) which
was collected from Uttarakhand of India.
Austroboletus appendiculatus Semwal, D. Chakr., K. Das,
Indoliya, D. Chakrabarty, S. Adhikari & Karunarathna, sp.
nov.
MycoBank number: MB 817672; Facesoffungi number:
FoF 2952, Figs. 147, 148 Etymology: referring to stipes (of
basidiomata) with appendages.
Holotype: CAL 1304
Diagnosis: is distinct from the closest species A. subvirens by larger and differently coloured pileus, longer and
wider stipe, presence of pleurocystidia and caulocystidia
and putative association with Shorea sp.
Pileus 75–90 mm. diam.; convex when young, to planoconvex, sometimes uplifted with age; surface areolate-
Fungal Diversity
Fig. 143 Phylogenetic relationships inferred from maximum parsimony analysis of ITS-rDNA sequences of 52 taxa. The percentages of
replicate trees in which the associated taxa clustered together in the
bootstrap test (1000 replicates) are shown next to the branches. The
tree is drawn to scale, with branch lengths in the same units as those
of the evolutionary distances used to infer the phylogenetic tree. The
new Thai record: Cyptotrama asprata having GenBank Accession
Number KY649460 and Herbarium Number MFLU 12-1892 is shown
in bold and blue. The topology is rooted with Marasmius epiphyllus.
Evolutionary analysis was conducted in PAUP 4.0b 10
squamose, then disrupted into several patches or segments
with the expansion exposing yellowish-orange to greyishyellow (4B7–4A7, 4B6) subpellis that becoming paler
towards margin as pastel yellow to butter yellow, light
yellow (3A4–4A5, 4A4); segments of suprapellis light
brown to brown, cottony, slightly viscid when wet; margin
plane, regular. Pore surface pinkish when young, gradually
on bruising or maturity greyish-orange to brownish-orange
(5B3–6C3), reddish-grey (12B2), or darker to brown, but
paler toward the stipe juncture as light orange grey (6B2),
depressed at junction to stipe, unchanging on bruising,
pores often up to 2 mm broad at maturity, angular to
123
Fungal Diversity
Fig. 144 Basidiocarps of Cyptotrama asprata in the field (MFLU
12-1892)
rectangular, attached. Tubes 12–16 mm long, concolorous
to slightly paler then pore surface. Stipe 80–120 9 20–30
mm, central, terete or more or less fusoid towards down,
surface distinctly covered with pale orange to light orange
(4–5A2–4) or sometimes yolk yellow (4B8) anastomosing
broad ridges or appendages forming elevated or flaring to
wing-like network (on yellowish-white background) which
gradually becoming thin (less elevated) reticulum towards
apex, but, never with typical lacerate surface, base with
white mycelia elements; ridges cottony, darker with age as
golden yellow to brownish-yellow (5B5–6, 5C6). Context
white, unchanging, fluffy, partly hollow in stipe. Odour
mushroomoid. Taste indistinct. Spore print rust brown to
reddish-brown (6E8–8E8). Basidiospores 14.2–14.8–16.5
9 7.3–8–9.1 lm (n = 20, Q = 1.68–1.83–1.97), broadly
fusoid to amygdaliform, depressed suprahilar part, surface
appears warted by disruption of the outer wall, deeply
pitted to form punctuate-warted; under SEM, pits minute at
both the fusoid ends, gradually with intricate furrows
towards mid region, at mid region with more broader and
deeper pits forming warty surface with truncate warts, with
disrupted adaxial surface. Basidia 22–32 9 12–16 lm, 4–
spored, clavate. Pleurocystidia rare. Tube trama boletoid,
gelatinous. Pileipellis trichoderm, composed of erect,
septate hyphae; terminal cells cylindrical with rounded
apex, width 6–9 lm. Stipitipellis fertile, composed of
caulocystidia and caulobasidia. Caulocystidia 40–70 9
11–20 lm, clavate to subclavate, subventricose to appendiculate. Caulobasidia 30–45 9 9–12 lm, 2- to 4-spored,
clavate.
Habitat and distribution: Under Shorea robusta Gaertn.
in subtropical broadleaf forest.
Fig. 145 Cyptotrama asprata (MFLU 12-1892). a Basidia. b Basidiospores. c Cheilocystidia. d Stipitipellis. e Pileipellis
123
Fungal Diversity
Fig. 146 Phylogenetic relationships inferred from neighbour-joining
analysis of ITS-rDNA sequences of 22 taxa. The optimal tree with the
sum of branch length = 0.82684195 is shown. The percentages of
replicate trees in which the associated taxa clustered together in the
bootstrap test (1000 replicates) are shown next to the branches. The
tree is drawn to scale, with branch lengths in the same units as those
of the evolutionary distances used to infer the phylogenetic tree. The
novel Indian species: Austroboletus appendiculatus having GenBank
Accession Number KX530028 (ITS-rDNA) is shown in bold and
blue. The topology is rooted with Boletus edulis. Evolutionary
analysis was conducted in MEGA 6.0 (Tamura et al. 2013)
Material examined: INDIA, Uttarakhand, Dehradun
District, Tapovan forest, 650 m, 17 July 2010,
K.C. Semwal, KCS 1401 (CAL 1304, holotype; isotype:
BSHC).
GenBank Numbers ITS:KX530028.
Notes: Austroboletus appendiculatus is typically featured by combination of characters: brownish areolatesquamose pileus with exposed yellowish subpellis, distinctively reticulate-appendicute and non-lacerate stipe
surface, pinkish pore surface that becomes brownish to
greyish on bruising or after maturity, partly hollow stipe,
almond-shaped deeply pitted basidiospores with distinct
suprahilar depression and prominently disrupted adaxial
surface and occurrence with putative association of Shorea
in subtropical broadleaf forest.
In the field, Austroboletus subvirens (Hongo) Wolfe
[:Porphyrellus subvirens Hongo], A. lacunosus (Kuntze)
T.W. May & A.E. Wood [:A. cookie (Sacc. & Syd.)
Wolfe] and A. novae-zelandiae (McNabb) Wolfe [:Porphyrellus novaezelandiae McNabb] are quite similar (as far
as the stipe surface is concerned) to the present taxon. But,
A. subvirens (probably the closest species being reported
earlier from Japan, Papua New Guinea, China, Americas
and Australia and labelled the Australian Material here
with GenBank Numbers KP242207, KP242214, KP242212
and KP242210 in Fig. 146) has distinctively smaller
(25–60 mm) and differently coloured pileus, shorter and
thinner stipe (50–90 9 4–10 mm), and lacks pleuro-,
cheilo- and caulocystidia (Wolfe 1979; Jian-Zhe 1985).
Moreover, A. subvirens are found to be associated with the
trees of Fagaceae in Australasia and Americas and Acacia,
Allocasuarina and Eucalyptus in Australia (Wolfe 1979;
Horak 1980a, c; Halling et al. 2006) whereas, A. appendiculatus (present species) grows under Shorea robusta
(Dipterocarpaceae). Austroboletus novae-zelandiae (reported from New Zealand and New Caledonia and labelled
with GenBank Numbers accession number HM060327 in
Fig. 146) is separated from the present species by distinctively shorter and slender stipe (70 9 8 mm), bigger
(17–19.5 9 8–10.4 lm) basidiospores (Wolfe 1979). Austroboletus lacunosus (reported from Australia and labelled
with GenBank Accession Number KC552014 in Fig. 146)
can also be separated from the present taxon by smaller
pileus (15–25 mm), and stipe (20–50 9 5–6 mm) and
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Fungal Diversity
Fig. 147 Austroboletus appendiculatus (CAL 1304, holotype). a–
c Fresh basidiomata in the field and base camp. d Hymenophoral
trama. e Terminal cells of hyphae of pileipellis. f Cross section
through stipitipellis showing anastomosing ridges. g, h Caulocystidia.
i Basidiospores. j, k SEM micrograph of basidiospores. Scale bars
a = 100 mm, d = 50 lm, e, g–j = 10 lm, f = 100 lm, k = 5 lm
smaller (32.5–56 9 10–15.5 lm) caulocystidia (Wolfe
1979).
Only two other species, hitherto known from India:
A. olivaceoglutinosus K. Das & Dentinger (labelled with
GenBank Numbers KM597478 and KM597479 in
Fig. 146) and A. gracilis (Peck) Wolfe can easily be differentiated in the field from the present novel species.
A. olivaceoglutinosus which is found in subalpine Himalayan region under Tsuga and Picea has distinguishingly
long and slender basidiomata (Pileus 30–55 mm diam.,
stipe 100–180 9 13–17 mm), highly glutinous olive
coloured pileus, lacerate (never with ridges) stipe surface
and narrower (12.7–19 9 5.9–7.7 lm) basidiospores
showing different ornamentation pattern (Das and
123
Fungal Diversity
Fig. 148 Austroboletus
appendiculatus (CAL 1304,
holotype). a Basidia.
b Caulocystidia. c Caulobasidia.
d Radial section through
pileipellis. Scale bars a–d = 10
lm
Dentinger 2015) whereas, A. gracilis has different pilear
surface (subtomentose, velvety or fibrillose), smaller
caulocystidia: ‘‘14–42(–57) 9 4–14 lm’’ and different
pattern (‘‘rugulose punctate’’) of spore ornamentation,
different host associations (Wolfe 1979; Lakhanpal 1996).
Considering the spore ornamentation, A. fusisporus
(reported from Japan and China and labelled with GenBank
Numbers AB509830 and JX889719 in Fig. 146) also
appears close to A. appendiculatus but the former has
altogether different and very small (pileus 5–20 mm, stipe
20–45 9 2–4 mm) basidiomata (Wolfe 1979).
Our ITS phylogeny (Fig. 146) were obtained by the
neighbour-joining analysis criterion considering the all
available ITS sequences available in GenBank and/or
appeared in BLAST search with the sequence of our Indian
material. The new sequence forming a separate clade is
clustered (80% bootstrap) with the sequences of A. subvirens derived from Australian Material (Fig. 146). These
two species in turn is clustered (80% bootstrap) with
A. festivus (sequence derived from specimen collected from
Guyana).
Unique combination of macro- and micromorphological
features coupled with ITS phylogeny clearly support the
novelty of Austroboletus appendiculatus.
Boletellus Murrill
Boletellus is a genus of mushrooms in the family Boletaceae (Binder and Hibbett 2006). The genus was first
123
Fungal Diversity
introduced by American mycologist William Alphonso
Murrill in 1909 (Murrill 1909). The genus contains about
50 species, has a widespread distribution, especially in
subtropical regions (Kirk et al. 2008) (Fig. 149).
Boletellus emodensis (Berk.) Singer, Annales Mycologici
40: 19 (1942)
Facesoffungi number: FoF 3141, Figs. 150, 151
Basidiomata (small to medium-sized. Pileus 3–9 cm in
diam., convex to applanate; surface smooth and dry, purple
to dull red colour, fading to pale brown with age, finely
tomentose when young, then peeling or cracking into large
squamules; at first margin extended into a fake veil and
covering the pores, then radially splitting, appendiculate
with fake veil remnants; context 0.5–1.2 cm thick in the
pileus center, yellowish, strongly and rapidly turn to blue
when bruised or injured. Hymenophore poroid, yellowish,
strongly and rapidly turn to blue when bruised or injured;
pores 0.1–0.2 cm diam., simple and angular. Stipe central,
6–8 9 0.7–1 cm, cylindrical, firm, solid; surface concolorous with the pileus, dry, fibrous but apex yellowish;
context yellow, strongly and rapidly turn to blue when
bruised or injured. Annulus not present. Basidia 33–55 9
10–18 lm, clavate, colourless to yellowish-brown in KOH,
mostly 4-spored. Basidiospores 18–24 9 8–11 lm,
Q = 1.9–2.8, Qm = 2.28, yellowish-brown in KOH, elongate-ellipsoid with longitudinal ridges, 6–9 ridges visible in
lateral view; ridges up to 0.9–1.6 lm in height. Tube trama
made up of thin-walled, colourless to yellowish hyphae
5–13 lm in width. Cheilocystidia 46–69 9 10–18 lm, thinwalled, subfusiform to fusiform, colourless or yellowishbrown in KOH. Pleurocystidia 68–92 9 10–20 lm, subfusiform to fusiform, colourless to yellowish-brown in
KOH. Pileipellis is an intricate trichoderm composed of
interwoven, thin-walled, yellowish-brown hyphae; terminal
cells 18–36 9 6–12 lm, with round to obtuse apex. Pileal
trama composed of thin walled, colourless to yellowish
hyphae 4–16 lm diam. Clamps absent.
Habit and habitat: Solitary or gregarious on soil, tree
stumps or rotten wood in forests dominated by Fagaceae
Distribution: originally described from India; then
reported from southeast Asia, east Asia and Australia, this
is the first report from Sri Lanka with morphological
characteristics and phylogenetic data.
Specimens examined: SRI LANKA, Kandy District,
Hanthana Mountains Range, 10 June 2012, Samantha C.
Karunarathna (MFLU 12-1214, new record).
GenBank Number ITS:KY649458.
Notes: Boletellus emodensis is characterized by its
purple to dull red pileus, yellowish context in both pileus
and stipe that rapidly and strongly turn to blue when
injured or bruised, ridged basidiospores with fine striations
on the ridges can be observed under the high power of light
123
microscope. This is the first report of B. emodensis with the
molecular phylogenetic confirmation (Fig. 149) from Sri
Lanka.
Rubroboletus Kuan Zhao & Zhu L. Yang
The genus Rubroboletus (Zhao et al. 2014), typified by
R. sinicus (W.F. Chiu) Kuan Zhao et Zhu L. Yang, was
established to accommodate Boletaceae with reddish pileal
surface, an orange, orange-red to blood red surface of the
hymenophore, yellow tubes, pink to red net or spots on the
stipe, a bluish colour-change when injured, a non-amyloid
context (even if the original diagnosis says ‘‘not
observed’’), smooth spores, olive-brown in deposit, and an
interwoven trichodermal pileipellis (Fig. 152).
Rubroboletus demonensis Vasquez, Simonini, Svetasheva,
Mikšı́k & Vizzini, sp. nov.
= Boletus rhodopurpureus f. polypurpureus s. Ruiz
Fernandez & Ruiz Pastor, Guı́a Micologica Tomo n. 4,
Supl. Orden Boletales en España: 62 (2006)
= Boletus rubrosanguineus s. Calzada Dominguez 2007,
Guı́a de los Boletos de España y Portugal: 163 (2007)
= Boletus legaliae s. Rodà, Funghi Aspromontani
Comparati Boletales: 96 (right), 97 (top), countercover
(2012)
Index Fungorum number: IF552526; Facesoffungi
number: FoF 2953, Figs. 153, 154, 155
Etymology: the specific epithet of the name of the taxon
comes from the Latin word ‘‘demonensis’’ and refers to the
ancient name ‘‘Valdemone’’, which was one of the three
valleys (‘‘valli’’) or real domains (‘‘reali dominii’’) in
which the region Sicily was subdivided from the Muslim
domination to the Bourbon period; the ‘‘Valdemone’’, with
respect to the ‘‘Val di Noto’’ and the ‘‘Val di Mazzara’’,
constitutes the north-eastern part of the Sicily, an area that
closely corresponds to the actual habitat of R. demonensis.
Furthermore, the epithet ‘‘demonensis’’ well reminds the
peculiar features of the species, i.e. the flaming red colour
of the cap and the pores, features shared with other close
‘‘devilish’’ species, belonging to the same genus (R. satanas). Moreover, unforeseeable circumstances would have
wanted that one of the first growth stations of R. demonensis had been the locality ‘‘Pizzo Inferno’’ (i.e.: Hell
Peak) on the Nebrodi chain, in the common of Floresta.
Holotype: MCVE 29081 (GS10244)
Pileus 60–150(–240) mm broad, initially hemispherical,
then convex, pulvinate, in the end almost applanate in
mature basidiomes. Margin ± regular in young, soon
uneven, ±undulate-lobate, sometimes distinctly lobate in
mature specimens. Colour at the very beginning whitish,
pale grey, pale ochre (4/A1–B2), soon, starting from the
margin, dirty pale pink-lilac (5–12/A1–B2), then strongly
variable from light grey (5–7/B1–C2), to pink-lilac (7–12/
Fungal Diversity
Fig. 149 Phylogenetic
relationships inferred from
maximum parsimony analysis
of ITS-rDNA sequences of 28
taxa. The percentages of
replicate trees in which the
associated taxa clustered
together in the bootstrap test
(1000 replicates) are shown next
to the branches. The tree is
drawn to scale, with branch
lengths in the same units as
those of the evolutionary
distances used to infer the
phylogenetic tree. The new Sri
Lankan record: Boletellus
emodensis having GenBank
Accession Number KY649458
and Herbarium Number MFLU
12-1214 is shown in bold and
blue. The topology is rooted
with Boletus sp. Evolutionary
analysis was conducted in
PAUP 4.0b 10
B2–C3), to purple-red (10–12/B3–C6, 11–12/C6–D8)
depending on environmental conditions; with wet weather
or in shady woods, the pileus shows particularly bright
colours, tending to purple-blood red, quite uniformly distributed on the pileus surface (11–12/C8–D8). In some
occurrences, red colour seems not to develop completely
both in pores and stipe, and in meantime also the pileus
colour stabilizes on a pink-lilac uniform colour (8–11/A3–
B4), that becomes blood red when bruised. Conversely, in
dry weather or in areas exposed to the sun, pileus colour is
much more variable, with flesh pink colours sometimes
fading to brownish-grey or to paler tone tending to creamwhitish (7–9/B2–D3). However, in the most of the basidiomes observed, large intensely reddish areas or wide blood
123
Fungal Diversity
Fig. 150 Basidiocarps of Boletellus emodensis in the field (MFLU
12-1214)
red spots persist, even more intense following to scrubbing,
touching or other manipulation. Surface typically tomentose and dry at the beginning, sometimes smooth and viscous in wet weather; the cuticle is not detachable from the
underlying flesh. Often, in mature large basidiomes, the
pileus appears pleasantly bright shining red, darker when
bruised. Looking closely at the pileus surface, also with the
aid of a lens, minute, in relief, concolorous scales can be
noticed. This feature is shared with the close species
R. rubrosanguineus, R. legaliae, R. pulcherrimus and
R. sinicus. Stipe 80–120(–150) 9 40–80(–90) mm, bulky,
robust and wide, cylindrical, often gradually enlarged or
clavate towards the base, sometimes also obese, not rooting. Surface in rare case yellow orange at the very beginning (4–6/A4–A6), with purplish base (9–10/A4–B6), then,
passing through intermediate orange-red tones (7–8/A4–
B6), in the end bright red, blood red, purple red, often
123
darker at the base (9–10B7–D8, 9–11D6–D8), often with
an evident deep yellow, or orange-yellow band (5–15 mm
wide) in the upper part. In some xanthoid aspects stipe does
not develop red tones in the upper half, remaining yellow
or orange yellow, but the stipe base is anyway red, purplered, blood red. Bitten areas often show the deep yellow
flesh colour, giving a chromatic effect reminiscent of
R. rhodoxanthus. A sharp net, with regular meshes, is well
distributed over at least of the height, darker than the
colour of the ground; the meshes are usually larger and the
ribs usually thicker than those of B. rubrosanguineus or
B. legaliae, giving to the net a general rougher appearance
with respect to the ones of the close species. The extreme
base of the stipe is covered with a whitish grainy, furfuraceous over layer, particularly evident in basidiomes
collected on moist grounds. Tubes 5–12(–20) mm, having
average length, free at the stipe, from deep yellow to olive
green, discolouring blue when cut. Pores small, round,
usually from the beginning purple-red, dark red (10–11/
C7–D8, 11–12/D7–F8), dull blue when touched, in aged
basidiomes orange-yellow towards the margin of the
pileus. In some collections, young specimens have yellow
pores, that with age develop only an orange, red–orange
colour, reminiscent that one of typical R. legaliae. Flesh
firm, yellow, deep yellow, lemon yellow (3/A4–A8), particularly intensive in both stipe and pileus wounds; when
exposed, quickly turning sky-blue, blue, deep blue (21–23/
A4–B6), due to discolouring not particularly intensive,
sometimes weak, then fading to a pale grey-cream colour.
In some cases, in the exposed pileus or mostly in the lower
part of the stipe some beetroot shades may appear. Subhymenophoral layer concolorous. Taste sweetish, slightly
acidic. Smell weak, fungal and pleasant. Spore print
tobacco-brown. Amyloid reaction in the flesh at the stipe
base, according to Imler’s procedure (Imler 1950): the
tissues at the stipe base are not amyloid in all the collections if examined microscopically. As in R. legaliae and
R. rubrosanguineus, sometimes the amyloid reaction could
be macroscopically ‘‘positive’’ (Cheype 1983) due to the
absorption of the Melzer’s regent in the tissues. Basidiospores smooth, fusiform ellipsoid, with a more or less
pronounced suprahilar depression, (12.5–)12.6–13.6(–13.9)
9 (4.6–)4.7–5.1(–5.1) lm (363/7/7), on average 13.1 ± 0.5
9 4.9 ± 0.2 lm, Q = (2.53–)2.57–2.79(–2.89),
Qm = 2.68 ± 0.11, from light yellow to yellow–brownish
in KOH. They may have different shape in different collections, despite the molecular investigation has confirmed
them as belonging to the same species. In some collections,
they are ellipsoid (Fig. 156b), more elongate with respect
to those of R. legaliae, smaller both in width and in length
with respect to those of R. rubrosanguineus but with almost
the same Q ratio; in other collections, they are more fusiform, with the apex opposed to apiculum more tapered and
Fungal Diversity
Fig. 151 Boletellus emodensis
(MFLU 12-1214).
a Pleurocystidia. b Basidia.
c Basidiospores.
d Cheilocystidia. e Pileipellis
Fig. 152 Phylogeny of the genus Rubroboletus based on a Bayesian
and Maximum Likelihood Inference analysis of a ITS dataset.
Bayesian posterior probability values (in bold) C0.7 and maximum
likelihood bootstrap (ML) values C50% are shown on the branches.
Thickened branches indicate BYPP C 0.95 and ML support C70%.
Dashed branches indicate BYPP C 0.95 and ML bootstrap support
\70%. Nodes that receive BYPP \ 0.95 but with C70% ML are
indicated by small black-filled squares. Newly sequenced collections
are in bold. Caloboletus calopus (HM347645) was chosen as the
outgroup taxon
acuminate (Fig. 156c). In some collections, lots of irregular
spores, with various, irregular protuberances, may also
occur (Fig. 156c(right)). Basidia mostly 4-spored, hyaline,
(23.8–)31.3–47.7(–62.6) 9 (4.2–)7.7–12.7(–13) lm (34/2/
2). Facial cystidia fusiform, versiform, sometimes
lageniform, hyaline, (15.5–)29.5–44.7(–50.7) 9 (4.2–)
5.3–7.7(–9.5) (64/2/2). Marginal cystidia similar to facial
cystidia but smaller, mostly versiform than fusiform or
lageniform, hyaline (15.8–)20–29 (37.2) 9 (3.5–)4.3–6.3
(–7.9) (46/2/2). Pileipellis an entangled trichodermium,
123
Fungal Diversity
Fig. 153 Rubroboletus demonensis (GS10244, holotype). Basidiomes from the main collections. a 9 October 2013, C.da Piano Torre
(Madonie), Castelbuono (PA), with Quercus pubescens s.l. and
Q. ilex. b 5 September 2013, Portella dell’Obolo (Nebrodi), Capizzi
(ME), with Q. pubescens s.l. c 4 August 2014, near Village Nimea,
Komotini, Rhodope Regional Unit, with Q. petraea ssp. medwediewii.
d 15 September 2013, Portella dell’Obolo (Nebrodi), Capizzi (ME),
with Q. pubescens s.l. e 15 September 2013, Portella dell’Obolo
(Nebrodi), Capizzi (ME), with Q. pubescens s.l. f 27 June 2009,
Portella Sella Maria (Nebrodi), Cesarò (ME), with Q. pubescens and
Q. virgiliana. g 30 June 2008, pizzo Inferno (Nebrodi), Floresta (ME),
with Castanea sativa and Pinus nigra ssp. calabrica. h GS10274.
i GS10244 (Holotype). j GS10708. k GS10278. l GS10242. Bars 50
mm
tending to a cutis, often gelatinized in mature individuals,
consisting of thin, cylindrical elements, collapsing gradually during development of the basidiomes. Terminal elements (33.3–)32.9–42.9(–44.3) 9 (5–)5.2–5.9(–6.9) lm
(138/5/5), Q = (6.6–)6.4–8.1, Qm = 7.3, with rounded or
tapered tip, sometimes clavate, rarely capitulate or pearshaped, with pale yellow vacuolar pigment and incrusted
pigmented here and there, mostly in the deeper and closer
to hypoderma hyphae. Hymenophoral trama boletoid in the
sense of Singer (1965, 1967). Clamp connections absent.
Habitat: Rubroboletus demonensis is typical in acid and
silicic soils, thermophilic, in summer. It grows in groups of
small basidiomes in warm deciduous forests, in mountains
or in some cases in mixed deciduous and coniferous woods
123
Fungal Diversity
Fig. 154 Rubroboletus
demonensis (GS10244,
holotype). Microscopic
structures. a Pileipellis in 3%
NH3 ? Congo Red, with
Differential Interference
Contrast (DIC) (GS10244).
b Spores in 3% NH3
(GS10244). c left Spores in 3%
NH3 (GS10274). c right
Abnormal spores in 3% NH3
(GS10243). d Basidia in 3%
NH3 ? Congo Red (GS10245).
e Facial cystidia in 3% NH3 ?
Congo Red, with DIC
(GS10245). f Marginal cystidia
in 3% NH3 ? Congo Red, with
DIC (GS10245). Bars 10 lm
(Pinus nigra and Taxus baccata), never with pure conifers.
It prefers mesic forests of deciduous oaks (Quercus pubescens sensu lato, Q. cerris, Q. congesta and Q. virgiliana),
rarely with holm (Q. ilex). It is common in chestnut woods
(Castanea sativa), pure or mixed, and at higher elevations
with beech (Fagus sylvatica). It is usually collected in June
after the late spring rains and, not liking the summer
drought, it disappears during the hottest period of July and
August, then appears again in September. During the
warmer autumns, it can be observed until the first half of
October.
Distribution: R. demonensis was found in a rather
delimited areal of the South Italy [(Sila Plateau (‘‘Altopiano della Sila’’) in Calabria] and Sicily, where it inhabits
the mesomediterranean and supramediterranean zone of the
mountain strip, characterized by a humid Mediterranean
123
Fungal Diversity
Fig. 155 Distribution of spore size of R. demonensis (7 collections),
compared with R. legaliae (13 collections) and R. rubrosanguineus
(20 collections), using ‘‘isoprobability ellipses’’. It is shown the
distribution of the average values of spore size for any of the
collections, at the confidence of 68% (corresponding to 1st deviation,
left side) and at the confidence of 95% (correspondent to 2 9 SD,
right side)
climate, with large rainfalls and Mediterranean humid
forest well represented by deciduous forests of oak
(Q. pubescens s.l.) and chestnut (C. sativa), some rare
patches of evergreen forest with Q. ilex, with the presence
of Pinus nigra ssp. calabrica, in an altitudinal range
between 1200 m and 1600 m a.s.l.. Sicily seems to be the
southern limit of distribution of the species. A unique
collection from Greece (04.08.2014, near Nimea Village,
Komotini, Rhodope Regional Unit, coord. 41120 42.200 N,
25310 16.500 E, alt. 650 m a.s.l., leg. D. Tragkos) with
Q. petraea ssp. medwediewii, proved only by photographic
material, represents the most eastern collection. The findings in Spain under Q. faginea and Q. ilex, until now
attributed to Imperator rhodopurpureus var. polypurpureus
(Ruiz Fernandez & Ruiz Pastor 2006) or R. rubrosanguineus (Calzada Dominguez 2007), seem likely referable
to R. demonensis, and, if confirmed, might represent the
western (Zamora, Salamanca) and northern (Navarra,
Palencia, Huelva) limits of the distribution of the species.
The main collection sites in Sicily are the north-eastern
mountain range (Appennino Siciliano), the Nebrodi chain
and the Madonie chains, but it has also been reported on
the Etna volcano massif, where it grows even at higher
altitudes. The most of the collections from Sicily comes
from the commons of Cesarò, Floresta, Maniace and
Troina in the Nebrodi Mountains Park, in the province of
Messina.
Edibility: like other closely related species in this genus,
it is probably poisonous if consumed raw.
Material examined: (GS refers to the personal herbarium
of G. Simonini): Rubroboletus demonensis. ITALY—
CALABRIA. 20 June 1999, Gambarie (RC), with C. sativa,
coord. 38100 N, 15500 E, alt. 1300 m a.s.l., leg. C. Lavorato,
GS2129. 9 June 2013, S. Domenica di Terreti (RC), with C.
sativa, coord. 38100 2100 N, 15470 4800 E, alt. 900 m a.s.l., leg.
P. Rodà, GS10708. SICILY. 26 September 2014, Portella
Sella Maria (Nebrodi), Cesarò (ME), with F. sylvatica and
Q. pubescens, coord. 37540 100 N, 14390 1700 E, alt.1400 m
a.s.l., leg. G. Vasquez, GS10400; 19 June 2015, Lago
Biviere di Cesarò (Nebrodi), Cesarò (ME), with F. sylvatica, coord. 37570 600 N, 14420 5000 E, alt. 1290 m a.s.l., leg.
M.G. Pulvirenti, GS10242; 22 June 2015, Portella Sella
Maria (Nebrodi), Cesarò (ME), with F. sylvatica and
Q. pubescens, coord. 37530 5700 N, 14390 1500 E, alt. 1470 m
a.s.l., leg. D. Milazzo, GS10243; 28 June 2015, Portella
Sella Maria (Nebrodi), Cesarò (ME), with F. sylvatica and
Q. pubescens, coord. 37540 100 N, 14390 1700 E, alt. 1420 m
a.s.l., leg. G. Vasquez, MCVE 29081, (GS10244,
123
Fungal Diversity
Fig. 156 Phylogram generated from neighbour-joining analysis
based on ITS-rDNA sequences: Phylogenetic relationships inferred
from neighbour-joining analysis of ITS sequences of 25 taxa. The
optimal tree with the sum of branch length = 0.79320854 is shown.
The percentages of replicate trees in which the associated taxa
clustered together in the bootstrap test (1000 replicates) are shown
next to the branches. The tree is drawn to scale, with branch lengths in
the same units as those of the evolutionary distances used to infer the
phylogenetic tree. The novel species having GenBank Numbers
KX364694 (ITS-rDNA) is shown in bold and blue font. Boletus edulis
(European material with GenBank Numbers AY278764) was considered as the out group. Evolutionary analysis was conducted in
MEGA6 (Tamura et al. 2013)
holotype); 20 August 2015, Piano della Cicogna, Cesarò
(ME), with Q. cerris, coord. 37520 5500 N, 14390 3200 E, alt.
1350 m a.s.l., leg. M.G. Pulvirenti, GS10274; 20 September 2015, Piano della Cicogna, Cesarò (ME), with Q. cerris, coord. 37520 5900 N, 14390 3800 E, alt. 1320 m a.s.l., leg.
M.G. Pulvirenti, GS10278.
Additional material examined (MM and JLC refer to the
personal herbaria of M. Mikšı́k and J.L. Cheype):
Rubroboletus legaliae. CZECH REPUBLIC—17 July
2005, Lednice, chateau park, with Q. robur, 48480 4400 N,
16480 2500 E, alt. 230 m a.s.l., leg. M. Mikšı́k, MM 17705B2;
26 July 2008, Bechov-Svobodı́n, Mladá Boleslav, with
Q. petraea and Carpinus betulus, 50260 700 N, 1540 5200 E, alt.
250 m a.s.l., leg. M. Mikšı́k, MM 26708B1; 6 August 2012,
Dlouhopolsko, Nymburk, ‘‘Kněžičky’’ reserve, with Q. petraea, 5090 4000 N, 15200 2400 E, alt. 240 a.s.l., leg. M. Mikšı́k,
MM 6812B2; 9 August 2012, Dlouhopolsko, Nymburk,
‘‘Kněžičky’’ reserve, with Q. petraea, 5090 4600 N,
15190 4700 E, alt. 250 a.s.l., leg. D. Kvasnička, MM 9812B3;
5 September 2015, Prodašice, Mladá Boleslav City, with
Q. petraea and C. betulus, 50210 4100 N, 1570 4400 E, alt. 250 m
a.s.l., leg. J. Schneider, MM 5915B4. SLOVAK REPUBLIC—8 July 2008, Hontianske Nemce, Krupina, ‘‘Štiavnické vrchy00 Natural reserve, with Q. pubescens,
48180 700 N, 18580 5800 E, alt. 337 a.s.l., leg. M. Mikšı́k, MM
8708B1. HUNGARY—27 July 1997, Pilisszentkereszt,
Budapest, with Tilia sp., Q. cerris, Q. petraea, coord.
47420 N, 18550 E, alt. 450 m a.s.l., leg. A. Kiss, GS1881.
123
Fungal Diversity
ITALY—EMILIA-ROMAGNA. 16 September 1983,
Pulpiano, Viano (RE), with Q. cerris, coord. 44300 3200 N,
10330 3100 E, alt. 514 m a.s.l., leg. unknown, GS12. SARDINIA. 31 July 1990, Case Floris, Desulo (NU), with
Q. pubescens and C. sativa, coord. 40000 N, 9120 E, alt. 1066
a.s.l., leg. G. Simonini, GS735; 9 November 1994, Cala
Gonone, Dorgali (NU), with Q. ilex, coord. 40160 2200 N,
9360 4700 E, alt. 187 m a.s.l., leg. A. Montecchi, GS1310.
SICILY. 21 October 2013, Portella dei Bufali (Nebrodi),
Cesarò (ME), with Q. cerris and Q. pubescens, coord.
37520 1700 N, 14410 1700 E, alt. 1180 m a.s.l., leg. G. Simonini,
GS10112; 3 October 2014, Portella dei Bufali (Nebrodi),
Cesarò (ME), with Q. pubescens and Q. cerris, coord.
37520 1700 N, 14410 1700 E, alt. 1180 m a.s.l., leg. G. Vasquez,
GS10304; 28 June 2015, Portella Sella Maria (Nebrodi),
Cesarò (ME), with F. sylvatica and Q. pubescens, coord.
37540 100 N, 14390 1700 E, alt. 1420 m a.s.l., leg. G. Vasquez,
GS10245. RUSSIA—23 July 1983, vic. of township
Soglasie, Penza Oblast, with Q. robur on the carbonaceous
soil, leg. A.I. Ivanov, Le 200041; 8 August 2008, Voronka
River Reservoir, vic. of Podgorodnie Datchi Village,
Leninsky District, Tula Oblast, with Q. robur, coord. 54
60 500 N, 37320 5300 E, alt. 150 m a.s.l., leg. T. Svetasheva,
GS10317; 14 July 2011 (also 16 August 2011, 27 July
2012, 5 August 2012), protected area ‘‘Nizovie Plushchan
river’’, ‘‘Galichia Gora’’ Reserve, Krasninsky District,
Lipetsk Oblast, with Q. robur, leg. T. Svetasheva, L. Sarycheva; 1 September 2012, vic. of village Podgorodnie
Datchi, Leninsky District, Tula Oblast, with Q. robur,
coord. 54 60 500 N, 37320 5300 E, leg. T. Svetasheva, GS10311;
21 September 2012, vic. of khutor Leshchevsky, Leninsky
District, Volga-Akthuba Floodplain, Volgograd Oblast,
with Q. robur, coord. 48310 37.4400 N, 44510 42.9000 E, alt. 30
m a.s.l., leg. T. Svetasheva, GS10308; 14 October 2013,
loc. Shutov Ugol, vic. of khutor Leshchevsky, Leninsky
District, Volga–Akthuba Floodplain, Volgograd Oblast,
with Q. robur, coord. 48310 50.2900 N, 44550 25.9400 E, alt. 20
m a.s.l., leg. T. Svetasheva, GS10304. Rubroboletus
rubrosanguineus. AUSTRIA—10 August 1998, Thurntaler, Sillian, with Picea abies, coord. 46450 5400 N,
12240 3900 E, alt. 1800 m a.s.l., leg. G. Simonini, GS1930.
CZECH REPUBLIC—17 August 2014, Francova Lhota,
Beskydy Mountains protected area, with P. abies and Abies
alba, alt. 650 m a.s.l., leg. J. Polcak, GS10285. FRANCE—
date unknown, loc. unknown, with P. abies, leg. J.L.
Cheype, HOLOTYPE JLC 83209. ITALY—EMILIAROMAGNA 20 July 1987, Le Borelle, Villa Minozzo
(RE), with F. sylvatica, coord. 44180 N, 10240 E, alt. 1300 m
a.s.l., leg. L. Cocchi, GS405; 31 July 1987, Le Borelle,
Villa Minozzo (RE), with F. sylvatica, coord. 44180 N,
10240 E, alt. 1300 m a.s.l., leg. L. Cocchi, GS410; 1
September 1991, Costa delle Veline, Villa Minozzo (RE),
with F. sylvatica, coord. 44160 N, 10250 E, alt. 1700 m a.s.l.,
123
leg. A. Montecchi, GS828; 8 August 1995, M. Prampa,
Villa Minozzo (RE), with F. sylvatica, coord. 44200 N,
10260 E, alt. 1300 m a.s.l., leg. L. Cocchi, GS1387.
TRENTINO–ALTO ADIGE 13 August 1997, Pochi, Salorno (BZ), with P. abies and F. sylvatica, coord. 46140 N,
11150 E, alt. 1400 m a.s.l., leg. K. Kob, GS1818; 15 August
1998, Fai della Paganella, Andalo (TN), with F. sylvatica
and P. abies, coord. 46110 N, 11040 E, alt. 1100 m a.s.l., leg.
M. Comuzzi, GS1960; 28 August 1999, Vipiteno (BZ),
with P. abies, coord. 46540 N, 11250 E, alt. 1150 m a.s.l., leg.
unknown, GS2131; 25 August 1999, Waldheim, Sarnonico
(TN), with A. alba, coord. 46250 N, 11100 E, alt. 1230 m
a.s.l., leg. G. Simonini, GS2282. FRIULI VENEZIA–
GIULIA, 27 June 1998, Bosco Taggers, Lauco (UD), with
P. abies, coord. 46250 N 12570 E, alt. 950 m a.s.l., leg.
G. Zecchin, GS1899; 27 June 1998, Bosco Taggers, Lauco
(UD), with P. abies, coord. 46250 N 12570 E, alt. 950 m
a.s.l., leg. G. Zecchin, GS1900. VENETO, 10 July 1998,
Val di Cadore (BL), with P. abies, coord. 46240 N, 12 190 E,
alt. 950 m a.s.l., leg. P. Gaggio, GS1911; 11 July 1998, Val
di Cadore (BL), with P. abies, coord. 46240 N, 12 190 E, alt.
950 m a.s.l., leg. P. Gaggio, GS1912; 11 July 1998, Val di
Cadore (BL), with P. abies, coord. 46240 N, 12180 E, alt. 920
m a.s.l., leg. P. Gaggio, GS1913; 12 July 1998, Val di
Cadore (BL), with P. abies, coord. 46240 N, 12190 E, alt. 930
m a.s.l., leg. P. Gaggio, GS1917; 12 July 1998, Val di
Cadore (BL), with P. abies, coord. 46250 N, 12180 E, alt. 750
m a.s.l., leg. P. Gaggio, GS1918; 21 August 2012, Padola
(BL), with A. alba and F. sylvatica, coord. 46350 N, 12280 E,
alt. 1550 m a.s.l., leg. A. Testoni, GS10097. RUSSIA—19
August 2009, Arkhys, Teberda Reserve, valley of Kyzgich
river, Karachay–Cherkessia Republic, Caucasus, with
Abies nordmanniana. and F. sylvatica, coord. 43400 N,
41530 E, alt. 1050 m a.s.l., leg. T. Svetasheva, A. Kovalenko, GS10315 and GS10316.
GenBank Number ITS:KY677920.
Notes: During mycological surveys and mapping of
Northeast of Sicily, in the territory between mountain
ranges of Nebrodi, Peloritani and Madonie and the massif
of Mount Etna, several collections of a Boletaceae apparently close to Rubroboletus rubrosanguineus (Cheype)
Kuan Zhao & Zhu L. Yang of Northern Italy were
observed. They grew exclusively with broadleaved trees
and in the first instance they were included in the large
colour variability of R. legaliae (Pilát & Dermek) Della
Maggiora & Trassin. In the meantime, other sporadic finds
of the same bolete occurred in the region Calabria, in the
areas of Gambarie and Sila Piccola (Vasquez 2014).
However, since the first findings, it was observed that these
collections differ from these two neighbouring taxa in
chorological-ecological, anatomical and macroscopic
characters. The sharp delineation of the new species has
become clear in additional collections (Vasquez
Fungal Diversity
2012, 2013), to which some microscopic and molecular
differences from the other close, but different species have
been later added. Based on the BLASTn results (sequences
were selected based on the greatest similarity) and the
recent phylogenetic study focused on Rubroboletus (Zhao
et al. 2014), sequences were retrieved from GenBank
number and UNITE (http://unite.ut.ee/) databases for the
comparative phylogenetic analysis with the newly generated sequences. Caloboletus calopus (HM347645) was
chosen as an outgroup species. Phylogenetic analyses were
performed using the Bayesian Inference (BI) (MrBayes v.
3.2.2, Ronquist et al. 2012) and maximum likelihood (ML)
(RAxML v. 7.0.4, Stamatakis 2006; RAxML version 8,
Stamatakis 2014) approaches. The ITS data set comprised
52 taxa (24 from GenBank number and 14 from UNITE).
BI and ML trees were congruent and only the Bayesian tree
with both BYPP (Bayesian Posterior Probabilities) and
MLB (Maximum Likelihood Bootstrap) values is shown in
Fig. 152. In the analysis, the six sequences of R. demonensis form a highly-supported clade (BYPP = 1,
MLB = 100); it resulted as a distinct species, sister (with
low support, BYPP = 89, MLB = 54) to R. eastwoodiae.
Distinctive features of R. demonensis are the overall purple
blood red colour (general appearance close to
R. rubrosanguineus with brighter colours), combined with
frequent orange-yellow collar in the upper part of the stipe,
growth with broadleaved trees (Quercus, Castanea, Fagus)
and spores narrower than in the closer species, with a
higher Q ratio (13.1 ± 0.5 9 4.9 ± 0.2 lm, Qm = 2.68 ±
0.11). As in the most of the boletes, red colours can be in
some situations attenuate: this originates xanthoid aspect,
whose appearance is quite close to R. legaliae. R. legaliae
has a paler, whitish, greyish-cream pileus tending to pink
or reddish [R. legaliae f. spinarii (Hlaváček) Mikšı́k], more
velutinous, and shares in many occurrences in Sicily the
same environment of R. demonensis. It has also shorter but
stouter in average spores (12.5 ± 0.4 9 5.5 ± 0.1 lm,
Q = 2.29 ± 0.05) (533/13/13) and generally paler yellowish-orange pores, even if some bright red pores form do
exist. It grows with broadleaved trees, mainly with oaks, in
neutral-acidocline or basicline soils (Lannoy and Estades
2001; Estades and Lannoy 2004; Muñoz 2005; Andersson
2013; Halama 2016). It is nevertheless much more widely
distributed in Europe and its areal extends North to Denmark, East to Penza region (Russia), South to Sicily (Italy)
and West to Baixo Alentejo (Portugal), including: Austria,
Bulgaria, Corse, Croatia, Czech Republic, Denmark,
France, Germany, Great Britain, Hungary, Ireland, Italy,
Montenegro, Norway, Portugal, Romania, Russia, Sardinia,
Serbia, Sicily, Slovakia, Slovenia, Spain, Sweden,
Switzerland, Turkey (Courtecuisse and Duhem 2011;
Halama 2016). Its occurrence in Russia, until now has been
overlooked (Courtecuisse and Duhem 2011; Halama 2016),
but this species is known in Russia as rather rare but typical
for the middle and southern oakeries, associated with
Quercus robur in regions of Penza, Tula (Svetasheva
2010), Volgograd, Stavropol, Lipetsk (Sarycheva and
Svetasheva 2015). R. rubrosanguineus is a rare bolete
growing in submontane and mountainous regions, on
alkaline soil, in spruce and firs (Abies, Picea) woods, also
mixed with beech (Fagus), rarely in pure Fagus woods and
if so in the areas formerly populated by Abies; it has pileus
colour more purplish than R. demonensis, without so
intensive and bright red tones (Walty 1969; Cheype 1983;
Lannoy and Estades 2001; Estades and Lannoy 2004;
Muñoz 2005). It is widespread along the Alpine arch but its
distribution western limit is constituted by Spain (Palazon
2006) and reaches the most northern distribution in the
West Carpathians (Beskydy Mountains, Czech Republic),
even if a doubtful record could set this limit to Belgium
(Noordeloos 2010); at East, it extends to Russian Caucasus
(Kiyashko 2012; Svetasheva and Kovalenko 2013). The
southern limit seems to be constituted by the Italian
Appennini chain, in Emilia-Romagna region, where it
grows with Fagus in the areas earlier populated by Abies
alba. So, its distribution includes Austria, Belgium, Bosnia
and Hercegovina, Bulgaria, Croatia, Czech Republic,
France, Germany, Greece, Italy, Monte Negro, Poland,
western part of Russia, Slovakia, Slovenia, Spain,
Switzerland (Courtecuisse and Duhem 2011). R. rubrosanguineus presents spores as slender as those of R. demonensis, but longer and broader (14 ± 0.8 9 5.4 ± 0.3 lm,
Qm = 2.58 ± 0.12, (618/20/20).
From both R. rubrosanguineus and R. legaliae, R. demonensis can be distinguished by the net (reticulum) with
larger meshes and thicker ribs. R. rubrosanguineus has also
significantly broader pileipellis terminal elements (47.7 ±
4.9 9 7.5 ± 0.5 lm, Qm = 6.73 ± 0.49 (42/4/4)) with
respect to the other two boletes (37.9 ± 5 9 5.5 ± 0.4 lm,
Qm = 7.24 ± 0.89 (138/5/5) and 32.8 ± 0.1 9 5.7 ± 0., 5
lm Qm = 6.15 ± 0.51 (49/2/2) respectively for R. demonensis and R. legaliae). R. pulcherrimus (Thiers & Halling)
D. Arora, N. Siegel & J.L. Frank is a North-American
species apparently very similar, growing in in the North
Western mixed forests of the Coastal Chain (Arora et al.
1999; Desjardin et al. 2015); it has much larger spores
(14.3 ± 1.1 9 5.8 ± 0.1 lm, Q = 2.48 ± 0.15 (106/4/4)
(Arora et al. 1999).
The phylogenetically closest related Boletus eastwoodiae (Murrill) Sacc. & Trotter differs in the pale greyish
pileus, becoming olive-buff-grey and developing pink
tones especially along the margin. The stipe is massive,
abruptly bulbous with a ventricose base, with pinkish-grey
colour and a concolorous or vinaceous reticulation (Bessette et al. 2000; Desjardin et al. 2015). Due to the
molecular evidence offered in our work, Suillellus
123
Fungal Diversity
eastwoodiae Murrill is here combined in the genus
Rubroboletus.
Rubroboletus eastwoodiae (Murrill) Vasquez, Simonini,
Svetasheva, Mikšı́k & Vizzini, comb. nov.
Index Fungorum number: IF552780; Facesoffungi
number: FoF 2813, Fig. 155
Basionym: Suillellus eastwoodiae Murrill, N. Amer. Fl.
(New York) 9 (3): 152 (1910)
It is worth noting that the combination R. eastwoodiae
(Murrill) Arora, C.F. Schwarz & J.L. Frank (Frank 2015) is
invalid (wrong citation of the basionym). Curiously,
R. demonensis shares both the growth area and edaphicenvironmental conditions of another Mediterranean species
very common in Sicily and Calabria, Boletus flavosanguineus Lavorato & Simonini: frequently these boletes are
observed growing together (pers. obs.).
The analysis of the spore size distribution with the method
of the ‘‘isoprobability ellipses’’ (Simonini 1992, 1998;
Consiglio and Simonini 2005) (Fig. 155) is based on average
values of 20 collections of R. rubrosanguineus, 13 collections of R. legaliae, and 7 collections of R. demonensis. This
comparison allows to analyse better the differences of the
spores both in size and shape. It could be observed that spores
of R. demonensis and R. legaliae are clearly separated each
other even at confidence level of 95%, having R. demonensis
clearly shorter and narrower spores. R. rubrosanguineus has
some overlapping with the other species only at confidence
level of 95%, while at 68% there is almost no overlapping:
R. demonensis has narrower and stouter spores with respect
to R. rubrosanguineus, and the latter has slender and longer
spores with respect to R. legaliae. The difference in width of
the spores of R. rubrosanguineus and R. legaliae is statistically not significant.
Strobilomyces Berk.,
The genus Strobilomyces Berk. is a member of the family
Boletaceae and characterized by greyish or blackish basidiomata; presence of fibrillose squamules or scales on pilear
surface; stipe dry, woolly to shaggy, with or without annulus;
ornamented basidiospores (Bessette et al. 2010; Das et al.
2014). This genus includes nearly 40 taxa (Gelardi et al. 2012;
Wu et al. 2014; Antonı́n et al. 2015) around the globe. During a
macrofungal survey to East District of Sikkim (a small Himalayan state in India), an interesting member this genus was
collected and here it is described as a novel taxon (S. longistipitatus) with its morphology and phylogeny (Fig. 157).
Strobilomyces longistipitatus D. Chakr., K. Das & S.
Adhikari, sp. nov.
MycoBank number: MB 817357; Facesoffungi number:
FoF 2954, Figs. 158, 159 Etymology: referring to the very
long stipe (of basidiomata)
123
Holotype: CAL 1336
Diagnosis: Distinct from the European species Strobilomyces strobilaceous by its exceptionally long stipe,
significantly short tubes and frequent occurrence of tonguelike appendages on the wall of the ridges of basidiospores.
Pileus 37–50 mm. diam.; convex when young, becoming
pyramidal to convex with maturity; surface densely squamulose to floccose in numerous conical clusters of somewhat repent pattern, greyish-brown to violet brown
(11E3–11F4) to blackish, reddish with KOH; margin wavy
with sterile flap of tissue, brownish-black. Pore surface
covered with thick cottony partial veil when young, white
to dingy white; depressed near stipe, yellowish-white
(3A2), turning rusty or darker after bruising; pore 2 mm,
simple, angular. Tube 6 mm long, sinuate, chalky white or
yellowish-white (3A2), turning reddish-orange after bruising. Stipe 110–200 9 17–28 mm, central, dark grey to
(11F4) or black; surface cotton like or floccose with white
basal mycelium. Context solid in pileus and stipe; context
in pileus and stipe chalky white but immediately turning
pastel red (8A4) and then greyish-red (8C5) or darker on
exposure, turning orange (6A7) with KOH, greyish-green
(25E5) with FeSO4, greyish-green (25D4) with guiacol.
Spore print blackish-brown. Odour like Boletus edulis.
Basidiospores 10–11.1–12.7 9 8–9.1–9.9 lm (n = 20,
Q = 1.16–1.22–1.33), broadly ellipsoid to ellipsoid, ornamentation composed of high ridges forming complete
reticulum; under SEM with broad and confluent ridges (of
irregularly wavy walls) forming complete reticulations and
with frequent tongue like erect appendages on the wall of
ridges, blackish-brown. Basidia 40–58 9 14–18 lm, 2 to 4
spored, clavate to subclavate. Pleurocystidia 34–75 9 8–17
lm, pigmented, emergent 15–30 lm, common, subfusiform to fusiform, ventricose to ventricose with somewhat moniliform or capitate head; content dense, slightly
fibrillose. Tube edge fertile, composed of basidia, cystidia.
Cheilocystidia 39–65 9 11–13 lm, subclavate to subventricose, some are appendiculate, mostly hyaline. Tube
trama divergent, tramal hyphae 5–13.5 lm wide, gelatinous, encrusted, septate, branched. Pileipellis 1000–1200
lm thick, trichododerm to palisadoderm, composed of
erect elements, sometimes septate; terminal cells up to 20
lm wide, slightly thick walled (up to 0.8 lm), mostly with
rounded to fusoid apices, brown pigmented, sometimes
faintly encrusted; finely warted when observed under SEM.
Stipitipellis trichoderm to palisadoderm, hyphae brown
pigmented, wall up to 1 lm thick, often minutely encrusted; fertile, with basidia and cystidia. Caulobasidia
4-spored, similar to tube basidia. Caulocystidia 23–65 9
14–20 lm, clavate to subclavate, or fusoid to ventricose.
Clamp connections not found.
Habitat and distribution: Under Abies densa in subalpine mixed (broadleaf and coniferous) forest.
Fungal Diversity
Fig. 157 Phylogram generated from neighbour-joining analysis
based on LSU-rDNA sequences: Phylogenetic relationships inferred
from neighbour-joining analysis of partial nuclear large subunit
ribosomal RNA gene sequences of 15 taxa. The optimal tree with the
sum of branch length = 0.27898332 is shown. The percentages of
replicate trees in which the associated taxa clustered together in the
bootstrap test (1000 replicates) are shown next to the branches. The
tree is drawn to scale, with branch lengths in the same units as those
of the evolutionary distances used to infer the phylogenetic tree. The
novel species having GenBank Numbers KX364695 (LSU-rDNA) is
shown in bold and blue font. Boletus edulis (European material with
GenBank Numbers AF050643) was considered as the out group.
Evolutionary analysis was conducted in MEGA6 (Tamura et al. 2013)
Material examined: INDIA, Sikkim, East District,
Memainchu, 3692 m, N27210 25.000 E88360 24.900 , 4 July
2015, D. Chakraborty, DC 15-010 (CAL 1336, holotype).
GenBank Numbers ITS:KX364694.
Notes: Characteristic features of S. longistipitatus are
presence of short pileus with almost blackish dense squamules on surface in numerous conical clusters arranged
repently (never erect), exceptionally long stipe (3–4 times
or more of pileus diameter) without any striation or reticulation on the surface, distinguishingly short white sinuate
tubes which becomes reddish-orange on bruising, basidiospores with complete reticulum with the confluent ridges
and frequent occurrence of tongue-like appendages on the
ridge-walls and occurrence in association with the coniferous trees.
Our ITS based phylogeny (Fig. 156) clearly supports the
existence of two distinct and strongly supported clades
(Clade A and Clade B) in Strobilomyces which are also
reported repeatedly by earlier workers (Gelardi et al. 2012;
Antonı́n et al. 2015) however, another clade represented by
the only species S. annulatus Corner from Malaysia (sequence of ITS region is not yet available in GenBank) is
also known when phylogeny is conducted based on partial
sequences of the largest subunit of RNA polymerase II
(RPB2) as shown by Gelardi et al. (2012) and Antonı́n
et al. (2015). Here, Clade A includes S. confusus Singer, S.
seminudus Hongo, S. verruculosus Hirot. Sato where
basidiospores are never reticulated as also stated by Gelardi
et al. (2012) and Antonı́n et al. (2015). Distinguishingly,
Clade B represents S. strobilaceous (Scop.: Fr) Berk.,
S. appendiculatus sp. nov., S. echinocephalus Gelardi &
Vizzini., S. pteroreticulosporus Antonı́n & Vizzini, S. mirandus Corner where basidiospores are typically reticulate
(Gelardi et al. 2012, Antonı́n et al. 2015). Basidiosporeornamentation pattern is known to be the reliable parameter
in the delimitation of major groups of Strobilomyces.
Recently, on the basis of morphology and ITS data Petersen et al. (2012) showed the existence of only a single
species of Strobilomyces in Europe, as S. strobilaceus.
Therefore, we considered (Fig. 156) only the sequences
derived from European collections as S. strobilaceus sensu
stricto in our phylogeny. Our LSU based phylogeny which
was conducted considering all the Strobilomyces LSU
sequences appeared on BLAST (or available in GenBank)
search with our specimen (GenBank Numbers KF112459,
KF112460, KF112463, KF030345, DQ534626) and some
other species of Boletaceae also clearly supports the
position of the present specimen amongst Strobilomyces
and confirms the novelty (Fig. 157). Here three sequences
labelled as Strobilomyces sp. (Wu et al. 2014) are placed in
123
Fungal Diversity
Fig. 158 Strobilomyces longistipitatus (CAL 1336, holotype). a, c
Fresh basidiomata in the field and basecamp. b Pore surface.
d Pileipellis. e Terminal and subterminal cells of hyphal elements
of pileipellis. f Transverse section through tube showing
hymenophoral trama and hymenium. g Pleurocystidia. h SEM image
of Basidiospore. Scale bars d = 500 lm; e = 50 lm; f = 100 lm;
g = 10 lm; h = 5 lm
the weakly supported sister clade of present species
(94–95% identity with S. longistipitatus for 91% query
coverage using BLAST).
Morphologically, this Indian specimen is quite similar to
S. strobilaceous (an European species represented here by
GenBank Numbers JQ319001, JQ318998, JQ318984,
JQ318975, JQ318977 and JQ318988 in Fig. 156) and S.
echinocephalus (reported from China) but, the former
(probably the closest relatives of present species) differs
from S. longistipitatus by its considerably shorter stipe
[60–140 mm and 80–150 mm as mentioned by Breitenbach
and Kränzlin (1991) and Knudsen and Vesterholt (2012)
respectively] and longer tubes (10–15 mm) whereas, the
latter is distinct from the present Indian specimen by its
much shorter stipe (74–105 mm), presence of striation on
stipe apex and longer tube (9 mm). Phylogenetically, both
123
Fungal Diversity
Fig. 159 Strobilomyces
longistipitatus (CAL 1336,
holotype). a Basidiospores.
b Basidia. c Cheilocystidia.
d Pleurocystidia.
e Caulocystidia. f Elements of
pileipellis. Scale bars a = 5
lm; b–f = 10 lm
the species are well distinct as shown in Fig. 156. Three
more species: S. alpinus, S. pteroreticulosporus, S. mirandus are also partly close to S. longistipitatus. Strobilomyces
alpinus can be separated in field by the presence of longitudinal striations on stipe and unchanging exposed context and larger spores (12.5–15 lm) (Gelardi et al. 2012).
Strobilomyces pteroreticulosporus is distant by its pileus
surface which is covered with pyramidal fibrillose dirty
whitish scales, longer (30 mm) tubes and shorter (100–140
mm) stipe (Antonı́n et al. 2015) whereas, S. mirandus has
yellow or golden orange to brownish-orange pileus, longer
(up to 10 mm) tubes and distinctly smaller (40–70 mm)
stipe (Sato et al. 2005).
Strobilomyces polypyramis Hook. f., another species
recently reported from the state Sikkim can easily be separated by presence of erect, small conical to pyramidal,
pointed or rarely spinoid black scales, longer tube (up to 13
mm), shorter stipe (80 mm), absence of complete reticulum
in basidiospores (Horak 1980a, c; Das et al. 2014). Strobilomyces dryophilus Cibula & N.S. Weber (96% identity
with S. longistipitatus for 95% query coverage using
BLAST) and S. floccopus were also appeared close (though
well separated) in the LSU phylogeny along with our
Indian material (Fig. 157). But, S. dryophilus is morphologically distinct from S. longistipitatus by its whitish
ground coloured pileus with coarse, woolly appressed or
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Fungal Diversity
erect, greyish-pink or pinkish-brown scales, 10–17 mm
long tubes, and much shorter (40–80 mm) stipe (Bessette
et al. 2010). Whereas, S. floccopus (sequence derived from
European material) is considered as the synonym (Petersen
et al. 2012; Gelardi et al. 2012) of S. strobilaceous (morphological comparison given in the earlier paragraph).
According to unique combination of morphological data
and phylogenetic analyses of ITS and LSU sequences, our
specimen from India should be treated as an independent
species, proposed as S. longistipitatus which is placed in
Clade B (Fig. 156) along with other species featured with
reticulate basidiospores.
Cantharellales Gäum.
Clavulinaceae Donk
Clavulinaceae was described in 1970 by Donk to
accommodate the genus Clavulina, based on bisterigmate,
secondarily septate, and stichic basidia. Currently, it
includes four genera and belongs to Cantharellales (Kirk
et al. 2008). It is characterized by having simple, branched
or resupinate basidiomata, monomitic hyphal system,
clamped or not hyphae, and cystidia that is present in a few
species. The basidia are clavate or subcylindric, with
sterigmata generally incurved and the basidiospores are
usually globose, subglobose to ellipsoid, smooth, guttulate
and inamyloid. They are reported as saprobes, ectomycorrhizal and lichenized (Corner 1950; Donk 1964; Moncalvo et al. 2006; Bernicchia and Gorjón 2010).
Clavulina J. Schröt.
Clavulina J. Schröt. accommodates species usually with
branched basidiomata, occasionally simple, infundibuliform, effused-coralloid or resupinate, typically with two
smooth, hyaline, guttulate basidiospores per basidia, and
cornute sterigmata. Additionally, a septum is sometimes
formed at the base of the basidia after the release of
basidiospores (Corner 1950, 1970; Petersen 1988; Thacker
and Henkel 2004; Henkel et al. 2005; Uehling et al. 2012a).
Clavulina is a genus of ectomycorrhizal fungi with about
75 species widely distributed mainly in tropical regions
(Uehling et al. 2012b). Studies about the phylogeny of the
genus show its monophyly within Cantharellales (Thacker
and Henkel 2004; Moncalvo et al. 2006; Olariaga et al.
2009; Uehling et al. 2012a, b).
Clavulina grisea Meiras-Ottoni & Gibertoni, sp. nov.
MycoBank number: MB 818101; Facesoffungi number:
FoF 2955, Fig. 161
Etymology: grisea (Greek) = greyish, referring to the
colour of the basidiomata.
Holotype: URM 89967
Basidiomata weakly branched or simple, solitary or in
small groups, 2.5–8 cm, branches polychotomous, tips
123
unique or bifurcated to cristae, consistency fleshy when
fresh, brittle to crumbling when dry, grey to brownish-grey
(5C1, 5C2, 5D1) when fresh, stipe well to not delimited
1–2.3 cm, white, beige to light brown (6A1, 6E3, 6E4) when
fresh, young specimens white turning ochraceus (5A1, 5C3,
5C4) towards the apex of the branches, hymenium
amphigenous developing just below the branching point and
extending to the apex of the branches. Basidiospores globose
to subglobose, 5.5–8 (L = 6.63) 9 4–6.5 (W = 5.67) lm,
Q = 1.18, smooth, hyaline in water, pale yellowish in KOH
3%, with one large oleiferous guttule, thin-walled, with short
apiculus (0.7–1 lm), IKI-. Hyphal system monomitic, tramal
hyphae smooth, parallel, hyaline, 2–10 lm, slightly inflated,
clamp connections abundant, subhymenium tightly interwoven. Cystidia hymenial hyaline, cylindrical with homogeneous to granular cytoplasmic contents, 85–98 9 5, 5–10
lm, no clamp connection at the base. Basidia elongate-clavate, approximately 31 9 5.5–6 lm, hyaline, guttulate, with
2 cornute sterigmata, 4–6 lm, clamped at the base, but
agglutinated and difficult to observe.
Material examined: BRAZIL, Acre: Sena Madureira,
Floresta Nacional de São Francisco, January 2015, A.
Meiras-Ottoni & S.O. Almeida (URM 89966); BRAZIL,
Acre: Sena Madureira, Reserva Extrativista Cazumbá-Iracema, January 2015, A. Meiras-Ottoni & S.O. Almeida
(URM 89967, holotype; isotype in O); BRAZIL, Acre:
Sena Madureira, Reserva Extrativista Cazumbá-Iracema,
January 2015, A. Meiras-Ottoni & S.O. Almeida (URM
89968, paratype).
GenBank Numbers URM 89967 ITS:KX811199, URM
89968 ITS:KX811184.
Notes: This species is characterized by the slender,
flexible, fleshy and greyish basidiomata when fresh, brittle
when dry, and by the hymenial, hyaline cystidia with
homogeneous to granular contents. Clavulina grisea composed of several solitary basidiomata growing on soil
covered by rotten leaves and branches.
Other species of Clavulina with greyish basidiomata and
cystidia have been reported by Corner et al. (1956), Corner
(1970) and Petersen (1983). Clavulina hispidulosa Corner,
however, has subhyaline to pale yellow or pale brownish,
subglobose to obovoid, larger basidiospores (8–12 9 7–10.5
lm), while C. griseopurpurascens Corner has hyaline, usually ellipsoid basidiospores (7.5–8.7 9 6–7 lm).
The specimens of C. grisea are grouped in a well-supported clade (100% ML/100% MP/1.00BYPP) sister of
C. cristata (80% ML) (Fig. 160). Clavulina cristata
(Holmsk.) J. Schröt. is characterized by white, often
becoming yellowish, ochraceous or fuliginous basidiomata,
branches with acute tips usually becoming cristate, and larger, slightly thick-walled basidiospores (7–11 9 6.5–10 lm).
Another greyish species in the phylogenetic analyses
(Fig. 160) are C. cinerea (Bull.) J. Schröt., C. craterelloides
Fungal Diversity
Fig. 160 Phylogenetic tree of the Clavulina obtained by analyses
from rDNA sequences. Taxa with two accession numbers were
analysed by concatenated ITS and partial LSU rDNA; both regions
ITS and partial LSU rDNA was used to analyse the other individuals.
Support values (from top) are maximum likelihood, maximum
parsimony 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 Hydnum albomagnum, H. repandum and
H. umbilicatum
Thacker & T.W. Henkel and C. griseohumicola T.W. Henkel, Meszaros & Aime, all without cystidia. Additionally,
C. cinerea has yellow or ochraceous, subglobose or broadly
ellipsoid, basidiospores (6.5–11 9 6–10 lm); C. craterelloides has infundibuliform basidiomata, and hyaline, subglobose to broadly ellipsoid, basidiospores (6.5) 7.5–8 9
(5.5) 6–7 (7.5) lm; and C. griseohumicola has dark bluishgrey, simple basidiomata, and subglobose basidiospores
(8–9.5 9 7–9 lm) (Fig. 161).
Etymology: ossea (Latin) = bone, referring to appearance basidiomata after drying.
Holotype: URM 89970
Basidiomata branched, solitary or in small groups, with
fleshy consistency, –10 cm, polychotomous branches, the
tips of branches are unique or rarely cristate, becoming
dark, stem visible, sterile, when fresh dark brown (8F6),
branches brownish-lilac grey (10C2, 16B2), hymenium
amphigenous starting just below the branch point and
extending to the apex of the branches. Basidiospores globose to subglobose, 6–9 (L = 7.66) 9 5–7.5 (W = 6.56)
lm, Q = 1.14, smooth, hyaline, with one large oleiferous
guttule, thin-walled, with short apiculus (0.2–1 lm), IKI-.
Clavulina ossea Meiras-Ottoni & Gibertoni, sp. nov.
MycoBank number: MB 818103; Facesoffungi number:
FoF 2956; Figs. 162
123
Fungal Diversity
Fig. 161 Clavulina grisea
(URM 89967, holotype). a, b
Basidiomata. c Hyphae with
clamps (KOH). d Basidiospores
with apiculus (KOH). e Cystidia
(in Melzer’s reagent). Scale
bars a, b = 5 cm, c–e = 10 lm
Hyphal system monomitic, tramal hyphae smooth sometimes verrucous, parallel, hyaline, 3–10 lm, some hyphae
are wide but not inflated, clamp connections abundant,
subhymenium tightly interwoven. Cystidia absent. Basidia
cylindrical, 21–44 9 5–6 lm, hyaline, multi-guttulate, with
2 cornute sterigmata, 4–5 lm, postpartal septa observed on
most basidia.
Material examined: BRAZIL, Paraı́ba: Areia, Parque
Estadual Mata do Pau-Ferro, June 2015, C.O. Mendonça
(URM 89970, holotype; isotype in O).
GenBank Number ITS:KX811197.
Notes: This species is characterized by the robust, fleshy, and pale lilac basidiomata when fresh, resembling
bones when dry, and hyphae occasionally incrusted. The
new species is composed of several solitary or in pairs
basidiomata, scattered on soil covered by rotten leaves and
branches. Of the species of Clavulina with similar colour
(vinaceous, lilac or purple tints), C. incrustata can be
distinguished by the basidiomata with very rare clamp
123
connections and ferruginous brown, incrusted hyphae
(Wartchow 2012); C. amethystina (Bull.) Donk by the
ovoid-ellipsoid or subglobose, slightly larger basidiospores
(7–11(–12) 9 6–8 lm), while C. amethystinoides (Peck)
Corner by the subglobose to broadly ellipsoid (7–10 9 6–8
lm). In all phylogenetic analyses, C. ossea comes an isolated position within the genus (Fig. 160).
Clavulina paraincrustata Meiras-Ottoni & Gibertoni, sp.
nov.
MycoBank number: MB 818102; Facesoffungi number:
FoF 2957, Figs. 163
Etymology: paraincrustata (Greek ? Latin) = similar to
Clavulina incrustata, referring to the extracellular incrustations on the hyphae of the context.
Holotype: URM 89969
Basidiomata branched in caespitose clusters, more
rarely solitary or in pairs, somewhat fused basally, initially
monopodial but branching early, with fleshy consistency,
Fungal Diversity
Fig. 162 Clavulina ossea (URM 89970, holotype). a, b Basidiomata. c, d Incrusted hyphae with clamps (Melzer, KOH ? floxine, respectively).
e basidiospores (KOH ? floxine, KOH, Melzer’s, respectively). Scale bars a = 1 cm, b = 2 cm, c–e = 10 lm
Fig. 163 Clavulina paraincrustata (URM 89969, holotype). a Basidiomata. b Incrusted hyphae (KOH). c Basidia (KOH). d Basidiospores
(KOH). Scale bars a = 2.5 cm, b, d = 10 lm, c = 20 lm
123
Fungal Diversity
-8 cm, branches occasionally hollow, tips of branches
cristate, stipe visible, sterile, when fresh brownish-orange
(6C7, 6C8), branches violet brown (11F8), hymenium
unilateral, appearing initially above first branching point.
Basidiospores globose to subglobose, 6–8 (L = 7.49) 9
5–7 (W = 6.32) lm, Q = 1.20, smooth, hyaline, with one
large oleiferous guttule, thin-walled, with short apiculus(–1
lm), IKI-. Hyphal system monomitic, tramal hyphae frequently incrusted, parallel, very compact, brownish, 2–10
lm, not inflated, without clamp connections. Cystidia
absent. Basidia elongate-clavate, 31–48 9 4, 5–7 lm,
hyaline or with granular cytoplasmic contents, with 2
cornute sterigmata, rarely 3, 4–5 lm, postpartal septa
observed on most basidia.
Specimen examined: BRAZIL, Paraı́ba: Rio Tinto e
Mamanguape, Reserva Biológica Guaribas, May 2015, A.
Meiras-Ottoni (URM 89969, holotype; isotype in O).
Additional specimen examined: Brazil, Pernambuco:
Igarassu, Refúgio Ecológico Charles Darwin, May 2010, F.
Wartchow (URM 82947).
GenBank Numbers ITS:KX811201; LSU:KX811196.
Notes: This species is characterized by the robust and
violet brown basidiomata when fresh, brittle when dry, and
incrusted hyphae. It is composed of several solitary or in
pairs basidiomata, scattered on soil covered by rotten
leaves and branches.
Wartchow (2012) described C. incrustata Wartchow,
the first species of Clavulina with incrusted hyphae, based
in material collected in Brazil. Despite being microscopically similar, it differs from C. paraincrustata by the less
robust, paler basidiomata, and the amphigenous hymenium.
DNA of the type was tentatively extracted, without
success.
Clavulina paraincrustata was grouped (94% ML/94%
MP/1.00BYPP) with C. craterelloides and C. rosiramea
Uehling, T.W. Henkel & Aime (Fig. 160), from which can
be distinguished by the lack of incrusted hyphae. Additionally, C. craterelloides has infundibuliform basidiomata,
subglobose to broadly ellipsoid basidiospores [(6.5) 7.5–8
9 (5.5) 6–7 (7.5) lm], while C. rosiramea has simple or
slightly branched basidiomata, with orange tints, subglobose to slightly triangular [(6.5) 7–9 (10) 9 (6) 7–8 (9)
lm].
Polyporales Gäum.
Fomitopsidaceae Jülich
Fomitopsidaceae is a family of Polyporales and
currently comprises 24 genera and 197 species (Kirk
et al. 2008), with Fomitopsispinicola (Sw.) P. Karst as
the type species. The species are characterized by the
basidiomata annual or perennial, resupinate, applanate,
effuse or pileate, woody, leathery or corky in texture.
The hymenium is poroid, and the hymenial surface is
123
usually pale or brownish. The hyphal system is di- or
trimitic and clamp connections are usually present
(Cannon and Kirk 2007). Most genera of Fomitopsidaceae such as Antrodia P. Karst. Fomitopsis P. Karst.,
Daedalea Pers., Laetiporus Murrill are of ecological,
evolutionary and economic importance because this
group has many forest pathogens and cause brown rot
which also play an important role in carbon sequestration (Fukami et al. 2010; Ortiz-Santana et al. 2013;
Zhou and Wei 2011).
Fomitopsis P. Karst.
Fomitopsis includes species with basidiomata sessile to
effused-reflexed, rarely annual to perennial, varying from
white to purple, and di- to trimitichyphal system with
clamped generative hyphae. The basidiospores are smooth
and hyaline, thin-walled, ranging from subglobose to
cylindrical (Gilbertson and Ryvarden 1986; Ryvarden and
Gilbertson 1993; Dai 2012). It is a cosmopolitan genus,
causing brown rot in the substrates, and several species
were used as biological models in genetic analyses, as well
as ecological, biotechnological and taxonomic studies
(Högberg et al. 1999; Chlebicki et al. 2003; Aime et al.
2007; Watanabe et al. 2010) (Fig. 164).
Fomitopsis flabellata Soares & Gibertoni, sp. nov.
MycoBank number: MB 817408; Facesoffungi number:
FoF 2958, Figs. 165
Etymology: flabella (Latin) = The name refers to the
fan-shaped basidiomata.
Holotype: URM 89405
Basidiomata annual, biannual, pileate, dimidiate, solitary or in small groups or clusters. Pileus 0.2–2 cm wide,
up to 2 cm long, 0.3 mm thick at base, upper surface purple
(9.1E purple) when fresh and dried, azonate to zonate,
glabrous to slightly velutinate. Margin obtuse, entire. Pore
surface pinkish-brown to lilac-violaceous (9.4D pinkishbrown, 11.5E lilac violaceous), pores round to angular (3–4
mm), dissepiment thin and entire. Context purple brownish
(9.1F brownish), 0.1 mm thick.
Basidiospores cylindrical to sub-cylindrical, 4–5 (6.5) 9
2–2.5 lm, smooth, thin-walled, IKI–. Basidia clavate with
four sterigmata, 8–10 (12) 9 4–5 lm. Hyphal system
trimitic, generative hyphae clamped, thin-walled, 2–3 lm
in diam.; skeletal hyphae dominant, thick-walled, hyaline,
3–5 lm in diam.; binding hyphae thick-walled, hyaline,
1.5–2.5 lm in diam. Cystidia absent.
Specimens examined: BRAZIL, AMAPÁ: Porto Grande,
Floresta Nacional do Amapá, October 2014, A. Soares
(URM 89405, holotype; Isotype in O). BRAZIL, Amapá:
Porto Grande, Floresta Nacional do Amapá, September
2013, A.M. Soares (URM 84210).
GenBank Numbers ITS:KX423688; LSU:KX423686.
Fungal Diversity
Fig. 164 Maximum likelihood phylogenetic tree of the concatenated
ITS1, 5.8S, ITS2 and partial 28S rDNA sequences. Bootstrap
supporting values (1000 replicates) and posterior probabilities (PP)
from Bayesian analysis to each node are shown from left to right.
Only bootstrap values above 50% and PP above 0.75 are provided.
The new species described in this study are highlighted in blue. The
tree was rooted with Grifola sordulenta and Perenniporia ochroleuca
123
Fungal Diversity
Fig. 165 Fomitopsis flabellata (URM89405, holotype). a Abhymenial surface. b Hymenial surface. c Pores. d Hyphae context. e Basidiospores.
Scale bars a, b = 1 cm, c = 2 mm, d = 20 lm, e = 5 lm (photos by R.L. Alvarenga)
Notes: Fomitopsis flabellate can be recognized by the
small, usually fan-shaped, coriaceous, brown-vinaceous
basidiomata. Fomitopsis lilacinogilva (Berk.) J.E. Wright
& J.R. Deschamps and F. cupreorosea (Berk.) Carranza &
Gilbn. have a similar colour, but differ by the larger and
thicker basidiomata (2–20 9 2–9 9 0.5 9 10 cm; 5–16.5 9
2.6–10.3 9 0.3–2.5 cm, respectively). Besides, the pores in
F. lilacinogilva are slightly labyrinthiform (1–2 mm) and
sub-daedaleoid, becoming sinuous-daedaleoid in F. cupreorosea (1–3 mm), while in F. flabellata they are round to
123
angular. Fomitopsis flabellata is placed as a basal taxon of
the Rhodofomitopsis sensu stricto, but with low statistical
support (Fig. 164). For the time being, we prefer to keep
this species in Fomitopsis until new sequences are added to
the Antrodia clade phylogeny.
Fomitopsis roseoalba Soares, Ryvarden & Gibertoni, sp.
nov.
MycoBank number: MB 812876; Facesoffungi number:
FoF 2959, Fig. 166
Fungal Diversity
Fig. 166 Fomitopsis roseoalba
(URM 86923, holotype). a, b
Fresh basidiomata. c Dried
basidiomata. d Basidia.
e Basidiospores. f Context.
Scale bars a–c = 1 cm, d,
e = 2 lm, f = 10 lm (photos
by T.B. Gibertoni)
Etymology: The name refers to the colour of basidiomata
when fresh, with pore surface white, and upper surface,
when present, pink to lilac.
Holotype: URM 86923
Basidiomata annual, pileate, resupinate to effused-reflexed. When resupinate, up to 32 cm long, 11 cm wide, 0.3
cm thick at centre. Pileus when present, 0.2–7 cm wide, 5
cm long, 0.3 cm thick at base, upper surface white to pink
(3.1A white, 7.4B pink) when fresh, becoming cream to
greyish when dried (3.3A pale yellow, 1.1C pastel grey),
slightly velutinate. Context white to cream (3.1A white,
3.3A pale yellow), 0.4 cm thick. Margin narrow in
123
Fungal Diversity
resupinate specimens, obtuse in pileate specimens. Pore
surface white to cream (3.1A white, 3.3A pale yellow)
when fresh and ochraceous when dried (4.4A light yellow),
pores circular to angular (4–6 mm), dissepiment thin and
entire.
Basidiospores ellipsoid to sub-cylindrical, 3–4 9 1.8–2
lm, smooth, thin-walled, IKI. Hyphal system trimitic,
generative hyphae clamped, thin-walled, 2.5–3 lm in
diam.; skeletal hyphae dominant, thick-walled, hyaline,
3.75–5 lm in diam.; binding hyphae thick-walled, hyaline,
1.5–2 lm in diam. (Fig. 166a–f). Cystidia absent. Basidia
clavate with four sterigmata, 10–12 9 5–7 lm.
Material examined: BRAZIL, Amapá: Porto Grande,
Floresta Nacional do Amapá, on dead hardwood, 00 58.
25.8 N and 051 41 73.6 W, February 2014, A. Soares, AS
1496 (URM 86923, holotype, Isotype O); Porto Grande,
Floresta Nacional do Amapá, on dead hardwood, September 2013, A. Soares (URM 86913), A. Soares (URM
86926); February 2014, A. Soares (URM 86918), A.
Soares, AS 1431 (URM 86920) A. Soares, AS 1457 (URM
86921), A. Soares, AS 1566 (URM 86922), A. Soares
(URM 86923), A. Soares (URM 86925); Brazil, Pará:
Melgaço, Floresta Nacional de Caxiuanã, July 2006, T.B.
Gibertoni (URM 79982, 79984, 79985, 79986, 79987,
79988, 79989, 79990), March 2007, T.B. Gibertoni (URM
79983, 79991, 79993, 79994, 79995, 79996), August 2007,
T.B. Gibertoni (URM 79997, 79998, 79999, 80000, 80001,
80002, 80003, 80004), February 2008, T.B. Gibertoni
(URM 79992, 80005, 80006, 80007, 80008), August 2013,
A. Soares (URM 86912) A. Soares (URM 86914) A.
Soares (URM 86915), A. Soares (URM 86916).
GenBank Numbers ITS:KT189139; LSU:KT189141.
Notes: This species is easily recognized by the white to
pink, pileate to effuse-reflexed basidiomata. Of the
neotropical species, F. nivosa (Berk.) Gilbertson and
Ryvarden has similar whitish colour and morphology
(pileate, and sometimes effused-reflexed), but is separated
by having cylindrical basidiospores (6–9 9 2–3 lm) and
smaller pores (6–8 mm). Despite herbaria revision and
several collections in other parts of the Brazilian Amazonia, F. roseoalba was only found in the Eastern Brazilian
Amazonia, what indicates that this species may be
restricted to this region. According to our molecular analyses, F. roseoalba grouped into a well-supported clade (81/
0.99) with one unidentified, endophytic Fomitopsis isolated
from Hevea brasiliensis sapwood collected in Brazil
(Fomitopsis sp. NHB31) (Martin et al. 2015). This isolate
may represent an endophytic stage of F. roseoalba. The
F. roseoalba clade grouped in a major clade (98/1.00) with
F. subtropica B.K. Cui, Hai J. Li and M.L. Han and two
unidentified, endophytic Fomitopsis isolated from Elaeis
guineensis from Thailand (Fomitopsis sp. 9 V 3.1 and
Fomitopsis sp. 7R 8.1) (Rungjindamai et al. 2008), all
123
included in the Fomitopsis sensu stricto (75/1.00).When in
a resupinate form, F. roseoalba is whitish as F. subtropica,
and both have similar basidiospores (cylindrical to oblongellipsoid 3–4.7 9 1.7–2.3 lm in F. subtropica). Fomitopsis
subtropica, however, has smaller pores (6–9 mm) and the
pileus, when present, is white to cream (Li et al. 2013b). Li
et al. (2013b) suggested that the two specimens of Fomitopsis sp. from Thailand (7R 8.1, 9 V 3.1) could be
F. subtropica, because of the high supporting values uniting them in their study (100/1.00). Although the phylogenetic relationship of these three Fomitopsis species was
strongly supported, our analyses indicates that F. subtropica, F. roseoalba and Fomitopsis sp. (7R 8.1, 9 V 3.1) are
clustered in an independent way (Fig. 164).
Polyporaceae Fr. ex Corda
Favolus (Fr.) Sotome & T. Hatt.
The genus Favolus, typified by F. brasiliensis (Fr.) Fr, is
a wood decaying fungus within the Polyporaceae, Polyporales. Previously, Núñez and Ryvarden (1995) treated
Favolus as one of the six infrageneric groups of the genus
Polyporus, and recently Sotome et al. (2013) proposed as a
new genus based on phylogenetic analyses of LSU and ITS
regions and morphological characters such as a radially
striate pileus, and lack of distinct cutis of agglutinated
hyphae. This genus is well supported by multi-gene phylogenies (Binder et al. 2013; Seelan et al. 2015) (Fig. 167).
Favolus gracilisporus H. Lee, N.K. Kim & Y.W. Lim, sp.
nov.
Index Fungorum number: IF552508; Facesoffungi
number: FoF 2626, Fig. 168
Etymology: referring to a slenderer spore relative to
other Favolus species.
Holotype: SFC20130704–40
Basidiocarps annual, lateral stipitate; pileus circular to
flabellate, up to 40 mm in diam., up to 3.5 mm thick; pileal
surface ivory to pale buff, with age becoming brown, fibrillose with flattened, radially striate, azonate, slightly
glabrous, margin concolorous, acute, lacerate. Hymonophore cream to pale brown when fresh, pores diamond-shaped, radially elongated, 2–3 mm long, 1–2 mm
wide near stipe to 2–3 per mm at margin; dissepiments thin
and lacerate with age. Context in pileus pale tan to ivory,
corky, brittle when dry, up to 0.5 mm thick. Tube continuous with the context, up to 2–3 mm thick. Stipe lateral,
buff, glabrous, up to 7 mm long and 6 mm thick. Hyphal
system dimitic; generative hyphae hyaline in KOH, thinwalled, 3–4 lm in diam., septa with clamps, skeletalbinding hyphae thick-walled, aseptate, much branched,
with tapering apices, 3–6 lm diam. Cystidia absent. Basidia clavate, 4–sterigmate (up to 4.5 lm), 27–35 9 5.7–8
lm, with a basal clamp. Basidiospores cylindric, hyaline,
Fungal Diversity
Fig. 167 Phylogram generated from maximum likelihood (RAxML)
analysis based on ITS sequence data of Favolus. Maximum likelihood
bootstrap support values greater than 70% are indicated above or
below the nodes and the new species is in bold. The tree is rooted with
Neofavolus mikawai and N. alveolaris
Fig. 168 Favolus gracilisporus
(SFC20130704–40, holotype).
a, b Fruiting bodies in the field
(SFC20130704–40). c Pores of
fruiting body
(SFC20130704–40).
d Microscopic structure;
basidiospores, basidia,
generative hyphae and skeletalbinding hyphae
123
Fungal Diversity
smooth, IKI-, 7.9–9.7 9 2.5–3.3 lm. L = 9 lm, W = 2.9
lm, Q = 3.11 (n = 20/1).
Habitat: solitary to gregarious on dead wood of
hardwood.
Material examined: KOREA, Gyeongsangbuk-do,
Yecheon-gun, Mt. Hagga, on rotten angiosperm stump, 4
July 2013, Y.W. Lim (SFC20130704–40, holotype).
GenBank Number ITS:KY038472.
Notes: Favolus gracilisporus shares similar characteristics with Favolus brasiliensis (Fr.) Fr., Favolusroseus
Lloyd, and Neofavolusalveolaris (DC.) Sotome & T. Hatt.
such as lateral stipitate basidiocarps with radially elongated
and hexagonal pores; however, F. gracilisporus has distinctly larger basidia (27–35 9 5.7–8 lm) compared to
F. roseus (18–23.8 9 4.5–7 lm, Sotome et al. 2013) and
N. alveolaris (17.5–26 9 4–7 lm, Sotome et al. 2013).
Microscopic features of F. brasiliensis are somewhat
similar to F. gracilisporus, but the latter species has a
larger basidiospore Q value (3.11) than F. brasiliensis
(Q = 2.75, Sotome et al. 2013). Phylogenetic analysis
based on the ITS region indicated that F. gracilisporus
separated distinctly from the other three species (Fig. 167).
Lentinus Fr.
Lentinus (Fr.) Quel is a cosmopolitan genus with an
estimated 63 species (Kirk et al. 2008) and 629 records
under the name of Lentinus in the Index Fungorum (2017)
and, species are able to survive over a wide temperature
range, are abundant in boreal, temperate and tropical
regions (Pegler 1983; Corner 1981; Karunarathna et al.
2011) (Fig. 169).
Lentinus sajor-caju (Fr.) Fr., Epicrisis Systematis Mycologici: 393 (1838)
Facesoffungi number: FoF 3142, Figs. 170, 171
Pileus 3–9 cm diam., coriaceous drying hard and rigid,
soft, convex with a deeply umbilicate centre then
cyathiform to infundiliform, or eccentric and flabelliform,
surface very variable in colour, cream colour, pale
ochraceous, more or less fuliginous, or umbrinous, dry
glabrous and smooth or sometimes with small, appressed,
darker squamules especially towards the centre, often
finely radially striate, rimose in old specimens; margin
initially incurved to involute soon straight, very thin,
smooth, undulating and lobed. Lamellae deeply decurrent,
not furcate, whitish, concolorous with the pileus, or
becoming darker towards the edge, often darkening on
drying, narrow to sub-linear, 0.5–3 mm broad, densely
crowded, with lamellulae of 4–6 lengths; edge entire or
finely denticulate. Stipe central, eccentric or lateral, short,
0.5–3 9 0.5–1.5 cm, cylindric with an abrupt base, solid;
surface concolorous with the pileus, at times blackening
123
especially towards the base. Annulus present, attached
towards stipe apex, firm, white to fulvous, either well
formed, slight or commonly weathered away. Context up
to 8 mm thick at the centre but very thin over the
hymenophore, tough and pliant drying hard and horny,
white, consisting of a very tightly woven, dimitic hyphal
system with skeleton-ligative hyphae. Generative hyphae
2–5 lm diam. not inflated, very thin-walled, frequently
branching with clamp connections. Skeleto-ligative
hyphae dominant, 2–8 lm diam., hyaline or pale yellowish, with a thick wall of 3 lm and often a very narrow
lumen, comprising a skeletal element, 100–400 lm long,
bearing two to several tapering ligative branches, up to
400 lm long, which are themselves dichotomously divided, sometimes the branches are very short, numerous and
nodulose to produce a coralloid appearance. Basidiospores 5–9 9 1.5–2.5 lm, Q = 3.23, narrowly cylindric, often curved, hyaline, thin-walled, with few
contents. Basidia 15–20 9 3.5–5 lm, very narrow, clavate, cylindric, bearing 4 sterigmata. Lamella-edge a
broad sterile zone, with scattered or clustered Cheilocystidia and numerous, emergent skeleto-ligative branches.
Cheilocystidia 20–25 9 4–6 lm, clavate, often sinuous or
nodulose, hyaline, thin-walled. Hyphal pegs abundant,
50–100 9 20–40 lm, truncate cylindric, consisting of
fascicles of up to 50 narrow hyaline hyphae, extending up
to 80 lm beyond the basidia. Hymenophoral trama
irregular, hyaline, broad, of radiate construction, very
compactly interwoven with few interhyphal spaces, and
ligative branches frequently penetrating the hymenium.
Subhymenial layer often indefinite or indistinct, becoming
pseudoparanchymatous when well developed. Pileipellis
an epicutis, 15–20 lm thick, of agglutinated, radiating
generative hyphae, often with a brown encrustation and
sometimes a brown membrane pigment. Smell mushroomy and an excellent edible species.
Habit, habitat and distribution: as a cluster, usually on
dead or decaying wood, May–August. Our collection was
collected on a decaying wood near Peradeniya Royal
Botanic Gardens.
Specimens examined: SRI LANKA, Kandy District, near
Peradeniya Royal Botanic Gardens, 23 June 2012,
Samantha C. Karunarathna (MFLU 12-1478).
GenBank Number ITS:KY649463.
Notes: Lentinus sajor-caju is one of the most common
macrofungi of the palaeotropical forests, with the distribution extending from equatorial and southern Africa,
although less frequent in West Africa to throughout southeast Asia and down to the north-east corner of Australia.
This is the first report of L. sajor-caju with the molecular
phylogenetic confirmation from Sri Lanka Pegler (1983)
has previously reported it (Fig. 169).
Fungal Diversity
Fig. 169 Phylogenetic
relationships inferred from
maximum parsimony analysis
of ITS-rDNA sequences of 36
taxa. The percentages of
replicate trees in which the
associated taxa clustered
together in the bootstrap test
(1000 replicates) are shown next
to the branches. The tree is
drawn to scale, with branch
lengths in the same units as
those of the evolutionary
distances used to infer the
phylogenetic tree. The new Sri
Lankan records: Lentinus sajorcaju, having GenBank Number
KY649463 and herbarium
number MFLU 12-1478;
Lentinus squarrosulus, having
GenBank Number KY649464
and Herbarium Number MFLU
12-1228; Lentinus velutinus,
having GenBank Number
KY649465 and Herbarium
Number MFLU 12-1123 are
shown in bold and blue. The
topology is rooted with Panus
lecomtei. Evolutionary analysis
was conducted in PAUP 4.0b 10
123
Fungal Diversity
Habit, habitat and distribution: as a cluster, usually on
dead or decaying wood, May–August. Our collection was
collected on a decaying wood near Peradeniya Royal Botanic
Gardens.
Specimens examined: SRI LANKA, Kandy District, near
Peradeniya Royal Botanic Gardens, 13 June 2012,
Samantha C. Karunarathna (MFLU 12-1228).
GenBank Number ITS:KY649464.
Notes: This is the first report of L. squarrosulus with the
molecular phylogenetic confirmation from Sri Lanka after
Pegler (1983) has previously reported it.
Fig. 170 Basidiocarps of Lentinus sajor-caju in the field (MFLU
12-1478)
Lentinus squarrosulus Mont., Annales des Sciences Naturelles Botanique 18: 21 (1842)
Facesoffungi number: FoF 3143, Figs. 172, 173
Basidiocarps moderate size. Pileus 2–8 cm diam., thin
and pliant, convex soon depressed; surface milk white,
fuscous brown cream coloured or pinkish-buff, radially
innate striate, with concentric zones of small squamules,
margin downcurved. Lamellae deeply decurrent, occasionally slightly interveined towards the stipe attachment,
white to pale buff, thin, 2–3 mm broad, moderately
crowded, 4–5 tiers of lamellulae, edge entire or finely
denticulate on large specimens. Stipe 1.5–5 cm 9 2–6 mm,
central or excentric, rarely lateral, tapering below, firm,
solid, cylindric; surface concolorous with pileus. Context
2–3 mm thick, fleshy coriaceous, white, consisting of a
dimitic hyphal system with generative and skeletal hyphae.
Generative hyphae 2–6 lm diam., not inflated or scarcely
inflated, hyaline, thin walled, frequently branched, with
prominent clamp connexions. Skeletal hyphae 2–10 lm
diam., with a hyaline, thickened wall, but always with a
continuous lumen, and normally branching. Basidiospores
5.5–7.5 9 1.8–2.7 lm [n = 40, (6.2 9 2.3 lm), Q = 2.69],
straight, cylindrical, hyaline, thin walled with few contents.
Basidia 15–21 9 4–6 lm, narrowly clavate, bearing 4
sterigmata. Hyphal pegs abundant.
123
Lentinus velutinus Fr., Linnaea 5: 510 (1830)
Facesoffungi number: FoF 3144, Figs. 174, 175 Pileus
2–10 cm diam., thin, coriaceous, deeply umbilicate to
broadly infundibuliform or cyathiform; surface uniformly
pale greyish-cinnamon to rufous or tawny brown, fuscous at
the centre, at times almost chestnut brown, dry, uniformly
velutinate to short hispid or subsquamulose-furfuraceous,
not zonate or obscurely so, neither strongly striate nor sulcate
except in very old specimens; margin thin, at first strongly
involute, reflexed at maturity, densely ciliate, at times fissile
and fenestrated. Lamellae arcuate, short decurrent, not
anastomosing, pale buff to cinereous brown, fulvescent,
often tinted dark violaceous brown, narrow, 1–1.5 mm wide,
moderately to densely crowded, with lamellulae of 3–4
lengths; edge entire. Stipe central, eccentric or lateral, slender, 2–25 cm 9 2–10 mm, cylindric, expanding slightly at
both apex and base, solid; surface concolorous with the
pileus or more often darker, uniformly and persistently
velutinate with the indumentums extending into the bases of
lamellae, generally arising from a pseudosclerotium. Pseudosclerotium 2–10 9 1–4 cm, rarely larger, irregularly
fusoid, comprising sclerified wood impregnated with
hyphae, surface pale greyish-brown, smooth. Context thin,
up to 1 mm thick, white, drying rigid, consisting of a dimitic
hyphal system with generative hyphae.
Generative hyphae 2–4 lm diam., narrow, not inflated,
hyaline, thin-walled, rather sparingly branched, with clamp
connections. Skeletal hyphae, 2–4 lm diam., cylindric,
hyaline, with a thickened wall of 1–2 lm and a narrow,
continuous lumen, unbranched, arising in either an intercalary or terminal position, and then with an obtusely
rounded apex. Basidiospore print pale buff to cream colour. Basidiospores 5–7 9 3–4 lm, Q = 1.87, short,
oblong-cylindric, hyaline, thin-walled, with few contents.
Basidia 18–22 9 4–5 lm, narrowly cylindric, bearing 4
sterigmata. Lamella-edge sterile, with numerous basidioles
and scattered sclerocystidia. Basidioles 16–30 9 4–8 lm,
sinuous clavate or fusoid, constricted, often nodulose,
hyaline, thin-walled. Sclerocystidia present on lamellaedge and more especially on sides of lamellae, fairly
numerous to very abundant, 20–64 9 3–12 lm, initially
Fungal Diversity
Fig. 171 Lentinus sajor-caju (MFLU 12-1478) a Cheilocystidia. b Basidiospores. c Basidia. d Skeletal hyphae. e Generative hyphae. Scale bars
a, c = 20 lm, b, d, e = 10 lm
123
Fungal Diversity
Fig. 172 Basidiocarps of Lentinus squarrosulus in the field (MFLU 12-1228)
clavate and thin-walled with refractive gloeo-contents,
soon developing a thickened wall of 2–2.5 lm, hyaline or
brownish, not or scarcely projecting beyond the basidia.
Hymenophoral trama irregular, of radiate construction,
hyaline, similar in structure to the context. Subhymenial
layer little developed, 5–12 lm wide, interwoven.
Pileipellis a trichodermal palisade producing erect, loose
fascicles, 30–1000 lm long, of brown, thick walled, generative hyphae, 3–6 lm diam., with clamp connections, and
an obtusely rounded apex, arising from a gelatinized
hypodermium of radially repent generative hyphae. Stipitipellis similar to the pileipellis. No special smell; edible
when it is young.
Habit, habitat and distribution: solitary, usually on dead
or decaying wood, May–August. Our specimen was collected on a decaying wood near Peradeniya Royal Botanic
Gardens.
Specimens examined: SRI LANKA, Kandy District, near
Peradeniya Royal Botanic Gardens, 11 June 2012,
Samantha C. Karunarathna (MFLU 12-1123).
GenBank Number ITS:KY649465.
Notes: The wide distribution of Lentinus velutinus complex has resulted in a range of variations and in some
instances constant morphological differences limited to
restricted geographical areas. This is the first report of
123
L. velutinus with the molecular phylogenetic confirmation
from Sri Lanka after Pegler (1983) has previously reported it.
Panus Fr.
The generic delineation and phylogenetic relationships
between Lentinus and Panus and their accepted allies Neolentinus, Heliocybe, and Pleurotus has been controversial
(Corner 1981; Pegler 1983; Redhead and Ginns 1985;
Singer 1975, 1986). Corner (1981) treated Lentinus and
Panus as separate genera on the basis of their hyphal
system and some species were placed in Pleurotus by
Singer (1975), whereas, Pegler (1983) treated Panus as a
subgenus of Lentinus and included several species of
Singer’s Pleurotus in Panus (Fig. 176).
Panus subfasciatus Thongbai, Karun., C. Richter & K.D.
Hyde
Index Fungorum number: IF 552827; Facesoffungi
number: FoF 02915, Fig. 177
Etymology: subfasciatus refers to the similarity to P.
fasciatus
Holotype: MFLU 16-2129
Basidiomata (Fig. 177a) small-sized, often short and
stocky, 1.2–2.5 9 0.5–1.0 cm in diameter, cylindric,
brownish violet to purple brown (11D5–6) when young,
Fungal Diversity
Fig. 173 Lentinus
squarrosulus (MFLU 12-1228)
a: Basidia b: Basidiospores c:
Skeletal hyphae d: Generative
hyphae Scale bars a = 20 lm,
b = 10 lm, c, d = 5 lm
greyish ruby to ruby (12D7–8) when old, leathery, solid,
dry; surface concolorous with the pileus, velutinate, fully
covered with short and soft hairs, brownish grey (11C2);
context white to pale purplish, consisting a dimitic hyphal
system with skeletal hyphae. Pileus 2.5–5.5 cm in diameter, thin, slightly umbilicate centre, coriaceous,
infundibuliform; surface brownish grey to violet brown
(10E3–4), often purplish fuscous especially towards the
margin, velutinate with numerous hispid squamules forming erect hairs or hispid-strigose; margin at first involute,
then hispid. Lamellae 0.3 mm broad, with 3–5 tiers of
lamellulae, deeply decurrent, purplish black, moderately
distant to moderately crowded.
Basidiospores (Fig. 177b–e) (5.03–)5.28–6.44(–6.56) 9
(3.04–)3.33–4.16(–4.44) lm (n = 40) Lm = 5.73 lm,
Wm = 3.61 lm, Q = (1.34–)1.43–1.92(–1.95), Qm = 1.60
lm, ellipsoid to elongate, hyaline, thin walled, with few
contents. Basidia (Fig. 177f–i) 17–21 9 7–10 lm, clavate
to elongate clavate, 4-spored or occasionally 2-spored, with
sterigmata up to 6 lm long; clamps present at base of
basidia. Lamella edge sterile with abundant cheilocystidia.
Cheilocystidia (Fig. 177l) 24–28 9 15–25 lm, clavate,
hyaline, thin-walled. Sclerocystidia abundant on the sides
of the lamellae, 32–40(–55) 9 6–9(–12) lm, initially clavate, thin- walled. Pileipellis an epicutis of impact, interwoven, slightly thick-walled; generative hyphae 2.4–5 lm
wide, branching, with clamp connections. Generative
hyphae (Fig. 177j) 2.5–3 lm, inflated, undifferentiated
hyphae, hyaline, thin-walled, branching, with clamp connections. Skeletal hyphae (Fig. 177k) 2.5–3.5 lm, cylindrical, hyaline, slightly thick-walled.
Habitat: On dead and rotting wood in forest of
Fagaceae.
Material examined: THAILAND, Chiang Rai Province,
along the road number 1149, Mae Fah Luang Sub-District,
Mae Fah Luang District, 17 June 2014, collector Benjarong
Thongbai (MFLU 16-2129, holotype).
GenBank Number ITS:LT614958.
Notes: Panus subfasciatus is characterized by its
brownish violet to purple brown young basidomata. Later
they turn greyish ruby, leathery, solid, dry and are fully
covered with short brownish grey hairs. In some aspects it
123
Fungal Diversity
undescribed Panus species (BS = 82%). The phylogenetic
tree suggests that the nrITS region may be useful for preliminary phylogenetic analyses, also because this is the
most frequent DNA locus from which references are
available. However, an expanded taxon sampling, best
supported by a multi gene genealogy, should be carried out
to redefine the boundaries between Lentinus and Panus.
The new fungus was recently referred to as Lentinus cf.
fasciatus in a report of a new secondary metabolite named
lentinulactam (Helaly et al. 2016). Nevertheless, after the
outcome of the phylogenetic study we preferred to place it
in the genus Panus because it is appears closer to the type
species of that genus, P. conchatus.
Polyporus P. Micheli ex Adans.
Polyporus is a well-known genus of wood rotting fungi,
which produce stipitate basidiocarps with dimitic hyphal
system that causes white rot (Ryvarden 1991; Ryvarden
and Gilvertson 1994; Núñez and Ryvarden 1995; Sotome
et al. 2007). Núñez and Ryvarden (1995) divided this genus
into six infrageneric groups based on macro-morphological
characters and recent phylogenetic analysis showed that
Polyporus sensu lato is separated into at least five welldefined clades as follows: Dendropolyporus, Favolus, Neofavolus, Polyporellus, and Polyporus (Sotome et al. 2008;
Krüger et al. 2006; Sotome et al. 2013; Binder et al. 2013;
Seelan et al. 2015). Morphological characteristics and
phylogenetic analysis of six new species from Korea are
described below. The phylogenetic tree is presented in
Fig. 178.
Fig. 174 Basidiocarps of Lentinus velutinus in the field (MFLU
12-1123)
bears a superficial resemblance to Panus fasciatus (Berk.)
Singer and Lentinus concentricus Karunarathna, K.D. Hyde
& Zhu L. Yang. Panus subfasciatus is easily distinguished
from P. fasciatus with its ochraceous basidiomata and
abundance of sclerocystidia, which are clavate or fusoid,
22–32 9 4–9 lm (Corner 1981; Pegler 1983). Panus
subfasciatus differs from L. concentricus by the latter
having yellowish brown-clay, velvety, concentrically
zonate of basidiomata and absence of sclerocystidia (Karunarathna et al. 2011). Panus subfasciatus also bears a
superficial resemblance to Panus conchatus (Bull.) Fr.
which also has purplish brown basidiomata but is lacking
cilia. The phylogenetic trees obtained from RAxML analyses are shown in Fig. 176. Panus subfasciatus was statistically supported in the Polyporaceae. In the major clade,
P. subfasciatus clustered in a subclade that predominantly
contained Panus species and Lentinus strigosus, while a
second subclade exclusively contained Lentinus species.
Panus subfasciatus appeared most similar to P. conchatus
(unpublished according to the entry in GenBank) and a yet
123
Polyporus brevibasidiosus H. Lee, N.K. Kim & Y.W. Lim,
sp. nov.
Index Fungorum number: IF552513; Facesoffungi
number: FoF 2631, Fig. 179
Etymology: referring to the characteristic short basidia.
Holotype: SFC20130808-24
Basidiocarps annual, central stipitate; Pileus circular,
plane then funnel-shaped, up to 30 mm diam., 1.5 mm
thick; pileal surface light buff in the center, brown near
margin, fibrillose with flattened, radially striate, glabrous,
margin reddish-black, undulate, enrolled. Hymonophore
white, cream to brown; pores fine, rounded to angular, 8–9
per mm. Context in pileus pale buff, corky, up to 1 mm
thick. Tube continuous with the context, up to 0.5 mm
thick. Stipe central, brown to dark brown, smooth to longitudinally wrinkled, apex distinct from the pore, up to 40
mm long and 2 mm thick. Hyphal system dimitic; generative hyphae hyaline in KOH, thin-walled, 2.5–3 lm in
diam., septa with clamps, skeletal-binding hyphae thickwalled, aseptate, much branched, with tapering apices, 3–6
lm diam. Cystidia absent. Basidia clavate, 4–sterigmate
(up to 2.3 lm), 15–16.6 9 6.3–7.4 lm, with a basal clamp.
Fungal Diversity
Fig. 175 Lentinus velutinus
(MFLU 12-1123) a Basidia.
b Basidiospores.
c Cheilocystidia.
d Sclerocystidia. e Generative
hyphae and Skeletal hyphae.
f Hairs on pileus Scale bars a, c,
d = 20 lm, b, f, e = 10 lm
Basidiospores cylindric, hyaline, smooth, IKI–, 6.3–7.4 9
2.8–3.3 lm. L = 6.77 lm, W = 2.86 lm, Q = 2.37
(n = 20/1).
Habitat: solitary to gregarious on dead wood of
hardwood.
Material examined: KOREA, Gyeongsangbuk-do,
Sangju-si, Mt. Noem, on fallen hardwood branch, 8 August
2013, H. Lee, W. J. Kim & S-Y. Oh (SFC20130808-24,
holotype). KOREA, Gangwon-do, Inje-gun, Baekdam
temple, on fallen hardwood branch, 9 July 1999, Y. W. Lim
(SFC19990709-26, paratype).
GenBank Number ITS:KY038474.
Notes: This species shares similar characteristics with
many Polyporus species including eccentric stipe, thin
pileus, and small pores. Although the smaller pileus (less
than 3 cm), pores (8–9 mm) and basidia (15–16.6 9
123
Fungal Diversity
Fig. 176 Phylogenetic tree
generated with RAxML (GRTGAMMA) based on ITS
sequence data aligned with
MAFFT. Bootstrap values
higher than 70% are shown
above or beneath the branches at
nodes. The tree is rooted with
Polyporus conifericola
KU189783, Polyporus
tubiformis AB587634 and
Polyporus melanopus
AF516569. Panus subfasciatus
LT614958 (Ex-type) is
highlighted in blue. GenBank
accession numbers are provided
after each species name
6.3–7.4 lm) of P. brevibasidiosus are distinguishable
characters, morphology alone is not sufficient for species
level identification. ITS sequence analysis is useful to
confirm its identity and phylogenetic position.
Polyporus koreanus H. Lee, N.K. Kim & Y.W. Lim, sp.
nov.
Index Fungorum number: IF552509; Facesoffungi
number: FoF00556; Fig. 180
Etymology: referring to the first reported species of this
genus found in Korea.
Holotype: SFC20150813-58
Basidiocarps annual, central stipitate; Pileus circular,
plane and somewhat depressed, funnel-shaped in the center, up to 70 mm in diam., 2 mm thick; pileal surface light
grey to pale buff, with age becoming brown and dark
brown, fibrillose with flattened, radially striate, slightly
glabrous; margin ochreous, sharp and undulate. Hymonophore white, cream, buff to brown; pores fine, rounded to angular, 6–7 per mm. Context in pileus pale buff,
corky, up to 1.5 mm thick. Tube slightly darker than context, up to 0.5 mm thick. Stipe central, brown, dark reddishbrown to black, smooth to longitudinally wrinkled, apex
distinct from the pore, upper portion of blackish stipe
123
usually covered by pores, up to 50 mm long and 9 mm
thick. Hyphal system dimitic; generative hyphae hyaline in
KOH, thin-walled, 2.8–3.5 lm in diam., septa with clamps,
skeletal-binding hyphae thick-walled, aseptate, much
branched, with tapering apices, 3–6 lm diam. Cystidia
absent. Basidia clavate, 4–sterigmate (up to 3.4 lm),
21–25.7 9 5.6–8 lm, with a basal clamp. Basidiospores
cylindric, hyaline, smooth, IKI–, 5.7–6.9 9 2.1–2.3 lm.
L = 6.34 lm, W = 2.24 lm, Q = 2.83 (n = 20/1).
Habitat: solitary on dead wood of hardwood or on the
ground.
Material examined: KOREA, Gyeonggi-do, Goyang-si,
West Five Royal Tombs, on fallen hardwood branch, 13
August 2015, N.G. Kim, J. E. Eom & M. J. So
(SFC20150813-58, holotype); KOREA, Gyeongsangbukdo, Ulleung-gun, Na-ri basin, on fallen hardwood branch,
15 August 1995, Y.W. Lim (SFC19950815-35, paratype); KOREA, Gyeongsangnam-do, Hamyang-gun, Mt.
Jiri, on rotten stump of Fraxinus mandshurica, 25
August 2004, J. S. Lee & K. M. Kim (SFC20040825187, paratype); KOREA, Gyeonggi-do, Uiwang-si, Mt.
Cheonggye, on fallen hardwood branch, 20 August
2015, N. G. Kim & H. J. Cho (SFC20150820-61,
paratype).
Fungal Diversity
Fig. 177 Panus subfasciatus (MFLU 16-2129, holotype). a Basidiomata. b–e Basidiospores. f, g Basidia. h Cheilocystidia. i, l Sclerocystidia. j,
k Pileipellis generative hypha. Scale bars b–e, h–i, l = 3 lm. f, g = 10 lm. j, k = 5 lm
GenBank Numbers SFC20150813-58 ITS:KY038479.
SFC19950815-35 ITS:KY038461. SFC20040825-187
ITS:KY038462. SFC20150820-61 ITS:KY038480.
Notes: This species is often confused with P. melanopus,
P. ulleungus and P. varius because of similar basidiocarps
and microscopic features; however, the smaller basidiospore of Polyporus koreanus (5.7–6.9 9 2.1–2.3 lm) is
distinguishable from P. melanopus (7–9 9 3–3.5 lm),
P. ulleungus (6.8–8 9 2.3–3.3 lm) and P. varius (9–12 9
2.5–3 lm) (Gilbertson and Ryvarden 1987).
Polyporus orientivarius H. Lee, N.K. Kim & Y.W. Lim,
sp. nov.
Index Fungorum number: IF552514; Facesoffungi
number: FoF 2632, Fig. 181
Etymology: referring to P. varius, but with Far East
Asian distribution.
Holotype: SFC20130704-20
Basidiocarps annual, central to laterally stipitate; Pileus
circular to dimidiate, plane then funnel-shaped, up to 70
mm diam., 2.5 mm thick; pileal surface cream to pale buff,
123
Fungal Diversity
123
Fungal Diversity
b Fig. 178 Phylogram generated from maximum likelihood (RAxML)
analysis based on ITS sequence data of Polyporus. Maximum
likelihood bootstrap support values greater than 70% are indicated
above or below the nodes and the new species is in bold. The tree is
rooted with two Neofavolus species
with age becoming brown and dark brown, fibrillose with
flattened, radially striate, slightly glabrous, margin ochreous, sharp and undulate. Hymenophore white, cream, buff
to brown; pores fine, rounded to angular, 6–7 per mm.
Context in pileus pale buff, corky, up to 1 mm thick. Tube
continuous with the context, up to 1.5 mm thick. Stipe
central, brown to dark brownish-black, longitudinally
wrinkled, apex distinct from the pore, up to 20 mm long
and 9 mm thick. Hyphal system dimitic; generative hyphae
hyaline in KOH, thin-walled, 2.8–3.5 lm in diam., septa
with clamps, skeletal-binding hyphae thick-walled, aseptate, much branched, with tapering apices, 3–6 lm diam.
Cystidia absent. Basidia clavate, 4-sterigmate, 21–25 9
5.7–8 lm, with a basal clamp. Basidiospores cylindric,
hyaline, smooth, IKI-, 5.1–6.8 9 2.3–2.5 lm. L = 5.7 lm,
W = 2.4 lm, Q = 2.4 (n = 20/1).
Habitat: solitary to gregarious on dead wood of
hardwood.
Material examined: KOREA, Gyeongsangbuk-do,
Yecheon-gun, Mt. Hagga, on fallen hardwood branch, 4
July 2013, Y.W. Lim (SFC20130704-20, holotype); ibid.
Fig. 179 Polyporus brevibasidiosus (SFC20130808-24, holotype). a,
b Fruiting bodies (SFC20130808-24). c fruiting bodies in the field
(SFC20130808-24). d Pores of fruiting body (SFC20130808-24).
(SFC20130704-06, paratype); KOREA, Gyeongsangbukdo, Sangju-si, Seongjubong Natural Recreation Forest, on
fallen hardwood branch, 9 August 2013, H. Lee, W. J. Kim
& S-Y. Oh (SFC20130809-44); KOREA, Jeju-do, Seogwipo-si, Hannam Experimental Forest, on rotten hardwood
stump, 7 August 2012, Y. W. Lim, H. Lee, W. J. Kim & J.
S. Gyeong (SFC20120807-12); KOREA, Chungcheongnam-do, Boryeong-si, Seongjusan Natural Recreation
Forest, on fallen hardwood branch, 21 August 2012, Y.
W. Lim, H. Lee, W. J. Kim & Y. J. Min (SFC2012082136); ibid. (SFC20120821-59); KOREA, Jeollabuk-do,
Namwon-si, Mt. Jiri, on fallen hardwood branch, 20 July
2005, H. S. Jung (SFC20050720-60); ibid., 22 July 2004, J.
S. Lee & S. H. Hong (SFC20040722-26, paratype).
GenBank Numbers SFC20130704-20 ITS:KY038470.
SFC20130704-06
ITS:KY038469.
SFC20130809-44
ITS:KY038475.
SFC20120807-12
ITS:KY038465).
SFC20120821-36 ITS: KY038468. SFC20050720–60
ITS:KY038463.
Notes: This species has variable basidiocarp shapes
similar to Polyporus varius. Therefore, many of specimens
in Korea had been misidentified as Polyporus varius. Two
types of P. varius sequences were reported in a previous
study (Sotome et al. 2008). The first included sequences
from the USA, Russia, and Japan (Sotome et al. 2011; Dai
et al. 2014) and the second was unique to Japan
(AB587635, Sotome et al. 2011). Phylogenetic analysis
e Microscopic structure; basidiospores, basidia, generative hyphae
and skeletal-binding hyphae
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Fungal Diversity
Fig. 180 Polyporus koreanus (SFC20150813-58, holotype). a, b
Fruiting bodies in the field (SFC20150820-61). c Fruiting bodies in
the field (SFC20150813-58). d Pores of fruiting body (SFC20150813-
58). e Microscopic structure; basidiospores, basidia, generative
hyphae and skeletal-binding hyphae
based on ITS sequences showed that P. orientivarius
formed a monophyletic group with the second type of
P. varius sequence and was distantly related to P. varius
(Fig. 3). In addition, P. varius has smaller pores (7–9 mm)
and longer basidiospores (9–12 9 2.5–3 lm) (Gilbertson
and Ryvarden 1987) than those of P. orientivarius.
Therefore, both morphological and molecular evidence
support P. orientivarius as a new species that is clearly
separate from P. varius.
usually covered by decurrent pores, up to 40 mm long and
2 mm thick. Hyphal system dimitic; generative hyphae
hyaline in KOH, thin-walled, 2.5–3 lm in diam., septa with
clamps, skeletal-binding hyphae thick-walled, aseptate,
much branched, with tapering apices, 3–6 lm diam. Cystidia absent. Basidia clavate, 4-sterigmate (up to 3.4 lm),
23.8–26.3 9 7.4–8.5 lm, with a basal clamp. Basidiospores cylindric, hyaline, smooth, IKI-, 7.4–8.6 9
3.5–4.5 lm. L = 7.52 lm, W = 4.25 lm, Q = 1.78
(n = 20/1).
Habitat: solitary to gregarious on dead wood of
hardwood.
Material examined: KOREA, Gyeongsangbuk-do,
Ulleung-gun, Na-ri basin, on fallen hardwood branch, 14
August 2012, Y. W. Lim, W. J. Kim & S-Y. Oh
(SFC20120814-33, holotype); KOREA, Gangwon-do,
Inje-gun, on rotten wood below litter layer, 1 July 2014, Y.
W. Lim & H. J. Cho (SFC2040701-19, paratype).
GenBank Numbers ITS:KY038466.
Notes: Polyporus parvovarius is similar to P. badius,
P. elegans and P. varius in its central stipitate basidiocarps,
blackish stipe, and microscopic features. Simple septate
generative hyphae separate P. badius from the other species. P. parvovarius has larger basidia (23.8–26.3 9
7.4–8.5 lm) than P. elegans (15–20 9 6–7 lm) and
P. varius (18–23 9 5–7 lm). Phylogenetic analysis of the
ITS region showed that P. parvovarius is closely related to
Polyporus parvovarius H. Lee, N.K. Kim & Y.W. Lim, sp.
nov.
Index Fungorum number: IF552511; Facesoffungi
number: FoF 2629, Fig. 182
Etymology: referring to the similar but smaller basidiocarps compared to P. varius.
Holotype: SFC20120814-33
Basidiocarps annual, central stipitate; Pileus circular,
plane then funnel-shaped, up to 35 mm diam., 1.5 mm
thick; pileal surface light buff to brown, fibrillose with
flattened, radially striate, glabrous; margin reddish-brown,
enrolled, undulate. Hymonophore white, cream to brown;
pores fine, rounded to angular, 6–7 per mm. Context in
pileus pale buff, corky, up to 1 mm thick. Tube continuous
with the context, up to 0.5 mm thick. Stipe central, brown
to dark brown, smooth to longitudinally wrinkled, apex
distinct from the pore, upper portion of blackish stipe
123
Fungal Diversity
Fig. 181 Polyporus orientivarius (SFC20130704-20, holotype).
a Fruiting bodies in the field (SFC20120807-12). b Fruiting bodies
in the field (SFC20130704-20). c Pores of fruiting body
(SFC20130704-20). e Microscopic structure; basidiospores, basidia,
generative hyphae and skeletal-binding hyphae
Fig. 182 Polyporus parvovarius (SFC20120814-33, holotype).
a Fruiting bodies in the field (SFC20120814-33). b, c Fruiting bodies
in the field (SFC20140701-19). d Pores of fruiting body
(SFC20120814-33). e Microscopic structure; basidiospores, basidia,
generative hyphae and skeletal-binding hyphae
123
Fungal Diversity
Fig. 183 Polyporus subdictyopus (SFC20130719-53, holotype).
a Fruiting bodies in the field (SFC20110818-02). b Fruiting bodies
in the field (SFC20130704-39). d Pores of fruiting body
(SFC20140723-08). e Microscopic structure; basidiospores, basidia,
generative hyphae and skeletal-binding hyphae
P. brevibasidiosus (Fig. 178). However, they are distinguishable by pore, basidia, and basidiospore size. P. parvovarius has larger than those of P. brevibasidiosus.
(up to 7.9 lm), 21–23 9 5.1–6.9 lm, with a basal clamp.
Basidiospores cylindric, hyaline, clear one or two oil drops,
smooth, IKI–, 6.8–9.1 9 2.2–2.8 lm. L = 7.39 lm,
W = 2.34 lm, Q = 3.16 (n = 20/2).
Habitat: solitary to gregarious or frequently concrescent
on dead wood of hardwood, occasionally on conifer
(SFC20110818–02).
Material examined: KOREA, Gyeongsangbuk-do,
Sangju-si, Mt. Noem, on fallen hardwood branch, 19 July
2013, H. Lee, W. J. Kim & Y. J. Min (SFC20130719-53,
holotype); KOREA, Gangwon-do, Wonju-si, Mt. Chiak,
on dead conifer, 18 August 2011, Y.W. Lim
(SFC20110818-02, paratype); KOREA, Gyeongsangbukdo, Yecheon-gun, Mt. Hagga, on fallen hardwood branch, 4
July 2013, Y.W. Lim (SFC20130704-39); KOREA, Jeollabuk-do, Jinan-gun, Unjangsan Natural Recreation Forest,
on fallen hardwood branch, 23 July 2014, J. Y. Park, H.
Lee & H. J. Cho (SFC20140723-08); KOREA, Jeju-do,
Jeju-si, Bijarim, on fallen hardwood branch, 30 June 2015,
Y. W. Lim, N. G. Kim & H. Lee (SFC20150630-48);
KOREA, Chungcheongbuk-do, Boeun-gun, Mt. Songni, on
dead trunk of oak, 25 July 2015, H. Lee (SFC2015072505).
GenBank Numbers SFC20130719-53 ITS:KY038473.
SFC20110818-02
ITS:KY038464.
SFC20130704-39
ITS:KY038471.
SFC20140723-08
ITS:KY038476.
SFC20150630-48
ITS:KY038477.
SFC20150725-05
ITS:KY038478).
Polyporus subdictyopus H. Lee, N.K. Kim & Y.W. Lim,
sp. nov.
Index Fungorum number: IF552512; Facesoffungi
number: FoF 2630, Fig. 183
Etymology: referring to similar morphology and close
phylogenetic relationship with P. dictyopus.
Holotype: SFC20130719-53
Basidiocarps annual, central to laterally stipitate; Pileus
circular to dimidiate, plane then funnel-shaped, up to 60
mm diam., 2.5 mm thick; pileal surface whitish when
juvenile to pale buff, brown and dark brown with age,
fibrillose with flattened, radially striate, slightly glabrous,
margin ochreous, sharp and undulate. Hymonophore white,
cream, buff to brown; pores fine, rounded to angular, 8–10
per mm. Context in pileus pale buff, corky, up to 2 mm
thick. Tube continuous with the context, up to 0.5 mm
thick. Stipe central, brown to dark brownish-black, smooth
to longitudinally wrinkled, apex distinct from the pore,
pores traces observed at upper portion of blackish stipe, up
to 15 mm long and 6 mm thick. Hyphal system dimitic;
generative hyphae hyaline in KOH, thin-walled, 1.7–3 lm
in diam., septa with clamps, skeletal-binding hyphae thickwalled, aseptate, much branched, with tapering apices, 3–5
lm diam. Cystidia absent. Basidia clavate, 4–sterigmate
123
Fungal Diversity
Fig. 184 Polyporus ulleungus (SFC20120814-41, holotype). a, b
Fruiting bodies in the field (SFC20120814-41). c Fruiting bodies in
the field (SFC20150902-99). d Pores of fruiting body (SFC20120814-
41). e Microscopic structure; basidiospores, basidia, generative
hyphae and skeletal-binding hyphae
Notes: Basidiocarps of this species are most similar to
those of P. badius. P. badius is easily distinguished by its
lack of clamp connection. In a phylogenetic analysis,
P. badius formed a separate clade within the Melanopus
clade, but P. subdictyopus is located in a sister clade
together with P. dictyopus (Fig. 178). Basidiocarps shape
and microscopic features of P. dictyopus are hard to distinguish except its pantropical distribution. One or two oil
drops in the basidiospores of P. subdictyopus differentiate
it from P. dictyopus.
usually covered by pores, up to 30 mm long and 8 mm
thick. Hyphal system dimitic; generative hyphae hyaline in
KOH, thin-walled, 1.7–3 lm in diam., septa with clamps,
skeletal-binding hyphae thick-walled, aseptate, much
branched, with tapering apices, up to 6 lm diam. Cystidia
absent. Basidia clavate, 4–sterigmate (up to 3.4 lm),
23.4–25.7 9 5.8–6.9 lm, with a basal clamp. Basidiospores cylindric, hyaline, smooth, IKI-, 6.8–8 9 2.3–3.3
lm. L = 7.06 lm, W = 2.85 lm, Q = 2.50 (n = 20/2).
Habitat: solitary to gregarious on dead wood of
hardwood.
Material examined: KOREA, Gyeongsangbuk-do,
Ulleung-gun, Na-ri basin, on fallen hardwood branch, 14
August 2012, Y.W. Lim, W. J. Kim & S-Y. Oh
(SFC20120814-41, holotype); KOREA, Gangwon-do,
Inje-gun, on fallen branch of Betula platyphylla var.
japonica, 2 September 2015, Y. W. Lim & H. Lee
(SFC20150902-99, paratype).
GenBank Numbers SFC20120814-41 ITS:KY038467.
SFC20150902-99 ITS:KY038481.
Notes: Polyporus ulleungus is commonly found in the
Na-ri Basin of Ulleung Island and has larger basidiocarps
(SFC20120814-41; 110 mm across) than other species
collected in inland area (SFC20150902-99; 35 mm across).
Polyporus ulleungus is most similar to P. koreanus and
P. melanpus. It has larger pore (5–6 mm) than those of
P. koreanus (6–7 mm) and P. melanpus (6–8 mm). In a
phylogenetic analysis, P. ulleungus formed a distinct clade
Polyporus ulleungus H. Lee, N.K. Kim & Y.W. Lim, sp.
nov.
Index Fungorum number: IF552510; Facesoffungi
number: FoF 2628, Fig. 184
Etymology: very common on Ulleung Island.
Holotype: SFC20120814-41
Basidiocarps annual, central stipitate; Pileus circular,
plane then infundibuliform, up to 110 mm wide, 3 mm
thick; pileal surface light brown to dark blackish-brown in
the center, fibrillose with flattened, radially striate, slightly
glabrous, margin ochreous, sharp and undulate. Hymonophore white to brown; pores fine, rounded to angular,
5–6 per mm. Context in pileus pale buff, corky, up to 2 mm
thick. Tube white and cream when young, ochreous with
age or dry, up to 1 mm thick. Stipe central, brown to dark
brownish-black, smooth to longitudinally wrinkled, apex
distinct from the pore, upper portion of blackish stipe
123
Fungal Diversity
Fig. 185 Maximum likelihood phylogenetic tree of the concatenated
ITS1, 5.8S, ITS2 and partial 28S rDNA sequences. Bootstrap
supporting values (1000 replicates) and posterior probabilities (PP)
from Bayesian analysis to each node are shown from left to right. The
new species Wolfiporia pseudococos described in this study is
highlighted in blue. The tree was rooted with Grifola sordulenta and
Laetiporus sulphureus
with strong support (100% ML, Fig. 179) and clearly
separated from P. koreanus and P. melanpus (Fig. 178).
cinnamon-buff when dry, not glancing; pores round, 1–3
per mm; dissepiments thick, even to slightly lacerate.
Subiculum cinnamon-buff, corky, up to 1.5 mm; tubes
corky to fragile, buff, up to 4.5 mm long. Hyphal system
dimitic in all parts, generative hyphae with simple septa,
skeletal hyphae dominant, all hyphae IKI-, CB-, weakly
inflated in KOH. Subicular hyphal structure homogeneous,
hyphae strongly interwoven; generative hyphae occasionally present, hyaline, thin-walled, occasionally branched,
frequently simple septate, 4–6 lm in diam.; skeletal hyphae
dominant, hyaline, thick-walled with a distinct wide lumen,
usually winding, frequently branched, occasionally simple
septate, 5–7 lm in diam. Tramal generative hyphae frequent, hyaline, thin-walled, occasionally branched, frequently simple septate, 3–5 lm in diam.; tramal skeletal
hyphae frequent, hyaline, thick-walled with a wide lumen,
winding, occasionally branched and simple septate, 4–6
lm in diam. Cystidia absent, cystidioles present; Basidia
clavate, with four sterigmata and a basal simple septum,
18–25 9 10–14 lm, basidioles in shape similar to basidia
but slightly smaller. Basidiospores ellipsoid to broadly
ellipsoid, tapering at apiculus, hyaline, thin-walled, IKI-,
Wolfiporia Ryvarden & Gilb.
Wolfiporia is a genus of mushrooms in the family
Polyporaceae. The genus was introduced by Ryvarden and
Gilbertson (1984) to contain the type species Wolfiporia
cocos (currently known as W. extensa) and W. dilatohypha
(Fig. 185).
Wolfiporia pseudococos F. Wu, J. Song & Y.C. Dai, sp.
nov.
MycoBank number: MB 818729; Facesoffungi number:
FoF 2961, Figs. 186, 187
Etymology: referring to the similarity of Wolfiporia
cocos.
Holotype: BJFC01938
Basidiocarp annual, resupinate, soft corky and without
odour or taste when fresh, corky to fragile when dry, up to
10 cm long, 4 cm wide, 6 mm thick at centre. Margin thin,
usually pores extend to the very edge. Pore surface cream
to ash-grey when fresh, becoming pinkish-buff to
123
Fungal Diversity
Fig. 186 Fruiting body of Wolfiporia pseudococos (BJFC01938, holotype) (photo by Y.C. Dai)
CB-, (7.6–)7.9–9.5(–9.7) 9 (2.9–)3–3.8 lm, L = 8.55 lm,
W = 3.17 lm, Q = 2.7 (n = 30/1).
Specimen examined: CHANA, Hainan Province, Ledong
County, Jianfengling Nature Reserve, On dead angiosperm
tree, 1 June 2015, Dai 15269 (BJFC019380, holotype).
GenBank Numbers ITS:KX354451.
Notes: The phylogenetic analysis of ITS sequence
(Fig. 186) shows that the new species, Wolfiporia pseudococos, is closely related to W. cocos (F.A. Wolf)
Ryvarden & Gilb, the type species of the genus Wolfiporia.
Sequence data from three specimens of W. Cocos also
clustered in a distinct lineage with strong support (100%
BS, 1 BYPP), and the sequence of the new species formed
a terminal lineage. Morphologically, Wolfiporia pseudococos is distinguished from W. cocos by the presence of
cystidioles and larger basidiospores (7.9–9.5 9 3–3.8 lm
vs. 6.5–8.1 9 2.8–3.1 lm). In addition, Wolfiporia cocos
usually grows on conifers in temperate areas, but Wolfiporia pseudococos has a distribution in tropics and growing
on angiosperm trees.
Psathyrellaceae Locq.
The family Psathyrellaceae is a family of dark-spored
agarics that generally have rather soft, fragile fruiting
bodies, and are characterized by black, dark brown, rarely
reddish, or even pastel-coloured spore prints.
Coprinopsis P. Karst., Acta Soc. Fauna Flora Fenn. 2 (1):
27 (1881)
During the last 25 years, molecular phylogenetic analyses have made great impact on our understanding of
phylogenetic relatonships among coprinoid fungi (Hopple
and Vilgalys 1994, 1999; Moncalvo et al. 2002). The
analyses have showed that the genus Coprinus Pers. sensu
lato is not monophyletic. A minority of Coprinus species
(including the type species C. comatus (O.F. Müll.) Pers.)
is clustered near the genus Agaricus L., while the great
majority of Coprinus species is grouped into three clades
closely related to the genus Psathyrella (Fr.) Quél. Based
on molecular studies, Redhead et al. (2001) transferred the
majority of Coprinus species into three genera (Coprinellus
P. Karst., Coprinopsis, and Parasola Redhead, Vilgalys &
Hopple) within the newly proposed family Psathyrellaceae
Vilgalys, Moncalvo & Redhead. Consequently, the genus
Coprinopsis includes five subsections of Coprinus sensu
lato, Alachuani Singer, Atramentarii (Fr.) Konrad &
Maubl., Lanatuli J.E. Lange, Narcotici Uljé & Noordel.,
and Nivei Citérin. Further phylogenetic research (Larsson
123
Fungal Diversity
Fig. 187 Microscopic
structures of Wolfiporia
pseudococos (BJFC01938,
holotype). a Basidiospores.
b Basidia and basidioles.
c Cystidioles. d Hyphae from
trama. e Hyphae from
subiculum (drawings by F. Wu)
and Örstadius 2008; Padamsee et al. 2008; Örstadius et al.
2015) revealed that the genus Psathyrella is not monophyletic either. Based on these studies, some Psathyrella
species belong to the genus Coprinopsis (Fig. 188).
Coprinopsis cerkezii Tkalčec, Mešić, I. Kušan & Matočec,
sp. nov.
MycoBank number: MB 818295; Facesoffungi number:
FoF 2962, Figs. 189, 190
Etymology: Described in honour of Mr. Milan Čerkez
for his great contribution to the study of coprinoid and
coprophilous fungi in Croatia.
Holotype: CNF 1/7253
123
Pileus 3–11 mm broad when expanded, spherical at first,
then ellipsoid, paraboloid, convex, at maturity plano-convex with depressed centre, applanate or plano-concave,
strongly plicate-sulcate up to 85% of the radius at maturity,
edge strongly crenate, sometimes radially splitting with
age, not hygrophanous, non-deliquescent, surface dry, at
first entirely covered with upper velar layer composed of
fibrils mostly directed downwards (more developed in the
lower part of the pileus), tufts and warts in the central zone,
upper velar layer tearing soon (mostly remaining in the
form of velar patches, tufts and fibrils) and exposing
whitish to cream granulose lower velar layer, fibrils white
to light orange-brown, tufts and warts pale to medium
Fungal Diversity
Fig. 188 Maximum likelihood
phylogenetic tree inferred from
the dataset of ITS1–5.8S-ITS2
gene sequences from
Coprinopsis cerkezii sp. nov.
and related species. The new
species is shown in blue.
Maximum likelihood bootstrap
values greater than 50% are
indicated at the nodes. The tree
is rooted with Coprinellus
xanthothrix and C. domesticus.
The bar indicates the number of
nucleotide substitutions per site
orange-brown to rusty-orange, darker at centre, often paler
towards the margin. Lamellae free, distant, L = 18–32,
l = 0–1, up to 1.5 mm broad, sometimes forked, often
deliquescent at the edge, white at first, becoming rustybrown or dark (chocolate) brown at maturity, sometimes
whitish to greyish till maturity, with whitish and flocculose
edge (when not deliquescent). Stipe 10–50 9 0.5–1.2 mm,
often with swollen base (up to 2 mm), slightly tapering
towards the apex, central, hollow, dry, densely hairy at
first, later floccose to flocculose, white to cream when
young, then subhyaline, orange-brown to rusty-orange at
the base, velar hairs sometimes forming volva-like basal
structure. Context fragile, very thin in the pileus, whitish.
Odour and taste not observed.
Basidiospores [300/6/4] (4.8–)5.3–6.8–8.3(–9.1) 9
(2.7–)2.8–3.2–3.6(–3.8) lm, average = 6.2–7.6 9 3–3.4
lm, Q = (1.48–)1.66–2.12–2.59(–2.90), Qav = 1.98–2.47
(very variable among different basidiocarps), narrowly
ellipsoid, elongate ellipsoid, cylindrical or subovoid in
frontal view, ellipsoid, amygdaliform, phaseoliform or
allantoid in side view, smooth, moderately thick-walled
(ca. 0.5 lm), with distinct and central (sometimes slightly
eccentric), 1–1.5 lm wide germ-pore, rusty-brown in H2O
and NH4OH, medium brown in KOH, non-amyloid and
non-dextrinoid. Basidia 8–24 9 4–8 lm, narrowly to
broadly clavate, 4-spored, thin-walled, hyaline, surrounded
by 3–5 hymenophysalides (pseudoparaphyses). Hymenophysalides 10–21 9 7–18 lm, subglobose, (broadly)
ellipsoid, broadly clavate or broadly oblong, thin-walled,
hyaline, fully developed only in mature basidiocarps.
Lamellar edge sterile with crowded cheilocystidia.
Cheilocystidia 10–65 9 10–35 lm, very variable in size
and shape, globose, subglobose, ellipsoid, clavate, ovoid,
obovoid, conical, obtuse-conical, utriform or broadly
lageniform, thin- to moderately thick-walled (up to 0.7
lm), hyaline. Pleurocystidia absent. Pileipellis a cutis,
composed of repent, hyaline, thin-walled, 1.5–7 lm wide
hyphae. Veil on pileus consists of two layers; lower layer
completely covering pileipellis and predominantly composed of globose to broadly ellipsoid, subhyaline to pale
123
Fungal Diversity
Fig. 189 Coprinopsis cerkezii (CNF 1/7253, holotype). a–e Basidiocarps (a–c CNF 1/7253 holotype, d CNF 1/7252, e CNF 1/5209). Scale bars
a = 10 mm, b–e = 2 mm
yellow–brown, thin- to thick-walled (up to 1.2 lm), often
minutely encrusted, 15–55 lm wide cells, intermixed with
some hyphoid elements; upper layer tearing during the
development, composed of long, (sub) cylindrical, nondiverticulate, hyaline to pale yellow–brown, thin-walled,
mostly minutely to coarsely encrusted, 2–18 lm wide cells,
± intermixed with inflated (ellipsoid, fusoid, ovoid, clavate, subglobose or globose) cells, subhyaline to light
yellow–brown, thin- to thick-walled (up to 1.2 lm), often
minutely to coarsely encrusted, up to 50 lm wide. Stipitipellis a cutis of parallel, repent, thin-walled, hyaline, 2–12
lm wide hyphae. Veil on stipe composed of (sub) cylindrical and inflated (ellipsoid, fusoid, ovoid, clavate, subglobose or globose) cells, hyaline to light yellow-brown,
thin-walled, minutely encrusted, 2–35 lm wide. Clamp
connections absent.
Ecology: In groups on deer dung heaps; among mosses
in forest of Abies alba, Picea abies, and Fagus sylvatica
123
and among grasses and sedges in a thicket of Salix aurita,
Betula sp., and Alnus glutinosa.
Known distribution: Croatia, mountainous area of Primorje-Gorski Kotar County, found at two localities 20 km
apart.
Material examined: CROATIA, Primorje-Gorski Kotar
County, near Skrad, about 1 km SE of Rogi, 45250 1300 N,
14520 5900 E, 725 m a.s.l., in forest of Abies alba, Picea
abies, and Fagus sylvatica, on deer dung, 14 May 2010 in
culture (substrate coll. 27 March 2010), leg. M. Čerkez
(CNF 1/7253, holotype); ibid., 45250 1800 N, 14520 5100 E, 695
m a.s.l., 22 April 2010 in culture (substrate coll. 27 March
2010), leg. M. Čerkez (CNF 1/7252); CROATIA, Primorje-Gorski Kotar County, about 4 km W-NW of Fužine,
45190 1600 N, 14400 1500 E, 745 m a.s.l., among grasses and
sedges, thicket of Salix aurita, Betula sp., and Alnus
glutinosa, on deer dung, 13 May 2008 in culture (substrate
coll. 30 April 2008), leg. M. Čerkez (CNF 1/5209); ibid.,
Fungal Diversity
Fig. 190 Coprinopsis cerkezii (CNF 1/7253, holotype). a Basidiospores. b, c Cheilocystidia (phase contrast). d Upper velar layer on pileus.
e Lower velar layer on pileus. Scale bars a = 5 lm, b, c = 20 lm, d, e = 30 lm
22 May 2008 in culture (substrate coll. 30 April 2008), leg.
M. Čerkez (CNF 1/5216).
GenBank Number ITS:KX869912.
Notes: Based on a megablast search of NCBIs GenBank
numbers nucleotide database, the closest hits using the
ITS1-5.8S-ITS2 sequence for our holotype collection (CNF
1/7253, GenBank Numbers KX869912) are Coprinopsis
(Coprinus) cortinata (J.E. Lange) Gminder (GenBank
Numbers
JF907847;
Identities = 619/702
(88%),
Gaps = 29/702 (4%)), Coprinopsis (Coprinus) bellula
(Uljé) P. Roux & Eyssart. (GenBank Numbers FN430682;
Identities = 621/705 (88%), Gaps = 30/705 (4%), and
Coprinopsis utrifer (Watling) Redhead, Vilgalys & Moncalvo (GenBank Numbers FN396410; Identities = 609/
688 (89%), Gaps = 28/688 (4%)). Other three collections
of C. cerkezii (CNF 1/5209, 1/5216, 1/7252) have identical
ITS and LSU sequences. The phylogenetic analysis using
maximum likelihood method was performed with MEGA6
software. The ITS sequence from the holotype of Coprinopsis cerkezii was combined with sequences of all
closely related species and some other taxa of the genus
Coprinopsis downloaded from GenBank number database.
Analysed sequences belong to the species from subsections
Alachuani, Atramentarii, Lanatuli, Narcotici, and Nivei
(Uljé 2005) of the genus Coprinus sensu lato, as well as to
a few Coprinopsis species transferred from the genus
Psathyrella (Örstadius et al. 2015). Sequences of Coprinellus species (C. xanthothrix (Romagn.) Vilgalys,
Hopple & Jacq. Johnson and C. domesticus (Bolton) Vilgalys, Hopple & Jacq. Johnson) were used as outgroup. In
phylogenetic analysis Coprinopsis cerkezii clustered with
other members of subsect. Nivei of the genus Coprinus
sensu lato (Fig. 188), in accordance with its morphological
characters.
123
Fungal Diversity
Coprinopsis cerkezii is primarily characterised by small
and delicate basidiocarps, a strongly plicate-sulcate pileus
with an orange-brown to rusty-orange upper veil, narrow
and elongated spores without myxosporium, an abundance
of spherical pileal velar elements lacking nipple-shaped
warts, an absence of pleurocystidia and clamp-connections,
and by living on dung. Comparison of C. cerkezii with
similar taxa is based on descriptions in the following literature: Bisby et al. (1929), Orton and Watling (1979),
Pegler (1986), Doveri (2004), Uljé (2005). Five species
morphologically similar to C. cerkezii, C. bellula, C. candidata (Uljé) Gminder & T. Böhning, C. coniophora
(Romagn.) Redhead, Vilgalys & Moncalvo, C. cortinata,
and Coprinus furfurellus (Berk. & Broome) Pegler, differ
by living on soil or wood, by presence of clamp-connections, and by absence of orange-brown tones in pileus.
From above mentioned species, Coprinus furfurellus possess the most similar basidiospores (6–7 9 3.5–4 lm), but
the average Q is smaller (1.73). Three other similar species
live on dung as Coprinopsis cerkezii. Coprinus parvisporus
Buller differs by having white pileus, less elongated spores
(6 9 3.5 lm), presence of pleurocystidia, and larger
spherical elements of pileal veil (60–80 lm). Coprinopsis
pseudcortinata (Cacialli, Caroti & Doveri) Doveri, Granito
& Lunghini differs by having white to greyish pileus,
broader (3.5–4.5 lm) and less elongated spores
(Q = 1.5–2.05), presence of pleurocystidia and smaller
number of lamellae (L = 6–12). Coprinopsis utrifer differs
by having broader (4–5.5 lm) and less elongated spores
(Q = 1.3–1.7) and by presence of clamp-connections and
pleurocystidia. ITS Sequences of Coprinopsis candidata,
C. pseudcortinata, Coprinus furfurellus, and C. parvisporus are not included in phylogenetic analysis since the
sequences of these species are not present in GenBank.
Russulales Pers.
Russulaceae Lotsy
Russulaceae is one of the dominant and morphologically
diverse families of mushrooms in the order Russulales,
with roughly 1900 known species and a worldwide distribution (Kirk et al. 2008). 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 genus Russula Pers. has a worldwide distribution
and can be easily identified with its colourful fruiting
bodies (Kirk et al. 2008). Russula is widely distributed
123
from Western Europe to North America in the northern
hemisphere (Romagnesi 1967; Singer 1986; Sarnari 1998;
Miller and Buyck 2002; Bau et al. 2008). The genus contains more than 200 species generally accepted in Europe
(Sarnari 1998, 2005) and more than 750 species worldwide
(Kirk et al. 2008). Keys to Russula being complex since of
this huge taxonomic diversity. They play a crucial beneficial role in forest ecosystems as ectomycorrhizal symbionts
(Li et al. 2013a). Russula species were used as traditional
food and medicine in China for a long history. Twenty-two
medicinal species and 82 edible species have been reported
in China (Dai et al. 2009; Li et al. 2010). Southwestern
China represents one of the world’s biodiversity ‘‘hotspots’’ and has a high diversity of macro-fungi (Yuan and
Dai 2008), but only a few russulacean species have been
identified from this region. However, some new species
and varieties have been reported from Southern and
Southwestern China (Wang et al. 2009). In this contribution one new species belong to genus Russula is introduced
from Guizhou Province, China (Fig. 191).
Russula yanheensis T.C. Wen, K. Hapuarachchi & K.D.
Hyde, sp. nov.
Index Fungorum number: IF 552563; Facesoffungi
number: FoF: 2707, Fig. 192
Etymology: refers to the type collecting site ‘‘Yanhe
County’’, Guizhou Province, China
Holotype: GACP13100308
Basidiocarps small-sized or medium-sized. Pileus 32–
70 mm diam., plano-convex, sometimes slightly depressed
above the stipe, surface smooth, sticky, broad, fleshy, firm,
hemispherical when young, then expanded with depressed
centre, depressed weakly in the center when mature,
extreme margin becoming somewhat sulcate with age,
glabrous, shining. Pileus peeling 1/2 to the disc; margin
even, crack when mature. Lamellae regular, adnate,
crowded, pale yellowish (2A2), brittle, edge entire. Stipe
23–42 9 13–25 mm, clavate (bottom slightly stigmas),
enlarged towards the base, centrally attached, orange white
(6A2) when young, pale coralline (9B7) when old, cylindrical to subcylindrical, stuffed, slightly longitudinally
wrinkled long and slender, flushed with pink Context
compact, 0.1–1.5 cm thick from stipe top to pileus center,
not firm, colour unchanged when bruised. Basidiospores
ornamentation amyloid, 5.87–8.26 9 4.88–7.31 lm
(
x = 6.23 9 7.11 lm, Q = 1.01–1.32, n = 28), subglobose to spherical, ellipsoid, colourless, without oil droplets,
moderately large and distant amyloid warts. Basidia,
broadly tapered towards the base, 48.7–77 9 10.3–12.05
lm (
x = 60 9 11.42 lm, Q = 4.32–5.58, n = 10), 4spored, clavate, hyaline, and smooth. Hymenophoral trama
cellular 18.52–33.20 9 16.02–28.58 lm (
x = 26.82 9
22.28 lm, Q = 0.99–1.45, n = 20), subglobose to
Fungal Diversity
Fig. 191 Phylogram generated from Maximum parsimony (PAUP)
analysis based on 5.8S-ITS rDNA sequence data. Bootstrap support
values for maximum likelihood (black) and maximum parsimony
(blue) greater than 75% are indicated above the nodes. The tree is
rooted with Albatrellus ovinus. The strain numbers are mentioned
after the species names. The new species is indicated in red and extype strains are indicated in black bold
spherical, ellipsoid, colourless, smooth. Cheilocystidia
73.52–103.79 9 10.29–13.57 lm (
x = 94.75 9 11.74 lm,
Q = 6.32 9 11, n = 8), clavate, fusiform to subfusiform,
papillae at the top, thin-wall, hyaline, smooth. Pilocystidium rare, emergent, 85.5–113 9 4.1–8.5 lm (
x = 95.28 9
6.7 lm, Q = 10.62–15.94, n = 7) clavate, thin-wall, hyaline, smooth, sometimes with a frayed small appendage and
dense crystal inclusions, projecting above the sub hymenium. Pileipellis an ixotrichoderm, composed of 4–6 lm
thick, cylindrical, rarely swollen hyphae with intracellular
reddish pigmentation; terminal cells. Often inflating up to
7–10 (12) lm thick and with brownish intracellular pigmentation, some fusiform and with zebroid-encrustations.
19.65–7.87 9 5.92–2.1 lm (
x = 16.77 9 3.05 lm,
Q = 2–11.2, n = 9). Stipitipellis a cutis with some hyphal
ends ascending; hyphae 4–6 lm diam., some hyphae near
surface yellowish-white; caulocystidia absent.
Habitat and distribution: On the ground, accompanied
in humus rich soil with over heavily rotted litter in forest,
mossy temperate mixed coniferous forests, solitary, producing basidiomata from summer to late autumn, only
found in ‘‘Yanhe County’’, Guizhou Province, China.
Material examined: CHINA, Guizhou Province, Coniferous mixed rainforest, 2800 N 108 ‘‘E, elev. 850 m, 3
October 2013, collector T.C Wen (GACP13100308, holotype). CHINA, Guizhou Province, Coniferous mixed
rainforest, 2800 N 108 ‘‘E, elev. 850 m, 3 October 2013,
collector T.C Wen (GACP13100319, paratype).
GenBank Number ITS:KY195927.
Notes: Russula yanheensis is morphologically and phylogenetically close to Russula pulchra Burl. Russula pulchra has 50–100 mm diameter pileus, stipe 30–70 mm
long, ellipsoid basidiospores 6.5–9 9 5.5–7.5 lm; and
pilocystidia absent (Binion et al. 2008), but R. yanheensis
has 32–70 mm diameter pileus, stipe 23–42 mm long,
ellipsoid to subglobose to globose basidiospores 5.8–8.3 9
4.8–7.3 lm, i.e. having smaller pileus, stipe and basidiospores, and the different shapes of basidiospores, presence of pilocystidia. Russula yanheensis is also
morphologically similar to R. flavisiccans Bills but the
latter species differs from our species by its pileus colour,
pilocystidia and cheilocystidia shapes and sizes, further,
the stipe never flushes with pink (Kuo and Methven 2014).
Further, R. yanheensis is phylogenetically closely related to
123
Fungal Diversity
123
Fungal Diversity
b Fig. 192 Russula yanheensis (GACP13100308, holotype). a Upper
surface. b Lower surface. c Section of the context layer (50um). d–
g Basidia/Basidiomes. Scale bars d = 50 lm, e–g = 10 lm, h–
k = 10 lm
R. sichuanensis G.J. Li & H.A. Wen, but its morphological
characteristics are very different from the latter species by
its colour, pilocystidia and cheilocystidia shapes and sizes.
Russula virescens (Schaeff.) Fr., Anteckningar öfver de i
Sverige växande ätliga svampar: 50 (1836)
Facesoffungi number: FoF 3145, Figs. 193, 194
Pileus 4–16 cm, when young round to convex, when
mature broadly convex to flat to uplifted with a shallow
depression, dry, velvety, surface soon cracking up into
small patches, green to yellowish-green, margin not lined;
the skin peeling towards the center. Lamellae attached to
the stem or at maturity nearly free, close or crowded, white
to cream. Stipe 2–11 cm long, 2–5 cm thick, dry, smooth,
white, brittle, discolouring brownish with age but not with
bruising or cutting. Context white, brittle, thick, not
changing when sliced. Odour and taste not distinctive, taste
mild. Spore print white. Basidiospores 6–9 9 5–7 lm,
elliptical to subglobose, warts 0.5 lm high, connectors
almost absent, scattered, or creating moderately reticulated
areas. Pleurocystidia very few. Pileipellis a cutis overlaid
with epithelium-like areas or crustose patches composed of
elements of chained cells diminishing in width from base to
tip, with the terminal cell projecting an extension that is
frequently elongated and tapered. Pileocystidia cylindric
with capitate apices, positive with sulphovanillin.
Habit, habitat and distribution: gregariously or solitary,
usually mycorrhizal with hardwoods, May–August. Our
collection was collected associated with Dipterocarpus sp.
at Peradeniya Royal Botanic Gardens.
Specimens examined: SRI LANKA, Kandy District, near
Peradeniya Royal Botanic Gardens, 2 June 2012, Samantha
C. Karunarathna (MFLU 12-1894, new record).
GenBank Number ITS:KY649467.
Notes: This has a widespread distribution in Asia, having been recorded from India, Malaysia, Korea, the
Philippines, Nepal, China, Thailand, and Vietnam. It is also
found in North Africa and Central America. This is the first
Fig. 193 Basidiocarps of
Russula virescens in the field
(MFLU 12-1894)
123
Fungal Diversity
Fig. 194 Russula virescens (MFLU 12-1894). a Basidia. b Pileipellis.
c Basidiospores. d Cheilocystidia. e Subcuticle. Scale bars a–
e = 10 lm
report of R. virescens with the molecular phylogenetic
confirmation from Sri Lanka.
Dacrymycetales Henn.
Dacrymycetaceae J. Schröt.
The family Dacrymycetaceae includes wood-decaying
fungi that cause brown-rot (Oberwinkler 1993; Kirk et al.
2001). This family includes eight genera: Calocera, Cerinomyces, Dacrymyces, Dacryopinax, Ditiola, Femsjonia,
Guepiniopsis and Heterotextus (Shirouzu et al. 2009).
Basidiomata of Dacrymycetaceae are characterized by
gelatinized yellow to orange basidiomata with Y-shaped
basidia (Kennedy 1958).
Dacrymyces Nees.
Dacrymyces is a genus in the order Dacrymycetales of
Dacrymycetes. The genus contains about 39 widely distributed species (Kirk et al. 2008) (Fig. 195).
123
Dacrymyces chiangraiensis Ekanayaka, S.C. Karunarathna, Q. Zhao & K D. Hyde, sp. nov.
Index Fungorum number: IF552549; Facesoffungi
number: FoF 2696, Fig. 196
Etymology: Name reflects the province, where the type
species was collected
Holotype: MFLU 16-0572
Saprobic on wood, stems and twigs. Sexual morph
Basidiomata 478–482 9 908–915 lm (
x = 480.8 9 911.6
lm, n = 10) superficial, mostly solitary, sometimes gregarious, pulvinate, yellow to orange. Sterile parts of
basidiocarps covered with simple or branched, tightly
packed, cylindrical, septate, hyaline, thin-walled marginal
hyphae. Internal hyphae branched, thin walled, gelatinous,
septate, hyaline, loosely packed, clamp connections absent.
Hymenium composed of dikaryophyses and basidia.
Dikaryophyses 2–5.5 lm (
x = 4 lm, n = 20) branched,
septate, thick-walled, hyaline. Hymenium hyaline, limited
to the upper surface of the basidiocarp. Probasidia 32–53
9 7–10 lm (
x = 41.4 9 8.5 lm, n = 20) hyaline, guttulate, cylindrical to clavate, thin-walled, becoming bifurcate. Basidiospore 19–23 9 6.5–8 lm (
x = 20.9 9 7.3 lm,
n = 20), allantoid, somewhat bent, hyaline, with many
transverse septa, smooth walled. Asexual morph
undetermined.
Material examined: THAILAND, Mae Fah Luang
University, Chiang Rai Province, 22 December 2014, A.H.
Ekanayaka, (MFLU 16-0572, holotype).
GenBank Number ITS:KY498587.
Notes: Dacrymyces chiangraiensis is phylogenetically
close to D.san-augustinii. However, they are separated
with strong bootstrap support. Morphologically D. chiangraiensis is similar to D. san-augustinii in having yellow to
orange pulvinate basidiomata, but they differ in the size of
the basidiomata and colour of probasidia and basidiospores. Dacrymyces san-augustinii has larger basidiomata (1–2 mm high, 1–3 mm diam.) with pale yellow
probasidia, and subhyaline, smaller basidiospores
(15.5–23.6 9 4.6–6.8 lm) (McNabb 1973). Dacrymyces
chiangraiensis differs from D. novae-zelandiae in having
yellow to orange non-gelatinized basidiomata with smaller
probasidia (Ellis and Ellis 1990; Shirouzu et al. 2009)
Femsjonia Fr.
The genus Femsjonia was established by Fries with the
type species Femsjonia luteoalba Fr. However, the systematic position of this genus was not clear and linked with
Peziza radiculata as the appearance of fruiting body is
similar to discomycetes (McNabb 1965). Currently this
genus classified within Dacrymycetaceae (Dacrymycetales,
Dacrymycetes, Agaricomycotina). The genus Femsjonia is
characterized by discoid, brightly coloured basidiomata
Fungal Diversity
Fig. 195 The best scoring RAxML maximum likelihood phylogenetic tree based on a combined ITS and SSU sequence dataset.
Strain/culture numbers are given following the taxon names. The
newly generated sequences are in blue. Bootstrap support values for
maximum likelihood [60% are given near the nodes. The tree is
rooted with Exidia uvapsassa (AFTOL ID 461)
with thick-walled internal hyphae and globose to sub-globose basidia (McNabb 1965; Shirouzu et al. 2009).
yellow. Internal hyphae branched, thin-walled, septate,
hyaline, loosely packed. Probasidia 100–150 9 7–10 lm
(
x = 139.6 9 8.2 lm, n = 20) cylindrical, yellow, thinwalled, becoming bifurcate. Basidiospores 25–30 9 10–15
lm (
x = 28.5 9 12.5 lm, n = 20), subglobose, guttulate,
aseptate, yellowish, smooth-walled.
Material examined: CHINA, Yunnan province, Kunming, 10 November 2015, S. C. Karunarathna (NB130),
(MFLU 16-0608, holotype).
GenBank Number ITS:KY498588.
Notes: Femsjonia monospora distinct from all the other
species in the genus by having larger probasidia and
aseptate, guttulate basidiospores (McNabb 1965; Liu et al.
Femsjonia monospora Ekanayaka, S.C. Karunarathna, Q.
Zhao & K.D. Hyde, sp. nov.
Index Fungorum number: IF552550; Facesoffungi
number: FoF 2697, Fig. 197
Etymology: Name reflects the aseptate basidiospores.
Holotype: MFLU 16-0608
Saprobic on wood, stems and twigs. Basidiomata
superficial, gregarious, discoid, sessile, bright yellow,
orange near the base, sterile surface rough tomentose,
whitish at the margins of the hymenial surface. Hymenium
123
Fungal Diversity
Fig. 196 Morphology of
Dacrymyces chiangraiensis
(MFLU 16-0572, holotype).
a Substrate. b Basidiomata on
wood. c Cross section of
basidiome. d Close up of the
hymenium layer and interscal
tissue. e Close up of hymenium
layer. f Septate, branched
hyphae. g, h Probasidia. i, j
Developing basidia. k, l
Ellipsoid basidiospores. Scale
bars a = 1000 lm, b = 200
lm, = 200 lm, c = 400 lm,
d = 100 lm, e = 100 lm,
f = 15 lm, g–j = 20 lm, k,
l = 5 lm
1988; Shirouzu et al. 2009). The phylogenetic comparisons
among Femsjonia species are not possible due to the lack
of sequence data.
Tremellales Fr.
Tremellaceae Fr.
This family was erected by Fries in 1821 based on
macromorphology of its members (Fries 1821). This family
currently comprises 18 genera and 250 valid species (Kirk
et al. 2008).
123
Tremella Pers.,
Tremella was one of the original genera created by
Linnaeus in his Species Plantarum of 1753. All Tremella
species are parasites of other fungi and most produce
anamorphic yeast states. Basidiocarps, when produced, are
gelatinous and are colloquially classed among the ‘‘jelly
fungi’’. Over 100 species of Tremella are currently recognized worldwide (Kirk et al. 2008). Two species, T. fuciformis and T. aurantialba, are commercially cultivated for
food (Fig. 198).
Fungal Diversity
Fig. 197 Morphology of Femsjonia monospora (MFLU 16-0608, holotype). a Basidiomata on wood. b Cross section of basidiome. c, d
Probasidia. e, f Developing basidia. g, h Immature and mature basidia. Scale bars b = 100 lm, c, d = 50 lm, e = 25 lm, f–h = 30 lm
Tremella fuciformis Berk., Hooker’s Journal of Botany
and Kew Garden Miscellany 8: 277 (1856)
Facesoffungi number: FoF 3146, Figs. 199, 200
Basidiocarps foliose, lobes caespitose, wet, gelatinous
but fairly hard, crisped, composed of graceful lobes, lobes
generally very thin, flower like, translucent whitish, up to
about 8 cm across and 5 cm high, surface smooth and
shiny, generally associated with Hypoxylon sp. Spore print
white. Basidiospores 7–15 9 5–9 lm, Qm = 1.2–1.6 lm,
broadly ellipsoid to ellipsoid, smooth, often germinating by
repetition, budding or sometimes germ tubes. Basidia
mostly 4–spored but occasionally 2 or 3 spored, becoming
longitudinally 4–septate with maturity, probasidia typically
clavate becoming subglobose to ellipsoid with maturity,
11–16 9 8–14 lm, with sterigmata up to 50 9 4 lm.
Conidia absent. Vesicles absent. Hyphidia absent. Hyphae
123
Fungal Diversity
Fig. 198 Phylogenetic
relationships inferred from
maximum parsimony analysis
of ITS-rDNA sequences of 40
taxa. The percentages of
replicate trees in which the
associated taxa clustered
together in the bootstrap test
(1000 replicates) are shown next
to the branches. The tree is
drawn to scale, with branch
lengths in the same units as
those of the evolutionary
distances used to infer the
phylogenetic tree. The new Sri
Lankan record: Tremella
fuciformis having GenBank
Number KY649461 and
Herbarium Number MFLU
12-1895 is shown in bold and
blue. The topology is rooted
with Filobasidium floriforme.
Evolutionary analysis was
conducted in PAUP 4.0b 10
123
Fungal Diversity
Fig. 199 Basidiocarps of Tremella fuciformis in the field (MFLU
12-1895)
2–6 lm diam. in inner part of basidiocarps, close to substrate 12 lm diam. Haustorial hyphae in basal part close to
substrate, often branched. Clamp connections present.
Habit, habitat and distribution: Tremella fuciformis,
following its host, fruit bodies are typically found on dead,
attached or recently fallen branches of broadleaf trees. This
fungus commonly prefers tropical and subtropical ecosystems, May–September. This species is mainly in tropical
and subtropical ecosystems, but does occur temperate areas
in Asia including Thailand; Laos and China, and North
America. It is found in South and Central America, the
Caribbean, parts of North America, sub-Saharan Africa,
southern and eastern Asia, Australia, New Zealand and the
Pacific Islands. Our collection was found on a fallen dead
tree branch near Hanthana Mountains Range, Peradeniya,
Sri Lanka.
Specimens examined: SRI LANKA, Kandy District,
Hanthana Mountains Range, 1 June 2012, Samantha C.
Karunarathna (MFLU 12-1895, new record).
GenBank Number ITS:KY649468.
Notes: The species is distributed mainly in tropical and
subtropical areas, but extends into temperate areas in Asia
and North America. It is known throughout South and
Fig. 200 Tremella fuciformis
(MFLU 12-1895). a Basidia and
attached basidiospores.
b Basidiospores. c Structure of
the inner part of the basidiocarp,
hyphae are swollen, thick
walled
123
Fungal Diversity
Central America, the Caribbean, parts of North America,
sub-Saharan Africa, southern and eastern Asia, Australia,
New Zealand, and the Pacific Islands. This is the first report
of T. fuciformis with the molecular phylogenetic confirmation from Sri Lanka.
Acknowledgements Saowaluck Tibpromma would like to thank the
Molecular Biology Experimental Center at Kunming Institute of
Botany for facilities for molecular work, the Mushroom Research
Foundation (MRF), Chiang Rai, Thailand for the financial support of
her study and Shaun Pennycook is thanked for nomenclatural advice.
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’’, the Chinese Academy of Sciences, Project Number
2013T2S0030, for the award of Visiting Professorship for Senior
International Scientists at Kunming Institute of Botany and National
Research Council of Thailand (Mae Fah Luang University) for a
grants ‘‘Biodiversity, phylogeny and role of fungal endophytes of
Pandanaceae’’ (Grant No.: 592010200112), ‘‘Diseases of mangrove
trees and maintenance of good forestry practice’’ (Grant No.:
60201000201) for supporting this study. 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
authors would like to extend their sincere appreciation to the
Deanship of Scientific Research at King Saud University for funding
this research group NO (RG-1436-025). Financial support by the
German Academic Exchange Service (DAAD) and the Thai Royal
Golden Ph.D. Jubilee-Industry program (RGJ) for a joint TRFDAAD 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
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 expresses sincere appreciation to The CAS President’s International Fellowship for Postdoctoral Researchers (Project No.
2017PB0072). 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). André Luiz Cabral Monteiro de
Azevedo Santiago, Carlos Alberto Fragoso de Souza, Diogo Xavier
Lima, Rafael José Vilela de Oliveira and Gladstone Alves da Silva
would like to thank the Coordenação de Aperfeiçoamento de Pessoal
de Nı́vel Superior (Coordination for the Improvement of Higher
Education Personnel) (CAPES) and the Fundação de Amparo à
Ciência e Tecnologia do Estado de Pernambuco (Foundation for the
support of Science and Technology of the state of Pernambuco)
(FACEPE) for the postgraduate scholarships awarded to Diogo X.
Lima and Carlos A. F. de Souza, respectively. We would also like to
thank the Conselho Nacional de Desenvolvimento Cientı́fico e Tecnológico (National Council for Scientific and Technological
123
Development) (CNPq) and FACEPE for financial support through
the projects: ‘Mucoromycotina in upland forests in the semi-arid
region of Pernambuco’ (CNPq-458391/2014-0), and ‘Diversity of
Mucoromycotina in different ecosystems of the Pernambuco Atlantic
Rainforest’ (FACEPE—APQ 0842-2.12/14). 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 Project on Discovery of Fungi
from Freshwater and Collection of Fungarium funded by 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. 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. Saranyaphat Boonmee thanks the National
Research Council of Thailand, project number 2560A30702021 and
the Thailand Research Fund, project number TRG5880152 for providing financial support. C.G. Lin and Y. Wang thank the grant from
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) and
Mr. Jingzu Sun thank for the National Natural Science Foundations
of China (No. 31600024). Wei Dong thanks the for National Natural
Science Foundation of China (Project ID: NSF 31500017 to Huang
Zhang). P.N. Singh, A. Baghela, S.K. Singh, and S. Aamir thank the
Director, MACS’ Agharkar Research Institute, Pune, India for providing facilities and Rajendra Singh (Department of Zoology, DDU
Gorakhpur University, UP, India) for identification of insect-host.
Saisamorn Lumyong and Rene K. Schumacher are thanked for
valuable suggestions and collecting specimens. K.N.A. Raj
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)). K.N.A. Raj also acknowledges 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 5 August
2009). K.P.D. Latha acknowledges the financial support from the
Kerala State Council for Science, Technology and Environment
(KSCSTE) in the form of a PhD fellowship (Grant No. 001/FSHP/
2011/CSTE). K.P.D. Latha also acknowledges the permission (No.
WL10-4937/2012, dated 3-10-2013) given to her by the Principal
Chief Conservator of Forests, Government of Kerala, to collect
agaric specimens from the forests of Kerala. Zdenko Tkalčec has
been partially supported by Croatian Science Foundation under the
project HRZZ-IP-11-2013-2202 (ACCTA) and is grateful to Milan
Čerkez for his great contribution to the study of coprinoid and
coprophilous fungi in Croatia. Vladimir Antonı́n thank the Moravian
Museum by the Ministry of Culture of the Czech Republic as part of
its long-term conceptual development programme for research
institutions (DKRVO, ref. MK000094862). T.C. Wen, Y.P. Xiao, C.
Norphanphoun and K.K. Hapuarachchi are grateful to the National
Natural Science Foundation of China (No. 31460012) and the Science and Technology Foundation of Guizhou Province (No.
[2016]2863). Y.W. Lim would like to thanks NIBR supporting the
Project on Survey and Discovery of Indigenous Fungal Species of
Korea. Kanad Das and Dyutiparna Chakraborty are thankful to the
Director, Botanical Survey of India (BSI) and Scientist-in-Charge,
BSI, Gangtok for providing facilities during this study. Sinchan
Adhikari, Joydeep Karmakar and Tapas Kumar Bandyopadhyay
would like to acknowledge DST-PURSE and DST-FIST for providing central instrumentation facilities and Alan JL Phillips
acknowledges the Biosystems and Integrative Sciences Institute
(BioISI, FCT/UID/Multi/04046/2013).
Fungal Diversity
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