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Fungal Diversity – March (2016)
Fungal diversity notes 253–366: taxonomic and phylogenetic contributions to
fungal taxa
Guo Jie Li1 ● Kevin D. Hyde2,3,4 ● Rui Lin Zhao1 ● Hongsanan Sinang2,3 ● Faten
Awad Abdel-Aziz5 ● Mohamed A. Abdel-Wahab4,5 ● Pablo Alvarado6 ●
Genivaldo Alves-Silva7 ● Joseph F. Ammirati8 ● Hiran A. Ariyawansa9 ●
Abhishek Baghela10 ● Ali Hassan Bahkali4 ● Michael Beug11.● D. Jayaram Bhat2
● Dimitar Bojantchev12 ● Thitiya Boonpratuang13 ● Timur S. Bulgakov14 ● Erio
Camporesi15,16,17 ● Marcela C. Boro18 ● Oldriska Ceska19 ● Dyutiparna
Chakraborty20 ● Jia Jia Chen21 ● K. W. Thilini Chethana2,22 ● Putarak
Chomnunti2 ● Giovanni Consiglio23 ● Bao Kai Cui21 ● Dong Qin Dai2● Yu
Cheng Dai21 ● Dinushani A. Daranagama1,2 ● Kanad Das20 ● Monika C.
Dayarathne2,3,24 ● Eske De Crop25 ● Rafael J. V. De Oliveira26 ● Carlos Alberto
Fragoso de Souza26 ● José I. de Souza18 ● Bryn T.M. Dentinger27,28 ● Asha J.
Dissanayake2,22 ● Mingkwan Doilom2,3 ● E. Ricardo Drechsler-Santos7 ●
Masoomeh Ghobad-Nejhad29 ● Sean P. Gilmore30 ● Aristóteles Góes-Neto31 ●
Michał Gorczak32 ● Charles H. Haitjema30 ● Kalani Kanchana Hapuarachchi2,33
● Akira Hashimoto34,35 ● Mao Qiang He1,36 ●John K. Henske30 ● Kazuyuki
Hirayama37 Maria J. Iribarren38 ● Subashini C. Jayasiri2 ● Timothy Y. James39
● Ruvishika S. Jayawardena2,22 ● Sun Jeong Jeon40 ● Gustavo H. Jerônimo18 ●
Ana L. Jesus18 ● E. B. Gareth Jones4 ● Ji Chuan Kang33 ● Samantha C.
Karunarathna2,3,24 ● Paul M. Kirk41 ● Sirinapa Konta2,3 ● Eric Kuhnert42,43 ●
Ewald Langer44 ● Haeng Sub Lee40 ● Hyang Burm Lee40 ● Wen Jing Li2,3 ● Xing
Hong Li22 ● Kare Liimatainen45 ● Diogo Xavier Lima26 ● Chuan Gen Lin2,46 ●
Jian Kui Liu9 ● Xings Zhong Liu1 ● Zuo Yi Liu9 ● J. Jennifer Luangsa-ard13 ●
Robert Lücking47 ● H. Thorsten Lumbsch48 ● Saisamorn Lumyong49 ● Eduardo
M. Leaño50 ● Agostina V. Marano18 ● Misato Matsumura34,35 ● Eric H. C.
McKenzie51 ● Suchada Mongkolsamrit13 ● Peter Mortimer3,24 ● Thi Thuong
Thuong Nguyen40 ● Tuula Niskanen27 ● Chada Norphanphoun2,3 ● Michelle A.
O’Malley30 ● Sittiporn Parnmen52 ● Julia Pawłowska32 ● Rekhani H. Perera2,3 ●
● Rungtiwa Phookamsak2,3 ● Chayanard Phukhamsakda2,3 ● Carmen L. A.
Pires-Zottarelli18 ● Olivier Raspé53,54 ● Mateus A. Reck7 ● Sarah C. O. Rocha18 ●
André L. C. M. de A. Santiago26 ● Indunil C Senanayake2 ● Ledo Setti55 ● Qiu
Ju Shang2 ● Sanjay K. Singh10 ● Esteban B. Sir56, 57 ● Kevin V. Solomon30 ● Jie
Song21 ● Prasert Srikitikulchai13 ● Marc Stadler42,43 ● Satinee Suetrong58 ●
Hayato Takahashi35 ● Takumasa Takahashi35 ● Kazuaki Tanaka35 ● Li Ping
Tang59 ● Kasun M. Thambugala2,9 ● Donnaya Thanakitpipattana13 ● Michael K.
Theodorou60 ● Benjarong Thongbai2 ● Tuksaporn Thummarukcharoen13 ● Qing
Tian2,3 ● Saowaluck Tibpromma2,3 ● Annemieke Verbeken25 ● Alfredo Vizzini61
● Josef Vlasák62 ● Kerstin Voigt63 ● Dhanushka N. Wanasinghe2,3 ● Yong
Wang46 ● Gothamie Weerakoon48 ● Hua An Wen1 ● Ting Chi Wen33 ● Nalin N.
Wijayawardene2 ● Sarunyou Wongkanoun13 ● Marta Wrzosek32 ● Yuan Pin
Xiao2,33 ● Jian Chu Xu3 ● Ji Ye Yan22 ● Jing Yang2,9 ● Shu Da Yang59 ● Yu Hu
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● Jin Feng Zhang2,9 ● Jie Zhao30 ● Li Wei Zhou64 ● Derek Peršoh65 ● Alan J.
L. Phillips 66● Sajeewa S. N. Maharachchikumbura67
1
R. L. Zhao, G. J. Li, D. A. Daranagama, M. Q. He, H. A. Wen, X. Z. Liu
State Key Laboratory of Mycology, Institute of Microbiology, Chinese Academy of
Sciences, No 3 1st West Beichen Road, Chaoyang District, Beijing 100101, P. R.
China
*Corresponding author: Rui Lin Zhao (zhaorl@im.ac.cn)
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2
K. D. Hyde, S. Hongsanan, D. Q. Dai, D. A. Daranagama, M. C. Dayarathne, K. W.
T. Chethana, A. J. Dissanayake, M. Doilom, K. K. Hapuarachchi, C. S. Jayasiri, R. S.
Jayawardena, S. C. Karunarathna, S. Konta, W. J. Li, C. G. Lin, Q. J. Shang, C.
Norphanphoun, R. H. Perera, R. Phookamsak, C. Phukhamsakda, I. C Senanayake, B.
Thongbai, K. M. Thambugala, Q. Tian, S. Tibpromma, D. N. Wanasinghe, N.
Wijayawardene, Y. P. Xiao, J. Yang, J. F. Zhang
Centre of Excellence in Fungal Research and School of Science, Mae Fah Luang
University, Chiang Rai, 57100, Thailand
3
K. D. Hyde, S. Hongsanan, D. Q. Dai, M. C. Dayarathne, S. C. Karunarathna, M.
Doilom, S. Konta, W. J. Li, P. Mortimer, C. Norphanphoun, R. H. Perera, R.
Phookamsak, C. Phukhamsakda, Q. Tian, S. Tibpromma, D. N. Wanasinghe, J.C. Xu
Key Laboratory for Plant Diversity and Biogeography of East Asia, Kunming Institute
of Botany, Chinese Academy of Science, Kunming 650201, Yunnan, P. R. China
4
K. D. Hyde, M. A. Abdel-Wahab, A. H. Bahkali, E. B. G. Jones
Department of Botany and Microbiology, College of Science, King Saud University,
P.O. Box: 2455, Riyadh 1145, Saudi Arabia
5
F. A. Abdel-Aziz, M. A. Abdel-Wahab
Department of Botany and Microbiology, Faculty of Science, Sohag University,
Sohag 82524, Egypt
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P. Alvarado
ALVALAB, C/ La Rochela nº 47, E-39012, Santander, Spain
7
G. Alves-Silva, M. A. Reck, E. R. Drechsler-Santos
Micolab, Programa de Pós-Graduação em Biologia de Fungos, Algas e Plantas,
Departamento de Botânica, Universidade Federal de Santa Catarina, Campus
Universitário Trindade, CEP: 88040-900, Florianópolis, Santa Catarina, Brazil
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J. Ammirati
Department of Biology, University of Washington, Box 351800, Seattle, Washington
98195–1800, U.S.A.
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H. A. Ariyawansa, J. K. Liu, Z. Y. Liu, S. S. N. Maharachchikumbura, K. M.
Thambugala, J. Yang, J. F. Zhang
Guizhou Key Laboratory of Agricultural Biotechnology, Guizhou Academy of
Agricultural Sciences, Guiyang 550006, Guizhou, P. R. China
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A. Baghela, S.K. Singh
National Fungal Culture Collection of India (NFCCI), Biodiversity and Palaeobiology
Group, MACS'Agharkar Research Institute, G.G. Agarkar Road, Pune 411004, India
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M. Beug
PO Box 116 Husum, WA 98623–0116, U.S.A.
12
D. Bojantchev
MushroomHobby.com, 345 Shipwatch Lane, Hercules, CA 94547 U.S.A.
13
T. Boonpratuang, J. J. Luangsa-ard, S. Mongkolsamrit, P. Srikitikulchai, D.
Thanakitpipattana, T. Thummarukcharoen, S. Wongkanoun
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
14
T. S. Bulgakov
Academy of Biology and Biotechnology,
Rostov-on-Don 344090, Rostov region, Russia
Southern
Federal
University,
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E. Camporesi
A.M.B. Gruppo Micologico Forlivese “Antonio Cicognani”, Via Roma 18, Forlì, Italy
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E. Camporesi
A.M.B. Circolo Micologico “Giovanni Carini”, C.P.314, Brescia, Italy
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E. Camporesi
Società per gli Studi Naturalistici della Romagna, C.P. 144, Bagnacavallo (RA), Italy
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M. C. Boro, G. H. Jerônimo, A. L. Jesus, J. I. de Souza, A. V. Marano, S. C. O.
Rocha, C. L. A. Pires-Zottarelli
Núcleo de Pesquisa em Micologia, Instituto de Botânica, Av. Miguel Stéfano 3687,
04301-912, São Paulo, SP, Brazil
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O. Ceska
1809 Penshurst Rd., Victoria, British Columbia, V8N 2N6 Canada
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D. Chakraborty, K. Das,
Cryptogamic Unit, Botanical Survey of India, Botanic Garden, Howrah 711103, India
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J. J. Chen, B. K. Cui, Y. C. Dai, J. Song
Institute of Microbiology, Beijing Forestry University, PO Box 61, Beijing 100083, P.
R. China
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K. W. T. Chethana, A. J. Dissanayake, R. S. Jayawardena, X. H. Li, J. Y. Yan
Institute of Plant and Environment Protection, Beijing Academy of Agriculture and
Forestry Sciences, No. 9 of ShuGuangHuaYuanZhongLu, Haidian District,
Beijing 100097, P. R. China
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G. Consiglio
Via C. Ronzani 61, I-40033, Casalecchio di Reno (BO), Italy
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M. C. Dayarathne, S. C. Karunarathna, D. N. Wanasinghe
Soil Biology Group, World Agro forestry Centre East and Central Asia Office, Lanhei
Road, Heilongtan, Kunming 650201, Yunnan, P. R. China
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E. De Crop, A. Verbeken
Research Group Mycology, Department of Biology, Ghent University, K.L.
Ledeganckstraat 35, B-9000 Gent, Belgium
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R. J. V. de Oliveira, D. X. Lima, C. A. F. de Souza, A. L. C. M. A. Santiago
PostGraduate Program in Biology of Fungi, Department of Mycology, Federal
University of Pernambuco, Av. Nelson Chaves, s/n, 50670-420, Recife, PE, Brazil
27
B. T. M. Dentinger, T. Niskanen
Jodrell Laboratory, Royal Botanic Gardens, Kew, Surrey TW9 3DS, UK
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B. T. M. Dentinger
Institute of Biological, Environmental and Rural Sciences, Cledwyn Building,
Aberystwyth University, Penglais, Aberystwyth, Ceredigion SY23 3DD, UK
29
M. Ghobad-Nejhad
Department of Biotechnology, Iranian Research Organization for Science and
Technology (IROST), P. O. Box 3353-5111, Tehran 3353136846, Iran
S. P. Gilmore, C. H. Haitjema, J. K. Henske, M. A. O’Malley, K. V. Solomon
Department of Chemical Engineering, University of California Santa Barbara, Santa
Barbara, CA, 93106, USA
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A. Góes-Neto
Laboratório de Pesquisa em Microbiologia (LAPEM), Departamento de Ciências
Biológicas, Programa de Pós-Graduação em Botânica, Feira de Santana, CEP:
44036-900, Bahia, Brazil
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M. Gorczak, J. Pawłowska, M. Wrzosek
Department of Molecular Phylogenetics and Evolution, Institute of Botany, Faculty of
Biology, University of Warsaw, Al. Ujazdowskie 4, 00-478 Warsaw, Poland
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K. K. Hapuarachchi, J. C. Kang, T. C. Wen, Y. P. Xiao
The Engineering and Research Center for Southwest Bio-Pharmaceutical Resources,
Ministry of Education, Guizhou University, Huaxi District, Guiyang 550025, P.R.
China
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A. Hashimoto, M. Matsumura
The United Graduate School of Agricultural Sciences, Iwate University, 18-8 Ueda 3
chome, Morioka 020-8550, Japan
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A. Hashimoto, M. Matsumura, K. Tanaka, H. Takahashi, T. Takahashi
Faculty of Agriculture and Life Sciences, Hirosaki University, 3 Bunkyo-cho,
Hirosaki, Aomori 036-8561, Japan
36
M. Q. He
Key Laboratory of Forest Disaster Warning and Control in Yunnan Province, Faculty
of Biology Conservation, Southwest Forestry University, Kunming 650224, P. R.
China
37
K. Hirayama
Apple Experiment Station, Aomori Prefectural Agriculture and Forestry Research
Center, 24 Fukutami, Botandaira, Kuroishi, Aomori 036-0332, Japan
38
M. J. Iribarren
Universidad Nacional de Luján (UNLu), Ruta 5 y Avenida Constitución, 6700, Luján,
Buenos Aires, Argentina
39
T. Y. James
Department of Ecology and Evolutionary Biology, University of Michigan, Ann
Arbor, MI 48109, USA
40
S. J. Jeon, H. S. Lee, H. B. Lee, T. T. T. Nguyen
Division of Food Technology, Biotechnology & Agrochemistry, College of
Agriculture and Life Sciences, Chonnam National University, Gwangju 61186, R.O.
Korea
41
P. M. Kirk
Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3DS, UK
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E. Kuhnert, M. Stadler
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Department of Microbial Drugs, Helmholtz-Zentrum für Infektionsforschung GmbH,
Inhoffenstrasse 7, 38124 Braunschweig, Germany
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E. Kuhnert, M. Stadler
Partner site Hannover-Braunschweig, German Centre for Infection Research (DZIF),
38124 Braunschweig, Germany
44
E. Langer
Department of Ecology, University of Kassel, Heinrich-Plett-Str. 40, D-34132 Kassel,
Germany
45
K. Liimatainen
Department of Biosciences, Plant Biology, University of Helsinki, P.O. Box 65,
FI–00014 University of Helsinki, Finland
46
C. G. Lin, Y. Wang
Department of Plant Pathology, College of Agriculture, Guizhou University, Guiyang,
Guizhou 550025, P.R. China
47
R. Lücking
Botanischer Garten & Botanisches Museum Berlin, Freie Universität Berlin,
Königin-Luise-Straße 6–8, 14195 Berlin, Germany
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H. T. Lumbsch, G. Weerakoon
Science & Education, The Field Museum, 1400 South Lake Shore, Drive, Chicago, IL
60605-2496, U.S.A
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S. Lumyong
Department of Biology, Faculty of Science, Chiang Mai University, Chiang Mai,
50200, Thailand
50
E. M. Leaño
Network of Aquaculture Centres in Asia-Pacific, Suraswadi Building, Kasetsart
University Campus, Jatujak, Ladyao, Bangkok 10900, Thailand
51
E. H. C. McKenzie
Landcare Research, Private Bag 92170, Auckland Mail Centre, Auckland 1142, New
Zealand
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S. Parnmen
Toxicology Center, National Institute of Health, Department of Medical Sciences,
Ministry of Public Health, Nonthaburi 11000, Thailand
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O. Raspé
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Botanic Garden Meise, Nieuwelaan 38, 1860 Meise, Belgium
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O. Raspé
Fédération Wallonie-Bruxelles, Service général de l'Enseignement Universitaire et de
la Recherche Scientifique, Rue A. Lavallée 1, 1080 Bruxelles, Belgium
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L. Setti
Via C. Pavese, 1, I-46029, Suzzara (MN), Italy
56
E. B. Sir
Fundación Miguel Lillo, Laboratory of Mycology, Miguel Lillo 251, San Miguel de
Tucumán 4000, Tucumán, Argentina
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E. B. Sir
Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET).
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S. Suetrong
Fungal Biodiversity Laboratory (BFBD), BIOTEC, National Science and Technology
Development Agency (NSTDA), 113 Thailand Science Park, Thanon Phahonyothin,
Tombon Khlong Nueng, Amphoe Khlong Luang, Pathum Thani 12120, Thailand
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L. P. Tang, J. Zhao, S. D. Yang, H. Yu
School of Pharmaceutical Sciences and Yunnan Key Laboratory of Pharmacology for
Natural Products, Kunming Medical University, 1168 West Chunrong Road, Yuhua
Avenue, Chenggong District, Kunming 650500, P. R. China
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M. K. Theodorou
Animal Production, Welfare and Veterinary Sciences, Harper Adams University,
Newport, Shropshire, TF10 8NB, United Kingdom
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A. Vizzini
Department of Life Sciences and Systems Biology, Università di Torino, Viale P.A.
Mattioli 25, I-10125, Torino, Italy
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J. Vlasák
Biology Centre of the Academy of Sciences of the Czech Republic, Institute of Plant
Molecular Biology, Branišovská 1160/31, CZ – 370 05 České Budějovice, Czech
Republic
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K. Voigt
Jena Microbial Resource Collection, Leibniz Institute for Natural Product Research
and Infection Biology, Adolf-Reichwein-Strasse 23, 07745 Jena, Germany
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L. W. Zhou
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Institute of Applied Ecology, Chinese Academy of Sciences, 72 Wenhua Road,
Shenyang 110016, P. R. China
D. Peršoh
Geobotany, Ruhr-Universität Bochum, Universitätsstraße 150, 44780 Bochum,
Germany
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A. J. L. Phillips
University of Lisbon, Faculty of Sciences, Biosystems and Integrative Sciences
Institute (BioISI), Campo Grande, 1749-016 Lisbon, Portugal
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S. S. N. Maharachchikumbura
Department of Crop Sciences, Sultan Qaboos University, Box 34, Al Khod 123,
Oman
Abstract Notes on 113 fungal taxa are compiled in this paper, including 11 new
genera, 89 new species, one new subspecies, three new combinations and xx reference
specimens. A wide geographic and taxonomic range of fungal taxa are detailed. In the
Ascomycota the new genera Angustospora (Testudinaceae), Camporesia
(Xylariaceae), Clematidis, Crassiparies (Pleosporales genera incertae sedis),
Farasanispora, Longiostiolum (Pleosporales genera incertae sedis), Multilocularia
(Parabambusicolaceae),
Neophaeocryptopus
(Dothideaceae),
Parameliola
(Pleosporales genera incertae sedis), and Towyspora (Lentitheciaceae) are
introduced. Newly introduced species are Angustospora nilensis, Aniptodera
aquibella, Annulohypoxylon albidiscum, Astrocystis thailandica, Camporesia
sambuci, Clematidis italica, Colletotrichum menispermi, C. quinquefoliae,
Comoclathris pimpinellae, Crassiparies quadrisporus, Cytospora salicicola, Diatrype
thailandica, Dothiorella rhamni, Durotheca macrostroma, Farasanispora avicenniae,
Halorosellinia rhizophorae, Humicola koreana, Hypoxylon lilloi, Kirschsteiniothelia
tectonae, Lindgomyces okinawaensis, Longiostiolum tectonae, Lophiostoma
pseudoarmatisporum,
Moelleriella
phukhiaoensis,
M.
pongdueatensis,
Mucoharknessia
anthoxanthi,
Multilocularia
bambusae,
Multiseptospora
thysanolaenae, Neophaeocryptopus cytisi, Ocellularia arachchigei, O. ratnapurensis,
Ochronectria thailandica, Ophiocordyceps karstii, Parameliola acaciae, P.
dimocarpi, Parastagonospora cumpignensis, Pseudodidymosphaeria phlei,
Polyplosphaeria thailandica, Pseudolachnella brevifusiformis, Psiloglonium
macrosporum, Rhabdodiscus albodenticulatus, Rosellinia chiangmaiensis,
Saccothecium rubi, Seimatosporium pseudocornii, S. pseudorosae, Sigarispora
ononidis and Towyspora aestuari. New combinations are provided for
Eutiarosporella dactylidis (sexual morph described and illustrated) and
Pseudocamarosporium pini. Descriptions, illustrations and / or reference specimens
are designated for Aposphaeria corallinolutea, Cryptovalsa ampelina, Dothiorella
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vidmadera, Ophiocordyceps formosana, Petrakia echinata, Phragmoporthe conformis
and Pseudocamarosporium pini. The new species of Basidiomycota are Agaricus
coccyginus, A. luteofibrillosus, Amanita atrobrunnea, A. digitosa, A. gleocystidiosa,
A. pyriformis, A. strobilipes, Bondarzewia tibetica, Cortinarius albosericeus, C.
badioflavidus, C. dentigratus, C. duboisensis, C. fragrantissimus, C. roseobasilis, C.
vinaceobrunneus,
C.
vinaceogrisescens,
C.
wahkiacus,
Cyanoboletus
hymenoglutinosus,
Fomitiporia
atlantica,
F.
subtilissima,
Ganoderma
wuzhishanensis, Inonotus shoreicola, Lactifluus armeniacus, L. ramipilosus,
Leccinum indoaurantiacum, Musumecia alpinaare, M. sardoa, Russula amethystina
subp. tengii and R. wangii are introduced. Descriptions, illustrations, notes and / or
reference specimens are designated for Clarkeinda trachodes, Dentocorticium
ussuricum, Galzinia longibasidia, Lentinus stuppeus and Leptocorticium tenellum.
The other new genera, species new combinations are Anaeromyces robustus,
Neocallimastix californiae and Piromyces finnis from Neocallimastigomycota,
Phytophthora estuarina, P. rhizophorae, Salispina, S. intermedia, S. lobata and S.
spinosa from Oomycota, and Absidia stercoraria, Gongronella orasabula,
Mortierella calciphila, Mucor caatinguensis, M. koreanus, M. merdicola and
Rhizopus koreanus in Zygomycota.
Keywords: Ascomycota, Basidiomycota, Neocallimastigomycota,
Zygomycota, Phylogeny, Taxonomy, new genus, new species
Oomycota,
Table of Contents
Ascomycota
Dothideomycetes
Botryosphaeriales
Botryosphaeriaceae
253.
Dothiorella rhamni Wanasinghe, Bulgakov, E.B.G. Jones & K.D. Hyde, in
Fungal Diversity 78: xx (2016)
254.
Dothiorella vidmadera W.M. Pitt et al.
255.
Eutiarosporella dactylidis (K.M. Thambugala, Camporesi & K.D. Hyde)
Dissanayake, Camporesi & K.D. Hyde, in Fungal Diversity 78: xx (2016)
256.
Mucoharknessia anthoxanthi Dissanayake, Camporesi & K.D. Hyde, in
Fungal Diversity 78: xx (2016)
Dothideales
392
393
394
395
396
397
398
399
400
401
402
403
404
405
406
407
408
409
410
411
412
413
414
415
416
417
418
419
420
421
422
423
424
425
426
427
428
429
430
431
432
433
434
Dothideaceae
257.
Neophaeocryptopus Wanasinghe, Camporesi, E.B.G. Jones & K.D. Hyde, in
Fungal Diversity 78: xx (2016)
258.
Neophaeocryptopus cytisi Wanasinghe, Camporesi, E.B.G. Jones & K.D.
Hyde, in Fungal Diversity 78: xx (2016)
Saccotheciaceae
259.
Saccothecium rubi Jayasiri, Wanasinghe, Camporesi & K.D. Hyde, in
Fungal Diversity 78: xx (2016)
Subclass Pleosporomycetidae
Hysteriales
Hysteriaceae
260.
Psiloglonium macrosporum Thambugala, Senan. & K.D. Hyde, in Fungal
Diversity 78: xx (2016)
Pleosporales
Didymosphaeriaceae
261.
Pseudocamarosporium pini Phukhamsakda, Camporesi & K.D. Hyde, in
Fungal Diversity 78: xx (2016)
Lentitheciaceae
262.
Towyspora Wanasinghe, E.B.G. Jones & K.D. Hyde, in Fungal Diversity 78:
xx (2016)
263.
Towyspora aestuari Wanasinghe, E.B.G. Jones & K.D. Hyde, in Fungal
Diversity 78: xx (2016)
Lindgomycetaceae
264.
Lindgomyces okinawaensis Tak. Takah. & Kaz. Tanaka, in Fungal Diversity
78: xx (2016)
Lophiostomataceae
265.
Lophiostoma pseudoarmatisporum Hay. Takah., K. Hiray. & Kaz. Tanaka,
in Fungal Diversity 78: xx (2016)
435
436
437
438
439
440
441
442
443
444
445
446
447
448
449
450
451
452
453
454
455
456
457
458
459
460
461
462
463
464
465
466
467
468
469
470
471
472
473
474
475
476
477
478
266.
Sigarispora ononidis Qing Tian, Thambug., Camporesi & K.D. Hyde, in
Fungal Diversity 78: xx (2016)
Melanommataceae
267.
Aposphaeria corallinolutea in Gruyter et al.
Parabambusicolaceae
268.
Multilocularia Phookamsak, Ariyawansa & K.D. Hyde, gen. nov., in Fungal
Diversity 78: XX (2016)
269.
Multilocularia bambusae Phookamsak, Ariyawansa & K.D. Hyde, in Fungal
Diversity 78: XX (2016)
270.
Multiseptospora thysanolaenae Phookamsak, Ariyawansa & K.D. Hyde, in
Fungal Diversity 78: XX (2016)
Phaeosphaeriaceae
271.
Parastagonospora cumpignensis Tibpromma, Camporesi & K.D. Hyde, in
Fungal Diversity 78: xx (2016)
Pleosporaceae
272.
Comoclathris pimpinellae Konta., Bulgakov & K.D. Hyde, in Fungal
Diversity 78: xx (2016)
Testudinaceae
273.
274.
Angustospora Abdel-Aziz in Fungal Diversity 78: xx (2016)
Angustospora nilensis Abdel-Aziz, in Fungal Diversity 78: xx (2016)
Tetraplosphaeriaceae
275.
Polyplosphaeria thailandica C.G. Lin, Yong Wang bis & K.D. Hyde, in
Fungal Diversity 78: xx (2016)
Pleosporales suborder Massarineae, incertae sedis
276.
Longiostiolum Doilom, Ariyawansa & K.D. Hyde, in Fungal Diversity 78:
xx (2016)
277.
Longiostiolum tectonae Doilom, D.J. Bhat & K.D. Hyde, in Fungal
Diversity 78: xx (2016)
278.
Pseudodidymosphaeria phlei Phukhamsakda, Camporesi & K.D. Hyde, in
Fungal Diversity 78: xx (2016)
479
480
481
482
483
484
485
486
487
488
489
490
491
492
493
494
495
496
497
498
499
500
501
502
503
504
505
506
507
508
509
510
511
512
513
514
515
516
517
518
519
520
521
522
Pleosporales genera, incertae sedis
279.
Clematidis Tibpromma, Camporesi & K.D. Hyde, in Fungal Diversity 78: xx
(2016)
280.
Clematidis italica Tibpromma, Camporesi & K.D. Hyde, in Fungal Diversity
78: xx (2016)
281.
Crassiparies Matsumura, K. Hiray. & Kaz. Tanaka, in Fungal Diversity 78:
xx (2016)
282.
Crassiparies quadrisporus Matsumura, K. Hiray. & Kaz. Tanaka, in Fungal
Diversity 78: xx (2016)
283.
Farasanispora Abdel-Wahab, Bahkali & E.B.G. Jones, gen. nov.
284.
Farasanispora avicenniae Abdel-Wahab, Bahkali & E.B.G. Jones, in Fungal
Diversity 78: xx (2016)
285.
Parameliola Hongsanan, Peršoh & K.D. Hyde, in Fungal Diversity 78: xx
(2016)
286.
Parameliola dimocarpi Hongsanan & K.D. Hyde, in Fungal Diversity 78: xx
(2016)
287.
Parameliola acaciae Hongsanan & K.D. Hyde, in Fungal Diversity 78: xx
(2016)
Dothideomycetes family, incertae sedis
Kirschsteiniotheliaceae
288.
Kirschsteiniothelia tectonae Doilom, D.J. Bhat & K.D. Hyde, in Fungal
Diversity 78: xx (2016)
Lecanoromycetes
Ostropales
Graphidaceae
289.
Ocellularia arachchigei Weerakoon, Lücking & Lumbsch, in Fungal
Diversity 78: xx (2016)
290.
Ocellularia ratnapurensis Weerakoon, Lücking & Lumbsch, in Fungal
Diversity 78: xx (2016)
291.
Rhabdodiscus albodenticulatus Weerakoon, Lücking & Lumbsch, in Fungal
Diversity 78: xx (2016)
Sordariomycetes
Chaetosphaeriales
523
524
525
526
527
528
529
530
531
532
533
534
535
536
537
538
539
540
541
542
543
544
545
546
547
548
549
550
551
552
553
554
555
556
557
558
559
560
561
562
563
564
565
566
Chaetosphaeriaceae
292.
Pseudolachnella brevifusiformis A. Hashim. & Kaz. Tanaka, in Fungal
Diversity 78: xx (2016)
Diaporthales
Gnomoniaceae
293.
Phragmoporthe conformis R.H. Perera & K.D. Hyde, in Fungal Diversity
78: xx (2016)
Valsaceae
294.
Cytospora salicicola C. Norphanphoun, Bulgakov & K.D. Hyde, in Fungal
Diversity 78: xx (2016)
Glomerellales
Glomerellaceae
295.
Colletotrichum menispermi Chethana, Jayawardena, Bulgakov & K.D.
Hyde, in Fungal Diversity 78: xx (2016)
296.
Colletotrichum quinquefoliae Jayawardena, Bulgakov & K.D. Hyde, in
Fungal Diversity 78: xx (2016)
Hypocreales
Bionectriaceae
297.
Ochronectria thailandica Q.J. Shang & K.D. Hyde, in Fungal Diversity 78:
xx (2016)
Clavicipitaceae
298.
Moelleriella phukhiaoensis Mongkol., Thanakitp. & Luangsa-ard, in Fungal
Diversity 78: xx (2016)
299.
Moelleriella pongdueatensis Mongkol., Thanakitp. & Luangsa-ard, in
Fungal Diversity 78: xx (2016)
Ophiocordycipitaceae
300.
Ophiocordyceps formosana Y.W. Wang et al.
567
568
569
570
571
572
573
574
575
576
577
578
579
580
581
582
583
584
585
586
587
588
589
590
591
592
593
594
595
596
597
598
599
600
601
602
603
604
605
606
607
608
609
610
301.
Ophiocordyceps karstii T.C. Wen, Y.P. Xiao & K.D. Hyde, sp. nov., in
Fungal Diversity 78: xx (2016)
Microascales
Halosphaeriaceae
302.
Aniptodera aquibella J. Yang & K.D. Hyde, in Fungal Diversity 78: xx
(2016)
Sordariales
Chaetomiaceae
303.
Humicola koreana Hyang B. Lee & T.T.T. Nguyen, in Fungal Diversity 78:
xx (2016)
Amphisphaeriales
Amphisphaeriaceae
304.
Seimatosporium pseudocornii Wijayaw., Camporesi & K.D. Hyde, in
Fungal Diversity 78: xx (2016)
305.
Seimatosporium pseudorosae Wijayaw., Camporesi & K.D. Hyde, in Fungal
Diversity 78: xx (2016)
Xylariales
Diatrypaceae
306.
Cryptovalsa ampelina (Nitschke) Fuckel
307.
Diatrype thailandica R.H. Perera, J.K. Liu & K.D. Hyde, in Fungal Diversity
78: xx (2016)
Xylariaceae
308.
Annulohypoxylon albidiscum J.F. Zhang, J.K. Liu, K.D. Hyde & Z.Y. Liu,
in Fungal Diversity 78: xx (2016)
309.
Astrocystis thailandica Daranagama and K.D. Hyde, in Fungal Diversity 78:
xx (2016)
310.
Camporesia W.J. Li & K.D. Hyde, in Fungal Diversity 78: xx (2016)
311.
Camporesia sambuci W.J. Li & K.D. Hyde, in Fungal Diversity 78: xx
(2016)
312.
Durotheca macrostroma Srikitik., Wongkanoun & Luangsa-ard, in Fungal
Diversity 78: xx (2016)
611
612
613
614
615
616
617
618
619
620
621
622
623
624
625
626
627
628
629
630
631
632
633
634
635
636
637
638
639
640
641
642
643
644
645
646
647
648
649
650
651
652
653
313.
Halorosellinia rhizophorae Dayarathne, E.B.G. Jones & K.D. Hyde, in
Fungal Diversity 78: xx (2016)
314.
Hypoxylon lilloi Sir, Lambert & Kuhnert, in Fungal Diversity 78: xx (2016)
315.
Rosellinia chiangmaiensis Daranagama and K. D. Hyde, in Fungal Diversity
78: xx (2016)
Ascomycota, genera incertae sedis
316.
Petrakia echinata (Peglion) Syd. & P. Syd.
Basidiomycota
Agaricomycetes
Agaricales
Agaricaceae
317.
Agaricus coccyginus M.Q. He & R.L. Zhao, in Fungal Diversity 78: xx
(2016)
318.
Agaricus luteofibrillosus M.Q. He, L.J. Chen & R.L. Zhao, in Fungal
Diversity 78: xx (2016)
319.
Clarkeinda trachodes (Berk.) Singer
Amanitaceae
320.
Amanita atrobrunnea Thongbai, Raspé & K.D. Hyde, in Fungal Diversity
78: xx (2016)
321.
Amanita digitosa Boonprat. & Parnmen, in Fungal Diversity 78: xx (2016)
322.
Amanita gleocystidiosa Boonprat. & Parnmen, in Fungal Diversity 78: xx
(2016)
323.
Amanita pyriformis Boonprat. & Parnmen, in Fungal Diversity 78: xx (2016)
324.
Amanita strobilipes Thongbai, Raspé & K.D. Hyde, in Fungal Diversity 78:
xx (2016)
Cortinariaceae
325.
Cortinarius albosericeus Ammirati, Beug, Liimat., Niskanen & O. Ceska, in
Fungal Diversity 78: xx (2016)
326.
Cortinarius badioflavidus Ammirati, Beug, Niskanen, Liimat. & Bojantchev,
in Fungal Diversity 78: xx (2016)
327.
Cortinarius denigratus Ammirati, Beug, Niskanen, Liimat. & O. Ceska, in
Fungal Diversity 78: xx (2016)
654
655
656
657
658
659
660
661
662
663
664
665
666
667
668
669
670
671
672
673
674
675
676
677
678
679
680
681
682
683
684
685
686
687
688
689
690
691
692
693
694
695
696
328.
Cortinarius duboisensis Ammirati, Beug, Niskanen & Liimat., in Fungal
Diversity 78: xx (2016)
329.
Cortinarius fragrantissimus Ammirati, Beug, Liimat., Niskanen & O.
Ceska, in Fungal Diversity 78: xx (2016)
330.
Cortinarius roseobasilis Ammirati, Beug, Liimat., Niskanen & O. Ceska, in
Fungal Diversity 78: xx (2016)
331.
Cortinarius vinaceobrunneus Ammirati, Beug, Liimat., Niskanen & O.
Ceska, in Fungal Diversity 78: xx (2016)
332.
Cortinarius vinaceogrisescens Ammirati, Beug, Liimat. & Niskanen, in
Fungal Diversity 78: xx (2016)
333.
Cortinarius wahkiacus Ammirati, Beug, Liimat. & Niskan, in Fungal
Diversity 78: xx (2016)
Tricholomataceae
334.
Musumecia alpina L.P. Tang, J Zhao & S.D. Yang, in Fungal Diversity 78:
xx (2016)
335.
Musumecia sardoa G. Consiglio, A. Vizzini & L. Setti, in Fungal Diversity
78: xx (2016)
Boletales
Boletaceae
336.
Cyanoboletus hymenoglutinosus D. Chakr., K. Das, A. Baghela, S.K. Singh
& Dentinger, in Fungal Diversity 78: xx (2016)
337.
Leccinellum indoaurantiacum D. Chakr., K. Das, A. Baghela, S.K. Singh &
Dentinger, in Fungal Diversity 78: xx (2016)
Polyporales genus, incertae sedis
338.
Galzinia longibasidia Hallenb.
Russulales genus, incertae sedis
339.
Leptocorticium tenellum Nakasone
Hymenochaetales
Hymenochaetaceae
340.
Fomitiporia atlantica Alves-Silva, Reck & Drechsler-Santos, in Fungal
Diversity 78: xx (2016)
697
698
699
700
701
702
703
704
705
706
707
708
709
710
711
712
713
714
715
716
717
718
719
720
721
722
723
724
725
726
727
728
729
730
731
732
733
734
735
736
737
738
739
740
341.
Fomitiporia subtilissima Alves-Silva, Reck, & Drechsler-Santos, in Fungal
Diversity 78: xx (2016)
342.
Inonotus shoreicola L.W. Zhou, Y.C. Dai & Vlasák, in Fungal Diversity 78:
xx (2016)
Polyporales
Ganodermataceae
343.
Ganoderma wuzhishanensis T.C. Wen, K. Hapuarachchi & K.D. Hyde, in
Fungal Diversity 78: xx (2016)
Polyporales genus, incertae sedis
344.
Dentocorticium ussuricum (Parmasto) M.J. Larsen & Gilb.
Polyporaceae
345.
Lentinus stuppeus Klotzsch
Russulales
Bondarzewiaceae
346.
Bondarzewia tibetica B.K. Cui, J. Song & Jia J. Chen, in Fungal Diversity
X78: XX (2016)
Russulaceae
347.
Lactifluus armeniacus De Crop & Verbeken, in Fungal Diversity 78: xx
(2016)
348.
Lactifluus ramipilosus Verbeken & De Crop, in Fungal Diversity 78: xx
(2016)
349.
Russula amethystina subsp. tengii G.J. Li, H.A. Wen & R.L. Zhao, in
Fungal Diversity 78: xx (2016)
350.
Russula wangii G.J. Li, H.A. Wen & R.L. Zhao, in Fungal Diversity 78: xx
(2016)
Neocallimastigomycota
Neocallimastigomycetes
Neocallimastigales
741
742
743
744
745
746
747
748
749
750
751
752
753
754
755
756
757
758
759
760
761
762
763
764
765
766
767
768
769
770
771
772
773
774
775
776
777
778
779
780
781
782
783
Neocallimastigaceae
351.
Anaeromyces robustus O’Malley, Theodorou & Henske, in Fungal Diversity
78: xx (2016)
352.
Neocallimastix californiae O’Malley, Theodorou & Solomon, in Fungal
Diversity 78: xx (2016)
353.
Piromyces finnis O’Malley, Haitjema & Gilmore, in Fungal Diversity 78: xx
(2016)
Oomycota
Peronosporales
Pythiaceae
354.
Phytophthora estuarina Marano, A.L. Jesus & Pires-Zottar., in Fungal
Diversity 78: xx (2016)
355.
Phytophthora rhizophorae Pires-Zottar., A.L. Jesus & Marano, in Fungal
Diversity 78: xx (2016)
Oomycota, incertae sedis
356.
Salispina Marano, A.L. Jesus & Pires-Zottar., in Fungal Diversity 78: xx
(2016)
357.
Salispina intermedia A.L. Jesus, Pires-Zottar. & Marano, in Fungal
Diversity 78: xx (2016)
358.
Salispina lobata (Fell & Master) A.L. Jesus, Marano & Pires-Zottar. , in
Fungal Diversity 78: xx (2016)
359.
Salispina spinosa (Fell & Master) Marano, A.L. Jesus & Pires-Zottar. , in
Fungal Diversity 78: xx (2016)
Zygomycota
Mucorales
Mortierellaceae
360.
Mortierella calciphila Wrzosek, in Fungal Diversity 78: xx (2016)
Cunninghamellaceae
361.
Absidia stercoraria Hyang B. Lee, H.S. Lee & T.T.T. Nguyen, in Fungal
Diversity 78: xx (2016)
784
785
786
787
788
789
790
791
792
793
794
795
796
797
798
799
800
801
802
803
804
805
806
807
808
809
810
811
812
813
814
815
816
817
818
819
820
821
822
823
824
825
826
827
362.
Gongronella orasabula Hyang B. Lee, K. Voigt, P.M. Kirk & T.T.T.
Nguyen, in Fungal Diversity 78: xx (2016)
Mucoraceae
363.
Mucor caatinguensis A.L. Santiago, C.A. de Souza & D.X. Lima, in Fungal
Diversity 78: xx (2016)
364.
Mucor koreanus Hyang B. Lee, S.J. Jeon & T.T.T. Nguyen, in Fungal
Diversity 78: xx (2016)
365.
Mucor merdicola C.A. de Souza & A.L. Santiago, in Fungal Diversity 78: xx
(2016)
366.
Rhizopus koreanus Hyang B. Lee & T.T.T. Nguyen, in Fungal Diversity 78:
xx (2016)
Introduction
This is the third paper in a series of complied notes on new fungal taxa, reference
specimens, new data, and other taxonomic changes.
Materials and methods
Specimens and cultures were photographed under daylight in the field or lamplight in
the laboratory. Macro- and microscopic characteristics were measured and recorded.
Codes of colours are cited from those of Kornerup and Wanscher (1978), Maerz and
Paul (1950), Ridgway (1912) and Seguy (1936). Fungal Names/Index
Fungorum/MycoBank accession numbers and Facesoffungi numbers were obtained as
detailed in Fungal Names (2016), Index Fungorum (2016), MycoBank (2016), and
Jayasiri et al. (2015).
Phylogenetic analyses were carried out based on holotypes, ex-types, and
sequence data available from GenBank. Genomic DNA samples were extracted from
growing mycelium, ascomata or basidiomata. Genetic markers applied for each genus
and family were based on historic references and have commonly been used in
corresponding families and genera. Multiple sequences were aligned in ClustalX v.
2.1 (Larkin et al. 2007), Maffet v. 7.215 (http://mafft.cbrc.jp/alignment/software/) or
Bioedit 7.0 (Hall 2004). The alignments were reviewed visually and adjusted
manually where necessary. All introns were deleted or aligned separately. Leading or
trailing regions containing many gaps were removed from the alignments prior to tree
building. Different single alignments were linked in needed of multi-gene backbone
tree constructions. The phylogenetic analyses were carried out for maximum
parsimony in PAUP v. 4.0b10 (Swofford 2002), maximum likelihood in RAxML v.
7.2.7 -HPC2, RAxML 7.4.2 Black Box (Stamatakis 2006; Stamatakis et al. 2008),
RAxML GUI (Stamatakis 2006; Silvestro and Michalak 2011), or PhyML 3.0
(Guindon et al. 2010), and Bayesian inferences in MrBayes v. 3.2 (Ronquist et al.
2012) as indicated in the legend of each phylogenetic tree. Data of the newly
828
829
830
generated sequences are listed in Table 1. The phylogenetic trees were constructed
and analyzed by authors of corresponding new taxa based on the selection of genes in
given publications.
831
Table 1 Newly generated sequence data from this study
Strain/Specimen
Taxon Name
ITS
nrLSU
nrSSU
TEF-1α
ACT
No.
Absidia stercoraria
EML-DG8-1
KU168828
KT921998
KT921996
KT922002
KT922000
Absidia stercoraria
EML-DG8-2
KU168829
KT921999
KT921997
KT922003
KT922001
Agaricus coccyginus
HMAS275416
KU245979
Agaricus coccyginus
HMAS275413
KU245984
Agaricus coccyginus
HMAS275412
KU245981
Agaricus coccyginus
HMAS275420
KU245983
Agaricus coccyginus
HMAS254484
KU245980
Agaricus luteofibrillosus
HMAS 254487
KU245972
Agaricus luteofibrillosus
HMAS275419
KU245978
Agaricus luteofibrillosus
HMAS 254486
KU245977
Agaricus luteofibrillosus
HMAS 275415
KU245973
β-tubulin
CHS
GADPH
RPB1
RPB2
COI
MFLU 15–1415
KT934314
Amanita atrobrunnea
Amanita digitosa
BBH 32154
KT213722
Amanita gleocystidiosa
BBH 31901
KT213717
Amanita gleocystidiosa
BBH 31902
KT213718
Amanita gleocystidiosa
BBH 31903
KT213719
Amanita pyriformis
BBH 38643
KT213723
Amanita strobilipes
MFLU 12–2246
Anaeromyces robustus
S4
Angustospora nilensis
MFLU 15–1511
KT944072
Aniptodera aquibella
MFLU 15–1140
KU556854 KU556853
Annulohypoxylon albidiscum
MFLU 15–3883
Annulohypoxylon annulatum
CBS 140775
KU159523
Annulohypoxylon moriforme
STMA 14065
KU159525
Annulohypoxylon nitens
MFLUCC 14-1232
KU159521
KT934313
KU057354
KT944071
KU852741
Annulohypoxylon stygium var.
annulatum
STMA 14066
KU159526
Annulohypoxylon truncatum
CBS 140777
KU159524
Aposphaeria corallinolutea
MFLU 15–3203
Astrocystis thailandica
MFLU 15–3525
KU246224
KU246225
Bondarzewia tibetica
BJFC Cui 12078
KT693202
KT693204
Bondarzewia tibetica
BJFC Yu 56
KT603203
KT693205
Camporesia sambuci
MFLU 15–3905
KU746392
KU746394 KU746396
Clarkeinda trachodes
MFLU10–0139
HM897839
Clematidis italica
MFLU 14–0669
KU842380
Colletotrichum menispermi
MFLU 14–0625
KU242357
KU242353 KU242354 KU242355 KU242356
Colletotrichum quinquefoliae
MFLU 14–0626
KU236391
KU236389 KU236392
Comoclathris pimpinellae
MFLU 15–0010
*****
Cortinarius albosericeus
K(M):200657
KU041721
KU243051 KU243052
KU243050
KU746390
KU842381 KU842382
KU236390
Cortinarius albosericeus
K(M):200658
KU041722
Cortinarius badioflavidus
K(M): 200672
KU041723
Cortinarius badioflavidus
DBB28196
KU041724
Cortinarius badioflavidus
DBB13504
KU041725
Cortinarius badioflavidus
K(M) 200689
KU041726
Cortinarius badioflavidus
K(M): 200673
KU041727
Cortinarius badioflavidus
K(M): 200686
KU041728
Cortinarius badioflavidus
01MWB032411
KU041729
Cortinarius badioflavidus
03MWB120308
KU041730
Cortinarius badioflavidus
K(M): 200690
KU041731
Cortinarius badioflavidus
02MWB043009
KU041732
Cortinarius badioflavidus
K(M): 200688
KU041733
Cortinarius denigratus
K(M): 200659
KU041734
Cortinarius duboisensis
K(M): 200660
KU041735
Cortinarius duboisensis
K(M): 200661
KU041736
Cortinarius duboisensis
K(M): 200662
KU041737
Cortinarius duboisensis
K(M): 200663
KU041738
Cortinarius fragrantissimus
K(M): 200664
KU041739
Cortinarius roseobasilis
K(M): 200665
KU041740
Cortinarius roseobasilis
K(M): 200666
KU041741
Cortinarius vinaceobrunneus
K(M): 200667
KU041742
Cortinarius vinaceogrisescens
K(M): 200668
KU041743
Cortinarius vinaceogrisescens
K(M): 200669
KU041744
Cortinarius wahkiacus
K(M): 200670
KU041745
Cortinarius wahkiacus
K(M): 200671
KU041746
Crassiparies quadrisporus
HHUF30409
LC100020
Creosphaeria sassafras
STMA 14088
Cryptovalsa ampelina
MFLU 16–0007
LC100025
LC100017
KU159533
KU550094
KU550096 KU550095
Cyanoboletus hymenoglutinosus
DC 14-010
KT907355
Cytospora salicicola
MFLU 14–0785
Dentocorticium ussuricum
TAA 42424
KU183718
Diatrype thailandica
MFLU 15–3662
KU315392
Dothiorella rhamni
MFLU 15–3541
KU246381
Dothiorella vidmadera
MFLU 15–3483
KU234792
Durotheca macrostroma
BBH39917
KT883901 KT883903
Durotheca macrostroma
BCC78380
KT883902 KT883904
Eutiarosporella dactylidis
MFLU 15–3502
Farasanispora avicenniae
MFLU
Fomitiporia atlantica
FLOR 58554
KU557528
Fomitiporia atlantica
FURB 47591
KU557529
Fomitiporia subtilissima
FURB 47557
KU557531
Fomitiporia subtilissima
FURB 47437
KU557530
KU246378
KT860060
KU246382
KU234806
KU246380
KT950962
KT950961
KU557526
KU557527
KU557532
KU557533
KU557534
Galzinia longibasidia
GB NH2417
KU183721
KU183722
Ganoderma wuzhishanensis
GZUH14081638
Gongronella orasabula
EML-QF12-1
KT936269
KT936263
KT936261
KT936267
KT936265
Gongronella orasabula
EML-QF12-2
KT936270
KT936264
KT936262
KT936268
KT936266
Halorosellinia rhizophorae
MFLU 15–0183
KU516688
KU516689 KU516690
Humicola koreana
EML-UD33-1
KU058192
KU058190
Humicola koreana
EML-UD33-2
KU058193
KU058191
Hypoxylon flavoargillaceum
STMA 14062
KU159532
Hypoxylon griseobrunneum
STMA 14052
KU159535
Hypoxylon haematostroma
STMA 14043
KU159527
Hypoxylon investiens
STMA 14058
KU159528
Hypoxylon lienhwacheense
MFLUCC 14-1231
KU159522
Hypoxylon lilloi
STMA 14142
KU159537
Hypoxylon lilloi
STMA 14143
KU159538
Hypoxylon lividipigmentum
STMA 14044
KU159529
Hypoxylon monticulosum
STMA14080
KU159534
Hypoxylon perforatum
STMA 14051
KU159531
Hypoxylon polyporus
STMA 14090
KU159530
Hypoxylon umbilicatum
STMA 15276
KU159536
Inonotus shoreicola
IFP LWZ 20140728-10
KT749418
Inonotus shoreicola
IFP LWZ 20140728-23
KT749419
Inonotus shoreicola
BJFC Dai13615
KT749417
IFP
LWZ
Inonotus shoreicola
20140729-1
KT749420
Inonotus shoreicola
BJFC Dai13614
KT749416
Kirschsteiniothelia tectonae
MFLU 15–1883
KU144916
KU764707
Kirschsteiniothelia tectonae
MFLU 15–1884
KU144924
KU764708
Lactifluus armeniacus
EDC 14-501
KR364127
Lactifluus ramipilosus
EDC 14-503
KR364128
Leccinellum indoaurantiacum
DC 14-019
KT907354
Lentinus stuppeus
MFLU10-0145
HM897840
Leptocorticium tenellum
GB NH16311
KU183719
KU183720
Lindgomyces okinawaensis
HHUF30498
LC100022
LC100027
Longiostiolum tectonae
MFLU 15–3532
KU712447
KU764700 KU712459
KU872759
LC100021
LC100026
LC100030
Lophiostoma pseudoarmatisporum HHUF 30497
KT860059
LC100019
LC100018
Moelleriella phukhiaoensis
BCC19769
KT880502
KT880506
Moelleriella phukhiaoensis
BCC19773
KT880503
KT880507
Moelleriella pongdueatensis
BCC31787
KT880500
KT880504
Moelleriella pongdueatensis
BCC31788
KT880501
KT880505
Mortierella calciphila
WA18944
KT964845
KT964846
Mucoharknessia anthoxanthi
MFLU 15–3477
KU246377
KU246379
Mucor caatinguensis
URM 7322
KT960377
KT960369
KT964847
Mucor caatinguensis
URM 7322
KT960376
KT960370
Mucor caatinguensis
URM 7322
KT960375
KT96037
Mucor koreanus
EML-QT1
KT936259
KT936253
KT936251
KT936257
KT936255
Mucor koreanus
EML-QT2
KT936260
KT936254
KT936252
KT936258
KT936256
Mucor merdicola
URM 7223
KT960373
Mucor merdicola
URM 7223
KT960374
KT960372
Multilocularia bambusae
MFLU 11–0216
KU693446
KU693438 KU693442
KU705656
Multiseptospora thysanolaenae
MFLU 11–0238
KU693439 KU693443
KU705658
Musumecia alpina
MHKMU 182
KR909102
KR909099
KR909096
Musumecia alpina
MHKMU 346
KR909100
KR909097
Musumecia alpina
MHKMU 347
KR909101
KR909098
Musumecia sardoa
AMB17139
KT122794
KT122795
Neocallimastix californiae
G1
KU057353
Neophaeocryptopus cytisi
MFLU 15–3542
KU248848
KU248849 KU248850
KR909095
Ochronectria thailandica
MFLU 16–0030
Ophiocordyceps formosana
MFLU 15–3888
Ophiocordyceps formosana
MFLU 15–3889
Ophiocordyceps karstii
KU564071
KU564069 KU564070
KU854949
KU854947
KU854951
KU854950
KU854948
MFLU 15–3884
KU854952
KU854945
KU854943
Ophiocordyceps karstii
MFLU 15–3885
KU854953
KU854946
KU854944
Parameliola acaciae
MFLU 15–0378
KU285142
Parameliola dimocarpi
MFLU 15–0045
KU285143
Parastagonospora cumpignensis
MFLU 15–1480
KU842388
Petrakia echinata
MFLU 15–7568
KU746391
KU746393 KU746395
Phragmoporthe conformis
MFLU 15–2662
KU315388
KU315389 KU315390
Phytophthora estuarina
CCIBt 4157
KT886034
KT886030
KT886051
Phytophthora estuarina
CCIBt 4116
KT886033
KT886029
KT886050
Phytophthora rhizophorae
CCIBt 4152
KT886031
KT886028
KT886048
Phytophthora rhizophorae
CCIBt 4121
KT886032
KU842389 KU842390
KU315391
KT886049
KU057352
Piromyces finnis
Polyplosphaeria thailandica
MFLU 15–3273
KU248766
KU248767
Pseudocamarosporium pini
MFLU 15−3290
KU764779
KU754540 KU754542
Pseudodidymosphaeria phlei
MFLU 15–3281
KU764780
KU754541 KU754543
Pseudolachnella brevifusiformis
HHUF 30495
LC100023
LC100028
Pseudolachnella brevifusiformis
HHUF 30496
LC100024
LC100029
Psiloglonium macrosporum
MFLU 14–0610
KU243048
KU243049
Rhizopus koreanus
EML-HO95-1
KU058202
KU058196 KU058194
KU058200
KU058198
Rhizopus koreanus
EML-HO95-2
KU058203
KU058197 KU058195
KU058201
KU058199
Rosellinia chiangmaiensis
MFLU 15–3524
KU246226
KU246227
Russula amethystina subsp. tengii HMAS253336
KT949399
Russula amethystina subsp. tengii HMAS271033
KT949400
Russula amethystina subsp. tengii HMAS253216
KT949401
Russula amethystina subsp. tengii HMAS253241
KT949402
Russula wangii
HMAS268809
KF851403
Russula wangii
HMAS269106
KT949396
Russula wangii
HMAS269308
KT949397
Russula wangii
HMAS269580
KT949398
Saccotheciumubi
MFLU 15–3400
KU290338
Salispina intermedia
CCIBt 4155
Salispina intermedia
CCIBt 4115
Salispina intermedia
CCIBt 4153
Salispina intermedia
CCIBt 4154
Salispina intermedia
CCIBt 4156
Salispina lobata
CBS 588.85
Salispina spinosa
CBS 591.85
Seimatosporium brunium
MFLU 14–0772
Seimatosporium pseudocornii
MFLU 13−0529
KU290336 KU290337
KT920432
KT886044
KT886053
KT886055
KT920431
KT886042
KT886052
KT886043
KT920433
KT920434
KT886045
KT886054
KT886046
KT886056
KT886047
KU359033 KU359034
Seimatosporium pseudorosae
MFLU 14−0468
Sigarispora ononidis
MFLU 15–2667
Stagonospora russa
MFLU 15–0012
Towyspora aestuari
MFLU 15–3543
KU359035
KU243128
KU243125 KU243126
KU248851
KU248852
KU243127
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Results and discussion
The new taxa are described and illustrated in alphabetical order as following. A total
of 74 genera in 44 families, 21 orders and five classes in the Ascomycota,
Basidiomycota, Oomycota, Neocallimastigomycota and Zygomycota are introduced.
Contributions to Ascomycota
Dothideomycetes
We follow Hyde et al. (2013) and Wijayawardene et al. (2014) for classification of
Dothideomycetes.
Botryosphaeriales
Members of the order Botryosphaeriales are commonly encountered as
endophytes or pathogens of various plant hosts and comprise six ecologically diverse
families; Aplosporellaceae, Botryosphaeriaceae, Melanopsaceae, Saccharataceae,
Phyllostictaceae and Planistromellaceae (Liu et al. 2012; Slippers et al. 2013).
Botryosphaeriaceae
The family Botryosphaeriaceae is found in all geographical and climatic areas of
the world, encompassing a range of morphologically diverse fungi that are either
pathogens, endophytes or saprobes (Phillips et al. 2013). Considerable interest in
Botryosphaeriaceae has arisen due to their association with plant diseases (Yan et al.
2013; Pitt et al. 2013b; Linaldeddu et al. 2015). The phylogenetic tree for
Botryosphaeriaceae is presented in Fig. 1.
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Fig. 1 Phylogram generated from Maximum Parsimony analysis based on combined ITS and
LSU sequence data of species of Botryosphaeriaceae. Parsimony bootstrap support values for
MP≥70 % are shown above the nodes and Bayesian posterior probabilities ≥95 % are
indicated in bold branches. The tree is rooted with Saccharata proteae. All ex-types and
reference strains are in bold and new isolates are in blue.
Dothiorella Sacc.
Based on morphology and molecular data, Phillips et al. (2005) revived
Dothiorella for species with conidia that become brown and 1-septate, while they are
still attached to the conidiogenous cells. Sexual morphs of Dothiorella have
pigmented, 1-septate ascospores (Phillips et al. 2005, 2013). With the exception of D.
sarmentorum and D. iberica, the sexual morphs of Dothiorella species are
infrequently found in nature and have never been reported in culture (Phillips et al.
2013). Although there are 350 species records in Dothiorella, Phillips et al. (2013)
revealed that cultures are available for only 17 species and of those four species have
yet to be named. Abdollahzadeh et al. (2014) introduced three species names for these
un-named taxa. Presently, 25 species are accepted in the genus (Abdollahzadeh et al.
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2014; Crous et al. 2015a; Li et al. 2014; Phillips et al. 2013; Pitt et al. 2013b, 2015;
Slippers et al. 2014). All species, except D. sarmentorum, have been introduced since
2005. A phylogenetic tree for Dothiorella is presented in Fig. 2.
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Fig. 2 Phylogram generated from Maximum Parsimony analysis based on combined ITS and
EF sequence data for species of Dothiorella. Parsimony bootstrap support values for
MP≥75 % and Bayesian posterior probabilities ≥0.9 % are shown above the nodes. The tree is
rooted with Spencermartinsia viticola CBS 117009. All ex-types and reference strains are in
bold and new isolates are in blue.
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253. Dothiorella rhamni Wanasinghe, Bulgakov, E.B.G. Jones & K.D. Hyde, sp. nov.
Index Fungorum number: IF 551784, Facesoffungi number: FoF 01668, Fig. 3
Etymology: Name reflects the host genus Rhamnus, from which the species was
isolated.
Holotype: MFLU 15–3541
Saprobic or weak pathogen on dead twigs of Rhamnus cathartica L. Sexual
morph Undetermined. Asexual morph Conidiomata 420–460 µm high × 590–660
µm diam. ( x = 443.3 × 623.2 µm, n = 10), pycnidial, stromatic, mostly solitary,
semi-immersed to immersed in the host, dark brown to black, ostiolate, apapillate.
Peridium 50–60 µm wide at the base, 70–90 µm wide in sides, comprising 8–10 layers,
heavily pigmented, thick-walled, comprising blackish to dark brown, angular cells,
becoming flattened towards the outer layers. Conidiogenous cells 8–12 µm high × 4–6
µm wide, holoblastic, cylindrical to subcylindrical, hyaline, the first conidium
produced holoblastically and subsequent conidia enteroblastically, forming typical
phialides with periclinal thickenings, swollen at the base, discrete, producing a single
conidium at the apex. Conidia 17–24 × 9–12 µm ( x = 20.7 × 10.4 µm, n = 50),
initially hyaline, unicellular, becoming cinnamon to sepia and 1-septate, while still
attached to conidiogenous cells; detached conidia, hyaline, sepia or dark brown,
unicellular or 1-septate, moderately thick-walled, wall externally smooth, roughened
on the inner surface, oval to ovoid, widest in the center, apex obtuse, base truncate or
rounded.
Material examined: RUSSIA, Rostov region, Oktyabrsky District, near natural
sanctuary «Persianovskaya steppe», Khoruli hollow, ravine grove (47.5006484° E,
40.1385927° N), on Rhamnus cathartica (Rhamnaceae), 26 April 2014, T.S.
Bulgakov (MFLU 15–3541, holotype); ex-type culture, MFLUCC 14–0902.
Notes: The genus Dothiorella was established by Saccardo (1880) to
accommodate D. pyrenophora (Berk.) ex Sacc., and is characterized by branched,
septate conidiophores, holoblastic conidiogenesis, and smooth to finely verruculose
but not striate, brown, 1-euseptate conidia (Crous and Palm 1999). Phillips et al.
(2005) re-introduced Dothiorella as a distinct Botryosphaeriaceae asexual morph with
brownish conidia, which become septate while still attached to the conidiogenous
cells. Dothiorella rhamni also has sepia to dark brown, 1-septate conidia, similar to
other members in Dothiorella. Phylogenetically D. rhamni clustered in a sister group
with D. sarmentorum (CBS 115038 and IMI 63581b) and Diplodia acerina (CBS
910.73), but D. rhamni separates from them with good statistical support.
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Fig. 3 Dothiorella rhamni (holotype) a Appearance of conidiomata on host substrate b
Vertical section through a conidioma c Close up of ostiole d Peridium of conidioma e, f
Mature and immature conidia attached to conidiogenous cells g Mature and immature conidia
h Germinated conidium. Scale bars: b = 100 µm, c, d = 20 µm, e–h = 10 µm.
254. Dothiorella vidmadera Pitt et al., Fungal Diversity 61: 216, 2013
Facesoffungi number: FoF 01326, Fig. 4
Saprobic on dead branch of Fraxinus ornus L. Sexual morph Ascostromata
320–410 µm diam., dark brown to black, globose, submerged in the substrate,
partially erumpent at maturity, ostiolate. Ostiole circular, central, papillate. Peridium
50–80 µm thick, composed of dark brown thick-walled cells of textura angularis,
becoming thin-walled and hyaline towards the inner region. Pseudoparaphyses 3–5
µm wide, thin-walled, hyaline. Asci 150–220 × 12–22 µm, 8-spored, bitunicate,
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cylindric-clavate, endotunica thick-walled, with a well-developed ocular chamber.
Ascospores 17–26 × 8–10 µm ( x = 22 × 9 µm, n = 20), obliquely uniseriate or
irregularly biseriate, initially hyaline and becoming dark brown, oblong to ovate,
widest in center, straight, 1-septate, constricted at the septum, moderately
thick-walled, surface smooth. Asexual morph Conidiomata 380 µm wide, globose,
pycnidial, stromatic, solitary, composed of dark brown, thick-walled cells of textura
angularis. Conidiogenous cells 8–14 × 3–6 µm, lining the pycnidial cavity,
holoblastic, hyaline, subcylindrical. Conidia 17–22 × 9–10 µm ( x = 21 × 10 µm, n =
20) initially hyaline and aseptate, becoming pigmented brown and 1-septate while
attached to conidiogenous cell, slightly constricted at the septum, ovoid with a
broadly rounded apex and truncate base.
Culture characteristics: Colonies on PDA, covering 20 mm diam. in Petri-dishes
after 30 days in the dark at 25°C; circular, initially white, after 1 week becoming
greyish brown to black; reverse grey to dark greyish green; flattened, fluffy, fairly
dense, aerial, surface smooth with crenate edge, filamentous and conidia produced on
pine needles after 3 weeks at 18°C.
Material examined: ITALY, Province of Forlì-Cesena [FC], Corniolo - Santa
Sofia, on dead branch of Fraxinus ornus (Oleaceae), 6 December 2013, Erio
Camporesi IT 1562 (MFLU 15–3483, reference specimen designated here), ex-type
living cultures MFLUCC 15–0759, KUMCC 15–0129, GZCC 15–0007.
Notes: The sexual morph of Dothiorella vidmadera is morphologically similar to
D. sarmentorum and D. iberica in having globose ascostromata with a central ostiole,
lined with hyaline cells, a wide peridium, bitunicate asci with a thickened endotunica,
and dull brown or dark reddish brown, septate, ellipsoid-obovoid, ascospores,
constricted at the septum. It however differs in spore dimensions and molecular
phylogeny. The asexual morph of Dothiorella vidmadera was observed in culture and
is similar to that described by Pitt et al. (2013b) and differs from the other asexual
morphs of Dothiorella species (Phillips et al. 2013; Abdollahzadeh et al. 2014; Crous
et al. 2015a). Our strains of D. vidmadera (MFLUCC 15–0759) clustered in the
Dothiorella clade with 94% bootstrap support (Fig. 1) and this is the first report of the
sexual morph for Dothiorella other than D. sarmentorum and D. iberica.
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Fig. 4 Dothiorella vidmadera (MFLU 15–3483, reference specimen) a Appearance of
ascostromata on host substrate b Cross section of ascoma c Peridium d–f Immature asci g–i
Mature asci j Mature brown ascospore k, l Culture on PDA m Immature and mature conidia
attached to conidiogenous cells n Immature hyaline conidia o Mature conidia. Scale bars: b, c
= 100 µm, d–g = 30 µm, h–j = 20 µm, k, l = 1 cm, m–o = 20 µm.
Eutiarosporella Crous
This genus was introduced by Crous et al. (2015b) named because of its
similarity to the genus Tiarosporella, and is distinguished from Tiarosporella by
having conidiomata with long necks, and holoblastic conidiogenesis. Tiarosporella
was introduced by Höhnel (1919), and is considered as an asexual genus in
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Botryosphaeriaceae (Jami et al. 2012; Phillips et al. 2013; Slippers et al. 2013) and
mainly occurs on grasses, conifers and members of Asteraceae and Zygophyllaceae
(Karadžić 2003; Jami et al. 2012). Thambugala et al. (2014a) introduced the sexual
morph of Tiarosporella, T. dactylidis and detailed descriptions and illustrations were
provided. The sexual morph of T. dactylidis which is illustrated here, is
morphologically similar to Botryosphaeria in having globose ascomata, with a central
ostiole, a two layered peridium, hyphae-like pseudoparaphyses and hyaline, aseptate,
fusoid to ovoid ascospores, with a mucilaginous sheath (Thambugala et al. 2014a).
Crous et al. (2015b) described Eutiarosporella tritici (B. Sutton & Marasas) as the
type species of the genus. Species of Eutiarosporella have been reported from Celtis
Africana N.L. Burm (Rosales), Triticum aestivum L. (Poales), Acacia karroo Hayne
(Fabales) and Dactylis glomerata L. (Poales) (Thambugala et al. 2014a; Crous et al.
2015b). Here we report the sexual morph of Eutiarosporella for the first time from
Avenella flexuosa L. (Poales).
255. Eutiarosporella dactylidis (K.M. Thambugala, E. Camporesi & K.D. Hyde)
Dissanayake, Camporesi & K.D. Hyde, comb. nov.
Basionym: Tiarosporella dactylidis Thambugala, E. Camporesi & K.D. Hyde,
Cryptog. Mycol.35: 359–367 (2014).
Index Fungorum number: IF 551751, Facesoffungi number: FoF 01650, Fig. 5
Saprobic on stem of grasses (Avenella sp.). Sexual morph Ascostromata
150–195 µm high × 175–240 µm diam., visible as black spots on host tissue,
uniloculate, scattered or gregarious, globose to subglobose, ostiolate. Ostiole circular,
central, papillate. Peridium up to 25–45 µm wide, comprising 2 layers: outer layer of
thin, small, brown to dark brown cells of textura angularis, inner layer of thick, large,
hyaline to lightly pigmented, cells of textura angularis. Hamathecium comprising 2–3
µm wide, hyphae-like, hyaline, sparse pseudoparaphyses. Asci 120–180 × 15–23 µm
( x = 145 × 19 µm, n = 30), 8-spored, bitunicate, fissitunicate, clavate to
cylindric-clavate, pedicellate, apically rounded, with an ocular chamber. Ascospores
22–28 × 7–8.5 µm ( x = 25 × 8 µm, n = 30), uni to bi-seriate in the upper half,
uniseriate at the base, hyaline, becoming olivaceous-brown at maturity, aseptate,
ellipsoidal to fusiform, usually wider in the center, thick-walled, smooth-walled,
surrounded by a mucilaginous sheath. Asexual morph see asexual morph description
in Thambugala et al. (2014a).
Material examined: ITALY, Province of Forlì-Cesena [FC], Montebello Modigliana, on dead stem of Avenella flexuosa L. (Poaceae), 24 November 2014,
Erio Camporesi IT 2251 (MFLU 15–3502), living cultures MFLUCC 15–0915.
Notes: The genus Tiarosporella was introduced by Höhnel (1919) and is
considered as an asexual genus in the family Botryosphaeriaceae. Thambugala et al.
(2014a) introduced a sexual morph for the genus Tiarosporella; T. dactylidis
Thambugala et al., based on the multi-gene phylogeny. Since the type species of
Tiarosporella; T. paludosa (Sacc. & Fiori ex P. Syd.) Höhn clusters in a distinct clade
in Botryosphaeriaceae apart from the species accommodated in Tiarosporella; Crous
et al. (2015b) introduced a new genus Eutiarosporella to accommodate
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tiarosporella-like taxa, based on E. tritici (B. Sutton &Marasas) on Triticum aestivum
L. from South Africa. The genus comprises three species, Eutiarosporella africana
Jami et al., E. tricti and E. urbis-rosarum Jami et al. Based on the multigene
phylogenetic analysis (Fig. 1), the ex-type strain of Tiarosporella dactylidis
(MFLUCC 13–0276) clusters with other species in Eutiarosporella. With the species
combined in Eutiarosporella here, the number of species in this genus increases to
four.
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Fig. 5 Eutiarosporella dactylidis sexual morph (MFLU 15–3502) a Appearance of
ascostromata on host surface b Section through ascostroma c, d Immature asci e-g Mature
bitunicate asci h, i Ascospores with inconspicuous mucilaginous sheath. Scale bars: b = 100
µm, c, d = 50 µm, e, f = 40 µm, g–i = 20 µm.
Mucoharknessia Crous, R.M. Sánchez & Bianchin.
The genus Mucoharknessia was introduced by Crous et al. (2015b) for a genus
resembling Harknessiaceae, in Diaporthales. Mucoharknessia can be distinguished
from Harknessiaceae in having pycnidia that lack furfuraceous tissue surrounding its
ostiole, and conidia that have a mucoid apical appendage. The type species is
Mucoharknessia cortaderiae (Crous et al. 2015b).
256. Mucoharknessia anthoxanthi Dissanayake, Camporesi & K.D. Hyde, sp. nov.
Index Fungorum number: IF 551752, Facesoffungi number: FoF 01651, Fig. 6
Etymology: Referring to the host Anthoxanthum odoratum L.
Holotype: MFLU 15–3477
Saprobic on dead stems of Anthoxanthum odoratum. Sexual morph
Undetermined. Asexual morph Conidiomata 240–320 µm high × 215–280 µm diam.,
globose, immersed to erumpent, brown, wall of 3–6 layers of brown textura angularis.
Conidiophores reduced to conidiogenous cells. Conidiogenous cells 7–14 × 4–5 µm
lining the inner cavity, hyaline, smooth, ampulliform to subcylindrical, proliferating
percurrently at the apex. Paraphyses 25–45 × 3–4 µm intermingled among
conidiogenous cells, hyaline to pale brown, smooth to verruculose, subcylindrical
with obtuse ends. Conidia 18–30 × 8–10 µm ( x = 24 × 9 µm, n = 30), solitary,
hyaline, smooth-walled, guttulate, fusoid-ellipsoid to subcylindrical, straight to curved,
apex apiculate, tapering at base, apex with flared mucoid appendage, up to 20 µm
long, 15 µm diam.
Material examined ITALY. Province of Forlì-Cesena [FC], Passo delle Forche Galeata, on dead stem of Anthoxanthum odoratum (Poaceae), 24 November 2012,
Erio Camporesi IT 981 (MFLU 15–3477), ex-type living cultures MFLUCC 15–0904,
CGMCC.
Notes: The genus Mucoharknessia was introduced in Botryosphaeriaceae by
Crous et al. (2015b) based on Cortaderia selloana L. from Argentina. Based on
multi-gene phylogenetic analyses (Fig. 1), our isolate clustered close to M.
cortaderiae. In this paper we introduce a new species, Mucoharknessia anthoxanthi
based on its distinct morphological characters.
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Fig. 6 Mucoharknessia anthoxanthi (holotype) a Appearance of conidiomata on host surface
b, c Sections through conidiomata d–f Immature conidia attached to conidiogenous cells g
Conidia with mucoid appendage h–l Conidia with mucoid appendage stained in Indian ink.
Scale bars: b, c = 100 µm, d = 50 µm, e–l = 25 µm.
Dothideales
The order Dothideales was introduced by Lindau (1897) to accommodate a
single family Dothideaceae Chevall. Subsequently, Theissen and Sydow (1917)
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introduced Dothioraceae Theiss. & Syd. in Dothideales. Dothideales however, has a
rather varied past as various authors treated this order with a number of different
families (Thambugala et al. 2014b). However, recently Thambugala et al. (2014b)
revised the order Dothideales and synonymized Dothioraceae under Dothideaceae,
and accepting only two families: Dothideaceae and Aureobasidiaceae K.M.
Thambugala & K.D. Hyde. Thambugala et al. (2014b) introduced Aureobasidiaceae
to accommodate Aureobasidium Viala & G. Boyer, Saccothecium and five other
genera, but this family is a homonym of Aureobasidiaceae Cif., which had been
previously introduced (Ciferri 1958). Later Saccotheciaceae Bonord. was proposed
(instead of Aureobasidiaceae) as Saccotheciaceae is the oldest available name for the
family that contains Aureobasidium and Saccothecium (Liu et al. 2015). The
phylogenetic tree for Dothideales is presented in Fig. 7.
Dothideaceae
The family Dothideaceae was introduced by Chevallier (1826) as ‘Dothideae’,
and later Fuckel (1870) established this family with Dothidea as the type genus and D.
gibberulosa (Fr.) Fr. as the type species. Dothideaceae is characterized by ‘immersed
to erumpent or superficial, uni or multi-loculate ascostromata, 8- or polyspored,
bitunicate asci and hyaline or brown, transversely septate, sometimes muriform
ascospores’ (Thambugala et al. 2014). Thambugala et al. (2014) revised the family
and included ten sexual genera (Phaeocryptopus, Sydowia, Pringsheimia,
Delphinella, Plowrightia, Stylodothis, Dictyodothis, Dothidea, Endodothiora and
Dothiora) and five asexual genera (Endoconidioma, Cylindroseptoria,
Neocylindroseptoria, Kabatina and Coleophoma).
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Fig. 7 Phylogram generated from maximum likelihood analysis based on analysis of
combined LSU, SSU and ITS sequence data of species of Dothideales. Maximum likelihood
bootstrap support values greater than 50 % and Bayesian posterior probabilities greater than
0.90 are near the nodes. The ex-type strains are in bold and the new isolates are in blue. The
scale bar indicates 0.03 changes. The tree is rooted with Elsinoe veneta and Elsinoe phaseoli.
257. Neophaeocryptopus Wanasinghe, Camporesi, E.B.G. Jones & K.D. Hyde, gen.
nov.
Index Fungorum number: IF 551785, Facesoffungi number: FoF 01669
Etymology: Named after its morphological similarity to the genus
Phaeocryptopus.
Type species: Neophaeocryptopus cytisi Wanasinghe, Camporesi, E.B.G. Jones
& K.D. Hyde
Saprobic on stems and twigs of herbaceous and woody plants in terrestrial
habitats. Sexual morph Ascostromata superficial, semi-immersed to erumpent,
solitary, scattered, broadly oblong, dark brown to black, coriaceous, uniloculate.
Peridium comprising 5–8 layers, outer part comprising heavily pigmented,
thick-walled, angular cells. Hamathecium lacking pseudoparaphyses. Asci 8-spored,
bitunicate, fissitunicate, clavate to broadly-clavate, short pedicellate, thickened and
rounded at apex, with an ocular chamber. Ascospores overlapping 1–2-seriate,
hyaline, broadly fusiform, rounded at both ends, 1-septate, with a median septum,
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constricted at the septum, smooth-walled, lacking a mucilaginous sheath. Asexual
morph Conidiomata stromatic, immersed in agar to superficial, uni- to multi-loculate,
globose to subglobose, glabrous, ostiole central, with minute papilla. Conidiomata
walls composed of several layers of hyaline to dark brown, pseudoparenchymatous
cells, organized in a textura angularis. Conidiophores arising from basal cavity of
conidiomata, mostly reduced to conidiogenous cells. Conidiogenous cells holoblastic,
phialidic, discrete, ampulliform to cylindric-clavate, hyaline, aseptate, smooth-walled.
Conidia solitary, one-celled, fusiform to falcate, with narrowed ends, initially hyaline,
becoming pale brown at maturity, aseptate, smooth and thin-walled, guttulate,
contents granular.
Notes: Phylogenetic analyses of LSU, SSU and ITS sequence data indicate that
Neophaeocryptopus is a distinct genus in Dothideaceae, which forms a clade sister to
the Coleophoma, Cylindroseptoria and Dothiora clades. Neophaeocryptopus,
however differs from these genera, having uni-loculate ascostromata, while
Coleophoma, Cylindroseptoria and Dothiora having multi-loculate ascostromata.
Neophaeocryptopus is morphologically most closely related to Phaeocryptopus which
has uniloculate ascostromata, cylindrical, clavate asci and partially overlapping,
hyaline, 1-septate ascospores, with rounded ends (Thambugala et al. 2014b).
However, this is not supported by sequence data, as Neophaeocryptopus forms a
remote clade from Phaeocryptopus (Fig. 7). Nevertheless, we could not include the
type species Phaeocryptopus abietis Naumov sequences in the phylogenetic analysis,
since they are not available. The type of Phaeocryptopus needs to be recollected and
sequenced in order to resolve the conformity of Neophaeocryptopus with
Phaeocryptopus in Dothideaceae.
258. Neophaeocryptopus cytisi Wanasinghe, Camporesi, E.B.G. Jones & K.D. Hyde,
sp. nov.
Index Fungorum number: IF 551786, Facesoffungi number: FoF 01670, Fig. 8
Etymology: Named after the host genus on which it occurs, Cytisus.
Holotype: MFLU 15–3542
Saprobic on dead and hanging branches of Cytisus sp. Sexual morph
Ascostromata 180–250 ×170–210 µm (x̅ = 212.8 × 187.1 µm, n = 10), superficial,
semi-immersed to erumpent, solitary, scattered, broadly oblong, dark brown to black,
coriaceous, uniloculate. Peridium 35–45 µm wide at the base, 30–40 µm wide at the
sides, comprising 5–8 layers, outer part heavily pigmented, thick-walled, comprising a
blackish to dark brown, amorphous layer, inner part composed of dark brown,
thick-walled, angular cells, becoming flattened and hyaline inwardly. Hamathecium
lacking pseudoparaphyses. Asci 70–90 × 20–30 µm (x̅ = 81.9 × 25.3 µm, n = 40),
8-spored, bitunicate, fissitunicate, clavate to broadly-clavate, short pedicellate,
thickened and rounded at apex, with an ocular chamber. Ascospores 25–35 × 7–10 µm
(x̅ = 29.1 × 9.2 µm, n = 50), overlapping 1–2-seriate, hyaline, broadly fusiform,
rounded at both ends, 1-septate, with a median septum, constricted at the septum,
smooth-walled, lacking a mucilaginous sheath. Asexual morph Conidiomata
stromatic, immersed in agar to superficial, uni- to multi-loculate, globose to
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subglobose, glabrous, ostiole central, with minute papilla. Conidiomata walls
composed of several layers of hyaline to dark brown, pseudoparenchymatous cells,
organized in a textura angularis. Conidiophores arising from basal cavity of
conidiomata, mostly reduced to conidiogenous cells. Conidiogenous cells holoblastic,
phialidic, discrete, ampulliform to cylindric-clavate, hyaline, aseptate, smooth-walled.
Conidia 25–35 × 6–9 µm (x̅ = 28.3 × 7.3 µm, n = 50), solitary, 1-celled, fusiform to
falcate, with narrowed ends, initially hyaline, becoming pale brown at maturity,
aseptate, smooth and thin-walled, guttulate, contents granular.
Material examined: ITALY, Arezzo Province: Croce di Pratomagno, dead and
hanging branches of Cytisus scoparius (L.) Link (Fabaceae), 30 June 2014, E.
Camporesi (MFLU 15–3542, holotype); ex-type culture, MFLUCC 14–0970, MUCL.
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Fig. 8 Neophaeocryptopus cytisi (holotype) a Appearance of ascostromata on host substrate
b, c Sections of the ascostromata d, e Asci f–i Ascospores j, k Conidiomata produced on PDA
l, m, n Mature and immature conidia attached to conidiogenous cells g Mature and immature
conidia. Scale bars: b, c = 50 µm, d, e = 20 µm, f–i, l = 10 µm, k = 500 µm, m, n = 20 µm.
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Saccotheciaceae Bonord. [as 'Saccotheciei'], Abh. naturforsch. Ges. Halle 8: 82
(1864)
= Aureobasidiaceae Cif., Man. Mic. Med., Edn 2 (Pavia) 1: 178 (1958)
= Aureobasidiaceae Thambugala & K.D. Hyde in Hyde et al., Fungal Diversity
68 (1): 133 (2014), isonym.
Type: Saccothecium Fr., Fl. Scan.: 349 (1836)
Notes: Saccotheciaceae was introduced by Bonorden (1864) in order to
accommodate Saccothecium Fr., while Theissen and Sydow (1917) introduced
Dothioraceae Theiss. & Syd. in Dothideales which was typified by Dothiora Fr.
Doweld (2012) suggested to conserve Dothioraceae against the older
Saccotheciaceae. However, Thambugala et al. (2014b) based on morphology and
molecular phylogeny introduced Aureobasidiaceae K.M. Thambugala & K.D. Hyde
to accommodate Aureobasidium Viala & G. Boyer, Saccothecium and five other
genera. The family Aureobasidiaceae had in fact already been introduced by Ciferri
(1958). However, Aureobasidiaceae should be synonymized under Saccotheciaceae
because the latter is the oldest available name for the family that contains
Aureobasidium and Saccothecium. The phylogenetic tree is presented in Fig. 7.
Saccothecium Fr.
Thambugala et al. (2014b) have discussed this genus with the new placement in the
order Dothideales. They have collected S. sepincola from Italy and directly isolated
DNA from the ascostromata. This collection of S. sepincola from Italy clustered in
Saccotheciaceae in the phylogenetic analysis. Therefore, they assigned Saccothecium
in family Saccotheciaceae.
259. Saccothecium rubi Jayasiri, Wanasinghe, Camporesi & K.D. Hyde, sp. nov.
Index Fungorum Number: IF 551769, Facesoffungi number: FoF 01663, Figs 9,
10
Etymology: In reference to host genus.
Holotype: MFLU 15–3400
Saprobic on dead spines of Rubus ulmifolius Schott. Sexual morph Ascostromata
94–110 µm high, 110–120 µm diam. ( x = 98 × 115 µm, n = 10), black, immersed to
erumpent, solitary or scattered, globose to subglobose, usually uniloculate, rarely
biloculate without a distinct ostiole. Peridium 20–30 µm ( x = 23 µm, n = 15) wide, a
single layer, composed of brown to inner hyaline cells of textura angularis, near the
base connected to the host tissue. Hamathecium lacking pseudoparaphyses. Asci
47–62 × 12–16 µm ( x = 50 × 15 µm, n = 20), 8-spored, bitunicate, saccate to broadly
clavate or cylindric-, with a short bifurcate pedicel and a distinct ocular chamber.
Ascospores 14–18 × 4–5 µm ( x = 16 × 4.5 µm, n = 25), overlapping biseriate,
hyaline, 3-septate, lacking vertical septate, asymmetric, obovoid, fusiform to clavate,
with broadly to narrowly rounded ends, with broad upper cells, smooth-walled.
Asexual morph Conidiomata acervular to sporodochial, amphigenous, substomatal,
subepidermal, pulvinate, dry or crystaline in appearance, dark brown to black,
discrete. Conidiogenous cells on hyaline hyphae, lateral, terminal or intercalary,
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cylindrical, clavate or globose, integrated, terminal, with holoblastic, polyblastic
conidiogenesis, with numerous synchronously produced conidia. Conidia blastic,
hyaline, smooth-walled, aseptate, straight, ellipsoidal to sphaerical, reniform to
sickle-shaped, sometimes cylindrical with obtuse ends and occasionally with a slightly
truncate base, rather variable in shape and size.
Material examined: ITALY, Province of Forlì-Cesena [FC], near Poderone –
Corniolo - Santa Sofia, on dead spines of Rubus ulmifolius (Rosaceae), 3 October
2014, Erio Camporesi IT 2136 (MFLU 15–3400, holotype), Ibid., (isotype in KUN);
ex-type living culture (MFLUCC 14–1171, KUNCC).
Culture characteristics: Colonies on MEA at 18 ºC attaining about 70–80 mm
diam. after 14 days, appearing smooth and slimy due to abundant sporulation, pinkish
white. Within first 6 weeks’ colonies filamentous and thereafter develop white,
setae-like mycelia, then turning to brown and then black at the irregular margin.
Notes: In this study we have collected a new species of this genus from Italy,
with different ascospore and ascus morphology, which also separates in the
phylogenetic tree. Wehmeyer (1957) and Holm (1957) proposed to lectotypify the
genus with Saccothecium sepincola. Saccothecium has been assigned to
Dothideaceae, Dothideales (Barr 1972, 1987 and 2001; Kirk et al. 2008; Lumbsch
and Huhndorf 2010; Thambugala et al. 2014b). In this study, we could obtain the
asexual morph of this species, which is similar to Aureobasidium pullulans (de Bary)
G. Arnaud var. (type species of genus Aureobasidium). Hence we can confirm
placement of Saccothecium with in family Saccotheciaceae. This is the first record of
species from host Rubus ulmifolius in the family Saccotheciaceae.
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Fig. 9 Saccothecium rubi (holotype) a, b Appearance of immersed ascostromata on the host
surface c, d Section through ascostromata e Arrangement of asci in ascostromata f–h Asci j–n
Spores o Germinating ascospore. Scale bar: c, d = 30 µm, e = 50 µm, f–i = 20 µm, j–o = 5 µm.
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Fig. 10 Saccothecium rubi asexual morph from the culture (ex-type) a, b Culture on MEA
incubated for 2 weeks, a from above, b from below c Asexual structures in the MEA d–g
Conidiophores and conidiogenesis h, i Conidia. Scale bars: a, b = 3 cm, c = 200 µm, d–i = 10
µm.
Subclass Pleosporomycetidae
Hysteriales
Hysteriaceae
The family Hysteriaceae was introduced by Chevallier (1826) and is
characterized by carbonaceous, immersed to erumpent to entirely superficial
hysterothecia, distinctly navicular in outline, bearing a pronounced longitudinal slit
running the length of the long axis and hyaline to pigmented, 1-multi-septate or
muriform ascospores (Boehm et al., 2009 a, b; Hyde et al., 2013; de Almeida et al.,
2014; Thambugala et al. 2016). Hyde et al. (2013) and Wijayawardene et al. (2014b)
accepted 13 genera including Actidiographium, Coniosporium, Gloniella, Gloniopsis,
Hysterium, Hysterobrevium, Hysterocarina, Hysteropycnis, Oedohysterium,
Ostreichnion, Psiloglonium, Rhytidhysteron and Sphaeronaema in the family, while
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de Almeida et al., (2014) introduced a new genus Hysterodifractum. The phylogenetic
tree is presented in Fig. 11.
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Fig. 11 Phylogram generated from Maximum Likelihood (RAxML) analysis based on LSU
sequence data of Hysteriaceae. Maximum likelihood bootstrap support values equal or greater
than 50 % are indicated above and below the nodes. New taxa are in blue and sequences
based on type material have names in bold. The tree is rooted with Delitschia winteri.
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Psiloglonium Höhn.
Psiloglonium was introduced by von Höhnel (1918) and Petrak (1923a)
designated P. lineare (Fr.) Petr as the type species. Zogg (1962) synonymised
Psiloglonium species which were introduced by von Höhnel (1918) and Petrak (1923
a, b) under the genus Glonium. von Arx & Müller (1975) reduced the genus
Psiloglonium to a synonym of Glonium. However, Boehm et al. (2009a)
re-established Psiloglonium within the Hysteriaceae, to accommodate non-subiculate
species with apically obtuse didymospores. Boehm et al. (2009b) introduced eight
new combinations for the genus Psiloglonium, to accommodate species previously
classified under the genus Glonium in Gloniaceae. Liu et al. (2015) introduced a new
Psiloglonium species, P. multi-septatum Phookamsak & K.D. Hyde, based on
morphological traits and phylogenetic placement and currently there are 19 epithets
listed in Index Fungorum (2016).
260. Psiloglonium macrosporum Thambugala, Senan. & K.D. Hyde, sp. nov.
Index Fungorum number: IF 551806, Facesoffungi number: FoF 01774, Fig. 12
Etymology: Referring to its relatively large ascospores
Holotype: MFLU 14–0610
Saprobic on decaying wood. Sexual morph Ascomata 600–1400 µm long ×
275–475 µm wide × 270–415 µm high ( x = 921 × 348 × 327 µm, n = 6),
hysterothecial, scattered, superficial, base immersed in the substrate, elongate and
depressed conchate, globose, surface black, shiny, longitudinally striate, apex
compressed, opening by a longitudinal slit. Peridium 30–60 µm ( x = 42, n = 15) wide,
carbonaceous, brittle, comprising heavily pigmented, small, prosenchymatous cells.
Hamathecium comprising 0.5–1 µm wide, hyaline, aseptate, branched, trabeculate
pseudoparaphyses, in a gelatinous matrix. Asci 168–215 × 50–60 µm ( x = 187 × 55
µm, n = 15), bitunicate, 8-spored, oblong to clavate, with a very short pedicel or
apedicellate, apically thickened, with a distinct ocular chamber. Ascospores 80–115 ×
25–31 µm ( x = 98 × 28.4 µm, n = 25), crowded to biseriate, fusiform when young,
oblong at maturity 80–113 × 25–31 µm ( x = 98 × 28.35 µm, n = 20), hyaline when
young and becoming brown at maturity, smooth-walled, ornamented, surrounded by a
mucilaginous sheath. Asexual morph Undetermined.
Material examined: THAILAND, Chiang Mai Province, Chom Thong District,
Doi Inthanon National Park, on dead twig, 2 November 2012, I.C. Senanayake TL026
(MFLU 14–0610, holotype); ibid (PDD, isotype), ex-type living culture (MFLUCC
13–0448, ICMP 20755).
Culture characteristics: Ascospores germinating on PDA within 24 h.
Colonies growing on PDA 2 cm diam. after 21 days at 25 °C, slow growing, circular,
effuse, dense, gray, smooth surface with entire to slightly undulate edge.
Notes: Psiloglonium macrosporum is introduced here as a new species based on
morphological traits and phylogeny. In the present phylogenetic analysis P.
macrosporum grouped with other Psiloglonium species (Fig. 11) and is closely related
to P. sasicola (N. Amano) E. Boehm & C. L. Schoch. Psiloglonium macrosporum
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differs from other Psiloglonium species in having 4-spored asci and relatively large,
brown ascospores with ornamentation
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Fig. 12 Psiloglonium macrosporum (holotype) a, b Hysterothecia on host c Vertical section
of hysterothecium d Apex of the hysterothecia e Peridium f Pseudoparaphyses g–i Asci j–m
Ascospores. Scale bars: c = 150 µm, d, e, g–i = 50 µm, f = 10 µm, j–m = 40 µm.
Pleosporales
For an account of Pleosporales see Hyde et al. (2013).
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Fig. 13 Phylogram generated from maximum likelihood analysis based on combined LSU,
SSU, RPB2 and TEF sequence data of Pleosporineae and Massarineae, Pleosporales,
Dothideomycetes. Maximum likelihood bootstrap support values greater than 50% are near
the nodes. The ex-type strains are in bold and the new isolates are in blue. The tree is rooted
with Halotthia posidoniae BBH 22481.
Didymosphaeriaceae
The family Didymosphaeriaceae was introduced by Munk (1953) and is 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. Ariyawansa et al. (2014a) detailed the family and
accepted 16 genera. Austropleospora, Cucubidothis, Munkovalsaria, Spegazzinia,
Sporidesmiella, Paracamarosporium, Pseudocamarosporium, Pseudopithomycetes,
Pseudotrichia, Verrucoconiothyrium, and Xenocamarosporium were later introduced
to the family based on morphology and phylogenetic analysis (Thambugala et al.
2014c, Wijayawardene et al. 2014a, Ariyawansa et al. 2015a, Crous et al. 2015a,
Tanaka et al. 2015). However, the strains of Munkovalsaria appendiculata Aptroot
that cluster with Montagnula species and Sporidesmiella fusiformis W.P. Wu were not
ex-type species. Therefore, Wanasinghe et al. (2016) synonymized Munkovalsaria
under Montagnula, when introducing a new genus, Laburnicola in
Didymosphaeriaceae. The family now contains 28 genera. A phylogenetic tree for the
family is presented in Wanasinghe et al. (2016) and in this paper we used the genera
closest to Pseudocamarosporium (Fig. 14).
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Fig. 14 Phylogram generated from maximum parsimony analysis based on combined LSU,
ITS and SSU sequenced data from species of Didymosphaeriaceae. Maximum
parsimony/likelihood bootstrap support values greater than 50 % and Bayesian posterior
probabilities greater than 0.50 are shown in above and below. The ex-type strains are in bold
and the new isolates is in blue. The tree is rooted with Pyrenochaeta protearum.
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Pseudocamarosporium Wijayaw. & K.D. Hyde
The genus Pseudocamarosporium is typified by P. propinquum and
Paracamarosporium is typified by P. psoraleae and were introduced to accommodate
camarosporium-like species that cluster in Didymosphaeriaceae (Wijayawardene et
al., 2014a). Based on morphology both genera are similar, but Paracamarosporium
has paraphyses and microconidia which are lacking in Pseudocamarosporium.
261. Pseudocamarosporium pini (Westend.) Phukhamsakda, Camporesi & K.D. Hyde,
comb. nov.
Index Fungorum number: IF 551896; Facesoffungi number: FoF 01817, Fig. 15
Basionym: Hendersonia pini Westend., Bull. Acad. R. Sci. Belg., Cl. Sci.: tab.
9, no. 7 (1857)
≡ Camarosporium pini (Westend.) Sacc., Syll. fung. (Abellini) 3: 465 (1884)
Saprobic on dead cone of Pinus nigra J.F. Arnold. Sexual morph
Undetermined. Asexual morph Conidiomata 105–174 µm high × 188–244 wide µm
(x̄ = 145 × 210 µm, n = 5) diam., pycnidial, solitary, uniloculate, scattered, immersed
to erumpent, subglobose, but sometimes irregular, brown to dark brown, ostiole
central. Pycnidial wall 14–28 µm (–40 µm at apex), composed of 5 layers of
brown-walled cells of textura angularis, hyaline inner layer lining bearing
conidiogenous cells. Conidiophores reduced to conidiogenous cells. Conidiogenous
cells 2–6 × 3–5 µm (x̄ = 4 × 4 µm, n = 20) diam., enteroblastic, phialidic, determinate,
smooth-walled, hyaline. Conidia 7–18 × 4–8 µm (x̄ = 14 × 6 µm, n = 50), oval to
oblong, curved at the apex, with 1–3 transverse septa, and 1–2 longitudinal septa at
the second and third cells, initially hyaline, brown to dark brown at maturity, narrowly
rounded at both ends, smooth-walled.
Culture characteristics: Colonies on PDA 60 mm diam. after 4 weeks at 16°C,
cream to white at the margins, pale-brown to yellowish at the center; reverse
yellowish to cream and orangish-white at the center, medium dense, circular,
umbonate, fairly fluffy, without diffusible pigments.
Material examination: ITALY, Forlì-Cesena Province, Monte Mirabello Predappio, on dead and terrestrial cone of Pinus nigra (Pinaceae), 13 Octorber 2014,
E. Camporesi (MFLU 15–3290, HKAS 91937, reference specimen designed here),
ex-type living culture, MFLUCC 14–1091, KUMCC 15–0550.
Note: Several Camarosporium species has been reported from Pinus spp., such
as C. propinquum (Sacc.) Sacc., C. brabeji Marincowitz et al., and C. pini (Westend.)
Sacc. (Grove 1937, Botella et al. 2010, Botella and Diez 2011). Wijayawardene et al.
(2014a) treated C. propinquum under Pseudocamarosporium typified by P.
propinquum. The strain clustered in Didymosphaeriaceae, separate from the type of
Camarosporium, C. quaternatum, which clustered in Pleosporinae. Crous et al.
(2015a) synonymized Camarosporium brabeji Marincowitz et al. under
Pseudocamarosporium brabeji as the molecular data placed them in
Didymosphaeriaceae. Camarosporium pini was originally described by Westendorp
(1857) as Hendersonia pini, and the species is recorded from Pinus silvestris (Grove
1937). When comparing the morphology of our species with C. pini, they are similar
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in the host and morphology. The conidiomata are similar in size, with thick walls up
to 40 µm wide. The dimension of conidia overlap and are oblong, rounded at both
ends, with one or two longitudinal septa in the middle cells. Based on phylogenetic
analysis (Fig. 14) our strain clusters within Pseudocamarosporium in
Didymosphaeriaceae with relative high support (92% MP /88 % ML /0.99 PP). We
therefore synonymize Camarosporium pini under Pseudocamarosporium pini based
on morphology and phylogeny, and designate our collection as a reference specimen
(sensu Ariyawansa et al. 2014c), which we illustrate here.
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Fig. 15 Pseudocamarosporium pini (MFLU 15–3290, reference specimen) a, b Appearance
of conidiomata on Pinus nigra cone c Vertical section of conidioma d Peridium e Ostiole f–i
Developing stages of conidia j–o Conidia p–q Culture characters on PDA. Scale bars: c = 100
µm, d–e = 20 µm, f–o = 5 µm, p–q = 30 mm.
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Lentitheciaceae
The family Lentitheciaceae was introduced to accommodate Lentithecium and
some other related taxa (Zhang et al. 2009a) with species occurring on herbaceous
plants and on submerged wood in freshwater environments (Zhang et al. 2012). There
have been several studies on Lentitheciaceae (Hirayama et al. 2010, Quaedvlieg et al.
2013, Wanasinghe et al. 2014, Ariyawansa et al. 2015b, Knapp et al. 2015, Liu et al.
2015, Phookamsak et al. 2015, Singtripopa et al. 2015, Tanaka et al. 2015,
Wijayawardane et al. 2015). Currently there are eleven accepted genera included
including the new genus introduced in this study (Darksidea, Katumotoa,
Keissleriella,
Lentithecium,
Murilentithecium,
Neoophiosphaerella,
Phragmocamarosporium, Poaceascoma, Setoseptoria, Tingoldiago and Towyspora).
The phylogenetic tree is presented in Fig. 16.
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Fig. 16 Phylogram generated from maximum likelihood analysis based on combined LSU,
SSU, TEF and ITS sequence data for species of Lentitheciaceae. Maximum likelihood
bootstrap support values greater than 50 % and Bayesian posterior probabilities greater than
0.90 are near the nodes. The ex-type strains are in bold and the new isolates are in blue. The
scale bar indicates 0.02 changes. The tree is rooted with Massarina eburnea and M. cisti.
262. Towyspora Wanasinghe, E.B.G. Jones & K.D. Hyde, gen. nov.
Index Fungorum number: IF 551787, Facesoffungi number: FoF 01671
Etymology: Named after the River Towy where this species was collected and
from the Latin, spora meaning spore.
Saprobic on dead shrubs in aquatic habitats. Sexual morph Undetermined.
Asexual morph Conidiomata pycnidial, stromatic, mostly solitary, semi-immersed to
immersed in the host, uni- to multi-loculate, globose to subglobose, dark brown to
black, ostiolate, apapillate. Peridium comprising 2–3 layers, pigmented, thin-walled,
comprise blackish to dark brown, angular cells. Conidiogenous cells phialidic,
discrete, ampulliform to cylindric-clavate, hyaline, aseptate, smooth. Conidia hyaline,
1-celled, oblong to cylindrical, with rounded or obtuse ends, aseptate, smooth-walled,
thin-walled, guttulate.
Type species: Towyspora aestuari Wanasinghe, E.B.G. Jones & K.D. Hyde
Notes: Towyspora gen. et sp. nov. is introduced in the family Lentitheciaceae to
accommodate, T. aestuari based on both morphology and phylogeny. Towyspora
shares most similarities with Setoseptoria in having hyaline, subcylindrical
conidiogenous cells and transversely euseptate, hyaline, smooth-walled,
subcylindrical conidia, with one large central guttule per cell. Towyspora however,
differs from Setoseptoria in having comparatively smaller conidia. This is also
supported phylogenetically as Towyspora aestuari forms a remote clade from
Setoseptoria with high bootstrap support (Fig. 16).
263. Towyspora aestuari Wanasinghe, E.B.G. Jones & K.D. Hyde, sp. nov.
Index Fungorum number: IF 551788, Facesoffungi number: FoF 01672, Fig. 17
Etymology: aestuari from estuary, the habit of the species
Holotype: MFLU 15–3543
Saprobic on Phragmites communis (Cav.) Trin. ex Steud. Sexual morph
Undetermined. Asexual morph Conidiomata 300–400 µm high × 200–250 µm diam.
(x̅ = 347.9 × 223.2 µm, n = 10), pycnidial, stromatic, mostly solitary, semi-immersed
to immersed in the host, uni- to multi-loculate, globose to subglobose, dark brown to
black, ostiolate, apapillate. Peridium 5–10 µm wide at the base, 7–12 µm wide in
sides, comprising 2–3 layers, pigmented, thin-walled, comprising blackish to dark
brown, angular cells. Conidiogenous cells 5–8 µm high × 2–4 µm wide, phialidic,
discrete, ampulliform to cylindric-clavate, hyaline, aseptate, smooth. Conidia 7–12 ×
2.5–3.5 µm (x̅ = 9.6 × 2.8 µm, n = 50), hyaline, 1-celled, oblong to cylindrical, with
rounded or obtuse ends, transversely euseptate, smooth and thin-walled, guttulate.
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Material examined: UK, Lanstephan, 8 July 2015, on Phragmites communis
(Poacaeae), E.B.G. Jones (MFLU 15–3543, holotype); ex-type culture, MFLUCC
15–1274, MUCL.
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Fig. 17 Towyspora aestuari (holotype) a Appearance of conidiomata on host substrate b
Vertical section through conidioma c–f Mature and immature conidia attached to
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conidiogenous cells g–i Mature and immature conidia j Germinated conidium k, l Culture on
PDA (note l reverse). Scale bars b = 50 µm, c = 20 µm, d–j = 5 µm.
Lindgomycetaceae
Lindgomyces K. Hiray. et al.
Lindgomyces K. Hiray. et al. (Lindgomycetaceae, Pleosporales,
Dothideomycetes) is a recently established ascomycetous genus from submerged
wood in freshwater habits (Hirayama et al. 2010). Lindgomyces is characterized by
globose to subglobose ascomata, fissitunicate, clavate to cylindrical asci, and clavate
to cylindrical, hyaline ascospores with a gelatinous sheath (Hirayama et al. 2010).
Lindgomyces currently includes eight species, viz. L. ingoldianus (Shearer & K.D.
Hyde) K. Hiray. et al. (type species), L. apiculatus K. Hiray. & Kaz. Tanaka, L.
breviappendiculatus (Kaz. Tanaka et al.) K. Hiray. & Kaz. Tanaka, L. cinctosporus
Raja et al., L. lemonweirensis Raja et al., L. rotundatus K. Hiray. & Kaz. Tanaka, L.
angustiascus Raja et al. and L. griseosporus Ying Zhang et al. (Hirayama et al. 2010;
Raja et al. 2011, 2013; Zhang et al. 2014). The phylogenetic tree is presented in Fig.
18.
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Fig. 18 Maximum-likehood tree of Lindgomyces okinawaensis based on SSU and LSU
sequence data. Bootstrap values greater than 50 % are presented at the nodes. New taxa are in
blue and ex-types in bold.
264. Lindgomyces okinawaensis Tak. Takah. & Kaz. Tanaka, sp. nov.
MycoBank number: MB 815296; Facesoffungi number: FoF 02022, Fig. 19
Etymology: In reference to the locality, Okinawa where the new species was
collected.
Holotype: HHUF 30498
Saprobic on submerged dead wood. Sexual morph Ascomata 260–290 µm high,
310–340 µm diam., globose to subglobose, black, scattered to grouped, immersed to
erumpent. Neck 50–60 µm long, 50–75 µm wide, short papillate, central. Peridium
35–41 µm thick, composed of an inner layer of polygonal to subglobose, hyaline to
pale brown, thin-walled, 8–12 × 6–7.5 µm cells, and an outer layer of brown-walled
cells. Hamathecium comprising numerous, 1.5–3 µm wide, anastomosed, branched,
cellular pseudoparaphyses. Asci 134.5–183(–208) × (18.5–)23–31(–40.5) µm ( x =
160.9 × 26.5 µm, n = 12), 8-spored, fissitunicate, clavate, rounded at the apex, with an
apical chamber. Ascospores (38–)40–48(–51) × (10–)12–19 µm ( x = 44.9 × 14.9 µm,
n = 50), L/W 2.2–4.3 ( x = 3.1, n = 50), overlapping biseriate to triseriate, hyaline,
pale brown with age, clavate with acute ends, straight or slightly curved, with the
primary septum almost submedian 0.46–0.58 ( x = 0.52, n = 50), filled with small
lipid droplets, slightly constricted at the primary septum, with a broad upper cell,
smooth-walled, becoming 3-septate with age. Asexual morph Undetermined.
Material examined: JAPAN, Okinawa, Kunigami, Aha, Tanagakumui, small
river, on submerged dead twigs of woody plant, 19 May 2015, collector K. Tanaka et
al., KT 3531 (HHUF 30498, holotype); ex-type living culture, MAFF 245410.
Notes: Lindgomyces okinawaensis has relatively wide ascospores. The
morphological features of ascospores are similar to those of L. cinctosporus
(Hirayama et al. 2010). However, the ascospores of L. okinawaensis do not have an
entire gelatinous sheath. The identities of ribosomal ITS sequences between L.
okinawaensis and L. cinctosporus were low [GenBank JF419905; Identities = 408/432
(94.4%), Gaps = 2/432 (0.5%)].
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Fig. 19 Lindgomyces okinawaensis a, b Appearance of ascomata on host surface c, d
Ascomata formed in culture e Ascoma in longitudinal section f Peridium in longitudinal
section g Pseudoparaphyses h–j Asci k–n Ascospores a, b, e, g, i from HHUF 30498
(holotype); c, d, f, h, j–n from MAFF 245410 (ex-holotype). Scale bars: a, c = 1 mm, b, d =
200 µm, e = 50 µm, f–n = 20 µm.
Lophiostomataceae
The family Lophiostomataceae was revisited by Thambugala et al. (2015a).
Based on morphology and phylogenetic analyses of the lophiostomataceous genera,
Lophiostomataceae is presently a large family comprising 16 genera. One new
species is each introduced in the genera Lophiostoma and Sigarispora in this study;
the phylogenetic trees for Lophiostomataceae are presented in Figs 20 and 21.
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Fig. 20 ML tree based on an analysis of combined LSU, SSU and TEF sequence data. Bootstrap
values greater than 70% are indicated at the nodes. New taxa are in blue and ex-type strains are
in bold.
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Fig. 21 Phylogram generated from Maximum likelihood (RAxML) analysis based on
combined LSU, SSU, ITS and TEF1 sequence data of species of Lophiostomataceae.
Maximum likelihood bootstrap support values greater than 50 % are indicated above or below
the nodes, and branches with Bayesian posterior probabilities greater than 0.90 are given.
New taxa are in blue and ex-type strains are in bold. The tree is rooted with Melanomma
pulvis-pyrius.
265. Lophiostoma pseudoarmatisporum Hay. Takah., K. Hiray. & Kaz. Tanaka, sp.
nov.
MycoBank number: MB 815298, Facesoffungi number: FoF 02023, Fig. 22
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Etymology: In reference to the similarity of the ascospore with that of
Lophiostoma armatisporum.
Holotype: HHUF 30497
Saprobic on dead wood. Sexual morph Ascomata 390–515 µm high, 555–645
µm diam., immersed, subglobose to ellipsoidal, black, with a slit-like ostiole.
Peridium in longitudinal section, 25–38 µm thick at sides, composed of 3–5 layers of
angular, hyaline to brown, 10–15 × 2.5–5 µm cells. Hamathecium comprising 1.5–2
µm wide pseudoparaphyses. Asci 105–152 × 15.5–25 µm ( x = 131.3 × 19.7 µm, n =
50), 8-spored, clavate, fissitunicate, pedicellate, with an ocular chamber. Ascospores
29–40 × 9.5–13 µm ( x = 34.4 × 11.3 µm, n = 100), 1–2-seriate, fusiform, hyaline,
with the primary septum mostly submedian (0.48–0.56; x = 0.52, n = 100), the cell
above the septum usually broader than the lower one, smooth-walled, with thin
mucilaginous appendages, 6–10 µm long. Asexual morph Undetermined.
Material examined: JAPAN, Kagoshima, Yakushima Island, Yakusugi land, on
dead twigs of unknown woody plant, 15 March 2007, collector K. Tanaka and H.
Yonezawa, KT 2237 (HHUF 30497, holotype); ex-type living culture, MAFF
245409.
Notes: Morphologically, this taxon has ascospores which are similar to
Lophiostoma armatisporum (Hyde et al. 1992). However, L. pseudoarmatisporum has
wider ascospores than those of L. armatisporum (vs. 28–39 × 7–9.8 µm; Hyde et al.
1992), and the ITS sequence similarity between these two taxa is rather low (405/544
= 74.4 %, with gaps 32/544 = 5.9 %; Liew et al. 2002). Multi-gene phylogenetic
analysis (Fig. 20) indicated that L. pseudoarmatisporum has a close relationship with
Lophiostoma alpigenum, but the latter has longer and slender ascospores (40–45 × 10
µm) with 9–11-septa (Holm and Holm 1988) than those of L. pseudoarmatisporum.
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Fig. 22 Lophiostoma pseudoarmatisporum a Ascoma on host surface b, c Ascoma formed in
culture d Ascoma in longitudinal section e Peridium f Pseudoparaphyses g Ascus apex h, i
Asci with 8 ascospores j–n Ascospores o Germinating ascospore a, d–h, n, o from HHUF
30497 (holotype); b, c, i–m from culture MAFF 245409 (ex-holotype). Scale bars: a–c = 500
µm, d = 100 µm, e–o = 10 µm.
266. Sigarispora Thambug. & K.D. Hyde, in Thambugala et al., Fungal Diversity:
199–266, [40] (2015)
Index Fungorum number: IF 551255, Facesoffungi number: FoF 00823
Notes: Sigarispora was introduced by Thambugala et al. (2015a) based on
morphological characters and phylogenetic analyses and is typified by S. ravennica
(Tibpromma et al.) Thambugala & K.D. Hyde. It is characterized by immersed to
semi-immersed ascomata, a small crest-like ostiole, and brown, cigar-shaped,
multi-septate ascospores. In this study, the new species clustered together with S.
arundinis (Pers.) Thambug. et al., S. ravennica (Tibpromma et al.) Thambugala &
K.D. Hyde, S. caudata (Fabre) Thambug. et al., S. coronillae Wanas. et al. and S.
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caulium (Fr.) Thambug. et al. and formed a distinct clade in Lophistomataceae (Fig.
21).
266. Sigarispora ononidis Qing Tian, Thambug., Camporesi & K.D. Hyde, sp. nov.
Index Fungorum number: IF 551729, Facesoffungi number: FoF 01639, Fig. 23
Etymology: In reference to its occurrence on Ononis sp., ononidis meaning “of
Ononis”.
Holotype: MFLU 15–2667
Saprobic on the dead stem of Ononis spinosa L. in terrestrial habtats. Sexual
morph Ascomata 240–311.5 µm diam. ( x = 287.2 µm, n = 10), perithecial, solitary,
scattered to gregarious, immersed or semi-immersed to erumpent, gregarious, circular,
globose or subglobose, coriaceous, black, ostiolate, smooth-walled. Ostiole central,
rounded, with a pore-like opening. Peridium 250–320 µm wide × 196–250 µm high ( x
= 279 × 220.5 µm, n = 10), two-layered, outer layer composed of irregular,
thick-walled, brown to dark brown cells of textura angularis and inner layer with
slightly, smaller cells of textura angularis. Hamathecium comprising 1–3 µm wide,
branched or simple, septate, cellular, pseudoparaphyses, embedded in agelatinous
matrix, between and above the asci. Asci 96–169 × 17–19 µm ( x = 120.6 ×18 µm, n =
10), 8-spored, bitunicate, fissitunicate, cylindrical to clavate or broader-clavate, long
pedicellate, apically rounded, with an ocular chamber. Ascospores 27–34 × 11–12 µm
( x = 29 × 11.7 µm, n = 10), overlapping uni-seriate or bi-seriate, yellowish brown to
dark brown, ellipsoid to fusiform or cigar-shaped, 3–5-septate or rarely muriform with
one vertical septa, slightly curved, constricted at the central septum, darkened, with
rounded ends, smooth-walled, without a sheath. Asexual morph Undetermined.
Material examined: ITALY, Province of Forlì-Cesena, Valbura-Premilcuore, on
dead stem of Ononis spinosa (Fabaceae), 18 June 2014, Erio Camporesi, IT1941
(MFLU 15–2667, holotype); ibid., (HKAS 92413, isotype); ex-type living cultures,
MFLUCC 14–0613, KUMCC 15–0524.
Notes: Sigarispora ononidis is introduced here as a new species which is
morphologically similar with species in Sigarispora, a genus established by
Thambugala et al. (2015a). Sigarispora ononidis differs from other species of
Sigarispora in having 3–5-septate or rarely muriform ascospores, without a
mucilaginous sheath (Fig. 23). Phylogenetic analyses of combined genes indicated
that the ex-type strain of S. ononidis clustered within the clade of Sigarispora (Fig.
21).
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Fig. 23 Sigarispora ononidis (holotype) a–c Appearance of ascomata semi-immersed in the
host d, e Vertical section of ascoma f Vertical section of peridium g Immature ascus h–j
Mature asci with ascospores k Hamathecium n Germinated ascospore o–r Ascospores l
Colony on MEA from above m Colony on MEA from below. Scale bars: a = 500 µm, b, c =
200 µm, d, e = 50 µm, f = 20 µm, g–k, n = 10 µm, o–r = 5 µm.
Melanommataceae
The family Melanommataceae was introduced by Winter (1885) and is
characterized by globose or depressed perithecial ascomata, bitunicate and
fissitunicate asci, hyaline or brown and 1 to multi-septate ascospores (Zhang et al.
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2012; Hyde et al. 2013; Tian et al. 2015). Barr (1990) reviewed the family and
included Ostropella, Keissleriella, Strickeria, Byssosphaeria and Melanomma.
Subsequently various authors had included and excluded different species in
Melanommataceae at various times. Tian et al. (2015) revised the family and accepted
20 genera, including seven asexual morphs. The phylogenetic tree is presented in Fig.
24.
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Fig. 24 Phylogram generated from Maximum Likelihood (RAxML) analysis based on
combined LSU and EF sequence data of taxa from Melanommataceae and Pleomassariaceae.
Maximum likelihood bootstrap support values greater than 50 % are indicated above and
below the nodes. New taxa are in blue and ex-type strains are in bold. The tree is rooted with
Massarina eburnea.
Aposphaeria Berk.
Aposphaeria is a poorly known genus and recent studies have been classified this
genus in Melanommataceae based on sequence data (De Gruyter et al. 2012; Tian et
al. 2015). Aposphaeria or aposphaeria”- like species have been reported for different
genera such as Chaetomastia, Massariosphaeria, Melanomma, Mytilinidion and
Rhytidhysteron (Sivanesan 1984; Barr 1989; Zhang et al. 2012; Hyde et al. 2013; Tian
et al. 2015). However, sequence data of the type species, A. pulviscula (Sacc.) Sacc.,
are essential to confirm the phylogeny of Aposphaeria in Melanommataceae. This
genus is characterized by pycnidial, unilocular conidiomata, short, cylindrical,
branched conidiophores and hyaline, aseptate, cylindrical or ellipsoidal conidia (Tian
et al. 2015).
267. Aposphaeria corallinolutea Gruyter et al., in Gruyter et al., Stud. Mycol. 75: 28
(2012)
Facesoffungi number: FoF 01647, Fig. 25
Saprobic on decaying wood. Sexual morph Undetermined. Asexual morph
Pycnidia 200–320 µm diam., superficial, globose to subglobose, black, shiny,
aggregated or solitary, with or without a distinct ostiole. Pycnidial wall comprising
several lightly pigmented to dark brown cells of textura angularis. Conidiophores
6–26 × 1–2 µm (x̅ = 14.4 × 1.5 µm, n = 25), branched, cylindrical, septate, hyaline and
formed from the inner wall cells of the pycnidial wall. Conidiogenous cells
enteroblastic, phialidic, determinate, ampulliform to filiform, hyaline, smooth.
Conidia 2.6–4.2 × 1–1.5 µm (x̅ = 3.8 × 1.2, n = 50), ellipsoidal, hyaline, aseptate,
eguttulate or with some small, polar guttules, smooth-walled.
Culture characteristics: Colonies on PDA 14–16 mm diam. after 9 d, margin
entire to somewhat lobate; colony white to pale white with white, felty aerial
mycelium; reverse brown to greenish olivaceous, greenish grey at centre, white near
margin.
Material examined: THAILAND, Chiang Rai Province, Mae Fah Luang
University Garden, 1 December 2014, Kasun M. Thambugala, TL 987 (MFLU
15–3203), living culture MFLUCC 14–0504.
Notes: Aposphaeria was introduced by Saccardo (1880) and currently there are
207 epithets listed in this genus (Index Fungorum 2016), but sequence data is
available for only a few species. Aposphaeria corallinolutea was introduced by de
Gruyter et al. (2012) and our strain clustered with the ex-type strain (CBS 131287) of
A. corallinolutea (Fig. 24). Aposphaeria corallinolutea has been reported on Kerria
japonica (Rosaceae) and Fraxinus excelsior (Oleaceae) in Netherlands (Gruyter et al.
2012). This is the first report of A. corallinolutea in Thailand.
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Fig. 25 Aposphaeria corallinolutea (MFLUCC 14–0504) a Pycnidia on PDA b
Section through stromatic pycnidia c Pycnidial wall d–e Conidiophores and
conidiogenous cells f Conidia. Scale bars: b = 100 µm, c = 20 µm, d–f = 10 µm.
Parabambusicolaceae
Parabambusicolaceae was introduced by Tanaka et al. (2015) and is typified by
Parabambusicola Kaz. Tanaka & K. Hiray. The family was introduced to
accommodate Massarina-like species from bamboo and grasses, and initially included
the sexual genera Aquastroma, Multiseptospora and Parabambusicola (Tanaka et al.
2015). Two unnamed Monodictys species also clustered in this family, but Monodictys
is obviously heterogenous (Tanaka et al. 2015). In this paper, we introduce a new
genus, Multilocularia to accommodate a single Dothideomycetes species, which was
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collected from bamboo in Thailand. Additionally, a new species of Multiseptospora,
M. thysanolaenae is introduced.
268. Multilocularia Phookamsak, Ariyawansa & K.D. Hyde, gen. nov.
Index Fungorum number: IF 551946, Facesoffungi number: FoF 01658
Etymology: The generic epithet “Multilocularia” refers to the multi-loculate
ascostroma
Saprobic on bamboo. Sexual morph Ascostromata gregarious, clustered,
immersed, visible as raised, black rows, on host surface, multi-loculate, elongate,
glabrous, ostiolate. Locules clustered, immersed in ascostromata, globose to
subglobose, or elongate hemisphaerical, ostiole individually central. Peridium thin- to
thick-walled, slightly thick at the rim, composed of several layers of dark brown to
black, pseudoparenchymatous cells, arranged in a textura angularis. Hamathecium
composed of dense, broad cellular pseudoparaphyses, filamentous, distinctly septate,
anastomosing among the asci, embedded in a hyaline, gelatinous matrix. Asci
8-spored, bitunicate, fissitunicate, clavate, long pedicellate, apically rounded, with
well-developed ocular chamber. Ascospores overlapping 1–2-seriate, hyaline,
ellipsoidal, with rounded ends, slighty curved, septate, slightly constricted at the
central septum, smooth-walled, with small guttules. Asexual morph Undetermined.
Type species: Multilocularia bambusae Phookamsak, Ariyawansa & K.D. Hyde
Notes: Multilocularia is introduced as a monotypic genus to accommodate the
Dothideomycetes species, forming elongate ascostromata with multi-loculate and
phragmosporous, hyaline, ellipsoidal ascospores. The genus is commonly found on
bamboo as saprobes, similar to the genus Munkovalsaria Aptroot in forming
ascostromata on the host, with asci have long pedicellate and ellipsoidal ascospores.
However, Multilocularia differs from Munkovalsaria in having a greater number of
locules than Munkovalsaria and ascospores are hyaline, while in Munkovalsaria
ascospores are brown. Multilocularia clusters with Aquastroma magniostiolata,
Pseudomonodictys tectonae and Monodictys species in Parabambusicolaceae in the
phylogenetic tree (Fig. 13), whereas, Munkovalsaria belongs in Didymosphaeriaceae
(Ertz et al. 2015) which is synonymized under Montagnula by Wanasinghe et al.
(2016). Multilocularia differs from Pseudomonodictys tectonae and Monodictys
species based on its phylogenetic distinctiveness. Pseudomonodictys and Monodictys
species are presently only know as asexual morphs, while Multilocularia is known in
its sexual morph. Aquastroma differs from Multilocularia in having globose
ascostromata, short pedicellate asci, clavate to fusiform, multi-septate ascospores and
an aquatic habitat.
269. Multilocularia bambusae Phookamsak, Ariyawansa & K.D. Hyde, sp. nov.
Index Fungorum number: IF 551947, Facesoffungi number: FoF 01659, Fig. 26
Etymology: The specific epithet “bambusae” refers to the host
Holotype: MFLU11–0216
Saprobic on bamboo. Sexual morph Ascostromata 200–240 µm high,
1100–1900 µm long, gregarious, clustered, immersed, raised, in black rows on host
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surface, multi-loculate, elongate, glabrous, ostiolate. Locules 130–240 µm high,
200–700 µm diam., clustered, immersed in ascostromata, globose to subglobose, or
elongate hemisphaerical, ostiole individually central. Peridium 10–40 µm wide, thinto thick-walled, slightly thick at the rim, composed of several layers of small, brown
to dark brown pseudoparenchymatous cells, arranged in a textura prismatica to
textura angularis, and arranged in textura porrecta at the sides among the locules.
Hamathecium composed of dense, 1.2–2 µm wide, cellular pseudoparaphyses,
distinctly septate, anastomosing among the asci, embedded in a hyaline gelatinous
matrix. Asci (64–)70–90(–94) × (10–)11–14(–17) µm ( x = 82.5 × 14.2 µm, n = 30),
8-spored, bitunicate, fissitunicate, clavate, long pedicellate (30–50 × 3–5 µm),
apically rounded, with well-developed ocular chamber. Ascospores (11–)12–15(–16)
× (3–)4–5 (–7) µm ( x = 14.2 × 4.7 µm, n = 30), overlapping 1–2-seriate, hyaline,
ellipsoidal, with rounded ends, slighty curved, 3-septate, rarely 1- to 4-septate,
slightly constricted at the central septum, smooth-walled, with small guttules. Asexual
morph Undetermined.
Culture characteristics: Colonies on PDA reaching 30–40 mm diam. after 4
weeks at 25–30°C, colony from above, dark greenish to black at the margin, white to
orange in the middle, white at the centre; from below, dark greenish to black; medium
dense, irregular, slightly raised to umbonate, surface slightly rough, dull with
umbonate edge, concave at the centre, fluffy to floccose, with white tufts at the centre;
producing brown pigmentation in agar.
Material examined: THAILAND: Chiang Rai Province, Mae Jun District, Huai
kang Pla Waterfall, on dead stem of bamboo (Poaceae), 25 October 2010, R.
Phookamsak, RP0096 (MFLU 11–0216, holotype), ex-type living culture, MFLUCC
11-0180, BCC.
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Fig. 26 Multilocularia bambusae (holotype) a Appearance of ascostromata on host surface b
Section through an ascostroma c Appearance of locules d Section through peridium e Asci
with pseudoparaphyses, stained in congo red f, g Asci h–l Ascospores m Ascospore stained
congo red n Spore germination on WA after 8 hours. Scale bars: b = 200 µm, c = 100 µm, d =
50 µm, e–g = 20 µm, n = 10 µm, h–m = 5 µm.
Multiseptospora Phookamsak & K.D. Hyde
The genus Multiseptospora Phookamsak & K.D. Hyde was introduced in Liu et
al. (2015) to accommodate a single species M. thailandica Phookamsak & K.D. Hyde,
which was collected on Thysanolaena maxima Kuntze. The genus was introduced in
the Pleosporales genera incertae sedis (Liu et al. 2015). However, Tanaka et al. (2015)
added the genus to Parabambusicolaceae when they introduced this family based on
their phylogenetic relationships. In this study, a new species, M. thysanolaenae is
introduced. The new species was also collected on Thysanolaena maxima in Thailand.
270. Multiseptospora thysanolaenae Phookamsak, Ariyawansa & K.D. Hyde, sp.
nov.
Index Fungorum number: IF 551948, Facesoffungi number: FoF 01660, Fig. 27
Etymology: The specific epithet “thysanolaenae” refers to the host.
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Holotypus: MFLU 11–0238
Saprobic on Thysanolaena maxima Kuntze. Sexual morph Ascostromata
190–270 µm high, 300–350 µm diam., gregarious, scaterred, immersed, visible as
raised, black dots on host surface, uni-loculate, globose to subglobose, glabrous,
ostiole central, with minute papilla. Peridium 12–40 µm wide, thin- to thick-walled,
slightly thick at the sides towards apex, composed of several layers of flattened,
pseudoparenchymatous cells, inner layers comprising flattened, hyaline cells,
arranged in a textura prismatica, outer layers comprising brown to dark brown cells,
arranged in a textura angularis. Hamathecium composed of dense, 1.8–4 µm wide,
cellular pseudoparaphyses, slightly constricted at the septum, anastomosing among
the asci, embedded in a hyaline gelatinous matrix. Asci (93–)100–120(–143) ×
(26–)28–32(–35) µm ( x = 114.3 × 30.4 µm, n = 30), 8-spored, bitunicate,
fissitunicate, broadly cylindric-clavate to clavate, subsessile to short pedicellate,
apically rounded, with an indistinct ocular chamber. Ascospores (55–)60–65(–73) ×
(8–)9–11(–13) µm ( x = 64.6 × 10.5 µm, n = 30), overlapping 3–4-seriate, initially
hyaline, becoming brown to dark brown at maturity, fusiform, with slightly rounded
ends, slighty curved, (6–)7-septate, not constricted at the septa, smooth-walled,
surrounded by thin, mucilaginous sheath, with small appendages at both ends.
Asexual morph Undetermined.
Culture characteristics: Colonies on PDA fast growing, reaching 70–80 mm
diam. after 4 weeks at 25–30°C, colony from above, light brown to dark brown; from
below: black; dense, circular, slightly raised to umbonate, surface smooth, dull with
entire edge, concave at the centre, fluffy to floccose, producing brown pigmentation in
agar.
Material examined: THAILAND, Chiang Mai, Doi Suthep-Pui, on dead leaf
sheath of Thysanolaena maxima (Poaceae), 5 June 2011, R. Phookamsak, RP0118
(MFLU 11–0238, holotype), ex-type living culture, MFLUCC 11–0202, BCC.
Notes: Multiseptospora thysanolaenae is similar to the type species, M.
thalandica in having multi-septate ascospores and is associated with Thysanolaena
maxima Kuntze. However, M. thysanolaenae differs from M. thailandica due to its
glabrous ascostromata, with brown ascospores. Multiseptospora thysanolaenae has
larger ascomata, asci and ascospores than M. thailandica, but has less ascospore
septation (ascospores septation: 10–11-septate in M. thailandica versus 6–7-septate in
M. thysanolaenae. Based on phylogenetic analysis, M. thysanolaenae clusters with M.
thailandica (Fig. 13).
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Fig. 27 Multiseptospora thysanolaenae (holotype) a Appearance of ascostromata on host
surface b Section through an ascostroma c Section through peridium d Pseudoparaphyses
stained in Indian ink e Asci with pseudoparaphyses f–h Asci i–l Ascospores m Ascospore
stained in Indian ink n Spore germination on WA after 8 hours. Scale bars: b = 100 µm, c, e =
50 µm, d, f–n = 20 µm.
Phaeosphaeriaceae
The family Phaeosphaeriaceae (Pleosporales) was introduced by Barr (1979a)
and is a heterogeneous group of taxa comprising plant pathogens, saprobes and
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endophytes, associated with a wide variety of plant hosts (Zhang et al. 2012; Hyde et
al. 2013; Phookamsak et al. 2014). The family is typified by Phaeosphaeria with P.
oryzae as the type species. Initially the family comprised 15 genera (Barr 1979a), and
now comprises more than 35 sexual and asexual genera (Hyde et al. 2013;
Phookamsak et al. 2014). Various phylogenetic studies have been carried out on
Phaeosphaeriaceae and several new genera has been introduced, while some has been
transferred to other families (Zhang et al. 2012; Hyde et al. 2013; Phookamsak et al.
2014; Trakunyingcharoen et al. 2014; Crous et al. 2015c, d; Ertz et al. 2015; Li et al.
2015c). In the present study, a backbone tree for the family is presented (Fig. 28) with
the genera Allophaeosphaeria, Ampelomyces, Chaetosphaeronema, Coniothyrium,
Dematiopleospora, Didymocyrtis, Edenia, Entodesmium, Galliicola, Leptospora,
Loratospora,
Muriophaeosphaeria,
Neosetophoma,
Neostagonospora,
Neosphaerellopsis, Nodulosphaeria, Ophiobolus, Ophiosphaerella, Paraphoma,
Parastagonospora, Phaeosphaeria, Phaeosphaeriopsis, Poaceicola, Populocrescentia,
Sclerostagonospora, Scolicosporium, Septoriella, Setomelanomma, Setophoma,
Sulcispora, Stagonospora, Vagicola, Vrystaatia, Wojnowicia, Wojnowiciella,
Xenophoma, and Xenoseptoria. The phylogenetic tree is presented in Fig. 28.
Notes: Our phylogenetic analyses of taxa of Phaeosphaeriaceae, uses combined
LSU and ITS sequence data, and comprises 106 strains, representing 37 genera, with
Didymella exigua (CBS 183.55) as the outgroup taxon. The phylogenetic analyses
provides good evidence for one new species, Parastagonospora cumpignensis (strain
MFLUCC 13–0573), which clusters with their respective genus with strong support.
Parastagonospora cumpignensis forms a distinct clade with P. dactylidis (strain
MFLUCC 13–0375), with a relatively high 100% MP and 96% ML bootstrap support,
and a high Bayesian posterior probability (1.0 PP).
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Fig. 28 Phylogram generated from maximum likelihood analysis based on combined LSU and
ITS sequenced data of species of Phaeosphaeriaceae. Branches of maximum parsimony and
maximum likelihood bootstrap support values greater than 50% and Bayesian posterior
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probabilities greater than 0.90 are indicated in bold. New taxa are in blue and ex-type strains
are in bold. The scale bar indicates 0.1 changes. The tree is rooted with Didymella exigua
CBS 183.55.
Parastagonospora Quaedvl. et al.
Parastagonospora was introduced by Quaedvlieg et al. (2013) with P. nodorum
(Berk.) Quaedvl. et al. as the type species. Parastagonospora is a plant pathogenic
genus accommodating taxa that were formerly placed in either Septoria/Stagonospora,
or Leptosphaeria/Phaeosphaeria (Quaedvlieg et al., 2011, 2013; de Gruyter et al.
2013; Ariyawansa et al. 2015c). The sexual and asexual characters of this genus were
described in Quaedvlieg et al. (2013).
271. Parastagonospora cumpignensis Tibpromma, Camporesi & K.D. Hyde, sp. nov.
Index Fungorum number: IF 551804, Facesoffungi number: FoF 01767, Fig. 29
Etymology: Name reflects the locality, Campigna, where this species was
collected.
Holotype: MFLU 15–1480
Saprobic on Dactylis glomerata L. in terrestrial habitats. Sexual morph
Ascomata 205–310 µm high × 197–217 µm diam. (x̅ = 245 × 207 µm, n = 5), scattered,
immersed in host tissue, globose to subglobose, thin-walled, solitary, with short neck,
dark brown to black. Peridium 14–19 µm, thin-walled, comprising 2 layers of hyaline
to brown cells of textura angularis. Hamathecium comprising numerous, 1.5–3 µm
wide, septate, branched, pseudoparaphyses. Asci 62–92 × 9–12 µm (x̅ = 78 × 10 μm, n
= 10), 8-spored, bitunicate, cylindrical to narrowly fusoid, short pedicellate, with a
relatively a small ocular chamber. Ascospores 26–31 × 6–7 µm ( x̅= 28 × 7 µm, n =
15), obliquely uniseriate, ellipsoid to narrowly obovoid, hyaline, becoming 3-septate
with age, constricted at each septum, cells above central septum often broader than the
lower ones, with acute rounded ends, constricted at the septa, with 1–2 distinct oil
droplets in each cell, smooth-walled, without a mucilaginous sheath. Asexual morph
Undetermined.
Culture characteristics: on MEA reaching 4 cm diam. after 2 weeks at 16°C,
later with dense mycelium, with entire edge, flat, smooth with raised elevation,
white-grey; hyphae septate branched, grey, thin-walled.
Material examined: ITALY, Campigna, Santa Sofia, Forlì-Cesena Province, on
dead stem of Dactylis glomerata (Poaceae), 23 June 2012, Erio Camporesi, IT458
(MFLU 15–1480, holotype); ex-type living culture, MFLUCC 13–0573, MUCL; Ibid.
(MFLU 16-0065bis, HKAS 92500tris, paratypes).
Notes: The phylogeny of the family Phaeosphaeriaceae is reconstructed based
on analysis combined LSU and ITS sequence data (Fig. 28). Parastagonospora
cumpignensis clusters with P. dactylidis W.J. Li et al. and P. minima W.J. Li et al. with
high support. Parastagonospora dactylidis and P. minima are asexual morphs with
3-septate, hyaline conidia, while P. cumpignensis is a sexual morph which shares
3-septate, hyaline ascospores with P. dactylidis and P. minima (Li et al. 2015c).
Parastagonospora cumpignensis is introduced as new species with an illustrated
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account and the phylogenetic trees of combined LSU and ITS sequence data confirm
its placement in Parastagonospora.
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Fig. 29 Parastagonospora cumpignensis (holotype) a Appearance of ascomata on host
substrate b Section of ascoma c Section of peridium d Paraphyses e, f Asci g–i Ascospores j
Germinated ascospore. Scale bars: a = 200 µm, b = 50 µm, c = 20 µm, d = 5 µm, e, f = 20 µm,
g–j = 10 µm.
Pleosporaceae
The family was recently detailed by Ariyawansa et al. (2015a) and this is followed
here.
Comoclathris Clem.
Comoclathris was introducing by Clements (1909) and is typified by
Comoclathris lanata Clem. Comoclathris is characterized by ascomata with circular
lid-like openings and applanate, reddish brown to dark reddish brown, muriform
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ascospores, with single longitudinal septa (Zhang et al. 2012; Ariyawansa et al. 2014b;
Crous et al. 2014a).
272. Comoclathris pimpinellae Konta, Bulgakov & K.D. Hyde, sp. nov.
Index Fungorum number: IF 551949, Facesoffungi number: FoF 01769, Fig. 30
Etymology: The specific epithet refers to the host genus Pimpinella.
Holotypus: MFLU 15–0010
Saprobic on dead stems of Pimpinella tragium Vill. subsp. titanophila
(Woronow) Tutin (syn. Pimpinella titanophila Woronow) appearing as black spots on
host surface, or small black lines arising from cracks in the epidermal cells. Sexual
morph Ascomata 155–135 wide × 88–95 µm high (x̅ = 149 × 95 μm, n = 10), solitary
or aggregated, semi-immersed or rarely somewhat superficial, globose to subglobose,
dark brown to black. Peridium 10–19 μm wide, comprising an outer layer of dark
brown cells of textura angularis and inner layer of mostly hyaline to pale brown cells
of textura angularis. Hamathecium comprising numerous, 1.3–2.1 μm wide, septate,
pseudoparaphyses. Asci 58–75 × 14–16 µm, (x̅ = 62 × 16 μm, n = 10), 8-spored,
bitunicate, fissitunicate, cylindrical-clavate, short-pedicellate, rounded at the apex,
with indistinct, shallow, ocular chamber. Ascospores 14–16 × 5–8 µm (x̅ = 15 × 7 μm,
n = 10), overlapping biseriate, yellow to light brown, transversely septate or muriform,
with 3 transverse septa, central segments with 2 longitudinal septa, end segments with
2 angular septa, surrounded by a thick, hyaline, a mucilaginous sheath. Asexual
morph Undetermined.
Culture characteristics: Colonies on MEA, reaching 5–6.5 cm diam. after 2
weeks at 16°C, smoky-grey to dark green, margins smooth, medium dense, with fairly
fluffy surface.
Material examined: RUSSIA, Rostov region, Shakhty City, near Grushevsky
Pond, stony steppe, dead stems of Pimpinella tragium Vill. subsp. titanophila
(Woronow) Tutin (syn. Pimpinella titanophila Woronow), 18 May 2014, T.S.
Bulgakov (MFLU 15–0010, holotype, HKAS, isotype); ex-type living culture,
MFLUCC 14–1159.
Notes: Comoclathris is characterised by ascomata with circular lid-like openings
and applanate, reddish brown to dark reddish brown, muriform ascospores, with
single longitudinal septa (Zhang et al. 2012). This genus includes 36 species names in
Index Fungorum (2016) and the type species is C. lanata Clem. In this paper we
introduce C. pimpinellae based on morphology and phylogeny. Maximum Likelihood
analysis of combined LSU, SSU, RPB2 and TEF sequence data (Fig. 13) indicates
that C. pimpinellae is closest to C. compressa with high bootstrap support (100% ML)
and groups in the Comoclathris clade, but is distinct with other species in this genus.
The sexual morph of C. pimpinellae differs from C. compressa, C. lanata (type) and
C. sedi in having ascomata not surrounded by radiating brown hypha (Fig. 30 a–c viz
Fig. 8 a and Fig. 9 a, d in Ariyawansa et al. 2015b), and yellow to light brown
ascospores with 3 transverse septa, with central segments with 2 longitudinal septa
and end segments with 2 angular septa (Fig. 30 j–m viz Fig. 8 g–i and Fig. 9 i in
Ariyawansa et al. 2015). No Comoclathris species have been described from
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Pimpinella. Therefore, we introduce C. pimpinellae as a new species based on
morphology, phylogeny and host association.
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Fig. 30 Comoclathris pimpinellae (holotype) a Appearance of ascomata on host substrate b
Close up of ascomata c Section of ascoma d Peridium e Pseudoparaphyses f–i Asci j–n
Ascospores. Scale bars: a = 500 µm, b = 200 µm, c = 50 µm, d–i = 20 µm, j–n = 10 µm.
Testudinaceae
A family of Pleosporales that was introduced by von Arx (1971) to
accommodate“astomatous ascomata with a dark peridium, which is often made up of
plates, with bitunicate asci, and dark 2-celled ascospores, about 10 µm long”. The
family contains five genera namely: Lepidosphaeria, Neotestudina, Testudina (type
genus), Ulospora and Verruculina. Species belonging to the family are either saprobic
in the terrestrial habitats (Lepidosphaeria, Testudina and Ulospora), dermatophytes
(Neotestudina) or marine fungi (Verruculina). Further information about the family is
available in Hyde et al. (2013).
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Fig. 31 Phylogram generated from maximum likelihood analysis (MEGA6) based on
combined dataset of SSU and LSU sequence data of the two new genera and related taxa in
Pleosporales. Representatives of the orders Mytilinidiales, Capnodiales and Dothideales are
used as outgroup taxa. Maximum Likelihood bootstrap values greater than 50 % are indicated
at the nodes. The new genera are in blue. Ex-type strains are in bold.
273. Angustospora Abdel-Aziz, gen. nov.
Index Fungorum number: IF 551714, Facesoffungi number: FoF 01632
Etymology: In reference to the striate ascospores.
Type species: Angustospora nilensis Abdel-Aziz
Saprobic on decayed wood in freshwater habitats. Sexual morph Ascomata
globose to subglobose, immersed to erumpent, solitary, ostiolate, papillate,
periphysate, coriaceous to sub-carbonaceous, dark-brown to black. Peridium
comprising two strata, outer stratum dark-brown to black, forming a textura
angularis, inner stratum comprising hyaline, thick-walled, flattened cells arranged in a
textura angularis. Hamathecium comprising numerous, 1–2.5 µm wide, distantly
septate, branched, trabeculate pseudoparaphyses, within a gelatinous matrix,
anastmosing above asci. Asci 8-spored, bitunicate, fissitunicate, clavate, short
pedicellate, apically rounded, with a wide, shallow ocular chamber and faint ring.
Ascospores overlapping biseriate, dark-brown to black, (3)–5–(7) septate, polar cells
lighter when young and apical cells with two-walls, surrounded by thin, gelatinous,
striate layer. Asexual morph Undetermined.
Notes: The phylogenetic analyses of both SSU and LSU sequence data place the
genus Angustospora within the family Testudinaceae (Fig. 31). This phylogenetic
placement was consistent with various degrees of bootstrap support in all the
phylogenetic analyses performed (data not shown). Arx (1971) established the family
Testudinaceae to accommodate four genera namely: Lepidosphaeria, Neotestudina,
Pseudophaeotrichum and Testudina (type genus). Suetrong et al. (2009) assigned the
monotypic marine genus Verruculina to the family Testudinaceae, based on
multi-gene analyses. Verruculina enalia (Kohlm.) Kohlm. & Volkm.-Kohlm. is
characterized by small ascomata (less than 500 µm in diam.), that are subglobose,
ampulliform or depressed ellipsoidal, immersed to erumpent, ostiolate, papillate,
clypeate, carbonaceous, black and solitary. Asci are 8-spored, cylindrical, pedicellate,
bitunicate, thick-walled, physoclastic, without apical apparatuses. Ascospores are
obliquely uniseriate, ellipsoidal, 1-septate, constricted at the septum, dark brown,
verrucose, with a hyaline tubercle at each apex which is probably a germ pore
(Kohlmeyer and Kohlmeyer 1979). Angustospora is not congeneric with Verruculina
as their morphology is quite different and they are phylogenetically distant (Fig. 31).
The genus Angustospora is reminiscent of Caryospora in having large ascospores
with a median septum and additional septa near poles of the ascospores. However,
Angustospora is different from species of Caryospora in having small ascomata and
8-spored, clavate asci (Barr 1979b, 1990; Hawksworth 1982; Abdel-Wahab and Jones
2000; Raja and Shearer 2008; Zhao and Zhao 2012; Ariyawansa et al. 2015b). Ten
species currently are recognized in the genus Caryospora, of which five were
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recorded from aquatic habitats (Abdel-Wahab and Jones 2000; Raja and Shearer
2008; Jones et al. 2015; Ariyawansa et al. 2015b). Ariyawansa et al. (2015)
established the new family Caryosporaceae for two species of Caryospora and the
marine genus Acrocordiopsis Borse & K.D. Hyde. The family Caryosporaceae
formed a basal clade to Testudinaceae (Ariyawansa et al. 2015b, Fig. 30).
Angustospora nilensis has smaller ascomata than most of the described Caryospora
species and different dimensions of asci and ascospores. Raja and Shearer (2008)
described C. obclavata Raja & Shearer from decayed wood in freshwater habitats,
with small ascomata, however, A. nilensis has larger asci and ascospores.
274. Angustospora nilensis Abdel-Aziz, sp. nov.
Index Fungorum number: IF 551715, Facesoffungi number: FoF 01633, Fig. 32
Etymology: In reference to the habitat where the fungus was first collected.
Holotype: CBS
Saprobic on decayed submerged wood in freshwater habitats. Sexual morph
Ascomata 225–420 µm high, 325–390 µm diam., globose to subglobose, immersed to
erumpent, solitary, ostiolate, papillate, periphysate, coriaceous to sub-carbonaceous,
dark-brown to black. Papilla 100–180 µm long, 110–160 µm wide, protruding above
the wood surface. Ostiolar canal 150–300 µm long, 80–160 µm wide, cylindrical to
triangular, filled with periphyses that are 30 to 50 µm long and 2–3 µm wide.
Peridium 57–85 µm thick, comprising two strata; outer stratum 39–54 µm thick,
dark-brown to black, forming a textura angularis; inner stratum 18–31 µm thick
comprising hyaline, thick-walled, flattened cells, arranged in a textura angularis.
Hamathecium comprising numerous, 1–2.5 µm wide, distantly septate, branched,
trabeculate pseudoparaphyses, embedded in a gelatinous matrix, anastmosing above
the asci. Asci 150–240 × 48–83 µm ( x = 193.9 × 59.9 µm, n = 10), 8-spored,
bitunicate, fissitunicate, clavate, semi-persistent, short pedicellate, apically rounded,
with a wide, shallow ocular chamber and faint ring. Ascospores 45–68 × 26–35 µm ( x
= 58.6 × 30 µm, n = 50), overlapping biseriate, dark-brown to black, (3)–5–(7)-septate,
polar cells are lighter when young and apical cells with two-walls, surrounded by thin
gelatinous, striate layer. Asexual morph Undetermined.
Culture characteristics: Colonies on PDA reaching a 20–30 mm diam. after 15
days at 25 °C, with gray to dark-brown aerial and immersed mycelium, dark-brown to
black in reverse, producing fertile ascomata after 40 to 60 days of incubation,
ascomata, asci and ascospores produced in culture with dimensions similar to those
recorded on natural wood.
Material examined: EGYPT, Sohag City, on decayed wood submerged in the
River Nile, 8 March 2005, F.A. Abdel-Aziz (CBS, holotype); ex-type living culture in
CBS.
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Fig. 32 Angustospora nilensis (holotype) a Vertical section of ascoma b Magnified part of
the vertical section of the ascoma showing the papilla and ostiolar canal c, d Immature asci e,
g Mature asci h Ocular chamber in ascus and faint ring i, k Variously shaped ascospores at
different stages of maturity with striate gel coating (evident in j). Scale bars: a = 100 µm, b =
50 µm, c = 40 µm, d–g = 30 µm, h–k = 12 µm.
Tetraplosphaeriaceae
The family Tetraplosphaeriaceae accommodates Tetraploa, Triplosphaeria,
Polyplosphaeria, Pseudotetraploa, and Quadricrura ( Tanaka et al. 2009; Hyde et al.
2013). Of these, the genera Tetraploa, Polyplosphaeria and Triplosphaeria have
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Massarina-like sexual morphs with almost hyaline 1(–3)-septate ascospores and
Tetraploa-like asexual morphs with several setose appendages (Tanaka et al. 2009;
Hyde et al. 2013;). The sexual morph of the genera Pseudotetraploa and Quadricrura
are undetermined. The phylogenetic tree is presented in Fig. 33.
Fig. 33 Phylogenetic tree generated from Maximum Likelihood (ML) analysis based on
combined ITS and LSU sequence data of family Tetraplosphaeriaceae. Bootstrap support
values for maximum likelihood (ML) and maximum parsimony (MP) greater than 50 % and
Bayesian posterior probabilities greater than 0.75 are indicated above or below the nodes as
MLBS/MPBS/PP. The ex-type strains are in bold; the new isolates are in blue. The tree is
rooted with Massarina arundinariae.
Polyplosphaeria Kaz. Tanaka & K. Hiray.
The genus was introduced by Tanaka et al. (2009) to accommodate
Polyplosphaeria fusca Kaz. Tanaka & K. Hiray. The asexual morph of
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Polyplosphaeria produces globose to subglobose conidia with several setose
appendages (Tanaka et al. 2009). Only one species was accepted in this genus, viz. P.
fusca Kaz. Tanaka & K. Hiray.
275. Polyplosphaeria thailandica C.G. Lin, Yong Wang bis & K.D. Hyde, sp. nov.
Index Fungorum number: IF 551791, Facesoffungi number: FoF 01676, Fig. 34
Etymology: Referring to the country where the fungus was first collected.
Holotype: MFLU 15–3273
Saprobic on bamboo culms. Mycelium superficial. Sexual morph Undetermined.
Asexual morph Conidiophores absent. Conidiogenous cells monoblastic. Conidia
solitary, dry, acrogenous, muriform, globose, obovoid, pyriform, ellipsoidal,
occasionally two conidia associated together at the basal cell, brown, 20.5–43 µm
long excluding the appendages, 17.5–54 µm wide at the broadest part, verrucose; with
2–5 appendages, grey to brown, straight, septate, 23–117 µm long, 2–4.5 µm thick,
rounded at the apex; basal cell usually cylindrical, obconical, dark brown,
smooth-walled.
Culture characteristics: Colonies on PDA slow growing, attaining a diam. of
0.5–0.8 cm at room temperature (25°C) in 7 days, effuse, hairy, olive green to gray on
above, green to gray yellow from below.
Material examined: THAILAND, Phetchaburi, Cha-am District, Kao Yai, Khao
Nang Panthurat Forest Park, 12°49'48.5"N 99°57'05.5"E, on decaying bamboo, 28
July 2015, Chuan-Gen Lin, KNP 8-2 (MFLU 15-3273, holotype; GZAAS 16-0001,
isotype); ex-type living culture, MFLUCC 15-0840, GZCC 16-0001.
Notes: This species belongs to family Tetraplosphaeriaceae, and its placement is
supported by morphological and phylogenetic analysis. Phylogenetic analysis of ITS
and LSU sequence data indicates that our new species belongs in the genus
Polyplosphaeria (Fig. 33). It differs from P. fusca Kaz. Tanaka & K. Hiray which has
globose to subglobose, 43–100(–125) µm diam. conidia (Tanaka et al. 2009; Hyde et
al. 2013).
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Fig. 34 Polyplosphaeria thailandica (holotype) a Host (decaying bamboo) b, c
Conidiophores on the host surface d–g Conidiophores, conidiogenous cell and conidia h
Germinating conidium i, j Colonies on PDA culture. Scale bars: b = 200 µm, c = 100 µm, d–h
= 20 µm.
Pleosporales suborder Massarineae, incertae sedis
Massarinaceae
The suborder was treated by Tanaka et al. (2015*) and this is followed here.
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276. Longiostiolum Doilom, Ariyawansa & K.D. Hyde, gen. nov.
Index Fungorum number: IF 551899, Facesoffungi number: FoF 01881
Etymology: Name refers to the long ostiole.
Saprobic on dead bark of Tectona. Sexual morph Ascostromata black, solitary
to gregarious, scattered, immersed to semi-immersed, locules visible as white
contents, uniloculate, globose to subglobose, with a central ostiole. Ostiole long,
circular, central, periphysate. Peridium comprising two types of cell layers, outer
layer black to brown, thick-walled cells of textura angularis, inner layer composed of
hyaline and thin-walled cells of textura angularis. Hamathecium comprising
numerous, hypha-like, filiform, septate, branched, cellular, pseudoparaphyses. Asci
8–spored, bitunicate, clavate, apically rounded with ocular chamber. Ascospores
mostly overlapping biseriate to 3-seriate, hyaline when young later pale brown, fusoid
to narrowly fusoid, with narrowly rounded ends, constricted at the center septa, with
7–10 transverse septa, smooth-walled. Asexual morph Undetermined.
Type species: Longiostiolum tectonae Doilom, Ariyawansa & K.D. Hyde
277. Longiostiolum tectonae Doilom, D.J. Bhat & K.D. Hyde, sp. nov.
Index Fungorum number: IF 551900, Facesoffungi number: FoF 01882, Figs 35,
36
Etymology: Name refers to the host genus Tectona.
Holotype: MFLU 15–3532
Saprobic on dead bark of Tectona grandis L.f. Sexual morph Ascostromata
(255–) 295–375 (–500) µm high × (230–) 275–335 (–385) µm diam. (x̅ = 340 × 300
µm, n = 10), black, solitary to gregarious, scattered, immersed to semi-immersed,
when cut horizontally, locules visible as white contents, uniloculate, globose to
subglobose, with a central ostiole. Ostiole 110–220 µm high, 100–170 µm diam.,
circular, long, central, periphysate. Peridium 58–85 µm thick, comprising two types of
cell layers, outer layer black to brown, thick-walled cells of textura angularis, inner
layer composed of hyaline and thin-walled cells of textura angularis. Hamathecium
comprising numerous, 1.8–2.9 µm wide, hypha-like, filiform, septate, branched,
cellular, pseudoparaphyses, embedded in a gelatinous matrix. Asci (105–) 135–150
(–195) × 22–33 µm (x̅ = 140 × 27 µm, n = 15), 8-spored, bitunicate, clavate, with a
short pedicel, apically rounded, with an ocular chamber. Ascospores (52–) 57–59
(–63) × 8–12 µm (x̅ = 57× 10 µm, n = 20), mostly overlapping biseriate to tri-seriate,
hyaline when young later pale brown, fusoid to narrowly fusoid, with narrowly
rounded ends, constricted at the central septum, slightly constricted at other septa,
with 7–10 transverse septa, smooth-walled. Asexual morph (see culture
characteristics).
Culture characteristics: Ascospores germinating on PDA within 24 h. Colonies
on MEA reaching 12–17 mm diam. after 7 days in the dark at 25 °C (x̅ = 14.1 mm, n
= 5), undulate, fluffy in the center of old mycelium plug, aerial, medium spare, flat or
effuse, initially white, becoming brown, grey (7D1) in the center and white (7A1) at
the edge from above, light brown (7D6–7D7) from below. Colonies producing yellow
to brown pigments on MEA and PDA. Mycelium 1–4.5 µm wide, white to pale brown,
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branched, septate. Conidia–like structures (3–) 6–8 (–11) × (4–) 6–7 (–9) µm (x̅ = 7 ×
6 µm, n = 30), produced on aerial mycelium, subglobose to ellipsoidal, aseptate,
initially hyaline, becoming olivacious brown and finally black, terminal and lateral,
thick-walled.
Material examined: THAILAND, Chiang Mai Province, Mae Tang District, on
dead bark of T. grandis (Lamiaceae), 22 May 2012, M. Doilom, (MFLU 15–3532,
holotype), ex-type living culture MFLUCC 12–0562, MKT 078, ICMP.
Notes: Longiostiolum is introduced as a monotypic genus in the suborder
Massarineae with L. tectonae as the type species. The genus has black, immersed to
semi-immersed, uniloculate, globose to subglobose ascostromata, with white contents,
with a long central ostiole and phragmosporous ascospores. Longiostiolum clearly
differs from other genera in suborder Massarineae based on phylogenetic analysis and
morphology. Although, in this study, the combined phylogeny of LSU, SSU, TEF1α
and RPB2 sequence data shows weak support, L. tectonae (isolate MFLUCC 12–0562)
however, grouped in a distinct lineage within the suborder Massarineae (Fig. 13).
Therefore, we introduce a new monotypic genus to accommodate the taxon.
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Fig. 35 Longiostiolum tectonae (holotype) a Ascostromata immersed in dead bark of
Tectona grandis b Ascostroma cut horizontally showing the white contents c
Peridium d Ascostroma in section e Pseudoparaphyses f, g Immature asci with
ascospores h Mature ascus i, j Ascospores. Notes: e–g, i stained with lactophenol
cotton blue. Scale bars: a = 500 µm, b = 200 µm, c, d = 100 µm, e = 10 µm, f–j = 20
µm.
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Fig. 36 Longiostiolum tectonae (holotype) on MEA a, b Colony on MEA after 7 days (a =
above view, b = below view) c Colony producing yellow pigment on MEA after 2 months d
Mycelia e–l Conidia–like structures. Notes: d, f stained with lactophenol cotton blue. Scale
bars: d, e, i = 20 µm, f, g = 10 µm, h, j–l = 5 µm.
Pseudodidymosphaeria Thambugala & K.D. Hyde
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Thambugala et al. (2015b) introduced Pseudodidymosphaeria, typified by P.
spartii (Fabre) Thambugala et al., and accommodated it in the family Massarinaceae.
In this paper a second species is introduced. The phylogenetic tree is presented in Fig.
13.
278. Pseudodidymosphaeria phlei Phukhamsakda, Camporesi, & K.D. Hyde, sp. nov.
Index Fungorum number: IF 551895, Facesoffungi number: FoF 01816, Fig.
37
Etymology: Names base on the host.
Saprobic on dead stem of Phleum pretense L. Sexual morph Ascomata
200–368 µm diam. (x̅ = 290.7 µm, n = 15), solitary, scattered or gregarious on host,
semi-immersed to superficial, globose to subglobose, base flattened, slightly tapering
to apex, lacking ostioles. Peridium 9–24 µm wide, composed of 2–3 wall layers, outer
layer of light brown to dark brown cells of textura prismatica, inner layer, 1–2 thin
gelatinous layers. Hamathecium comprising numerous, long, 2–5 µm wide (x̅ = 2.5
µm, n = 50), transversely septate, branched, cellular pseudoparaphyses, embedded in a
gelatinous matrix. Asci 60–100 × 10–20 µm (x̅ = 73.49 × 13.86 µm, n = 20), 8-spored,
bitunicate, fissitunicate, clavate to sub-cylindrical, short pedicellate, ocular chamber
clearly visible when immature. Ascospores 15–21 × 6–10 µm (x̅ = 16.8 × 7.5 µm, n =
50) bi-seriate or overlapping, ovoid to sub-oval, slightly narrow at the apex,
1-transversely septate, slightly constricted at the septa, mucilaginous sheath clearly
visible, immature spores hyaline, light brown to brown when mature, smooth-walled.
Asexual morph Undetermined.
Culture characteristics: Ascospore geminating on PDA within 48 hours, germ
tubes developed from both ends of the ascospores. Colonies on PDA reaching 30 mm
diam. after 4 weeks. Culture incubated at 16 ̊C, at first white, after 2 weeks pale green
from center and bottom of colonies. After four weeks olive-green. Colonies
morphology, umbonate, with dense mycelium, slightly papillate on the surface,
circular, with dentate margin.
Material examined: ITALY, Forlì-Cesena Province, Monte Fumaiolo –
Verghereto, on a dead stem of Phleum pretense (Poaceae), 31 July 2014, E.
Camporesi (MFLU 15–3281, holotype; isotype HKAS 91937), ex-type living culture,
MFLUCC 14–1061, KUMCC 15–0551.
Notes: Pseudodidymosphaeria phlei is introduced from vertical dead stems of
Phleum pretense L. (Poaceae). Pseudodidymosphaeria phlei is closely related to the
type species, P. spartii (Fabre) Thambugala et al., as in phylogenetic analysis they
form sister clades with high support values (100 % ML). Pseudodidymosphaeria phlei
nevertheless is distinct in having semi-immersed to superficial ascomata, larger
peridium cell walls, with 2–3 wall layers, and ascospores with less distinctly rounded
ends. Therefore, we introduce Pseudodidymosphaeria phlei as a new species.
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Fig. 37 Pseudodidymosphaeria phlei (holotype) a, b Appearance of ascomata on host surface
c Section throught ascoma on host d Section of peridum e Hyaline cellular pseudoparaphyses
f Immature asci g–h Mature asci i–l Ascospores m Ascospores stained in Indian ink to show
sheath. Scale bar: b = 200 µm, c = 100 µm, d = 50 µm, e–h, m = 20 µm, i–l = 10 µm.
Pleosporales genera, incertae sedis
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Fig. 38 Phylogram generated from maximum likelihood analysis based on combined LSU and
SSU sequence data of Pleosporales. Maximum likelihood bootstrap support values greater
than 50% are near the nodes. New isolates are in blue. The tree is rooted with Hysterium
angustatum CBS 236.34
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279. Clematidis Tibpromma, Camporesi & K.D. Hyde, gen. nov.
Index Fungorum number: IF 551867, Facesoffungi number: FoF 01813
Etymology: named for its occurrence on the host plant genus (Clematis)
Saprobic on Clematis vitalba L. in terrestrial habitats. Sexual morph Ascomata
solitary or scattered on the host surface, superficial, globose to subglobose, with
flattened base, ostiole in the center, black. Peridium composing several layers of
brown to dark brown, flattened pseudoparenchymatous cells arranged in a textura
angularis. Hamathecium of 1.3–1.7 µm wide, long, cylindrical, cellular, anastomosed,
guttulate, septate, pseudoparaphyses. Asci 8-spored, bitunicate, cylindrical to
cylindric-clavate, short pedicellate or sessile. Ascospores overlapping 2–3-seriate,
hyaline, fusiform, 1-septate in center, swollen with large guttules in each cell, lacking
a mucilaginous sheath.
Type species: Clematidis italica Tibpromma, Camporesi & K.D. Hyde
Notes: Clematidis italica is morphologically similar to Lophiotrema
(Lophiotrema nucula). Clematidis can be distinguished morphologically from
Lophiotrema nucula (Fr.) Sacc. 1878 by having fusiform, 1-septate, straight or
slightly curved and hyaline ascospores, but L. nucula has elliptic-fusiform brown
ascospores with 3-septa (Tanaka and Harada 2003). Clematidis italica is introduced as
new genus based on morphology and combined LSU and SSU sequence phylogenetic
support (Fig. 38).
280. Clematidis italica Tibpromma, Camporesi & K.D. Hyde, sp. nov.
Index Fungorum number: IF 551868, Facesoffungi number: FoF 01814, Fig. 39
Etymology: Name reflects the country, where this species was collected
Holotype: MFLU 14–0669
Saprobic on Clematis vitalba L. in terrestrial habitats. Sexual morph Ascomata
170–182 µm high × 137–168 µm diam. ( x = 174 × 149 μm, n = 5), superficial,
solitary or scattered on the host surface, globose to subglobose, with flattened base,
ostiole in the center, not easy to removed, black, without papilla. Peridium 14–20 μm
wide, composed of several layers of brown to dark brown, flattened
pseudoparenchymatous cells, arranged in a textura angularis. Hamathecium of
1.3–1.7 µm wide, long cylindrical, cellular, anastomosed, septate, pseudoparaphyses.
Asci 79–114 × 13–18 μm ( x = 93 × 15 μm, n = 15), 8-spored, bitunicate, cylindrical
to cylindric-clavate, rounded at the apex, short pedicellate or sessile. Ascospores
21–30 × 5–8 μm ( x = 26 × 6 μm, n = 20), overlapping 2–3-seriate, hyaline, fusiform,
straight or slightly curved, 1-septate in center, slightly constricted at the median septa,
swollen with large guttules in each cell, lacking a mucilaginous sheath, smooth-walled.
Asexual morph Undetermined.
Culture characteristics: on MEA reaching 2 cm diam. after 2 weeks at 16°C,
later with dense mycelium, with irregular colony, edge undulate, surface smooth with
raised elevation, white-gray; hyphae septate branched, grey, thin-walled.
Material examined: ITALY, Corniolino, Santa Sofia, Forlì-Cesena Province, on
dead stem of Clematis vitalba (Ranunculaceae), 2 March 2013, Erio Camporesi,
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IT1086 (MFLU 14–0669, holotype); ex-type living culture, MFLUCC 15–0084); Ibid.
(HKAS92499 bis, paratypes).
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Fig. 39 Clematidis italica (holotype) a Appearance of ascomata on host substrate. b
Section of ascoma c Section of peridium d Pseudoparaphyses e–h Ascus with minute
pedicel i–k Ascospores l Germinated spore. Scale bars: a = 200 µm, b = 50 µm, c = 10
µm, d = 2 µm, e–h = 20 µm, i–l = 5 µm
281. Crassiparies Matsumura, K. Hiray & Kaz. Tanaka, gen. nov.
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MycoBank number: MB 815294, Facesoffungi number: FoF 02024
Etymology: From the Latin crassi-, meaning thick, and paries, meaning wall, in
reference to the thick ascomatal wall.
Saprobic on dead twigs of Acer sp. Sexual morph Ascomata scattered,
sometimes in groups of 2–3, immersed to superficial, hemisphaerical, ostiolate.
Peridium composed of 2 strata; outer stratum composed of brown, angular cells; inner
stratum composed of hyaline, prismatic cells. Hamathecium comprising numerous,
cellular, septate pseudoparaphyses. Asci 4-spored, fissitunicate, cylindrical to clavate,
pedicellate. Ascospores 1–2-seriate, hyaline, broadly fusiform, straight, thick-walled,
with a submedian septum, 1-septate, smooth-walled. Spermatia subglobose to elliptic,
hyaline, smooth-walled. Asexual morph Undetermined.
Notes: Crassiparies is similar to Massarina typified by M. eburnea (Tul. & C.
Tul.) Sacc. in that both have cylindrical, bitunicate asci and broadly fusiform,
1-septate, hyaline ascospores (Bose 1961; Aptroot 1998). Crassiparies, however,
differs from Massarina in having thick ascomatal walls, ascomatal necks without
clypei, and 4-spored asci. Massarina belongs to Massarinaceae, Massarineae (Hyde
et al. 2013), but Crassiparies nests between Massarineae and Pleosporineae (Fig.
40).
In phylogenetic analysis based on a combined dataset of SSU and LSU sequence
data, this genus formed a sister clade to Medicopsis (Fig. 40). However, sequence
similarity of ITS region between Crassiparies and Medicopsis romeroi (Borelli)
Gruyter et al., the type species of Medicopsis (CBS 252.60) was rather low (426/480
= 88.8 %), with 1.7 % gaps (8/480). Crassiparies occurs on woody plants (Acer),
while Medicopsis is known as a human pathogen (Borelli 1959; Ahmed et al. 2014).
Therefore, Crassiparies is introduced as a new genus.
Type species: Crassiparies quadrisporus Matsumura, K. Hiray. & Kaz. Tanaka
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Fig. 40 Maximum-likelihood tree of Crassiparies based on analysis of combined SSU and
LSU sequence data of Pleosporales. Bootstrap values greater than 50 % are presented at the
nodes. The ex-types are in bold. New species is annotated in blue.
282. Crassiparies quadrisporus Matsumura, K. Hiray. & Kaz. Tanaka, sp. nov.
MycoBank number: MB 815295, Facesoffungi number: FoF 02025, Fig. 41
Etymology: In reference to the 4-spored asci.
Holotype: HHUF 30409
Saprobic on dead twigs of Acer sp. Sexual morph Ascomata 300–590 µm high,
400–820 µm diam., scattered, sometimes in groups of 2–3, immersed to superficial,
hemisphaerical in section, with a central ostiole. Peridium 63–125 µm thick at the
base, 75–150 µm thick at sides, composed of 2 strata; outer stratum composed of
brown, angular cells (7.5–11 × 5–10 µm); inner stratum composed of hyaline,
prismatic cells. Hamathecium comprising numerous, 2–3 µm wide, septate, branched,
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cellular pseudoparaphyses. Asci 87–110(–124.5) × 17.5–22.5 µm ( x = 101.3 × 20.3
µm, n = 20), 4-spored, fissitunicate, cylindrical to clavate, pedicellate [(17–)22.5–37.5
µm long]. Ascospores 27–37 × 9–15 µm ( x = 31.4 × 12 µm, n = 30), L/W (2–)2.4–3
( x = 2.6, n = 30), 1–2-seriate, hyaline, broadly fusiform, straight, thick-walled, with
a septum mostly submedian (0.48–0.56; x = 0.52, n = 30), 1-septate, smooth-walled.
Spermatia 3–5.5 × 2–2.5 µm, subglobose to elliptic, hyaline, smooth-walled. Asexual
morph Undetermined.
Material examined: JAPAN, Mie, Tsu, Mie University, on dead twigs of Acer sp.,
30 May 2008, collector K. Tanaka, KH 111 (HHUF 30409, holotype); ex-type living
culture, MAFF 245408.
Fig. 41 Crassiparies quadrisporus a, b Appearance of ascomata on host surface c Ascoma in
longitudinal section d Peridium e Pseudoparaphyses f, g Asci h–k Ascospores l Germinating
ascospore m Spermogonia formed in culture n Spermatia a–l from HHUF 30409 (holotype);
m, n from culture MAFF 245408 (ex-holotype). Scale bars: a, b, m = 500 µm, c = 100 µm,
d–l, n = 10 µm.
283. Farasanispora Abdel-Wahab, Bahkali & E.B.G. Jones, gen. nov.
Index Fungorum number: IF 551712, Facesoffungi number: FoF 01634
Etymology: In reference to the Farasan Island where it was recorded.
Saprobic on submerged mangrove wood. Sexual morph Ascomata globose to
subglobose, immersed to erumpent, solitary, ostiolate, papillate, coriaceous,
dark-brown to black. Peridium thick at the upper part, two-layered; outer layer
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comprising polygonal, brown to dark-brown, thick-walled cells; inner layer 12–15 µm
wide, comprising hyaline, thin-walled, flattened cells, hard to distinguish from the
host cells. Hamathecium comprising numerous, 1.5–3 µm wide, septate, branched,
trabeculate pseudoparaphyses, within a gelatinous matrix, anastomosing above asci
and emerging through the ostiolar canal. Asci 8-spored, bitunicate, fissitunicate,
clavate, short pedicellate, apically rounded, with an ocular chamber. Ascospores
overlapping biseriate, hyaline, 1-septate, senescent ascospores light brown, flattened,
striate, rough, 2–3-septate. Asexual morph Undetermined.
Notes: During an ongoing study of marine fungi from Saudi Arabia (Hodhod et
al. 2012; Abdel-Wahab et al. 2014) an undescribed Massarina-like fungus was
recorded on decaying intertidal wood of Avicennia marina from Farsan Island
mangroves. Phylogenetic analyses of SSU and LSU sequence data placed the new
taxon in the order Pleosporales with affinities to the marine families:
Trematosphaeriaceae, Ascocylindricaceae and Morosphaeriaceae however, it did not
group with any known family and form a distant clade and it is described in here as a
new genus and species (Fig. 31). The genus Farasanispora closely resembles species
of Massarina in having hyaline, 1-septate ascospores, that become light brown and
rough-walled when senescent (Aptroot 1998). The genus Massarina is polyphyletic
and several new genera have been named to accommodate Massarina species, e.g.,
Halomassarina to accommodate M. thalassiae Kohlm. & Volkm.-Kohlm. (Suetrong
et al. 2009); Lindgomyces to accommodate M. ingoldiana Shearer & K.D. Hyde
(Hirayama et al. 2010); Morosphaeria to accommodate M. ramunculicola K.D. Hyde
and M. velatispora K.D. Hyde & Borse (Suetrong et al. 2009).
Type species: Farasanispora avicenniae Abdel-Wahab, Bahkali & E.B.G. Jones
284. Farasanispora avicenniae Abdel-Wahab, Bahkali & E.B.G. Jones, sp. nov.
Index Fungorum number: IF 551713, Facesoffungi number: FoF 01635, Fig. 42
Etymology: In reference to the host, Avicennia marina.
Holotype: CBS
Saprobic on submerged intertidal mangrove wood. Sexual morph Ascomata
180–270 µm in diam., globose to subglobose, immersed to erumpent, ostiolate,
solitary, coriaceous, dark-brown to black. Peridium 25–35 µm thick at the upper part,
two-layered, forming textura angularis; outer layer 10–15 µm comprising polygonal,
brown to dark-brown thick-walled cells; inner layer 12–15 µm wide, comprising
hyaline thin-walled flattened cells; peridium at the lower part of the ascomata is
one-layered, hyaline to light brown comprising of 10–15 µm diam. polygonal
flattened cells. Hamathecium comprising numerous, 1.5–3 µm wide, septate,
trabeculate pseudoparaphses, branched, within a gelatinous matrix, anastomosing
above the asci and emerging through the ostiolar canal. Asci 115–162 × 23–34 µm ( x
= 37.2 × 29.3 µm, n = 25), 8-spored, bitunicate, fissitunicate, clavate, short
pedicellate, apically rounded, with an ocular chamber. Ascospores 30–39 × 9–13 µm
( x = 34.9 × 11.4 µm, n = 60), overlapping biseriate, hyaline, 1–septate, the septum is
sub-median, upper cell longer and wider, slightly curved, guttulate; senescent
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ascospores are larger 38–43 × 11–14 µm ( x = 40.5 × 12.5 µm, n = 15), light brown,
flattened, striate, verrculose, 2–3-septate. Asexual morph Undetermined.
Culture characteristics: Colonies on PDA reaching a 25–30 mm radius after 22
days at 25°C, with white to gray aerial and immersed mycelium, from below brown.
Material examined: SAUDI ARABIA, Jizan City, Farasan Island, 16° 44′ 22′′ N
42° 4′ 41′′ E, on decayed wood of Avicennia marina at a mangrove stand, 8 March
2012, M.A. Abdel-Wahab (CBS, holotype); ex-type living culture, MF 1207.
Notes: Farasanispora avicenniae has ascospore dimensions that overlap with
Halomassarina thalassiae (Kohlm. & Volkm.-Kohlm.) Suetrong et al., however,
Farasanispora avicenniae have smaller ascomata without a clypeus or papillae and
the ostiolar canal is not periphysate. Ascospores in Halomassarina thalassiae has a
prominent and larger gelatinous sheath (Kohlmeyer and Volkmann-Kohlmeyer 1987).
Phylogenetically H. thalassiae and Farasanispora avicenniae are distantly related,
where the latter formed a basal clade to the families Morosphaeriaceae and
Trematosphaeriaceae and its phylogenetic placement is not well-resolved (Fig. 31).
Fig. 42 Farasanispora avicenniae (holotype) a, b Vertical section of ascomata c Ascus
dehiscence d–e Mature asci f Senescent ascospore. Scale bars: a–b = 40 µm, c = 15 µm, d–e =
20 µm, f = 5 µm.
285. Parameliola Hongsanan, Peršoh & K.D. Hyde, gen. nov.
Index Fungorum number: IF 551765, Facesoffungi number: FoF 01664
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Etymology: From Greek Para meaning near or beside, meliola is from the genus
name, in reference to the occurrence on Meliolaceae.
Hyperparasite on the surface of hyphae of Meliola thailandicum Hongsanan &
K.D. Hyde. On superficial hyphae of M. thailandicum, growing on the lower surface
of living leaves, branched, septate, darker at the septum, brown to dark brown, with
hyphopodia, later forming outwardly radiating black colonies with capitate
hyphopodia, mostly alternate or sometimes opposite on hyphae, near to hyphal septum,
2-celled, brown and hyphal setae 5 µm diam., aseptate, brown to reddish brown, pale
brown to hyaline at the apex. Conidiomata of Parameliola superficial, solitary,
globose to subglobose, attached to the superficial hyphae of Meliola thailandicum,
ostiole absent, thin-walled, brown to dark brown. Peridium comprising 2 layers of
textura angularis, inner layer very thin and hyaline, outer layer dark brown.
Hamathecium lacking pseudoparaphyses.
Conidiophores not observed.
Conidiogenous cells holoblastic in cavity of conidiomata, cylindrical, hyaline,
smooth-walled. Conidia borne singly at the apex of the conidiophore, ellipsoid to
cylindrical, both ends broadly rounded, aseptate, hyaline, smooth-walled.
Notes: Parameliola was found on the surface of leaves based of a black sooty
mould collected in northern Thailand. The species develops among the setae and on
the hyphae of Meliola thailandicum Hongsanan & K.D. Hyde, as a hyperparasite. The
morphology of Parameliola is typical of Coniothyrium in having globose, black
conidiomata and unicellular hyaline conidia. It is distinct from Coniothyrium and
other genera in Pleosporales in being hyperparasitic on the thallus or hyphae of
Meliola species. DNA extraction of Parameliola dimocarpi and P. acaciae were made
directly from dry fruiting bodies which contained many conidia to obtain sequence
data. Molecular analyses of LSU and SSU sequence data indicate that these two
species are separated from other known genera in Pleosporales. Therefore,
Parameliola should be a new genus in Pleosporales, typified by P. dimocarpi.
Furthermore, Parameliola species do not clusterd in any family of Pleosporales in
phylogenetic tree. More collections are needed to confirm their placement which is
possibly a new family in Pleosporales.
Type species: Parameliola dimocarpi Hongsanan & K.D. Hyde
286. Parameliola dimocarpi Hongsanan & K.D. Hyde, sp. nov.
Index Fungorum number: IF 551927, Facesoffungi number: FoF 01962, Fig. 43
Etymology: dimocarpi referring to the host.
Holotypus: MFLU15–0045
Hyperparasite on the surface of hyphae of Meliola thailandicum. Conidiomata
90–98 µm diam. ( x = 96 µm, n = 10), superficial, solitary, globose to subglobose,
attached to the superficial hyphae of M. thailandicum, ostiole absent, thin-walled,
brown to dark brown. Peridium 7–10 µm ( x = 8 µm, n = 10), comprising cell layers
of textura angularis, inner layer hyaline, outer layer dark brown. Hamathecium
lacking pseudoparaphyses. Conidiophores reduesed to conidiogenous cells.
Conidiogenous cells 5–4 × 2–3 µm ( x = 4.5 × 3 µm, n = 5), holoblastic in cavity of
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conidiomata, cylindrical, hyaline, smooth-walled. Conidia 6–9 × 2–3 µm ( x = 7 ×
2.5 µm, n = 10), borne singly at the apex of the conidiophore, ellipsoid to cylindrical,
both ends broadly rounded, aseptate, hyaline, smooth-walled.
Material examined: THAILAND, Chiang Rai, Amphoe Thoeng, on the living
leaves of Dimocarpus longan Lour. (Sapindaceae), 18 January 2015, S. Hongsanan
(MFLU15–0045 holotype; KIB, isotype).
Fig. 43 Parameliola dimocarpi (holotype) a, b Conidiomata developing as hyperparasites on
the thallus or hyphae of Meliola thailandicum c Hyphae of M. thailandicum with hyphopodia
d Section through conidioma in 10% lactic acid e Peridium of conidiomata f Conidiogenous
cell g Conidia in 10% lactic acid Scale bars: c–e = 10 µm, f, g = 5 µm
287. Parameliola acaciae Hongsanan & K.D. Hyde, sp. nov.
Index Fungorum number: IF 551928, Facesoffungi number: FoF 01963, Fig. 44
Etymology: acaciae referring to the host.
Holotypus: MFLU15–0378
Hyperparasite on the surface of hyphae of Meliola thailandicum. Conidiomata
84–88 µm diam. ( x = 85 µm, n = 10), superficial, solitary or gregarious, globose to
subglobose, attached to the superficial hyphae of Meliola thailandicum, ostiole absent,
thin-walled, brown to dark brown. Setae 5 µm diam., aseptate, brown to reddish
brown, pale brown to hyaline at the apex. Peridium 10 µm ( x = 8 µm, n = 10),
comprising 2 layers of textura angularis, inner layer hyaline, outer layer dark brown.
Hamathecium lacking pseudoparaphyses. Conidiophores reduced to conidiogenous
cells. Conidiogenous cells 4–5 × 1–2 µm ( x = 4.5 × 1.5 µm, n = 5) wide, holoblastic
in cavity of conidiomata, cylindrical, hyaline, smooth-walled. Conidia 7–10 × 3–4 µm
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( x = 9 × 3.5 µm, n = 10), borne singly at the apex of the conidiophore, cylindrical to
oblong, both ends broadly rounded, aseptate, hyaline, smooth-walled.
Material examined: THAILAND, Chiang Rai, Mueang, Agricultural Research
Center, on living leaves of Acacia auriculiformis A. Cunn. ex Benth. (Fabaceae), 23
January 2015, S. Hongsanan (MFLU 15–0378; KIB, isotype).
Notes: Parameliola acaciae is similar to P. dimocarpi, however, it differs in
having cylindrical to oblong ascospores, which are slightly larger than those of P.
dimocarpi. Parameliola acaciae was found among the colonies of Meliola
thailandicum growing on dead leves of Acacia auriculiformis, while Parameliola
dimocapi was found among the colonies of Meliola thailandicum growing on leaving
leaves of Dimocarpus longan (Hongsanan et al. 2015). Phylogenetic analyses indicate
that Parameliola. acaciae is closely related to the type species of Parameliola, but is
a distinct species, therefore the placement of Parameliola in Pleosporales is
supported.
Fig. 44 Parameliola acaciae (holotype) a, b Conidiomata developing as hyperparasites on
the thallus or hyphae of Meliola thailandicum c Hyphae of M. thailandicum with hyphopodia
d Section through conidiomata in 10% lactic acid e Conidiogenous cell f Conidia. Scale bars:
a, b = 100 µm, c = 10 µm, d = 50 µm, e, f = 5 µm.
Dothideomycetes family, incertae sedis
Kirschsteiniotheliaceae
Boonmee et al. (2012) established the new family Kirschsteiniotheliaceae based
on morphological features and phylogenetic analysis. The family is typified by
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Kirschsteiniothelia aethiops (Berk. & M.A. Curtis) D. Hawksw. and its asexual morph
is Dendryphiopsis atra (Corda) S. Hughes (Kirk et al. 2008; Su et al. 2016), and
Wijayawardene et al. (2014b) proposed the correct name for the type species as
Kirschsteiniothelia atra (Corda) D. Hawksw. Two species of Kirschsteiniothelia, K.
elaterascus Shearer and K. maritima (Linder) D. Hawksw. have been transferred to
Morosphaeria (Morosphaeriaceae) and a new genus Halokirschteiniothelia
(Mytilinidiaceae) by Boonmee et al. (2012) respectively. Kirschsteiniothelia comprises
19 species according to Index Fungorum (2016). Kirschsteiniothelia tectonae is
introduced as a new species in Kirschsteiniotheliaceae. The phylogenetic tree is
presented in Fig. 45.
Fig. 45 Phylogram generated from combined LSU, SSU and ITS sequence data. The tree is
rooted to Dothidea insculpa CBS 189.58. Maximum parsimony bootstrap values ≥50%,
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2623
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Bayesian posterior probabilities ≥ 0.95, (MPBS/PP) are given at the nodes. The ex-type
strains are in bold and the new isolates are in blue.
288. Kirschsteiniothelia tectonae Doilom, D.J. Bhat & K.D. Hyde, sp. nov.
Index Fungorum number: IF 551992, Facesoffungi number: FoF 01883, Fig. 46
Etymology: Name refers to the host genus Tectona on which the fungus was
collected.
Holotype: MFLU 15–1883.
Saprobic on dead branches and twigs of Tectona grandis (L. f.). Sexual morph
Undetermined. Asexual morph Colonies on natural substrate, superficial, hairy, dark
brown, scattered, single or in groups. Conidiophores up to 200 µm long, 4–8 µm wide at
the swollen base, superficial on host surface, macronematous, mononematous, simple,
erect to slightly curved, unbranched or branched, septate, slightly constricted at septa,
pale brown to dark brown, cylindrical. Conidiogenous cells 7.5–9.5 × 3.5–5 µm,
monoblastic,
integrated,
terminal,
cylindrical,
determinate.
Conidia
(85–)135–150(–212) × (15–)16–17(–19) µm ( x = 137 × 16 µm, n = 30), 9–25 or more
transverse septa, cylindric-obclavate, elongate, straight or slightly curved, rounded at
the apex and slightly paler, with sheath at apex, obconically truncate at the base, dark
reddish brown, thick–walled, smooth; secession schizolytic.
Culture characteristics: Conidia germinating on PDA within 24 h. Colonies on
MEA reaching 14–16 mm diam. after 7 days in the dark at 25 °C ( x = 14.9 mm, n = 5),
entire edge, circular, flat or effuse, raised at the edge, superficial at the center, dense,
fluffy, grey (5E1) from above, brownish (5F2) from below. Mycelium 1.5–4.7 µm wide,
aerial, reddish brown to dark brown, septate, branched hyphae, slightly constricted at
septa. Conidiophores up to 45 µm long, 3.5–8 µm wide, semi-macronematous,
mononematous, erect to slightly curved, indeterminate, branched, reddish brown to
dark brown. Conidiogenous cells holoblastic, doliiform, integrated, terminal. Conidia
(33–) 70–110 (–200) μm long × (7–) 11–13 (–18) µm thick at the broadest part ( x = 83
× 12 µm, n = 30), produced on aerial mycelium, initially subglobose and acellular,
becoming cylindric-obclavate, 1–29 or more transverse septa, flexuous, slightly curved,
rounded at the apex and slightly paler, obconically truncate at the base, dark reddish
brown, thick-walled.
Material examined: THAILAND, Phrae Province, Denchai District, Ban Maejour
Subdistrict, on dead branches of Tectona grandis (Lamiaceae), 29 October 2011, M.
Doilom (MFLU 15–1883, holotype), ex-type living culture MFLUCC 12–0050, MKT
016, MUCL55897; Chiang Rai Province, Mae Chan District, on dead twigs of T.
grandis, 3 March 2013, M. Doilom, MFLU 15–1884, living culture MFLUCC 13–0470,
MKT 111.
Notes: Kirschsteiniothelia tectonae was found only in its asexual morph, while K.
thujina is known only as the sexual morph. Thus, a morphological comparison could
not be made, and K. tectonae is only compared to K. atra, K. emarceis and K. lignicola.
These three species have been reported with asexual morphs both on natural substrates
and cultures. It differs from these species in size and shape of conidiophores and
conidia both on natural substrates and cultures. The conidia of K. tectonae are longer
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2641
than those of the other three species (Table. 2). Based on its morphology (Fig. 46) and
the fact it is phylogenetically separate from other species in Kirschsteiniothelia (Fig.
45), we introduce it a new species. The combined LSU, SSU and ITS sequence analysis
shows that K. tectonae isolate MFLUCC 12–0050 and MFLUCC 13–0470 grouped
close to, but is distinguishable from K. thujina with strong bootstrap support 100%
MPBS and 1.00 PP (Fig. 45).
Table. 2 Comparison of morphological characters of asexual morph of
Kirschsteiniothelia
Species
Morphology on natural substrate
Conidiophore
Conidia (µm)
s (µm)
Morphology on MEA culture
Conidiophore
Referenc
Conidia (µm)
e
s (µm)
K.
Up to 200, 4–8
(85–) 135–150
up to 45, 3.5–8
(33–) 70–110
This
tectonae
wide at the
(–212) long × (15–)
wide
(–200) long × (7–)
study
swollen base
16–17 (–19) thick
11–13 (–18) thick
in broadest part,
in broadest part,
9–25 or more
1–29 or more
transverse septa,
transverse septa,
cylindric–obclavate
cylindric–obclavat
, elongate
e
K.
Up to 500
aethiops
long, 8–11
40–80 × 12–25
Not reported
Not reported
Ellis 1971
(40–)45– 56(–67) ×
32–92 long,
(21–)27–28(–36) ×
Boonmee
thick.
K.
162–271 ×
emarcei
7–14
s
(10–)14–15(–17),
5–7 thick,
9– 13(–15),
et al.
3–4(–5) septate,
branched at
1–2(–3) transverse
2012
oblong to obclavate
apex
septate, fusiform
to obclavate
K.
287–406 ×
39–48(–52) ×
39–148 long,
24.5–35(–41) ×
Boonmee
lignicol
11–13
21–25(–28), 1–2
4–7 thick
14–16(–19), 1–2
et al.
transverse septa,
transverse septa,
2012
obovoid to broadly
broadly obovoid
a
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Fig. 46 Kirschsteiniothelia tectonae (holotype) a Conidia host surface (arrows) b–d Conidia
e Colony on MEA for 7 days (above and below views) f Colony on PDA for 2 months (above
and below views) g Mycelia h Immature conidia attached to conidiophore i Conidia j–m, o
Conidia attached to conidiophores with mycelia n, p Conidia attached to conidiophores a–d
Morphology on host g–p Morphology on MEA culture. Scale bars: a = 200 µm, b–d, g, j, n–p
= 20 µm, h, i, l, m = 10 µm, k = 50 µm.
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Lecanoromycetes
Ostropales
Graphidaceae
Graphidaceae is the second largest family of lichenized fungi, with
approximately 2,100 species in nearly 80 genera and an additional 1,800 species
predicted (Rivas Plata et al. 2012a; Lücking et al. 2014; Jaklitsch et al. 2016). Here
we described three new species of Graphidaceae discovered in the course of
collaborative inventory work in Sri Lanka and adjacent areas (Weerakoon 2015;
Weerakoon et al. 2012a, b, c, 2014, 2015; Weerakoon & Aptroot 2013, 2014). All
belong to the Ocellularia clade, a clade that has been recognized as hyper diverse in
recent molecular and revisionary studies (Rivas Plata et al. 2012b; Cáceres et al. 2014;
Lücking 2014, 2015; Kraichak et al. 2015), surpassing the genus Graphis and
relatives in species richness. Since Ocellularia and relatives are mostly found in
well-preserved tropical forests (Rivas Plata et al. 2008), it is predicted that the
remaining forest ecosystems still yield a high number of undiscovered species
(Lücking et al. 2014). This is also true for Sri Lanka, which has only begun to be
studied systematically with regard to its lichen biota (Weerakoon & Aptroot 2014;
Weerakoon 2015), but where tropical forest has largely been degraded, leaving a few
pristine, highly diverse areas. Although we were unable to generate molecular data for
the newly described species, our broad molecular framework of the family (Rivas
Plata et al. 2012b, 2013) has helped us to establish a much refined species concept in
the Ocellularia clade, leading to numerous recent discoveries (Lücking 2014, 2015;
Lücking & Pérez-Ortega 2015), including the three species described here.
Ocellularia
289. Ocellularia arachchigei Weerakoon, Lücking & Lumbsch, sp. nov.
MycoBank number: MB 815548, Facesoffungi number: FoF 02026, Fig. 47a
Etymology: In honor of the collector of the type specimen, Mr. Omal Selika
Arachchige.
Holotype: O. S. Arachchige 107A (F).
Diagnosis: Differing from Ocellularia papillata and O. rongklaensis in the grey
thallus with large internal clusters of calcium oxalate crystals and the whitish cover of
the columella.
Thallus corticolous, epiperidermal, up to 5 cm diam., continuous; surface smooth
to uneven, light grey; prothallus absent. Thallus in section 70–100 µm thick, with
prosoplectenchymatous cortex, 15–20 µm thick, photobiont layer 30–50 µm thick, and
medulla 30–50 µm thick, strongly encrusted with numerous clusters of calcium
oxalate crystals, thicker near apothecial margin (up to 100 µm). Photobiont
Trentepohlia; cells rounded to irregular in outline, in irregular groups, yellowish green,
8–10 5–7 µm. Ascomata immersed-erumpent, with complete thalline margin,
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2705
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0.8–1.2 mm diam.; disc covered by 0.2–0.5 mm wide pore; proper margin distinct,
entire, visible as thin, white rim around the pore; thalline margin entire, smooth, white.
Excipulum entire, yellowish to orange-brown (difficult to separate from modified
periderm), together with periderm 50–100 µm wide, fused with thalline margin;
laterally covered by algiferous, corticate thallus containing periderm and large crystals
of calcium oxalate crystal layers up to 100–150 µm. Columella present, finger-like to
barrel-shaped, becoming irregular, 150–200 µm broad, yellowish brown with whitish
cover. Hypothecium prosoplectenchymatous, 10–15 µm high, colourless. Hymenium
150 µm high, colourless, clear. Epithecium indistinct, 10–15 µm high, colourless.
Paraphyses unbranched, apically smooth; periphysoids absent. Asci cylindrical,
120–140 20–25 µm. Ascospores 8 per ascus, ellipsoid, 7–9-septate, 30–35 9–10
µm, 3–4 times as long as wide, hyaline, distoseptate with lens-shaped lumina, I+
violet-blue. Secondary chemistry: No substances detected by TLC.
Material examined: SRI LANKA, Central Province, Matale District,
Gammaduwa; 7° 31' N, 80° 40' E, 360 m; low altitude, on tree bark of home garden;
January 2015, O. S. Arachchige 107A (PDA holotype and F Isotype).
Distribution and ecology: The new species was collected from a home garden in
the central region of Sri Lanka. It is thus far only known from the type locality.
Notes: This new species keys out close to Ocellularia papillata (Leight.) Zahlbr.
and O. rongklaensis (Homchant. & Coppins) Lücking. All three agree in lacking
secondary metabolites, having a non-carbonized excipulum, a smooth to uneven
thallus, immersed to erumpent ascomata, and ascospores over 20 µm long. However,
O. papillata differs in the pale olive thallus lacking large clusters of calcium oxalate
crystals, the completely immersed apothecia, and the distinctly brown apothecial rim
and columella, with the columella becoming more distinctly irregular. In contrast, O.
ronklaensis has a pale olive, indistinctly verrucose thallus, due to clustered
distribution of calcium oxalate crystals, more erumpent apothecia, and the columella
appears dark with only a thin whitish pruina. Other similar species are O.
laeviusculoides Sipman & Lücking, differing chiefly in its carbonized columella, and
O. bonplandii (Fée) Müll. Arg. and O. auberianoides (Nyl.) Müll. Arg., which both
produce protocetraric acid and the columella becomes distinctly irregular in the latter.
290. Ocellularia ratnapurensis Weerakoon, Lücking & Lumbsch, sp. nov.
MycoBank number: MB 815549, Facesoffungi number: FoF 02027, Fig. 47b
Etymology: Referring to the type locality.
Holotype: G. Weerakoon 1005 (F).
Diagnosis: Differing from Ocellularia guptei in the larger ascospores and the
only partially (upper half) carbonized columella.
Thallus corticolous, epiperidermal, up to 5 cm diam., continuous; surface uneven
to verrucose, brownish yellow; prothallus absent. Thallus in section 50–80 µm thick,
with paraplectenchymatous cortex, 5–10 µm thick, photobiont layer 30–60 µm thick,
and medulla 30–50 µm thick, strongly encrusted with clusters of calcium oxalate
crystals, near apothecial margins much thicker, up to 150 µm; in addition with
numerous small, grey granules. Photobiont Trentepohlia; cells rounded to irregular in
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outline, in irregular groups, yellowish green, 8–11 5–8 µm. Ascomata rounded,
erumpent to prominent, with complete thalline margin, 0.7–1.2 mm diam., 0.2–0.3
mm high; disc covered by 0.15–0.25 mm wide pore; proper margin distinct, entire,
visible as brownish rim around the pore; thalline margin entire, smooth, light
yellowish. Excipulum entire, yellowish to orange-brown, upper half carbonized,
50–70 µm thick, covered by periderm layer, 70–100 µm thick, orange, fused with
thalline margin. Columella present, finger-like to barrel-shaped, 100 µm broad, upper
half carbonized; hypothecium prosoplectenchymatous, 10–15 µm high, colourless.
Hymenium 300 µm high, colourless, clear; epithecium indistinct, 10–15 µm high,
colourless. Paraphyses unbranched, apically smooth; periphysoids absent. Asci
cylindrical, 200–250 50–70 µm. Ascospores 1–2 per ascus, muriform, 200–250
40–50 µm, about 5 times as long as wide, hyaline, distoseptate with lens-shaped
lumina, I+ violet-blue. Secondary chemistry: No substances detected by TLC.
Material examined: SRI LANKA. Sabaragamuwa Province: Ratnapura District,
Coolbone Tea Estate, on tree bark, 7° 02' N, 80° 23' E, 1288 (PDA holotype and F
Isotype).
Distribution and ecology: The new species was collected from montane forest
patches in a Tea estate in the Sabaragamuwa region of Sri Lanka. It is thus far only
known from the type locality.
Notes: Ocellularia ratnapurensis belongs in a small group of species with
carbonized excipulum and columella, large, muriform ascospores, and absence of
secondary substances. Among these, the neotropical O. sanfordiana Zahlbr. differs by
the carbonization of the excipulum and columella reaching down to the base, the
larger apothecia, and the smaller ascospores (130–170 25–35 µm). The
paleotropical Ocellularia kalbii Mangold also differs in the basal carbonization of
excipulum and columella and in addition has less erumpent apothecia with gently
sloping sides and much longer ascospores (300–600 25–50 µm). Ocellularia guptei
(Nagarkar, Sethy & Patw.) D. D. Awasthi, from India, apart from a fully carbonized
columella, differs in its smaller ascospores (100–180 15–30 µm). All other similar
species differ in their chemical components, mostly producing hypoprotocetraric or
isonotatic and norisonotatic acid.
Rhabdodiscus
291. Rhabdodiscus albodenticulatus Weerakoon, Lücking & Lumbsch sp. nov.
MycoBank number: MB 815550, Facesoffungi number: FoF 02028, Fig. 47c, d
Etymology: Referring to the white teeth-like apothecial columella.
Holotype: G. Weerakoon 880 (F).
Diagnosis: Differing from Rhabdodiscus integer by the thicker, verrucose thallus
and the smaller, more immersed apothecia.
Thallus corticolous, up to 5 cm diam., continuous, olive-grey to olive- green,
uneven-verrucose; prothallus not observed. Thallus in section 200–300 µm thick, with
prosoplectenchymatous cortex 10–20 µm thick, photobiont layer 50–70 µm thick, and
medulla 150–200 µm thick, strongly encrusted with numerous large crystals of
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2801
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calcium oxalate, forming clusters that cause the verrucae. Photobiont Trentepohlia;
cells rounded to irregular in outline, in irregular groups, pale green, 7–11 4–6 µm.
Apothecia erumpent, 0.8–1.2 mm diam.; disc partially covered by 0.2–0.4 mm wide
pore, rim around the pore whitish to pale yellowish, pore mostly filled by columella.
Columella present, broad-stump-shaped but becoming ruptured in to 3–6 irregular
teeth, 100–200 µm broad, carbonized but with whitish top. Excipulum 30–50 µm
broad, carbonized; periphysoids absent. Hymenium 120 µm high, clear. Paraphyses
unbranched. Asci 100 20 µm, fusiform. Ascospores 8 per ascus, submurifom, 3
transverse septa and 0–1 longitudinal septa, 15–18 7–8 µm, ellipsoid, with thick
septa and lens-shaped lumina, brown, I+ violet-blue (amyloid). Secondary chemistry:
Psoromic, subpsoromic and 2'-O-demethylpsoromic acids.
Material examined: SRI LANKA, Central Province, Matale district,
Siyabalabokka-Rattota, 7° 31' N, 80° 40' E, 360 m, low altitude, on tree bark of home
garden; January 2015, G. Weerakoon 880 (PDA holotype and F Isotype); Along
Karagastanna road, 7° 34' N, 80° 42' E, 990 m, mid elevation, January 2015, G.
Weerakoon 205, 237 (F); Meepiliyamana -Nuwaraeliya, 6° 56' N, 80° 47' E, 1350 m,
high elevation, January 2015, Weerakoon 732 (F).
Distribution and ecology: The new species was collected in high elevation
disturbed montane forest patches.
Notes: This new species is most similar to Rhabdodiscus integer (Müll. Arg.)
Rivas Plata & Lumbsch, which which it shares the submuriform, brown ascospores,
the columella rupturing into teeth, and the psoromic acid chemistry. However, R.
integer has a thinner, smooth to uneven thallus and much larger, strongly prominent
apothecia. Rhabdodiscus marivelensis (Vain.) Rivas Plata & Lumbsch differs in the
minutely grainy thallus caused by columnar clusters of calcium oxalate crystals, the
thicker apothecial margin, and the larger ascospores (20–30 8–18 µm).
2809
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Fig. 47 Ocellularia arachchigei (holotype) a Thallus with ascomata. Ocellularia
ratnapurensis (holotype) b Thallus with ascomata. Rhabdodiscus albodenticulatus (c
holotype, d paratype) c, d Thallus with ascomata. Scale bars: a–d = 1 mm.
Sordariomycetes
Chaetosphaeriales
Chaetosphaeriales was established as distinct order in the Class Sordariomycetes
based on phylogenetic analysis of LSU sequence data (Huhndorf et al. 2004). At
present, two families, Chaetosphaeriaceae (Réblová et al. 1999) and
Helminthosphaeriaceae (Samuels et al. 1997) are recognized as members of this order
(Maharachchikumbura et al. 2015).
Chaetosphaeriaceae
Species of Chaetosphaeriaceae are widely distributed and are saprobic on
various plants (Fernández and Hundorf 2005). The representative genus
Chaetosphaeria is characterized by non-stromatic perithecia, cylindrical asci, and
transversely septate ascospore in its sexual morph, but the genus has been reported to
have morphologically diverse asexual morphs (Réblová and Winka 2000).
Phylogenetic studies also suggest that the genus is polyphyletic (Fernández et al. 2006;
Jeewon et al. 2009). To date, 32 asexual hyphomycetous genera have been reported in
this family (Wijayawardene et al. 2012). Additionally, eight coelomycetous genera,
Brunneodinemasporium,
Dendrophoma,
Dinemasporium,
Infundibulomyces,
Neopseudolachnella, Pseudodinemasporium, Pseudolachnea and Pseudolachnella are
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known as members of Chaetosphaeriaceae (Somrithipol et al. 2008; Crous et al.
2012; Wijayawardene et al. 2012; Hashimoto et al. 2015a, b; Liu et al. 2015).
Pseudolachnella
The genus Pseudolachnella was established by Teng (1936) to segregate species
with multi-septate conidia from Pseudolachnea. The generic concept has been
recently revised based on detailed morphological observations and molecular analysis
(Hashimoto et al. 2015b). It is characterized by setose condiomata with thin basal
stroma and less-developed excipulum, and condia bearing appendages. Sixteen
species of Pseudoalchnella have been described from bamboo (Nag Raj 1993; Zhao et
al. 2004; Sato et al. 2008; Hashimoto et al. 2015b), but P. guaviyunis occurred on
Myrcianthes pungens (Myrtaceae) (Crous et al. 2014b). The phylogenetic tree for
Pseudolachnella is presented in Fig. 48.
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Fig. 48 Maximum-likehood tree of Pseudolachnella spp. based on analysis of ITS sequence
data. Bootstrap values greater than 50 % are presented at the nodes. New taxa are in blue
extypes in bold.
292. Pseudolachnella brevifusiformis A. Hashim. & Kaz. Tanaka, sp. nov.
MycoBank number: MB 815299, Facesoffungi number: FoF 02029, Fig. 49
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Etymology: named after its resemblance to Pseudolachnella fusiformis, but with
smaller conidia.
Holotype: HHUF 30495
Saprobic on dead sheath of bamboo. Sexual morph: Undetermined. Asexual
morph: Conidiomata stromatic, acervular, setose, shallow-cupulate, superficial,
globose to oval, up to 295 µm high, (325–)450–700(–895) µm diam., scattered to
occasionally 2–5 grouped, conical in sectional view; basal stroma 6.5–15 µm thick,
composed of brown, globose, thick-walled, 2–2.5 µm diam. cells; excipulum
30–44.5(–50) µm thick, poorly developed, composed of globose, pale brown cells.
Setae marginal, cylindrical, straight to slightly curved, aseptate, brown to dark brown,
thick-walled, (315–)380–520 µm long, acute and 2–3.5 µm wide at the apex, 3–4 µm
wide at the base. Conidiophores absent. Conidiogenous cells phialidic, cylindrical to
lageniform, hyaline, smooth, 6.5–14 × 1.5–2.5 µm. Conidia (9.5–)10.5–18(–19) ×
2–3.5 µm ( x = 14 × 2.9 µm, n = 78), L/W 3.4–7.6(–8.7) ( x = 5, n = 78),
(1–)3-septate, clavate to cylindrical, obtuse at the apex, truncate at the base, hyaline,
smooth, bearing (2–)3–6 unbranched appendages at each end; apical appendage
(2.5–)3–6 µm long ( x = 4.3 µm, n = 61), central; basal appendage (2.5–)3–5.5(–6.5)
µm long ( x = 4 µm, n = 61), eccentric.
Material examined: JAPAN, Okinawa, Kunigami, Yona, Mt. Fuenchiji, on dead
sheath of Pleioblastus linearis, 19 May 2015, collector K. Tanaka et al., KT 3536
(HHUF 30495, holotype); ex-type living culture, MAFF 245411; ibid., KT 3537
(HHUF 30496, paratype); ex-paratype living culture, MAFF 245412.
Notes: In terms of the similar conidial size and multiple conidial appendages,
Pseudolachnella brevifusiformis resembles P. fusiformis, but can be distinguished
from the latter by its smaller conidia (vs. 15–20 × 4–6.5 µm; Hashimoto et al. 2015b)
in addition, there were 25 base differences with 12 gaps in their ITS sequence data.
Pseudolachnella brevifusiformis was collected from Pleioblastus linearis.
Pseudolachnella ryukyuensis was also recorded from same host plant (Hino and
Katumoto 1958; Nag Raj 1993). Morphologically, P. brevifusiformis has smaller
conidia, as compared with those of the latter (vs. 30–40 × 2.5–3 µm; Nag Raj 1993).
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Fig. 49 Pseudolachnella brevifusiformis a, b Appearance of conidiomata on substrate c
Conidioma in longitudinal section d Excipulum of conidioma e Conidiomatal setae f, g
Conidiogenous cells and immature conidia h–l Conidia m Germinating conidium a–d, i, k, m
from HHUF 30495 (holotype); e–h, j, l from HHUF 30496 (paratype). Scale bars: a = 1 mm,
b = 250 µm, c = 50 µm, d, e = 20 µm, f–h, m = 10 µm, i–l = 5 µm.
Diaporthales
Gnomoniaceae
The family Gnomoniaceae was established by Winter (1886) based on the genus
Gnomonia. Gnomoniaceae is simialr with Obryzaceae, which is considered to be a
lichenicolous family, while Gnomoniaceae is a well-known plant pathogenic family
(McNeill et al. 2006). Hawksworth and Eriksson (1988) proposed that the name
Obryzaceae should be rejected to conserve Gnomoniaceae and the proposal was
accepted (McNeill et al. 2006). Gnomoniaceae is characterised by immersed, rarely
erumpent or superficial astromatic ascomata, arranged solitary, or aggregated with a
rudimentary stroma, dark brown to black, and generally soft-textured, and
pseudoparenchymatous and thin-walled, with necks. Generally the asci have a distinct
apical ring (Sogonov et al. 2008). Species of this family are found in herbaceous plant
material, especially in leaves, twigs or stems, rarely in bark or wood (Sogonov et al.
2008).
Phragmoporthe Petr.
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2912
2913
2914
2915
The genus Phragmoporthe was introduced based on P. ploettneriana (Henn.)
Petr. as the type species (Petrak 1934). Phragmoporthe is characterised by
multi-septate ascospores and 8-spored asci (Sogonov et al. 2008). The closest genus to
Phragmoporthe is Ditopella, which differs from Phragmoporthe in having 1-septate,
rarely aseptate ascospores and polysporous asci (Sogonov et al. 2008). The
phylogenetic tree is presented in Fig. 50.
2916
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2918
2919
2920
2921
2922
2923
2924
2925
2926
Fig. 50 Maximum Likelihood tree resulting from analysis of combined LSU, ITS and TEF-1α
sequence data for taxa of the family Gnomoniaceae. Maximum likelihood bootstrap support
values greater than 50% are shown near the nodes. New taxa are in blue and ex-type strains in
bold. The tree is rooted with Valsella salicis and Leucostoma niveum.
293. Phragmoporthe conformis (Berk. & Broome) Petr., Annls mycol. 39(4/6): 285
(1941)
Facesoffungi number: FoF 01794, Fig. 51
Basionym
Sphaeria conformis Berk. & Broome, Ann. Mag. nat. Hist., Ser. 2 9: 325 (1852)
2927
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2933
2934
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2939
2940
2941
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2948
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2950
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2968
Synonym
= Gnomonia conformis (Berk. & Broome) Ferd. & Winge
= Metasphaeria conformis (Berk. & Broome) Sacc., Miscell. mycol. 1: 6 (1884)
= Calospora conformis (Berk. & Broome) Starbäck, Bih. K. svenska
VetenskAkad. Handl., Afd. 3 15(no. 2): 16 (1889)
= Sphaeria ditopa f. octospora Cooke
= Valsa alnicola Cooke & Massee, Grevillea 16(no. 78): 47 (1887)
= Calospora alnicola (Cooke & Massee) Sacc., Syll. fung. (Abellini) 9: 872
(1891)
= Phragmoporthe alnicola (Cooke & Massee) Petr., Annls mycol. 38(2/4): 209
(1940)
= Sphaerulina alni A.L. Sm., Trans. Br. mycol. Soc. 6(2): 151 (1918)
Saprobic on Alnus glutinosa L. Sexual morph Appearing as conical, pustules on
the host surface. Ascomata perithecial, minutely stromatic, immersed, erumpent.
Perithecia 700–770 µm diam. (n = 20), solitary, immersed in or directly below the
host epidermis, globose, membranous, dark brown to black, with a periphysate
ostiole. Peridium 14–38 µm (x̅ = 22 µm, n = 10) wide, comprising 7–15 cell layers,
outer layers heavily pigmented, thin-walled, comprising dark brown cells of textura
angularis, inner layers composed of hyaline to brown, thin-walled, flat cells of textura
angularis. Hamathecium lacking paraphyses. Asci 60–80 × 17–24 µm (x̅ = 72 × 19.5
µm, n = 30), 8-spored, unitunicate, clavate, straight, short pedicellate, apically
rounded or truncate, with a refractive, J- apical ring. Ascospores 19–24 × 6.5–8 µm (x̅
= 22 × 7 µm, n = 50), multi-seriate, fusiform, mainly with 3 transverse septa,
occasionally constricted at septum, hyaline, smooth and thick-walled, without a
mucilaginous sheath or appendages. Asexual morph Undetermined
Culture characteristics: Colonies growing on MEA, slow growing, reaching 4
cm diam. in 21d at 16 °C on MEA, white, dense, moderate aerial mycelium on the
surface, underneath similar in colour, margins even.
Material examined: ITALY, Forlì-Cesena Province, Lago Pontini-Bagno di
Romagna, dead branches of Alnus glutinosa (L.) Gaertn. (Betulaceae), 26 May 2014,
Erio Camporesi, IT 1892 (MFLU 15–2662 reference specimen designated here),
also in HKAS 92498, living culture, MFLUCC 14–0567.
Notes: The putatively named strain of Phragmoporthe conformis (CBS 109793)
clustered with our newly collected strain (MFLU 14–0567), collected from Italy, on a
dead a stem of Alnus glutinosa. Berkeley and Broome (1852) originally described
Phragmoporthe conformis as Sphaeria conformis on Alnus spp. from the UK. Later
Petrak (1941) synonymized Sphaeria conformis under Phragmoporthe conformis. The
ascomata, size of asci and ascospores of our strain are typical of P. conformis (Petrak
1941) and the molecular data is identical to CBS 109793. We therefore designate our
collection as a reference specimen of P. conformis to stabilize the taxonomy of the
genus.
2969
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2973
2974
Fig. 51 Phragmoporthe conformis (MFLU 15–2662, reference specimen) a, b Appearance
of ascomata on host substrate c Section of ascoma d Transverse section through ostiole e, f
Periphyses g Close up of peridium h–j Asci k Close up of apical ascus strained in Melzer’s
reagent l–n Ascospores o Germinating spore p, q Colonies growing on MEA. Scale bars: c =
500 µm, d, e = 100 µm, f–j = 50 µm, k = 20 µm, l–o = 10 µm.
2975
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3001
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Valsaceae
The family Valsaceae was introduced by Tulasne and Tulasne (1861) and placed
in Diaporthales by Barr (1978). Most of Valsaceae species are plant pathogens
causing canker and dieback disease, with damage to several economic crops
worldwide (Adams et al. 2005; Fan et al. 2014a, b, 2015a, b; Ariyawansa et al.
2015b). Valsaceae was restricted to Cytospora (asexual morph), Valsa, Leucostoma,
Valsella, and Valseutypella; sexual morph for the last four genera (Fries 1823;
Saccardo 1884; Gvritishvili 1982; Spielman 1985; Adams et al. 2002, 2005;
Castlebury et al. 2002; Bulgakov 2010; Yang et al. 2015). However, all sexual genera
were synonymized under Valsa as a subgenus or species without additional
infrageneric rank (Adams et al. 2005). According to the International Code of
Nomenclature for Algae, Fungi, and Plants (ICN) in 2011, a single name is needed for
a biological species and for genera, the older and more commonly encountered genus
Cytospora (1818) was chosen over that of its sexual morph, Valsa (1849), for
placement on the list of protected fungi (Adams et al. 2005; Fotouhifar et al. 2010,
Fan et al. 2015a; Wingfield et al. 2012; Crous et al. 2015e; McNeill et al. 2012;
Rossman et al. 2015). Cytospora is characterized by single or labyrinthine locules,
filamentous conidiophores (or clavate to elongate obovoid asci), and allantoid, hyaline
conidia (Spielman 1983, 1985; Adams et al. 2005). In moist conditions, conidia
emerge from the fruiting bodies as yellow masses, and become orange to red
gelatinous tendrils later (Adams et al. 2005, 2006). The genus Cytospora comprised
110 species (Kirk et al. 2008), however, 572 epithets are recorded in Index Fungorum
(2015). Ex-type sequence data, is however, available for a few species. Thus it is
difficult to identify species (Liu et al. 2015; Ariyawansa et al. 2015b). A systematic
account of the genus Cytospora is needed to clarify cryptic species in Cytospora
(Adams et al. 2002; Fotouhifar et al. 2010; Hyde et al. 2010, 2014; Fan et al. 2015a, b;
Liu et al. 2015; Ariyawansa et al. 2015b, Yang et al. 2015). The phylogenetic trees for
Cytospora are presented in Figs 52 and 53.
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3009
3010
3011
Fig. 52 Maximum Parsimony (MP) majority rule consensus tree for the analyzed Cytospora
isolates based on a combined dataset of ACT, ITS and LSU sequence data. MP bootstrap
support values higher than 50% and Bayesian posterior probabilities (PP) above 95%
(MP/PP). The tree is rooted with Diaporthe vaccinii (CBS 160.32). The strain numbers are
mentioned after the species names. The species obtained in this study is in blue bold and
ex-type strains in black bold.
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3013
3014
3015
3016
Fig. 53 Phylogenetic tree based on an alignment of the sequences of the ITS sequence data for
Cytospora, Leucostoma, and Valsa species, which was generated using the MP and Bayesian
posterior probabilities (PP) in PAUP. Numbers separated by a slash represent MP bootstrap
values >50% and Bayesian posterior probabilities (PP) above 95% are given at the nodes
3017
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3019
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3021
3022
3023
3024
3025
3026
3027
3028
3029
3030
3031
3032
3033
3034
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3037
3038
3039
3040
3041
3042
3043
3044
3045
3046
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3050
(MP/PP). The tree is rooted in outgroup taxon Diaporthe vaccinii (CBS 160.32). New strains
are in blue bold and ex-type strains are in black bold.
294. Cytospora salicicola C. Norphanphoun, Bulgakov & K.D. Hyde, sp. nov
Index Fungorum number: IF 551803, Facesoffungi number: FoF 01768, Fig. 54
Etymology: Named after the host genus on which the fungus occurs.
Holotype: MFLU 14–0785
Pathogen causing dieback of twigs and branches of Salix alba L. Conidiomata
500–300 µm diam. (x̅ = 400 × 350 µm, n = 10), pycnidial, solitary, immersed in host
tissue, unilocular, dark brown, ostiolate. Ostiole 150–40 µm diam. (x̅ = 145 × 40 µm,
n = 10), at the same level as the disc surface. Peridium comprising a few to several
layers of cell of textura angularis, with inner most layer thin, hyaline, outer layer
brown to dark brown. Conidiophores reduced to conidiogenous cells. Conidiogenous
cells blastic, enteroblastic phialidic, formed from the inner most layer of pycnidial
wall, hyaline, smooth. Conidia (3.4–) 4.3–5.3 × 0.7–0.8 (–1) µm (x̅ = 4.3 × 0.8 µm, n
= 30), unicellular, allantoid to subcylindrical, hyaline, smooth-walled. Sexual morph
Undetermined.
Culture characteristics: Colonies on PDA, reaching 3.5 cm diam. after 10 days at
25 °C, producing dense mycelium, circular, rough margin white, after 5 days, flat or
effuse on the surface, without aerial mycelium.
Material examined: RUSSIA, Rostov Region, Krasnosulinsky District,
Donskoye forestry, riparian forest, on dead twigs and branches of Salix alba L.
(Salicaceae), 21 May 2014, T.S. Bulgakov (MFLU 14–0785, holotype; PDD,
isotype); ex-type-living cultures, MFLUCC 14–1052, ICMP.
Notes: Cytospora salicicola belongs in Valsaceae based on morphology and
phylogeny. The new species has immersed, uniloculate conidiomata, with a single
ostiole and shares common walls with the host tissue. Cytospora salicicola is most
similar to C. schulzeri Sacc. & P. Syd. in conidia size [4.5–8(6.3) × 0.9–1.3(1.1) µm].
It however, differs in having a single locule, while C. schulzeri has multiple locules
with 2-11 ostioles per disc (Mehrabi et al. 2011). Phylogenetic analyses, using ITS
sequence data (Fig. 53) indicate that C. salicicola can be distinguished from other
species within the genus Cytospora. The tree using ACT, ITS and LSU sequence data
(Fig. 52) demonstrate that C. salicicola separates from other sequenced species in
Cytospora, and should be introduced as a new species.
3051
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3053
3054
3055
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3057
3058
3059
3060
3061
3062
3063
3064
Fig. 54 Cytospora salicicola (holotype) a Appearance of fruiting bodies in wood b Fruiting
bodies on substrate c Surface of fruiting bodies d Cross section of the conidioma e Peridium f
Ostiole of conidioma g Conidia h–g Colonies on PDA (P from below). Scale bars: a = 2 mm,
b–c = 1 mm, d = 100 µm, e = 10 µm, f = 50 µm, and g = 20 µm.
Glomerellales
Chadefaud (1960) proposed the order “Glomerellales” to accommodate a group
of endophytic and pathogenic fungi with ascomata varying from endostromatal to
apostromatal and ascospores that are often unicellular and hyaline. Réblová et al.
(2011) validated this order and accepted three families namely Australiascaceae,
Glomerellaceae and Reticulasceae in the class Sodariomycetes. This introduction was
based on analysis of ITS, LSU, and SSU datasets, and a combined data set of LSU
SSU and RPB2. Maharachchikumbura et al. (2015) included Plectosphaerellaceae in
3065
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3085
to this order based on a combined data set of LSU SSU TEF and RPB2. The
phylogenetic tree for Colletotrichum is presented in Fig. 55.
Glomerellaceae
The family Glomerellaceae was invalidly published by Locquin (1984),
validated in Zhang et al. (2006), and it was accepted as one of the three families of
Glomerellales in Réblová et al. (2011). Glomerellaceae is a monotypic family
characterized by the Glomerella sexual morph and the Colletotrichum asexual morph
(Maharachchikumbura et al. 2015).
Fig. 55 Phylogram generated from parsimony analysis based on combined ITS, GADPH,
CHS, ACT and β-tubulin sequence data of Colletotrichum. Parsimony bootstrap support
values greater than 50 % are indicated above or below the nodes, and branches with Bayesian
posterior probabilities greater than 0.95 are given in bold. The ex-type strains are in bold; the
new isolates are in blue. The tree is rooted with Monilochaetes infuscans CBS 869.96.
Colletotrichum Corda
Réblová et al. (2011) placed Colletotrichum in Glomerellaceae, and its
placement has been further confirmed by the study of Maharachchikumbura et al.
(2015). In the latter study the use of the name Colletotrichum over its sexual name
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3116
Glomerella was suggested. Hyde et al. (2009), Cai et al. (2009) and Cannon et al.
(2012) have treated this genus subsequently, and the most recent treatment is of Hyde
et al. (2014). This genus comprises plant pathogens, endophytes and saprobes
(Cannon et al. 2012).
295. Colletotrichum menispermi Chethana, Jayawardena, Bulgakov & K.D. Hyde, sp.
nov.
Index Fungorum number: IF 551744, Facesoffungi number: FoF 01648, Fig. 56
Etymology: The specific epithet menispermi is named after the host genus
Menispermum from which the taxon was collected.
Holotype: MFLU 14–0625
Saprobic on dead twigs of Menispermum dauricum DC. Sexual morph
Undetermined. Asexual morph Conidiomata 180–265 µm (x̅ = 229 µm, n = 10) diam.,
solitary, acervulus, black, oval. Setae 59–109 µm long, pale to dark brown,
smooth-walled, straight, 2–3-septate, base cylindrical, 4–9 µm diam. and rounded
apex. Conidiophores simple, to 33 µm long, hyaline to pale brown, smooth-walled.
Conidiogenous cells reduced. Conidia 6–18 × 2–5 µm (x̅ = 12 × 4 µm, n = 20), L/W
ratio 3.0, hyaline, aseptate, smooth-walled, both sides gradually tapering towards the
round to slightly acute apex, truncate base and guttulate. Appresoria not observed.
Material examined: RUSSIA, Rostov region, Rostov-on-Don city, Botanical
Garden of Southern Federal University, introductional nursery, on dead twigs of
Menispermum dauricum (Menispermaceae), 5 March 2014, T.S. Bulgakov, (MFLU
14–0625, holotype), (isotype in GZAAS, under the code of GZAAS 15–0102).
Note: Based on phylogenetic analyses and morphological comparison
Colletotrichum menispermi clusters in the Colletotrichum dematium species complex,
forming a separate branch with 100 % bootstrap support and 1.00 Bayesian posterior
probabilities. Colletotrichum menispermi separates from C. quinquefoliae with 100 %
bootstrap support and 1.00 Bayesian posterior probabilities. Morphologically it differs
from C. quinquefoliae in having larger conidiomata with minute 2–3-septate setae
which cannot be observed by unaided eye.
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3119
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3123
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3125
3126
Fig. 56 Colletotrichum menispermi (holotype) a Appearance of the conidiomata on the host
substrate b Close up of black conidioma c Brown 4-septate setae d Hyaline conidiogenous
cells e Hyaline conidia. Scale bars: a, b = 100 µm, c–e = 10 µm.
296. Colletotrichum quinquefoliae Jayawardena, Bulgakov & K.D. Hyde, sp. nov.
Index Fungorum number: IF 551745, Facesoffungi number: FoF 01649, Fig. 57
Etymology: The specific epithet quinquefoliae is named after the host
Parthenocissus quinquefolia (L.) Planch. from which the taxon was collected.
Holotype: MFLU 14–0626
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3149
3150
3151
Saprobic and weak pathogen on dying and dead leafstalks, twigs and tendrils of
Parthenocissus quinquefolia. Sexual morph Undetermined. Asexual morph
Conidiomata 267–517 µm ( x = 410 µm, n = 10) diam., black, acervulus, oval,
solitory, gregarious. Setae straight or ± bent, abundant, dark brown, becoming paler
towards the apex, opaque, smooth-walled, septa difficult to distinguish, 1–5-septate,
58–258 µm long, base cylindrical, 6.8–10.5 µm diam., tip somewhat acute.
Conidiophores medium brown, smooth–walled, simple, to 35 µm long.
Conidiogenous cells 7.3–12.8 × 1.4–3.3 µm ( x = 8.5 × 2.5 µm, n = 20), hyaline to
pale brown, smooth–walled, cyllindrical to slighty inflated, opening 0.5–1 µm diam.,
collarette or periclinal thickening not observed. Conidia 5.9–15.8 × 2.2–5.2 µm ( x =
9.9 × 3.3 µm, n = 40), L/W ratio 3.0, hyaline, smooth or verruculose, aseptate, curved,
both sides gradually tapering towards the round to slightly acute apex and base,
guttulate.
Material examined: RUSSIA, Rostov region, Rostov-on-Don city, Botanical
Garden of Southern Federal University, Higher Park, underwood, on Parthenocissus
quinquefolia (Vitaceae), 5 March 2014, T.S. Bulgakov (MFLU 14–0626, holotype),
(isotype in GZAAS, under the code of GZAAS 15–0101).
Notes: Colletotrichum dematium species complex is mainly characterized by
having curved conidia (Damm et al. 2009). Colletotrichum quinquefoliae falls within
the Colletotrichum dematium species complex and forms a separate clade which is
supported by 100% bootstrap value and 1.00 Bayesian posterior probability (Fig. 55).
This species differ from C. menispermi in having larger conidiomata, 1–5-septate,
long setae, with a larger base and conidia with an acute base. This species differs from
C. circinans and C. spinaceae in having longer setae with 1–5 septa and simple
conidiophores.
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3154
3155
3156
Fig. 57 Colletotrichum quinquefoliae (holotype) a Conidiomata on host b Black acervuli
with setae c Acute tip of the setae d Base of the setae e Seta f Conidiophores g Conidiogenous
cell h Conidium Scale bars: b = 200 µm, c = 50 µm, d = 5 µm, e = 150 µm, f = 20 µm, g–i = 5
µm.
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3168
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Hypocreales
Bionectriaceae
Fig. 58 Phylogram generated from maximum likelihood analysis based on LSU sequence data
of the family Bionectriaceae. New taxa are in blue ex-type strains are in bold. The tree is
rooted with Trichoderma viride.
Ochronectria
Ochronectria was established by Rossman and Samuels (1999) and is typified by
Ochronectria calami (Henn. & E. Nyman) Rossman & Samuels. The genus has
subglobose to globose ascomata, that are cupulate when dry, a three layered peridium,
clavate asci and fusiform ascospores with guttules (Rossman et al. 1999; Lechat 2010).
Ochronectria includes two species epithets (Index Fungorum 2016).
297. Ochronectria thailandica Q.J. Shang, D.Q. Dai & K.D. Hyde, sp. nov.
Index Fungorum number: IF 551918, Facesoffungi number: FoF 01815, Fig. 59
Etymology: The specific epithet “thailandica” refers to the country where the
fungus was first collected.
Holotype: MFLU 16–0030
Saprobic on bark. Sexual morph Ascomata 71–189 µm high, 78–223 diam.,
solitary to gregarious, superficial, black, globose, cup-like, or collapsing laterally
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3186
3187
3188
3189
3190
3191
3192
3193
3194
3195
3196
3197
3198
3199
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3201
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3209
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3214
3215
when dry. Ostioles brown to dark brown, 28–32 µm diam., with paraphyses. Peridium
31–52 µm wide, composed of three layers, inner 1–3 layers, comprising of hyaline,
thin-walled, elongated cells, central 3–4 layers of yellow to brown cells arranged in a
textura angularis, outer 5–6 layers, comprising dark brown to black, thick-walled
cells of textura angularis to globosa, having yellow oily droplets between the cells.
Hamathecium comprising 1.2–3 µm wide, hyaline, aseptate paraphyses. Asci 34–56 ×
6–9 µm ( x = 45 × 7 µm, n = 30), 8-spored, unitunicate, clavate, with short pedicel,
slightly rounded to truncate at the apex. Ascospores 12–17 × 3–4 µm ( x = 14 × 3 µm,
n = 50), overlapping 2-seriate, fusiform, hyaline, 2-celled, straight to curved,
smooth-walled, with small guttules. Asexual morph Undetermined.
Culture characteristics: Ascospores germinating on MEA within 24 h. Germ
tubes produced from any cell. Colonies on MEA reaching 1.5–2 mm diam. after 7 d in
the dark at 25 °C, edge entire, flat or effuse or umbonate, sparse, forming ascomata on
MEA in the centre. After 7 d colonies white (n) above, from below reddish yellow (o).
Material examined: THAILAND, Chiang Rai, Mae Sai, Pong Ngam Village,
Tham Pla Cave, on unidentificated wood in the water, 25 November 2014, Qiu Ju
Shang, SHTM02–4 (MFLU 16–0030, holotype), ex-type living culture, MFLUCC
15–0140, (isotype in KUN-HKAS, under the code of KUN-HKAS 93730), ex-living
culture KUMCC 16-0001).
Notes: Based on phylogenetic analyses and morphological comparison, our
isolate belongs to the genus Ochronectria in the family Bionectriaceae. The
morphology of Ochronectria thailandica fits well with the description provided by
Rossman and Samuels (1999). It differs from the type, O. calami (Henn. & E. Nyman)
Rossman & Samuels and O. courtecuissei Lechat based on the size and colour of
ascomata, peridium colour and number of septa and dimensions of the ascospores.
Ochronectria thailandica has black ascomata, a peridium composed of black brown
outer layers and yellow middle layers, while, O. calami has white or yellow to orange
ascomata, a peridium composed of hyaline middle and outer layers, and O.
courtecuissei has yellow to brown ascomata and a peridium composed of yellow to
orange outer layers and hyaline middle layers (Rossman et al. 1999; Rossman et al.
2001; Lechat 2010). Furthermore, O. thailandica, which has 1-septate ascospores can
be distinguished from O. calami which forms multi-septate ascospores. Phylogenetic
analysis based on LSU sequence data of the family Bionectriaceae showed that O.
thailandica is closely related to Ochronectria calami, forming a distinct lineage
within the sclade (Fig. 58).
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3221
3222
Fig. 59 Ochronectria thailandica (holotype) a Host b, c Ascomata on host d Vertical section
of ascoma e Periphysate ostiole f Section of peridium g Paraphyses and asci h–j Asci; note i
stained in Melzer’s reagent k, l Ascospores m Germinating ascospore n, o Culture on MEA.
Scale bars: b, c = 100 µm, d = 25 µm, e, g–l = 10 µm, f, m= 20 µm.
Clavicipitaceae
3223
3224
3225
3226
3227
3228
3229
3230
3231
3232
3233
3234
3235
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3237
3238
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3243
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3245
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3247
3248
3249
3250
The family Clavicipitaceae (Hypocreales) is a very heterogeneous group of fungi
that are associated with a broad range of invertebrate animals, plants and occasionally
with other fungi (Sung et al. 2007, Schardl et al. 2014, Kepler et al. 2012). The
plant-associated Clavicipitaceae includes mutualistic symbionts, such as the grass
endophytes Epichloë and Balansia, as well as plant pathogens, many of which
produce alkaloids (e.g. Claviceps purpurea) with diverse neurotropic effects on
vertebrate and invertebrate animals, with important implications for human health,
agriculture and food security (Spatafora et al. 2007). The invertebrate-associated
Clavicipitaceae comprises many pathogens of scale insects and whiteflies, such as
Conoideocrella, Hypocrella, Moelleriella, Orbiocrella, Regiocrella, and Samuelsia
(Chaverri et al. 2008). Paecilomyces, Pochonia and Metarhizium are also other
invertebrate-pathogens that infect a wide range of insect hosts (Kepler et al. 2014).
The sexual morphs in this family produce various types of stromata and colors, but all
produce filiform asci with ascospores that may or may not disarticulate into
part-spores.
Moelleriella infects scale insects and white flies and was recently separated from
the genus Hypocrella together with Samuelsia (Chaverri et al. 2008). The delimitation
and separation of Moelleriella was based on molecular data and morphology: its
ascospores disarticulate inside the ascus. The asexual morph of Moelleriella is
aschersonia-like, i.e., it is similar to Aschersonia sensu stricto (sexual morph
Hypocrella sensu lato; Chaverri et al. 2008). Species in Aschersonia sensu lato are
characterized mostly by the shape and colour of the stromata that cover the hosts,
pycnidium-like conidiomata, phialides, and presence or absence of paraphyses. These
characters have been useful in distinguishing between subgenera of Aschersonia
(Petch 1921; Mains 1959a, b; Chaverri et al. 2008). The combined analysis of LSU
and RPB1 in comparison with related species, support M. phukhiaoensis and M.
pongdueatensis as new species from Thailand. The phylogenetic tree is presented in
Fig. 60.
3251
3252
3253
3254
3255
3256
3257
3258
3259
3260
3261
3262
3263
3264
3265
Fig. 60 Phylogenetic relationships between Moelleriella phukhiaoensis, M. pongdueatensis
and related species generated from a combined LSU and RPB1 gene dataset using maximum
parsimony and Bayesian analysis. The numbers on each branch represent the bootstrap
values/Bayesian PP. New taxa are in blue and species for ex-type strains in bold.
298. Moelleriella phukhiaoensis Mongkol., Thanakitp. & Luangsa-ard, sp. nov.
Index Fungorum number: IF 551609, Facesoffungi number: FoF 02030, Fig. 61
Etymology: The specific epithet refers to Phu Khiao Wildlife Sanctuary, the
collection location
Holotype: BBH 17305
Specimens were found on the underside of dicotyledonous leaves. Hosts are
scale insect nymphs (Hemiptera). Stromata flattened pulvinate, sometimes surrounded
by a membranous hypothallus; up to 5 mm diam. and 2 mm high, dark orange to
golden yellow. Sexual morph Perithecia 400–520 × 150–200 µm, crowded,
3266
3267
3268
3269
3270
3271
3272
3273
3274
3275
3276
3277
3278
3279
3280
3281
3282
3283
3284
3285
3286
3287
3288
3289
3290
3291
3292
immersed, elongate flask-shaped, ostioles slightly projecting, translucent. Asci
195–220 × 8–12 µm, cylindrical, with cap approx. 4–6 thick. Ascospores
disarticulating into 12.5–17.5 × 2–3 µm part-spores inside the ascus, cylindrical with
somewhat rounded ends. Asexual morph Conidiomata orifice scattered or circularly
arranged, ultimately hidden by the orange-yellow mass or extruded conidia, oval or
elongate flask shaped, up to 430 µm deep, up to 100 diam. Conidiogenous cells
cylindrical, up to 25 µm long, 1–2 µm wide. Conidia 16–17 µm × 2.5–3.5 µm,
cylindrical narrow, tapering slightly towards the ends. Paraphyses present, linear,
filiform, up to 90 µm long; 1–2 µm wide.
Culture characteristics: Cultures were obtained from germinating ascospores and
conidia. The ascospores and conidia germinated within 48 h on PDA. The colonies on
PDA grew slowly, to approx. 5 mm diam. after 4 wk at 20°C. The stromatic colonies
derived from germinating ascospores or conidia formed a compact mycelium. The
conidial mass yellow to orange yellow appearing as abundant slimy masses scattered
over the surface of stromatic colonies.
Material examined: THAILAND, Chaiyaphum Province, Bueng Pan Protect
Forest Unit, Phu Khiao Wildlife Sanctuary, 15 October 2005, S. Mongkolsamrit, R.
Ridkaew, B. Thongnuch, K. Tasanathai (BBH 17305, holotype); ex-type living
culture, BCC19769.
Notes: The sexual morph of M. phukhiaoensis is rarely found when compared
with the asexual morph. The asexual morph of M. phukhiaoensis was compared with
the Thai material of Aschersonia placenta (sexual morph M. raciborskii) based on the
pale yellow to light orange stromata. Although the asexual morph of M.
phukhiaoensis morphologically resembles A. placenta, it differs significantly from the
latter in having longer conidia (12–14 × 2–2.5 µm) as reported for A. placenta by
Luangsa-ard et al. (2007). Moelleriella phukhiaoensis has only been collected in the
Phukhiao Phu Khiao Wildlife Sanctuary.
3293
3294
3295
3296
3297
3298
3299
3300
3301
3302
3303
3304
3305
3306
3307
3308
3309
3310
3311
3312
Fig. 61 Moelleriella phukhiaoensis (holotype) a, b Fungi on hosts c Culture derived from
ascospores on PDA (sporulation present) d Side view of ascostroma showing flask-shaped
perithecia (arrows) e Ascus showing a thickened cap f Part of ascus showing ascospores g
Ascospores h Longitudinal section through the stroma showing conidiomata with conidia
(arrows) i Conidiogenous cells and paraphysis j Conidia. Scale bars: b, c = 1 mm; d = 500 µm;
e, f = 10 µm, g, j =20 µm, h, i = 100 µm.
299. Moelleriella pongdueatensis Mongkol., Thanakitp. & Luangsa-ard, sp. nov.
Index Fungorum number: IF 551610; Facesoffungi number: FoF 02031, Fig. 62
Etymology: The specific epithet refers to Pong Dueat Pa Pae Geyser, the
collection location
Holotype: BBH 24730
Specimens were found on the underside of bamboo leaves. Hosts are scale insect
nymphs (Hemiptera). Stromata usually discoid, distinctly stud-shaped, up to 4 mm
diam. and 2 mm high, pale yellow, base surrounded by a membranous hypothallus.
Sexual morph No stromata observed. Asexual morph Conidiomata scattered around
a narrow neck, extruding an orange yellow mass of conidia. Conidiogenous cells
cylindrical, up to 23 µm long, 1–2 µm wide. Conidia fusoid, 9–12.5 µm × 1.5–2.5 µm.
Paraphyses present, linear, filiform, up to 110 µm long; 1–2 µm wide. The
3313
3314
3315
3316
3317
3318
3319
3320
3321
3322
3323
3324
3325
3326
3327
3328
3329
3330
3331
3332
3333
3334
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3336
3337
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3339
3340
3341
3342
3343
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3346
3347
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3349
hirsutella-like synanamorph is scattered on the upper surface of the stroma, phialides
with a long thin neck, up to 20 µm, 1–2 µm wide, conidia citriform, 2–3 × 1–2.5 µm.
Culture characteristics: Cultures were obtained from germinating conidia. The
conidia germinated within 24 h on PDA. The colonies on PDA grew slowly, to approx.
5 mm diam. after 2 wk at 20°C. The stromatic colonies formed a compact mycelium.
The cream to pale yellow conidial mass covers the stromatic colonies.
Material examined: THAILAND, Chiang Mai Province, Pong Dueat Pa Pae
Geyser, 5 July 2008, S. Mongkolsamrit, B. Thongnuch, K. Tasanathai, P.
Srikitikulchai, A. Khonsanit (BBH 24730, holotype); ex-type living culture,
BCC31787
Notes: The sexual morph of this species was not found in the field although
several attempts to find it were made throughout the year. The asexual state of
Moelleriella pongdueatensis is similar to Aschersonia basicystis Berk. & M.A. Curtis
(sexual morph Moelleriella basicystis P. Chaverri & K.T. Hodge) reported from Costa
Rica, Cuba and Panama by Chaverri et al. (2008) based on stud-shaped and pale
yellow stroma, and yellow mass of extruded conidia, around a narrow neck. The
conidia of M. pongdueatensis, however, are somewhat smaller; the conidia are fusoid,
9–12.5 × 1.5–2.5 µm, with paraphyses up to 110 µm long. In contrast, Aschersonia
basicystis conidia are ventricose, (11–)13–13.5 (–15.5) × (3–)4–4.2(–5) µm, with
acute ends, paraphyses are absent. Based on our study, Moelleriella pongdueatensis is
the second species that show the presence of hirsutella-like synanamorphs
simultaneously occurring on stromata in nature. Tadych et al. (2009) first reported M.
zhongdongii having both Aschersonia and hirsutella-like synanamorphs on stromata
in nature along with the Moelleriella sexual morph.
Phylogenetic analysis
Independent maximum parsimony analyses were done for each gene.
Comparisons of the bootstrap supports for the nuclear large subunit rRNA gene (LSU)
and RNA polymerase II subunit one (RPB1) gene datasets showed no significant
contradictory nodes, and where the bootstrap supports were ≥70% the strains were
prepared to make a combined data set for both LSU and the RPB1 for analysis. The
combined dataset for the LSU and RPB1 sequence data consisted of 1447 characters,
986 of which are constant, 50 are variable and parsimony-uninformative, while 411
are parsimony-informative. Maximum parsimony analysis of the combined dataset of
LSU and RPB1 resulted in 12 most parsimonious trees. Maximum parsimony
analyses of this data set yielded one parsimonious tree (tree length 1540; CI = 0.455,
RI = 0.802, RC = 0.365, HI = 0.545) as shown in Fig. 60.
3350
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3352
3353
3354
3355
3356
3357
3358
3359
3360
3361
3362
3363
3364
3365
3366
3367
3368
3369
3370
Fig. 62 Moelleriella pongdueatensis (holotype) a, b Fungi on hosts c Culture derived from
conidia on PDA (sporulation present) d Side view of stroma showing stud-shaped e
Paraphyses f Conidiogenous cells and paraphysis g Conidia h SEM derived from stroma i
SEM of hirsutella-like on stroma. Scale bars: a–d, h = 1 mm, e = 50 µm, f, g, i = 20 µm.
Ophiocordycipitaceae
The family Ophiocordycipitaceae (order) was introduced by Sung et al. (2007)
based on phylogenetic analyses and later emended by Kirk et al. (2013) and Quandt et
al. (2014). Kirk et al. (2013) listed eleven genera under this family, while Quandt et al.
(2014) refined it and proposed six genera, including Drechmeria, Harposporium,
Ophiocordyceps, Polycephalomyces, Purpureocillium and Tolypocladium.
Maharachchikumbura et al. (2015) confirmed this system and Spatafora et al. (2015)
introduced some necessary species combinations based on this classification. Most
species of Ophiocordycipitaceae are known to produce dark pigmented, tough to
pliant stromata, that often possess aperithecial apices (Sung et al. 2007). The main
distinguishing characters of this genus are that the ascospores do usually not break
into part-spores at maturity and asci have thin apical caps (Petch 1931, 1932). The
phylogenetic tree is presented in Fig. 63.
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3373
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3375
3376
3377
3378
3379
3380
3381
3382
3383
3384
3385
3386
3387
3388
3389
3390
3391
3392
3393
Fig. 63 Phylogram of Ophiocordyceps generated from Maximum likelihood analysis
of SSU, rpb1 and tef1-α sequence data. Simplicillium lanosoniveum (J.F.H. Beyma)
Zare & W. Gams is used as outgroup taxon. Maximum likelihood bootstrap values
greater than 50 % and Bayesian posterior probabilities over 0.90 are indicated above
or below the nodes. The new species are indicated in blue.
300. Ophiocordyceps formosana Y.W. Wang et al. in Wang et al., Evidence-Based
Complementary and Alternative Medicine (no. 189891): 4 (2015)
Facesoffungi number: FoF 01796, Fig. 64
Parasitic in larva of Coleoptera (Superfamily Tenebrionoidea), forming yellow
to orange ascostromata. Sexual morph Ascomycetous. Stromata 14 mm long, 2-5
mm wide, growing from the head and the tail of Coleoptera larva, simple or branched,
yellow to orange, stipitate. Stipe 1.9–3.7 cm long, 2–4 cm wide, yellow, cylindrical,
surface rough. Fertile head 30 mm long, 2–2.5 mm wide, orange, mostly elliptic
barely branched, head-like, with orange, pseudoparenchymatous, epidermal tissues,
surface mastoid, differentiated from stipe. Ascomata 453–546 × 265–298 µm ( x =
479 × 270, n = 30), completely immersed, orange, flask-shaped to oval, with the
ostioles opening on the surface of the head. Peridium 26–38 µm wide ( x = 30, n =
60), comprising three layers. Asci 366–498× 8–11 µm ( x = 437 × 10, n = 60),
8-spored, hyaline, cylindrical, with apical cap, breaking into secondly ascospores.
Secondary ascospores 2–6 × 1–3 µm ( x = 4 × 2, n = 60), hyaline, cylindrical.
Asexual morph Undetermined.
3394
3395
3396
3397
3398
3399
3400
3401
3402
3403
3404
3405
Material examined: CHINA. Province of Hunan, on dead larva of
Tenebrionoidea, 23 October 2014, Ping Zhang, ZP8282 (MFLU 15–3888); ZP828i
(MFLU 15–3889, MFLU 15–3890, MFLU 15–3891).
Notes: Ophiocordyceps formosana was introduced by Kobayashi (1979) as
Cordyceps formosana Kobayasi & Shimizu. Wang et al. (2015a) revised it as
Ophiocordyceps formosana. This species is frequently used in Traditional Chinese
Medicine and has a long history of use as tonics and folk medicines that can be used
as anticancer and diabetes treatments and contains antioxidants (Wang et al. 2015a).
This species was previously known from Fujian and Taiwan (Wang et al. 2015a). We
collected this species in Hunan Province, China, which is a new record for the
province. We also provide a colour figure for this species which includes asci and cap
and entire ascospores, which are illustrated for the first time.
3406
3407
3408
3409
3410
Fig. 64 Ophiocordyceps formosana (MFLU 15–3888) a Stromata appearing from the tree b
Yellow, superficial stroma appearing from host head c Overview of the stromata and the host
d, f Apical part of the stroma e Vertical section of stroma g Cross section showing the
complete perithecia h–j Asci at immature and mature stages k Entire ascospore l, m Asci with
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3445
apical cap n, o Secondly ascospores. Scale bars: d = 1000 µm, e, f = 200 µm, g = 100 µm, h, k
= 20 µm, i, j = 50 µm, l–o =5 µm.
301. Ophiocordyceps karstii T.C. Wen, Y.P. Xiao & K.D. Hyde, sp. nov.
Index Fungorum number: IF 551814, Facesoffungi number: FoF 01795, Fig. 65
Etymology: Name referring to the location which the specimen was collected.
Holotype: MFLU 15–3884
Parasitic in larva of Hepialus jianchuanensis, brown to dark brown, forming
yellow to brownish stromata. Sexual morph Thallus within host white, composed of
intercalary hyphal bodies. Stromata mostly single, 140–145 × 2–4 mm, stipitate
arising from head of the host. Stipe 12 cm long, 2 mm wide, clavate, with a fertile
apex, becoming golden yellow to brownish yellow when mature. Fertile head 20–25
mm long, 2–4 mm diam., clavate, light yellow to yellow-brown, upper surface
roughened, covered with white non compact mycelium. Ascomata 600–765 ×
247–323 µm ( x = 683 × 285 µm, n = 30), superficial, yellow to brown, flask-shaped,
thick-walled, ostiole on the top. Peridium 63–42 mm ( x = 52 µm, n = 60) wide, three
layers. Asci 186–228 × 8–12 µm ( x = 207 × 10 µm, n = 60), 8-spored, hyaline,
narrow cylindrical, with a thickened apex. Apical cap 5–7 µm ( x = 6 µm, n = 60)
diam. Ascospores 173–202 × 3–5 µm ( x = 188 × 4 µm, n = 60) fasciculate, fusiform,
smooth, as long as asci, hyaline, 10–18 septate, not breaking into secondly spores.
Asexual morph Undetermined.
Material examined: CHINA. Guizhou Province, Chishui, on dead larva of
Hepialus jianchuanensis, 23 October 2014, TingChi Wen CS2014102301 (MFLU
15–3884, holotype); CS2014102304 (MFLU 15–3885, MFLU 15–3886, MFLU
15–3887, paratype).
Notes: Ophiocordyceps was introduced by Petch (1931) with Ophiocordyceps
blattae (Petch) Petch as the type species and used by Sung et al. (2007) as the type
genus of Ophiocordycipitaceae. According to morphological and phylogenetic
analysis, Ophiocordyceps karstii closedly matches O. lanpingensis Hong Yu bis &
Z.H. Chen bis and O. robertsii (Hook.) G.H. Sung et al. This species is characterized
by fusiform ascospores (173–202 × 3–5 µm, x = 188 × 4 µm, n = 60), which do not
break into secondly ascospores and superficial ascomata. Phylogenetic analysis of th
of combined TEF1, RPB1 and SSU sequence data (Fig. 63) confirms that
Ophiocordyceps karstii clusters with O. robertsii in Ophiocordyceps with high
bootstrap support. Therefore, we proposed O. karstii as a new species.
3446
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3449
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3452
Fig. 65 Ophiocordyceps karstii (holotype) a Overview of the host and stromata b Host:
Hepialus jianchuanensis c Stroma d Vertical section of stroma e Vertical section showing the
superficial perithecia f, g Perithecia h–k Asci at immature to mature stages l–n Ascospores.
Scale bars: c = 2 mm, d, f = 500 µm, e–g = 200 µm, h–n = 50 µm.
Table 3 Synopsis of Ophiocordyceps species discussed in the paper
Species
Stromata Ascomata Asci
Ascospor Secondl Referenc
(mm)
(µm)
(µm)
es (µm)
y spores e
(µm)
O. karstii
140–150
× 2–4
600–765 × 186–228
247–323
× 8–12
O.
lanpingensis
50–160
×
0.2–1.3
100–380
× 3–4
310–370 × 240–300 240–300
200–240
× 5.1–6.5 × 1.4
O. robertsii
O. sinensis
O.
xuefengensis
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3466
40–110
140–460
× 2–7
173–202
× 3–5
600–880 × 280–400
300–400
× 9–10
280 × 3
380–550 × 240–485
140–240
× 12–16
160–470
× 5–6
Not
breakin
g
Not
breakin
g
5–6 × 3
Not
breakin
g
416–625 × 191–392 130–380 Not
161–318
× 4.5–8.9 × 1.4–5.2 breakin
g
This
study
Chen et
al. 2013
Cunning
ham
1921
Liang et
al. 2007
Wen et
al. 2013
Microascales
Halosphaeriaceae
The family Halosphaeriaceae was introduced by Müller and von Arx (1962)
with Halosphaeria as the type genus (Barghoorn and Linder 1944). Morphological
characters include the perithecioid ascomata, presence of catenophyses that generally
deliquesce, clavate to fusiform, unitunicate thin-walled asci; hyaline, septate
ascospores sometimes with polar appendages (Jones 1995, Sakayaroj et al. 2011,
Jones et al. 2015). Members of Halosphaeriaceae constitute the largest group of
marine Ascomycota mainly found in marine habitats, with few transitional species
found in freshwater and brackish water (Jones 1995; Pang et al. 2003; Jones et al.
2009, Sakayaroj et al. 2011). The phylogenetic tree is presented in Fig. 66.
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3477
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3479
3480
3481
3482
Fig. 66 Maximum likelihood (ML) majority rule consensus tree for the analyzed
Halosphaeriaceae isolates based on a dataset of combined LSU and SSU sequence data.
RAxML bootstrap support values (ML) are given at the nodes (ML). The scale bar represents
the expected number of changes per site. The tree is rooted with Microascus trigonosporus
and Petriella setifera. The original isolate numbers are noted after the species names. The
new strain is in blue bold and other strains in Aniptodera are in black bold.
302. Aniptodera aquibella J. Yang & K.D. Hyde, sp. nov.
Index Fungorum number: IF 551897, Facesoffungi number: FoF 01818, Fig. 67
Etymology: from the Latin aqua = water, bellus = lovely, referring to the
freshwater habitat.
Holotype: MFLU 15–1140
Saprobic on decaying, submerged twigs in freshwater habitats, shining on the
host surface. Sexual morph Ascomata 130–160 × 150–200 µm, superficial or
immersed, globose or subglobose, scattered, hyaline or greyish, membranous. Neck
3483
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3485
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3487
3488
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3497
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3500
3501
3502
3503
3504
3505
3506
3507
3508
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3517
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3524
80–110 × 40–60 µm, cylindrical to conical, hyaline, with periphyses. Peridium 7–15
µm thick, composing several layers of hyaline-walled cells of textura globosa.
Catenophyses sparse, hyaline, septate, consisting of elongated cells, slightly
constricted at the septa. Asci 60–110 × 25–45 µm (x = 90 × 30, n = 20), 8-spored,
thin-walled, clavate, becoming balloon-shaped or swollen, flattened at apex, tapering
to a pointed pedicel, unitunicate, wall thickened at the apex, subapical cytoplasm
retracted, mostly persistent, with a J-, apical thickening, which has an apical pore.
Ascospores 25–30 × 7–10 µm (x = 28 × 8, n = 50), 1-euseptate, slightly constricted at
the septa, thin-walled, hyaline, smooth-walled, ellipsoidal, 2–3-seriate, guttulate,
sometimes with indistinct appendages at both ends. Asexual morph Undetermined.
Culture characteristics: Ascospores germinating on PDA within 24 h and germ
tubes produced from the poles of both cells. Colony on MEA slow-growing, reaching
5–10 mm diam. at 14 days, dark brown in the middle, conspicuous paler and sparser
at edge, with dense white mycelium on surface in the middle of colony; in reverse
with a dark brown middle and olive-green smooth margin. Mycelium immersed and
superficial in the media, composed of branched, septate, smooth-walled, hyaline aerial
hyphae and dark brown hyphae near or within the media.
Habitat and distribution: On submerged wood in freshwater, Thailand.
Material examined: THAILAND, Prachuap Khiri Khan Province, Hua Hin,
Kaeng Krachan, near Pala-U Waterfall, stream outside national park, on submerged
wood, 25 December 2014, Jaap van strien (MFLU 15–1140, holotype), ex-type living
culture, MFLUCC 15–0605, GZCC 15–0055.
Notes: The genus Aniptodera was established by Shearer and Miller (1977) with
A. chesapeakensis Shearer & M.A. Mill. as the type species. The genus was described
as having hyaline or light colored ascomata, catenophyses, apically thickened
persistent asci with a distinct pore and subapical retraction of cytoplasm, and hyaline,
thick-walled, 1-septate ascospores with or without appendages (Shearer and Miller
1977; Raja and Shearer 2008). Aniptodera aquibella fits well within Aniptodera. It is
most similar to A. chesapeakensis, except that the ascospores are smaller and the
ascospore walls are thinner than those of A. chesapeakensis (Shearer and Miller 1977).
Aniptodera aquibella differs from other species in the genus by conspicuous
differences in the size and shape of asci and ascospores. Aniptodera intermedia K.D.
Hyde & Alias has the shortest asci (46–62 × 16–19 µm) and smallest ascospores
(10.5–13 × 7–8 µm), while A. longispora K.D. Hyde has the longest asci (145–201 ×
24–31 µm) and larger ascospores (39–51 × 9–13.5 µm) in the genus (Hyde 1990,
1999). Aniptodera megaloascocarpa Raja & Shearer differs distinctly from A.
aquibella because it has the largest ascomata (1060–1360 × 430–530 mm) of all the
species in Aniptodera (Raja and Shearer 2008). Aniptodera margarition Shearer and A.
mangrovei K.D. Hyde lack any apical thickening and the subapical retraction of
cytoplasm and the former also lacks a distinguishable apical pore characteristic of all
Aniptodera species (Shearer 1989). Aniptodera triseptata K.D. Hyde is the only
species with 3-septate ascospores in the genus (Hyde 2002).
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Fig. 67 Aniptodera aquibella (holotype) a, b Appaerance of ascomata on submerged wood c
Section of an ascoma d Section through peridium e Peridium in surface view f Surface of
periphysate neck g–k Asci l–q Ascospores r Catenophyses s Germinated spore t–u Culture
on MEA t from above. Scale bars: a = 100 µm, b–c = 50 µm, d–e, h = 20 µm, f–g, i–k, s = 30
µm, l–q = 15 µm, r = 10 µm.
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Sordariales
The order Sordariales was detailed by Maharachchikumbura et al. (2015) and this is
followed here.
Chaetomiaceae
Fig. 68 Phylogenetic tree for Humicola koreana EML-UD33-1 and EML-UD33-2 and related
species based on Maximum likelihood analysis of a ITS, b LSU sequence data. Sequences of
Penicillium griseofulvum, Mucor indicus and Rhizomucor pusillus were used as outgroups.
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.
Humicola Traaen
The genus Humicola was established by Traaen (1914) for two species, H.
fuscoatra Traaen and H. grisea Traaen which were isolated from Norwegian soil.
Species belonging to this genus are slow growing and form solitary, dark, globose to
elongate, single-celled conidia (Omvik 1955; De Bertoldi et al. 1972). However, until
now, the taxonomy of the genus have not yet been studied in detail. About six species
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including some varieties are recognized in this genus (Ko et al. 2011). The genus is
likely to be polyphyletic with some species being the asexual morphs of Chaetomium.
Several species of the genus, Humicola are rich in organic matter and are able to
produce strong cellulolytic enzymes and may have important economical application
(White and Downing 1953; Sharma et al. 2008; Du et al. 2013). Species may also
reduce disease caused by Aspergillus flavus, Phytophthora capsici and Alternaria
brassicicola (Wicklow et al. 1998; Ko et al. 2011). Thus, the purpose of this study
was to investigate the morphological charateristics of a Humicola species isolated
from soil and to conduct molecular phylogenetic analyses to establish their placement
in Ascomycota.
During a study on the Sordariales from a soil sample of Ulleung-do island which
is about 161 km far from the mainland of Korea, a Humicola species that differs
morphologically and phylogenetically from the other species of the genus was isolated
and is described as new to science.
303. Humicola koreana Hyang B. Lee & T.T.T. Nguyen, sp. nov.
MycoBank number: MB 814402, Facesoffungi number: FoF 02068, Fig. 69
Etymology: koreana. Referring to the country which from the species was first
isolated (Korea)
Holotype: EML-UD33-1
Colonies of strain grow slowly on PDA, initially nearly buff and then changing
to luteus, reaching 59–61 mm in diam. at 25oC after 7 days of incubation. The reverse
of colonies is yellow in the center with a lighter margin and irregular zonation.
Conidia are formed laterally, the shapes are commonly round, ovovoid to some
ellipsoid, and measure 8–10.7 µm in diam. The conidia have outwardly melanized
thick wall layers. At maturity, conidia are detached from the conidiophores having
scars.
Notes: Humicola koreana is morphologically similar to H. fuscoatra and H.
grisea, but differs from the related species in having smaller spores and producing
yellow pigment when cultivated on PDA.
Material examined: REPUBLIC OF KOREA, from a soil sample from
Ulleung-do island; EML-UD33-1 (EML-UD33-1, holotype a dried culture, stored at
Division of Food Technology, Biotechnology & Agrochemistry, College of
Agriculture & Life Sciences, Chonnam National University, Gwangju 61186, Korea)
ex-type living culture at the Culture Collection of National Institute of Biological
Resources (NIBR), Incheon, preserved as glycerol stock at -80oC in the CNUFC and
deposited at Jena Microbial Resource Collection (University of Jena and Leibniz
Institute for Natural Product Research and Infection Biology, Jena, Germany)
(JMRC:SF:012183).
The isolate was observed to grow over a wide range of temperatures with varying
growth rates on PDA, MEA (malt extract agar), and CDA (czapek dox agar). The
average growth rates of EML-UD33-1 on PDA, MEA, and CDA were 7 mm, 6 mm,
and 7.5 mm per 24 hours, respectively. Optimal growth was observed around
25–27°C, slow growth was observed at below 20°C, and no growth at 37°C.
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Humicola koreana appears to be phylogenetically related to H. fuscoatra, the type of
the genus Humicola (Fig. 68).
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Fig. 69 Humicola koreana (holotype) a, b Yellow colonies in potato dextrose agar (PDA) (a
from above, b from below) c–f, i–l Two different types of conidiophores (white arrows) and
aleuriconidia, vase-shaped conidiophore c Column-shaped conidiophore e Ground to obovoid
conidia with scar (purple arrow) and thick wall layer h Scar on the conidiophore after
detachment (yellow arrow). Scale bars: c–h = 20 µm, i, k = 10 µm, j, l = 15 µm.
Amphisphaeriales
Amphisphaeriaceae
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Fig. 70 Phylogram generated from maximum likelihood analysis (ML) based on combined
LSU and ITS sequence data of Seimatosporium. Maximum likelihood bootstrap support
values greater than 50% are near the nodes. New taxa are in blue and ex-type strains are in
bold. The tree was rooted to Pseudopestalotiopsis theae (MFLUCC 12–0055).
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Seimatosporium
The genus Seimatosporium was introduced by Corda (1833) with S. rosae as the
type species, and Shoemaker (1964), Shoemaker and Muller (1964), Sutton (1980)
and Nag Raj (1993) revisited the genus. Barber et al. (2011), Tanaka et al. (2011),
Norphanphoun et al. (2015) and Senanayake et al. (2015) re-visited the genus and
discussed the taxonomic placement based on sequence analyses.
Nag Raj (1993) and Okane et al. (1996) stated Discostroma was the sexual
morph of Seimatosporium. Recent publications also showed that both Seimatosporium
and Discostroma grouped in a monotypic clade (Barber et al. 2011; Tanaka et al. 2011;
Norphanphoun et al. 2015; Senanayake et al. 2015). Norphanphoun et al. (2015)
designated the epitype for Seimatosporium rosae, the type species of Seimatosporium.
304. Seimatosporium pseudocornii Wijayaw., Camporesi & K.D. Hyde, sp. nov.
Index Fungorum number: IF 551754, Facesoffungi number: FoF 01653, Fig. 71
Etymology: Named as its morphological similarity to Seimatosporium rosae
Holotype: MFLU 15–3558
Saprobic on dead branches and stems of Cornus sp. Sexual morph
Undetermined. Asexual morph Conidiomata 320–350 µm diam., 50–120 µm high,
acervular, unilocular, subglobose, superficial to subepidermal, solitary to gregarious,
dark brown to black, non papillate ostiole. Conidiomata wall multi-layered, outer wall
thick, composed of brown cells of textura angularis, inner wall thin, hyaline.
Conidiophores 5–30 × 2–4 µm, long, cylindrical, branched, hyaline, smooth-walled.
Conidiogenous cells holoblastic, annellidic, simple, integrated, determinate, hyaline.
Conidia 31–42 × 5–7 µm ( x = 38.1 × 6.1 µm, n = 20), obovoid to fusiform,
occasionally truncate base, obtuse apex, straight to slightly curved, 3-transverse
septate, brown to dark brown septa, constricted at the septa, often guttulate at
immaturity, medium brown, hyaline to sub-hyaline basal cell, smooth-walled,
appendage absent.
Culture characteristics: On PDA slow growing, attaining a diam. of 2 cm in 7
days at 18 ºC, white to pale brown from top, greyish white from below, with sparse
mycelium, flat, uneven margin.
Material examined: ITALY, Forlì-Cesena [FC] Province, near Monte Riccio Bagno di Romagna, on dead branch of Cornus sp. (Cornaceae), 5 January 2013, Erio
Camporesi, IT 1000 (MFLU 15–3558, holotype); (HKAS isotype), ex-type living
cultures MFLUCC 13–0529, GUCC IT 1000, KIB.
Notes: Farr and Rossman (2015) reported Seimatosporium lichenicola (Corda)
Shoemaker & E. Müll. (conidial dimensions 13–15 × 5.5–6.5 µm fide Sutton 1980)
and S. salicinum (Corda) Nag Raj (11–17 × 4–6 µm fide Nag Raj 1993) from Cornus
spp. Senanayake et al. (2015) reported Seimatosporium corni Wijayawardene et al.
(conidial dimensions 21–29 × 9–11 µm). In morphology our new collection is distinct
from these species, thus we introduce a new species based on morphology, host
association and phylogenetic analyses.
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Fig. 71 Seimatosporium pseudocornii (holotype) a Appearance of conidiomata on dead
branch of Cornus sp. b, c Cross sections of conidiomata d–h Different stages of
conidiogenesis i–o Conidia p Germinating conidium. Scale bars: b = 50 µm, c–o = 25 µm, p =
30 µm.
305. Seimatosporium pseudorosae Wijayaw., Camporesi & K.D. Hyde, sp. nov.
Index Fungorum number: IF 551753, Facesoffungi number: FoF 01652, Fig. 72
Etymology: Named as its morphological similarity to Seimatosporium rosae
Holotype: MFLU 15–3559
Saprobic or endophytic on living branches and stems of Rosa villosa (Rosaceae).
Sexual morph Undetermined. Asexual morph Conidiomata 175–250 µm diam.,
200–250 µm high, acervular, unilocular, subglobose, superficial to subepidermal,
solitary, dark brown to black, with apapillate ostiole. Conidiomata wall multi-layered,
with thick outer wall, composed of brown walled-cells of textura angularis, with thin,
hyaline, inner wall. Conidiophores 10–60 × 2–4 µm, long, cylindrical, branched,
hyaline, smooth-walled. Conidiogenous cells holoblastic, annellidic, simple,
integrated, determinate, hyaline. Conidia 12–17.5 × 3–6 µm ( x = 13.54 × 4.79 µm, n =
20), obovoid to fusiform, truncate at base, obtuse at apex, straight, with 3-transverse
septa, brown to dark brown at septa, constricted at the septa, eguttulate, medium
brown, hyaline to sub-hyaline at basal and apical cell, smooth-walled, with or without
tubular basal and apical appendages; basal appendage when present 6–15 µm,
unbranched; apical appendage when present unbranched, 8–25 µm.
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Culture characteristics: On PDA slow growing, attaining a diam. of 1.5 cm in 7
days at 18 ºC, white to light brown from above, pale brown from below, with sparse
mycelium, flat, uneven margin.
Material examined: ITALY, Province of Trento [TN], Marilleva 900 - Val di
Sole, on dead branch of Rosa villosa L. (Rosaceae), 29 July 2013, Erio Camporesi, IT
1392 (MFLU 15–3559, holotype); (HKAS isotype), ex-type living cultures
MFLUCC 14–0468, GUCC IT1392
Notes: Farr and Rossman (2015) list several Seimatosporium species which were
recorded from Rosa spp. Among these, only Seimatosporium rosae shows both apical
and basal appendages (Sutton 1980; Nag Raj 1993). Crous et al. (2014a) introduced S.
pistaciae Crous & Mirab which also has apical and basal appendages. Our collection
is morphologically distinct from both these species and the key is provided below to
distinguish the three species.
Molecular analysis shows our collection groups with S. pseudorosarum
(MFLUCC 14–0466), but the latter species lacks apical appendages. Norphanphoun et
al. (2015) introduced Seimatosporium physocarpi C. Norphanphoun et al. from
Physocarpi sp. (15–16 × 3.5–4.8 µm) which has both apical and basal appendages and
has conidial dimensions similar with our collection. However, our collection has
longer conidiophores (10–60 µm), while in S. physocarpi conidiophores are only up to
20 µm. The new taxon is phylogenetically distinct from Seimatosporium physocarpi
(Fig. 70) and it is thus introduced as a new species.
Key to distinguish Seimatosporium spp. with apical and basal appendages
1. Conidia longer than 17 µm................................................................................... S. pistaciae
1. Conidia shorter than 15 µm .................................................................................................. 2
2. Conidia12–17.5 × 3–6 µm ................................................................................................... 3
2. Conidia 10–15 × 3–4 µm .......................................................................................... S. rosae
3. Conidiophores up-to 20 µm ............................................................................. S. physocarpi
3. Conidiophores 10–60 µm ............................................................................... S. pseudorosae
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Fig. 72 Seimatosporium pseudorosae (holotype) a–c Appearrance of conidiomata on dead
branches of Rosa villosa d Cross section of conidiomata e Conidia baring conidiophore and
paraphyses f–h Different stages of conidiogenesis i–m Conidia. Scale bars: d = 200 µm, e–h =
20 µm, i–m = 10 µm.
Xylariales
Diatrypaceae
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Fig. 73 Phylogram generated from maximum likelihood analysis based on ITS sequence data
of the family Diatrypaceae. The new isolates are in red and ex-type strains are in bold. The
tree is rooted with Xylaria hypoxylon.
Cryptovalsa Ces. & De Not. ex Fuckel
Cryptovalsa is a common diatrypaceous genus known to occur on grapevines in
the family Diatrypaceae which was typified by C. protracta (Pers.) De Not. (Mostert
et al. 2004; Mehrabi et al. 2015). The genus was characterized by eutypoid
ascostromata, polysporous asci and allantoid ascospores (Spooner 1981; Vasilyeva
and Stephenson 2005; Trouillas et al. 2011). Currently, there are 58 epithets in Index
Fungorum (2016), while four species have been transferred to other genera in
Diatrypaceae, Massariaceae and Xylariaceae (Index Fungorum 2016). Molecular
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data are only available for C. ampelina (Nitschke) Fuckel and C. rabenhorstii
(Nitschke) Sacc. (Trouillas et al. 2011; Mehrabi et al. 2015; EBI 2016; NCBI 2016).
Cryptovalsa ampelina is the most studied species in Cryptovalsa (Nitschke 1867;
Mostert et al. 2004; Vasilyeva and Stephenson 2005; Luque et al. 2006; Martín et al.
2009; Trouillas et al. 2010; Trouillas et al. 2011; Mehrabi et al. 2015).
306. Cryptovalsa ampelina (Nitschke) Fuckel, Jb. nassau. Ver. Naturk. 23-24: 212
(1870) [1869-70]
Basionym: Valsa ampelina Nitschke, Pyrenomycetes Germa-nici 1, p. 156,
1867.
Index Fungorum number: IF 241474, Facesoffungi number: FoF 01800, Fig. 74
Saprobic on bark. Sexual morph Stromata poorly developed, immersed in bark, with
occasionally protruding perithecial necks, single or in groups, irregularly scattered.
Ascomata 510–580 µm high, 340–440 µm diam. (x̅ = 530 × 391 µm, n = 8), solitary to
gregarious, immersed, dark brown to black, arranged in a single layer, singly arising,
in rows globose to subglobose, often compressed, ostiolate, with cylindrical necks,
raising above the epidermis and forming black, blister-like areas, periphysate.
Peridium 35–45 µm wide, composed of two layers; outwardly comprising several
layers of thick-walled, dark brown to black cells of textura angularis, inwardly
comprising 3–5 layers of thin-walled, hyaline cells of textura angularis to textura
prismatica. Hamathecium comprising dense, 2–4 µm wide, hyaline, aseptate,
anastomosing paraphyses. Asci (98–)118–133(–146) × (7–)7–11(–14) µm ( x = 119 ×
9 µm, n = 30), polysporous, unitunicate, cylindric-clavate, long pedicellate, apically
rounded to truncate with indistinct, amyloid apical annulus. Ascospores (7–)7.5–9(–10)
× (1–)2–2.5(–3) µm, ( x = 8.3 × 2.4 µm n = 60), crowded, pale yellowish to pale
brown at maturity, allantoid–reniform, 1-celled, smooth-walled, with small guttules.
Asexual morph Coelomycetous, forming on MEA. Conidiomata 150–260 mm diam.,
pycnidial, superficial, solitary or aggregated, dark brown to black, globose to
subglobose, covering by yellow to light brown interwoven, thick-walled, hyphae.
Conidiophores 10–22 × 1.5–2 µm (x̅ = 18 × 2 µm, n = 10), septate, bicellately to
verticillately branched, arranged in dense palisades, cylindrical, hyaline, smooth,
arising from the base. Conidiogenous cells 8–14 × 1–2 µm (x̅ = 11 × 1.5 µm, n = 20),
holoblastic, sympodial to synchronous, straight or curved, subcylindrical, hyaline,
apically distorted on conidial secession. Conidia 16.5–20 × 1–1.5 µm (x̅ = 18.6 × 1.3
µm, n = 55), hyaline, cylindrical to filiform, unicellular, slightly curved, apically
rounded, with truncate base.
Culture characteristics: Ascospores germinating on MEA within 24 hours, germ
tubes produced at both ends cell, colonies on MEA reaching 4 mm diam. after 7 days
in darkness condition at 25 °C, medium dense, raised, circular with fimbriate edge,
fluffy to fairy fluffy, white from above, light yellowish from below, forming asexual
morph, with black, stromatic after 15 days.
Material examined: ITALY, Fiumana di Predappio, Province of Forlì-Cesena
[FC], on dead branch of Vitis vinifera L. (Vitaceae), 5 January 2015, E. Camporesi,
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(MFLU 16–0007, KUN-HKAS 93731, reference specimen designate here), living
culture, MFLUCC15–0139, KUMCC 16-0003).
Notes: Cryptovalsa ampelina is a pathogen of grapevines (Vitis vinifera L.) and
is abundant on pruned canes and necrotic wood of living plants (Mostert et al. 2004;
Luque et al. 2006; Trouillas et al. 2010; Pitt et al. 2013a). The species was reported as
a pathogen from South Africa, Australia, North East of Spain, California and Eastern
United States (Mostert et al. 2004; Vasilyeva and Stephenson 2005; Luque et al. 2006;
Trouillas et al. 2010; Pitt et al. 2013a). Cryptovalsa ampelina causes internal wood
discoloration, similar to that caused by Eutypa lata (Pers.) Tul. & C. Tul (Ferreira
1987; Mostert et al. 2004). However, the species can be distinguished from E. lata in
having polysporous asci and pigmented allantoid ascospores (Luque et al. 2006).
The asexual morph of Cryptovalsa ampelina has been reported in the
coelomycetous genus Libertella, which is characterized by sporodochium-like
conidiomata, hyaline, branched conidiophores, with hyaline, subcylindrical,
conidiogenous cells which proliferate sympodially and hyaline, filiform, slightly
curved to hamate, unicellular conidia, with a truncate, flattened base (Mostert et al.
2004; Luque et al. 2006). In this study, the asexual morph formed in culture on MEA
after 20 days. The characters of our taxon are similar to previous studies, although our
taxon differs due to its slightly smaller conidia.
Based on phylogenetic analysis of ITS gene dataset (Fig. 73), Cryptovalsa
ampelina clearly separates from Eutypa lata and clusters with Quaternaria quaternata
(GNF13, EL60C). However, C. ampelina can be distinguished from Q. quaternata by
its polysporous asci. Our strain (MFLU 15-0139) forms a well-supported clade (100%
ML) with other strains of C. ampelina (KHJ 20 and A 001) in the family
Diatrypaceae (Fig. 73). Our isolate is similar to the protolgue described by Nitschke
(1867) as well as Trouillas et al. (2010). Nevertheless, it differs from the type
protoloque in having larger asci (from Saccardo (1882), 75–90 × 8–9 versus
(98–)118–133(–146) × (7–)7–11(–14), this study) and slightly smaller ascospores
(from Saccardo (1882), 9–10 × 2.5 versus, (7–)7.5–9(–10) × (1–)2–2.5(–3), this
study). Therefore, we propose our new collection as a reference specimen.
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Fig. 74 Cryptovalsa ampelina (MFLU 16–0007, reference specimen) a Appearance of
stromata on host surface b Longitudinal section through stromata showing globose ascomata
embedded in stromatal tissues c Ostiole with periphysate ostiolar neck d Peridium e
Paraphyses f, g Asci h Immature ascus i Ascus with apical apparatus inconspicuously bluing
in Melzer’s reagent j–l Ascospores m Germinating ascospore n, o Culture in MEA, note n is
from above and o is from below. Scale bars: a = 200 µm, b = 500 µm, c = 30 µm, d = 50 µm,
e–g = 20 µm, h, i, m= 10 µm, j–l = 5 µm.
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Fig. 75 Culture of Cryptovalsa ampelina in PDA (MFLU 15–0139) a, b Conidiomata on the
culture c Hyphae on conidiomatal surface d Section of conidiomata e Conidiophore with
young conidia f Conidiogenous cells with conidia g Conidiophores h–j Conidia. (Note: f, i, j
with cotton blue) Scale bars: b = 200 µm, d= 20 µm, c, e–j = 5 µm.
307. Diatrype thailandica R.H. Perera, J.K. Liu & K.D. Hyde, sp. nov.
Index Fungorum number: IF 552008, Facesoffungi number: FoF 01797, Figs 76,
77
Etymology: The specific epithet thailandica refer to the country in which the
fungus was first collected.
Holotype: MFLU 15–3662
Saprobic on wood. Sexual morph Stromata 1–1.2 mm wide, scattered on host,
erumpent, arising through the cracks in bark epidermis, with 4 ascomata immersed in
a single stromata, comprising an outer, dark brown to black, small, tightly packed,
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thin parenchymatous cell layer, inner layer yellowish, loosely packed, with
parenchymatous cells, with ostioles opening to outer surface, appearing as black spots.
Ascomata 226–336 µm high, 177–235 µm diam., (x̅ = 282 × 209 µm, n = 20),
perithecial, immersed in stromatic tissues, aggregated, globose to subglobose,
narrowing towards the apex, pale brown, ostiolate. Ostiolar necks emerging separately,
short, immersed in only dark outer layer of stromata, conical, periphysate. Peridium
6.5–15 µm wide (x̅ = 11 µm, n = 20), comprising strata of 4–8 layers of cells of
hyaline to dark brown cells of textura angularis. Hamathecium comprising 2.2–4.5
µm wide (x̅ = 3 µm, n = 20), aseptate, paraphyses, longer than the asci, wider at the
apex. Asci 55–80 × 5–7 µm (x̅ = 67 × 6 µm, n = 25), 8-spored, unitunicate, with
narrow, long, thin-walled pedicel, with cylindrical, thick-walled, swollen upper
portion, apex flat, with J-, conspicuous apical apparatus. Ascospores 3.8–6.9 × 1–1.4
µm (x̅ = 5.4 × 1.2 µm, n = 20), multi-seriate to overlapping pale brown, allantoid to
cylindrical, unicellular, with small, fat globules at the ends, smooth-walled. Asexual
morph Coelomycetous, libertella-like, Mycelial clumps white. Conidiomata pycnidial,
0.4–1 mm diam., brownish yellow, becoming dark brown when mature, watery,
bubble-like, rounded, conidial masses forming from mycelial clumps. Pycnidia
superficial, solitary or aggregated, subconical, globose to subglobose, shiny, with
smooth surface, yellow, dark brown, comprising brown, thick-walled cells of textura
angularis. Conidiophores 12–16 µm high, 1.8–2.3 µm wide (x̅ =14 × 2.1 µm, n = 20)
branched, arising from pseudoparenchymatous cells or interwoven hyphae.
Conidiogenous cells 5.9–10 µm high, 1.1–1.8 µm wide (x̅ = 8.4 × 1.6 µm, n = 20),
cylindrical, in dense palisades, straight or curved, apically distorted or bearing
annellations. Conidia 14.2–18 × 0.7–1 µm (x̅ = 16.7 × 0.9 µm, n = 20), filiform,
curved or rarely straight, with flattened base and blunt apex, hyaline.
Culture characteristics: Fast growing, reaching 6.7 cm within 14 days on PDA,
at 25 °C, circular, flat, with diffuse margin, white, and becoming yellowish-white,
dull yellow to brownish with age.
Material examined: THAILAND, Doi Mae Salong, on stems of unidentified
plant, 12 March 2015, R.H. Perera, RHP 27 (MFLU 15–3662, holotype); ibid.,
HKAS 92497, isotype), ex-type living culture, MFLUCC 14–1210, CUMCC 15-0019.
Notes: Based on the phylogenetic analysis of ITS sequence data, Diatrype
thailandica form a separate branch as a sister group with Diatrypella and Diatrype
species. Previous studies by Trouillas et al. (2011) and Acero et al. (2004) suggested
that both Diatrypella and Diatrype are polyphyletic within the family. However
Diatrype thailandica is morphologically similar to the members of the genus Diatrype
in both sexual and asexual morph characteristics. In the phylogenetic analysis it has a
close relationship with D. macowaniana which was isolated from dead branches of
Cassina capensis in South Africa. Diatrype thailandica is different from D.
macowaniana in having yellow inner cells in the stromata, with smaller, pale brown,
mostly allantoid ascospores, and longer asci, while D. macowaniana is characterized
by stromata with white inner cells, larger, cylindrical ascospores and smaller asci.
Diatrypaceae is a taxonomically confused family and it is presently difficult to
segregate genera (Trouillas et al. 2011; Vasilyeva et al. 2006; Liu et al. 2015).
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Therefore, the placement of this isolate into the genus Diatrype may require
reconsideration in the future together with a revision for the entire family.
Fig. 76 Diatrype thailandica (holotype) a Herbarium material b–d Appearance of stromata
on host substrate. e Longitudinal section through stroma f Vertical section through stroma
showing ascomata g Close up of the ostiole h Close up of the peridium i Paraphyses j
Arrangement of asci k Ascus in Melzer’s reagent l Immature and mature asci m Ascospores n
Germinating ascospore. Scale bars: b = 2 mm, c–e = 500 µm, f = 50 µm, g–j = 20 µm, k = 100
µm, l = 10 µm, m = 20 µm, n, o = 10 µm.
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Fig. 77 Diatrype thailandica (holotype) a, b Conidiomata on PDA c Cross section of
conidioma d Conidia attached to conidiophores e Conidia f Sporulation on one month old
culture on PDA, 25 °C. Scale bars: a, b = 1 mm, c–e = 20 µm.
Xylariaceae
The family Xylariaceae is defined as one of the largest families of
pyrenomycetous fungi with unitunicate asci and pigmented ascospores. This family
comprises about 85 genera (Maharachchikumbura et al. 2015, 2016) with more than
1300 accepted species (Stadler et al. 2013). The majority of Xylariaceae are
saprotrophs on decaying wood, animal dung, fruits and seeds, leaves and herbaceous
stems, while some are endophytes of vascular plants and some are even associated
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with termite nests (Rogers 2000; Stadler 2011). Morphological characteristics of the
sexual morph, such as the stromata, perithecia, asci, ascospore, apical apparatus and
germ slit, or of the asexual morph, such as nodulisporium-like and
geniculosporium-like are used to delineate species. Phylogenetic analysis of
multi-gene sequence data (ITS, LSU, RPB2 and β-tubulin) has shown that
Xylariaceae comprising two major groups representing the subfamilies Xylarioideae
and Hypoxyloideae. The Xylarioideae comprises the genus Xylaria, and the asexual
morph is known to be geniculosporium-like. The Hypoxyloideae comprises four
subclades with the major subclade containing the genera Hypoxylon and
Annulohypoxylon and the second subclade consists of Daldinia, Entonaema and
Ruwenzoria and two small subclades of Rhopalostroma and Phylacia clustering
separately in the poorly supported tree. The asexual morphs are either
nodulisporium-like or virgariella-like (Stadler et al. 2013). The phylogenetic tree is
presented in Fig. 78.
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Fig. 78 Phylogram generated from RAxML analysis based on combined ITS, LSU, RPB2 and
β-tubulin sequenced data of species of Xylariaceae. Maximum Likelihood values equal or
greater than 50 are indicated above or below the nodes and branches. The tree is rooted to
Sordaria fimicola. New taxa are in blue and ex-type strains in bold.
Annulohypoxylon Y.M. Ju, J.D. Rogers & H.M. Hsieh
The genus Annulohypoxylon was introduced by Hsieh et al. (2005) with the type
species Annulohypoxylon truncatum (Schwein.) Y.M. Ju, J.D. Rogers & H.M. Hsieh
and 53 species are listed to date (Index Fungorum 2016). Annulohypoxylon is
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characterized by effused-pulvinate or pulvinate, glomerate stromata, sphaerical or
obovoid perithecia with a carbonaceous stromata layer, with KOH-extractable
pigments in most cases, cylindrical, stipitate asci with an apical apparatus and light- to
dark-coloured, ellipsoid or short fusoid, nearly equilateral ascospores, with narrowly
of broadly rounded ends and a germ slit, and perispore dehiscence or indehiscence in
KOH 10% (Hsieh et al. 2005). Molecular analysis showed this genus is closely related
with Hypoxylon with strong support. However, it differs from the Hypoxylon in
having a carbonaceous stromata layer, discretely enclosing each perithecium, and the
ostioles are always higher than the surrounding stromatal surface, usually encircled
with a distinct annulate disk (Hsieh et al. 2005). The phylogenetic tree for
Annulohypoxylon is presented in Fig. 79.
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Fig. 79 Phylogram generated from RAxML analysis based on ITS sequence data of
species of Annulohypoxylon. Maximum Likelihood values equal or greater than 50 are
indicated above or below the nodes and branches. The tree is rooted to Xylaria
hypoxylon. Newly introduced taxa in this study are highlighted in blue and ex-types
are in bold.
308. Annulohypoxylon albidiscum J.F. Zhang, J.K. Liu, K.D. Hyde & Z.Y. Liu, sp.
nov.
Indexfungorum number: IF 551809, Facesoffungi number: FoF 01812, Fig. 80
Holotype: MFLU 15–3883
Etymology: From the Latin albus referring to white, and discus meaning disc, in
reference to the morphology of stromata, which have a white, flattened
truncatum-type disc, encircling the ostioles.
Saprobic on decorticated wood. Sexual morph Stromata 1.5–7.5 × 1–4.5 ×
0.2–0.5 cm, ( x = 5.2 × 2.8 × 0.35 cm), glomerate, pulvinate to effused-pulvinate,
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with conspicuous perithecial mounds, surface shiny black, sphaerical to
hemisphaerical, carbonaceous, blackish granules immediately beneath surface and
between perithecia, with KOH-extractable pigments greenish-olivaceous (90).
Ostioles conical, papillate, encircled with a white, flattened truncatum-type disc.
Perithecia 0.4–0.8 mm diam., sphaerical. Peridium laterally 43–51 µm thick,
composed of carbonaceous, thick-walled, dark brown to black cells of texura
angularis. Hamathecium comprising long, septate paraphyses, 4.5–5.3 µm wide at the
base, 1.5–2.5 µm wide at the apex, with hyaline, guttulate cells. Asci (61–)77–87(–97)
3.5–5 µm ( x = 83.5 4.3 µm, n = 20), 8-spored, unitunicate, cylindrical, long
pedicellate, with a wedge-shaped, J+, subapical apparatus, 0.7 1.6 µm. Ascospores
7.1–7.9(–8.4) (3.4–)3.6–4.2(–4.8) µm ( x = 7.7 3.8 µm, n = 30), uniseriate, 1-celled,
inequilaterally ellipsoidal, with narrowly rounded ends, light brown to brown, with or
without guttules when young, germ slit straight, running along the entire spore-length
on flattened side Asexual morph Undetermined.
Culture characteristics: Ascospores germinating on WA within 12 h and germ
tubes produced from ends. Colonies growing fast on PDA, reaching 7 cm in 7 days at
25–28 °C, whitish colonies, azonate with diffuse margins, reverse at first whitish and
turning light brown after 5 days.
Material examined: THAILAND, Chiang Rai, Muang District, Mae Chang Hot
Spring, on limestone outcrops, on decorticated wood of unidentified host, 25
November 2014, JinFeng Zhang, ZJF–16 (MFLU 15–3883, holotype), ex-type living
culture, MFLUCC 15–0645.
Notes: This is a typical Annulohypoxylon species with pulvinate to
effused-pulvinate stromata, long cylindrical asci and pale brown, inequilaterally
ellipsoidal ascospores. As well it is reminiscent to A. stygium (Lév.) Y.M. Ju et al. and
A. nitens (Ces.) Y.M. Ju et al., regarding the stromatal characters. However, A.
albidiscum differs from A. stygium in having larger perithecia (0.4–0.8 mm vs.
0.2–0.3 mm), a wider ascal apical apparatus (1.6 µm vs. 0.7 µm) and having white,
flattened truncatum-type disc encircling the ostioles. In addition, the KOH-extractable
pigments of this specimen is greenish-olivaceous (90), whereas, the latter is greenish
olivaceous (90) or dull green (70) (Ju and Rogers 1996). Annulohypoxylon albidiscum
is distinct from A. nitens (Ces.) because the latter has a vinaceous reddish tone in the
younger stages (Ju and Rogers 1996), as well as the asci of A. albidiscum are
significantly shorter than the latter (77–87 µm long vs. 110–140 µm long). The
phylogenetic analysis showed that the A. albidiscum clustered with other
Annulohypoxylon species and is phylogenetically closely related to A. bovei var.
microspora (J.H. Mill.) Y.M. Ju et al., A. moriforme var. microdiscus (Y.M. Ju & J.D.
Rogers) Y.M. Ju et al. and A. purpureonitens (Y.M. Ju & J.D. Rogers) Y.M. Ju et al.,
but they have different morphological characters.
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Fig. 80 Annulohypoxylon albidiscum (holotype) a Stromata habit on wood b Stromata in
side view c Cross section of the stromata showing perithecia d Section of peridium e
Germinating ascospore f Long, hyaline paraphyses g–h Asci with ascospores in water i Ascus
in Melzer’s reagent, showing the J+, subapical ring j–o Ascospores. Scale bars: a = 500 µm, b,
c = 200 µm, d, f–i = 10 µm, j–o = 3 µm.
Astrocystis Berk. & Broome
Astrocystis was introduced based on A. mirabilis Berk. & Broome, a
bamboo-inhabiting xylariaceous taxon. The stellate or coronate appearance of the
stromata is characteristic feature of the Astrocystis species (Læssøe and Spooner
1994). The genus is mostly confined to monocotyledons such as bamboo and has
superficial, uniperitheciate stromata, which may develop beneath the host cuticle. The
asci are relatively short-stipitate, with a relatively small, amyloid and stopper-shaped
ascal apparatus (Smith et al. 2001). Index Fungorum (2015) listed 24 Astrocystis
species epithets.
309. Astrocystis thailandica Daranagama & K. D. Hyde, sp. nov.
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4010
4011
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Indexfungorum Number: IF 551727, Facesoffungi number: FoF 01637, Fig. 81b
Etymology: Referring to the country, Thailand where the species was collected.
Holotype: MFLU 15–3525
Saprobic on bamboo clumps. Sexual morph Stromata superficial, gregarious,
black, shiny, smooth, carbonaceous, multi-peritheciate, with 2–3 perithecia, 650–1075
× 250–375 µm ( x = 720 × 310 µm, n = 10), globose to hemisphaerical, carbonaceous,
with black, stellate area of mixed host and stromatic material encircling the base of
stromata. Ostioles papillate, black. Peridium >50 µm wide, comprising several thick
layers of compressed cells, black. Hamathecium comprising numerous, 2 µm wide,
filamentous, septate, paraphyses, embedded in a gelatinous matrix. Asci 88–125 ×
8.2–12.2 µm (x = 93.5 × 10.5 µm, n = 25), 8–spored, unitunicate, cylindrical–clavate,
short pedicellate, apically rounded, with a J+, wedge-shaped apical apparatus, 4.5–5 ×
2.5–3 µm. Ascospores 17–24 × 6.2–7.5 µm (x = 20 × 6.8 µm, n = 25), overlapping
uniseriate, dark brown, equilaterally ellipsoidal, unicellular, germ slit full-length or ¾
of the length, with a conspicuous mucilaginous sheath, forming slimy caps at both
ends. Asexual morph Undetermined.
Culture characteristics: Colonies on Difco OA plates at 25–28 oC reaching 5 cm
edge Petri-dish in 2 weeks, at first whitish, felty, azonate, with diffuse margins, after 3
weeks become citrine; reverse turning light brown.
Material examined: THAILAND, Chaing Mai Province, road to Wat Pa Dang,
on clumps of fallen bamboo clumps, 14 August 2014, Anupama Daranagama AXL
323 (MFLU 15–3525, holotype, HKAS 92485, isotype), living culture, MFLUCC
15–0009, KIBCC.
Notes: Astrocystis thailandica displayed a close relationship with A. eleiodoxae
A. Pinnoi et al., which was also encountered in Thailand on submerged petioles of
Eleiodoxa conferta (Pinnoi et al. 2010). However A. thailandica differs from A.
eleiodoxae because of its unique characters such as, superficial stromata with black,
stellate stromatic material encircling the base, shorter and wider asci and ascospores
with a thick, conspicuous mucilaginous sheath forming slimy caps at both ends.
According to the phylogenetic analysis the species clustered with other Astrocystis
species with 93 bootstrap support forming a monophyletic clade. As well as the
phylogenetic analysis of the genus (Fig. 81a) confirmed the placement of Astrocystis
thailandica with a high bootstrap support, as a distinct species from other Astrocystis
species.
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Fig. 81a Phylogram generated from RAxML analysis based on ITS sequenced data of
Astrocystis. Maximum Likelihood values equal or greater than 50 are indicated above
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or below the nodes and branches. The tree is rooted to Xylaria hypoxylon. Newly
introduced taxa in this study are highlighted in blue.
Fig. 81b Astrocystis thailandica (holotype) a Stromata on host surface b Multi-peritheciate
ascomata c Papillate ostiole d, e Mature asci f Apical apparatus bluing in Melzer’s reagent g
Ascospore with straight germ slit h, i Developmental stages of ascospores with sheath. Scale
bars. a = 2000 µm, b, c = 500 µm, d–j = 10 µm.
310. Camporesia W.J. Li & K.D. Hyde, gen. nov.
Index Fungorum number: IF 552005; Facesoffungi number: FoF 01822
Etymology: Named after the collector Erio Camporesi
Type species: Camporesia sambuci W.J. Li & K.D. Hyde
Saprobic on dead stems of plant host. Sexual morph Undetermined. Asexual morph
Coelomycetous. Conidiomata pycnidial, globose, superficial to subepidermal,
separate, unilocular, thick-walled, ostiolate. Peridium composed of cells of texura
angularis, with inner layers hyaline gradually merging with the outer dark brown
layers. Conidiophores short, unbranched, hyaline, formed from the innermost layer of
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wall cells. Conidiogenous cells hyaline, phialidic, ampuliform, smooth-walled, with a
periclinal wall thickening at the tip. Conidia pale brown, fusiform, rounded at both
ends, 2–3-septate, smooth-walled.
Notes: The asexual morph of Xylariaceae has mainly been linked to
hyphomycetous (i.e. genicolosporium-like and nodulisporium-like) (Ju and Rogers
1996). Subsequently, the asexual structures were extended to libertella-like
coelomycetous genera (Ju et al. 1993, Stadler et al. 2013, Senanayake et al. 2015).
Camporesia sambuci was collected form Sambucus ebulus L. and is characterized by
globose pycnidia and pale brown, fusiform conidia with 2–3-septa. Camporesia
sambuci is morphologically distinct from libertella-like species, which have hyaline,
long slender falcate conidia. The phylogeny of the family Xylariaceae is reconstructed
based on combined gene (LSU, ITS, RPB2 and β-tubulin) analysis, showing that
Camporesia sambuci clusters away from any other genera in Xylariaceae (Fig. 78).
Thus Camporesia is introduced as a novel genus in this study.
311. Camporesia sambuci W.J. Li & K.D. Hyde, sp. nov.
Index Fungorum number: IF 552006 Facesoffungi number: FoF 01823, Fig. 82
Etymology: Named after the host genus Sambucus
Saprobic on dead stems of Sambucus ebulus. Sexual morph Undetermined.
Asexual morph Coelomycetous. Conidiomata 100–150 µm high, 200–250 µm diam.,
pycnidial, globose, superficial to subperidermal, separate, unilocular, thick-walled,
ostiolate. Peridium 30–50 µm wide, composed of 6-8 layers, with outer 4–5 layers of
dark brown and inner 2-3 layers of pale brown to hyaline cells texura angularis.
Conidiophores short, unbranched, hyaline, formed from the innermost layer of wall
cells. Conidiogenous cells 10–15 × 2–4 µm, phialidic, ampuliform, hyaline, smooth,
with a periclinal wall thickening at the tip. Conidia 8–15 × 4–5 µm ( x = 10 × 4.5 µm;
n = 20), pale brown, fusiform, rounded at both ends, 2–3-septate, smooth.
Culture characteristics: Colonies fast growing on PDA, reaching 20 mm diam.
after one week at 20–25 oC, with circular margin, whitened, flattened, felt-like, with
filamentous, dense, aerial mycelium on the surface, reverse similar in colour.
Material examined: ITALY, Province of Arezzo [AR], near Passo della
Consuma, on dead stem of Sambucus ebulus (Adoxaceae), 19 June 2012, Erio
Camporesi, IT–450 (MFLU 15–3905, holotype); ex-type living culture, MFLUCC
13–0203, ICMP 20775.
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Fig. 82 Camporesia sambuci (holotype). a Herbarium specimen b Appearance of black
coniodiomata on the host c, d Vertical sections of conidiomata h Section of peridium f–j
Conidiophores, conidiogenous cells and developing conidia k Germinated spore l–p Conidia.
Scale bars c–d = 100 µm, e = 20 µm, f–j = 5 µm, k = 10 µm, 1–p = 5 µm.
Durotheca Læssøe et al.
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The genus Durotheca was introduced by Læssøe et al. (2013) with D. depressa
Læssøe & Srikitik. as type species and D comedens (Ces.) Læssøe & Srikitik. and D.
rogersii (Y.M. Ju & H.M. Hsieh) Læssøe & Srikitik. transferred from Theissenia
based on morphology and molecular phylogeny. Durotheca is characterized by
stromata which are erumpent through bark or wood, initially covered in white pruina,
highly carbonaceous tissue, globose to cylindrical perithecia, with or without
columella, and filiform and distantly septate paraphyses. Mature asci deliquescent
early and young asci are clavate, without an apical apparatus. Ascospores are
moderate to very thick-walled, pale to medium brown, ellipsoid–oblong to allantoid,
and with or without a germ slit. The phylogenetic tree is presented in Fig. 83.
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Fig. 83 One of four MPTS inferred from combined β-tubulin and α-actin gene dataset
generated with maximum parsimony and Bayesian analysis. Maximum parsimony bootstrap
value greater than 50% and Bayesian posterior probabilities greater than 0.95 are given above
and below each clade, respectively. The internodes that are highly supported by bootstrap
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(100%) and posterior probabilities (1.00) are shown as a thicker line. New taxa are in blue and
ex-type strains in bold.
312. Durotheca macrostroma Srikitik., Wongkanoun & Luangsa-ard, sp. nov.
Index Fungorum number: IF 551628, Facesoffungi number: FoF 02033, Fig. 84
Etymology: based on the large stroma when compare with other Durotheca
species.
Holotype: BBH39917
Saprobic on bark of dead Castanopsis acuminatissima (Blume) A.DC. Sexual
morph Stromata superficial, solitary, subglobose 1 cm thick × 2.3–2.4 cm diam.,
stromata surface smooth, chalky white, creamy, owing to the presence of a thin pruina,
when mature surface greyish green (28C3), crust and tissue highly carbonaceous, with
beveled margin. Perithecia completely immersed, usually monostichous,
globose-ovoid, 1.8–2 mm high × 0.8–1 mm diam. Ostioles umbilicate/lower than
stromatal surface. Paraphyses not observed. Asci 8-spored, deliquescing, mature asci
not observed, young asci 77–93.5 × 11–13 µm, cylindrical, and long stalked, apical
apparatus lacking, and no reaction with Melzer’s reagent. Ascospores light brown,
unicellular, oblong to allantoid in side view, smooth-walled, (13–) 14–16 (–17.5) ×
(5–) 6–7 (–8) µm (xˉ = 15.03 × 6.67 µm, n = 54), germ slit lacking; perispore
non-dehiscent in 10% KOH. Asexual morph Undetermined.
Culture characteristics: Colony on PDA reaching 49–51 mm diam. in 10 days,
the culture produced botryose structures from the type and paratypes after 4 weeks.
Mycelia initially white and fluffy, turning to yellow brown after 2 weeks.
Material examined: THAILAND, Chaiyaphum, Phu Khiao Wildlife Sanctuary,
12 August 2015, on Castanopsis acuminatissima wood (Fagaceae), P. Srikitikulchai
& S. Wongkanoun (BBH39917, holotype); ex-type living culture, BCC78380.
Distribution: Only known from a single site in Phu Khiao Wildlife Sanctuary in
northeastern Thailand.
Notes: Molecular phylogenetic analyses of combined β-tubulin and α-actin gene
datasets based on maximum parsimony and Bayesian analysis has placed D.
macrostroma in Durotheca. Durotheca macrostroma differs from other Durotheca
species in having a large stroma; the shape of D. macrostroma is subglobose, 10 mm
thick, while other species are widely effused-pulvinate and are not over than 2.5 mm
thick. The ascospores of D. macrostroma are smaller than other Durotheca species. In
addition, the phylogenetic tree supported the position of D. macrostroma as closely
related to D. rogersii with 100% bootstrap support. They differ in the shape of
stromata and ascospore shape and size. Durotheca rogersii has a widely
effused-pulvinate 2.5 mm thick stroma (Ju et al. 2007), while that of D. macrostroma
is very thick (10 mm) and subglobose. Ascospores of D. rogersii have very thick
walls (3–4.5 µm) and are larger (25–36 × 19–24 µm) than D. macrostroma, but all
lack a germ slit.
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Fig. 84 Durotheca macrostroma (holotype) a Stroma on bark b Stroma surface and ostiole,
arrow: ostioles C Ascospore release on apex of ostioles, arrow: black spore mass d Young
asci e Perithecium f Botryose structures produced in culture g–i Ascospores j Colony on PDA
plate after 2 weeks. Scale bars: d = 5 µm, e = 0.25 mm, g–i = 5 µm, f = 10 µm, j = 1 cm.
Halorosellinia Whalley et al.
Halorosellinia is a monophyletic genus with a single species Hypoxylon
oceanicum S. Schatz which is characterized by uniperitheciate ascomata which are
immersed in a pseudostroma (Whalley et al. 1999).
313. Halorosellinia rhizophorae Dayarathne, Jones E.B.G. & K.D. Hyde, sp. nov.
Index Fungorum number: IF 551858, Facesoffungi number: FoF 01811, Fig. 85
Etymology: Name referring to the host genus Rhizophora.
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Holotype: MFLU 15–0183
Saprobic on dead root of Rhizophora sp. submerged in marine habitats. Sexual
morph Pseudostromata 1.5–2.5 × 0.9–1 mm ( x = 2 × 0.8 mm; n = 10),
semi-immersed, pulvinate to hemisphaerical, in clusters of up to 20 uni peritheciate
pseudostromata, surface black, carbonaceous, lacking ascomatal projections. In
section pseudostromata comprises host cells, filled with amorphous black fungal
material. Ascomata 350–380 × 96–114 µm ( x = 365 × 105 µm; n = 10), immersed in
pseudostroma, subglobose to hemisphaerical, black, ostioles papillate. Peridium
25–38 µm wide, two-layered, outer layer of cells of textura angularis, black, fusing at
the outside with the pseudostromata, inner layer of elongate cells, dark brown to black.
Paraphyses 1–3 µm wide, hyaline, abundant, persistent, aseptate. Asci 165–270 ×
12–18 µm ( x = 217.5 × 15 µm; n = 20), overlapping, 6–8-spored, cylindrical, long
pedicellate, unitunicate, with J+, rectangular apical ring. Ascospores 24–36 × 10–15
µm ( x = 30 × 12.5 µm; n = 20) overlapping uniseriate, light brown when immature,
dark to opaque brown when mature, more or less equilaterally ellipsoid, ventral side
varying in degree of convex curvature, upper end broadly rounded, lower end slightly
pointed, 1-celled, 1–2-guttulate, without appendages, germ slit on the ventral side,
straight, ¾ total length of spore. Asexual morph Undetermined.
Culture characteristics: Colonies on PDA at 25–28 °C reaching 5 cm in 7 days,
whitish, zonate with diffuse margins, reverse at first whitish and turning light brown
after 3–4 days.
Material examined: THAILAND, Krabi Province, Krabi, 8°25'52" N, 98°31'42"
E, 0 m asl., on submerged root of Rhizophora sp., 7 December 2014, Monika
Dayarathne, KRB018 (MFLU 15–0183, holotype, HKAS 92496 isotype); ex-type
living culture, MFLUCC 15–1281, KUMCC 16-0004.
Notes: Distinctive features of Halorosellinia include a poorly developed
pseudostromata which lack extractable pigments in KOH, asci with a relatively large
apical apparatus, that become dark blue in Melzer’s reagent and ascospores with a
prominent, straight germ slit on the ventral side (Whalley et al. 1999). The new
species, Halorosellinia rhizophorae is clearly different from the type, Halorosellinia
oceanica (S. Schatz) Whalley et al. in lacking ascomatal projections (Table 4). They
are approximately similar in ascospore morphology being 1-celled, light brown to
opaque brown, more or less equilaterally ellipsoid, with the ventral side varying in the
degree of convex curvature, the upper end broadly rounded, lower end slightly
pointed, and with 1–2 guttules. A Geniculosporium-like asexual morph was reported
from the ex-type culture of H. oceanica (Whalley et al. 1999). However, an asexual
morph was not found associated with H. rhizophorae on host substrate or in culture
media. Halorosellinia rhizophorae also has morphological affinities to Nemania
maritima having more or less inequilaterally ellipsoid ascospores with germ slits.
However, ascospores of H. rhizophorae are larger than that of N. maritima [9–12 ×
5–6(–6.5) μm]. When considering the differences between these taxa, in H.
rhizophorae the ascomata are immersed in a pseudostroma, asci have a long stipe with
a well-developed apical ring. In N. maritima ascomata are aggregated and submerged
in the carbonaceous stroma and asci are short-stalked. Maximum likelihood analysis
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of combined ITS and LSU sequence data confirmed the placement of H. rhizophorae
within the family Xylariaceae, where it forms a sister clade to the type, H. oceanica
with 81% bootstarp support (Fig. 78). However, H. rhizophorae is distantly placed
from Nemania spp. in the phylogenetic analyses.
Table 4 Comparison of the measurments of Halorosellinia oceanica and H.
rhizophorae.
Characters
H. oceanica
H. rhizophorae
Pseudostromata
0 ± 4–0 ± 8 mm in diam.
1.5–2.5 × 0.9–1 mm ( x
= 2 × 0.8 mm; n = 10)
Peridium
25–35 µm
25–38 µm
Asci
177–219 µm
165–270 µm
Ascospores
(17 ± 9–)18 ± 7–26(–28)¬7 ± 24–36 × 10–15 µm ( x
5–13(–13 ± 5) µm
= 30 × 12.5 µm; n = 20)
Paraphyses
2–2 ± 5 µm wide
1–3 µm wide
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Fig. 85 Halorosellinia rhizophorae (holotype) a, b Appearance of pseudostromata on host b
Horizontal section through pseudostroma c Section through pseudostromata d Peridium e
Apical apparatus stained blue in Melzer’s reagent f–h Asci i Paraphyses j–m Ascospores.
Scale bars: b = 200 µm, c = 100 µm, d, e = 20 µm, f–i = 50 µm, j–m = 20 µm.
Hypoxylon Bull.
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The genus Hypoxylon is one of the largest genera within the family Xylariaceae
with currently 159 accepted taxa. Its species are distributed world-wide with the
highest diversity in the tropics. Their sexual morph is usually associated with dead
hardwood and can often be found along with the respective asexual morph. The
generic concept is mainly based on the monograph by Ju & Rogers (1996), which was
later improved by Hsieh et al. (2005). In most cases the stromata contain large
quantities of secondary metabolites, which show characteristic colour reactions in
potassium hydroxide solutions, a feature that is used to discriminate between species.
Moreover, Stadler and coworkers employed analytical chromatographic methods
(HPLC) to identify the stromatal compounds and to generate respective secondary
metabolite profiles (Kuhnert et al. 2014). These chemical profiles are often species
specific and help to validate the erection of new species. The phylogenetic tree is
presented in Fig. 86.
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Fig. 86 Phylogenetic relationships among Hypoxylon lilloi and related Xylariaceae as inferred
from β-tubulin gene sequences. Likelihood (ML) bootstrap support values above 50%, from
1000 RAxML replicates are assigned to the tree topology of the most likely tree found by
RAxML. The tree is rooted to Creosphaeria sassafras. Species names are followed by strain
numbers. Ex-type strains are highlighted in bold and new isolates are in blue.
314. Hypoxylon lilloi Sir, Lambert & Kuhnert, sp. nov.
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4294
Mycobank number: MB 814982, Facesoffungi number: FoF 02034, Figs 87–89
Etymology: In honor of Dr. Miguel Lillo, a pioneer biologist in Tucuman
province (Argentina).
Holotype: ARGENTINA, Salta, Depto. Anta, Parque Nacional El Rey, 30 April
2014, Sir & Hladki 739 (LIL, ex-type culture STMA 14142)
Differs from Hypoxylon vogesiacum by livid purple stromatal pigments in 10%
KOH, as well as in having an amyloid apical apparatus and smaller ascospores.
Sexual morph Stromata effused-pulvinate, 14–30 mm long × 5–26 mm broad ×
1 mm thick; plane or with inconspicuous perithecial mounds; surface Purplish Gray
(128) or Vinaceous Grey (116); pruinose; brown to dark red granules immediately
beneath surface and between perithecia; with KOH-extractable pigments Livid Purple
(81), the tissue below the perithecial layer inconspicuous, black, 0.2–0.5 mm thick.
Perithecia obovoid to lanceolate-tubular 0.5–0.8 mm high × 0.2–0.3 mm diam;
ostiolar openings lower than the stromatal surface, umbilicate with white area
surrounding ostioles. Paraphyses 2–4 µm wide at base, tapering above asci. Asci
8-spored, cylindrical, 92–134.5 µm total length, the spore-bearing parts 56–46 µm
long × 5–6.5 µm broad, the stipes 40–82.5 µm long; with amyloid, discoid apical
apparatus 0.7–0.9 µm high × 1.9–2.3 µm broad. Ascospores brown to dark brown,
unicellular, ellipsoid-inequilateral, with narrowly rounded ends, slightly curved,
7.4–8.9 (9.7) × 3.2–4.2 µm (n = 60, Me = 8.3 × 3.8 µm); with straight germ slit
spore-length on convex side; perispore dehiscent in KOH; with inconspicuous
coil-like ornamentation by light microscopy, revealing reticulate ornamentation by
SEM (5000×); epispore smooth. Asexual morph In culture, Conidiophores with
virgariella-like branching pattern, usually borne on aerial hyphae, hyaline, smooth.
Conidiogenous cells hyaline, smooth, 10–27 × 1–2.5 µm. Conidia 4–5 × 1.5–2.5 µm,
ellipsoid, hyaline, smooth-walled.
Culture: Colonies on OA covering Petri dish in 2 week, at first whitish,
becoming Olivaceous Grey (121) to Dull Green (70), felty, zonate, with entire margin;
reverse Apricot (42), later turning Dark Green (21) in places. Sporulating regions
scattered over entire surface of colony.
Secondary metabolites: Stromata of this species contain two unknown major
metabolites in its stromatal extracts (Fig. 89) in addition to some other yet unknown
minor metabolites, besides binaphthalene tetrol (BNT).
Additional material examined: ARGENTINA, Jujuy Province, Depto. Santa
Bárbara, Reserva provincial Las Lancitas, 13 May 2012, Sir & Hladki 278 (LIL);
Salta, Depto. Anta, Parque Nacional El Rey, 30 April 2014, Sir & Hladki 744 (LIL,
culture STMA 14143).
Notes: Hypoxylon lilloi, which was found in the course of a study on Xylariaceae
of the Argentine cloud forest “Las Yungas” (Sir et al. 2016) might be confused with H.
vogesiacum (Pers. ex Curr.) Sacc. due to their similar purplish gray or vinaceous grey
stromatal surfaces. However, H. lilloi differs in having livid purple KOH-extractable
pigments, smaller ascospores and in lacking a dotted band in the centre of the
ascospores. This new taxon resembles the group of species with purplish
KOH-extractable pigments, such as H. lienhwacheense Y.M. Ju & J.D. Rogers, H.
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lividicolor Y.M. Ju & J.D. Rogers, H. lividipigmentum F. San Martín et al. and H.
texcalense F. San Martín et al. Those can be easily differentiated from H. lilloi by the
colour of the stromatal surface and granules. In addition H. lienhwacheense has
smaller ascospores (6–7.5 × 3–3.5 µm vs. 7.4–9.7 × 3.6–4.6 µm) and a smooth
perispore. Hypoxylon lividicolour differs in having longer perithecia (0.5–1.3 ×
0.2–0.4 mm vs. 0.5–0.8 × 0.2–0.3 mm), larger ascospores (11–12.5 × 4.5–5 µm vs.
7.4–9.7 × 3.6–4.6 µm) and sporothrix-like conidiogenous structures and H.
lividipigmentum can be differentiated by its larger ascospores (10–15 × 4.5–6 µm vs.
8.5–10 × 4–4.5 µm) and nodulisporium-like conidiogenous structures. In comparison
with H. texcalense, the latter has also much larger ascospores (17–24 × 6.5–9.5 µm vs.
7.4–9.7 × 3.6–4.6 µm), and lack ascal apical rings and nodulisporium-like
conidiogenous structures.
The type of secondary metabolites produced in the stromata seems to be a unique
feature of the species, because they were not detected in more than 1000 studied
specimens. Only BNT could be identified, which is common in hypoxyloid genera of
the Xylariaceae.
In the phylogenetic reconstruction based on β-tubulin gene sequences (Fig. 86),
H. lilloi forms a separated clade. The latter is located between the H. fragiforme clade
and H. lenormandii clade. Besides huge morphological differences of those species
compared to H. lilloi, they can be easily distinguished by their orange
KOH-extractable pigments due to the production of azaphilones such as the
mitorubrins (H. cinnabarinum Henn.) Y.M. Ju & J.D. Rogers, H. fragiforme (Pers.) J.
Kickx f., H. jecorinum Berk. & Ravenel, H. rickii Y.M. Ju & J.D. Rogers) and the
lenormandins (H. lenormandii Berk. & M.A. Curtis; cf. Kuhnert et al. 2016).
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Fig. 87 Hypoxylon lilloi (holotype) a Stromatal habit b Close-up view of stromatal surface
with white area surrounding umbilicate ostioles (black arrow) c Bown granules beneath
surface and between perithecia (white arrow) d Asci e extractable pigments in 10% KOH f
Section through stroma showing perithecia and dark red granules (white arrow) g Apical ring
bluing in melzer´s iodine reagent (black arrow) h Ascospores showing germ slit (white arrow)
i Ascospores showing perispore dehiscent in KOH (black arrow) j Perispore showing
inconspicuous ornamentation k, l Ascospores showing reticulate ornamentation on perispore
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4334
under SEM. Scale bars: a = 5 mm, b, c and f = 0.5 mm, d = 20 µm, g, h, i and j = 10 µm, k = 2
µm, l = 200 nm.
Fig. 88 Hypoxylon lilloi (ex-type) Culture of on OA after 3 weeks a top view b reverse c, d
Conidiophores with virgariella-like branching patterns e Conidia. Scale bars: c–e = 5 µm).
200
Intens.
[mAU]
250
300
350
400
450
500
550
Wavelength [nm]
218
600
200
Intens.
[mAU]
250
300
350
400
450
500
550
Wavelength [nm]
550
Wavelength [nm]
216
500
258
500
400
400
1
3+4
256
300
300
200
200
300
0
200
Intens.
[mAU]
1000
300
100
100
0
250
300
350
400
450
500
550
226
Wavelength [nm]
200
Intens.
[mAU]
250
300
350
400
450
500
500
800
400
2
600
400
5
300
200
296
200
100
434
0
0
Intens.
[mAU]
x105
4
1
2
3
2
3
1
BNT
4
5
0
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0
5
10
15
20
25
Time [min]
Fig. 89 Stromatal HPLC-UV profiles of H. lilloi derived from EBS278 and corresponding
DAD spectra of the unknown main metabolites.
Rosellinia De Not.
The genus is typified by Rosellinia aquila (Fr.) Ces. & De Not. and was
introduced to accommodate species with uniperitheciate, superficial, ostiolate
stromata seated on a subiculum with cylindrical, stipitate asci usually with an amyloid
apical apparatus and produce dark brown ascospores (Petrini 1992). Rosellinia is a
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relatively large genus in Xylariaceae. Index Fungorum (2016) includes 496 records
under the name However according to the world monograph by Petrini (2013) only
142 species are accepted, of which 37 species are described as new species.
315. Rosellinia chiangmaiensis Daranagama & K. D. Hyde, sp. nov.
Index Fungorum Number: IF 551728, Facesoffungi number: FoF 01638, Fig. 90b
Etymology: Referring to the province Chiang Mai, where the species was
encountered.
Holotype: MFLU 15–3524
Saprobic on dead dicotyledonous wood. Sexual morph Stromata globose, with a
pointed top, 1220–1400 × 800–1080 µm ( x = 72.5 × 4.8 µm, n = 20), chestnut brown,
shiny, smooth, solitary, in small groups, uniperitheciate, surrounded by woolly to felty,
pale yellow subiculum, confined to the stroma base, black entostroma, reduced at the
base. Ostioles black, distinctively papillate, pointed. Ascomata globose, 400–500 ×
500–600 µm ( x = 467 × 560 µm, n = 20). Peridium thick-walled, > 70 µm,
carbonaceous. Hamathecium comprising long, dehiscent, filamentous, few paraphyses,
2µm wide, longer than asci. Asci 150–200 × 4.5–6.4 µm ( x = 172 × 5.2 µm, n = 20),
8-spored, unitunicate, cylindrical, short pedicellate, apical narrowly rounded, with a
J+, inverted hat-shaped, apical apparatus, upper width 4–6 µm, lower width 2–3 µm,
with rounded bulge at upper rim. Ascospores 70–90 × 7–10 µm ( x = 84 × 9 µm, n =
20), overlapping uniseriate, dark brown, elongate fusiform, with acute ends, with thin
mucilaginous sheath, germ slit and appendages absent.
Culture characteristics: Colonies on Difco OA plates at 25–28°C reaching 5 cm
edge of Petri-dish in 2–3weeks, at first citrine, felty, azonate, with diffuse margins,
reverse turning yellow.
Material examined: THAILAND, Chiang Mai Province, garden of Mushroom
Research Center, on decorticated bark of a fallen log, 17 August 2014, Anupama
Daranagama, AXL 342 (MFLU 15–3524, holotype, HKAS 92486, isotype), ex-type
living culture, MFLUCC 15–0015, KIBCC.
Notes: Rosellinia chiangmaiensis is reminiscent to R. macrosperma Speg. and R.
procera Syd. & P. Syd. because its large length: width ascospore ratio, lacking germ
slits and generally large stromata more than 1 mm high (Petrini 2013). However the
new species possess longer ascospores with thin mucilaginous sheath with acute ends
and a white to pale yellow subiculum restricted to the stromatal base. These characters
make this species unique from other known, morphologically similar species.
According to the description by Petrini (2013) this new species belongs to the R.
emergens group, which is a phylogenetically heterogeneous group. The reconstructed
phylogenetic trees for the family Xylariaceae (Fig. 83) and the genus Rosellinia (Fig.
90a) confirmed the placement of Rosellinia chiangmaiensis with high bootstrap
support.
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Fig. 90a Phylogram generated from RAxML analysis based on ITS sequenced data of
Rosellinia. Maximum Likelihood values equal or greater than 50 are indicated above
or below the nodes and branches. The tree is rooted with Xylaria hypoxylon. Newly
introduced taxa in this study are highlighted in blue and ex-type strains are in bold.
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Fig. 90b Rosellinia chiangmaiensis (holotype) a Ascomata in host surface b Papillate
ostioles c Side view of ascomata d Cross section through stroma e Vertical section of stroma f,
g Asci with J+, apical apparatus in Melzer’s reagent h, i Asci in water j, k Ascospores in
water. Scale bars: a = 500 µm, b, c = 1000 µm, d, e = 200 µm, f, g = 10 µm, h, i = 50 µm, j, k
= 30 µm.
Ascomycota, genera incertae sedis
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Petrakia Syd. & P. Syd.
Petrakia is typified by Petrakia echinata and characterized by having dark
brown, rounded to oval, muriform conidia bearing cellular, long, hyaline appendages.
Butin et al. (2013) described sexual morph of P. echinata based on field collections,
culture studies and ITS sequence data and assigned it to the genus Mycodidymella.
Following the rulings of the current ICN, we propose to use the oldest name, Petrakia
over Mycodidymella. The phylogenetic tree is presented in Fig. 91 which shows that
Petrakia probably belong in Dothideomycetes genera, incertae sedis.
Fig. 91 Best scoring RAxML tree of Petrakia echinata and related species obtained from
analysis of LSU sequence data. RAxML bootstrap support values (equal to or greater than
50% based on 1.000 replicates) are shown at the nodes. The tree is rooted to Mycosphaerella
punctiformis CBS 113265. New taxa are in blue and species for which obtained sequences are
based on type material have names in bold.
316. Petrakia echinata (Peglion) Syd. & P. Syd., Annls mycol. 11(5): 406 (1913)
Index Fungorum number: IF 192652, Facesoffungi number: FoF 01821
≡ Epicoccum echinatum Peglion, Malpighia 8: 459 (1895)
Parasitic on living leaves of Acer pseudoplatanus L., forming numerous, conspicuous
rounded, black, sporodochia. Sexual morph Mycodidymella (Butin et al. 2013).
4420
4421
4422
4423
4424
4425
4426
4427
4428
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4449
4450
4451
4452
Asexual morph Sporodochia 90–110 µm high, 100–150 µm diam., dark brown to
black, solitary, scattered to gregarious, occasionally confluent, superficial, erumpent,
elliptical or irregular in outline, with a basal stroma variably developed, 20–30 µm
thick, composed of cells of textura angularis to textura globulosa. Conidiophores
reduced to conidiogenous cell arising from the uppermost cells of the basal stroma.
Conidiogenous cells 12–35 × 3–10 µm, hyaline to pale yellow, integrated, annellidic,
with 2–3 annellations, cylindrical, thick-walled, smooth. Conidia 22–45 × 12–32 µm
( x = 32 × 25 µm, n = 30), rounded to oval or broadly ellipsoidal, muriform, with
multi-transverse and longitudinal septa or oblique septa in the central zone,
constricted at septa, thick-walled, smooth, at first hyaline, later becoming brown or
dark brown, bearing 8–33 × 3–9 µm, cellular, long appendages; appendages, arising as
a tubular extension of the body of the conidium, unbranched, narrow and attenuated,
subhyaline, cylindrical, smooth-walled.
Culture characteristics: Colonies on PDA slow growing, reaching 15 mm diam.
after one week, circular, white to pale grey, velvety, felty, sparse, aerial, surface
smooth with crenate edge, filamentous; reverse black at the central zone, white at the
margin.
Material examined: ITALY, Province of Forlì-Cesena [FC], Camposonaldo,
Santa Sofia, on living leaves of Acer pseudoplatanus L. (Sapindaceae), 20 February
2013, Erio Camporesi IT-1570 (MFLU 15–7568, reference specimen designated
here), living culture MFLUCC 15–0582.
Notes: In the phylogenetic analysis, strain MFLUCC 15–0582 is closely related
to Petrakia echinata (Fig. 92). The comparisons of ITS sequence data from both
strains show 100% similarity. Morphologically, strain MFLUCC 15–0582 has similar
sporodochia and conidia characteristics to those of P. echinata, and the only
distinguishing character is the dimension of the conidia. Strain MFLUCC 15–0582
has slightly larger conidia (22–45 × 12–32 µm, versus 16–28 × 18–22 µm) than P.
echinata. However, the differences noted here similarly reflect reasonable
intraspecific variation. Petrakia echinata has been reported as an pathogen in Austria,
Caucasus, Germany Switzerland and the Czech Republic (Kirisits 2007, Butin et al.
2013), and this is first record of the species in Italy. Details of the conidiogenous cells
are also provided.
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Fig. 92 Petrakia echinata (MFLU 15–7568, reference specimen) a Herbarium specimen b, c
Appearance of black sporodochia on the host d Vertical section of sporodochia e–h
Conidiogenous cells and developing conidia i Germinating conidium j–m Conidia n, o
Culture on PDA note o reverse. Scale bars: b = 200 µm, c = 100 µm, d = 50 µm, e, f = 5 µm, g,
h, m = 10 µm, i–l = 20 µm, n, o = 10 mm.
Contributions to Basidiomycota
Agaricomycetes
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Agaricales
Agaricaceae
Agaricaceae is the type family of the order Agaricales, which is distributed
widely around the world. This family contains 1340 species in 85 genera (Kirk et al.
2008). Species in this family mostly have a fleshy basidiome, with pileus and stipe,
some of them also have an annulus, such as the genera Agaricus and Micropsalliota.
Besides the agaricoid, secotioid and gasteroid taxa are also included in this family.
The phylogenetic tree for Agaricaceae is presented in Fig. 96.
Agaricus L.
The genus Agaricus (Agaricaceae) is a well known group with many cultivable
species. Its systematics has been well-studied in recent years (Parra 2008, 2013; Zhao
et al. 2011; Chen et al. 2012, 2015a; Wang et al. 2015b; Zhao et al. 2016). There are
some sections of this genus, such as sections Sanguinolenti and Sppisicaules, that
have been revealed to be polyphyletic (Zhao et al. 2011; 2016). However, section
Minores has been stable since it was introduced by Fries (1874), based on its
morphology and molecular phylogeny (Zhao et al. 2011; Parra 2013; Lebel 2013).
Section Minores is characterized by relatively small-sized basidiomes, a simple
annulus, the surfaces of the pileus and stipe often discolouring yellow on scratching, a
context yellow discolouring on exposure, and a pleasant odour (Heinemann 1978;
Parra 2013). Historically the species of section Minores have been limited in number.
Recent research has revealed a high biodiversity of species in Europe (Parra 2013),
Thailand (Liu et al. 2015), Australia (Lebel 2013) and China (He et al. 2015). Herein
we add two more new species of this section from China. The phylogenetic tree for
Agaricus is presented in Fig. 93.
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Fig. 93 Phylogeny of species of Agaricus section Minores generated from Bayesian analysis
of ITS sequence data rooted with Agaricus arvensis. Bayesian posterior probability (PP)
values above 90% and parsimony bootstrap support (BS) above 50% are given at the
internodes (PP/BS). New taxa are in blue ex-types in bold.
317. Agaricus coccyginus M.Q. He & R.L. Zhao, sp. nov.
Fungal Names number: FN 570238, Facesoffungi number: FoF 02035, Fig. 94
Etymotogy: the epithet “coccyginus” refers to the purple red squamules on the
cap of this species.
Holotype: HMAS 275416
Macroscopical characters: Pileus 35–110 mm in diam., umbonate at disc,
parabolic when young, then convex, finally plane with age; margin straight, decurved,
sometimes little exceeding; surface dry, covered by tiny fibrils on the whole cap,
appressed, denser on the disc and broken into radially triangular squamules toward
margin, purple red, brown, or reddish brown on the lighter background. lamellae free,
crowded, 3–8 mm broad, white or pink at first, then grayish brown, brown finally.
Context white, fresh, 2–6 mm thick at disc, white, turns yellow on cutting first, then
reddish brown after several minutes. Annulus membranous or cortinate-membranous,
simple, pendant, white, 4–10 mm in diam., smooth on both sides of surface. Stipe
64–160 × 4–9 (base 9–21) mm, cylindrical or slightly clavate, hollow, white, smooth
or fine fibrils below the annulus, always with rhizomorphs. Basidiome surface
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strongly discolouring yellow when touching or bruising, then reddish brown after
several minutes. Odour of strong almond.
Macrochemical reaction: KOH reaction strongly yellow; Schäffer’s reaction
orange.
Microscopical characters: Basidiospores 5.5–6.5 (– 6.8) × 4.3–4.5 µm, [x = 6 ±
0.3 × 3.8 ± 0.2, Q = 1.4–1.8, Qm = 1.6 ± 0.1, n = 20], ellipsoid to elongate, smooth,
thick-walled, brown, no germ pore. Basidia 14.1–19 × 5.6–7.8 µm, clavate, hyaline,
4-spored, smooth. Cheilocystidia 16–60 × 9.2–22 µm, mostly pyriform and clavate,
sometimes oblong, pheropedunculate, rarely septa at base, smooth, hyaline, with
yellow pigment inside. Pleurocystidia absent. Pileipellis a cutis composed of 5.9–14.5
µm in diam. hyphae, smooth, cylindrical, brown, constricted at septa.
Habitat: Solitary on the soil of forest.
Material examined: CHINA, Tibet, Bomi, Baga Village, 26 July 2012,
Su-ShengYu, ZRL2012485 (HMAS 275416, holotype); Tibet, Milin County,
Nanyigou, Li Guang-Ping ZRL2012597 (HMAS275413,); Yunnan Province, Weixi
County, 4 August 2014, He Mao-Qiang, Dai Rong-Chun, Su Sheng-Yu, ZRL2014354
(HMAS 275412,), ZRL2014364 (HMAS275414), ZRL2014415 (HMAS275420),
ZRL2014430 (HMAS 254484).
Notes: see under Agaricus luteofibrillosus.
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Fig. 94 Agaricus coccyginus a, b Basidiome c, e Annulus d Discoloration on stipe f
Cheilocystidia g Basidia h Basidiospores i Hyphae of Pileipellis Scale bars: a from holotype
ZRL2012485, b, d from ZRL2012597, c from ZRL2014415, e from ZRL2012576. Scale bars:
a = 3 cm, b–d = 2 cm, e = 1 cm, f, g, i = 10 µm, h = 5 μm.
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318. Agaricus luteofibrillosus M.Q. He, L.J. Chen & R.L. Zhao, sp. nov.
Fungal Names number: FN 570234, Facesoffungi number: FoF 02036, Fig. 95
Etymotogy: the epithet “luteo” refer to the yellow colour; and “fibrillosus” refers
to the fibrils on the pileus and stipe.
Holotype: HMAS 254487
Marcoscopical characters: Pileus 35–94 mm in diam., parabolic at first, then
convex, finally plane, sometimes with slightly subumbonate disc with age; margin
slightly decurved when young, then straight; surface dry, fibrillose, yellowish brown
against white to light brown background, appressed, denser at disc, then broken into
triangular fibrillose squamules towards the margin. Context 3–8 mm thick at disc,
fresh, white, and yellow discolouring on exposure. Lamellae 4–8 mm broad, free,
crowded, pink when young, then brown when mature. Annulus simple, membranous,
pendant, white, lower surface floccose with light brown tiny squamose. Stipe 60–141
× 5–14 (base 8–25) mm, white, cylindrical, base clavate or subbulbose, surface
smooth and white above the annulus, fibrillose squamose or floccose and light brown
below the annulus, hollow. Basidiome surface yellow discolouring on touching or
bruising. Odour of almond.
Macrochemical reaction: KOH reaction strongly yellow; Schäffer’s reaction
orange.
Microscopical characters: Basidiospores 5–6.5 (–7.2) × 3–4.2 µm [x = 5.8 ± 0.4
× 3.4 ± 0.2, Q = 1.5–2, Qm = 1.7 ± 0.1, n = 20], ellipsoid to cylindric, smooth,
thick-walled, brown, no germ pore. Basidia 14–18 × 5.6–7.3 µm, clavate, hyaline,
4-spored, smooth. Cheilocystidia 9.4–28 × 6.4–17 µm, mostly globose and clavate,
sometimes pyriform and pheropedunculate, septa at base sometimes, smooth, hyaline,
some with yellow pigment inside. Pleurocystidia absent. Pileipellis a cutis composed
of hyphae of 3.2–13.2 µm in diam., smooth, cylindrical, light brown, constricted at
septa. Annulus composed of hyphae with 3–9.5 µm in diam., hyaline, cylindrical, not
constricted at septa.
Habitat: Solitary on soil of forest.
Material examined: CHINA, Yunnan Province, Baoshan, Gaoligong Mountain,
Wanzi Village, He Mao-Qiang ZRL 2013484 (HMAS 254487, holotype); Yunnan
Province, Yongde County, Pingtian Village, Li Guang-Ping ZRL 2012359 (HMAS
275419); Yunnan Province, Cangyuan County, Nanban Village, Zhao Rui-lin ZRL
2012121 (HMAS 254486), ZRL 2012200 (HMAS 275415).
Notes: In the phylogenetic tree (Fig.3), the proposed new species A. coccyginus
and A. luteofibrillosus are represented by two clades respectively with strong PP and
BS support. Their phylogenetic positions are also clearly distinguished from other
known species in section Minores. In morphology, they both have related larger
basidiomes which the cap reaching 110 mm in diam. There are only two species with
such large-sized basidiomes in section Minores, one is A. brunneolus (J.E. Lange)
Pilát and the other is A. megalosporus J. Chen et al. Agaricus brnneolus is the most
similar species to A. coccyginus. They both have the same shape of cap, stipe and
same colour of fibrils on the basidiome. Also, under the microscope they have the
similar cheilocystidia. There are some distinguishable autapomorphies between these
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two species. Agaricus coccyginus has the longer basidiospores than those of A.
brunneolus (length 4.5–6.2 µm). The yellow pigment of cheilocystidia in A.
coccyginus is also another difference from A. brunneolus. Agaricus megalosporus is
the most similar species to A. luteofibrillosus, because both species have similar
basidiomes, they both have coloured fibrils on the cap, annulus and stipe. Both have
the same size of basidospores, but Agaricus luteofibrillosus has a yellowish brown cap,
while in A. megalosporus it is purplish brown. Under the microscope they have
different cheilocystidia: in A. megalosporus they are broadly clavate to pyriform, and
white, while in A. luteofibrillosus they are pheropedunculate, septa at base and contain
yellow pigment.
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Fig. 95 Agaricus luteofibrillosus a, b Basidiome c Annulus d Fibrils on cap e Basidiospores f
Cheilocystidia g Basidia h Hyphae of pileipellis. Scale bars: a from holotype ZRL2013484, b
from ZRL2012359, c, d from ZRL2012121. Scale bars: a = 5 cm, b = 3 cm, c, d = 1 cm, e =
5μm, f–h = 10 µm.
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Clarkeinda Kuntze
The genus Clarkeinda belongs to the family Agaricaceae, and was circumscribed
by Kuntze (1891). According to the Dictionary of the Fungi the widespread genus
contains five species and Index Fungorum lists 14 records (Kirk et al. 2008; Clements
1909; Index Fungorum 2016). Species in this genus, especially Clarkeinda trachodes,
are only distributed in south and southeast Asia (Yang 1991; Kuntze 1891; Hosen and
Ge 2011).
Fig. 96 Phylogeny of Clarkeinda trachodes and satellite genera in the Agaricaceae based on
analysis of ITS sequence data, inferred by maximum likelihood (ML) analysis. Numbers at
internodes refer to confidence estimates based on 100 rapid ML bootstraps (only those >50
are indicated). Clarkeinda trachodes from Sri Lanka is highlighted. Leucoagaricus barssii
and Leucoagaricus leucothites are outgroup taxa. New sequences are in blue and ex-type and
reference specimens are in bold.
319. Clarkeinda trachodes (Berk.) Singer, Lilloa 22: 413, 1951.
Facesoffungi number: FoF 01844, Figs 97, 98
Description: Basidiomes medium to large, fleshy. Pileus 120 mm in diam.,
hemisphaerical when young, and becoming convex to applanate at maturity; pellicle
on the cap brown to coffee or chocolate brown, thin when young and thick when
mature, and brown to grayish brown at maturity; the whole surface except the pellicle
area covered with grayish brown to vinaceous brown squamules, with numerous,
small, loosely floccose, brown squamules; context up to 8–9 mm thick in the center of
the pileus, white, instantly turning reddish with exposure. Lamellae free and distant
from the stipe, white to dirty white when young, turning to olive brown when mature,
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becoming reddish brown after bruised, crowded with lamellulae, entire margin,
concolorous. Stipe 140 × 45 mm, central, subcylindrical, fistulose in mature
specimens; surface dirty white to white at the apex, light brown to brown towards the
base, glabrous above the annulus, lower half densely covered with minute, brown,
furfuraceous squamules. Annulus present on the upper part of the stipe but not the top,
up to 20 mm, thick, membranous and remaining up to maturity, adaxial part glabrous
with fine longitudinal striate but abaxial part rough with squamules. Volva presents,
grayish, dirty white to white, membranous, usually closely appressed to stipe and
eventually inconspicuous. Basidiospore deposit not obtained. Basidia 17–28 × 5.5–9
μm, mostly clavate to subclavate, thin-walled, tetrasporic, but seldom 1-, 2- or
3-spored, bearing four short sterigmata, hyaline, smooth, lacking incrustations, clamp
connections absent. Basidioles narrowly clavate to clavate. Hymenophoral trama
interwoven, hyphae cylindrical to slightly inflated, up to 14 μm wide, thin-walled,
hyaline, and without clamp connections. Basidiospores (Fig. 98b) (5–)5.5–6 (–7) ×
(3.5–)3.9–4(–4.5) μm, mean Q = 1.4–1.5, ovoid, occasionally broadly ellipsoid to
ellipsoid, glabrous, thick-walled, apiculus eccentric, apex or germinating pore
prominent and truncate with slightly depressed, olive brown to dark, umber brown in
deposit, dextrinoid in Melzer’s solution, not metachromatic in Cresyl blue.
Cheilocystidia 25–33 × 10.5–15.5 μm, abundant, scattered to more or less crowded,
narrowly clavate, clavate to broadly clavate, obpyriform, hyaline, thin-walled, smooth,
lacking incrustations, sometimes with long pedicel and narrow. Pleurocystidia absent.
Pileipellis consisting of short branching chains of 4–7 cells, slightly interwoven,
terminal cells 12–23 × 8–14.5 μm, dull brown vacuolar pigment inside the cells in
glycerin, water and 5% KOH solutions, thin-walled, clavate, cylindrical, obpyriform
to fusiform or spindle-shaped in rare cases, occasionally branching with lateral cells
that are mostly clavate, basal cells nearly subglobose to clavate or cylindrical.
Habit, habitat, distribution: The basidiomes of C. trachodes normally fruit as
isolated individuals or in groups of two in disturbed habitats and at forest edges. Our
collection was collected on grassland in Royal Botanic Gardens, Peradeniya, Sri
Lanka. It is also known from China, India, Indonesia, Bangladesh and Malaysia.
Material examined: Sri Lanka. Central Province: Peradeniya, Royal Botanic
Gardens, 7°15'35.03"N 80°36'4.07"E, elev. 590 m, 15 July 2009, Samantha C.
Karunarathna (MFLU 10–0139, reference specimen designated here)
Notes: Clarkeinda trachodes is distinguished by its large basidiome size,
prominent chocolate or coffee brown to dark brown pellicle on the pileus disc surface,
presence of an annulus, olive brown to umber brown spore deposit, slightly
thick-walled basidiospores with a truncate apex, and a context that changes from
white to reddish brown when exposed. Since Berkeley (1847) first described the
species from Sri Lanka, it has been reported from south and Southeast Asia by Petch
and Bisby (1950, as Chitoniella), Leelavathy et al. (1981), and Pegler (1985, 1986).
Yang (1991) has also reported it from the tropical region of Yunnan, China. This is
the first report with the molecular phylogenetic confirmation after Berkeley (1847)
first described this from Sri Lanka. We therefore designate it as a reference specimen.
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Fig. 97 Clarkeinda trachodes (MFLU 10–0139, reference specimen) a Basidiomes in the
field b Pellicle on the cap c Longitudinal section of the basidiome d Veil. Scale bars: a–d = 10
cm.
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Fig. 98 Clarkeinda trachodes (MFLU 10–0139, reference specimen) a Basidia with
basidioles b Basidiospores c Cheilocystidia d Pileipellis. Scale bars: a = 30 µm, b = 10 µm, c
= 15 µm, d = 20 µm.
Amanitaceae
Amanita Pers.
Amanita Pers. is a widespread basidiomycete genus, with about 700 described
species (Tulloss and Yang 2016, http://www.amanitaceae.org). It is divided into two
subgenera, Amanita and Lepidella (E.-J. Gilbert) Veselý. The subgenus Amanita
includes sections Amanita, Caesareae Singer, and Vaginatae (Fr.) Quél., while the
subgenus Lepidella includes sections Amidella (E.-J. Gilbert) Konrad & Maubl.,
Lepidella, Phalloideae (Fr.) Quél., and Validae (Fr.) Quél. (Yang 1997; Yang et al.
2004). Most Amanita species are known to form ectomycorrhizal (ECM) associations
with trees. The phylogenetic tree of Amanita is presented in Figs 99 and 100.
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Fig. 99 Phylogram inferred by Maximum Likelihood analysis of LSU sequences. Bootstrap
support values greater than 50% are indicated above the nodes. New taxa are in blue and
species for which obtained sequences are based on type material have names in bold. The tree
is rooted with Limacella glioderma.
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Fig. 100 Maximum likelihood tree depicting infrageneric relationships of Amanita based on
nuclear ITS dataset. ML and MP bootstrap values ≥ 70% are shown above branches.
Sequences derived from three new toxic species are in bold.
320. Amanita atrobrunnea Thongbai, Raspé & K.D. Hyde, sp. nov.
Index Fungorum number: IF 551652, Facesoffungi number: FoF 02070, Fig. 101
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Etymology: the epithet refers to the dark brown colour of the pileus
Holotype: MFLU 15–1415
Pileus 120 mm in diam., conic to paraboloid when young, then plano-convex,
becoming convex and broadly umbonate when mature, dark brown to chestnut brown
(6F7, 6F8), darker in the center, paler and becoming teak brown to leather brown (6F5)
towards the margin, minutely rimose, sub-viscid when wet; margin lacking striations,
slightly appendiculate, sometimes with scattered annulus remnants; context 1 mm
thick at mid-radius, white. Lamellae free, white, crowded, up to 8 mm high;
lamellulae attenuate, with two to three series. Stipe 170 × 15 mm, slender, slightly
tapering upwards, white to pale yellowish, finely fibrillo-squamulose; context white,
solid, unchanging when bruised. Bulb 15–25 mm wide, inconspicuous, subfusiform,
white (1A1). Volva limbate, slightly firm, up to 20 mm high, white (1A1). Annulus
membranous, easily broken, white. Odour absent.
Lamellar trama bilateral; mediostratum 30–35 µm wide, composed of ellipsoid
to fusiform, 35–45 × 10–18 µm cells, mixed with abundant, filamentous 3–6 μm wide,
branching hyphae. Subhymenium 20–35 µm thick, with two to three layers of
subglobose to irregularly-shaped cells, 12–25 × 10–15 µm. Basidia 36–41 × 9–12 µm,
4-spored, clavate, thin-walled; sterigmata 4–6 µm long. Basidiospores 7.3–8.3–9.5 ×
5.4–6.6–7.8 µm, Q = 1.15–1.26–1.46 (N = 40), broadly ellipsoid to ellipsoid,
thin-walled, colourless, amyloid, smooth, with small apiculus. Lamellar edge
composed of numerous, subglobose, (15–25 × 8–18 µm) cells, and rare filamentous,
thin-walled, hyaline, 3–9 µm wide hyphae. Pileipellis 90–100 µm thick, composed of
two distinct layers, the upper layer gelatinized, made up of radially arranged,
thin-walled, filamentous, 3–8 µm wide, colourless hyphae, with inflated, sometimes
cylindrical, rarely subglobose to elliptical terminal cells; the lower layer mostly
non-gelatinized, composed of filamentous, sometimes branching, 4–10 μm wide
hyphae with pale brown pigment, mixed with abundant inflated cells. Velar remnants
from stipe base composed of thin-walled to slightly thick-walled, filamentous, 3–8 µm
wide hyphae, mixed with abundant inflated cells, with yellowish to pale brown
intracellular pigments. Annulus composed of thin to slightly thick-walled,
filamentous, 3–8 μm wide, branching hyphae, mixed with ellipsoid to subglobose,
hyaline, inflated, thin-walled cells. No clamps observed in any tissue.
Habitat: Terrestrial in forest dominated by Fagaceae species.
Material examined: THAILAND, Chiang Mai Province, Doi Saket District,
Sub-District Tepsadet, N18° 57’ 1.0016” E99° 20’ 1.0452”, 30 June 2014, collector
B. Chuankid, BZ–2014–09 (MFLU 15–1415, holotype)
Notes: Amanita atrobrunnea is a member of Amanita subgenus Lepidella (J.-E.
Gilbert) Veselý, section Lepidella (Bas 1969). Remarkable features of A. atrobrunnea
are the dark brown pileus, the broad umbo at the disc, the slender basidiocarp, the
absence of membranous velar remnants on the pileus, even when young, the abundant
inflated cells in the pileal surface and the broadly elipsoid to ellipsoid basidiospores.
The most morphologically similar species are A. manginiana sensu W.F. Chiu and A.
pseudoporphyria Hongo, which share several characters with A. atrobrunnea, such as
an inconspicuous bulb, dark pileus, and velar remnants on the pileus consisting of
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inflated cells (Zhang et al. 2010). However, A. atrobrunnea can easily be
distinguished from A. manginiana and A. pseudoporphyria by its distinctive umbonate
pileus at maturity. In addition, the inflated cells of the pileipellis, a key character of A.
atrobrunnea, are not present in the other species. Like A. atrobrunnea, A.
pallidorosea P. Zhang & Zhu L. Yang possesses a conspicuous umbo, but the pallid
rose colour of latter is very different. Amanita manginiana and A. pseudoporphyria
were initially placed in section Phalloideae (Hongo 1982, Yang 1997, Zhang et al.
2004, Zhang et al. 2010). However, recent phylogenetic analyses clearly showed that
both species belong to section Lepidella (Cai et al. 2014). Our molecular phylogenetic
analysis indicates that A. atrobrunnea is a sister species to A. manginiana and A.
pseudoporphyria.
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Fig. 101 Amanita atrobrunnea (holotype) a–c Basidiome d Radial section of pileipellis e, f
Basidiome g Basidia and subhymenium h Basidiospores. Scale bars: a, b = 8 cm, d = 20 µm, e,
f = 30 mm, g = 20 µm, h = 10 µm
321. Amanita digitosa Boonprat. & Parnmen, sp. nov.
Index Fungorum number: IF 551619, Faceoffungi number: FoF 02069, Fig. 102
Etymology: The specific epithet refers to Amanita with abundant digitate cell
types among other elements of the volva.
Holotype: BBH 32154
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Pileus 13.5–29 mm, paraboloid when young, convex to applanate with age,
smooth, yellowish brown 5(D–E) 8 at disc, towards half of pileus and pale yellow
3(A)4 in the middle of the pileus to margin, or the whole pileus yellowish brown
3(A)4, smooth from disc towards the half of pileus and striate from the middle of
pileus towards the margin, with dry and even margin. Pileus context off white, soft.
Lamellae free, unequal, subsistent, broad, fimbriate, lamella edge and face pale
yellow 3(A)4. Stipe 4.5–6 × 21–53 mm, central, cylindrical to tapering from base to
apex, yellowish white 1(A)2, soft, context reaction yellow with 3%KOH, base
bulbous: width 12–16 mm. Annulus not observed. Volva white membranous saccate.
Basidiospores 8–10 × 7–9 [x = 8 ± 0.65 × 9 ± 0.65 µm, Q = 1.13 ± 0.01, n = 25
spores, 1 collection] subglobose, smooth, hyaline, inamyloid, thin-walled, sometimes
with wart-like to network-like interior ornamentation. Basidia 30–37.5 × 10.5–12.5
µm, clavate with 2 and 4-spores, clamp connection absent, smooth, hyaline, inamyloid,
thin-walled. Basidioles 16–28 × 4.7–9.5 µm, clavate, smooth, hyaline, inamyloid,
thin-walled. Pleurocystidia 31–34 × 5.5–9.4 µm, clavate, smooth, hyaline, inamyloid,
thin-walled. Cheilocystidia absent. Lamellae trama divergent, composed of broadly
clavate to broadly ellipsoid cells, smooth, hyaline, dextrinoid, thin-walled, base of
hymenial layer directly arising from a few layers of cellular cells connected to trama
element. Pileipellis composed with cutis of repent hyphae, cylindrical, smooth,
hyaline, inamyloid, thin-walled. Stipilipellis composed with cutis of repent hyphae,
3–5 µm diam., smooth, hyaline, inamyloid, thin-walled. Stipe trama composed of two
types of element: repent hyphae and broadly clavate to broadly ellipsoid hyphae,
smooth, hyaline, inamyloid, thin-walled. Volva composed of three types of elements:
apex 19–21 × base 6 µm of digitate cells, 16–68 × 2.5–8.9 µm of clavate cells and
21–32 × 10.5–23 µm of broadly clavate to broadly ellipsoid cells, smooth, hyaline,
inamyloid, thin-walled. Clamp absent in all parts of basidiomata.
Notes: Amanita digitosa differs from A. subfrostiana Zhu L. Yang (Yang 1997)
in having brown and smaller basidiomata, while in A. subfrostiana they are red over
the disc to orange at the margin. Micro-characters include pleurocystidia, while these
are absent in the protologue of A. subfrostiana.
Habitat: Terrestrial in mixed forest.
Material examined: THAILAND, Si Sa Ket Province, Phu Sing District, Khok
Tan Tambon, 3 September 2012, collector SRRT Team, Bureau of Epidemiology,
Department of Disease Control Ministry of Public Health (BBH 32154, holotype).
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4806
Fig. 102 Amanita digitosa (holotype) a Basidiomata b Basidiospores c(i-ii) Basidia d Basidia
with basidioles e Pleurocystidia f(i-ii) Veil trama. Scale bars: a = 10 mm, b-f = 10 µm.
322. Amanita gleocystidiosa Boonprat. & Parnmen, sp. nov.
Index Fungorum number: IF 551614, Faceoffungi number: FoF 02071, Fig. 103
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Etymology: The specific epithet refers to Amanita with abundant of yellow
gleocystidium, ‘gleocystidium’ (n, neuter = versiform cystidia which have granular
content) + ‘-osus’ (adjA suffix = abundant)
Holotype: BBH31903
Pileus 22–45 mm diam. at first, first convex to parabolic when young, expanding
to applanate with age, sometimes depressed, sulcate, sticky, moist, colour ranges from
dark brown 8(F)5–8 at disc to grayish yellow 1(A)3–5 at margin when young; olive
yellow 2–3(C–E)6–8 at disc to yellowish white 2–3(A)2 at margin with age,
sometimes dark brown 8(F)5–8 at disc to grayish yellow 1(A)3–5 at margin with age,
with striate and even margin. Pileus context off white, 2–3 mm thick, soft and moist.
Lamellae free, broad, average, 3 series, sub-distant, yellowish white 2–3(A)2. Stipe
75–100 × 6–9 mm, central, tapered from base to apex, clavate-bulbous base, fistulose,
longtitudinal striate, pale orange to orange white 5(A)2–3 with grayish orange striate
5(B)3–6 after bruising. Annulus with single layer, pale yellow to brown, apical and
partial veil still intact when young, many of disappearing with age but few present at a
center of stipe. Volva constricted, adherent with flaring margin, white.
Basidiospores 7–10 (–11) × 7–10 µm [x = 8.76 ± 0.91 × 8.12 ± 0.13 µm, Q =
1.07 ± 0.10, n = 25 spores per collection, 2 collections], globose subglobose, smooth,
hyaline, inamyloid, thin-walled. Basidia 27–41 × 9.5–12.5 µm, clavate 2-spored,
clamp connection absent, smooth, hyaline, inamyloid, thin-walled. Basidioles 18–21 ×
6.5–7.5 µm, clavate, smooth, hyaline, inamyloid, thin-walled. Pleurocystidia with two
types of clavate and lanciolate, smooth, hyaline, inamyloid, thin-walled, clavate
pleurocystidia 30–35 × 7.5–12.5 µm, lanciolate pleurocystidia 35–50 × 8.5–12.5 µm.
Cheilocystidia apex 12–13 µm × middle 5–7 × base 3–4 µm, broadly clavate to
pyriform, smooth, hyaline, inamyloid, thin-walled. Gleocystidia abundant among
basidioles, pleurocystidia and cheilocystidia, shape and size dependent on the position
of appearance, contains yellow granules, smooth, hyaline, inamyloid, thin-walled.
Lamellae trama divergent, broadly clavate to broadly ellipsoid, smooth, hyaline,
dextrinoid, thin-walled, base of hymenial layer directly arising from a few layers of
cellular cells, which connects to the trama element. Pileipellis composed of cutis of
repent hyphae, smooth, hyaline, inamyloid, thin-walled. Stipilipellis composed with
cutis of repent hyphae, 2.5–5 µm diam., smooth, hyaline, inamyloid, thin-walled.
Stipe trama composed of two types of element: repent hyphae and broadly clavate to
broadly ellipsoid hyphae 73–105 × 31–34 µm, smooth, hyaline, inamyloid,
thin-walled. Volva composed of two types of elements: 22–31 × 3.5–7 µm of clavate
cells and 14–28 × 6.3–11.5 µm of broadly clavate to broadly ellipsoid cells, smooth,
hyaline, inamyloid, thin-walled. Clamp absent in all parts of basidiomata.
Habitat: Terrestrial in mixed forest.
Material examined: THAILAND, Phetchabun Province, Lom Kao District, Na
Sang Tambon, 28 May 2012, collector SRRT Team, Bureau of Epidemiology,
Department of Disease Control Ministry of Public Health (BBH31903, holotype);
Ibid., BBH31901, BBH31902 and BBH31908, paratypes, all collections were from
Phetchabun Province, Lom Kao District, Na Sang Tambon, collector SRRT Team,
4850
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4859
Bureau of Epidemiology, Department of Disease Control Ministry of Public Health,
28 May 2012, specimen scattered around temple.
Notes: Amanita gleocystidiosa is similar to A. sychnopyramis f. subannulata
Hongo (Yang et al. 2001) in having a similar macroscopic morphology and
basidiospore shape and size, but A. gleocystidiosa differs from A. sychnopyramis f.
subannulata in having pleurocystidia and cheilocystidia, while those two types of
cystidia were absent in A. sychnopyramis f. subannulata. The most important feature
in A. gleocystidiosa are gleocystidia containing yellow granular cells, abundantly
dispersed among cells in the hymenial layer.
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Fig. 103 Amanita gleocystidiosa (holotype) a (i–ii) Basidiomata b Basidiospores c(i) Basidia
c(ii) Basidia with basidioles d Basidioles e(i) Pleurocystidia e(ii) Pleurocystidia and
basidioles mixed with gleocystidia in the different shapes f(i) Cheilocystidia f(ii)
Cheilocystidia with basidioles g Veil trama. Scale bars: a = 10 mm, b–g = 10 µm.
323. Amanita pyriformis Boonprat. & Parnmen, sp. nov.
Index Fungorum number: IF 551620, Faceoffungi number: FoF 02072, Fig. 104
Etymology: The specific epithet refers to a type of pleurocystidia ‘pyriformis’ =
pear-shaped, narrowly obovoid with a tapering base.
Holotype: BBH 38643.
Pileus 33–55 mm, convex when young, plane with age, rugulose, umbonate, the
whole pileus grayish yellow 1(B)3–7 and yellowish orange 4(A–B)7–8 at margin, dry,
striate 1/8 from margin toward to disc, margin even. Pileus context off white, soft.
Lamellae free, unequal, subdistant, broad, eroded, grayish yellow 1(B)3–7. Stipe
79–112 × 3–7.5 mm, central, cylindrical, enlarged base, grayish yellow 1(B)3–7 with
yellowish orange 4(A–B)7–8 at stipe base near volva, soft. Annulus cream, hanging
about 1/3 of pileus from stipe apex, single, sheathing, smooth, white, thin, apical
attachment 19–27 mm from base toward the apex. Volva constricted, adherent with
flaring margin, white.
Basidiospores (7–) 8–10 × (6–) 7–9 µm [x = 9.12 ± 0.97 × 7.76 ± 0.83 µm, Q =
1.18 ± 0.14, n = 25 spores, 1 collection] broadly ellipsoid, smooth, hyaline, inamyloid,
thin walled. Basidia 29.5 × 11.5 µm, clavate with 4-spores, clamp connection absent,
smooth, hyaline, inamyloid, thin-walled. Basidioles 16–26 × 6.5–11 µm, clavate to
broadly clavate, sometimes pyriform, smooth, hyaline, inamyloid, thin-walled.
Pleurocystidia 28–30 × 7–8 µm, clavate to pyriform, smooth, hyaline, inamyloid,
thin-walled. Cheilocystidia absent. Lamellae trama divergent, composed with broadly
clavate to broadly ellipsoid cells, smooth, hyaline, dextrinoid, thin-walled, base of
hymenial layers directly arising from a few layer of cellular cells which connects to
trama element. Pileipellis composed with cutis of repent hyphae, cylindrical, smooth,
hyaline, inamyloid, thin-walled. Stipilipellis composed with cutis of 3.5–7.5 µm diam.
of repent hyphae, sometime obclavate cells, found among simple cylindrical cells,
smooth, hyaline, inamyloid, thin-walled. Stipe trama composed of two types of
element: repent hyphae and broadly clavate to broadly ellipsoid, smooth, hyaline,
inamyloid, thin-walled. Volva composed of three types of element: 2–6.8 µm diam. of
repent hyphae, broadly clavate to broadly ellipsoid cells 32–52 × 11.5–26 µm and
branching of repent hyphae, smooth, hyaline, inamyloid, thin-walled. Clamp absent in
all parts of basidiomata.
Habitat: Terrestrial in mixed forest.
Material examined: THAILAND, Chiang Mai Province, Omkoi District, Mae
Tun Tumbon, 27 June, 2014, collector SRRT Team, Bureau of Epidemiology,
Department of Disease Control Ministry of Public Health (BBH38643, holotype)
Notes: Amanita pyriformis is similar to A. orientigemmata Zhu L. Yang &
Yoshim. Doi (Yang and Doi 1999) in having broadly ellipsoid basidiospores, but
differs from A. orientigemmata in having smaller, umbonate, pale yellow basidiomata
4904
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and presence of pleurocystidia, while A. orientigemmata has larger basidiomes, up to
100 mm wide, floccose patches on the pileus and the absence of pleurocystidia.
ITS sequence data belonging to core taxa of different sections of Amanita were
selected based on current classification and phylogeny of the genus Amanita (Zhang
et al. 2004). A matrix of 1,005 unambiguously aligned nucleotide characters was
constructed and 276 characters were constant. The topology of the trees from the
maximum likelihood (ML) and maximum parsimony (MP) analyses did not show any
conflict and hence, only the ML tree is shown here (Fig. 100). The boundary of each
section is supported as monophyletic. In this study, we focused on the toxic
mushroom samples from the outbreaks of mushroom poisoning cases in 2012 and
2014. These samples clustered in section Amanita. In our phylogenetic analysis based
on ITS sequence data, Amanita gleocystidiosa, A. digitosa and A. pyriformis were
placed near A. sychnopyramis f. subannulata (Yang et al. 2001), A. subfrostiana
(Yang 1997) and A. orientigemmata (Yang and Doi 1999), respectively. Only
Amanita gleocystidiosa contains a high quality of toxic amanitin.
4919
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Fig. 104 Amanita pyriformis (holotype) a Basidiomata b Basidiospores c Basidia d
Basidioles e Pleurocystidia f Stipilipellis g Veil trama. Scale bars: a = 10 mm, b–f = 10 µm.
324. Amanita strobilipes Thongbai, Raspé & K.D. Hyde, sp. nov.
Index Fungorum number: IF 551651, Facesoffungi number: FoF 02073, Fig. 105
Etymology: Refers to base of stipe like a pine cone.
Holotype: MFLU 12–2246
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Pileus 105 mm in diam., slightly convex then plane, pale gray or grayish white
(1A2, 1C1) with dark gray (1E1, 1F1) conical or pyramidal warts over the center,
progressively becoming brownish gray (5D2, 5D3) squamules towards the margin,
slightly pulverulent-flocculose, margin paler, lacking striations, slightly appendiculate,
edge fibrillose, dry; context 1.5 mm thick at mid-radius, white. Lamellae sub-free to
free, crowded; lamellulae attenuate, with more than 4 series, white to very pale
ochraceous (1A1, 1A2). Stipe 120 × 20 mm, subcylindrical, bulbous, inflated near the
pileus, surface mostly white to smoke gray, pale grayish below, covered with
cottony-fibrillose pulverulence all over, which is easily lost when touched; context
white, solid, unchanging when bruised. Bulb maximum 30 mm wide, spindle-shaped,
covered with white to slightly ochraceous (1A1, 1A2) curved scales. Annulus
membranous, fibrillose, fragile, white. Odour absent.
Lamellar trama bilateral; mediostratum 25–40 µm wide, mainly consisting of
filamentous, 2–5 µm wide, branching hyphae; lateral stratum made up of intercalary
inflated, 25–45 × 5–20 µm, connected with subhymenium. Subhymenium 20–30 µm
thick, with three to four layers of subglobose to broadly ellipsoid cells. Basidia 30–55
× 9–11 µm, 4-spored, clavate, thin to slightly thick-walled, sterigmata 4–6 µm long,
clamps absent at base. Basidiospores (6.8) 7–8.2–9.8 (10.1) × (4.4) 5.2–5.6–6 (8.5)
µm, Q = 1.04–1.46–1.87, (N = 40), ellipsoid to elongate, colourless, amyloid, smooth,
thin-walled, with apiculus. Lamellar edge sterile, mainly consisting of subglobose to
clavate, 12–20 × 4–8 µm, thin-walled cells, mixed with filamentous, 2–3 celled,
brownish hyphae. Pileipellis 250–300 µm thick, composed of filamentous,
subcylindric, occasionally branching, 3–8 µm wide, slightly gelatinized to gelatinized,
hyphae, with pale yellow vacuolar pigments. Velar remnants from pileus consisting of
abundant globose to ellipsoid, 30–60 × 25–65 µm cells, sometimes mixed with
cylindrical, branching, thin-walled, filamentous 1.5–7 μm wide, hyaline or with
brownish to yellowish pigments hyphae with terminal inflated cells. Annulus
composed of clavate, 42–71 × 16–32 μm to cylindrical, 36–50 × 9–15 μm cells, with
brownish to yellowish pigments. No clamps observed in any tissue.
Habitat: Terrestrial in forest with Fagaceae species.
Material examined: THAILAND, Chiang Mai Province, Mae Taeng District,
Mushroom Research Center, N19° 07.20’ E98°44.04’, 25 June 2012, collector B.
Thongbai, BZ–2012–22 (MFLU 12–2246, holotype)
Notes: Amanita strobilipes is a member of Amanita subgenus Lepidella (J.-E.
Gilbert) Veselý emend section Lepidella (Bas 1969) subsection Solitariae. The pale
gray or grayish white pileus with brownish gray squamules on the surface, pyramidal
dark gray warts over the center, whitish stipe covered with white gray to grayish
cottony-fibrillose pulverulence, white to slightly ochraceous, recurved scales on the
spindle-shaped bulb, and amyloid, ellipsoid to elongate basidiospores characterize this
species. Within the subsection Solitariae, the morphologically most similar species is
Amanita griseoverrucosa Zhu L. Yang, originally described from China. Both species
share some similarities, namely the pale gray or grayish white pileus. However, A.
griseoverrucosa produces larger basidiomes, wider basidiospores and the pileus of A.
strobilipes is more distinctively covered with dark gray pyramidal warts to brownish
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gray squamules. Amanita strobilipes also can easily be differentiated from A.
griseoverrucosa by its distinctively spindle-shaped bulb, covered with white to
slightly ochraceous, curved scales, whereas A. griseoverrucosa has a rather ventricose
to subglobose, subradicate bulb, with the upper part covered with grey to greyish
warts or irregularly formed velar remnants. Amanita cinereopannosa Bas, originally
described from the USA, resembles A. strobilipes in the ellipsoid to elongate
basidiospores, a subcylindric stipe and grayish white pileus. However, in A.
cinereopannosa the pileus is covered with rather abundant, soft, pulverulent-subfelty,
low irregular warts, to flat or more angular patches. Additionally, the upper part of
bulb of A. cinereopannosa is usually covered with a few transverse bands or
concentric rows of greyish flocculose-pulverulent patches. Another species that shares
some similarities is A. heishidingensis Fang Li & Qing Cai, originally described from
China, which also shows dark gray pyramidal warts on the pileus, a whitish stipe
covered with white-gray to grayish cottony-fibrillose pulverulence, but its pileus is
rather dirty white to whitish and viscid, the bulb is larger and napiform, subclavate to
ventricose. Moreover, A. heishidingensis appears not to be very closely
phylogenetically related to A. strobilipes.
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Fig. 105 Amanita strobilipes (holotype) a, b Basidiome c Basidia and subhymenium d
Basidiospores e Longitudinal section of velar remnants from pileus. Scale bars: a, b = 20 mm,
c = 20 µm, d = 10 µm.
Cortinariaceae
The limits of the family Cortinariaceae remain unclear at this time. The majority
of the species are in the genus Cortinarius. Many genera formerly placed in the
Cortinariaceae, e.g., Phaeocollybia, Hebeloma, Galerina, and some others have been
moved to other families in Agaricales. On the other hand, the sequestrate genera,
Thaxterogaster, Quadrispora, Protoglossum and Hymenogaster p.p., as well as
Cuphocybe, Rapacea and species of Rozites, once thought to be genera within the
Cortinariaceae, are currently included in the genus Cortinarius (Peintner et al. 2001,
2002). The basidiocarps range from agaricoid to sequestrate, and many have poorly to
well-developed veils. The basidiospores are typically ornamented and cinnamon
brown in deposit.
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Cortinarius (Pers.) Gray
Cortinarius is the largest genus of Agaricales with a cosmopolitan distribution
and over 2000 described species. The species are important ectomycorrhizal fungi and
are associated with different trees and shrubs, belonging to the families Fagaceae,
Salicaceae, Caesalpiniaceae, Cistaceae, Dipterocarpaceae, Myrtaceae, Rhamnaceae,
Rosaceae and Pinaceae, as well as some herbaceous plants in the Cyperaceae and
Polygonaceae. Some species form arbutoid mycorrhizae with Arbutus, Arctostaphylos,
and Comarostaphylis. Revealing the true diversity of species using only
morphological and ecological characteristics has proven to be a difficult if not an
impossible task. The use of sequence data has made it possible to elucidate
phylogenetic relationships within the genus, to show patterns of speciation, and to
help define new, convergent and cryptic species.
In recent years several workers have investigated Cortinarius species associated
with oak and mixed oak-conifer forests and woodlands along the Pacific coast from
California north to Victoria, British Columbia (Bojantchev & Davis 2011, Bojantchev
2013, Bojantchev 2015, Ceska 2013, Garnica et al. 2011, Harrower et al. 2011,
Liimatainen 2015). In most instances, the studies show that the species in these
habitats are new to science and often represent unique and/or significant additions to
our understanding of the phylogenetic relationships in Cortinarius.
Below we introduce nine new species of Cortinarius, subgenus Telamonia that
represent a number of evolutionary lineages. The majority of the specimens were
collected in Quercus garryana Dougl. dominated woodlands of southwestern
Klickitat County, Washington. All collecting was carried out in a 44 km long region,
immediately north of the Columbia River. Elevations ranged from 30 meters to 427
meters. Average annual rainfall is 790 mm/year at the west end and 365 mm/year at
the east end of the oak study area. Further west, oak forests are replaced by Douglas
fir (Pseudotsuga menziesii (Mirb.) Franco) dominated forests, with oaks only found
on the very steep, warmer and dryer south-facing slopes. Mushrooms were rarely seen
on these steep, dry slopes. Further east, the oaks are mainly found in the colder and
wetter north facing slopes where fewer of these mushrooms were found. Nor were
Cortinarius species found under higher elevation oaks where nighttime temperatures
were much colder. In some portions of the oak woodlands, the oaks may be mixed
with Grand fir (Abies grandis (Douglas ex D. Don) Lindley) and Ponderosa pine
(Pinus ponderosa Douglas ex C.Lawson). Specimens collected on British Columbia,
Canada are from Vancouver Island and Salt Spring Island. The average total annual
precipitation is about 880 mm near Victoria. Quercus garryana reaches its global
distribution limit south of the 50° parallel on Vancouver Island near Courtenay and on
Savory Island, northwest of Powell River. Its distribution on Vancouver Island and
adjacent Gulf Islands is determined by the rain shadow of the Olympic and Vancouver
Island Mountains. On the other hand, Arbutus menziesii Pursh that is missing in
Klickitat Co. is a common associate of Quercus garryana in British Columbia. It is an
important co-dominant of Quercus garryana stands on shallow soil and a constant
species at the Quercus garryana/Pseudotsuga menziesii ecotone. The phylogenetic
tree for Cortinarius is presented in Fig. 106.
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Several factors appear to explain why such high portions of the Cortinarius
species in these oak woodlands are new to science. In past decades few Cortinarius
collectors visited these relatively dry habitats. Poison oak (Toxicodendron
diversilobum (Torr. & A. Gray) Greene) may have prevented some people from
entering these woodlands. The fall fruiting often occurs after leaves have fallen from
the trees, thickly covering the ground. The fungi themselves tend to fruit from deep in
the soil and often only just barely stick up above the soil. These two factors make the
mushrooms hard to find. The fruiting window can be very narrow and occurs after
mushrooms have largely ceased fruiting in other near-by areas. The mushrooms often
fruit in small hot spots, often with five to ten species appearing within 30 meters of
each other and no Cortinarius species elsewhere in the oak grove. In some years
fruiting is poor or completely absent in these rather dry habitats.
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Fig. 106 Phylogram resulting from the RaXML (Stamatakis 2014) analysis of ITS regions.
Bootstrap values greater than 50% are indicated above branches. The names in blue represent
the new species of Cortinarius and the specimens in boldface the type specimens of the
species. The tree is rooted with section Cyanites.
325. Cortinarius albosericeus Ammirati, Beug, Liimat., Niskanen & O. Ceska, sp.
nov.
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Index Fungorum number: IF 551701, Facesoffungi number: FoF 02037, Fig. 107
Etymology: Name based on white thinly sheathing veil of stipe and pileus.
Holotype: Michael Beug 01MWB112013 (WTU)
Pileus 30–40 mm diam., convex to broadly umbonate, silky dry, Mahogany Red
to chestnut brown becoming Amber Brown then Tilleul Buff, margin white,
hygrophanous. Lamellae adnate, subdistant, light pinkish cinnamon to cinnamon or
cinnamon brown when mature, edge pale. Stipe 55–80 mm long, 5–7 mm thick, ±
equal, slightly rooting, dry, apex sometimes with bluish tints, light vinaceous
cinnamon to whitish buff. Universal veil white fibrillose, thinly sheathing the surface
of the stipe with indistinct belts. Basal mycelium white. Context pale brown. Odour
fungoid or slightly of radish. Taste slight fungoid to mild. Macrochemical reaction
(40 % KOH): pileus context and surface clove brown, raw umber, bronze, stipe apex
pinkish cinnamon to light ochraceous buff, stipe base warm buff to fuscous black.
Exsiccatae: pileus margin light brown, disc darker brown, lamellae rust brown from
spores, stipe pallid to light brown, white basal mycelium, context pallid to light
brownish. Basidia 4-spored, 7–8.1 × 28–31 µm, clavate, hyaline or slightly brownish.
Basidiospores (7) 7.4–8.5 × 4.6–5.5 µm (20 spores, holotype specimens), ellipsoid,
broadly ellipsoid, or some amygdaloid, slightly to somewhat curved apiculus,
moderately to coarsely verrucose, slightly to moderately or strongly dextrinoid.
Lamella trama hyphae hyaline to yellowish brown or brownish, walls yellow
refractive, encrusted in KOH. Pileipellis in KOH: Surface layer thin, hyphae
cylindrical, 4–6 µm wide, hyaline or rarely yellowish. Subtending layer of ± enlarged
hyphae 8–30 µm wide, hyaline, walls yellow refractive, hyaline to somewhat
yellowish. Beneath a light yellow brown to light brown pigmented layer of cylindrical
to enlarged hyphae, mostly 4–20 µm wide that gradually grade into trama hyphae. ITS
sequence distinct from the other known members of the Decipientes, and differs from
them in the ITS region by more than 15 substitutions and indel positions.
Ecology and distribution: In mixed forests of Quercus garryana and Pinus
ponderosa or Quercus garryana, Pseudotsuga and Arbutus menziesii. Producing
basidiomata in late autumn. Known from British Columbia, Canada and Washington
USA, Western North America.
Material examined: CANADA, British Columbia, Observatory Hill, Saanich,
behind smaller dome, 48.52° N, 123.416° W, margin of mixed forest (Quercus
garryana., Pseudotsuga menziesii, Arbutus menziesii) and open mossy rock outcrops
on SW slope, 26 Nov 2005, leg. Oluna Ceska OC188, F17260 (UBC). USA,
Washington. Klickitat County, Land Trust property, N45°44’20.65” W121°13’11.9”,
Quercus garryana, 20 Nov 2013, leg. Michael Beug 01MWB112013 (holotype,
WTU), (isotype, K(M):200657). Klickitat County, Wahkiacus, N45°49’20.6”
W121°05’38.5”, Quercus garryana and Pinus ponderosa, 20 Nov 2013, leg. Michael
Beug 05MWB112013 (WTU, K).
Notes: Based on the phylogenetic analysis C. albosericeus belongs to clade
Decipientes (Fig. 106). The species in this clade are small and have chestnut brown to
blackish brown pileus, white universal veil, and often some kind of smell in lamellae
(cedar wood-like, spicy, or raphanoid). The stipe apex in almost all species sometimes
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has bluish tints. Cortinarius albosericeus is most similar to C. ohlone Bojantchev, but
C. ohlone has cedar wood-like smell, nondextrinoid spores and occurs with coast live
oak (Quercus agrifolia) and interior live oak (Q. wislizenii) in California.
Fig. 107 Cortinarius albosericeus (05MWB112013, reference specimen) a Basidiomata b
Basidiospores. Photograph a Michael Beug, b Joseph Ammirati. Scale bars: a = 10 mm, b =
10 µm.
326. Cortinarius badioflavidus Ammirati, Beug, Niskanen, Liimat. & Bojantchev, sp.
nov.
Index Fungorum number: IF 551702, Facesoffungi number: FoF 02038, Fig. 108
Etymology: Name based on coloration of pileus and stipe.
Holotype: Joseph Ammirati JFA13668 (WTU)
Pileus 20–60 mm diam., rounded conic, convex to plano–convex, umbonate or
broadly umbonate or uplifted, margin incurved to decurved then plane, non-striate to
striate, expanded, finely pale yellowish to white silky, silky fibrillose or fibrillose
scaly, more or less glabrescent, colour some shade of red brown (brown Russet,
Xanthine Orange, Dresden Brown, Mars Brown, Prout’s Brown, Cinnamon Brown,
Vinaceous Cinnamon), faded more medium brown, edge pale (faded) in older pilei,
disc paler brown at times, hygrophanous. Lamellae distinctly adnexed, subdistant to
distant, sometimes intervenose, moderately broad, moderately thick then thicker in
age, light medium brown, becoming rich brown (brownish Chamois, Cinnamon,
Buckthorn Brown, Tawny Olive, Sudan Brown, Brussels Brown, Amber Brown,
Argus Brown, Carob Brown), edges even to uneven in age, remaining pale for some
time, then concolor. Stipe 43–88 mm long, apex 5.5–15 mm thick, equal or strongly
tapered to base, tough, rigid, yellowish Cream Colour, Light Ochraceous Buff, Light
Buff, Colonial Buff to Chamois, buff and yellow becoming mixed with brown, lower
stipe developing watery red brown areas, often dull watery red brown to watery dull
yellow brown., stipe surface longitudinally fibrillose, fibrils white to faintly yellowish
or orange buff. Universal veil white, forming a ring and incomplete girdles or almost
a sock-like sheath on the stipe. Basal mycelium white. Context rather thin in pileus,
watery concolour with surface, above stipe apex yellowish white or sometimes
pinkish cinnamon, in stipe central area stuffed whitish to yellowish white, cortex
watery yellow brown to brown or dark brown or red brown (Sudan Brown, Brussels
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Brown, Argus Brown, Antique Brown) in base the cortex somewhat darker brown.
Odour sharply fragrant to that of green corn. Taste slightly unpleasant or astringent.
Macrochemical reaction (40 % KOH): pileus cuticle Xanthine Orange, context pale
yellow orange, stipe apex Xanthine Orange, stipe base Seal Brown. Exsiccatae: pileus
light brown to dark brown or somewhat blackish, lamellae rich medium brown, stipe
pallid to brownish or somewhat yellowish, with some blackish area, context dull
whitish to pallid or slightly brownish. Basidia 4-spored, 8.7–9.2 × 29–31 µm, clavate,
hyaline or commonly rich orange brown to yellow brown. Basidiospores 8.1–10.5 ×
5.8–6.5 µm (20 spores, holotype specimens), broadly ellipsoid to broadly amygdaloid,
very coarsely verrucose, moderately to strongly dextrinoid. Lamella trama hyphae
heavily pigmented, red brown, orange brown, yellow brown, strongly encrusted in
KOH. Pileipellis in KOH: Surface hyphae ± cylindrical to broadly cylindrical, 6–11
µm wide, hyaline or yellowish; subtending layer of ± enlarged hyphae 7–24 µm wide,
hyaline to yellowish brown; beneath a yellow brown to orange brown pigmented layer
of cylindrical to enlarged hyphae, 6–25 µm wide adjacent to trama hyphae. ITS
sequence distinct from the other known members of the section Hinnulei, and differs
from them in the ITS region by more than 6 substitutions and indel positions.
Ecology and distribution: Collections have been made in mixed forests of
Quercus garryana, Q. douglasii, Pseudotsuga menziesii, Abies grandis, and Pinus
ponderosa, Salix scouleriana has also been present in some areas. Producing
basidiomata in late autumn-winter and spring. Known from Western North America,
from California to Washington.
Material examined: USA, California, Contra Costa County, Tilden Park,
N37°54'14.5" W122°15'32.1", Quercus agrifolia and Pseudotsuga menziesii, 23 Nov
2009, leg. Dimitar Bojantchev DBB28196. Marin County, Marin Watershed,
N37°56'44.1", W122°35'32.6", Quercus agrifolia and Pseudotsuga menziesii, 09 Jan
2009, leg. Dimitar Bojantchev DBB13504. Yuba County, Southern Sierra Research
Station src94, Quercus douglasii woodland, 14 Feb 2001 leg. Matthew Smith (UCB),
Washington, Klickitat County, 45°48’36.71“N, 121°30”55.72 “W, Quercus garryana,
19 Feb 2010, leg. Michael Beug 01MWB021910 (WTU, K). Beug Farm, near air field,
Quercus garryana, Pseudotsuga menziesii, Salix sp., 20 Nov 2010, leg. Joseph
Ammirati JFA13668 (holotype, WTU) (isotype, K(M): 200672), JFA13669. Beug
Property, 45° 48.607 N, 121° 30.986 W, Quercus garryana, 3 April 2009 leg. Michael
Beug 02MWB040309 (WTU, K). Oak grove (Quercus garryana, Pseudotsuga
menziesii and Abies grandis) behind Beug house, 45° 48.606 N, 121° 30.973 W, 8
March 2009, leg. Michael Beug 01MWB030809 (WTU, K). Behind house, 194
Spring Creek, Husum, 24 March 2011, Quercus garryana, leg. Michael Beug
01MWB032411 (WTU, K). One thousand feet west of Beug property, 45°48.430 N,
121°31.135 W, Quercus garryana, 3 Dec 2008, leg. Michael Beug 03MWB120308
(WTU, K). Lindserth Old Road, 45° 48.419 N, 121°31.122 W, Quercus garryana,
Pseudotsuga menziesii and Abies grandis, 9 Nov 2010, leg. Michael Beug
01MWB110910 (WTU, K). 45° 48.611 N, 121° 30.936W, Quercus garryana,
Pseudotsuga menziesii and Abies grandis, 30 Apr 2009 leg. Michael Beug
02MWB043009 (WTU, K). SDS west of Beug Farm, N45°48’24”, W121°31’06”,
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Quercus garryana, Pseudotsuga menziesii, 19 Nov 2013, leg. Michael Beug
11MWB111913, (WTU, K).
Notes: Cortinarius badioflavidus looks like a typical member of section Hinnulei
(Fig. 106). The overall coloration of the basidiocarp is red brown to brown, the stipe
is equal or tapered, lamellae are distant and the smell of lamellae is green corn-like.
The broadly ellipsoid to broadly amygdaloid spores differentiate it from C. hinnuleus
collections which have subglobose to obovoid-subglobose spores. European
Cortinarius hinnuleoarmillatus is otherwise very similar to C. badioflavus, but it has
orange red universal veil.
Fig. 108 Cortinarius badioflavidus (holotype) a Basidiomata b Basidiospores. Photographs
Joseph Ammirati. Scale bars: a = 10 mm, b = 10 µm.
327. Cortinarius denigratus Ammirati, Beug, Niskanen, Liimat. & O. Ceska, sp. nov.
Index Fungorum number: IF 551703, Facesoffungi number: FoF 02039, Fig. 109
Etymology: Name based on blackening of the basidiocarps on drying.
Holotype: Michael Beug 02MWB043014 (WTU)
Pileus 10–20 mm diam., papillate umbo, dry, Dresden Brown to Mars Yellow,
edge blackens dried (in sun), minutely fibrillose, hygrophanous. Lamellae adnexed, ±
distant, tan rusty. Stipe 30–40 mm long, 2–4 mm thick, ± equal, minutely fibrillose,
buckthorn brown. Universal veil not recorded. Basal mycelium white. Context in stipe
context ochraceous buff to yellowish tan. Odour indistinct. Macrochemical reaction
(40 % KOH): all parts instantly black. Exsiccatae: pileus brown to blackish, lamellae
dark dull brown to blackish, stipe brown to blackish, some white mycelium at base,
context brown. Basidia 4-spored, 9–10 × 27–37 µm, clavate, hyaline, light brown or
dark brown in KOH. Basidiospores 9–11.2 × 4.8–6 µm (20 spores, holotype
specimens), narrowly to broadly amygdaloid, distinct apiculus, moderately to very
coarsely verrucose, apex ± extended and less ornamented, slightly to somewhat
moderately dextrinoid. Lamella trama hyphae hyaline, light brown or very dark
brown, walls yellow refractive, heavily brown encrusted and with brown interhyphal
plaques in KOH. Pileipellis in KOH: Surface layer thin, somewhat compressed,
hyphae ± cylindrical, 4–12 µm wide, hyaline or yellowish, some encrusted.
Subtending layer of cylindrical to enlarged hyphae 7–25 µm wide, yellow brown to
orange brown, walls yellow refractive, some heavily encrusted with brown pigment.
Beneath a darker brown pigmented layer of encrusted hyphae with interhyphal brown
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plaques, cylindrical to enlarged, mostly 7–22 µm wide, that gradually grade into trama
hyphae. ITS sequence distinct from other species of Cortinarius subgenus Telamonia.
Ecology and distribution: Found from forests of Quercus garryana and Pinus
ponderosa, and Pseudotsuga menziesii and Arbutus menziesii. Producing basidiomata
in spring in April. Known from British Columbia, Canada and Washington USA,
Western North America.
Material examined: CANADA, British Columbia, Salt Spring Island, Mt. Tuam,
48.72° N 123.485° W, along the trail through mixed forest (Pseudotsuga, Arbutus), 19
April 2007, leg. Oluna Ceska OC155, F17227 (UBC). USA, Washington, Klickitat
County, Beug Farm, N45°48’36.6” W121°30’59.04”, Quercus garryana and Pinus
ponderosa, 30 April 2014, leg. Michael Beug 02MWB043014 (holotype, WTU),
(isotype, K(M): 200659).
Notes: Cortinarius denigratus is easily recognized since it produces fruitbodies
in the spring when not that many other Cortinarius species are fruiting. Characteristic
for the species are small, brown basidiomata, highly brown pigmented lamella trama
hyphae, and amygdaloid, rather large spores with ± extended apex. Cortinarius
denigratus is not very closely related to any of the known Telamonia species, but
groups together with other small Telamonias in our phylogenetic analysis (Fig. 106).
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Fig. 109 Cortinarius denigratus (holotype) a Basidiomata b Basidiospores. Photograph a
Michael Beug, b Joseph Ammirati. Scale bars: a = 10 mm, b = 10 µm.
328. Cortinarius duboisensis Ammirati, Beug, Niskanen & Liimat, sp. nov.
Index Fungorum number: IF 551704, Facesoffungi number: FoF 02040, Fig. 110
Etymology: Named for DuBois Lake, the original name of Roland Lake in
Washington, USA
Holotype: Joseph Ammirati JFA13311(WTU)
Pileus 50–135 mm diam., broadly obtuse-umbonate to plano-umbonate then ±
plane to uplifted, margin decurved at first, mature becoming irregular and lacerated,
easily broken, surface moist to dry, not striate, center often with whitish bloom,
margin in places silky or with thin coating of whitish fibrils, colour variable, when
moist watery dark brown to watery grey brown, faded areas ochraceous tawny, light
brown, brownish buff or light buff, center sometimes very pale, margin frequently
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finely rivulose-variegated or streaked with brown colors, sometimes with darker areas
or blotches, hygrophanous. Lamellae adnexed, close to subdistant, very broad,
moderately thick to thick, somewhat easily broken, pale brown at first or in non-spore
covered areas, rich deep brown when mature, edges irregular. Stipe 45–135 mm long,
10–32 mm thick above, base 23–35 mm thick, varies from narrow clavate to bulbous,
upper surface shiny, sometimes twisted striate, whitish to pallid-white, with some thin
darker watery buff brown streaks, without veil remains above. Universal veil white.
Basal mycelium white and extends up onto base of the stipe. Context whitish to pallid
or brownish white, darkening with age and with exposure, watery brown streaked in
stipe, cortex rather tough, lower stipe flesh soon grayish then much darker brown,
especially in stipe base. Odour strong fungoid to mildy woodsy. Taste mild, fungoid.
Macrochemical reaction (40 % KOH): on pileus surface raw umber, pileus context
bronze, stipe apex, pinkish buff exterior, interior of stipe including stipe base, fuscous
black. Exsiccatae: pileus grey brown to rather dark grey brown, lamellae dark brown,
stipe whitish to pallid or greyish with a few blackish areas, context similar to stipe
surface, basal mycelium white. Basidia 4-spored, 8.5–9 × 29–48 µm, clavate, hyaline
or pale brownish in KOH. Basidiospores 8.9–10.2 × 5–6.2 µm (20 spores, holotype
specimens), ellipsoid, broadly ellipsoid or somewhat amygdaloid, moderately
verrucose, somewhat to strongly dextrinoid. Lamella trama hyphae smooth, not
encrusted in KOH. Pileipellis in KOH: Surface hyphae ± cylindrical, 4–14 µm wide,
hyaline or brownish, some encrusted. Subtending layer of cylindrical to enlarged
hyphae 4–20 µm wide, hyaline, not encrusted. Beneath a brown pigmented layer of
cylindrical to enlarged hyphae 8–22 µm wide, grading into trama hyphae. ITS
sequence distinct from the other known members of the subgenus Telamonia, and
differs from them in the ITS region by more than 15 substitutions and indel positions.
Ecology and distribution: Collections have been made under Quercus garryana,
Pinus ponderosa or a mixture of Quercus garryana, Pinus ponderosa and Abies
grandis. Producing basidiomata in late autumn. Known from Washington, Western
North America.
Material examined: USA, Washington, Klickitat County, Roland Lake, 47.36N
122.73W, ecology, Quercus garryana and Pinus ponderosa, 28 Nov 2008, leg. Joseph
F. Ammirati JFA13308 (WTU, K), JFA13311 (holotype, WTU), (isotype, K),
JFA13312 (WTU). Klickitat County, Lower Staats Road, N45°50’38.5”, W121°24’
44.1”, Quercus garryana, Pinus ponderosa and Abies grandis, 18 Nov 2013, leg.
Michael Beug 01MWB111813 (WTU, K).
Notes: Cortinarius duboisensis is a rather large species with dark brown to
watery grey brown pileus with a whitish bloom in the center and clavate to bulbous
stipe. Typical are also exsiccatae with grey brown to rather dark grey brown pileus
and whitish to greyish stipe. Cortinarius duboisensis is related to C. crassisporus
Kytöv., Niskanen & Liimat. which also has basidiomata with bulbous stipe and brown
pileus (Fig. 106). Cortinarius crassisporus, however, has larger spores (10.7–13.6 x
7.5–9.1 µm) and occurs in hemiboreal–boreal and mountain coniferous forests on
calcareous soil. The species are morphologically most reminiscent to those of section
Bovini, but do not seem to belong to that section based on our phylogenetic analysis.
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Fig. 110 Cortinarius duboisensis (holotype) a Basidiomata b Basidiospores. Photographs
Joseph Ammirati. Scale bars: a = 10 mm, b = 10 µm.
329. Cortinarius fragrantissimus Ammirati, Beug, Liimat., Niskanen & O. Ceska, sp.
nov.
Index Fungorum number: IF 551705, Facesoffungi number: FoF 02041, Fig. 111
Etymology: Name based on fragrant Odour.
Holotype: Michael Beug 10MWB111913 (WTU)
Pileus 15–30 mm, rounded-umbonate to obtuse umbonate to plano-umbonate,
umbo ± acute, surface silky, dry, fuscous to pale ochraceous salmon, hygrophanous.
Lamellae adnexed, subdistant, dark vinaceous purple when young, buffy brown when
mature. Stipe 55–70 mm long, 3–5 mm thick above, ±equal, dry, pale pinkish buff,
hollow. Universal veil white. Basal mycelium white. Odour slightly fragrant sweet.
Taste mild. Macrochemical reaction (40 % KOH): pileus cuticle raw umber, stipe
apex bronze, base fuscous black. Exsiccatae: pileus pallid to brown, greyish brown
and some blackish areas, lamellae brown, stipe pale shiny at apex, below pallid to
light brownish or blackish with whitish areas from universal veil, basal mycelium
white. Basidia 4-spored, 8.7–9.2 × 29–31 µm, clavate, hyaline or commonly brown in
KOH. Basidiospores (7.4) 7.8–9 (9.3) × 4.8–6 µm (20 spores, holotype specimens),
ellipsoid to broadly ellipsoid, coarsely verrucose, apiculus ±curved, slightly to
somewhat dextrinoid. Lamella trama hyphae hyaline or more commonly brown
pigmented, commonly brown encrusted, many brown plaques in KOH. Pileipellis in
KOH: Surface hyphae ± cylindrical, 4–11 µm wide, hyaline, walls refractive.
Subtending distinct layer of ± enlarged hyphae 7–26 µm wide, hyaline to yellowish
brown, walls refractive. Beneath a brown to yellow brown pigmented layer of
cylindrical to enlarged hyphae, 7.5–22 µm wide, encrusted and with pigment plaques,
grading into trama hyphae. ITS sequence distinct from the other known members of
the section Paleacei, and differs from them in the ITS region by more than 8
substitutions and indel positions.
Ecology and distribution: Collections have been made from mixed forests of
Quercus garryana and Abies grandis, and Pseudotsuga menziesii and Arbutus
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menziesii. Producing basidiomata in late autumn. Known from British Columbia,
Canada and Washington USA, Western North America.
Material examined: CANADA, British Columbia, Cobble Hill, off Thain Rd.,
48.686° N, 123.6° W, mixed forest (Pseudotsuga menziesii, Arbutus menziesii), 25
Nov 2000, leg. Oluna Ceska OC66, F17138 (UBC). Skulow Lake, forest soil from the
long-term soil productivity (LTSP) site, Aug 2007, environmental sample. USA,
Washington. Klickitat County, SDS west of Beug Farm, N45°48’24”, W121°31’06.5”,
Quercus garryana and Pseudotsuga menziesii, 19 Nov 2013, leg. Michael Beug
10MWB111913 (holotype, WTU), (isotype, K(M): 200664).
Notes: Cortinarius fragrantissimus belongs to section Paleacei (Fig. 106).
Typical for the species of the section is the fragrant smell in lamellae, often
reminiscent of that of Pelargonium, as well as small basidiomata and white universal
veil. Several species also have purplish tints in lamellae and/or stipe apex. Cortinarius
fragrantissimus can be separated from the other species of the section by the
combination of smooth cap with more or less acute umbo and ellipsoid 8–9 ×5–6 µm,
coarsely verrucose spores.
Fig. 111 Cortinarius fragrantissimus (holotype) a Basidiomata b Basidiospores. Photograph
a Michael Beug, b Joseph Ammirati. Scale bars: a = 10 mm, b = 10 µm.
330. Cortinarius roseobasilis Ammirati, Beug, Niskanen & Liimat., sp. nov.
Index Fungorum number: IF 1551706, Facesoffungi number: FoF 02042, Fig.
112
Etymology: Name based on reddish stipe base.
Holotype: Michael Beug 20MWB111813 (WTU)
Pileus 42–75 mm diam., obtuse-umbonate to plano-umbonate then
uplifted-irregular umbonate, margin decurved to straight, becoming lacerate-split in
age, non–striate or only short striate at edge in a few places, very little veil materials
on edge, silky dry, colour Blackish Brown (1) to Dusky Brown or Dresden Brown
streaked with light ochraceous buff where faded, edge grayish to greyish brown,
hygrophanous. Lamellae adnexed with a decurrent line, distant, thick, becoming
irregular, deep brown with Vinaceous Drab mixed in, becoming Dresden Brown.
Stipe up to 80 mm (often 70–80 mm) long, above up to 8 (or sometimes to 20) mm
thick, equal above, strongly tapered to base, with some dull whitish fibrillose areas,
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otherwise buffy brown to lighter brown then watery brown to watery vinaceous brown.
Universal veil white, sparse. Basal mycelium white. Context of pileus thin, watery and
concolor, faded whitish, in stipe drab with pale drab gray streaks, hollow. Odour not
distinctive. Taste mild or not distinctive. Macrochemical reaction (40 % KOH): pileus
cuticle fuscous, pileus context buffy brown, stalk apex pale ochraceous salmon, stipe
base fuscous black. Exsiccatae: pileus dark brown to blackish, lamellae brown to dark
brown, stipe pallid, brownish or blackish, lower stipe whitish in one, context pallid to
darkened in lower stipe. Basidia 4-spored, 6.5–8 × 28–31 µm, clavate, hyaline or
slightly brownish. Basidiospores 6.7–8.9 × 4.5–4.8 (5.5) µm (20 spores, holotype
specimens), ellipsoid, broadly ellipsoid, rarely subglobose, slightly curved apiculus,
coarsely verrucose, moderately to very strongly dextrinoid. Lamella trama hyphae
hyaline to yellowish brown or brownish, somewhat encrusted in KOH. Pileipellis in
KOH: Surface layer thin, hyphae cylindrical, 5.2–9 µm wide, hyaline or yellowish,
some slightly encrusted. Subtending layer of ± enlarged hyphae 8.9–26 µm wide,
hyaline, walls refractive, somewhat yellowish beneath a light yellow brown to light
brown pigmented layer of cylindrical to enlarged hyphae, 4.5–25 µm wide adjacent to
trama hyphae. ITS sequence distinct from the other known members of the /Castanei,
and differs from them in the ITS region by more than 7 substitutions and indel
positions.
Ecology and distribution: Gregarious under Quercus garryana or in mixed
forests of Q. garryana, Crataegus, and Populus tremuloides. Producing basidiomata
in late autumn. Known from Washington USA, Western North America.
Material examined: USA, Washington. Klickitat County, Balch Farm,
45°42.896N, 121°18.939W, Quercus garryana with Populus and Crataegus, 20 Nov
2010, leg. Joseph F. Ammirati JFA13666 (WTU, K). Klickitat County Lower Staats
Road, N45°50’39”W121°24’50”, Quercus garryana, 18 Nov 2013, leg. Michael Beug
20MWB111813 (holotype, WTU), (isotype, K).
Notes: In our phylogenetic analysis C. roseobasilis is placed in Castanei
although the group is not well-supported (Fig. 106). However, the species in the group
are morphologically similar. They have dark brown to blackish brown pileus;
reddening, but first white, universal veil and/or stipe base; and indistinctive smell in
lamellae. Cortinarius rosebasilis is most reminiscent of European C. erubescens M.M.
Moser, but the spores of C. erubescens are narrowly ellipsoid and almost smooth.
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Fig. 112 Cortinarius roseobasilis (holotype) a Basidiomata b Basidiospores. Photograph a
Michael Beug, b Joseph Ammirati. Scale bars: a = 10 mm, b = 10 µm.
331. Cortinarius vinaceobrunneus Ammirati, Beug, Liimat., Niskanen & O. Ceska,
sp. nov.
Index Fungorum number: IF 551707, Facesoffungi number: FoF 02043, Fig. 113
Etymology: Named for the colour of the pileus and stipe.
Holotype: Joseph Ammirati JFA13301 (WTU)
Pileus 47–60 mm diam., obtusely rounded to obtuse-uplifted, with slight umbo at
times, margin incurved to straight but often folded and irregular, opaque, edge whitish
fibrillose from veil, colour evenly deep vinaceous brown with a pale sheen from thin
layer of silky fibrils, hygrophanous. Lamellae deeply adnexed, intervenose, close to
subdistant, thick, deep rich brown with paler brown edges, light medium brown
viewed from edges, edges uneven. Stipe 82–100 mm long, apex 10–14 mm thick,
tapered below, deeply inserted in soil, shiny, silky streaky, dull watery light vinaceous
brown ground color. Universal veil white, sparse, forming a few surface fibrils on
stipe, no zones. Basal mycelium white, sparse. Context watery brown in cortex (rather
thick and tough), interior of stipe pale brownish white, dark watery brown in pileus
cuticle, flesh thin, brownish white above stipe apex. Odour pleasant, like parsley.
Taste mild. Macrochemical reaction (40 % KOH): not recorded. Exsiccatae: pileus
blackish with slight purplish cast, lamellae rich brown or a few blackish, stipe
blackish or with some pallid greyish or brownish areas, context is a light bright
cinnamon brown. Basidia 4-spored, 8.1–8.5× 28–35 µm, clavate, hyaline to light
brown in KOH. Basidiospores 8.1–9.6 × 4.8–5.9 µm (20 spores, holotype specimens),
ellipsoid to broadly ellipsoid or somewhat amygdaloid, distinct, ± curved apiculus,
moderately to coarsely verrucose, slightly to moderately (a few darker) dextrinoid.
Lamella trama hyphae hyaline to brown, walls yellow refractive, some encrusted (not
heavily so) in KOH. Pileipellis in KOH: Surface layer of ± cylindrical hyphae, 3–11
µm wide, hyaline or yellowish, wall refractive, some encrusted; subtending layer of ±
cylindrical to enlarged hyphae 7–26 µm wide, colourless to yellowish or slightly
brownish, walls yellow refractive, some encrusted. Beneath a somewhat darker brown
layer of cylindrical to enlarged hyphae, mostly 8–22 µm wide, hyaline or with brown
pigments, grading into trama hyphae; hyaline to dark brown lactiferous hyphae
scattered throughout trama. ITS sequence distinct from other species of Cortinarius
subgenus Telamonia and deviating from them by more than 15 substitutions and indel
positions in the ITS region.
Ecology and distribution: With Quercus garryana. Producing basidiomata in late
autumn. Known from British Columbia, Canada and Washington USA, Western
North America.
Material examined: CANADA, British Columbia, Elkington property Reserve,
Duncan, 48.805° N, 123.622° W, Quercus garryana stand, 25 Nov 2001, leg. Oluna
Ceska OC78, F17150 (UBC). USA, Washington, Klickitat County, Balch Farm,
45°42.896N,121°18.939W, Quercus garryana, 2 Nov 2008, leg. Joseph Ammirati
JFA13301 (holotype, WTU), (isotype, K(M): 200667).
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Notes: Cortinarius vinaceobrunneus is a small to medium-sized species of
subgenus Telamonia. From many other similar looking species it can be distinguished
by the combination of vinaceous brown pileus, silky white rooting stipe almost
without veil remnants, and the parsley-like smell in lamellae. The exact phylogenetic
position of the species is not known, but in our analysis it is grouped in the same large
clade with section Hinnulei and many small Telamonias (Fig. 106).
Fig. 113 Cortinarius vinaceobrunneus (holotype) a Basidiomata b Basidiospores.
Photographs Joseph Ammirati. Scale bars: a = 10 mm, b = 10 µm.
332. Cortinarius vinaceogrisescens Ammirati, Beug, Liimat. & Niskanen, sp. nov.
Index Fungorum number: IF 551708, Facesoffungi number: FoF 02044, Fig. 114
Etymology: Name based on coloration of the stipe.
Holotype: Michael Beug 03MWB111913 (WTU)
Pileus 30–65 mm diam., convex, at times subumbonate, becoming uplifted silky,
red brown to Light Pinkish Cinnamon, hygrophanous. Lamellae adnexed, subdistant,
reddish brown to dark brown (Natal Brown) when mature. Stipe 60–100 mm long,
5–10 mm thick, equal, dry, at first white, later pale greyish vinaceous brown (Tilleul
Buff) at apex, lower down grey vinaceous brown (Wood Brown). Universal veil white.
Basal mycelium white. Odour very slightly fragrant, pleasant. Taste mild.
Macrochemical reaction (40 % KOH): pileus cuticle and stipe base Chaetura Black,
context and stipe apex Chamois. Exsiccatae: pileus light brown to brown with
blackish areas, lamellae brown, stipe brownish to blackish with whitish veil covering
above base, basal mycelium white, context pallid to brownish. Basidia 4-spored,
8.5–9 × 35–42 µm, clavate, hyaline or brownish. Basidiospores 8.5–10 × 5.4–6.4 µm
(20 spores, holotype specimens), broadly ellipsoid, moderately to coarsely verrucose,
somewhat to moderately dextrinoid. Lamella trama hyphae hyaline to brown, some
encrusted. Pileipellis in KOH: surface hyphae cylindrical, 5–9.5 µm wide, hyaline or
yellowish to brownish, some encrusted; subtending layer of cylindrical to enlarged
hyphae 5–18 µm wide, hyaline, walls refractive, interhyphal and encrusted pigment
common, grading into trama hyphae. ITS sequence distinct from other species of
Cortinarius subgenus Telamonia. With an isolated position and deviating from the
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other members of the subgenus in the ITS region by more than 20 substitutions and
indel positions.
Ecology and distribution: Collections have been made in mixed forests of
Quercus garryana and Pseudotsuga menziesii. Producing basidiomata in late autumn.
Known from Washington and Oregon, Western North America. The Oregon record is
based on a sequence (GenBank no. JQ393038) from a mycorrhizal root tip of Arbutus
menziesii (Ericaceae). It differs by some bases from the type material but the
differences might be artificial.
Material examined: USA, Washington, Klickitat County, Beug Farm, N 45
48.624, W 121 30.969, mixed forest of Quercus garryana and Pseudotsuga menziesii,
20 Nov 2010, leg. Joseph F. Ammirati JFA13674 (WTU, K). Klickitat County, SDS
west of Beug Farm, N45°48’24”, W121°31’06”, mixed forest of Quercus garryana
and Pseudotsuga menziesii, 19 Nov 2013, leg. Michael Beug 03MWB111913
(holotype, WTU), (isotype, K(M): 200668).
Notes: Cortinarius vinaceogriseus can be recognized by a combination of brown
pileus, first white, later vinaceous brown stipe, rather large, broadly ellipsoid spores
and rather dark exsiccatae. It is not very closely related to any previously known
species/sections of Telamonia (Fig. 106).
Fig. 114 Cortinarius vinaceogrisescens (holotype) a Basidiomata and b Basidiospores.
Photograph a Michael Beug, b Joseph Ammirati. Scale bars: a = 10 mm, b = 10 µm.
333. Cortinarius wahkiacus Ammirati, Beug, Liimat. & Niskanen, sp. nov.
Index Fungorum number: IF 551709, Facesoffungi number: FoF 02045, Fig. 114
Etymology: Named for Wahkiacus Washington, USA
Holotype: Michael Beug 09MWB111813 (WTU).
Pileus 45–60 mm diam., convex to ± plane, silky dry, streaked with yellow
brown (Raw Umber) and cinnamon buff or umber brown on light vinaceous cinnamon.
Lamellae adnexed, distant to subdistant, cinnamon or light brown to yellowish brown
(Buckthorn Brown) when mature. Stipe 70–80 mm long, 8–12 mm thick above, ±
equal down to an ± enlarged base inserted in soil, surface dry, honey yellow to
cinnamon buff or light vinaceous cinnamon. Universal veil white, sheathing lower
stipe, forming inferior ring. Basal mycelium white, with white rhizomorphs. Taste
mild. Odour slightly musty to fishy. Macrochemical reaction (40 % KOH): on pileus
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surface fuscous to olivaceous black, stipe apex olive to fuscous, interior citrine drab to
olive, stipe base fuscous black to dark olive, rhizomorphs white. Exsiccatae: Pileus
dark brown to dark reddish brown, one with large, central white veil patch. Lamellae
dark brown. Stipe surface light brown to pale in some places above but often blackish.
Basal mycelium, sheathing veil above base and rhizomorphs white. Context pale to
brownish discolored blackish. Basidia 4-spored, 8–10 × 31–38 µm, clavate, hyaline or
brownish in KOH. Basidiospores 10–11.6 × 5.4–6.6 µm (20 spores, holotype
specimens), amygdaloid to ± ellipsoid, moderately to coarsely verrucose, apiculus
somewhat curved, somewhat to strongly dextrinoid. Lamella trama hyphae smooth,
not encrusted in KOH. Pileipellis in KOH: Surface hyphae cylindrical to broadly
cylindrical, 8–10 µm wide, hyaline or brownish, some encrusted. Subtending layer,
hyphae 8–21 µm wide, hyaline, walls refractive, some encrusted, gradually grading
into trama hyphae. ITS sequence distinct from the other known members of the
section Bovini, and differs from them in the ITS region by more than 20 substitutions
and indel positions.
Ecology and distribution: Found from forests of Quercus garryana or Q.
garryana and Pinus ponderosa. Producing basidiomata in late autumn. Known from
Washington, Western North America.
Material examined: USA, Washington, Klickitat County, Lower Staats Road, N
45°50’36.4”, W121°24’ 33.7”, under Quercus garryana, 18 Nov 2013, leg. Michael
Beug 09MWB111813 (holotype, WTU), (isotype, K(M): 200670). Klickitat County,
Wahkiacus, under Quercus garryana and Pinus ponderosa, N 45°49’20.6”,
W121°05’ 38.9”, 20 Nov 2013, leg. Michael Beug 03MWB112013 (WTU, K).
Notes: Cortinarius wahkiacus is a medium-sized, brown species with a white,
sheath-like universal veil covering the lower part of the stipe, and with rather large,
amygdaloid, moderately to coarsely verrucose, dextrinoid spores. It belongs to section
Bovini (Fig. 106) and as other members of the group has exsiccatae with dark brown
to blackish brown pileus. Cortinarius eldoradoensis Bojantchev is another species of
section Bovini encountered in Western North America, but it fruits in the spring and
has somewhat shorter spores, 8.5–10.5 × 5–6 µm.
Fig. 115 Cortinarius wahkiacus (holotype) a Basidiomata b Basidiospores. Photograph a
Michael Beug, b Joseph Ammirati. Scale bars: a = 10 mm, b = 10 µm.
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Tricholomataceae R. Heim ex Pouzar
The family Tricholomataceae, as traditionally circumscribed (Singer 1986),
includes 98 genera with a pale spore print (white, cream, light pink, pale violet, light
green, or pale greyish), lamellae variously attached to the stipe (rarely free, adnate,
sinuate, or decurrent); hymenophoral trama regular to subregular, irregular,
interwoven, bilateral; spores amyloid or inamyloid; clamp-connections present or
absent; mainly saprotrophic or symbiotic. The family was demonstrated to be
polyphyletic in several molecular analyses (Hofstetter et al. 2002; Moncalvo et al.
2000, 2002; Matheny et al. 2006; Garnica et al. 2007). Some taxa previously included
in Tricholomataceae have been placed in other families such as Lyophyllaceae Jülich
(Hofstetter et al. 2002), “Marasmiaceae” (Wilson and Desjardin 2005), Mycenaceae
Overeem (Moncalvo et al. 2002), Omphalotaceae Bresinsky (Moncalvo et al. 2002),
“Physalacriaceae” (Binder et al. 2006), and Hygrophoraceae Lotsy (Lodge et al.
2014).
Based on a multi-gene analysis, Sánchez-García et al. (2014) recognized a
Tricholomataceae sensu stricto which encompasses only seven genera, Albomagister
Sánchez-García, Birkebak & Matheny, Corneriella Sánchez-García, Dennisiomyces
Singer, Leucopaxillus Boursier, Porpoloma Singers.str., Pseudotricholoma (Singer)
Sánchez-García & Matheny, Tricholoma (Fr.) Staude. Vizzini et al. (2016) added to
the family the genus Pseudoporpoloma Vizzini & Consiglio.
Pseudoclitocybe-Musumecia clade
Binder et al. (2010) and Vizzini et al. (2011) showed a well-supported
phylogenetic relationship between Infundibulicybe Harmaja and Pseudoclitocybe
(Singer) Singer at the base of the Tricholomatoid clade. Vizzini et al. (2011) and
Sánchez-García et al. (2014) found also a significant relationship between Musumecia,
Pseudoclitocybe, and the genus Pogonoloma (Singer) Sánchez-García (= Porpoloma
subgen. Pogonoloma Singer), while Aspropaxillus Kühner & Maire and Notholepista
Vizzini & Contu were found also to represent basal lineages to the Tricholomatoid
group. This clade is characterized by the absence or scarce number of cystidia and
clamp connections in most species, as well as the cutis-like pileipellis, elongated
basidia
and
acyanophilous
spores.
The
phylogenetic
tree
for
Pseudoclitocybe-Musumecia clade is presented in Figs 116 and 117.
Musumecia Vizzini & Contu
The genus Musumecia is a small genus in the so called Tricholomatoid clade
(Matheny et al. 2006, Sánchez-García et al. 2014). Its name was originally proposed
by Vizzini et al. (2011) to honor the Swiss mycologist Enzo Musumeci, who was the
first to collect this genus from Alsace (France). Molecular data revealed that this
genus is closely related to Pseudoclitocybe. The genus Musumecia was established to
encompass clitocyboid fungi phylogenetically close to Pseudoclitocybe with a
hygrophoroid habit (non-depressed convex pileus and distant thick lamellae), a
cutis-like pileipellis, regular hymenophoral trama, elongated basidia, smooth
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acyanophilous and inamyloid spores, absence of hymenial cystidia and
clamp-connections (Vizzini et al. 2011). The type species, M. bettlachensis Vizzini &
Contu (Vizzini et al. 2011), is whitish and grows caespitose in Abies alba, Fraxinus
sp., and Fagus sp. forests, while the only other known taxon, M. vermicularis
Musumeci (Musumeci 2014), has a zonate brownish dark pileus, is gregarious but not
caespitose, grows under Carpinus betulus, and produces rhizomorphs. Although the
genus Musumecia was originally described with inamyloid spores (Vizzini et al.
2011), the spores of M. bettlachensis (holotypus TO HG2284) examined under a
standardized procedure by some of the authors turned out to be weakly amyloid in
grey colour. Moreover, the spores of M. sardoa are clearly amyloid. Thus, the
amyloidity feature should not be used to qualitatively discriminate spores of
Musumecia and Pseudoclitocybe, and so, the generic diagnosis has to be amended
accordingly.
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Fig. 116 Phylogenetic relationships of Musumecia based on LSU sequences. Bayesian
posterior probabilities (PP ≥ 0.90) and RAxML bootstrap values (BP ≥ 70%) are shown above
or below the branches. New taxa are in blue ex-type specimens in bold.
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Fig. 117 Phylogenetic relationships of Musumecia based on ITS sequences data. Bayesian
posterior probabilities (PP ≥ 0.90) and RAxML bootstrap values (BP ≥ 70%) are shown above
or below the branches. New taxa are in blue and ex-type specimens in bold.
Key to the known species of Musumecia
1. Pileus ivory-white to cream-white………………………………M. bettlachensis
1. Pileus dark coloured…………………………………………………………….2
2. Spores minutely ornamented, presence of hymenial cystidia and clamp-connections
abundant in all tissues………………………………………………….M. alpina
2. Spores smooth, absence of hymenial cystidia and clamp-connections rare and
scattered…………………………………………………………………………..3
3. With abundant white rhizomorphs at the stipe base; pileipellis with cystidioid
terminal elements…………………………………………………M. vermicularis
3. Without rhizomorphs; pileipellis without cystidioid elements …………M. sardoa
334. Musumecia alpina L.P. Tang, J. Zhao & S.D. Yang, sp. nov.
MycoBank number: MB 812873, Facesoffungi number: FoF 02046, Figs
118–120
Etymology: Derived from latin alpinus, relative to the Alps, in reference to their
preference for mountain habitats.
Holotype: MHKMU 182
Colour codes follow Kornerup and Wanscher (1981).
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Habit mycenoid. Pileus 3–4 cm in diam., applante or slightly depressed around
umbo, dark grey (1E1-2, 4E1) to grey-black (4E2, 7E2, 8E2) over centre, paler
towards the margin, greyish-black (1D3, 2D1–2) to greyish-white (2C2, 3C2), with a
vague to evident, greyish to pale grey zone at margin; surface covered with dense
tomentum or pubescence; margin slightly inflexed or involute, greyish-white (2B1,
3B1–2, 4B1); pileus context colour not changing when injured. Lamellae 0.4–0.6 cm
in width, adnate, crowded to subdistant, sinuous, grey (3C1–2) to greyish white
(1B1), interspersed with lamellulae. Stipe 5–7.5 × 0.5–0.6 cm, single, central attached
to subcentral, subcylindrical to cylindrical, slightly narrowing upwards, greyish (3B1)
to white (3A1); surface slightly smooth; base slightly enlarged, with whitish (2A1,
3A1) to white (1A1) mycelium or rhizomorphs; solid to loose when young, then
fistulose; stipe context fibrous, consistent when handled, greyish-white (2B13, B1–2)
to cream-white or white (3A1, 2A1). Smell and taste faint, not distinct. Spores
[80/4/3] (6.5–) 7.5–9 (–10) × (3.5–) 4–5 (–5.5) µm, Q= (1.35–) 1.58–2.16 (–2.49),
Qm= 1.89 ± 0.22, ellipsoid to oval, with a small apiculus, inamyloid, thin-walled,
hyaline, colourless in KOH, densely covered with irregular rugulose ornaments
(ornaments not clearly in KOH, but clearly observed in Cotton Blue and under SEM).
Basidia 35–38 × 4–5 µm, clavate, hyaline, colourless in KOH, thin-walled, 2–4
spored, predominantly 2-spored, sterigmata 6–8 µm in length. Cheilocystidia and
pleurocystidia clustered or scattered, quite similar in shape and size, 24–30 × 3–5 µm,
clavate, thin-walled, hyaline, clamped. Hymenophoral trama composed of subparallel
filamentous hyphae, 3–7 µm wide, thin-walled, hyaline, colourless in KOH.
Pileipellis made up of subparallel filamentous hyphae, 6–8 µm wide, thin-walled,
hyaline, clamped. Stipitipellis composed of subparallel filamentous hyphae, 5–7 µm in
diametre, slightly thick-walled (up to 1 µm), hyaline. Clamp-connections abundant in
every part of basidioma.
Habitat and known distribution: Alpine mountain in southwestern China.
Material examined: CHINA, Yunnan Province, Eryuan County, Ma’an
mountain, N 26°15′21.74", E100°06′04.02", alt. 3500m asl, in broad leaved forest
with Ericaceae (Rhododendron anthosphaerum, R. fictolacteum, and R. irrotatum
subsp. irrotatum) and Fagaceae (Quercus monimotricha), 22 August 2014, L.P. Tang
1778 (MHKMU 182, holotype). Yunnan Province, Eryuan County, Ma’an mountain,
N 26°15′21.74", E 100°06′04.02", alt. 3560m, in broad leaved forest with Ericaceae
(Rhododendron anthosphaerum, R. fictolacteum, and R. irrotatum subsp. irrotatum)
and Fagaceae (Quercus monimotricha), 22 August 2014, S.D. Yang 89 (MHKMU
346). Ibid. S.D. Yang 90 (MHKMU 347).
Notes: see under M. sardoa.
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Fig. 118 Musumecia alpina a, b Basidiomes from L.P. Tang 1778 (holotype) a Mature
basidiomes with a tomentose-fibrillose to pubscent pileus b Clustered basidiomes with base
enlarged stipe and white rhizomorphs at the base of stipes c, d Basidiomes from S.D. Yang 90
(MHKMU 347) c Single basidiome d Basidiomes with grey-whitish, curving lamellae and
hollow stipe. Scale bars = 1 cm.
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Fig. 119 Musumecia alpina (holotype) a Basidia, cheilocystidia, and pleurocystidia b Spores
c Pileipellis d Stipitipellis.
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Fig. 120 Spores under SEM of Musumecia alpina (holotype MHKMU 182) a–d
Basidiospores under SEM.
335. Musumecia sardoa G. Consiglio, A. Vizzini & L. Setti, sp. nov.
MycoBank number: MB 812779, Facesoffungi number: FoF 02047, Fig. 121
Etymology: Derived from latin sardous, relative to the Sardinia, the region where
it was first found.
Holotype: AMB n. 17139
Colour codes follow Kornerup and Wanscher (1981).
Habit mycenoid. Pileus 2–4 cm in diam., funnel shaped or infundibuliform, dark
reddish brown (9E3, 10E3); surface pubescent; margin strongly involute; colour not
changing when injured. Lamellae interspersed with lamellulae, decurrent, about
0.4–0.6 cm in width, rather broad, rather crowded, whitish cream. Stipe 3–5 × 0.8–1.5
cm, single, centrally attached to subcentral, subcylindrical to obclavate, slightly
widening upwards, whitish cream to slightly brownish (5A2, 5C6); surface smooth;
solid when young, becoming hollow or fistulose when mature; flesh fibrous, whitish
cream (3A1, 2A1) or slightly brownish (5B3, 6B3). Odour faintly herbaceous. Spores
[60/1/1] (5.8–) 5.9–8.5 (–9.5) × (3.5–) 3.8–4.2 (–4.9) µm, Q= (1.43–) 1.53–2.00
(–2.17), Qm= 1.76 ± 0.18, long ellipsoid to cylindrical, sometimes dacryoid, with an
apiculus up to 1 µm, thin-walled, hyaline; containing small refractive droplets
greenish in 5% ammonia, cyanophilous in Cotton Blue; smooth; amyloid, in Melzer’s
reagent the spore contour stains blackish blue, including the apiculus which stains
more intensely. The basidiospores show a tendency to form tetrads. Basidia 25–32 ×
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6–8 µm, subcylindrical to subclavate, hyaline, containing small droplets greenish in
5% ammonia, thin-walled, 4-spored, sterigmata up to 5 µm long; basidioles more or
less cylindrical, rare septa with clamps at the base of basidia and basidioles. Hymenial
cystidia absent. Hymenophoral trama subregular to irregular, composed by cylindrical
hyphae, 3.5–10 µm wide, hyaline, septate, sometimes the septa slightly contracted,
some hyphae with plates of encrusting parietal pigment. Subhymenium composed by
short elements, 3–6 µm wide. Pileipellis made up of a thin layer of periclinal
cylindrical hyphae, 4–10 µm wide, slightly entangled, with rare septa, with an evenly
grey cytoplasmic content and covered with plates of ochraceous parietal encrusting
pigment. Scattered superficial hyphae forming small erect tufts and small more or less
hemisphaerical warts. Stipitipellis composed by cylindrical, more or less parallel,
septate hyphae with a pale ochraceous citoplasmatic pigment and small plates, 4.5–10
µm wide, of parietal encrusting pigment. At the stipe apex short tufts of hyaline
smooth septate hyphae, with a rounded and reclined apex. Thromboplerous hyphae
present in pileipellis and stipitipellis, 3–7 µm wide, with an evenly greenish yellow
content. Context dextrinoid (more or less deep orange). Clamp-connections rare and
scattered, present in subpellis and in pileitrama.
Habitat and known distribution: Only known from Sardinia (Italy).
Material examined: ITALY, Sardinia, Desulo (NU), in a Pinus halepensis forest,
2 November 2004, leg. G. Consiglio, F. Franceschetti, A. Garbellotto & C. Orlandini
(Holotype Herbarium AMB n. 17139, holotype).
Notes: Species in the genus Musumecia are characterized by their clitocyboid
basidiomata, stipe more or less enlarged at the base, and more or less amyloid
basidiospores. However, M. alpina has a dark grey, zonate, and fibrous-tomentose or
pubescent pileus with the disc subumbonate in age, slightly larger basidiospores with
granular decorations on the surface, often 2-spored basidia, and this species has
cheilocystidia, pleurocystidia, and abundant clamps. Three European species have
slightly larger basidiomata, shallowly depressed or infundibuliform or pileus in age,
commonly 4-spored, smooth basidiospores without any decorations on the surface,
absence of cheilocystidia and pleurocystidia. Additionally, there are no or rare
clamp-connections in their basidiome. Musumecia bettlachensis has an ivory-white or
cream-white glabrous pileus sometimes with a small umbo when young, somewhat
smaller basidiospores (5.5–8.5 × 3.5–5 µm), and lacks rhizomorphs (Vizzini et al.
2011). Musumecia sardoa has an infundibuliform, dark reddish brown, pubescent
pileus, and amyloid spores. Musumecia vermicularis has an infundibuliform minutely
tomentose pileus lacking umbo, and smaller basidiospores (6.5–7.5 × 3.5–5 µm)
(Musumeci 2014). The discovery of M. alpina in southwestern China suggests that
Musumecia has a much wider geographical distribution ranging from East Asia to
Europe. Musumecia alpina is here reported from an alpine region of southwestern
China, growing in very different climate conditions.
Musumecia alpina and M. sardoa introduce some aberrant features for the genus:
the first displays minutely ornamented spores, abundant clamp-connections, and
hymenial cystidia; the second is characterized by its amyloid spores. As a matter of
fact, the genus Musumecia shows a marked macro- and micromorphological
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heterogeneity while evident shared morphological features are currently unknown. In
contrast, its molecular homogeneity is very high and all Musumecia species so far
known appear as a well supported monophyletic clade. Future work will be necessary
to assess the presence of yet undescribed unifying morphological and/or physiological
characters. To date, four taxa were reported in this genus. A key to the known species
in Musumecia is provided above.
Yunnan region is one of the major biodiversity hotspots in the world. Over 4000
species of fungi from different groups have been identified in this area during the last
decades (Zhang et al. 2005; Li et al. 2009, 2011b, 2014; Yang et al. 2012, 2013, 2015;
Zeng et al. 2013, 2014; Hao et al. 2014; Song et al. 2014; Tang et al. 2014; Zhao et al.
2014). Research is needed to confirm if this is a truly disjoint distribution or else there
exist specimens of M. alpina or other related taxa in the intermediate regions of
Central Asia.
Fig. 121 Musumecia sardoa (holotype) a Basidiomes b Lamella edge (interferential contrast)
c Basidiospores (interferential contrast) d Basidiospores in Melzer’s (light fase).
Boletales
Boletaceae
The mushroom family Boletaceae is composed of >1000 species in ~70 genera.
They are distributed worldwide primarily as obligate ectomycorrhizal mutualists with
vascular plants. Species in this family are characterised by producing soft, fleshy
stipitate-pileate basidiomata with a tubulose or sometimes lamellate to loculate fertile
layer (hymenophore), gasteroid basidiomata (truffles), and few secotioid basidiomata.
Cyanoboletus Gelardi, Vizzini & Simonini
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The genus Cyanoboletus was erected in 2014 to accommodate three existing
species that were phylogenetically shown as a clade distinct from Boletus (Wu et al.
2014, Vizzini 2014). It is typified by the European Cyanoboletus pulverulentus (Opat.)
Gelardi, Vizzini & Simonini. All three species exhibit an intense bluing colour
reaction in the flesh when exposed to air, which, although not unique to the group, is a
distinctive field character uniting them. They associate with both coniferous and
broadleaf trees worldwide. According to Species Fungorum (www. speciesfungorum.
org) and this report, five species are currently accepted for the genus. The
phylogenetic tree for Cyanoboletu is presented in Fig. 122.
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Fig. 122 Phylogenetic placement of the new species Cyanoboletus hymenoglutinosus. a Best
maximum likelihood circle phylogram recovered using RAxML of an LSU dataset including
the new species Cyanoboletus hymenoglutinosys (DC14-010) and the alignment of Wu et al.
(2014). Tree is rooted with Suillus spp. (HKAS57622 and HKAS57748), following the
topology of Wu et al. (2014). The clade containing C. hymenoglutinosus is magnified to the
left. Numbers on branches are percent nonparametric bootstraps. b Best maximum likelihood
circle phylogram recovered using RAxML of an ITS dataset including the new species
Cyanoboletus hymenoglutinosys (DC14-010) and the 100 best hits on GenBank identified
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using blastn. Tree is arbitrarily rooted using Xerocomus badius. The Cyanoboletus clade
containing C. hymenoglutinosus is magnified at right. Numbers on branches are percent
nonparametric bootstraps.
336. Cyanoboletus hymenoglutinosus D. Chakr., K. Das, A. Baghela, S.K. Singh &
Dentinger, sp. nov.
Index Fungorum number: IF 551541, Facesoffungi number: FoF 02048, Figs 123,
124
Etymology: Named after characteristic highly glutinous hymenium layer
Holotypus: D. Chakraborty & K. Das DC 15-010 (H).
Diagnosis: Distinguished from American species: Cyanoboletus pulverulentus
by its highly glutinous hymenium layer, pileipellis and differently coloured (yellowish
orange to brownish orange) pore surface.
Pileus 16–25 mm. diam.; hemisphaerical when young, becoming convex with
maturity; surface rough, highly glutinous, brown (6E5–6) or brownish orange (6C6–7),
mostly darker after maturity; margin entire with narrow sterile flap of tissue. Pore
surface narrowly depressed near stipe, yellowish orange, orange to greyish orange or
brownish orange (5B5–6, 6C5) instantly becoming bluish black (20F4–5) on bruising;
pore 2–3/mm, simple, rounded, mostly stuffed. Tube 3–5 mm long, narrowly
adnate-sinuate, pale yellow to pastel yellow (1A3–4), becoming bluish black (20F4–5)
after bruising. Stipe 50–65 ×5–8 mm, central, cylindrical, with slightly bulbous base,
yellow at apex (near pileus juncture), greyish red (7–8B5) or darker up to black on
bruising, surface scaly-pruinose with longitudinal striations on the upper half, highly
glutinous. Context solid (in pileus and stipe); context in pileus pale yellow to pastel
yellow (1A3–4), immediately becoming blue on exposure, reddish yellow to melon
yellow (4A7–5A6) with FeSO4, but, unchanging colour change with guiacol and KOH.
Spore print not found.
Basidiospores 11.6–12.8–14.8 × 4.8–5.2–5.8 µm (n = 20, Q = 2.31–2.71–2.79),
inequilateral, smooth under light microscope and SEM. Basidia 34–49 × 6–8 µm, 2–4
spored, clavate to subclavate, covered by very thick gluten. Hymenial cystidia 34–50
× 5–8 µm, emergent 15–20 µm, cylindrical to subfusiform or fusiform, content mostly
hyaline, some brown pigmented, mostly associated or partly to completely submerged
in gluten. Hymenophoral trama divergent. Pileipellis 65– 100 µm thick, ixotrichoderm,
composed of erect elements, terminal cell 17–36 × 6–7 µm, mostly with oval to
subfusoid apices, brown pigmented, heavily encrusted, wall up to 0.7 µm. Stipitipellis
320–340 µm, somewhat ixocutis, composed of subrepent to loosely interwoven
hyphae submerged under moderately thick gluten, fertile, with caulobasidia and
caulocystidia in groups. Caulocystidia 19–48 × 8–10 µm, encrusted, gelatinous.
Habitat and distribution: Under Castanopsis sp. in upper Phadamchen area,
humid temperate mixed (broadleaf and coniferous) forests dominated by species of
Cryptomeria, Pinus, Castanopsis and bamboos. Producing basidiomata in the rainy
season. Uncommon, found in East district of Sikkim (India).
Material examined: INDIA, Sikkim, East district, Upper Phadamchen, 29 July
2014, D. Chakraborty & K. Das, DC 14-010 (holotype, CAL; isotype, AMH).
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Notes: LSU sequence data from the holotype (DC 14-010) was added to a dataset
consisting of all LSU used in Wu et al. (2014). Multiple sequence alignment was
achieved using the Practical Alignment using Sate and TrAnsitivity (PASTA)
algorithm (Mirarab et al. 2014). The resulting alignment was used for maximum
likelihood analysis implemented in RAxML v8.1.17 (Stamatakis 2006, 2014; Ott et al.
2007) using a GTRGAMMA model and branch support assessed using rapid
bootstrapping set to terminate automatically based on the MRE criterion. The LSU
sequence of DC 14-010 was strongly supported (94% bootstrap) in a clade with
Cyanoboletus pulverulentus and three unidentified taxa (Fig. 122a). The ITS sequence
of DC 14-010 was queried against GenBank (Benson et al. 2013) using blastn
(Altschul et al. 1990). The top 100 best hits in GenBank were downloaded and
combined with the newly generated sequence. Multiple sequence alignment and
phylogenetic analysis were carried out as above. Similar to the LSU dataset, DC
14-010 was strongly supported (97% bootstrap) in a clade composed of multiple
sequences from Cyanoboletus pulverulentus, C. sinopulverulentus, and two
environmental sequences (Fig. 122b). Taken together, independent phylogenetic
analyses of LSU and ITS sequences unequivocally place DC 14-010 with close
affinity to Cyanoboletus spp.
Cyanoboletus hymenoglutinosus is characterized by highly glutinous basidiomata
(always associated with mud particles on gluten), yellow- to brown-orange pore
surface with stuffed pores, instantaneously changing (to blue-black) pore surface and
context, typically highly glutinous hymenial layer, basidia distinctly covered by thick
gluten and the apparent association with Castanopsis. Morphologically, Cyanoboletus
pulverulentus (Opat.) Gelardi, Vizzini & Simonini (Europe, North America), C.
sinopulverulentus (Gelardi & Vizzini) Gelardi, Vizzini & Simonini (similar
distribution: China, adjacent to Sikkim, India) and C. rainisii (Bessette & O.K. Mill.)
Gelardi, Vizzini & Simonini (North America) look very similar to the present species.
But, both C. pulverulentus and C. rainisii lack the typical glutinous pileipellis (cutis in
C. pulverulentus and trichoderm in C. rainisii). All three earlier species never shows
entirely glutinous hymenial layer and gluten-covered basidia, which is the striking
feature of the present species i.e. C. hymenoglutinosus (Smith and Thiers 1971,
Bessette et al. 2010, Gelardi et al. 2013, Vizzini 2014). Moreover, C. pulverlentus is
separated by differently coloured pore surface (“yellow when young, darkening to
golden yellow to brownish yellow when mature” as in Bessette et al. 2010). Similarly,
C. sinopulverulentus has deep yellow pore surface (never with orange pore surface
like C. hymenoglutinosus) and unstuffed pores (Gelardi et al. 2013) whereas, in C.
rainisii the pore surface becomes dark green (not blue-blak) when bruised and the
spores are much larger (10–17 × 4.2–7 µm as mentioned in Bessette et al. 2010).
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Fig. 123 Cyanoboletus hymenoglutinosus (holotype) a, c Fresh basidiomata b Pore surface
before and after bruising d Longitudinal section through plugged tubes e Hymenial layer
submerged in gluten f, g Basidia covered by thick gluten h Tube edge i Transverse section
through pileipellis j Transverse section through stipitipellis k Caulocystidia l, m
Basidiospores. Scale bars: a, b = 1cm, d = 100 µm, e–m = 10 µm.
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Fig. 124 Cyanoboletus hymenoglutinosus (holotype) a Basidiospores b Basidia c Hymenial
cystidia d Caulocystidia e Transverse section through pileipellis. Scale bars: a–e = 10 µm.
Leccinellum Bresinsky & Manfr. Binder
In the family Boletaceae (Basidiomycota, Agaricomycetes, Boletales), leccinoid
members (boletes with scabrous stipe surfaces) are some of the dominant
ectomycorrhizal fungi that associate with coniferous and broadleaf trees in the
Himalayas. Leccinellum was segregated from Leccinum to accommodate taxa with a
pileipellis composed of a palisade of swollen hyphal tips and a yellow hymenophore,
but not including three taxa with similar features, now belonging to Hemileccinum
Šutara (Šutara 2008), based on phylogenetic evidence (Bresinsky and Besl 2003).
Together, these two new genera represent Leccinum sect. Luteoscabra Singer, who
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separated these boletes with scabrous stipe surfaces but yellow hymenophores from
the remainder of Leccinum (Singer 1947). According to Species Fungorum
(www.speciesfungorum.org) and this report, 9 species are currently accepted for the
genus. The phylogenetic tree for Leccinellum is presented in Fig. 125.
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Fig. 125 Phylogenetic placement of the new species Leccinellum indoaurantiacum a Best
maximum likelihood circle phylogram recovered using RAxML of an LSU dataset including
the new species Leccinellum indoaurantiacum (DC 14-019) and the alignment of Wu et al.
(2014). Tree is rooted with Suillus spp. (HKAS57622 and HKAS57748) following the
topology of Wu et al. (2014). The clade containing L. indoaurantiacum is magnified to the
left. Numbers on branches are percent nonparametric bootstraps b Best maximum likelihood
circle phylogram recovered using RAxML of an ITS dataset including the new species
Leccinellum indoaurantiacum (DC14-019) and related leccinoid taxa. Tree is rooted with
Harrya chromapes following the topology of Wu et al. (2014). The clade containing L.
indoaurantiacum is magnified to the right. Numbers on branches are percent nonparametric
bootstraps.
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337. Leccinellum indoaurantiacum D. Chakr., K. Das, A. Baghela, S.K. Singh &
Dentinger, sp. nov.
Index Fungorum number: IF 551569, Facesoffungi number: FoF 02049, Figs 126,
127
Etymology: Named after leccinoid specimens (collected from India) with an
orange pileus like in Leccinum aurantiacum (Bull.) Gray.
Holotypus: D. Chakraborty & K. Das DC 14-019 (H).
Diagnosis: Distinguished from the Chinese species Boletus sinoaurantiacus M.
Zang & R.H. Petersen by its context (pileus and stipe) that quickly becomes pinkish
white to light pink on exposure and presence of encrusted hymenial cystidia.
Pileus 22–45 mm. diam.; hemisphaerical to convex; surface irregularly ridged
and wavy, slightly glutinous in young fruitbodies, reddish orange (7B8) gradually
paler (4A8) towards margin, orange to deep orange or light yellow to yellowish
orange (5A7–8/ 4A5–6), turning deep orange to reddish orange (5–7A8) with KOH;
margin entire with narrow sterile flap of tissue. Pore surface slightly depressed near
stipe, pastel yellow (2A4) to lemon yellow, unchanging when bruised; pore 2–3/mm,
rounded, compound. Tube 11 mm long, adnate-sinuate, light yellow (1A4),
unchanging when bruised. Stipe 80–105 ×10–13 mm, central, often curved, with
white basal mycelia, surface longitudinally striate-lacerate to squamulose or scabrate,
with brownish yellow (5–6C8) squamules on yellowish background (2–3A4–5).
Context solid in pileus and stipe; context (pileus and stipe) pale yellow (1A3), soon
becoming distinctly pinkish white to light pink when exposed. Pileus context turning
deep yellow (4A8) with KOH, reddish grey (12D2) with FeSO4 but, unchanging with
guiacol. Stipe context turning reddish grey (12D2) with FeSO4, unchanging with
KOH and guiacol. Odour and taste indistinct.
Basidiospores 13.6–16.2–19 × 5.8–6.4–7 µm (n = 20, Q = 2.19–2.52–2.92),
oblong to subfusoid, inequilateral, smooth under light microscope, olive brown.
Basidia 33–53 × 11–16 µm, 4-spored, clavate to subclavate; sterigmata 4–7 ×1–1.5
µm. Hymenial cystidia 27–75 × 8.5–12 µm, common, subcylindrical, subfusiform to
subappendiculate, content insignificant, often encrusted, incrustations distinct, mainly
located in concentric zones on neck. Tube edge fertile. Hymenophoral trama
intermediate type. Pileipellis 110–150 µm thick, ixotrichoderm, composed of erect
septate hyphae, sometimes slightly interwoven; terminal cells 10–42 × 7–16 µm,
cylindrical to subfusoid to fusoid or ventricose, subclavate to clavate or rarely
irregular, content slightly dense. Stipitipellis 100–130 µm thick, fertile, composed of
hyphae, basidia and cystidia; caulocystidia 47–85 × 10–21 µm, subfusoid, fusoid,
ventricose, ventricose-rostrate to appendiculate; caulobasidia similar to tube basidia
but less in number. Clamp connections absent in all tissues.
Habitat and distribution: Under Betula sp. in Memainchu and Kyangnosla areas,
humid subalpine mixed (broadleaf and coniferous) forests dominated by species of
Abies, Betula and Acer. (Pseudotsuga, Tsuga, Abies). Producing basidiomata in the
rainy season. Uncommon, Found in East district of Sikkim (India).
Material examined: INDIA, Sikkim, East district, Memainchu area, 2 August
2014, D. Chakraborty & K. Das, DC 14-019 (holotype, CAL; isotype, AMH); ibid.,
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East district, Kyangnosla alpine sanctuary, 7 August 2014, D. Chakraborty & K. Das,
DC 14-030, (CAL); ibid., East district, Memainchu area, 4 July 2015, D. Chakraborty,
DC 15-007, (CAL).
Notes: LSU sequence data from the holotype (DC 14-019) was added to a dataset
consisting of all LSU used in Wu et al. (2014). Multiple sequence alignment was
achieved using the Practical Alignment using Sate and TrAnsitivity (PASTA)
algorithm (Mirarab et al. 2014). The resulting alignment was used for maximum
likelihood analysis implemented in RAxML v8.1.17 (Stamatakis 2006, Ott et al. 2007)
using a GTRGAMMA model and branch support assessed using rapid bootstrapping
set to terminate automatically based on the MRE criterion. The LSU sequence of DC
14-019 was strongly supported (93% bootstrap) in a clade with Leccinellum,
Rossbeevera, Chamonixia, Octaviania, and Leccinum (Fig. 125a). The ITS sequence
from the holotype (DC14-019) was combined with sequences from related taxa
downloaded from GenBank (Benson et al. 2013). Relevant GenBank sequences were
downloaded following queries using search terms including the target taxon followed
by “AND internal transcribed spacer”, with model organisms excluded, including
Octaviania (75 sequences), Chamonixia (21 sequences), Rossbeevera (92 sequences),
and Leccinum (178 sequences). After adding the sequence of DC14-019 and removing
duplicate sequences, the final dataset consisted of 367 sequences. One sequence
(AB848541) was on the complementary strand and was corrected before alignment.
Multiple sequence alignment was achieved using the Practical Alignment using Sate
and TrAnsitivity (PASTA) algorithm (Mirarab et al. 2014). The resulting alignment
was used for maximum likelihood analysis implemented in RAxML v8.1.17
(Stamatakis 2006, Ott et al. 2007) using a GTRGAMMA model and branch support
assessed using rapid bootstrapping set to terminate automatically based on the MRE
criterion. The sequence of DC 14-019 was weakly supported (43% bootstrap) with a
clade composed of Leccinellum crocipodium, L. carpini, L. spp., and unnamed
sequences (Fig. 125b). Although support was weak, the sequence clearly did not
cluster with Leccinum s.s., and so we have provisionally included it within
Leccinellum due to its putative phylogenetic affinities with other member of this
genus.
Leccinellum indoaurantiacum is characterized by yellow-orange to orange-red
typically hemisphaerical or convex pileus, yellow unchanging pore surface, striate
squamulose to scabrate stipe with white basel mycelia, context quickly becoming
pinkish white to light pink on exposure and presence of encrusted hymenial cystidia.
In the field Boletus sinoaurantiacus M. Zang & R.H. Petersen appears to be similar
with the present species but, the earlier grows on considerably lower altitudinal zone
(1550–1680 m) and can be separated from the latter by showing unchanging context
(pileus/stipe) and absence of encrusted hymenial cystidia. Moreover, the association
of B. sinoaurantiacus with the members of Fagaceae is quite distinct (Zang et al.
2001).
Two other superficially similar species with an orange red pileus, Leccinum
aurantiacum (Bull.) Gray (reported from North America) and L. insigne A.H. Sm.,
Thiers & Watling (reported from North America and also from India), may also create
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confusion with Leccinellum indoaurantiacum. However, the context of the Leccinum
species are distinctly different, showing other colour reactions: context white initially
becoming intermediate pinkish to wine-red then finally purple gray to blackish on
exposure and pale blue with FeSO4 in L. aurantiacum; context white initially
becoming purplish gray and then blackish without any intermediate reddening on
exposure and bluish with FeSO4 in L. insigne (Bessette et al. 2010, Das &
Chakraborty 2014). Moreover, L. aurantiacum has larger basidiomata (pileus 50–205
mm, stipe 100–160 × 20 mm) and a pore surface that becomes brownish on bruising.
Similarly, in L. insigne, basidiomata are more robust (pileus up to 15 cm diam., stipe
7–12 × 1–2 cm) with smaller (11–16 × 4–5 µm) spores.
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Fig. 126 Leccinellum indoaurantiacum (holotype) a, b Fresh basidiomata c Pink context on
exposure d Tube trama e Tube edge f Basidia g Hymenial cystidia h Transverse section
through pileipellis i Caulocystidia j Basidiospores k SEM image of a basidiospore. Scale bars:
a, b = 1 cm, d = 100 µm, e, h, i = 50 µm, f, g, j = 10 µm, k = 5 µm.
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Fig. 127 Leccinellum indoaurantiacum (holotype) a Basidiospores b Basidia c Hymenial
cystidia d Caulocystidia e Transverse section through pileipellis. Scale bars: a–e = 10 µm.
Polyporales genus, incertae sedis
Galzinia Bourdot
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Galzinia is a small genus of corticiod fungi typified with G. pedicellata Bourdot.
The genus currently comprises nine species (Biodin and Gills 1990, Index fungorum
2016), but the boundaries of the genus is not well-defined and its taxonomy needs to
be revised. In our Galzinia type studies (unpublished), we noticed that several types
are in poor condition. Morphologically, Galzinia is mainly characterized by
cylindrical to urniform basidia sometimes with internal repetition, and allantoid
basidiospores (Bernicchia and Gorjón 2010). Except for G. incrustans (Höhn. &
Litsch.) Parmasto, most of the other species produce scanty basidiomes which are
difficult to see. The generic type G. pedicellata is not yet sequenced; the species is
known only from very few collections and our attemps to get sequence data from this
species have failed until now. Galzinia incrustans is the only member of the genus
sequenced, and nests in the order Corticiales, within the family Corticiaceae
(Ghobad-Nejhad et al. 2010).
338. Galzinia longibasidia Hallenb., Mycotaxon 11(2): 454, 1980.
MycoBank number: MB 112942, Facesoffungi number: FoF 02050
This is a little know species described by Hallenberg (1980) from Iran, and is
characterized by its long basidia and relatively large, subcylindrical basidiospores.
Here, we obtained ITS and LSU sequence data from the holotype material of G.
longibasidia. Blast searches at NCBI shows the new sequences as close to
Phanerochaete P. Karst. and Phlebia Fr. spp., with the highest similarity to uncultured
and insufficently identified isolates. Galzinia is shown to be a polyphyletic genus, and
G. longibasidia is nested within Polyporales but its closest relatives could be verified
via thorough phylogentic analyses of Polyporales, mainly the phlebioid clade.
Material examined: IRAN, Golestan Province, Gorgan, Golestan National Park,
on a fallen branch of a deciduous tree, 4.V.1978, Hallenberg NH2417 (GB, holotype).
Leptocorticium Hjortstam & Ryvarden
The corticioid genus Leptocorticium was typified with L. cyatheae (S. Ito & S.
Imai) Hjortstam & Ryvarden and is characterized by monomitic hyphal system with
clamps, dendrohyphidia, subulate leptocystidia, and fusiform to navicular
basidiospores (Bernicchia and Gorjón 2010). The genus currently contains seven
species and was recently discussed by Gorjón and Saitta (2014). Because no sequecne
data is avaliable from the type, the phylogenetic position of the genus is not clear.
Based on morphology, Larsson (2007) proposed that the genus may belong to the
order Corticiales. Leptocorticium tenellum is the first member of the genus for which
we present sequence level data.
Russulales genus, incertae sedis
339. Leptocorticium tenellum Nakasone, Mycol. Progr. 4(3): 253, 2005.
MycoBank number: MB 341582, Facesoffungi number: FoF 02051
The species was recently re-described by Hallenberg (2012) who studied its type
and reported some new material he collected in Chile; from one of those material we
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could obtain ITS and LSU sequence data. A megablast search of GenBank nucleotide
database at NCBI (as of 20 November 2015) using the new LSU showed that the best
hits were isolates of Aleurodiscus Rabenh. ex J. Schröt. spp. and Lentinellus ursinus
(Fr.) Kühner. Blast searches of the new ITS (only 392 bp recovered) showed the best
three hits to be Lentinellus subargillaceus (Kauffman) R.H. Petersen, and L.
tridentinus (Sacc. & P. Syd.) Singer, with 99% over 41% query coverage. Therefore,
Leptocorticium tenellum is shown to be a member of the order Russulales.
Material examined: CHILE, Los Lagos, Parque Nacional Puyehue, Trail Los
Rapidos, Circuito, 40° 44' 01.4'' S, 72° 18' 44.1'' W, elev. 496 m, on bamboo,
22.II.2010, Hallenberg (GB NH16311, reference specimen designate here).
Hymenochaetales
Hymenochaetaceae
Hymenochaetaceae, belonging to Hymenochaetales, is one of the most important
families in Basidiomycota. This family accommodates some serious forest pathogens
(Cui et al. 2015) and important medicinal species (Zhou et al. 2016a). In the last two
decades, molecular phylogeny extremely improved the knowledge of
Hymenochaetaceae at the generic level. At least seven genera were newly erected
(Niemelä et al. 2001, Dai 2010, Rajchenberg et al. 2015, Zhou 2015a, Zhou et al.
2016a) and some old genus names were also reused (Dai 2010). Meanwhile, studies
on global diversity of certain genera in Hymenochaetaceae extremely increased
known species number (Zhou 2015b, Zhou and Dai 2012, Zhou et al. 2016a, b).
However, there are still some undescribed species that need to be introduced. The
phylogenetic tree for Fomitiporia is presented in Fig. 130.
Fomitiporia Murril
Fomitiporia is characterized by pileate to resupinate basidiomata, hymenial setae
present in some species, dextrinoid basidiospores, and a dimitic hyphal system
through all the basidioma (Decock et al. 2007). The genus has about 40 species
described, many of those have been collected on live tree hosts, suggesting some
levels of host-specificity (Amalfi et al. 2012; Dai et al. 2008). Historically, due to low
morphological variation, several taxa represent morphological complexes of cryptic
species, thus the phylogenetic reconstructions based on molecular data have been
playing a crucial role in the discovery of unknown lineages (Decock et al. 2007;
Vlasák and Kout 2011). Neotropical region presents a high diversity unknown
(Amalfi and Decock, 2013; Amalfi et al. 2014), mainly because there are many areas
without records of collections. Two new pileate species of Fomitiporia from south
Brazil are described in this study. The phylogenetic tree for Fomitiporia is presented
in Fig. 128.
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Fig. 128 Phylogram generated from Maximum Likelihood (RAxML) analysis based on
combined nrLSU, nrITS, EF and RPB2 sequence data of Fomitiporia. Maximum Likelihood
bootstrap support values greater than 70 % and Bayesian posterior probalities (BPP) greater
than 0.98 are indicated above and below the nodes (BS/BPP). In the BI analysis average
standard deviation of split frequencies = 0.005 and the bootstopping criteria of RAxML
indicated 204 pseudoreplications as sufficient to access the internal branch support. New taxa
are in blue and species for which obtained sequences are based on type material have names
in bold. The tree is rooted with Phellinus uncisetus.
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340. Fomitiporia atlantica Alves-Silva, Reck & Drechsler-Santos, sp. nov.
Index Fungorum number: IF 551915, Facesoffungi number: FoF 01831, Fig. 129
Etymology: referring to the vegetacional type where the fungus was found, the
Atlantic forest.
Holotype: FLOR 58554.
Basidiomata perennial, pileate, sessile and mostly broadly attached, semicircular,
solitary to imbricate, then with the different pilei fusing, with a nodulous aspect when
emerging from the wood, obtriquetrous to obungulate, also triquetous, projecting
12.5–51 mm, 21–66 mm wide and 20–82 mm thick at the base, woody consistency
when dried; pileus glabrous, concentrically zonated with multiple narrow bands,
slightly sulcate, faintly cracked when old, dull, when fresh pilear surface greyish
brown 11E3, violet brown11F4 to dark brown [7 F(6–8)], upon dried brown[6
E(5–8)]to olive brown [6 F(4-8)] when young [6 E(5–8)], becoming dark brown [6
F(5–8)]; margin finely velutinous, round, folded, thick, 3.5–19 mm thickness, sterile,
olive brown [6 F(4–8)], yellowish brown to brown [5 DEF(6–8)]; pore surface light
greyish brown (5D8) when young, greyish brown to cinnamon; pores rounded to
angular, 6–8 (–9) per mm, (60–) 70–110 (–120) µm diam. (mean = 89 µm);
dissepiments entire, (30–) 40–120 µm (mean = 67 µm) thick; tubes distinct to mostly
indistinctly stratified, with several layers (up to 15 layers in the oldest basidioma),
those interleaved with context layers usually thicker (up to five times), individual tube
layers relatively thin, sometimes difficult to distinguish, up to 2 mm tall, brown [5 EF
(4–5)] to grayish brown (5E3), the older layers filled with whitish mycelium; context
simple, up to 20 mm thick, concentrically zonate, sometimes constituted by extremely
thin black lines (invisible to the unaided eye) that made the separation between
growth layers of the context, with dense texture and woody consistency, golden to
brownish yellow [5 BC (7–8)], with a distinct dark line at the surface, which is dark
brown when young, becoming black, sometimes with a resinous aspect. Hyphal
system dimitic in all parts; generative hyphae simple septate, hyaline to pale yellow,
sparingly branched, 2–3 µm diam; skeletal hyphae golden brown to reddish brown,
unbranched, thick-walled, rarely with local swelling up to 8 µm, in the context
4–5(–5.5) µm diam., the lumen 1.5–3 µm wide, in the hymenophoral trama 4–5(–6)
µm diam., the lumen 1.5–3(–4) µm wide. Hymenium: hymenial setae absent, other
sterile elements presents (as basidioles), hyaline, thin-walled; basidia subglobose to
globose, hyaline, tetrasporic, 9–11 × 7–8 (mean = 9.5 × 8 µm) Q = 1–1.3 µm (meanQ
= 1.18 µm); basidiospores subglobose, globose to obovoid, with the wider portion
displaced towards the apex,(4.5–) 5–5.5 (–6) × 4–5.5 µm (mean = 5.1 × 4.8 µm) Q =
1–1.25 µm (meanQ = 1.08 µm) (n = 40), hyaline, strong to weakly dextrinoid,
cyanophilous, thick-walled, smooth.
Material examined: BRAZIL, Santa Catarina, Blumenau, Parque Natural
Municipal São Francisco de Assis, 26°55'17"S 49°04'18"W, on dead cut tree, 21
November 2014, G. Alves-Silva 640, (FLOR 58554, holotype); Ibid., on dead
standing trunk, 15 September 2015, F. Bittencourt 507 (FURB 47591).
Notes: Fomitiporia atlantica is mainly characterized by the nodulose aspect of
basidiomata with thick-rounded margin and darkness aspect of basidiomata when
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fresh, the narrowly zonated pilear surface, the zonation of the context (with variable
presence of concentric thin black lines invisible to the unaided eye) and by the
irregular layers of tubes (Fig. 2i); microscopically, the new species presents dimitic
hyphal system and globose, subglobose to obovoid basidiospores with variable
dextrinoid reaction. Fomitiporia atlantica shares with F. castilloi Decock & Amalfi
the nodulous basidiomata (better observed in young specimens). However, F. castilloi
is described by Amalfi and Decock (2013) from French Guiana as presenting distinct
hymenial setae and slightly larger basidiospores in range and average (6.2 × 5.2 µm),
besides having a wider pilear zonation as well as an azonated context. Fomitiporia
gabonensis Amalfi & Decock also presents imbricate basidiomata and variable
dextrinoid basidiospores. Nevertheless, F. gabonensis was described by Amalfi et al.
(2010) from Africa (Gabon) as presenting smaller basidiospores (4.7 × 4.1 µm) and
acute thinner margin. Besides the morphological evidences, F. atlantica is also
supported by molecular results. The phylogenetic analysis (Fig. 128) showed the two
specimens clustered together in a strong supported clade (BS = 100, BPP = 1).
Fomitiporia atlantica forms a more inclusive clade with other two species, F.
subtilissima (described below) and another undescribed species from Brazil (FLOR
58555). This clade displays nested vicinity to F. apiahyna sensu lato clade (Amalfi et
al. 2014), appearing as a sister clade of this lineage. Fomitiporia atlantica differs from
F. apiahyna (Speg.) Robledo, Decock & Rajchenb. sensu Amalfi and Decock (2013)
mainly by its slightly smaller basidiospores (F. apiahyna = 5.9 × 5.1 µm) and pileus
slightly sulcate and cracked, conspicuous features in F. apiahyna.
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Fig. 129 Fomitiporia subtilissima (FURB 47437) a Basidiomata in situ c Abmenial surface
showing the concentric zonation and spathulate aspect of basidioma. Fomitiporia subtilissima
(holotype) f, g Details of context and tubes f Black line at the surface g Context and tube
layers j Hymenophoral surface m Basidiospores. Fomitiporia atlantica (FURB 47591) b
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Darkness aspect of basidiomata in situ e Abhymenial surface h Black line at the surface k
Nodulous basidioma l Hymenophoral surface n Basidiospores. Fomitiporia atlantica
(holotype) d Detail of slightly cracked abhymenial surface i Context and tube layers. Scale
bars: a, b = 50 mm, c–e, g and i, l = 20 mm, f, h = 2 mm, m, n = 5 µm.
341. Fomitiporia subtilissima Alves-Silva, Reck, & Drechsler-Santos, sp. nov.
Index Fungorum number: IF 551916, Facesoffungi number: FoF 01832, Fig. 129
Etymology: referring to the relatively thin basidiomata.
Holotype: FURB 47557.
Basidiomata perennial, pileate; sessile, subdimidiate to pseudostipitate, the
pseudostipe formed by successive deposited layers, single or with distinct pilei
developing from the same point, semicircular, flabelliform to slightly spathulate,
aplanate to convex, in section and near the base slightly obtriquetrous, projecting
18–162 mm, 17–96 mm wide and 6–40 mm thick at the base, soft, woody consistency;
pileus glabrous,concentrically zonated with multiple narrow bands, moderately
sulcate, light brown [6 D(5–8)], brownish orange [6 C(7–8)] to brown [6 E(5–8)],
becoming dark brown [6 F(5–8)] to black; margin acute to obtuse, sterile, light brown
[5 D(6–8)] to yellowish brown [6 E(5–8)]; pore surface grayish brown [6 F(3)] to
cinnamon, near the base the newest tube layer presents an wider sterile yellowish
brown [5 D( 6–8)] margin (up to 2 mm), contrasting with the precedent layer
recovered by brown [6 E(6–8)]context; pores rounded to angular, (4–) 5–9 per mm,
(70–) 80–131 (–170) µm diam. (mean = 107 µm); dissepiments entire, 30–76 (–100)
µm (mean = 45 µm) thick; tubes distinctly stratified, up to 9 layers, individual layers
0.3–3 mm thick, with context among it, up to 2 mm thick, greyish brown [6 F(3–4)] to
brown [6 E (6–8)], but the youngest (active) layer greyish brown (6E4) to cinnamon,
the older layers filled with whitish mycelium; context simple, concentrically zonate,
up to 6mm thick, with soft and hard to woody consistency, light golden brown to light
brownish yellow [5 BC (7–8)], with a distinct dark line at the surface. Hyphal system
dimitic in all parts; generative hyphae simple septate, hyaline to pale yellow, mildly
branched, 1.5–2 (–2.5) µm diam; skeletal hyphae golden brown to reddish brown,
unbranched, thick-walled, occasionally with constrictions uncompleted becoming
local swellings up to 8 µm diam., in the context 3–5 µm diam., the lumen 1–3 µm
wide, in the hymenophoral trama 3.5–4.5 µm diam., the lumen 1–3 µm wide.
Hymenium: hymenial setae absent, other sterile elements presents (as basidioles),
hyaline, thin-walled; basidia subglobose to globose, hyaline, tetrasporic, 9–10 × 7–9
(mean = 9.2 × 8.1 µm) Q = 1–1.3 µm (meanQ = 1.14 µm); basidiospores subglobose,
globose to obovoid, the wider portion displaced towards the apex, 4–5 × 4–4.5(–5)
µm (mean = 4.5 × 4 µm) Q = 1–1.25 µm (meanQ = 1.13 µm) (n = 40), hyaline,
slightly to moderately dextrinoid and cyanophilous, thick-walled and smooth.
Material examined: BRAZIL, Santa Catarina, Blumenau, Parque Natural
Municipal São Francisco de Assis, 26°55'17"S 49°04'18"W, growing on dead root of
living Sloanea guianensis (Aubl.) Benth. (Elaeocarpaceae), 28 July 2015, F.
Bittencourt 493 (FURB 47557, holotype, isotype in FLOR); Ibid., in the base of a
living unidentified angiosperm, 13 May 2015, F. Bittencourt 428 (FURB 47437).
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Notes: The flabelliform to spathulate, thin and aplanate basidiomata, with the
presence of a pseudostipe, and the narrow concentrically zonated and sulcate
abhymenial surface characterizes this species as unique in the genus. The pseudostipe
is explained by its successive depositing forward tube layers that do not cover the
precedent layer near the base. Besides, F. subtilissima has slightly to moderately
dextrinoid basidiospores, which are relatively small when compared with other
Fomitiporia species. Due to its macroscopic features, F. subtilissima resembles some
Phylloporia species, but this genus is characterized by monomitic hyphal system and
IKI- basidiospores. Variably dextrinoid small basidiospores are also found in F.
gabonensis and F. ivindoensis Decock, Amalfi & Yombiyeni (Amalfi et al. 2010),
both described from Gabon, Africa. Fomitiporia gabonensis has thick, obtriquetrous
and broadly attached basidiomata, while F. ivindoensis has cushion-shaped to
aplanate basidiomata, but they do not have pseudostipe. The morphologic and
molecular data (BS = 100, BPP = 1) high support the new species. The phylogenetic
analysis (Fig. 128) recovered F. subtilissima in a clade nested with F. atlantica and
another undescribed species from Brazil.
Inonotus P. Karst.
Inonotus, typified by I. hispidus (Bull.) P. Karst., is one of the largest genera
within the Hymenochaetaceae; in a wide sense, this genus, accommodating more than
100 species, is distinct from other genera in Hymenochaetaceae by its annual,
non-stipitate or rarely laterally stipitate basidiocarps, poroid hymenophores and a
monomitic hyphal system (Ryvarden 2005). According to phylogenetic results, four
narrowly defined genera segregated from Inonotus sensu lato, viz. Inocutis Fiasson &
Niemelä, Inonotopsis Parmasto, Mensularia Lázaro Ibiza, Onnia P. Karst., are
accepted, whereas some species with perennial basidiocarps and/or a dimitic hyphal
system were also transferred to Inonotus (Wagner and Fischer 2002, Dai 2010, Wu et
al. 2012, Vlasák et al. 2013). Recently, Zhou (2015a) introduced a monotypic genus
Cylindrosporus L.W. Zhou & Y.C. Dai for species previously belonging to Inonotus,
while Zhou et al. (2016a) segregated Sanghuangporus Sheng H. Wu, L.W. Zhou &
Y.C. Dai and Tropicoporus L.W. Zhou, Y.C. Dai & Sheng H. Wu from Inonotus and
proposed that the remain species in Inonotus still have polyphyletic origins (Zhou et al.
2016a). Herein, a new species of Inonotus is described from Chiang Mai, Thailand.
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Fig. 130 Phylogenetic position of Inonotus shoreicola inferred from nLSU sequence data.
Topology is from maximum likelihood (raxmlGUI 1.2) analysis, and the statistical values
simultaneously above 50% for bootstrap values and 0.80 for Bayesian posterior probabilities
are indicated at the nodes. New taxa are in blue and species for which obtained sequences are
based on type material have names in bold.
342. Inonotus shoreicola L.W. Zhou, Y.C. Dai & Vlasák, sp. nov.
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Index Fungorum number: IF 551555, Facesoffungi number: FoF 02052, Fig. 131
Etymology: refers to the host genus Shorea.
Holotypus: LWZ 20140728-10 (IFP)
Basidiocarps perennial, sessile, single, ungulate, woody hard, without Odour and
taste when dry. Pileus dimidiate, projecting up to 7 cm, 20 cm wide and 8 cm thick at
base. Pileal surface pale mouse-grey to vinaceous grey, radially cracked,
concentrically zonate and sulcate; margin obtuse, ash-grey. Pore surface dark brown,
slightly glancing; sterile margin distinct, yellowish brown, up to 5 mm; pores circular
to angular, 7 per mm; dissepiments thin, entire. Context dark brown, woody hard, up
to 4 mm. Tubes yellowish brown, woody hard, tube layers distinctly stratified, annual
layer about 5 mm long; white mycelial strands present in old tubes.
Hyphal system monomitic; generative hyphae simple septate; tissue darkening
but otherwise unchanged in KOH. Contextual generative hyphae yellowish,
thick-walled with a wide lumen, rarely branched, simple septate, interwoven,
acyanophilous, 1–2.5 µm in diam. Tramal generative hyphae yellowish, slightly thickto thick-walled with a wide lumen, occasionally branched, simple septate, parallel
along the tubes, acyanophilous, 2–3 µm in diam. Hyphoidsetae absent; hymenialsetae
occasionally present, subulate to ventricose, dark brown, thick-walled, sharp pointed,
sometimes with an elongated base, 15–38 × 8–20 µm; cystidia and cystidioles absent;
basidia and basidioles not seen; rhomboid crystals present in hymenium.
Basidiospores broadly ellipsoid, yellowish, slightly thick-walled, neither amyloid
nor dextrinoid, cyanophilous, (4.4–)4.6–5.1(–5.4) × (3.2–)3.5–3.9(–4) µm, L = 4.86
µm, W = 3.75 µm, Q = 1.29–1.3 (n = 60/2).
Material examined: THAILAND, Chiang Mai Province, Sri Lanna National Park,
Mae Taeng Forests, on living tree of Shorea, 28 July 2014, LWZ 20140728-10 (IFP,
holotype), LWZ 20140728-23 (IFP); Ibid., 21 October 2013, Dai 13614 (BJFC), Dai
13615 (BJFC), 29 July 2014, LWZ 20140729-1 (IFP).
Notes: Zhou et al. (2016a) identified three clades within Inonotus and also
several species of Inonotus outside the three clades. The clade, including the generic
type Inonotus hispidus, was considered to be Inonotus sensu stricto, while the other
two clades were introduced as new genera Sanghuangporus and Tropicoporus; the
species outside any clade were also accommodated in Inonotus sensu stricto for they
have key characters of Inonotus (Zhou et al. 2016a). The current phylogeny (Fig. 130)
shows that Inonotus shoreicola is close to Inonotus sensu stricto, Sanghuangporus
and Tropicoporus. Moreover, Inonotus shoreicola fits well with the morphological
concept of Inonotus sensu Dai (2010). Therefore, we place species in Inonotus.
Inonotus shoreicola resembles the pileate members of Sanghuangporus and
Tropicoporus in its perennial basidiocarps, cracked pileal surfaces and colored
basidiospores; however, these two genera are distinguished by having dimitic hyphal
system at least in trama (Zhou et al. 2016a), while Inonotus shoreicola has a
monomitic hyphal system in both context and trama. Inonotus pachyphloeus (Pat.) T.
Wagner & M. Fisch. also has perennial basidiocarps and a monomitic hyphal system
as I. shoreicola, but differs in the presence of hyphoid setae (Fidalgo 1968, Dai 2010).
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Some pileate species of Fomitiporia Murrill, such as F. hartigii (Allesch. &
Schnabl) Fiasson & Niemelä and F. robusta (P. Karst.) Fiasson & Niemelä, also have
cracked pileal surfaces (Dai 2010), which make them similar to I. shoreicola
especially in the field. However, in micromorphology, Fomitiporia is characterized by
a dimitic hyphal system and hyaline, dextrinoid basidiospores (Dai 2010).
Inonotus shoreae (Wakef.) Ryvarden, originally described from India, also
inhabits Shorea like I. shoreicola, and is a serious parasite on roots and butts of
Shorea (Sharma 1995). These two species could be easily differentiated in the field: I.
shoreae has annual basidiocarps with much larger pores (2–4 per mm, Sharma 1995).
Moreover, Inonotus shoreae has shorter basidiospores (3.5–5 × 2.5–3 µm) than I.
shoreicola (Sharma 1995). It is noteworthy that I. shoreicola is relatively common
and has been considered to be a medicinal fungus in Thailand (Fig. 131).
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Fig. 131 Inonotus shoreicola a Basidiocarps in situ (holotype) b Cultivations c
Basidiospores d Hymenial setae e Hyphae from trama f Hyphae from context.
Polyporales
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Ganodermataceae
Ganoderma P. Karst
The genus Ganoderma was established by Karsten (1881) with Ganoderma
lucidum (W.Curt:Fr.) as the only species (Moncalvo and Ryvarden 1997). Ganoderma
species are distributed all over the world, in tropical and temperate regions, although
usually found in subtropical and tropical regions, since it can withstand \hot and
humid conditions (Pilotti 2004). Ganoderma species are not classified as edible
mushrooms, as the fruiting bodies are always thick, corky and tough and do not have
the fleshy texture characteristic of true edible mushrooms (Singh et al. 2013).
Ganoderma has long been regarded as one of the most important medicinal fungi
worldwide (Paterson 2006), and laccate species of Ganoderma, have been used as
medicinal fungi in traditional Chinese medicine for over two millennia (Anon 1955).
China is very rich in Ganoderma species, with at least 80 species names (Zhao and
Zhang 2000; Wang et al. 2009a; Cao et al. 2012, 2013), although part of them are
synonyms. Ganoderma P. Karst. (Ganodermataceae, Polyporales) is characterized by
its double-walled basidiospores with interwall pillars, bears an apical umbo, often
shrunk, and the apex appears then truncate (Li et al. 2013c). The taxonomy of the
genus is, however, poorly circumscribed, not universally accepted, and has been
described as being in a state of chaos (Ryvarden 1991). The objective of the present
study is to introduce a new Ganoderma sp.with a description from Hainan Province,
China and compare it with similar taxa.
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Fig. 132 Phylogram generated from Maximum likelihood (RAxML) analysis based on ITS
and RPB2 sequence data. Maximum likelihood bootstrap support values greater than 50% are
indicated above the nodes, new species is in red and ex-type specimens in bold. The tree is
rooted with Tomophagus colossus.
343. Ganoderma wuzhishanensis T.C. Wen, K. Hapuarachchi & K.D. Hyde, sp. nov.
Index Fungorum number: IF 551681, Facesoffungi number: FoF 00915, Fig. 133
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6422
Etymology: refers to the type collecting site “Wuzshishan Mountain”, Hainan,
China
Holotype: GACP 14081689
Basidiocarp annual, sessile, woody, Pileus 3–5.5 × 1–3 cm, up to 1.5 cm thick at
the base, suborbicular, plano convex, sub applante. Upper surface; hard, several layers
thick, deep buff (460) to leaf brown (489), crust overlies the pithy context, not
cracking, containing fibrous pithy context, strongly laccate, no concentrically sulcate
zones, no differentiated zones. radially rugose, margin soft or with numerous
undulations and irregularities, 5 mm thick, rounded and concolorous with the pileus.
Lower surface light straw (384) basidiospores. Pore surface light straw (384), tubes
up to 0.7 mm long in total, middle buff (359) to middle brown (411), pores circular or
sub circular or isodiametric. Context up to 1.5 cm thick, dry, triplex, lower layer;
golden brown (414), fibrous/pithy, composed of coarse loose fibrils, soft, middle
layer; red oxide (446), upper layer: dark camouflage red (436), woody, not cracking,
composed of tightly interwoven, finer fibrils, dulling when cut, trimitic hyphal
system, generative hyphae; 0.8 –2(–3) µm ( x =1.4, n = 30) in width, thin walled,
colourless, hyaline, Skeletal hyphae; (–2)3–3.5(–4) µm ( x = 3, n = 40) in width,
Golden brown (414) to Light brown (320) in 5% KOH, dextrinoid, thick walled,
ligative hyphae; (–0.5)1– 2(–3) µm ( x =1.8, n = 40) in width, Dark camouflage red
(436), 2) to Light brown (320), thick walled, branched, intertwined the skeletal
hyphae. Basidiospores 7–9 (–10) × (–3)4–6 µm ( x = 8.4 × 5, n = 30, Q = 1.3–2.7, Q =
1.7, with myxosporium). 5– 7 (–8) × (–2)3–4 µm ( x = 6.2 × 3.3, n = 30, Q =
1.43–3.18, Q = 1.99, without myxosporium), elongate, Dark camouflage red (436) to
Light brown (320), eusporium bearing fine, short and distinct echinulae, overlaid by a
hyaline myxosporium, bitunicate. Cuticle hymeniodermiformic, Light brown (320),
composed of apically acanthus like branched cells, dextrinoid.
Habitat and distribution: On a decaying wood log, accompanied in humus rich
soil with over heavily rotted litter in forest,.mossy coniferous forests, producing
basidiomata from late summer to late autumn, only found in Hainan Province, China.
Material examined: CHINA, Hainan Province, Wuzhishan Mountain, Coniferous
rainforest, 18°"N 110 "E, elev. 1350 m, 16 August 2014, collector T.C Wen,
(GACP14081689, holotype).
Notes: Ganoderma wuzhishanensis is a new member of Genus Ganoderma (Fig.
132) and it clustered with G. multi-pileum Hou, which is characterized by two kinds
of pilei, one from the stalk with some of the stipes and pilei growing together, and the
other growing from the lower pilei; a thin crust, composed of enlarged and bulbous
ends of hyphae, 16.5 × 2–6 µm; and basidiospores 8–9 × 4 µm, ovoid, truncate, with
numerous and minute echinulae 4–6 µm (Wang et al. 2009a). Ganoderma
wuzhishanensis is morphologically similar to Ganoderma tropicum (Jungh.) Bres.
which is characterized by its laccate surface, large spores (distinctly larger than for
most species in the G. lucidum–complex), 11–14 × 7.5–10 µm and the slightly small
pileus size, deep buff (460) to leaf brown (489) pileus colour, grow as invidual but
live as a group, without concentrically sulcate zones, small tube size, triplex context,
basidiospores 7–9 × 4–6 µm, elongate, dark camouflage red (436) to light brown
6423
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(320) and bitunicate. The species is currently only known from the type locality,
Wuzshishan Mountain, Hainan, China.
Fig. 133 Ganoderma wuzhishanensis (holotype) a Upper surface b Lower surface c Cutting
surface d Pores in the lower surface e–f Spores g Vegetative hyphae h Skeletal hyphae i
Ligative hyphae. Scale bars: a–c = 1 cm, d = 0.5 cm, f =10 µm, g–i = 5 µm.
6430
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6464
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6468
Polyporales genus, incertae sedis
Dentocorticium (Parmasto) M.J. Larsen & Gilb.
The genus Dentocorticium, typified with D. ussuricum, currently comprises
seven species of corticioid fungi with resupinate, smooth to dentate hymenophore,
monomitic hyphal system with clamps, and smooth, non-amyloid basidiospores. The
species possess dendrohyphidia (dendrophyses) and lack cystidia (Boidin and Gilles
1998, Duhem and Michel 2009).
344. Dentocorticium ussuricum (Parmasto) M.J. Larsen & Gilb., Norw. Jl Bot. 21(3):
226, 1974.
MycoBank number: MB 312868
We studied the type material of D. ussuricum and obtained ITS sequence of an
authentic material conforming to the type, to deduce the relationships of the type of
the genus. A megablast search of GenBank nucleotide database at NCBI (as of 16
November 2015) using the new ITS showed that the best hits were isolates of
Dentocorticium sulphurellum (Peck) M.J. Larsen & Gilb. with 95–96% identity over
98% query coverage, followed by isolates of Trametes Fr. spp. According to Binder et
al. (2013), Trametes and Dentocorticium sulphurellum reside in the core polyporoid
clade (Polyporaceae, Polyporales). Here, the position of generic type D. ussuricum
within the family Polyporaceae, and congeneric relationship of D. sulphurellum with
D. ussuricum are established.
Material examined: RUSSIA, Primorsk, Insula Petrova, on Actinidia arguta, 1
September 1961, leg. A. Raitviir (TAA 42424, holoype). CHINA, Jilin Province,
Antu County, Erdaobaihe, south of Erdaocun town, ca. 30 km from Erdaocun towards
Changbaishan Mountain and Lake; forest mainly with Abies, Picea, Larix, Acer spp.,
also Betula, Populus, Tilia amurensis, and Pinus; 42.205 Lat., 128.165 Long., elev. ca.
1100 m; on hanging branch of cf. Acer, 3 cm in diam.; 11 September 2011;
Ghobad-Nejhad 2465 (Ghobad-Nejhad ref. collection, and BJFC).
Polyporaceae
Lentinus
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 (Index
Fungorum 2016) and, species are able to survive over a wide temperature range, are
abundant in boreal, temperate and tropical regions (Corner 1981; Pegler 1983;
Karunarathna et al. 2011). The phylogenetic tree for Lentinus is presented in Fig. 134.
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Fig. 134 Phylogeny of Lentinus stuppeus and related species in the genus based on nrITS
sequences, inferred by maximum likelihood (ML) analysis. Numbers at internodes refer to
confidence estimates based on 100 rapid ML bootstraps (only those >50 are indicated).
Lentinus stuppeus from Thailand is in blue. Leucoagaricus barssii and Leucoagaricus
leucothites are outgroup taxa.
345. Lentinus stuppeus Klotzsch [as 'stuppens'], Linnaea 8(4): 480, 1833.
≡ Pocillaria stuppea (Klotzsch) Kuntze [as 'stupea'], Revis. gen. pl. (Leipzig) 2:
866, 1891.
≡ Panus stuppeus (Klotzsch) Pegler & R.W. Rayner [as 'stupeus'], Kew Bull.
23(3): 385, 1969.
Facesoffungi number: FoF 02054, Fig. 135
Basidiomes very small to medium. Pileus 1–5.5 cm in diam., coriaceous, deeply
umbilicate to deeply infundibuliform; margin inflexed, entire, thin at first reflexed,
surface mahogany red, dark purplish brown to almost black, dry, densly villose,
covered with curled, hispid, fibrillose hairs up to 7–8 mm long, glabrescent and finely
rimose at the centre; margin strongly and persistently involute, densely pilose.
Lamellae short decurrent, usually with some anastomosing at the stipe apex, pale
yellowish buff, narrow, up to 3–4 mm wide, moderately crowded, with 4–5 tiers of
lamellulae, edge strongly denticulate. Stipe 1.5–4.5 cm × 2–4.5 mm, central, rarely
lateral, cylindrical, slender, solid, expanding above, surface dull yellowish brown,
often with deeply purple tints, covered by cinnamon brown tomentum at the apex,
elsewhere with small, blackish, apprised squamules becoming hispid at the base;
context 2–3 mm, white to dull white in color, fibrous, consisting of a dimitic hyphal
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system with generative and skeletal hyphae. Generative hyphae 2–4 µm diam.,
hyaline, very thin walled, frequently branched, with prominent clamp connexions.
Skeletal hyphae 3–7 µm diam., hyaline with a thickened wall, with wide dichotomous
branching. Spore print cream color. Basidiospores (Fig. 135a) 6–9 × 2.3–3.4 µm [n =
30, (7.5 × 2.8 µm), Q = 2.78], cylindric, hyaline, thin walled. Basidia (Fig. 135b)
20–24 × 5–6.5 µm, clavate, bearing 4 sterigmata. Lamella-edge sterile. Cheilocystidia
(Fig. 135c) 16–36 × 4–8 µm, sinuous clavate, hyaline, thin-walled. Hyphal pegs
abundant. Hymenophoral trama hyaline, irregular, similar to context. Subhymenial
layer narrow. Pileipellis an epicutis, with reddish brown walls. Hairs comprising
fascicles of unbranched hyphae, with thickened, pigmented wall.
Habitat: On dead wood, in clusters, in rain forest dominated by Castanopsis
armata, and Lithocarpus sp.
Material examined: THAILAND, Chiang Mai Prov., Mae Taeng Dist., Ban Pha
Deng village, N19°17.123’ E 98°44. 009’, elev. 900 m, rainforest dominated by
Castanopsis armata and Pinus kesiya. 18 June 2013, (MFLU 10–0667, reference
specimen designate here).
Distribution: Ghana, Nigeria, West Cameroons, Zaire Republic, Uganda, Kenya,
Madagascar, Mauritius, Zimbabwe, South Africa (Pegler 1986), new record to
Thailand (this study).
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Fig. 135 Lentinus stuppeus (MFLU 10–0667, reference specimen) a Basidiocarps b
Basidiospores c Basidia d Cheilocystidia e Hyphal pegs f Generative hyphae g Skeletal
hyphae. Scale bars: a = 5 cm, b = 10 µm, c–g = 20 µm.
Russulales
Bondarzewiaceae
Bondarzewia
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Bondarzewia Singer was established by Singer (1940) based on B. mesenterica
(Schaeff.) Kreisel, originally described from Abies in Germany. It is a remarkable
genus because the species usually have huge and imbricate basidiocarps. Some
species are edible and medicinal mushrooms (Dai et al. 2009), while others are
pathogens on their host trees (Dai et al. 2007). The genus is characterized by an
annual growth habit, pileate basidiocarps with poroid hymenophores and it is
morphologically a polypore genus. However, it has strongly amyloid and ornamented
basidiospores and phylogenetic analysis showed that it belongs to Russulales (Larsson
and Larsson 2003). The phylogenetic tree is presented in Fig. 136.
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Fig. 136 Phylogeny of species in Bondarzewia and related species generated by maximum
likelihood based on ITS+nLSU sequence data. Branches are labeled with bootstrap
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proportions (before the slash markers) higher than 50% and Bayesian posterior probabilities
(after the slash markers) more than 0.95. New taxa are in blue and ex-type specimens in bold.
346. Bondarzewia tibetica B.K. Cui, J. Song & Jia J. Chen, sp. nov.
MycoBank number: MB 815274, Facesoffungi number: FoF 02055, Figs 137,
138
Etymology: tibetica (Lat.), referring to the locality of the type specimen.
Holotype: BJFC 016992
Basidiocarps annual, pileate, broadly attached to the substrate, imbricate, soft
corky and watery when fresh, becoming fragile upon drying; pileus fan-shaped,
projecting up to 16 cm long, 25 cm wide and 2 cm thick; pileal surface cream to
orange brown when fresh, olivaceous buff to deep olive when dry, azonate, glabrous;
margin white when fresh, becoming deep olive when dry; pore surface white to cream
when fresh, becoming cream to buff when dry; pores irregular to angular, 1–3 per
mm, mostly 1 per mm; dissepiments thin, entire to slightly lacerate; context white
when fresh, up to 0.8 cm thick; tubes concolorous with the pore surface, up to 1.2 mm
long. Hyphal system dimitic; generative hyphae simple septate; skeletal hyphae IKI–,
CB–; tissues unchanged in KOH. Contextual generative hyphae seldom, hyaline,
thick-walled, simple septate, 4–8 µm in diam; contextual skeletal hyphae dominant,
hyaline, thick-walled with a narrow to wide lumen, rarely branched, flexuous,
interwoven, 4–10 µm in diam. Tramal generative hyphae dominant, hyaline, slightly
thick-walled to thick-walled, simple septate and numerous branched, 2–3.5 µm in
diam; tramal skeletal hyphae rarely, hyaline, thick-walled with a narrow to wide
lumen, rarely branched, flexuous, interwoven, 2–4 µm in diam. Cystidia and
cystidioles absent; basidia clavate, with a simple basal septum and four sterigmata,
35–58 × 9–11 µm; basidioles in shape similar to basidia, but distinctly shorter.
Basidiospores subglobose, hyaline, thick-walled, with obvious ridges, strongly
amyloid, CB+, (5.5–)5.8–7 × 5–6.5(–6.8) µm, L = 6.4 µm, W = 5.8 µm. Ridges of
spores blunt, up to 1 µm long.
Type of rot: White rot.
Material examined: CHINA: Xizang Autonomous Region (Tibet), Milin County,
Nanyigou Park, on fallen trunk of Picea, 16 Sep 2014, Cui 12078 (holotype, BJFC
016992); ibid, Linzhi County, Bayi, on fallen trunk of Picea, 16 Aug 2004, Yu 56
(paratype, IFP 000968); Milin County, Nanyigou Park, on fallen trunk of Picea, 16
Sep 2014, Cui 12079 (paratype, BJFC 016993).
Notes: Bondarzewia tibetica is found on Picea in Xizang Autonomous Region of
China. It is characterized by its cream to orange brown pileal surface, white to cream
pore surface, small pores, a dimitic hyphal system, and large basidiospores with blunt
spines. Bondarzewia dickinsii (Berk.) Jia J. Chen, B.K. Cui & Y.C. Dai, B.
occidentalis Jia J. Chen, B.K. Cui & Y.C. Dai and B. podocarpi Y.C. Dai & B.K. Cui
are morphologically similar to B. tibetica; they all produce similar pileal surface and
pore surface; however, B. dickinsii is different by producing a monomitic hyphal
system and sharp basidiospore spines; B. occidentalis is separated by its larger
basidiospores and pores; B. podocarpi is different by producing sharp basidiospore
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spines (Chen et al. 2016; Dai et al. 2010). Phylogenetically, all species of
Bondarzewia formed a monophyletic lineage belonging to Russulales (Fig. 136).
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Fig. 137 Bondarzewia tibetica (holotypes) a, b Basidiocarps c, d Basidiospores. Scale bars: a,
b = 1 cm, c = 7 µm, d = 2 µm.
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Fig. 138 Bondarzewia tibetica (holotype) a Basidia and basidioles b Hyphae from trama c
Hyphae from context. Scale bars: a–c = 10 µm.
Russulaceae
Within the Russulales order, members of the Russulaceae family display a large
diversity in sporophore morphology. Sporophores range from resupinate to agaricoid,
pleurotoid or sequestrate types, with hymenophores that can be poroid or lamellate
(Miller et al. 2006). The vast majority of the known species are mainly agaricoid and
belong to the genera Lactifluus (Pers.) Roussel, Lactarius Pers., Multi-furca Buyck &
V. Hofstetter and Russula Pers. (Buyck et al. 2008, Buyck et al. 2010). These genera
are all ectomycorrhizal and have representatives in Thailand. Next to these genera, the
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Russulaceae family also contains three mainly corticoid genera: Boidinia Stalpers &
Hjortstam, Gloeopeniophorella Rick and Pseudoxenasma K.H. Larss. & Hjortstam
(Larsson and Larsson 2003, Miller et al. 2006).
Lactifluus
The ectomycorrhizal genus Lactifluus is the smallest of the two milkcap genera
(Russulaceae). The genus is mainly distributed in the tropics and is well-represented
in Thailand (Le et al. 2007; Stubbe et al. 2010; Van de Putte et al. 2010; De Crop et al.
2014). In a recent study (De Crop et al. subm.), the genus is revised and four
subgenera are proposed: L. subg. Lactariopsis, L. subg. Rugati, L. subg. Gymnocarpi
and L. subg. Lactifluus. The two species from Thailand that are presented here belong
to L. subg. Lactariopsis and L. subg. Rugati. The phylogenetic tree is presented in
Fig.139.
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Fig. 139 Maximum likelihood tree of Lactifluus subg. Lactariopsis and L. subg. Rugati, based
on ITS-LSU sequence data. Maximum likelihood bootstrap values >70 are shown. New taxa
are in blue and species for which obtained sequences are based on type material have names
in bold.
347. Lactifluus armeniacus De Crop & Verbeken, sp. nov.
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MycoBank number: MB 815137, Facesoffungi number: FoF 02056, Figs 140,
141
Etymology: Referring to the apricot-coloured basidiocarps.
Holotype: MFLU E. De Crop 14–501
Diagnosis: A medium-sized, warm apricot-coloured species which is
microscopically characterized by septated lamprocystidia, low ornamented spores and
a lampropalisade as pileipellis structure, with small to medium-sized, thick-walled
hairs in the suprapellis and a thick layer of sphaerical cells in the subpellis.
Pileus 69–72 mm diam., planoconvex with central depression to slightly
infundibuliform; margin sometimes slightly striate, sometimes concentrically
wrinkled; edge rather irregular, sometimes crenulate or locally undulate; surface
chamois leather-like, locally wrinkled but smooth in the centre, pruinose, bright
orange(as 5B5/6, but more yellow), unicolourous. Lamellae adnate with decurrent
tooth to subdecurrent, distant (2L + 1l / cm – 4L + 3l / cm), bright orange to yellow
(4A3 to 4/5A4), very broad, rather thick and brittle, slightly intervenose; edge entire
and concolourous. Stipe 27–28 × 11–18 mm, cylindrical to slightly tapering
downwards, sometimes curved, centrally attached to pileus; surface very soft,
pruinose and finely striate, concolourous with pileus (bright orange 5B5/6 with a more
yellowish tinge). Context solid and quite firm, white, unchanging; taste sweet, mild;
smell not distinctive. Latex abundant, white, unchanging; taste sweet.
Basidiospores broadly ellipsoid, sometimes subglobose, sometimes ellipsoid,
6.4–7.7–9 × 5.1–6.2–6.7 µm (n = 20, Q = 1.11–1.24–1.41); ornamentation amyloid,
forming an almost complete reticulum, composed of very low warts connected by fine
ridges, up to 0.2 µm high; plage inamyloid. Basidia 4-spored, sometimes 2-spored,
59–71 × 8–9 µm, cylindric to subclavate, with refringent to slightly thickened walls;
content guttate to granular. Pleurolamprocystidia abundant, slightly emergent up to
17 µm, cylindrical, septate, 50–80 × 4–8 µm, with slightly thickened walls (<1 µm).
Pleuropseudocystidia very scarce, 7–9 µm, cylindrical, mostly collapsed at apex;
content
granular.
Lamellae-edge
sterile;
completely
composed
of
cheilolamprocystidia which are 41–45 × 4–7 µm, cylindrical, septate, thick-walled.
Hymenophoral trama cellular, with abundant lactifers and sphaerocytes. Pileipellis a
lampropalisade; elements of the suprapellis 28–64 × 3–5 µm, cylindrical, obtuse,
thick-walled; subpellis 132–174 µm thick, sphaerical cells 9–22 µm diam., with
thickened wall. Stipitipellis hymeniderm; elements of the suprapellis 15–26 × 5–11
µm, cylindrical to clavate, sometimes with strong congophilous content, thick-walled.
Material examined: THAILAND, Chiang Mai Province, Mae Taeng district,
Baan Tapa (22km marker along road 1095), N19°7'45" E98°46'1", alt. 766.8 m, on
soil in mixed forest, with Dipterocarpus sp., Castanopsis sp., Lithocarpus sp. and
Quercus sp., 31 July 2014, E. De Crop 14–501 (holotype in MFLU, isotype in
GENT).
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Fig. 140 a Lactifluus armeniacus (holotype) b Lactifluus ramipilosus (holotype)
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Fig. 141 Lactifluus armeniacus (holotype) a Section through pileipellis b Basidiospores c
pleuropseudocystidia d Pleurolamprocystidia e Marginal cells f Bsidia g Terminal elements of
the pileipellis. Scale bars: a–g = 10 µm.
348. Lactifluus ramipilosus Verbeken & De Crop,
sp. nov.
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MycoBank number: MB 815138, Facesoffungi number: FoF 02057, Figs 140,
142
Etymology: with branched (rami-) hairs (-pilosus), referring to the striking hairs
in the pileipellis structure.
Holotypus: MFLU E. De Crop 14–503
Diagnosis: A medium-sized, warm yellowish orange species which is
microscopically characterized by the very lowly and indistinctly ornamented spores,
the absence of true cystidia and ramified thick-walled hairs in the pileipellis structure.
Pileus 55 mm diam., convex to planoconvex with undeep depression in the
center; surface soft, chamois-leather like and pruinose, almost smooth but slightly
irregular, yellowish orange (5A3-4A4); margin entire, straight to slightly deflexed.
Stipe 25 × 17 mm, strongly tapering downwards; surface pale yellow (4A2), slightly
paler towards the lamellae, very finely fibrillose. Lamellae broadly adnate to
decurrent, up to 4 mm broad, medium thick, brittle, yellow (4A3). Context whitish
yellow. Latex not observed.
Spores 5.6–7.2–8.9(9.1) × 5.5–6.2–7.2(7.3) µm, Q = 1.03–1.16–1.32, broadly
ellipsoid, sometimes subglobose; ornamentation amyloid but very low and weakly
developed, composed of low and irregular warts that are often connected by very fine
ridges forming a partial reticulum; plage mostly not amyloid, but sometimes with a
very weak central amyloid spot. Basidia 4-spored, with some rare 2-spored basidia
present, 45–55 × 8–10 µm, subcylindrical to subclavate, with guttate contents. True
cystidia absent. Pleuropseudocystidia abundant, not emergent to slightly but distinctly
abundant, 6–8 µm diam., cylindric but often swollen at the apex, with rounded apex,
with needle-like to granular content. Hymenophoral trama mixed with some hyphae
present but especially abundant sphaerocytes of up to 25 µm diam., with abundant
lactifers. Subhymenium cellular. Lamellar edge sterile; marginal cells 15–28 × 6–8 µm,
subclavate to irregular, mostly hyaline, sometimes with refringent walls, sometimes
with slightly needle-like content. Pileipellis lamprotrichoderm-like, composed of a
layer of hyphae with 3-5 µm diam., which are mainly horizontally arranged and often
terminating in remarkable thick-walled hairs which are pericline to oblique; hairs
thick-walled, 35–125 × 3–5 µm, often branched, sometimes septate, sometimes
tapering near paex, sometimes with rounded apex.
Material examined: THAILAND, Chiang Mai Province, Mae Taeng district,
Baan Tapa (22km marker along road 1095), N19°8'0" E98°46'15", alt. 829.6 m, on
soil in mixed forest, with Dipterocarpus sp., Castanopsis sp., Lithocarpus sp. and
Quercus sp., 31 July 2014, E. De Crop 14–503 (holotype in MFLU, isotype in
GENT)
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Fig. 142 Lactifluus ramipilosus (holotype) a Section through pileipellis b Marginal cells c
Basidiospores d Basidia e Pleuropseudocystidia f Terminal elements of the pileipellis. Scale
bars: a–f = 10 µm.
Russula
Russula is a genus of high species diversity with a comprehensive wide
distribution from frigid to tropical forests (Kundsen and Borgen 1982; Singer 1986;
Buyck 1989; Buyck et al. 1996; Miller et al. 2012). Russula is evidenced from ITS,
nLSU and rpb2 to be a monophyletic genus (Buyck et al. 2008, 2010), but it contains
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stipitate epigeous, hypogeous, and pleurotoid-formed fruiting bodies (Buyck and
Hoyak 1999; Miller et al. 2001; Larsson and Larsson 2003; Lebel and Tonkin 2007).
Nine subgenera have been introduced in Russula based on morphological
characteristics, such as taste of fruiting bodies, colour of spore print, shape of
pileipellis hyphal extremities, existence of lamellulae, dermatocystidia and primordial
hyphae (Romagnesi 1967, 1985, 1987), and phylogenetic data is needed in
classification (Eberhardt 2002; Li and Wen 2009; Li 2014, Li et al. 2015a). A total of
22 Russula taxa have been described from China and the adjacent Himalayan
Mountain in recent years (Das et al. 2005, 2006a, 2006b, 2010, 2013, 2014; Wang et
al. 2009b; Li et al. 2011, 2012, 2013a, 2013b, 2015a, 2015b). Two taxa are newly
described from Tibet Plateau based on morphological characters and phylogenetic
analyses. The phylogenetic tree for Russula is presented in Fig. 143.
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Fig. 143 Phylogram generated from maximum parsimony (PAUP* v.4.01) analysis based on
ITS sequence data of Russula. Sequences used in this study have been sampled from previous
studies to represent the major phylogenetic clades of Russula (Eberhardt 2002; Miller and
Buyck 2002). Maximum parsimony bootstrap support values above 50% and Bayesian
posterior probabilities greater than 0.9 are indicated above or below the nodes (BS/PP), new
taxa are shown in blue. Holotype are shown in bold and blue.
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349. Russula amethystina subsp. tengii G.J. Li, H.A. Wen & R.L. Zhao, subsp. nov.
Fungal Names number: FN 570231, Facesoffungi number: FoF 02058, Fig. 144.
Etymology: named after Prof. S.C. Teng, in honor of his contribution to the
taxonomy of Russula.
Holotype: HMAS 253336
Basidiomata small- to medium-sized. Pileus 43–52 mm in diam., hemispheric
when young, plano-convex, expanding to applanate when mature, rarely center
slightly depressed with age, not striate, sometimes cracked, slightly viscid when wet,
peeling 1/4–1/3 from the edge, lilac to vinous tinged with intermixed with brown
vinous tinged with Brownish Vinaceous (XXXIX5′′′b), Light Russet-Vinaceous
(XXXIX1′′′d) to Haematite Red (XXXIX5′′m), reddish tinge of Pompeian Red
(XIII3′i) in center, Dark Vinaceous-Brown (XXXIX5′′′k) and Vinaceous-Brown
(XXXIX5′′′i) intermixed with Pale Brownish Drab (XLV5′′′′d) towards the margin
when dry. Lamellae slightly subfree, 2–5 mm in height, 13–16 pieces per cm in the
edge, rarely forked near the stipe or in the middle, interveined, with ocherous,
yellowish tinged with Light Ochraceous-Salmon (XV13′b), Light Ochraceous-Buff
(XV15′d) to Ochraceous-Buff (XV15′b); lamellulae absent. Stipe 5.5–6.8 × 0.9–1.5
cm, subcylindrical, surface dry, rugulose longitudinally, dull, without annulus,
slightly attenuate upwards, White (LIII), a tinge of Pale Yellow–Orange (III15f) when
injured and dry, stuffed first, becoming hollow when old. Context up to 1–2 mm at the
center of the pileus, White (LIII), fragile, with iodoform Odour; taste mild. Spore
print Ocher (Romagnesi IIId–IVa).
Basidiospores [100/10/8] 7.4–8.7 (–9.2) × 6.2–7.5 (–8) µm, Q = (1.06–)
1.10–1.28 (–1.34), (Q = 1.20 ± 0.06), hyaline, mostly broadly ellipsoid, rarely
subglobose or ellipsoid; ornamentation cristulate to subreticulate, composed of
amyloid warts that linked as small crests and ridges, forming a nearly complete to
complete network, rarely intermixed with isolated verrucae, warts 0.5–0.8 µm in
height; suprahilar area distinctly amyloid. Basidia 30–40 × 7–10 µm, 4-spored,
sterigmata 3–6 µm long, hyaline, sometimes yellowish in KOH, subclavate to clavate,
rarely cylindrical. Pleuroystidia scattered, 55–100 × 8–13 µm, distinctly projecting
20–60 µm beyond the basidia, subfusoid to subcylindrical, sometimes clavate to
subclavate, apex obtuse, thin–walled, contents rare, granular to crystal, weakly gray in
sulphovanillin (SV). Cheilocystidia not observed; lamellar edge sterile. Subhymenium
a cellular layer 20–35 µm thick composed of inflated cells 7–13 µm in diam., hyaline,
sometimes pale yellowish in KOH. Pileipellis composed of epipellis and subpellis;
epipellis a trichoderm 125–150 µm thick, composed of thin-walled, cylindrical
hyaline hyphae 3–6 µm wide; primordial hyphae 4–7 µm wide, with
heteromorphous-opalescent inclusions and acid-resistant incrustations, septate, clavate
to cylindrical, apex obtuse; subpellis a cutis 100–120 µm thick, composed of
gelatinized, interweaved hyaline hyphae 2–6 µm wide, pileocystidia not observed.
Stipitipellis a cutis, composed of filamentous hyphae 3–6 µm in diam., interweaved
with inflated cells 10–15 µm in diam., hyaline, some hyphae pale yellow in KOH;
caulocystidia absent. Clamp connections and lacticiferous hyphae absent from all
tissues.
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Habit and habitat. Single or small groups in coniferous forest (dominated by e.g.
Pinus densata var. pygmaea, P. yunnanensis, Picea likiangensis var. likiangensis and
P. likiangensis var. linzhiensis) at 2000–3500 m altitude. Distribution. China (Xizang
and Yunnan). Season. July and August.
Material examined: CHINA, Yunnan Province, Lijiang City, Yulong County,
Lijiang Alpine Botanic Garden, N27°05' E100°10', elevation 3447 m., 17 July 2014,
collector Guojie Li and Yaning Wang, 14252 (HMAS 253336, holotype); Ibid.,
elevation 3258 m., collector Guojie Li and Yun Yu, 14075 (HMAS 271033); Ibid.,
elevation 3471 m., collector Guojie Li and Mingjun Zhao, 14188 (HMAS 271034);
Ibid., elevation 3274 m., collector Guojie Li and Shuhua Jiang, 14088 (HMAS
271161); Ibid., collector Guojie Li and Yunlong Li, 14187 (HMAS 271048);
Chuxiong City, Nanhua County, Zixishan Forest Park, N25°01' E101°32', elevation
2134 m., 20 August 2013, collector Weilai Lu, Tiezheng Wei and Zhenping Yang,
354 (HMAS 252864); Xizang Autonomous Region, Nyingchi Prefecture, Mainling
County, roadside of National Road 318 to Nang County, N29°12' E94°11', elevation
2994 m., 12 August 2013, collector Tiezheng Wei, Xiaoyong Liu, Jianyun Zhuang
and Tian zhou Li, 3701 (HMAS 253216); Ibid., 3698 (HMAS 253241).
Notes: The combination of a violet-tinged pileus without olive green, bluish gray
or entirely yellow tinges, deep yellow spore print, pileipellis with primordial hyphae,
absence of pileocystidia, and coniferous habitat assign this taxon into Russula
amethystina Quél. of Russula subgenus Incrustatula Romagn, Russula section
Amethystinae Romagn. (Romagnesi 1987). The phylogenetic result (Fig. 143) also
supports the placement (BS 97% and PP 1.00). These suggest R. amethystina subsp.
tengii is very closely related to R. amethystina subsp. amethystina, which however,
has easily distinguishable higher basidiospore ornamentation composed of mostly
isolated. (Romagnesi 1967; Sarnari 2005; Kränzlin 2005). Because the basidiospore
of R. amethystina subsp. tengii is almost the same as that of R. turci Bres., the
morphological distinction between the two closely related species is becoming blurred.
Yet it is clear in the distinction among R. amethystina, R. turci, and R. roseipes Secr.
ex Bres. phylogenetic analyses. The high phylogenetic BS/PP values and the only
obvious morphological difference effectively supported that R. amethystina subsp.
tengii is a subspecies of R. amethystine (Fig. 143).
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Fig. 144 Basidiocarps and microcharacters of Russula amethystina subsp. tengii (holotype)
a Basidiocarps b Basidiospores c Basidia d Pleurocystidia e Epipellis. Scale bars: a = 1 cm,
b–e = 10 µm.
350. Russula wangii G.J. Li, H.A. Wen & R.L. Zhao, sp. nov.
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Fungal Names number: FN 570232, Facesoffungi number: FoF 02059, Fig. 145.
Etymology: named after Prof. Y.C. Wang, in honor of his contribution to the
study of fungi from China.
Holotype: HMAS 268809
Basidiomata small- to medium-sized. Pileus 38–56 mm in diam., hemispheric
when young, plano-convex, expanding to applanate when mature, rarely center
slightly depressed with age, not striate, sometimes cracked, viscid when wet, peeling
1/3–1/2 from the edge, brownish vinous to violet tinged with Pecan Brown
(XXXVIII11′′i) to Cacao Brown (XXXVIII9′′i), intermixed with darker tinge of
Walnut Brown (XXXVIII9′′k) to Rood's Brown (XXXVIII11′′k) in center, sometimes
completely Dark Bull Bluish Violet (X57m), Prussian Red (XXVII5′′k) and Dark
Indian Red (XXVII3′′m) intermixed with Deep Cortinthian Red (XXVII3′′i) towards
the margin when dry. Lamellae slightly subfree, 2–5 mm in height, 13–17 pieces per
cm in the edge, not forked, interveined, with ocherous, yellowish tinged with Salmon
Buff (XIV11′d), Salmon Colour (XIV9′d) to Apricot Buff (XIV11′b); lamellulae
absent. Stipe 4.4–6.5 × 0.8–1.7 cm, subcylindrical, surface dry, rugulose
longitudinally, dull, without annulus, slightly attenuate upwards, White (LIII), a tinge
of Pale Yellow–Orange (III15f) when injured and dry, stuffed first, becoming hollow
when old. Context up to 3 mm at the center of the pileus, White (LIII), fragile, no
distinct Odour; taste acrid. Spore print Yellow (Romagnesi IVd–IVe).
Basidiospores [100/10/7] (6.3–) 6.8–8.2 × 7–8 (–8.5) µm, Q = (1.06–) 1.13–1.30
(–1.34), (Q = 1.38 ± 0.06), hyaline, broadly ellipsoid to ellipsoid, rarely subglobose;
ornamentation cristulate to subreticulate, composed of amyloid warts that linked as
small crests and ridges, forming a nearly complete network, often intermixed with
isolated verrucae, warts 0.5–1 µm in height; suprahilar area amyloid. Basidia 30–40 ×
8–10 µm, mostly with four sterigmata 4–7 µm long, hyaline, sometimes yellowish in
KOH, subclavate to clavate, rarely cylindrical. Pleuroystidia scattered, 60–80 × 8–13
µm, projecting 20–55 µm beyond the basidia, subfusoid to subcylindrical, sometimes
clavate to subclavate, apex obtuse, often with a moniliformto papillate appendage,
thin–walled, contents granular to crystal, blackish gray in SV. Cheilocystidia not
observed; lamellar edge sterile. Subhymenium a cellular layer 20–35 µm thick
composed of inflated cells 7–13 µm in diam., hyaline, sometimes pale yellowish in
KOH. Pileipellis composed of epipellis and subpellis; epipellis a trichoderm 125–150
µm thick, composed of thin-walled, diverticulate, cylindrical hyaline hyphae 3–6 µm
wide; pileocystidia 6–8 µm wide with refractive contents blackened in SV, abundant,
septate, diverticulate, clavate to cylindrical, apex obtuse, sometimes inflated; subpellis
a cutis 100–150 µm thick, composed of gelatinized, interweaved hyaline hyphae 2–6
µm wide. Stipitipellis a cutis, composed of filamentous hyphae 3–5 µm in diam.,
interweaved with inflated cells 15–25 µm in diam., hyaline, some hyphae yellowish to
pale ocher in KOH; caulocystidia absent. Clamp connections and lacticiferous hyphae
absent from all tissues.
Habit and habitat. Single or scattered in coniferous forest (dominated by e.g.
Pinus densata var. pygmaea, P. yunnanensis, Picea purpurea and P. likiangensis var.
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balfouriana) at 3000–4000 m altitude. Distribution. China (Qinghai and Sichuan).
Season. July and August.
Material examined: CHINA, Sichuan Province, Garzê Autonomous Prefecture,
Dawo County, Geka Township, Geka Village, N30°59' E101°08', elevation 3471 m.,
12 August 2013, collector Weilai Lu, Lan Jiang and Guojie Li, 13279 (HMAS 268809,
holotype); Ibid., 13278 (HMAS 268808); Zamtang County, N32°19' E100°59',
elevation 3930 m., 28 July 2013, collector Binbin Li, Xiaoying Li and Ruiheng Yang,
180 (HMAS 269580); Ngawa Tibetan Qiang Autonomous Prefecture, Ngawa County,
N32°53' E101°42', elevation 3457 m., 24 July 2013, collector Binbin Li, Xiaoying Li
and Ruiheng Yang, 48 (HMAS 269308); Qinghai Province, Golog Autonomous
Prefecture, Baima County, Hongjungou, N32°57' E100°42', elevation 3516 m., 26
July 2013, collector Binbin Li, Xiaoying Li and Ruiheng Yang, 197-1 (HMAS
269106); Ibid., 243 (HMAS 269398); Ibid., 383 (HMAS 269143).
Notes: The violet tinged pileus, acrid tasted context, yellow spore print,
sulphoaldehyde sensitive pileocystidia, diverticulate epicutis hyphal ends and
pileocystidia clearly place R. wangii within Russula section Urentinae Maire ss. str. of
Russula subgenus Insidiosula Romagn. Ten species, R. adulterina Secr., R. cristata
Romagn., R. cuprea J.E. Lange, R. cupreoaffinis Sarnari, R. cupreola Sarnari, R.
firmula Jul. Schäff., R. gigasperma Romagn. ex Romagn., R. juniperina Ubaldi, R.
subcristulata Romagn., R. transiens (Singer) Romagn. and R. urens Romell, have
been recognized in Russula section Urentinae. Russula adulterina differs in larger
basidospores 7.5–12 × 7–9.5 µm with higher ornamentations composed of isolated
warts up to 1.6–2 µm, and non-diverticulate pileocystidia (Romagnesi 1967). Russula
cristata can be distinguished from R. wangii in brownish grey staining context, lower
basidiospore ornamentations up to 0.75 µm, nonseptate and non-diverticulate
pileocystidia (Romagnesi 1967). Russula cuprea discriminates from R. wangii in
larger basidospores 8.5–12 × 6.7–8.5 µm with higher ornamentations composed of
isolated warts up to 1.5 µm (Romagnesi 1967; Sarnari 1998). Russula cupreoaffinis
differs in larger basidiospores variable pileus colors, larger basidiospores 7.7–10 ×
6.2–7.4 µm, and habitat of broad-leaved forest dominated by Quercus spp. (Sarnari
1998). Russula cupreola can be distinguished from R. wangii in longer and wider
basidia 42–64 × 10.5–14 µm, longer and wider pleurocystidia 60–100 × 9–16 µm, and
habitat of alpine dwarf shrubs associated with Salix herbacea and S. reticulata
(Sarnari 1998). Russula firmula discriminate from R. wangii in larger basidospores
8–10.5 × 6.8–8.4 µm with ornamentations composed of mostly isolated warts, and
non-diverticulate pileocystidia (Romagnesi 1967; Sarnari 1998). Russula gigasperma
is different from R. wangii in larger basidospores 10–12 × 8–10 µm with higher
ornamentations composed of isolated warts up to 1.4 µm, and habitat of hardwood
forest (Romagnesi 1967; Sarnari 1998). Russula juniperina discriminates from R.
wangii in brightly red pileus, larger basidiospores 8–11 × 7.2–9 µm, and habitat of
broad-leaved forest dominated by Quercus ilex or Q. pubescens (Sarnari 1998).
Russula subcristulata can be distinguished from R. wangii in in longer basidia 42–57
× 9–12 µm, longer and wider pleurocystidia 65–105 × 10–15.7 µm, nonseptate and
non-diverticulate pileocystidia (Romagnesi 1967). Russula transiens differs in larger
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basidiospores 7.5–10 × 6.7–10 µm with ornamentations up to 1.25 µm, and
non-diverticulate pileocystidia (Romagnesi 1967; Sarnari 1998). Russula urens
discriminate from R. wangii in large green to yellowish green tinged pileus up to 12
cm with strongly tuberculate-striated margin (Sarnari 1998). Russula olivina Ruots. &
Vauras from Russula section Laricinae Romagn. of Russula subgenus Tenellula
Romagn. and R. olivobrunnea Ruots. & Vauras from Russula section Integroidinae
Romagn. of Russula subgenus Polychromidia Romagn., cluster together with R.
wangii by support of BS 71% and PP 0.98 in phylogenetic tree. However, R. olivina
differs in larger basidospores 9–11.2 × 7.2–9.5 µm with higher ornamentations
composed of isolated warts up to 1.5 µm, longer and wider basidia 37–71 × 13–20 µm,
and longer and wider pleurocystidia 65–105 × 10–19 µm. Russula olivobrunnea can
be distinguished from R. wangii in larger basidospores 9–12.8 × 7.4–10.4 µm with
higher ornamentations composed of isolated warts up to 1.6 µm, longer and wider
basidia 37–58 × 9–15 µm, and longer, wider pleurocystidia 45–98 × 9–15.5 µm, and
non-diverticulate pileocystidia (Sarnari 2005).
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Fig. 145 Basidiocarps and microcharacters of Russula wangii (holotype) a Basidiocarps b
Basidiospores c Pleurocystidia d Basidia e Epipellis. Scale bars: a = 1 cm, b–e =10 µm.
Contributions to Neocallimastigomycota
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Neocallimastigales
Neocallimastigaceae
Neocallimastigomycota or anaerobic fungi represent a special group of
microorganisms inhabiting the digestive tract ecosystem of large mammalian
herbivores, including ruminants and non-ruminants. Anaerobic fungi release a broad
range of polysaccharide-degrading enzymes that, to date, are among the most
effective reported for the breakdown of plant material. Their active role in the
degradation of plant structural material has simulated considerable worldwide interest
both in terms of their place in fungal evolution and in their potential for industrial
exploitation. The phylogeny of the Neocallimastigomycota is illustrated in Figs 146,
147.
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Fig. 146 Molecular phylogeny generated by maximum likelihood analysis of ITS1 sequence
data from the Neocallimastigomycota. Representative species from all known eight genera
(indicated) are shown. Bootstrap values above 50% are indicated above each branch. Ex-types
(reference strains) are bolded and new isolates are indicated in blue.
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Fig. 147 Molecular phylogeny generated by maximum likelihood analysis of partial large
subunit (28S) ribosomal DNA sequence data from the Neocallimastigomycota. Bootstrap
values above 50% are indicated above each branch. New isolates are indicated in with a filled
shape.
Anaeromyces Breton et al.
The genus Anaeromyces was described using morphological characteristics by
Breton et al. (1991). Following isolation of Anaeromyces mucronatus from the rumen
of a cow. The type culture, Anaeromyces mucronatus (NR_111156.1) was obtained
from faeces of an American bison by Fliegerova et al. (2004). This group isolated a
number of different polycentric fungi belonging to the genera Orpinomyces and
Anaeromyces. These two genera are morphologically very similar, but Fliegerová et al.
(2004) used molecular methods (analysis of ITS1 fragments) in addition to
morphology to distinguish between them. From a descriptive perspective, the genus
Anaeromyces contains species of strictly anaerobic fungi, which are characterized by
a polycentric thallus, a polynuclear rhizomycelium of extensively branched hyphae,
zoosporangia that are sometimes mucronate with an acuminate apex and
uniflagellated zoospores. The rhizomycelium contains hyphae that can be tubular and
uniform or very wide, sometimes with constrictions. Sporangia can develop
intercalary as swellings in hypha or on sporangiophores. Some cultures fail to produce
mature sporangia and zoospores are rarely seen making classification by molecular
means the only sure way of assigning them to the genus.
351. Anaeromyces robustus O’Malley, Theodorou & Henske, sp. nov.
Index Fungorum number: IF 551676, Facesoffungi number: FoF 02060, Fig. 148
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Etymology: The specific epithet refers to the physical similarities between some
fungal zoosporangia and the tails of gray whales (Eschrichtius robustus) that travel
the Californian coast near where the fungus was isolated.
Holotype: Anaeromyces sp. S4 (O’Malley Lab, University of California, Santa
Barbara, NCBI Taxon ID: 1642509), JMRC:SF:12178.
An obligate anaerobic fungus isolated from the feces of a sheep (Ovisaries) at the
Santa Barabara Zoo (www.sbzoo.org) in 2013. The species is polycentric, producing
many zoosporangia per fungal thallus and therefore has an indeterminate (infinite) life
cycle. The fungus exhibits exogenous zoosporangial development (i.e., the encysted
zoospore does not retain the nucleus, which can migrate and by mitosis populate the
developing zoosporangium and the rhizomycelium). The zoosporangia are typically
club-shaped (≥ 50 µm long × 30 µm wide at their widest point). Occasionally they
fuse to form a shape like a whale’s tail. Upon maturity, each zoosporangium can
liberate ≥ 60 zoospores. The rhizomycelium does contain nuclei (as seen under DAPI
staining) and is highly branched and tapering. The zoosporangium is typically
attached to the rhizomycelium via one or several main rhizoids and is capable of
vegetative reproduction by fragmentation. Free swimming zoospores are typically
sphearical (ca. 10 µm diam.) and the species is characterized by the presence of
several posteriorly directed flagella that are in length up to 3–fold the diam. of the
zoospore. When swimming the flagella beat together as if they were a single
flagellum and thus propel the zoospore forward in a spiral or helical motion.
The reference culture is maintained by continual passage at the University of
California, Santa Barbara (S4, JMRC:SF:12178, holotype), and under
cryopreservation in repositories at the O’Malley Lab, University of California, Santa
Barbara, and University of Jena and Leibniz Institute for Natural Product Research
and Infection Biology, Jena, Germany (Jena Microbial Resource Collection JMRC:
SF: 012178 – ex-type). Fixed glutaraldehyde preparations are also kept by the
O’Malley Lab.
The internal transcribed spacer regions of the ribosomal RNA were amplified
with primers JB206/JB205 (Tuckwell et al. 2005). Phylogenetic analysis of the ITS1
regions of several cultured anaerobic fungal specimens spanning all 8 known genera,
firmly place S4 within Anaeromyces as a distinct, previously unclassified species
comparable in age to the type culture A. mucronatus JF1 (Fig. 146). The partial 28s
rRNA sequence of A. robustus, however, appears as a unique outgroup, perhaps due
to its incompleteness (Fig. 147). The ~72 Mbp genome has been sequenced by the US
Department of Energy’s Joint Genome Institute (JGI). The genome will be made
available
at
Mycocosm
in
2016
(http://genome.jgi.doe.gov/Anasp1/Anasp1.home.html) and aid in the discovery of
novel biomass degrading enzymes that may be engineered or heterologously
expressed for the production of lignocellulosic biofuels and other value added
chemicals. Furthermore, the genome will enable future –OMICs based
characterization of these organisms, including insight into their unique organelles and
biomass-degrading enzyme complexes.
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Fig. 148 Aneromyces robustus (holotype) a Multiple sporangia of A. robustus displaying a
range of morphologies b A whale-tale shaped sporangia, which inspired the name of this
species, with a single zoospore c A zoospore with multiple flagella visible d Navajo-Churro
sheep host from which the species was isolated e Multiple sporangia demonstrating club-like
morphology, with several sharing the same mycelial structure.
Neocallimastix Vávra & Joyon ex I.B. Heath
The genus Neocallimastix was described by Vavra and Joyon (1912). At that
time, the propensity for nutritional microbiologists to work with rumen fluid and
discard rumen solids meant that the solids-associated vegetative stage of the fungus
was not recognized. The zoospores of the fungus evident in rumen fluid were
therefore mistakenly identified as polyflagellated protozoans. The correct
identification of these zoospores by Orpin’s pioneering studies showed that the
‘flagellates’ were liberated from a benthic, vegetative stage of a ‘chytrid-like’ fungus
(Orpin 1975). Soon after his initial observations, chitin was identified in the fungal
cell walls and by 1989 a new classification had emerged to accommodate these
obligately anaerobic (oxygen intolerant) fungi (Orpin 1977a, b; Barr 1989). Anaerobic
fungi from this genera are among the most studied of all the anaerobic fungi.
Numerous isolates have been obtained and at least three species, N. frontalis, N.
patriciarum and N. hurleyensis have been classified (Heath et al. 1983; Orpin and
Munn 1986; Webb and Theodorou 1991). The original classification of these species
used classical taxonomy whereby morphological characteristics were used to identify
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the genus and zoospore ultrastructure was used for the species recognition. Brookman
et al. (2000) under took a molecular characterization of the gut fungi based on
ribosomal ITS1 and 18S rRNA. Their analysis revealed that N. hurleyensis and N.
frontalis were very similar, but that they differed from N. patriciarum. From a
morphological perspective the genus Neocallimastix contains species of strictly
anaerobic fungi characterized by a monocentricthallus, which consists of a network of
branched, tapering rhizoids devoid of nuclei. The zoosporangia are variable, often
oval or sphaerical in shape supported on a sporangiophore developed from one or
more main rhizoids. As the life cycle of a monocentric fungus is determinate (finite),
each thallus (the rhizoidal network) supports just one zoosporangium. Zoospores are
uninucleate, and either monoflagellated or often polyflagellated.
352. Neocallimastix californiae O’Malley, Theodorou & Solomon, sp. nov.
Index Fungorum number: IF 551675, Facesoffungi number: FoF 02061, Fig. 149
Etymology: The specific epithet refers to the state of California where the fungus
was isolated.
Holotype: Neocallimastix sp. G1 (O’Malley Lab, University of California, Santa
Barbara, NCBI Taxon ID: 1550276), JMRC:SF:12176.
An obligate anaerobic fungus isolated from the feaces of a goat (Capra
aegagrushircus) housed at the Santa Barbara Zoo (www.sbzoo.org) in 2013. The
species is monocentric and has a determinate (finite) life cycle. The fungus exhibits
endogenous zoosporangial development (i.e., the encysted zoospore retains the
nucleus). The encysted zoospore germinates to form a rhizoidal system and a single
typically sphaerical zoosporangium (≥ 120 µm diam.) that on maturity liberates ≥ 100
zoospores. The rhizoidal system is devoid of nuclei (as seen under DAPI staining) and
is highly branched and tapering. The zoosporangium is typically attached to the
rhizoidal system via one main rhizoid or sporangiophore. A septum is often visible in
mature zoosporangia, separating the zoosporangium from the sporangiophore. Free
swimming zoospores are typically sphaerical (ca. 10 µm diam.) and the species is
characterized by the presence of ca. 16 or more posteriorly directed flagella that are in
length up to 3–fold the diam. of the zoospore. When swimming the flagella beat
together as if they were a single flagellum and thus propel the zoospore forward in a
spiral or helical motion.
The reference culture is maintained by continual passage at the University of
California, Santa Barbara (G1, JMRC:SF:12176, holotype), and under
cryopreservation in repositories at the O’Malley Lab, University of California, Santa
Barbara, and University of Jena and Leibniz Institute for Natural Product Research
and Infection Biology, Jena, Germany (Jena Microbial Resource Collection JMRC:
SF: 012176 – ex-type). Fixed glutaraldehyde preparations are also kept by the
O’Malley Lab.
The internal transcribed spacer regions of the ribosomal RNA were amplified
with primers JB206/JB205 (Tuckwell et al. 2005). Phylogenetic analysis of the ITS1
regions of several cultured anaerobic fungal specimens spanning all eight known
genera and partial 28s reads, firmly place G1 in the genus Neocallimastix as a distinct,
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previously unclassified sister species to established cultures such as N. frontalis (Figs
146, 147). The ~190 Mbp genome has also been sequenced by the US Department of
Energy’s Joint Genome Institute (JGI) to reveal that G1 is a polyploid organism. The
genome
will
be
made
available
at
Mycocosm
in
2016
(http://genome.jgi.doe.gov/programs/fungi/index.jsf).
Fig. 149 Neocallimastix californiae (holotype) a Sphearical zoospores with multiple flagella
which are splayed out b Multiple sporangia, demonstrating the predominantly sphearical to
ovoid structure c Goat host from which N. californiae was isolated d Large sphearical
sporangia that is characteristic of this species.
Piromyces J.J. Gold et al.
Monoflagellated protozoans found in the rumen were assigned to the genus
Piromonas (Liebetanz 1910; Braune 1913). Orpin concluded that these flagellated
cells were in fact zoospores of anaerobic fungi (Orpin 1977a). Orpin retained the
generic name on the assumption that the fungi he isolated from the sheep rumen were
the same as Liebetanz’s ‘protozoans’ (Liebetanz 1910). However, Gold et al. (1988)
questioned this assumption because Liebetanz’s isolates were anteriorly flagellated,
obtained nutrition by phagocytosis and divided by binary fission, whereas Orpin’s
isolates were posteriorly flagellated, rhizoid producing saprobes that did not undergo
binary fission. For these reasons, and to stress fungal affinity, Piromonas was
renamed Piromyces (Gold et al. 1988). Piromyces appears the most heterogeneous
genus among anaerobic fungi, covering up to eight species. Species of Piromyces
isolated to date include P. communis, P. mae, P. dumbonica, P. rhizinflata, P. minutus,
P. spiralis, P. citronii, P. polycephalus and P. cryptodigmaticus (Gold et al. 1988; Li
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et al. 1990; Breton et al. 1991; Ho et al. 1993a, 1993b; Gaillard-Martinie et al. 1995;
Chen et al. 2002; Fliegerová et al. 2010). While some of these species appear to have
morphologically distinct characteristics, relationships with each other and indeed with
other gut fungal genera remains unclear. Just one named but uncultured species (P.
cryptodigmaticus GQ850355.1, GQ850368.1, and GQ850318.1) has been categorized
according to their molecular characteristics (Fliegerová et al. 2010). Piromyces sp. E2
Teunissen et al. (1991) has been sequenced by the JGI and sequence data is available
on request. From a morphological perspective the genus Piromyces contains species
of strictly anaerobic fungi characterized by a monocentricthallus, which consists of a
network of branched, tapering rhizoids devoid of nuclei. The zoosporangia are
variable, sphearical, oval or club-shaped and are supported by a sporangiophore,
which develops from one or more rhizoids. As the life cycle of the monocentric fungi
is determinate (finite), each thallus (the rhizoidal network) supports just one
zoosporangium. Zoospores are uninucleate, sometimes bi- or quadri-flagellate
(Gruninger et al. 2014). The phylogenetic relatedness of the rhizoidal genera with
monoflagellated zoospores (Piromyces and Anaeromyces) is unclear and as observed
by Brookman et al. (2000), it seems likely that the genus Piromyces is polyphyletic
and in need of reappraisal.
353. Piromyces finnis O’Malley, Haitjema & Gilmore, sp. nov.
Index Fungorum number: IF 551677, Facesoffungi number: FoF 02062, Fig. 150
Etymology:”‘Piromyces of Finn”/”Finn’s Piromyces.’”The specific epithet refers
to the animal host, a horse named “Huckleberry Finn”, from which the fungus was
isolated.
Holotype: Piromyces sp. finn (O’Malley Lab, University of California, Santa
Barbara, NCBI Taxon ID: 1577477), JMRC:SF:12177.
An obligate anaerobic fungus isolated in 2011 at MIT from the feaces of the
award-wining show jumping horse Huckleberry Finn, owned by Susan Huyett of
Concord, MA. The species is monocentric and has a determinate (finite) life cycle.
The fungus exhibits endogenous zoosporangial development (i.e., the encysted
zoospore retains the nucleus). The encysted zoospore geminates to form a rhizoidal
system and a single oval or club shaped zoosporangium (≥ 100 µm long and 30–60
µm wide), which on maturity liberates ≥ 100 zoospores. The rhizoidal system is
devoid of nuclei (as seen under DAPI staining) and is highly branched and tapering.
The zoosporangium is typically attached to the rhizoidal system via one main rhizoid
or sporangiophore. A septum is often visible in mature zoosporangia, separating the
zoosporangium from the sporangiophore. Free swimming zoospores are typically
sphearical (ca. 10 µm diam.) and the species is characterized by the presence of a
single posteriorly directed flagella that is in length up to 3–fold the diam. of the
zoospore. When swimming the flagella beats posteriorly and thus propel the zoospore
forward in a spiral or helical motion.
The reference culture is maintained by continually passage at the University of
California, Santa Barbara (JMRC:SF:12177, holotype), and under cryopreservation in
repositories at the University of Jena and Leibniz Institute for Natural Product
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Research and Infection Biology, Jena, Germany (Jena Microbial Resource Collection
JMRC:SF:012177, ex-type). Fixed glutaraldehyde preparations are also kept by the
O’Malley Lab.
The internal transcribed spacer regions of the ribosomal RNA were amplified
with primers JB206/JB205 (Tuckwell et al. 2005). Phylogenetic analysis of the ITS1
regions of several cultured anaerobic fungal specimens spanning all eight known
genera and partial 28s reads, firmly place Finn within the Piromyces as a distinct,
previously unclassified species (Figs 146, 147). The ~56 Mbp genome has been
sequenced by the US Department of Energy’s Joint Genome Institute (JGI). The
genome
will
be
available
at
Mycocosm
in
2016
(http://genome.jgi.doe.gov/Pirfi3/Pirfi3.home.html).
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Fig. 150 Piromyces finnis (holotype) a Multiple sporangia of P. finnis exhibiting a range of
morphological features from club-like to ovoid b A group of young sporangia, not much
larger than zoospores beginning to form c Mature zoosporangia d Several zoospores of P.
finnis.
Contribution to Oomycota
The Oomycota are a highly diverse group of heterotrophic fungal-like eukaryotes
that are placed within the kingdom Straminipila, in the supergroup SAR (Adl et al.
2012). The major components of their cell walls are cellulose and β-1,3-glucans and
unlike fungal cell walls, only small amounts of chitin are present in some species
(Kamoun 2003, Rossman and Palm 2006). They reproduce asexually by heterokont
biflagellate zoospores (Hardham 2009) and when sexuality is present, by forming in
most cases oogonia and antheridia that mate, producing thick-walled oospores
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(Judelson 2009). They are cosmopolitan and ubiquitous, playing key roles in a wide
range of ecosystems as saprotrophs and parasites of a variety of host organisms such
as algae, oomycetes, fungi, plants, invertebrates and vertebrates (Marano et al. 2014).
They were informally classified into two lineages or “galaxies”, the
“peronosporaleans” and the “saprolegnialeans” until recently when Beakes et al.
(2014) have designated these lineages as classes, the Peronosporomycetes and
Saprolegniomycetes in the phylum Oomycota.
Peronosporales
Pythiaceae
Phytophthora de Bary
Phytophthora includes mainly ecologically and economically important plant
pathogens (e.g. Kroon et al. 2004; Brasier et al. 2005; Balci et al. 2007), but also
species that have not been yet associated with disease (Hansen et al. 2012) and that
are abundantly distributed in forested streams (Reeser et al. 2011). The genus is
currently subdivided into 10 well-recognized ITS clades (Kroon et al. 2012), plus the
novel lineage represented by P. stricta (Yang et al. 2014a). Multi-gene phylogenies
have shown that most of the 10 clades are monophyletic, except the Clades 4 and 9
(Blair et al. 2008). Clade 9 is the most rapidly expanding, with most of its species
recently described (Hong et al. 2010, 2012; Naher et al. 2011; Rea et al. 2011; Yang
and Hong, 2013; Yang et al. 2014a, b). Members of this clade generally produce
non-papillate and non-caducous zoosporangia. A well-defined subclade of species
within Clade 9 have a relatively high-temperature optima, ca. 30–32°C, and are able
to tolerate up to 40°C (Yang et al. 2014a). In this contribution, we describe two new
species for the Phytophthora ITS Clade 9, which both fall into this high-temperature
optima subclade (Fig. 151).
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Fig. 151 Phylogram generated from Maximum likelihood (ML) analysis (PhyML 3.1,
Guindon & Gascuel 2003) based on entire ITS rDNA sequences showing the phylogenetic
placement of Phytophthora rhizophorae and P. estuarina within Phytophthora Clade 9. ML
bootstrap support values < 50% are marked with (-). Clades that do not appear in the Bayesian
analysis are indicated with a zero. Bayesian posterior probability values (MrBayes 3.2,
Ronquist et al. 2012) > 0.50 are labelled numerically. Scale bar indicates the average number
of substitutions per site. New taxa are in blue and species for which obtained sequences are
based on type material have names in bold.
354. Phytophthora estuarina Marano, A.L. Jesus & Pires-Zottar., sp. nov.
Index Fungorum number: IF 551608, Facesoffungi number: FoF 01275, Fig. 152
Etymology: “estuarina” refers to the estuarine habitat in where this species was
isolated.
Holotype: SP 466380
Mycelium well-developed on PYGs, aerial mycelium scanty, hyaline, branched,
aseptate, hyphae 3.75–5 µm thick (av. 4.85 µm); hyphal swellings sphaerical, globose,
tubular to irregular. Zoosporangiophores undifferentiated of the vegetative hyphae,
long, simple or sympodially branched, bearing one terminal zoosporangium.
Zoosporangia produced abundantly in water cultures, non-caducous, semipapillate or
apapillate, ovoid to obpyriform, 55–83 × 43–63 µm (av. 77 × 54 µm), internally
proliferating in both a nested and extended way; secondary lateral zoosporangia
regularly formed; transparent lens-shaped plug material prior to zoospore
differentiation; wall rough after zoospore discharge; basal-plug present. Zoospores
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formed inside the zoosporangium and discharged by an elongate, vase-shaped, and
semi-persistent vesicle, 33–80 µm long when expanded, through which zoospores
swim away; encysted zoospores 7.5–12.5 µm diam. (av. 10.1 µm). The vesicle shrinks
completely in length and width in up to 1 hour after zoospore release.
Chlamydospores and sexual structures not observed. Gametangia not produced in
single culture or when paired with tester strains of P. capsici A1 (CBS 111334) and
A2 (CBS 370.72). Auto-sterile when the isolates were paired with each other. Radial
growth rates on PYGs (photoperiod: 12 h) at near the optimum temperature (30°C) =
12 ± 1 mm/d (n = 10); at near the maximum temperature (35°C) = 2 ± 1 mm/d (n =
10); no growth during five days at 40ºC and even after subsequent incubation at room
temperature (~20°C).
Culture characteristics: colonies cottony, with scanty aerial mycelium and no
defined growth pattern on PYGs.
Material examined: BRAZIL, São Paulo, Cananéia, “Parque Estadual da Ilha do
Cardoso” (PEIC), 25°03’05’’–25°18’18’’S; 47°53’48’’–48°05’42’’W, Perequê river
(salinity 1.3%), from leaves of Laguncularia racemosa onto PYGs medium, 27 Feb
2013, A.L. Jesus, C.L.A. Pires-Zottarelli & A.V. Marano (SP 466380, holotype),
ex-types CCIBt 4157, MMBF 14/15; Ibid., permanent shallow lagoon (salinity 2.8%),
from leaves of Rhizophora mangle, on Sorghum sp. seeds, 30 Aug 2012 A.L. Jesus,
C.L.A. Pires-Zottarelli & A.V. Marano (SP 466372, paratype), ex-paratypes CCIBt
4116, MMBF 06/15.
Notes: The isolates of P. rhizophorae and P. estuarina were recovered from
mangrove swamps, which exhibited salinity concentrations between 0.8–2.8% and,
therefore, the habitat of the ITS Clade 9 members is expanded to include estuaries.
Both P. rhizophorae and P. estuarina appear as well-delimited species and along with
ten other species, they consistently form a high temperature-tolerant subclade within
Clade 9, supported by strong bootstrap (100%) in our ITS phylogeny (Fig. 151).
Phytophthora rhizophorae is phylogenetically related with P. virginiana and P.
parsiana, while P. estuarina appear as closely related to P. macilentosa and P.
irrigata in our ITS phylogeny. The two new species share the presence of ovoid to
obpyriform, apapillate and non-caducous zoosporangia, which are terminal and
internally proliferating in both a nested and extended way (Figs 152 and 153). These
characteristics appear to be common to most members of Clade 9. Phytophthora
estuarina has additionally semi-papillate zoosporangia, a characteristic that is present
in a few species from this clade, such as P. constricta (Rea et al. 2011). During
zoospore discharge, it develops an elongate and semi-persistent vesicle, which
completely retracts in length and width in up to 1 h after zoospore release and
acquires a characteristic morphology after shrinkage (Fig. 153). The zoosporangia
have wrinkled walls after zoospore release and the shrunken vesicle remains
constricted at the apex of the zoosporangium. This process of vesicle development is
peculiar and has not been previously reported for Phytophthora species.
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Fig. 152 Phytophthora estuarina (holotype) a, b Zoospore differentiation inside the
zoosporangium and discharge of zoospores through an elongate semi-persistent vesicle After
shrinkage, the vesicle acquires a characteristic morphology (arrows) c Secondary lateral
zoosporangium and empty zoosporangium with characteristic rough walls after zoospore
discharge d Internal proliferation of the zoosporangium e Hyphal swellings f Colony with
scanty aerial mycelium and no defined growth pattern onto PYGs culture medium (CCIBt
4116). Scale bars: a–e = 10 µm.
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355. Phytophthora rhizophorae Pires-Zottar., A.L. Jesus & Marano, sp. nov.
Index Fungorum number: IF 551607, Facesoffungi number: FoF 01274, Fig. 153
Etymology: “rhizophorae” refers to Rhizophora mangle, the substrate from
where this species was isolated.
Holotype: SP 466375
Mycelium well-developed on PYGs, aerial mycelium scanty, hyaline, branched,
non-septate, hyphae 3.5–6.3 µm thick (av. 5.1 µm); hyphal swellings sphearical,
globose, tubular, obpyriform to irregular. Zoosporangiophores undifferentiated from
the vegetative hyphae, long, simple, bearing one terminal zoosporangium.
Zoosporangia internally proliferating, ovoid to obpyriphorm, non-papillate to
semi-papillate, non-deciduous, 35–58 × 20–45 µm (av. 45–32 µm); basal-plug
present. Zoospores formed inside the zoosporangia and discharged by a globose
vesicle; encysted zoospores 7.5–12.5 µm diam. (av. 9.3 µm). Chlamydospores and
sexual structures absent. Gametangia not produced in single culture or when paired
with tester strains of P. capsici A1 (CBS 111334) and A2 (CBS 370.72). Auto-sterile
when the isolates were paired with each other. Radial growth rates on PYGs
(photoperiod: 12 hs) at near the optimum temperature (30°C) = 14 ± 2 mm/d (n = 10);
at near the maximum temperature (35°C) = 7 ± 2 mm/d (n = 10); no growth was
observed during five days at 40ºC but the growth was reactivated after subsequent
incubation at room temperature (~20°C).
Culture characteristics: colonies petaloid on PYGs.
Material examined: BRAZIL, São Paulo, Cananéia, “Parque Estadual da Ilha do
Cardoso” (PEIC), 25°03’05’’–25°18’18’’S; 47°53’48’’–48°05’42’’W, Perequê river
(salinity 0.8%), from leaves of Rhizophora mangle, on Sorghum sp. seeds, 30 Aug
2012, A.L. Jesus, C.L.A. Pires-Zottarelli & A.V. Marano (SP 466375, holotype),
ex-holotypes CCIBt 4152, MMBF 09/15; Idem (SP 466374, paratype), ex-paratypes
CCIBt 4121, MMBF 08/15.
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7324
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Fig. 153 Phytophthora rhizophorae (holotype) a Apapillate zoosporangia during different
stages of zoospore differentiation b, c Empty zoosporangium with internal proliferation c
General aspect of the zoosporangiophore with both nested and extended internal proliferation
d, e Nested proliferation of the zoosporangium f–h Different morphologies of hyphal
swellings i Petaloid colony pattern onto PYGs culture medium (CCIBt 4121). Scale bars: a–i
= 10 µm.
Oomycota, incertae sedis
Salispina Marano, A.L. Jesus & Pires-Zottar., gen. nov.
In the last few years, increasing molecular evidence has shown that the genus
Halophytophthora (Peronosporales, Oomycota) as currently circumscribed is
polyphyletic, being composed by an assemblage of species that belong to related
peronosporalean genera, i.e. Salisapilia, Phytophthora and Phytopythium, and to yet
undescribed genera (Marano et al. 2016). Several phylogenetic studies have shown
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7347
that Halophytophthora spinosa falls into a new clade, commonly referred as “spinosa”
clade, which appears to represent a basal lineage phylogenetically more closely
related to Sapromyces elongatus (Rhipidiales) than to the “Halophytophthora sensu
stricto” clade (Nakagiri 2002; Nakagiri and Izumi 2005; Beakes et al. 2014; Marano
et al. 2014). Additional sequences of Rhipidiales are not available at GenBank to
putatively test this hypothesis and place the members of this clade into a higher-level
taxonomic category. Therefore, based on the phylogenetic analyses of the SSU (Fig.
154) and COI (Fig. 155) mtDNA regions we propose to establish Salispina gen. nov.
in an incertae sedis order for accommodating H. spinosa var. spinosa and H. spinosa
var. lobata, both elevated to species level, and the new species S. intermedia, until its
relatedness with other members of the Rhipidiales and Peronosporales could be tested
in a multi-gene phylogeny and its taxonomic placement confirmed.
Fig. 154 Phylogram generated from Bayesian inference analysis (MrBayes 3.2, Ronquist et al.
2012) based on SSU rDNA sequences showing the phylogenetic placement of Salispina gen.
nov. in a well-defined clade (indicated in bold). Maximum likelihood (ML) bootstrap support
values (PhyML 3.1, Guindon and Gascuel 2003) < 50% are marked with (-). Clades that do
not appear in the ML analysis are indicated with a zero. Bayesian posterior probability values
> 0.50 are labelled numerically. Scale bar indicates the average number of substitutions per
site. New taxa are in blue and ex-type strains are in bold.
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7354
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7356
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7358
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7361
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7374
Fig. 155 Phylogram generated from Bayesian inference analysis (MrBayes 3.2, Ronquist et al.
2012) based on cytochrome oxidase I (COI mtDNA) sequences showing the phylogenetic
placement of Salispina gen. nov. in a well-defined clade (indicated in bold). Maximum
likelihood (ML) bootstrap support values (PhyML 3.1, Guindon & Gascuel 2003) < 50% are
marked with (-). Clades that do not appear in the ML analysis are indicated with a zero.
Bayesian posterior probability values > 0.60 are labelled numerically. Scale bar indicates the
average number of substitutions per site. New taxa are in blue and ex-type strains in bold.
356. Salispina Marano, A.L. Jesus & Pires-Zottar., gen. nov.
Index Fungorum number: IF 551605, Facesoffungi number: FoF 01276
Etymology: salis (salt) indicating its presence under saline conditions (estuarine
and marine habitats), and spina (spine) because of the common presence of spines on
the zoosporangia.
Type species: Salispina intermedia A.L. Jesus, Pires-Zottar. & Marano
Holotypus: SP 466378
Colonies petaloid, with scanty aerial mycelium on agar culture media; mycelium
well-developed, hyaline, tortuous and highly branched, non-septate to few septate,
hyphae irregular. Zoosporangiophores undifferentiated from the vegetative hyphae,
long, simple, bearing one terminal zoosporangium. Zoosporangia with vacuolated
protoplasm, sphaerical, globose, obovate, obpyriform, or elongated, thick-walled,
from smooth to with spines showing variable degree of coverage on the zoosporangia;
basal-plug hyaline, slightly below the zoosporangia. Zoospore release takes place
through the formation of a persistent, short to long, dehiscence tube; vesicle absent.
Chlamydospores absent. Sexual reproduction unknown.
Notes: Salispina forms a well-defined lineage phylogenetically distant from the
Halophytophthora s.s. clade (Figs 154 and 155) and appears as closely related to
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7378
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7386
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7392
7393
7394
7395
7396
7397
7398
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7401
7402
7403
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7409
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Sapromyces elongatus (Fig. 154; Nakagiri 2002; Nakagiri and Izumi 2005; Beakes et
al. 2014; Marano et al. 2014). Fatty acid profiles evidenced that most members of the
Halophytophthora s.s. clade produce both arachidonic (ARA) and eicosapentaenoic
(EPA) acids while Salispina spinosa (H. spinosa var. spinosa) seems to be able to
produce only ARA (Pang et al. 2015). Fell and Master (1975) observed that
zoosporangial size and degree of spine coverage are nutritionally determined.
Zoosporangia formed on a rich substrate are larger and completely covered with
spines while those formed on poorer substrates are smaller and have only a few distal
spines or are even smooth. The dehiscence tube appears to be hydrotropic, being its
development conditioned by the presence of water (Fell and Master 1975).
357. Salispina intermedia A.L. Jesus, Pires-Zottar. & Marano, sp. nov.
Index Fungorum number: IF 551603, Facesoffungi number: FoF 01277, Fig. 156
Etymology: “intermedia” refers to the presence of intermediate morphological
features between S. spinosa and S. lobata.
Holotypus: SP 466378
Mycelium well-developed on PYGs, aerial mycelium scanty, hyaline, irregular,
branched, few septate, hyphae 2.5–10 µm thick. Zoosporangiophores undifferentiated
from the vegetative hyphae, long, simple, bearing one terminal zoosporangium,
6.25–12.5 µm (av. 9.8 µm). Zoosporangia of variable morphology, ranging from
obovate, obpyriform, globose to elongate, thick-walled, 33–197 × 25–183 µm (av. 86
× 62 µm); with vacuolated content; smooth to spiny; spines with variable degree of
coverage on the zoosporangium, from only one spine at the tip of the zoosporangium
to completely spiny; spines (5–)7.5–35 µm long. (av. 17 µm); basal plug hyaline,
2.5–7.5 µm thick (av. 5.5 µm). Zoospores discharged through a persistent tube, long
or short, 15–30 × (7.5–)12.5–15(–20) µm (av. 23 × 15 µm); vesicle absent; encysted
zoospores, 6.3–12.5 µm diam. (av. 9 µm), germination by one germ tube.
Chlamydospores absent. Sexual reproduction unknown.
Culture characteristics: colonies petaloid on PYGs; no growth on Sorghum sp.
(L.) seeds.
Material examined: BRAZIL, São Paulo, Cananéia, “Parque Estadual da Ilha do
Cardoso” (PEIC), 25°03’05’’–25°18’18’’S; 47°53’48’’–48°05’42’’W, Perequê river
(salinity 2.2%), from leaves of Rhizophora mangle, 8 Nov 2012, A.L. Jesus, C.L.A.
Pires-Zottarelli & A.V. Marano (SP 466378, holotype), ex-holotypes CCIBt 4155,
MMBF 12/15; Ibid., Perequê river (salinity 0.8–2.8%), from leaves of R. mangle and
Laguncularia racemosa, 30 Aug and 8 Nov 2012, 27 Feb and 5 Jun 2013, A.L. Jesus,
C.L.A. Pires-Zottarelli & A.V. Marano (SP 466373, SP 466376, SP 466377,
SP466379, paratypes), ex-paratypes CCIBt 4115 = MMBF 07/15, CCIBt 4153 =
MMBF 10/15, CCIBt 4154 = MMBF 11/15, CCIBt 4156 = MMBF 13/15.
Notes: Salispina intermedia appears as morphologically and phylogenetically
intermediate between S. spinosa CBS 591.85 (KT886057) and S. lobata CBS 588.85
(KT886056), Figs 154, 155 and 156. Its zoosporangial morphology resembles S.
spinosa, although their zoosporangia are considerably larger than those observed for
the ex-type cultures of S. spinosa (CBS 591.85) and S. lobata (CBS 588.85). Onto
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PYGs, S. intermedia forms clusters of zoosporangia visible at naked eye. This species
was particularly abundant and frequently recovered during spring (Nov) and summer
(Feb) samplings, when water temperature was higher (25–28°C) than in the other
samplings (18–22°C). We were not able of sequencing the ITS region of Salispina
using the primers ITS4 and ITS6 (Cooke et al. 2000) and UN-up18S42 and
UN-up28S22 (Robideau et al. 2011).
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Fig. 156 Salispina intermedia (holotype) a–d Zoosporangia of different morphologies, from
smooth (a) to with various degree of spines coverage (b–d) e–i Formation of a persistent tube
through which zoospores swim away (no vesicle is formed) f Detail of the zoosporangial
basal plug. Bars: a, h = 20 µm, b–g, i = 10 µm.
358. Salispina lobata (Fell & Master) A.L. Jesus, Marano & Pires-Zottar., comb. &
stat. nov.
Index Fungorum number: IF 551606
Basionym: Phytophthora spinosa var. lobata Fell & Master, Can. J. Bot. 53:
2919 (1975).
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7440
7441
7442
7443
7444
7445
7446
7447
7448
7449
7450
7451
7452
7453
7454
7455
7456
7457
7458
7459
7460
7461
7462
7463
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7471
7472
7473
7474
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7476
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7478
7479
= Halophytophthora spinosa var. lobata (Fell & Master) H.H. Ho& S.C. Jong,
Mycotaxon 36: 381 (1990).
Holotype: ATCC 28291 (Fell & Master 1975); ex-holotypes CBS 588.85, IFO
32592, IMI 33018.
Distribution: Malaysia, Seychelles, Singapore, Taiwan, Thailand, USA, Vietnam
(Fell and Master 1975; Marano et al. 2012).
359. Salispina spinosa (Fell & Master) Marano, A.L. Jesus & Pires-Zottar., comb. &
stat. nov.
Index Fungorum number: IF 551604
Basionym: Phytophthora spinosa var. spinosa Fell & Master, Can. J. Bot. 53:
2917 (1975).
= Halophytophthora spinosa var. spinosa (Fell & Master) H.H. Ho& S.C. Jong,
Mycotaxon 36: 381 (1990).
Holotype: ATCC 28294 (Fell & Master 1975); ex-holotypes CBS 591.85, IFO
32593, IMI 330187.
Distribution: Bahamas, Colombia, Grand Cayman, Haiti, Japan, Philippines,
Thailand, The Netherlands Antilles, Trinidad and Tobago, USA (Fell and Master
1975; Marano et al. 2012).
Zygomycota
Mortierellales
Mortierellaceae
Mortierella Coem.
The genus Mortierella, the type of the order Mortierellales, was described in
1863 by Coemans and the first described species was Mortierella polycephala. The
order Mortierellales is one of the largest basal fungal lineages. It is currently
classified either within the subphylum Mucoromycotina (Hibbett et al. 2007) or within
its own subphylum Mortierellomycotina (Hoffmann et al. 2011). Traditionally the
genus Mortierella was divided into nine sections (Gams 1977). However, recent
phylogenetic analyses do not support this classification. However some groups may
be distinguished: ‘selenospora and parvispora’, ‘verticillata-humilis’, ‘lignicola’,
‘mutabilis, globulifera and angusta’, ‘strangulata and wolfii’, ‘alpina and
polycephala’, ‘gamsii’. It was also shown that the genera Dissophora, Gamsiella,
Lobosporangium and Modicella (Smith et al. 2013) are placed within the genus
Mortierella. The morphology of Mortierellales is quite simplified and it seems to
depend on culture condition, explaining the incompatibility between morphological
and phylogenetic studies. The representatives of this group are mostly soil inhabiting
saprotrophs (Wagner et al. 2013). The phylogenetic trees for Mortierella are presented
in Figs 157, 158.
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Fig. 157 Maximum likelihood analysis based on the ITS1-5.8S-ITS2 dataset for clade
lignicola (as defined by Wagner et al. 2013). The phylogram is constructed from a muscle
alignment of 616 nucleotides of 42 strains. Node support above 75% is given. New taxa are in
blue and ex-type strains in bold.
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7493
7494
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Fig. 158 Maximum likelihood analysis based on the D1/D2 domain of the large subunit (LSU,
28S) dataset for selected species of Mortierellales. The phylogram is constructed from a
muscle alignment of 670 nucleotides of 27 strains. Node support above 75% is given. New
taxa are in blue and ex-type strains in bold.
360. Mortierella calciphila Wrzosek, sp. nov.
MycoBank number: MB 814918, Facesoffungi number: FoF 02063, Figs 159,
160
Etymology: refers to the type of soil where the species was found (limestone soil)
Holotype: WA18944
Radiate colonies fast-growing (6–9 mm per day on PDA), without characteristic
zonate growth nor garlic odour. Sporangiophores arising from the substratum with
2–4 (7) basal sympodial ramification or formed on aerial hyphae (then 0–1
ramification), slender, 2–3 µm under sporangium, 600–1400 µm long, without any
cross wall. Sporangia (27–) 70 (–80) µm in diam., many-spored, with early
deliquescent wall. Columella strongly reduced, without apophysis sometimes with
tiny projection (up to 1 µm) on the top. Spores broadly ellipsoidal, smooth-walled,
regular in shape (8–) 9 (–11) µm (SD = 0.8) × (6–) 7 (–9) µm (SD = 0.8). Gemmae
abundantly produced in substratum or aerial hyphae, in chains or irregular clusters
often connected by anastomosis, globose, hyaline to pale ochraceous (11–) 18 (–25)
µm (SD = 3.44) in diam.
Habitat and distribution: humid soil in beech forest on limestone, Northern
Poland
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Material examined: POLAND, West Pomeranian Voivodeship, Polanów Forest
District, Wapienny Las area (‘forest on limestone soil’), 53º59’59.16 N, 16º42’47.75
E,elev. 110 m, 26 August 2015, collector Marta Wrzosek; holotype WA18944 (dried
culture), ex-holotype CBS 140728 (lyophylised culture); ex type (living culture) is
deposited in Jena Microbial Resource Collection (University of Jena and Leibniz
Institute for Natural Product Research and Infection Biology, Jena, Germany)
(JMRC:SF:012173).
Notes: The phylogenetic analyses (Figs 157 and 158) show that this species
belongs to group lignicola as defined by Wagner et al. (2013). However, the similarity
of ITS sequence to any previously described taxa in this group is low: BS = 85% to
M. beljakovae, BS = 84% to M. paraensis, BS = 83% to M. formicicola, BS = 81% to
M. gemmifera and M. kuhlmanii.
The species is morphologically most similar to Mortierella zychae Linn., which
is placed by Wagner et al. (2013) in Clade 7 grouping some species from former
section “elongata” and to M. parazychae from Clade 5, containing M. wolfii and
relatives. The most characteristic feature of this fungus is formation of gemmae in
clusters, both in substrate and on aerial mycelium. Gams (1976) use the term
“chlamydospores” for gemmae, what seems to be not always proper because the cell
wall of these structures is very thin and it could be easily mechanically damaged. In
young cultures of M. calciphila the clusters of gemmae are quite loose, regularly
placed, and globose with enlargements. Sometimes they are arranged in chains with
thin liaisons (Fig. 160c, d). The cross walls were observed sporadically. Large
clusters, up to 0.8 mm diam. built by dense layers of gemmae, with very short liaison,
and with numerous anastomosae could be observed in older cultures (Fig. 160e). The
arrangement of the gemmae/chlamydospores in chains and clusters has been observed
also in M. zychae, M. parazychae, M. beljakowae, M. kuchlmanii and others (Gams
1976). The gemmae of M. calciphila (as well as these of M. parazychae) are usually
completely rounded, in contrast with M. zychae Linnem., where the gemmae outline
merges gradually into the connecting hyphal parts (Gams 1976). We suggest that
gemmae are organs of symbiotic associations with bacteria, which seem to be quite
widespread among Mortierellales, rather than resting structures (Fujimura et al. 2014,
Ogawa et al. 2012).
The sporangiophores of M. calciphila are more slender than in M. beljakovae and
do not have an apophysis, nor collerate. The sporangiophores of M. calciphila are
often larger than sporangiophores of M. zychae and others species of section
“elongata”. The spores of M. calciphila resemble those of M. zychae, but they are
colourless and some granules in cytoplasm could be seen. They are much more
regular than spores of M. parazychae, M. beljakowae and M. kuhlmanii.
The most closely related species to M. calciphila was M. formicicola D.S. Clark
& W. Gams. The ITS and LSU sequences of that species were generated by Wagner
et al. (2013) for phylogenetic studies, but the description of this fungus is not
available and probably this species is not validly published.
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Fig. 159 Mortierella calciphila (holotype) a Branching sporophore emerging from substrate
b Typical sporophore with sporangium on aerial mycelium c Loose cluster of young gemmae
d Top of sporophore e Spores from sporangium, and germinating spore. Scale bars: a = 100
µm, b = 50 µm, c = 20 µm, d, e = 10 µm.
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Fig. 160 Mortierella calciphila (holotype) a Type of growth (24 h colony) b Branching
sporophore c, d, e Gemmae forming loose (c, d) or dense (e) clusters f Small sporangium
formed on short sporophore emerging from aerial hyphae g, h Spores. Scale bars b = 500 µm,
c–e = 20 µm, f = 50 µm, g, h = 20 µm.
Mucorales
Cunninghamellaceae
Absidia Tiegh.
The genus Absidia (Cunninghamellaceae, Mucorales) was originally described
by van Tieghem (1876) with type species A. reflexa Tiegh. (Hesseltine and Ellis
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7607
1964). To the best of our knowledge, 21 species of Absidia have been reported thus
far (Kirk et al. 2008).
The species belonging to this genus are characterized by the production of
stolons and sporangiophores bearing pyriform columellate sporangia with
deliquescent walls with a septum below the apophysis; the sporangiophores of Absidia
never arise opposite the rhizoids as found in Rhizopus (Hesseltine and Ellis 1964).
Species of Absidia typically exhibit rapid growth at temperatures ranging from 25oC
to 34oC, although some species are able to grow at temperatures between 12oC and
37°C (Hoffmann et al. 2007). They are frequently isolated from soil and dead or dying
plant tissue (Hesseltine and Ellis 1964; Ho et al. 2004; Benny 2008). Several species
of Absidia are implicated in diseases such as mucormycosis in humans and animals
(Ribes et al. 2000; Santos 2003; Hoffmann and Voigt 2009; Alastruey-Izquierdo et al.
2010). Since first described, some species of Absidia have been transferred to other
genera, for example, Tieghemella Berl. & De Toni, Mycocladus Beauverie, and
Proabsidia Vuill. However, with the exception of Lichtheimia, all are regarded as
synonyms of Absidia (Hesseltine and Ellis 1964; Schipper 1990; Kirk et al. 2008).
Recently, Hoffmann et al. (2007) revised the classification of the genus based on
physiological, phylogenetic, and morphological characteristics. They observed
different growth patterns under different temperature conditions, and divided the
species into three groups, namely, thermotolerant (species that exhibited optimum
growth between 37oC and 45oC), mesophilic (species that exhibited optimum
growth between 25oC and 34oC), and mycoparasitic (species that are potentially
parasitic on other fungi within the order Mucorales and exhibit optimum growth
below 30oC). Although the identification of species based on morphological
characteristics is important in traditional taxonomy, the delimitation of species of
mucoralean fungi requires the addition of molecular data (O’Donnell et al. 2001;
Hoffmann et al. 2013; Walther et al. 2013). In a previous study, a new species,
Absidia koreana was reported from a soil sample from Dokdo island, Korea
(Ariyawansa et al. 2015b). The phylogenetic tree for Absidia is presented in Fig. 161.
While evaluating the diversity of fungi of the order Mucorales isolated from a
sample of rat dung from Gwangju, Korea a new species, based on morphological
characteristics and multi-gene phylogenetic analyses, was isolated and is described
here.
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Fig. 161 Phylogenetic tree for Absidia stercoraria EML-DG8-1 and EML-DG8-2 and related
species based on Maximum likelihood analysis of multi-genes including 18S and 28S rDNA,
actin (Actin-1) and translation elongation factor (EF-1α). Sequences of Umbelopsis nana and
U. isabellina were used as outgroups. Bootstrap support values >50% are indicated at the
nodes. The bar indicates the number of substitutions per position. New taxa are in blue and
ex-type strains in bold.
361. Absidia stercoraria Hyang B. Lee, H.S. Lee & T.T.T. Nguyen, sp. nov.
MycoBank number: MB 814409, Facesoffungi number: FoF 02064, Fig. 162
Etymology: stercoraria. Named for rat dung from which the species was first
collected.
Holotype: EML-DG8-1, deposited at the Environmental Microbiology
Laboratory Fungarium, Chonnam National University, Gwangju, Korea. Living
culture CNUFC-EML-DG8-1, in Chonnam National University Fungal Collection,
Gwangju, Korea.
Colonies exhibit rapid growth on SMA attaining a diam. of 85–90 mm after 5
days at 25oC, initial colour white, later changing to grayish-white or smoky-gray, the
reverse white and irregularly zonate. Sporangiophores are 4–6 µm wide and arise as
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7649
1–5 sporangiophores (average 2–3) per whorl from a single point on the stolons.
Sporangia 19–30 × 20–31 µm, globose to pyriform, multi-spored, frequently with a
bell-shaped apophysis. Columellae are 9–13 × 12–13.5 µm, hemisphaerical.
Collarette appearing after sporangium maturation. Sporangiospores mostly short
cylindrical, 4–5 × 2–3 µm. Zygospores not observed and rhizoids not well developed.
Notes: Absidia stercoraria is morphologically similar to A. koreana, but
apparently differs from the related species by having a bell-shaped apophysis when
cultivated on SMA, and by multi-gene sequence data. It is currently known from a
single collection.
Material examined: REPUBLIC OF KOREA, Division of Food Technology,
Biotechnology & Agrochemistry, College of Agriculture & Life Sciences, Chonnam
National University, Gwangju 61186, Korea, from rat dung sample from Gwangju,
Korea; EML-DG8-1 (ex-type) at Culture Collection of National Institute of Biological
Resources (NIBR), Incheon, and preserved as glycerol stock at -80oC in the CNUFC;
living culture(ex-type) deposited at Jena Microbial Resource Collection (University of
Jena and Leibniz Institute for Natural Product Research and Infection Biology, Jena,
Germany) (JMRC:SF:012179) (ex-type).
The isolate was observed to grow over a wide range of temperatures with varying
growth rates of 18 mm, 14 mm, and 13 mm per 24 hours on SMA, PDA and MEA,
respectively. Optimal growth was observed around 25–27oC, slow growth was
observed down to 20oC, and no growth above 35oC. Absidia stercoraria appears to be
phylogenetically related to A. koreana, both clustering in the same clade together with
other Absidia spp. within the family Cunninghamellaceae (Fig. 161).
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Fig. 162 Absidia stercoraria (holotype) a, b Colony in synthetic mucor agar (a obverse view,
b reverse view) c, d Young sporangia with sporangial net wall e Young sporangium with a
bell-shaped apophysis (red arrow) f, g Mature sporangia with bell-shaped apophysis h Mature
sporangium without bell-shaped apophysis i Columellae with collarette and a single
projection (yellow arrow), and septum (white arrow) below the apophysis j, k Rod-shaped
sporangiospores. Scale bars: c, d = 10 µm, f–i = 20 µm, j, k = 5 µm.
Gongronella Ribaldi
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Mucorales comprise ubiquitous, mostly saprotrophic organisms and are one of
the most ancient groups of fungi. They can be easily isolated from soil, dung, water,
stored grains, plants, as well as other fungi due to their rapid growth rate and ability to
colonize and sporulate on diverse, carbohydrate-rich, terrestrial substrates (Benny
2008; O'Donnell et al. 2001). Some species are responsible for a number of
opportunistic infections in immunocompromised humans and other mammals
(Hoffmann et al. 2013). The genus Gongronella (Cunninghamellaceae, Mucorales)
was established in 1952 by Ribaldi, for a single species, Gongronella urceolifera
Ribaldi (Ribaldi 1952). The primary reason for introducing a separate genus to
accommodate this species was its distinct urn-shaped apophyses and columellae.
Three years later, based on the presence of an identical apophysis, Peyroneland Dal
Vesco (1955) and Pici (1955) transferred Absidiabutleri Lendn. to Gongronella, both
studies indicating that the type species, G. urceolifera, was identical to G. butleri
(Lendn.) Peyronel & Dal Vesco. Hesseltine and Ellis (1961) added an additional
species, G. lacrispora Hesselt. & J.J. Ellis, differing from G. butleri by forming
circinate sporangia and teardrop-shaped sporangiospores. To date Gongronella
includes only these two species: G. butleri and G. lacrispora (Kirk et al. 2008).
Recently, Walther et al. (2013) showed that Hesseltinella vesiculosa H.P. Upadhyay
and Circinellala crymispora Aramb. & Cabello belong to the Gongronella clade, but
their morphological characteristics differ from those of the other species of
Gongronella. In general, species of Gongronella grow slowly between 25ºC and 27ºC
(Hesseltine and Ellis 1964) and are frequently found in soil (Hesseltine and Ellis
1961; Upadhyay 1969; Ho and Chen 1990). Several studies have reported that species
of Gongronella have important biotechnological applications, such as the production
of enzymes and antifungal proteins (Zhou et al. 2008; Wang et al. 2008; Wei et al.
2010). The taxonomy of Gongronella has been determined on the basis of
morphological characteristics including the size and shape of sporangia,
sporangiospores and columellae. Benny (1995) alluded to the limitations in the usage
of morphological characters for species delineation in certain zygomycetes, and has
suggested the use of molecular tools for solving existing controversies surrounding
taxonomic classification. O’Donnell et al. (1998) also suggested that the traditional
classification scheme for Zygomycota did not reflect the phylogenetic relationships
among these taxa. Recently, molecular identification has been evaluated for
Mucorales. O’Donnell et al. (2001) performed a comprehensive study of Mucorales
with partial nucleotide sequences of nuclear 18S ribosomal RNA small subunit (SSU),
nuclear large subunit 28S ribosomal RNA (LSU), and translation elongation factor-1α
(EF-1α) gene exons. The phylogeny of Mucorales was also studied by White et al.
(2006), who used the combined rRNA operon (18S + 28S + 5.8S gene) to infer
relationships. In recent years, several studies based on multi-loci analysis (18S, 28S,
EF-1α, actin, RNA polymerase II) have been conducted (Tanabe et al. 2003;
Hoffmann et al. 2013; Walther et al. 2013). Different molecular targets have been
used to characterize phylogenetic genera. In a previous study, a new species, G.
koreana, isolated from forest soil from Jeonnam, Korea, was reported (Ariyawansa et
al. 2015b). The phylogenetic trees are presented in Figs 163 and 164.
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While evaluating the diversity of fungi of the order Mucorales isolated from a
soil sample collected at Gwangan beach, Busan, Korea, an isolate showing
morphological variation compared to other species of Gongronella was identified and,
based on subsequent multi-gene phylogenetic analyses is described here as a new
species.
Fig. 163 Phylogenetic tree for Gongronella orasabula EML-QF12-1 and EML-QF12-2 based
on Maximum likelihood analysis of ITS rDNA sequence. Sequences of Gongronella
lacrispora was used as outgroup. Bootstrap support values >50% are indicated at the nodes.
The bar indicates the number of substitutions per position. New taxa are in blue and ex-type
strains in bold.
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Fig. 164 Phylogenetic tree for Gongronella orasabula EML-QF12-1 and EML-QF12-2 and
related species based on Maximum likelihood analysis of multi-genes including 18S and 28S
rDNA, actin (Actin-1) and translation elongation factor (EF-1α). Sequences of Umbelopsis
nana and U. isabellina were used as outgroups. Bootstrap support values >50% are indicated
at the nodes. The bar indicates the number of substitutions per position.
362. Gongronella orasabula Hyang B. Lee, K. Voigt, P.M. Kirk & T.T.T. Nguyen,
sp. nov.
MycoBank number: MB 814447, Facesoffungi number: FoF 02065, Fig. 165
Etymology: orasabula. Referring to beach soil from which the species was first
isolated (Busan, Korea).
Holotype: EML-QF12-1, deposited at the Environmental Microbiology
Laboratory Fungarium, Chonnam National University, Gwangju, Korea. Living
culture CNUFC-EML-QF12-1, in Chonnam National University Fungal Collection
(CNUFC), Gwangju, Korea.
Colonies exhibit fast growth on SMA attaining a diam. of 33–35 mm after 5 days
at 25oC, initial colour white, later off-white, in reverse white with an irregular margin.
Sporangiophores 35–200 × 2.5–4 µm, erect, either unbranched or with 2–3 branches.
Sporangia 12–20 × 12.5–22 µm, globose to subglobose or calabash vase-shaped,
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multi-spored, with a thin wall having a purplish tinge and deliquescent at maturity.
Columellae 2–3 × 3–4 µm, hemisphaerical, with a collarette. Apophysis of diverse
shape, globose, subglobose to pyriform, 5–10 × 4.5–8.5 µm. Sporangiospores mostly
bean-shaped, 2–3.5 × 2–2.5 µm. Chlamydospores absent in aerial mycelia.
Zygospores not observed; rhizoids not well developed.
Notes: Gongronella orasabula is morphologically similar to G. koreana, but
differs from related species by having larger and differently shaped sporangia. The
apophysis is also larger, mainly globose, subglobose or pyriform or rarely long
conical. Furthermore, the isolate has two septa below the apophysis.
Material examined: REPUBLIC OF KOREA, Division of Food Technology,
Biotechnology & Agrochemistry, College of Agriculture & Life Sciences, Chonnam
National University, Gwangju 61186, Korea, from a soil sample collected at Gwangan
beach, Busan, Korea; EML-QF12-1 (ex-type) at Culture Collection of National
Institute of Biological Resources (NIBR), Incheon, and preserved as glycerol stock at
-80oC in the CNUFC; living culture (ex-type) deposited at Jena Microbial Resource
Collection (University of Jena and Leibniz Institute for Natural Product Research and
Infection Biology, Jena, Germany) (JMRC:SF:012180).
The isolate was observed to grow over a wide range of temperatures with varying
growth rates of 7.3 mm, 6.7 mm, and 6 mm per 24 hours on SMA, PDA (potato
dextrose agar), and MEA (malt extract agar), respectively. Optimal growth was
observed at 27oC, slow growth was observed at 20oC, and no growth at 37oC.
Gongronella orasabula appears to be phylogenetically related to G. koreana, both
clustering in the same clade together with G. butleri which is the type of the genus
Gongronella (Figs 163, 164).
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Fig. 165 Gongronella orasabula (holotype) a, b Colony in synthetic mucor agar (SMA) (a
from above, b reverse view) c–g Mature sporangia with variously shaped apophysis (red
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arrows) and sporangia h Columellae with collarette and two septa (blue arrows). Scale bars =
20 µm.
Mucor Fresen.
The zygomycota is an artificial grouping of related basal clades comprising the
subphyla Mortierellomycotina Kerst. Hoffm. et al., Mucoromycotina Benny,
Kickxellomycotina Benny and Zoopagomycotina Benny (Muszewska et al. 2014). The
genus Mucor is the largest within the Mucoromycotina and includes more than 50
species several of which have important economical application, including the
production of enzymes, fumaric acid, fatty acid, and also antifungal agents for plants
(Dexter and Cooke 1984; Alves et al. 2002; Roa Engel et al. 2008). It is characterized
by fast-growing colonies, simple or branched sporangiophores without basal rhizoids,
non-apophysate sporangia, and zygospores which are borne from opposed suspensors,
possess a thick pigmented and ornamented zygosporangium and are seldom produced
(Schipper and Samson 1978; Benny 2013). This genera has a worldwide distribution,
with most species described as saprobes commonly isolated from soil, stored grains,
fruits, vegetables and the excrement of herbivores (Schoenlein-Crusius et al. 2006;
Jacobs and Botha 2008; Santiago et al. 2011, 2013).
According to Álvarez et al. (2011) Mucor has the greatest number of described
species among Mucorales. In a series of studies, Schipper (1973, 1975, 1976, 1978)
monographed this genus and described 39 species, four varieties and 11 forms.
Subsequently, 17 species have been proposed (Mehrotra and Mehrotra 1978; Mirza et
al. 1979; Subrahamanyam 1983; Chen and Zheng 1986; Schipper and Samson 1994;
Watanabe 1994; Zalar et al. 1997; Pei 2000; Alves et al. 2002; Jacobs and Botha 2008;
Hermet et al. 2012; Madden et al. 2012).
Molecular studies have shown that Mucor is polyphyletic (O’Donnell et al. 2001;
Kwasna et al. 2006; Jacobs and Botha 2008; Budziszewska and Piatkowska 2010;
Álvarez et al. 2011). Based on phylogenetic relationships inferred from data of LSU
and ITS regions (rDNA), and morphological characteristics, Walther et al. (2013)
concluded that Mucor and Backusella Hesselt. & J.J. Ellis species represents a natural
group characterized by transitorily recurved sporangiophores. Therefore, all Mucor
species with this feature were transferred to Backusella [B. grandis (Schipper &
Samson) G. Walther & de Hoog, B. indica (Baijal & B.S. Mehrotra) G. Walther & de
Hoog, B. oblongielliptica (H. Nagan., Hirahara & Seshita ex Pidopl. & Milko) G.
Walther & de Hoog, B. oblongispora (Naumov) G. Walther & de Hoog, B. recurva
(E.E. Butler) G. Walther & de Hoog, B. tuberculispora (Schipper) G. Walther & de
Hoog, and B. variabilis (A.K. Sarbhoy) G. Walther & de Hoog]. Considering that
some of the characteristics traditionally used to separate Zygorhynchus Vuill. from
Mucor, such as the unequal suspensors of the zygospores and the Zygorhynchus
zygospore production pattern (two suspensors originating from the same hypha) do
not represent synapomorphies of the genus Zygorhynchus, and seem to be convergent
characters within Mucor, Walther et al. (2013) recombined all Zygorhynchus species
in Mucor as follows: M. exponens (Burgeff) G. Walther & de Hoog, M. fusiformis G.
Walther & de Hoog, M. heterogamus Vuill., M. japonicus (Komin.) G. Walther & de
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Hoog, M. megalocarpus G. Walther & de Hoog, M. moelleri (Vuill.) Lendn. and M.
multiplex (R.Y. Zheng) G. Walther & de Hoog. Non-thermophilic Rhizomucor
endophyticus and Circinella rigida were reclassified as M. endophyticus (R.Y. Zheng
& H. Jiang) J. Pawłowska & G. Walther and M. durus G. Walther & de Hoog,
respectively.
Recently, molecular data have been used to evaluate mucoralean species
(Hoffmann et al. 2013; Walther et al. 2013). During studies on the Mucorales from
Brazil and Korea, taxa of Mucor that differs morphologically and molecularly from
the other species was isolated and are thus described as new. The phylogenetic tree for
Mucor are presented in Figs 166–169.
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Fig. 166 Phylogenetic tree of Mucor constructed using the large subunit (LSU) rDNA
sequence data. Circinella species were used as outgroup. Sequences are labeled with their
database accession numbers. Support values are from Bayesian inference and maximum
likelihood analyses (values above and below the branches, respectively). The sequences
obtained in this study are annotated in blue.
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Fig. 167 Phylogenetic tree of M. amphibiorum group constructed using the ITS rDNA
sequences. Mortierella parvispora was used as outgroup. b Phylogenetic tree of Mucor
hiemalis group constructed using the ITS rDNA sequences. Mucor gigasporus was used as
outgroup. Sequences are labeled with their database accession numbers. Support values are
from Bayesian inference and maximum likelihood analyses (values above and below of the
branches, respectively). Sequences with only ITS1 and 5.8s rDNA are marked with *. New
taxa are in blue and ex-type strains in bold.
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Fig. 168 Phylogenetic tree for Mucor koreanus EML-QT1 and EML-QT2 based on
Maximum likelihood analysis of ITS rDNA sequence. Sequence of Syncephalastrum
racemosum was used as outgroup. Bootstrap support values >50% are indicated at the nodes.
The bar indicates the number of substitutions per position. New taxa are in blue and ex-type
strains in bold.
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Fig. 169 Phylogenetic tree for Mucor koreanus sp. nov. EML-QT1 and EML-QT2 and related
species based on Maximum likelihood analysis of multi-genes of 18S and 28S rDNA, actin
(Actin-1) and translation elongation factor (EF-1α). Sequences of Umbelopsis nana and U.
isabellina were used as outgroups. Numbers at the nodes indicate the bootstrap values (>50%)
from 1000 replications. The bar indicates the number of substitutions per position.
363. Mucor caatinguensis A.L. Santiago, C.A. de Souza & D.X. Lima, sp. nov.
Index Fungorum number: IF 551680, Facesoffungi number: FoF 01328,
Fig. 170
Etymology: caatinguensis. Referring to the biome where the species was
first isolated.
Holotype: URM 7322
Fast growing colonies, 9 cm diam. after 72 hours in MEA at 25 ºC, firstly white
then turning cream with grey spots (MP 18A1), touching the plate lid in the central
region. Reverse yellow (MP 10H2). Sterile mycelium abundant. Sporangiophores
coenocytic, simple or slightly sympodially branched with long branches,
(5–)7.5–15(–17) µm diam. with or without yellowish contents, slightly roughed-wall.
Some sporangiophores show a globular swelling distant from the columellae.
Sporangia first yellow then becoming light brown, globose, subglobose, 25–65 µm
diam., subglobose to slightly flattened, 30–60 × 32–55 µm with a slightly echinulate
wall. Columellae light gray, smooth-walled, globose, subglobose, (20–)2–45 (–60) µm
in diam., ellipsoid, obovoid with a truncated base (mostly) and piriform
(–25)30–60(–75) × (20–)27–45(–55) µm. Collar evident. Columellae cylindrical with
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or without a constriction in the central part, 24.5–35 × 30–55 µm where rarely
observed. Sporangiospores hyaline, smooth-walled, regular in size and containing
granules at each end, mostly ellipsoid, 5–6(–7) × 3–5 µm and cylindrical ellipsoid,
5–6 × 3–4 µm., some subglobose and globose, 3–5 µm diam. Chlamydospores
abundant, globose, subglobose and doliform, sometimes produced in the
sporangiophores. Zygosporangia not observed.
Material examined: BRAZIL, Buíque: Parque Nacional do Catimbau
(8º31’55.8’’S, 37º15’34.2’’W), in soil samples. Soil, 11.III.2014, leg. C. Lira (URM
7322) and deposited in the Jena Microbial Resource Collection (University of Jena
and Leibniz Institute for Natural Product Research and Infection Biology, Jena,
Germany) (JMRC:SF:012174).
Media and temperature tests: On MEA. At 10ºC – very limited growth (2 cm in
diam. in 120 hours); total lack of reproductive structures. At 15ºC – low colonies (< 1
mm in height) with slow growth (4 cm in diam. after 120 hours); poor sporulation. At
20ºC – low colonies (<1mm diam.) with slow growth (5 cm in 120 hours); good
sporulation. At 25ºC – better growth (9 cm in 72 hours); excellent sporulation. At
30ºC – good growth (8 cm in 72 hours); excellent sporulation. At 35ºC – better
growth than at 15 and 20ºC (9 cm in 120 hours); rare sporangiophores production and
poor sporulation. At 40ºC – lack of growth and sporulation. The growth of M.
caatinguensis on PDA was slightly slower than on MEA at all tested temperatures.
However, at 35ºC, on PDA, the production of reproductive structures was good, and
the sporangiophores were more sympodially branched (up to seven times) than in at
other temperatures. The columellae were mostly applanate and bizarrely shaped
sporangiophores were also observed.
Notes: Mucor caatinguensis is distinguished from the other species of the genus
as it simultaneously produces numerous chlamydospores in mycelia (sometimes in
sporangiophores), unbranched or weakly branched sporangiophores, columellae and
sporangiospores that are variable in shape and size. At first, Mucor caatinguensis
could be confused with M. silvaticus Hagem because of the unbranched or weakly
sympodially branched sporangiophores, the small size of the sporangia (up to 70 µm
diam.) and by the production of cylindrical ellipsoid sporangiospores. However,
colonies of M. silvaticus are pale olive gray, and it produces blackish brown sporangia
(Schipper 1973), in contrast to the cream colonies of the new species, which show
shows light brown sporangia. The former only produces sphaerical columellae, which
are rarely ellipsoidal, never obovoid with a truncated base or piriform, as observed in
M. caatinguensis. Additionally, the sporangiospores of M. silvaticus are 3.5–5.2 ×
2.6–3.7 µm, smaller than the M. caatinguensis sporangiospores, and no
chlamydospores where reported in M. silvaticus
The abundant production of chlamydospores, sometimes observed in
sporangiophores, is also very common in M. racemosus f. racemosus Fresen.
(Schipper 1976), although we did not observed these structures were not observed
inside the columellae of M. caatinguensis. Nevertheless, the sporangiophores of M.
caatinguensis are not as branched as those of M. racemosus f. racemosus which may
be sympodially and monopodially branched. Additionally, the sporangiospores of M.
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racemosus f. racemosus are broadly ellipsoidal to subglobose, and the colonies of M.
racemosus f. racemosus are pale smoke gray, whereas the colonies of the new species
are cream with grey spots.
Our molecular analysis (LSU and ITS rDNA, Figs 166, 167, respectively)
showed that M. caatinguensis is genetically different from the other species of the
genus, and placed the new species within the M. amphibiorum group, close to M.
indicus Lendn. (Walther et al. 2013). In fact, the colour of both colonies of M. indicus
and M. caatinguensis may be similar, but the sporangiophores of M. indicus are
repeatedly sympodially branched (with long branches) and the columellae are mostly
applanate and subglobose. We found repeatedly sympodial branches in M.
caatinguensis at 35ºC on PDA. According to Schipper (1978), chlamydospores of M.
indicus are also abundant in cultures grown in darkness at 20 ºC, but only in substrate
hyphae, and the sporangiospores are ellipsoidal to globose.
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Fig. 170 Mucor caatinguensis (holotype) a Colony surface b, b1 Simple sporangiophore
with chlamydospores c Simple sporangiophore with sporangia d Sporangiophore branch e–g
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Simple sporangiophores with columellae with different shapes h Chlamydospores i
Sporangiospores.
364. Mucor koreanus Hyang B. Lee, S.J. Jeon & T.T.T. Nguyen, sp. nov.
MycoBank number: MB 814424, Facesoffungi number: FoF 02066, Fig. 171
Etymology: koreanus. Referring to the country which from the species was first
isolated (Korea).
Holotype: EML-QT1, deposited at the Environmental Microbiology
Laboratory Fungarium, Chonnam National University, Gwangju, Korea. Living
culture CNUFC-EML-QT1, in Chonnam National University Fungal Collection,
Gwangju, Korea.
Colonies growing fast on PDA, dark brown in the center, with a lighter margin,
grayish-white in reverse, reaching 70–72 mm diam. at 23oC after 2 days of incubation.
Sporangiophores 21–44 µm wide, erect, unbranched or branched sympodially.
Sporangia globose, subglobose, yellow to golden brown, multi-spored, reaching
129–159 × 137–165 µm; at maturity the sporangial wall fully deliquesces, leaving a
small collar. Columellae globose, cylindrical-ellipsoidal, reniform or pyriform, 67–82
× 71–87 µm. Sporangiospores of diverse shape, ellipsoidal, globose, sometimes
asymmetrically globose or bean-shaped, 9–14 × 6.5–11.5 µm. Zygospores not
observed.
Notes: Mucor koreanus was similar in morphology and closely related to M.
piriformis A. Fisch., but differs by larger sporangiospores, their different shapes, and
colour of sporangia. Columellae are diverse in shape. Sometimes, the collar is not
seen below the columellae. The sporangiospores have thick walls.
Material examined: REPUBLIC OF KOREA, Division of Food Technology,
Biotechnology & Agrochemistry, College of Agriculture & Life Sciences, Chonnam
National University, Gwangju 61186, Korea, from a tangerine fruit purchased from
the grocery store in Korea; EML-QT1 (ex-type) at Culture Collection of National
Institute of Biological Resources (NIBR), Incheon, Korea, and preserved as glycerol
stock at –80oC in the CNUFC; living culture (ex-type) deposited at Jena Microbial
Resource Collection (University of Jena and Leibniz Institute for Natural Product
Research and Infection Biology, Jena, Germany) (JMRC:SF:012181).
The isolate was observed to grow over a wide range of temperatures with varying
growth rates on PDA, MEA (malt extract agar), and CDA (czapek dox agar) of 35
mm, 17 mm and 28 mm per 24 hours, respectively. Optimal growth was observed
around 20–23oC, slow growth was observed at 5oC, and no growth at 27oC. Mucor
koreanus appears to be phylogenetically related to M. piriformis, both clustering in
the same clade together with M. mucedo which is the type of the genus Mucor (Figs.
168, 169).
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Fig. 171 Mucor koreanus (holotype) a Colony on potato dextrose agar b–d Young sporangia
e Mature sporangium f–k Columellae with clear collar present at the apex of the
sporangiophore l Sporangiospores. Scale bars: b, c = 50 µm, d–l = 20 µm.
365. Mucor merdicola C.A. de Souza & A.L. Santiago, sp. nov.
Index Fungorum number: IF 551679, Facesoffungi number: FoF 01327,
Fig. 172
Etymology: merdicola. Merda-dung, cola-dwelling.
Holotype: URM 7223
Colony initially white then becoming yellowish to cream (MP 19D1) with
yellowish reverse (MP 11J6), reaching 9.5 cm in diam. and 9 mm in height after 4
days in MEA at 25ºC. Sporangiophores simple or repeatedly sympodially branched,
erect, some slightly curved, arising from aerial hyphae (3–) 5–15.5 (–18) µm diam.,
hyaline, smooth walled, with or without yellowish contents. Sporangia globose (16–)
17.5–60 (–85) µm diam., initially yellow becoming greyish brown with diffluent wall,
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smooth-walled. Columellae globose (12.5–)15–45(–50) µm, subglobose and applanate
(15–)20–29 × 30–34(–35) µm, hyaline or light to grey, smooth-walled; collar absent
or little evident. Sporangiospores smooth-walled, hyaline, mostly ellipsoid to
fusiform (2.5–)5–7 × 5–8.5(–10.5), but also ellipsoid 4–7.5 × 3–7.5(–10) µm or
subglobose (2.5–)4–7.5(–8.5) µm diam., rarely globose. Rhizoids poorly developed.
Chlamydospores globose, subglobose, doliform, some bizarre in shape.
Zygosporangia not observed.
Media and temperature tests: On MEA. At 5ºC – lack of growth and sporulation.
At 10ºC – slow growth colonies, reaching 5.9 cm in diam. after 168 hours; poor
sporulation. At 15ºC – Slow growth (9 cm in 192 hours); good sporulation. At 20ºC –
Better growth than at 10 and 15ºC (9 cm in 120 hours); good sporulation. At 25ºC –
better growth (9 cm in 72 hours); excellent sporulation. Growth reasonably good at
30ºC (9 cm in 144 hours); good sporulation. At 35ºC – limited growth (3.3 cm in 168
hours); rare sporophores production and poor sporulation. At 40ºC – lack of growth
and sporulation. The growth of M. merdicola on PDA was a slightly slower than on
MEA at all tested temperatures.
Material examined: BRAZIL, Arcoverde: Instituto Agronômico de Pernambuco
(IPA) (8°25'00"S; 37°04'00"W), in dung samples, Bos taurus L., breed Holandesa.
Dung, 05.IX.2014, leg. C.A.F de Souza (URM 7223) and deposited in the Jena
Microbial Resource Collection (University of Jena and Leibniz Institute for Natural
Product Research and Infection Biology, Jena, Germany) (JMRC:SF:012175).
Habitat: Dung.
Notes: Mucor merdicola is morphologically similar to M. circinelloides f.
circinelloides Tiegh. The former is distinguished from M. circinelloides f.
circinelloides as it produces globose, subglobose and applanate columellae, differing
from the obovoid to ellipsoidal columellae of M. circinelloides f. circinelloides as
described by Schipper (1976). The author describes globose columellae in M.
circinelloides f. circinelloides only in the small sporangia. Additionally, M. merdicola
presents sporangiospores smooth-walled, mostly ellipsoid to fusiform, 5–7 × 5–8.5
µm, but also ellipsoid, subglobose and rarely globose, whereas M. circinelloides f.
circinelloides sporangiospores are only ellipsoidal, mostly 5.4 × 4 µm (Schipper,
1976). Our molecular analysis (ITS and LSU rDNA shown in Figs 166 and 167,
respectively) revealed that M. merdicola is genetically different from the other species
of the genus and placed the new species within the M. hiemalis group (Figs 166 and
167), in which species are characterized as producing tall sporangiophores that are
weakly sympodially branched, and small sporangia that do not exceed 80 µm diam.,
while M. merdicola produces sporangiophores that are repeatedly sympodially
branched, similar to the ones produced by the species from the M. circinelloides group
(Fig. 4b).
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Fig. 172 Mucor merdicola (holotype) a Colony surface b A sympodially branched
sporangiophore c, d Simple sporangiophores with sporangia e–g Simple sporangiophores with
columellae with different shapes h Sporangiospores.
Rhizopus Ehrenb.
The classification system in the genus Rhizopus was previously revised based on
physiological and morphological characteristics such as size of sporangia and
sporangiophore and branching of rhizoids (Schipper 1984; Schipper and Stalpers
1984). Recently, however, molecular identification has been employed by analyses of
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rDNA ITS, small subunit (SSU), large subunit (LSU), actin (Actin-1) and translation
elongation factor (EF-1α) genes (Abe et al. 2007, 2010; Hoffmann et al. 2013;
Walther et al. 2013).
The genus Rhizopus, one of the genera of Mucoromycotina, includes many
species that are often used as starters in food fermentation. In Asia especially, some
species of Rhizopus are used to make Tempe, a fermented food based on soybeans
(Schipper 1984; Schipper and Stalpers 1984). However, several species of Rhizopus
are also implicated in diseases such as mucormycosis in humans and animals (Frye
and Reinhardt 1993).
During a study on the Mucorales from a persimmon fruit in Korea, a species of
Rhizopus was isolated that differs morphologically and molecularly from other
species and is described here as new. The phylogenetic trees for Rhizopus are
presented in Figs. 173, 174.
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Fig. 173 Phylogenetic tree for Rhizopus koreanus EML-HO95-1 and EML-HO95-2 based on
Maximum likelihood analysis of ITS rDNA sequence. Sequence of Phycomyces
blakesleeanus was used as outgroup. Bootstrap support values >50% are indicated at the
nodes. The bar indicates the number of substitutions per position. New taxa are in blue and
ex-type strains in bold.
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Fig. 174 Phylogenetic tree for Rhizopus koreanus EML-HO95-1 and EML-HO95-2 and
related species based on Maximum likelihood analysis of multi-genes including 18S and 28S
rDNA, actin (Actin-1) and elongation factor (EF-1α). Sequences of Umbelopsis nana and U.
isabellina were used as outgroups. Bootstrap support values >50% are indicated at the nodes.
The bar indicates the number of substitutions per position.
366. Rhizopus koreanus Hyang B. Lee & T.T.T. Nguyen, sp. nov.
MycoBank number: MB 814406, Facesoffungi number: FoF 02067, Fig. 175
Etymology: koreanus. Referring to the country which from the species was first
isolated (Korea).
Holotype: EML-HO95-1, deposited at the Environmental Microbiology
Laboratory Fungarium, Chonnam National University, Gwangju, Korea, as dried
fungal mass from culture (PDA), isolated from persimmon fruit, August 2014, by
H.B. Lee. Living culture CNUFC-EML-HO95-1, in Chonnam National University
Fungal Collection, Gwangju, Korea.
Colonies growing fast on PDA, reaching 73–77 mm diam. at 23oC after 1 day of
incubation, initially white, later grayish-black, reverse white, irregularly zonate.
Sporangia globose to oval, reaching 88–215 × 84–193 (mean 123 × 126) µm in diam.
Columellae 20–62 × 26–80 µm, conical, hemisphaerical or globose. Sporangiospores
globose to ellipsoidal, sometimes asymmetrically ovoid, 12.5–17 × 14–19 (mean 14.6
× 15.4) µm. Zygospores are seldom observed in the artificial media.
Notes: Rhizopus koreanus is similar in morphology and closely related to R.
stolonifera (Ehrenb.) Vuill., however the columellae were smaller, diverse in shape,
reaching 20–61 × 26–79 µm, forming a separate clade as a new species in a
phylogenetic tree.
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Material examined: REPUBLIC OF KOREA, Division of Food Technology,
Biotechnology & Agrochemistry, College of Agriculture & Life Sciences, Chonnam
National University, Gwangju 61186, Korea, from a persimmon fruit purchased from
the grocery store in Korea; EML-HO95-1 (ex-type) at Culture Collection of National
Institute of Biological Resources (NIBR), Incheon, and preserved as glycerol stock at
-80oC in the CNUFC; living culture (ex-type) deposited at Jena Microbial Resource
Collection (University of Jena and Leibniz Institute for Natural Product Research and
Infection Biology, Jena, Germany) (JMRC:SF:012182).
The isolate was observed to grow over a wide range of temperatures with varying
growth rates on PDA, MEA (malt extract agar), and OA (oatmeal agar) of 74 mm, 52
mm, and 47 mm per 24 hours, respectively. Optimal growth was observed around
20–25oC, slow growth was observed at 5oC, and no growth at 35oC. Rhizopus
koreanus appears to be phylogenetically related to R. stolonifer which is the type of
the genus Rhizopus (Figs 173, 174).
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Fig. 175 Rhizopus koreanus (holotype) a Colony in potato dextrose agar b Rhizoids (white
arrow) c–e Young sporangia f, g Mature sporangia h–k Different shapes of columella l, m
Sporangiospores with asymmetrically oval to globose shapes. Scale bars: b = 200 µm, c–g =
50 µm, d, e = 50 µm, h–k = 20 µm, l = 30 µm, m = 5 µm.
Acknowledgements
G.J. Li, H.A. Wen, X.Z. Liu, M.Q. He, and R.L. Zhao thank the National Natural
Science Foundation of China (No. 30770013, No. 31500013, No. 31000013, No.
31360014, No. 31470152), the Special Program of Basic Science of the Ministry of
Science and Technology (No. 2012FY111600), the Technology of and International
Cooperation Program of the Ministry of Science and Technology (No.
2009DFA31160) of the People’s Republic of China, and the opening funding of State
key Laboratory of Mycology, Institute of Microbiology, Chinese Academy of
Sciences for funding. They are also grateful to S.P. Jiang, A.G. Xu (Xizang Institute
of Plateau Biology), Wangmu (Agriculture and Animal Husbandry College of
Xizang), R. Wang, S.P. Wan (Kunming Institute of Botany, Chinese Academy of
Sciences), T.Z. Wei, W.L. Lu, R.H. Yang, X.Y. Liu, X.Y. Li, L. Jiang, B.B. Li, Y. Li,
Y. Yu, M.J. Zhao, H. Li, S.H. Jiang, Z.X. Zhu, and J. Bing for assistance in specimen
collecting; to Y.J. Yao, H.M. Lü, Q.R. Yan, and L. Yang for loans of herbarium
specimens; to X.F. Zhu for inking in line drawings; to X.L. Wang for providing
advice and suggestions in the phylogenetic analysis; to C.L. Li, X.L. Zhang, and J.N.
Liang (Institute of Microbiology, Chinese Academy of Sciences) for providing help
with SEM photography. S. Mongkolsamrit, D. Thanakitpipattana and J.J. Luangsa-ard
thank National Center for Genetic Engineering and Biotechnology (BIOTEC) for
project "Surveys and Collection Invertebrate-Pathogenic Fungi and Xylariaceae on
Forests Conservation of Thailand" grant number P-14-51240. They would also like to
thank M. Tanticharoen, K. Kirtikara, L. Eurwilaichitr and R. Tantalakha for their
support of the program "Biodiversity studies of entomopathogenic fungi in Thailand".
They are grateful to the Department of National Parks, Wildlife and Plant
Conservation for their cooperation and support to our project research. A.LCM de A.
Santiago thanks the Fundação de Amparo à Ciência e Tecnologia do Estado de
Pernambuco (FACPE - APQ 0842-2.12/14), the Conselho Nacional de
Desenvolvimento Científico e Tecnológico (CNPq - 458391/2014-0) and the
Programa de Pesquisa em Biodiversidade do semiárido (MCT/CNPQ/PPBio 457498/2012-9). T. Boonpratuang, S. Parnmen, and T. Thummarukcharoen thank the
National Science and Technology Development Agency (NSTDA) Cluster and
Management Program Office (CPMO) for the flagship project “Raising the quality
and standards of microorganism data for use in biotechnology” grant number
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P-12-01829 and the Thailand Research Fund (TRG5780217) and Department of
Medical Sciences, Ministry of Public Health. They also express appreciation to T.
Flegel for encouragement of working on mushroom taxonomy and especially on
poisonous mushrooms, L. Eurwilaichitr for funding support for molecular
identification through BBH, and always support on mushroom taxonomy and M.
Tanticharoen for forever support in biodiversity and taxonomy study in Thai fungi.
They also thank the SRRT Team, Bureau of Epidemiology, Department of Disease
Control Ministry of Public Health for collecting a specimen of poisonous mushroom
in Thailand. Moreover, and toxicology center staffs (NIH: Department of Medical
Sciences, Ministry of Public Health) for their suggestion and knowledge of peptide
toxins identification. D. Chakraborty, K. Das, A. Baghela, S.K. Singh, and B.T.M.
Dentinger thank the Director, Botanical Survey of India, Kolkata and Agharkar
Research Institute, Pune for providing facilities during this study. Two of them (D.
Chakraborty & K. Das) are thankful to the entire forest department of Sikkim for
allowing them to undertake the macrofungal exploration in the restricted subalpine
areas of North Sikkim. K. Das are indebted to Z.L. Yang (Chinese Academy of
Sciences, China) for his indispensible suggestions and literature help in this regard.
Field assistance rendered by S. Pradhan (BSI, Gangtok) is also duly acknowledged.
A.V. Marano, A.L. Jesus, J.I. de Souza, G.H. Jerônimo, T.Y. James, M.C. Boro,
S.C.O. Rocha, E.M. Leaño, M.J. Iribarren, and C.L.A. Pires-Zottarelli thank “Instituto
Florestal” for the permission for sampling at the PEIC, M.O.Neves Junior for his
valuable help during sampling, and to C.C. Aparecido, curator of the culture
collection MMBF, for accepting our voucher cultures. They also thank São Paulo
Research Foundation (FAPESP) for the fellowships given to A.L. Jesus (Process Nº
2013/01409-0) and for the financial support given to C.L.A. Pires-Zottarelli (Process
Nº 2012/50222-7), CAPES (“Coordenação de Aperfeiçoamento de Pessoal de Nível
Superior”) for the fellowship and support given to A.V. Marano (“Ciência Sem
Fronteiras” Program, “Atração de Jovens Talentos” DRI-CAPES Process Nº
006/2012) and CNPq (“Conselho Nacional de Desenvolvimento Científico e
Tecnológico”) for the grant given to C.L.A. Pires-Zottarelli (Process
Nº304411/2012-4). L.W. Zhou, Y.C. Dai and J. Vlasák thank O. Euatrakool and A.
Auetragul for their help in field trip. They also thank National Natural Science
Foundation of China (Project No. 31200015). T. Niskanen, K. Liimatainen, M. Beug,
and J. Ammirati are grateful to D.E. Stuntz Memorial Foundation. M.A.
Abdel-Wahab, A.H. Bahkali, E.B.G. Jones and F.A. Abdel-Aziz thank the National
Plan for Science, Technology and Innovation (MAARIFAH), King Abdulaziz City for
Science and Technology, Kingdom of Saudi Arabia, Award Number
(12-BIO2840-02). E. Kuhnert, E.B. Sir, and M. Stadler are grateful to A.I. Hladki for
her contribution to the collection of Argentinian specimens. They thank C. Lambert
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and S. Heitkämper for obtaining the cultures and performing the molecular work. M.
Rohde is thanked for SEM recordings. Funding from the DAAD and the Argentina
Ministerio de Ciencia, Tecnología e Innovación Productiva for an academic exchange
program involving E. Kuhnert, E.B. Sir and M. Stadler is gratefully acknowledged.
T.C. Wen, K.K. Hapuarachchi and K.D. Hyde thank the National Natural Science
Foundation of China (No. 31460012) and the Science Foundation of Guizhou
University (No. 201309). E. De Crop is supported by the “Special Research Fund
Ghent University” (BOF). The survey in Thailand was part of the Northern Thailand
mushroom diversity workshop prior to the 10th International Mycological Congress
and was financially supported by the Research Foundation Flanders (FWO, grant
K1A7614N). F. Hampe is thanked by M. Verbeken for conducting laboratory work. K.
Tanaka would like to thank the Japan Society for the Promotion of Science (JSPS,
25440199 and 26291084) and Hirosaki University Grant for Exploratory Research by
Young Scientists and Newly appointed Scientists for financial support. B.K. Cui, J.
Song and J.J. Chen are grateful to H.S. Yuan (IFP, China) for loan of specimens. The
research was supported by the National Natural Science Foundation of China (Project
No. 31422001). B. Thongbai was financial supported by the Royal Golden
Jubilee-Industry Ph.D (Ph.D/0138/2553 in 4.S.MF/53/A.3). We thank the
International Research Group Program (IRG-14-27), Deanship of Scientific Research,
King Saud University, Saudi Arabia for partially supporting this research. K.D. Hyde
thanks the Chinese Academy of Sciences, [project number 2013T2S003], for the
award of Visiting Professorship for Senior International Scientists at Kunming
Institute of Botany. MFLU [grant number 56101020032] is thanked for supporting
studies on Dothideomycetes. We are grateful to the Mushroom Research Foundation,
Chiang Rai, Thailand. C.G. Lin is grateful to J.Z. Sun (Mae Fah Luang University,
Thailand) for comments on the manuscript and S.F. Ran (Guizhou University,
Guizhou, China) for assistance in molecular work, and to the support by the National
Natural Science Foundation of China (No. NSFC 31560489). N.N. Wijayawardene
thanks Guizhou University for helping to carryout DNA sequencing. T.T.T. Nguyen,
S.J. Jeon, H.S. Lee, P.M. Kirk, K. Voigt, and H.B. Lee were 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 of the
Ministry of Environment (MOE), Republic of Korea. M. 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. C. Phukhamsakda would like to thank the Royal Golden
Jubilee PhD Program under Thailand Research Fund, for the award of a scholarship
no. PHD/0020/2557 to study towards a PhD. M. Gorczak was supported by Polish
Ministry of Science and Higher Education under grant no. DI2014012344. J.
Pawłowska was partially supported by the National Science Center of Poland under
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grant no. 2015/17/D/NZ8/00778. M. Abdel-Wahab was funded by the National Plan
for Science, Technology and Innovation (MAARIFAH), King Abdulaziz City for
Science and Technology, Kingdom of Saudi Arabia, Award Number
(12-BIO2840-02). J. Zhao, G. Consiglio, P. Alvarado, S.D. Yang, L. Setti, Y. Hu, A.
Vizzini, and L.P. Tang wish to express their sincere gratitude to Swiss mycologist E
Musumeci (Basel, Switzerland) for providing valuable information (including
photographs) on Musumecia vermicularis. They are grateful to J.F. Liang (Research
Institute of Tropical Forestry, Chinese Academy of Forestry) for helping to scan
basidiospores. This work was financially supported by the National Natural Science
Foundation of China (No. 31560004), Yunnan applied basic research projects-joint
special project (No. 2014FB016), the Science Research Foundation of department of
education, Yunnan Province (No. 2015Y147), and the Open Research Foundation of
Yunnan Key Laboratory of Pharmacology for Natural Products (No. 2015G003). G.
Alves-Silva, A. Góes-Neto and E.R. Drechsler-Santos thank the Parque Natural
Municipal São Francisco de Assis for permission to sample collections and F.
Bittencourt for specimen collected and pictures in situ; herbaria mentioned (FURB
and FLOR); Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES)
for providing a master’s scholarship to G. Alves-Silva; Fiocruz for performing the
molecular sequencing; PPGFAP/UFSC and BrBOL for partial financing of the
research.
M.A.
Reck
thanks
CAPES
(PNPD
Institucional
2011—23038.007790/2011-93) for scholarship and funding. P. Chomnunti would like
to thank the National Research Council of Thailand (NRCT), for the project
“Biodiversity, phylogeny and biological activity of Dothideomycetes” grant number
58201020010 and Thailand Research Fund (TRF) grant number TRG5780008 for
partial funding. R. Phookamsak sincerely appreciates The Royal Golden Jubilee Ph. D.
Program (PHD/0090/2551) under the Thailand Research Fund for financial support.
S.C. Karunarathna would like to thank the World Agroforestry Centre, East and
Central Asia Office, and the Kunming Institute of Botany for financial support,
Humidtropics, a CGIAR Research Program that aims to develop new opportunities for
improved livelihoods in a sustainable environment and the National Research Council
of Thailand (NRCT), projects - Taxonomy, phylogeny and cultivation of Lentinus
species in northern Thailand (NRCT/55201020007) are thanked, for partially funding.
K.V. Solomon, J.K. Henske, C.H. Haitjema, S.P. Gilmore, M.K. Theodorou, and M.A.
O’Malley thanks the Office of Science (BER), U.S. Department of Energy
(DE-SC0010352) and the Institute for Collaborative Biotechnologies through grant
W911NF-09-0001, and the Mellichamp Academic Initiative in Sustainability at UC
Santa Barbara. A portion of their research was performed under the JGI-EMSL
Collaborative Science Initiative and used resources at the DOE Joint Genome Institute
and the Environmental Molecular Sciences Laboratory, which are DOE Office of
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Science User Facilities. Both facilities are sponsored by the Office of Biological and
Environmental Research and operated under Contract Nos. DE-AC02-05CH11231
(JGI) and DE-AC05-76RL01830 (EMSL).
References
Abdel-Wahab MA, Jones EBG (2000) Three new marine ascomycetes from driftwood
in Australian sand dunes. Mycoscience 41:379–388
Abdel-Wahab MA, Hodhod MS, Bahkali AHA, Jones EBG (2014) Marine fungi of
Saudi Arabia. Bot Mar 57:323–335
Abe A, Oda Y, Asano Y, Sone T (2007) Rhizopus delemar is the proper name for
Rhizopus oryzae fumaric-malic acid producers. Mycologia 99:714–722
Abe A, Oda Y, Asano Y, Sone T (2010) A molecular phylogeny based taxonomy of
the genus Rhizopus. Biosci Biotechnol Biochem 74:1325–1331
Abdollahzadeh J, Javadi A, Zare R, Phillips AJL (2014) A phylogenetic study of
Dothiorella and Spencermartinsia species associated with woody plants in Iran,
New Zealand, Portugal and Spain. Persoonia 32:1–12
Acero FJ, González V, Sánchez-Ballesteros J, Rubio V, Checa J, Bills GF, Salazar O,
Platas G, Peláez F (2004) Molecular phylogenetic studies on the Diatrypaceae
based on rDNA-ITS sequences. Mycologia 96:249–259
Adams GC, Surve-Iyer RS, Iezzoni AF (2002) Ribosomal DNA sequence divergence
and group I introns within the Leucostoma species L. cinctum, L. persoonii, and
L. parapersoonii sp. nov., ascomycetes that cause Cytospora canker of fruit trees.
Mycologia 94:947–967
Adams GC, Wingfield MJ, Common R, Roux J (2005) Phylogenetic relationships and
morphology of Cytospora species and related teleomorphs (Ascomycota,
Diaporthales, Valsaceae) from Eucalyptus. Stud Mycol 52:1–144
Adams GC, Roux J, Wingfield MJ (2006) Cytospora species (Ascomycota,
Diaporthales, Valsaceae): introduced and native pathogens of trees in South
Africa. Australasian Plant Pathology 35:521–548
Adl SM, Simpson AGB, Lane CE, Lukeš J, Bass D, Bowser SS, Brown MW, Burki F,
Dunthorn M, Hampl V, Heiss A, Hoppenrath M, Lara E, Le Gall L, Lynn DH,
McManus H, Mitchell EA, Mozley-Stanridge SE, Parfrey LW, Pawlowski J,
Rueckert S, Shadwick RS, Schoch CL, Smirnov A, Spiegel FW (2012). The
revised classification of Eukaryotes. J Eukaryot Microbiol 59:429–493
Ahmed SA, van de Sande WWJ, Stevens DA, Fahal A, van Diepeningen AD, Menken
SBJ, de Hoog GS (2014) Revision of agents of black-grain eumycetoma in the
order Pleosporales. Persoonia 33:141–154
Alastruey-Izquierdo A, Hoffmann K, de Hoog GS, Rodriguez-Tudela JL, Voigt K,
Bibashi E, Walther G (2010) Species recognition and clinical relevance of the
zygomycetous genus Lichtheimia (syn. Absidia pro parte, Mycocladus). J Clin
Microbiol 48:2154–2170
Altschul SF, Gish W, Miller W, Myers EW, Lipman DJ (1990) Basic local alignment
search tool. Journal of Molecular Biology. 215:403–410
8292
8293
8294
8295
8296
8297
8298
8299
8300
8301
8302
8303
8304
8305
8306
8307
8308
8309
8310
8311
8312
8313
8314
8315
8316
8317
8318
8319
8320
8321
8322
8323
8324
8325
8326
8327
8328
8329
8330
8331
8332
8333
8334
Álvarez E, Cano J, Stchigel AM, Sutton DA, Fothergill AW, Salas V, Rinaldi MG,
Guarro J (2011) Two new species of Mucor from clinical samples. Med Mycol
49:62–72
Alves MH, Campos-Takaki GM, Porto ALF, Milanez AI (2002) Screening of Mucor
spp. for the production of amylase, lipase, polygalacturonase and protease. Braz J
Microbiol 33:325–330
Amalfi M, Yombiyeni P, Decock C (2010) Fomitiporia in sub-Saharan Africa:
morphology and multi-gene phylogenetic analysis support three new species
from the Guineo-Congolian rainforest. Mycologia 102(6):1303–1317
Amalfi M, Decock C (2013) Fomitiporia castilloi sp. nov. and multiple clades around
F. apiahyna and F. texana in Meso- and South America evidenced by multi-loci
phylogenetic inferences. Mycologia 105(4):873-887
Amalfi M, Raymundo T, Valenzuela R, Decock C (2012) Fomitiporia cupressicola sp.
nov., a parasite on Cupressus arizonica, and additional unnamed clades in the
southern USA and northern Mexico, determined by multi-locus phylogenetic
analyses. Mycologia 104(4):880–893
Amalfi M, Robledo G, Decock C (2014) Fomitiporia baccharidis comb. nov., a little
known species from high elevation Andean forests and its affinities within the
Fomitiporia Neotropical lineages. Mycological Progress. 13:995
Anon (1955) Shen Nong materia medica 102–200 A.D (E. Han). Reprinted. People
Hygiene Press, Beijing
Aptroot A (1995). Redisposition of some species excluded from Didymosphaeria
(Ascomycotina). Nova Hedwigia (60):325–379
Aptroot A (1998) A world revision of Massarina (Ascomycota). Nova Hedwigia
66:89–162
Ariyawansa HA, Tanaka K, Thambugala KM, Phookamsak R, Tian Q, Camporesi E,
Hongsanan S, Monkai J, Wanasinghe DN, Chukeatirote E, Kang JC, Xu JC,
McKenzie EHC, Jones EBG, Hyde KD (2014a) A molecular phylogenetic
reappraisal of the Didymosphaeriaceae (= Montagnulaceae). Fungal Divers
68:69–104
Ariyawansa H.A., Phookamsak R., Tibpromma S., Kang J.C., Hyde K.D (2014b) A
molecular and morphological reassessment of Diademaceae. The Scientific
World Journal Article ID 675348, 11 pages
Ariyawansa HA, Hawksworth DL, Hyde KD, Jones EBG, Maharachchikumbura SSN,
Manamgoda DS, Thambugala KM, Udayanga D, Camporesi E, Daranagama A,
Jayawardena R, Liu JK, McKenzie EHC, Phookamsak R, Senanayake IC,
Shivas RG, Tian Q, Xu JC (2014c) Epitypification and neotypification:
guidelines with appropriate and inappropriate examples. Fungal Divers
69(1):57 – 91
Ariyawansa HA, Thambugala KM, Manamgoda DS, Jayawardena R., Camporesi E,
Boonmee S, Wanasinghe DN, Phookamsak R, Hongsanan S, Singtripop C,
Chukeatirote E, Kang JC, Gareth Jones EB, Hyde KD (2015a) Towards a natural
classification and backbone tree for Pleosporaceae. Fungal Diversity 71:85–139
8335
8336
8337
8338
8339
8340
8341
8342
8343
8344
8345
8346
8347
8348
8349
8350
8351
8352
8353
8354
8355
8356
8357
8358
8359
8360
8361
8362
8363
8364
8365
8366
8367
8368
8369
8370
8371
8372
8373
8374
8375
8376
8377
8378
Ariyawansa HA, Hyde KD, Jayasiri SC, Buyck B, Chethana KWT, Dai DQ, Dai YC,
Daranagama DA, Jayawardena RS, Lücking R, Ghobad-Nejhad M, Niskanen T,
Thambugala KM, Voigt K, Zhao RL, Li GJ, Doilom M, Boonmee S, Yang ZL,
Cai Q, Cui YY, Bahkali AH, Chen J, Cui BK, Chen JJ, Dayarathne MC,
Dissanayake AJ, Ekanayaka AH, Hashimoto A, Hongsanan S, Jones EBG,
Larsson E, Li WJ, Li QR, Liu JK, Luo ZL, Maharachchikumbura SSN, Mapook
A, McKenzie EHC, Norphanphoun C, Konta S, Pang KL, Perera RH,
Phookamsak R, Phukhamsakda C, Pinruan U, Randrianjohany E, Singtripop C,
Tanaka K, Tian CM, Saowaluck Tibpromma, Mohamed A. Abdel-Wahab,
Dhanushka N. Wanasinghe, Nalin N. Wijayawardene, Zhang JF, Zhang H,
Abdel-Aziz FA, Wedin M, Westberg M, Ammirati JF, Bulgakov TimurS, Lima
DX, Callaghan TM, Callac P, Chang CH, Coca LF, Dal-Forno M, Dollhofer V,
Fliegerová K, Greiner K, Griffith GW, Ho HM, Hofstetter V, Jeewon R, Kang J
C, Wen TC, Kirk PM, Kytövuori I, Lawrey JD, Xing J, Li H, Liu ZY, Liu XZ,
Liimatainen K, H. Lumbsch T, Matsumura M, Moncada B, Nuankaew S,
Parnmen S, Santiago ALCMDA, Sommai S, Song Y, de Souza CAF, de
Souza-Motta CM, Su HY, Suetrong S, Wang Y, Wei S Fong, Yuan HS, Zhou
LW, Réblová M, Fournier J, Camporesi E, Luangsa-ard JJ, Tasanathai K,
Khonsanit A, Thanakitpipattana D, Somrithipol S, Diederich P, Millanes AM,
Common RS, Stadler M, Yan JY, Li XH, Lee HW, Nguyen TTT, Lee HB,
Battistin E, Marsico O, Vizzini A, Vila J, Ercole E, Eberhardt U, Simonini G,
Wen HA, Chen XH, Miettinen O, Spirin V, Hernawati (2015b) Fungal diversity
notes 111–252—taxonomic and phylogenetic contributions to fungal taxa.
Fungal Diversity 75:27–274
Ariyawansa HA, Phukhamsakda C, Thambugala KM, Wanasinghe DN, Perera RH,
Mapook A, Camporesi E, Kang JC, Jones EBG, Bahkali AH, Bhat JD, Hyde KD
(2015c) Revision and phylogeny of Leptosphaeriaceae. Fungal Divers 74(1):19 –
51
Arx JA von (1971) Testudinaceae, a new family of Ascomycetes. Persoonia
6:365–369
Arx JA von, Müller E (1975). A re-evaluation of the bitunicate Ascomycetes with
keys to families and genera. Studies in Mycology 9:1–159
Balci Y, Balci S, Eggers J, MacDonald WL, Juzwik J, Long RP, Gottschalk KW
(2007). Phytophthora spp. associated with forest soils in eastern and
north-central US oak ecosystems. Plant Dis 91:705–710
Barber PA, Crous PW, Groenewald JZ, Pascoe IG, Keane P (2011) Reassessing
Vermisporium (Amphisphaeriaceae), a genus of foliar pathogens of Eucalypts.
Persoonia 27:90–118
Barghoorn ES, Linder DH (1944) Marine fungi: their taxonomy andbiology. Farlowia
1:395–467
Barr ME (1972) Preliminary studies on the Dothideales in temperate North America.
Contributions from the University of Michigan Herbarium 9:523–638
Barr ME (1978) The Diaporthales in North America with emphasis on Gnomonia and
its segregates. Mycol Mem7:1–232
8379
8380
8381
8382
8383
8384
8385
8386
8387
8388
8389
8390
8391
8392
8393
8394
8395
8396
8397
8398
8399
8400
8401
8402
8403
8404
8405
8406
8407
8408
8409
8410
8411
8412
8413
8414
8415
8416
8417
8418
8419
8420
8421
Barr ME (1987) Prodomus to class Loculoascomycetes. Publ. by the author, Amherst
Barr ME (1989) The genus Chaetomastia (Dacampiaceae) in North America.
Mycotaxon 34:507–515
Barr ME (1979a) A classification of Loculoascomycetes. Mycologia 71:935–957
Barr ME (1979b) On the Massariaceae in North America. Mycotaxon 9:17–37
Barr ME (1990) North American Flora. Melanommatales (Loculoascomycetes). New
York Botanical Garden Series II Part 13:129
Barr ME (2001) Revisionary studies on the Dothioraceae. Harvard Papers in Botany
6:25–34
Barr DJS, Kudo H, Jakober KD, Cheng KJ (1989) Morphology and development of
rumen fungi: Neocallimastix sp., Piromyces communis, and Orpinomyces bovis
gen. nov., sp. nov. Can. J. Bot 67:2815–2824
Bas C (1969) Morphology and subdivision of Amanita and monograph of its section
Lepidella. Persoonia 5:285–579
Beakes GW, Honda D, Thines M (2014). Systematics of the Straminipila:
Labyrinthulomycota, Hyphochytriomycota, and Oomycota. In: McLaughlin DJ,
Spatafora JW (eds.) TheMycota VII Part A – Systematics and Evolution.
Springer-Verlag, Berlin Heidelberg, pp 39–97
Benson DA, Cavanaugh M, Clark K, Karsch-Mizrachi I, Lipman DJ, Ostell J, Sayers
EW. (2013) GenBank. Nucleic Acids Research 41:D36–42
Berkeley MJ (1847) Decades of fungi XII–XIV. Lond J Bot 6:312–326
Berkeley MJ, Broome CE (1852) Notices of British Fungi (615-639). Annals and
Magazine of Natural History 9:317–329
Bernicchia A, Gorjón SP (2010) Fungi Europaei 12 - Corticiaceae s.l. Edizioni
Candusso, Italy
Benny GL (1995) Classical morphology in Zygomycetes taxonomy. Can J Bot
73:S725–S730
Benny GL (2008) The methods used by Dr. R. K. Benjamin, and other mycologists, to
isolate Zygomycetes. Aliso 26:37–61
Benny GL (2013) Zygomycetes. Published on the internet at www.zygomycetes.org
Bessette AE, Roody WC, Bessette AR (2010) North American Boletes: A colour
guide to the Fleshy Pored Mushrooms. First paperback edition. Syracuse
University Press, USA, 396
Binder M, Hibbett DS, Wang Z, Farnham W (2006) Evolutionary relationships of
Mycaureola dilseae (Agaricales), a basidiomycete pathogen of a subtidal
rhodophyte. American Journal of Botany 93:547–556
Binder M, Larsson KH, Matheny PB, Hibbett D (2010) Amylocorticiales ord. nov.
and Jaapiales ord. nov.: Early diverging clades of Agaricomycetidae dominated
by corticioid forms. Mycologia 102:865–880
Binder M, Justo A, Riley R, Salamov A, Lopez-Giraldez F, Sjokvist E, Copeland A,
Foster B, Sun H, Larsson E, Larsson KH, Townsend J, Grigoriev IV, Hibbett DS
(2013) Phylogenetic and phylogenomic overview of the Polyporales. Mycologia
105:1350–1373
8422
8423
8424
8425
8426
8427
8428
8429
8430
8431
8432
8433
8434
8435
8436
8437
8438
8439
8440
8441
8442
8443
8444
8445
8446
8447
8448
8449
8450
8451
8452
8453
8454
8455
8456
8457
8458
8459
8460
8461
8462
8463
8464
8465
Blair JE, Coffey MD, Park SY, Geiser DM, Kang S (2008). A multi-locus phylogeny
for Phytophthora utilizing markers derived from complete genome sequences.
Fungal Genet Biol 45:266–277
Boehm EW, Schoch CL, Spatafora JW (2009a). On the evolution of the Hysteriaceae
and Mytilinidiaceae (Pleosporomycetidae, Dothideomycetes, Ascomycota) using
four nuclear genes. Mycol Res 113(4), 461–479.
Boehm EWA, Mugambi G, Miller AN, Huhndorf S, Marincowitz S, Schoch CL,
Spatafora JW (2009b) A molecular phylogenetic reappraisal of the Hysteriaceae,
Mytilinidiaceae and Gloniaceae (Pleosporomycetidae, Dothideomycetes) with
keys to world species. Stud Mycol 64:49–83
Boidin J, Gilles G (1998) Contribution à l'étude des genres Dendrocorticium,
Dendrodontia et Dentocorticium (Basidiomycotina). Cryptogam Mycol
19(3):181–202
Bojantchev D (2013) Cortinarius of California: eight new species in subg. Telamonia.
Mycotaxon 123:375–402
Bojantchev D (2015) Nomenclatural novelties, Cortinarius. Index Fungorum no. 247
Bojantchev D, Davis RM (2011) Cortinarius xanthodryophilus sp. nov. – a common
Phlegmacium under oaks in California. Mycotaxon 116:317–328
Boonmee S, Hyde KD, KoKo TW, Chukeatirote E, Chen H, Cai L, McKenzie EHC,
Jones EBG, Kodsueb R, Hassan BA (2012) Two new Kirschsteiniotheliaceae
species with Dendryphiopsis anamorphs cluster in Kirschsteiniotheliaceae fam.
nov. Mycologia 104:698–714
Bonorden HF (1864) Abhandlungen aus dem Gebiete der Mykologie. Abhandlungen
der Naturforschenden Gesellschaft zu Halle. 8:1–168
Borelli D (1959) Pyrenochaeta romeroi n. sp. Dermatologia Venezplana 1:325–326
Bose SK (1961) Studies on Massarina Sacc. and related genera. Phytopathol Z
41:151–213
Botella L, Diez JJ (2011) Phylogenic diversity of fungal endophytes in Spanish stands
of Pinus halepensis. Fungal Divers 47(1):9–18
Botella L, Santamaría O, Diez JJ (2010). Fungi associated with the decline of Pinus
halepensis in Spain. Fungal Divers 40(1):1–11
Brasier CM, Beales PA, Kirk SA, Denman S, Rose J (2005). Phytophthora kernoviae
sp. nov., an invasive pathogen causing bleeding stem lesions on forest trees and
foliar necrosis of ornamentals in the UK. Mycol Res 109:853–859
Braune RA (1913) Untersuchungen über die im Wiederkäuermagen vorkommenden
Protozoen. Arch. Für Protistenkd 32:111–170
Bresinsky A, Besl H (2003) Beiträge zu einer Mykoflora Deutschlands: Schlüssel zur
Gattungsbestimmung der Blätter-, Leisten- und Röhrenpilze: mit
Literaturhinweisen zur Artbestimmung. Regensburger mykologische Schriften.
Regensburg: Verlag der Gesellschaft
Breton A, Dusser M, Gaillard-Martine B, Guillot J, Millet L, Prensier G (1991)
Piromyces rhizinflata nov. sp., a strictly anaerobic fungus from faeces of the
Saharian ass: a morphological, metabolic and ultrastructural study. FEMS
Microbiol Lett 82:1–8
8466
8467
8468
8469
8470
8471
8472
8473
8474
8475
8476
8477
8478
8479
8480
8481
8482
8483
8484
8485
8486
8487
8488
8489
8490
8491
8492
8493
8494
8495
8496
8497
8498
8499
8500
8501
8502
8503
8504
8505
8506
8507
8508
8509
Brookman JL, Mennim G, Trinci APJ, Theodorou MK, Tuckwell DS (2000)
Identification and characterization of anaerobic gut fungi using molecular
methodologies based on ribosomal ITS1 and 18S rRNA. Microbiology
146:393–403
Budziszewska J, Piatkowska J (2010) Taxonomic position of Mucor hiemalis f. luteus.
Mycotaxon 111:75–85
Bulgakov TS (2010) Microfungi of Leucostoma and Valsa genera and their Cytospora
anamorphs on arboreal plants in the steppe zone of Southern Russia // Actual
problems of ecology: Mater. IV All-Russian Scien. Conf. in Vladikavkaz, North
Ossetian State University:40–45 (in Russian)
Butin H, Holdenrieder O, Sieber TN (2013) The complete life cycle of Petrakia
echinata. Mycol Prog 12(2):427–435
Buyck B (1989) Revision du genre Russula Persoon en Afrique Centrale. PhD
dissertation, Rijksuniversiteit Gent
Buyck B, Hoyak E (1999) New taxa of pleurotoid Russulaceae. Mycologia
91(3):532–537
Buyck B, Thoen D, Watling R (1996) Ectomycorrhizal fungi of the Guinea-Congo
region. Proceedings of the Royal Society of Edinburgh. Section B. Biological
Sciences 104:313–333
Buyck B, Hofstetter V, Eberhardt U, Verbeken A, Kauff F (2008) Walking the thin
line between Lactarius and Russula: the dilemma of Russula sect.
Ochricompactae. Fungal Divers 28:15–40
Buyck B, Hofstetter V, Verbeken A, Walleyn R (2010) Proposal 1919: To conserve
Lactarius nom. cons. (Basidiomycota) with a conserved type. Mycotaxon
111:504–508
Cáceres MES, Aptroot A, Parnmen S, Lücking R (2014) Remarkable diversity of the
lichen family Graphidaceae in the Amazon rain forest of Rondônia, Brazil.
Phytotaxa 189:87–136
Cai L, Hyde KD, Taylor PWJ,Weir BS,Waller J, AbangMM, Zhang JZ, Yang YL,
Phoulivong S, Liu ZY, Prihastuti H, Shivas RG, McKenzie EHC, Johnston PR
(2009) A polyphasic approach for studying Colletotrichum. Fungal Divers
39:183–204
Cai Q, Tulloss RE, Tang LP,Tolgor B, Zhang P, Chen ZH, Yang ZL (2014)
Multi-locus phylogeny of lethal amanitas: Implications for species diversity and
historical biogeography. BMC Evol Biol 14(143):1–16
Cannon PF, Damm U, Johnston PR, Weir BS (2012) Colletotrichum-current status
and future directions. Stud Mycol 73:181–213
Cao Y, Wu SH, Dai YC. (2012) Species clarification of the prize medicinal
Ganoderma mushroom ‘‘Lingzhi’’ Fungal Divers 56:49–62
Cao Y, Yuan. HS (2013) Ganoderma mutabile sp. nov. from southwestern China
based on morphological and molecular data Mycol Progress 12:121–126
Castlebury LA, Rossman AY, Jaklitsch WJ, Vasilyeva LN (2002) A preliminary
overview of the Diaporthales based on large subunit nuclear ribosomal DNA
sequences. Mycologia 94:1017–1031
8510
8511
8512
8513
8514
8515
8516
8517
8518
8519
8520
8521
8522
8523
8524
8525
8526
8527
8528
8529
8530
8531
8532
8533
8534
8535
8536
8537
8538
8539
8540
8541
8542
8543
8544
8545
8546
8547
8548
8549
8550
8551
8552
Castlebury LA, Rossman AY, Sung GH, Hyten AS, Spatafora JW (2004) Multi-gene
phylogeny reveals new lineage for Stachybotrys chartarum, the indoor air fungus.
Mycol. Res. 108(8):864–872
Ceska O (2013) The Macrofungi of Observatory Hill: Long–term Survey and
Inventory
November
2004
–
March
2013.
http://www.goert.ca/activities/2013/05/macrofungi–observatory–hill/
Chadefaud M (1960) Les végétaux non vasculaires (Cryptogamie). In: Chadefaud M,
Emberger L (eds) Traité de botanique systématique, vol 1. Tome I, Masson et
Cie, Paris, p 1016
Chaverri P, Liu M, Hodge KT (2008) A monograph of the entomopathogenic genera
Hypocrella, Moelleriella and Samuelsia gen. nov. (Ascomycota, Hypocreales,
Clavicipitaceae) and their aschersonia-like anamorphs in the Neotropics. Stud
Mycol 60:1–66
Chen GQ, Zheng RY (1986) A new species of Mucor with giant spores. Acta Mycol
Sin 1:56–60
Chen J, Zhao RL, Parra LA, Guelly AK, Kesel AD, Rapion S, Hyde KD, Chukeatirote
E, Callac P. (2015a) Agaricus section Brunneopicti: a phylogenetic
reconstruction with descriptions of four new taxa. Phytotaxa 192:145–168.
Chen J, Zhao RL, Karunarathna SC, Callac P, Raspe´ O, Bahkali AH, Hyde KD (2012)
Agaricus megalosporus: a new speciesin section Minores. Cryptogam Mycol
33:145–155.
Chen JJ, Cui BK, He SH, Cooper JA, Barrett MD, Chen JL, Song J, Dai YC (2016)
Molecular phylogeny and global diversity of the remarkable genus Bondarzewia
(Basidiomycota, Russulales). Mycologia (In press)
Chen YC, Hseu RS, Chien CY (2002) Piromyces polycephalus (Neocallimastigaceae),
a new rumen fungus. Nova Hedwig 75:409–414
Chen ZH, Dai YD, Yu H, Yang K, Yang Z L, Yuan F, Zeng WB (2013) Systematic
analyses of Ophiocordyceps lanpingensis sp. nov., a new species of
Ophiocordyceps in China. Microbiol Res 168(8):525–532
Chethana T, Liu M, Ariyawansa HA, Konta S, Wanasinghe DN, Zhou Y, Yan J,
Camporesi E, Bulgakov TS, Chukeatirote E, Hyde KD, Bahkali AH, Liu J, Li X
(2015)
Splanchnonema-like
species
in
Pleosporales:
Introducing
Pseudosplanchnonema gen. nov. in Massarinaceae. Phytotaxa, 231(2):133–144
Chevallier FF (1826) Flore Générale des Environs de Paris 1:1–674
Ciferri R (1958) Manuale di micologia medica. edn 2 (Pavia) 1:1–178
Clements FE (1909). The Genera of Fungi (1st ed.). Minneapolis, Minnesota: H.W.
Wilson. 114pp
Cooke DEL, Drenth A, Duncan JM, Wagels G, Brasier CM (2000). A molecular
phylogeny of Phytophthora and related Oomycetes. Fungal Genet Biol 30:17–32
Corda ACI (1833) Die Pilze Deutschlands. In: Sturm J (ed) Deutschlands Flora in
Abbildungen nach der Natur mit Beschreibungen. Sturm, Nürnberg 12:33–64
Corner EJH (1981) The agaric genera Lentinus, Panus, and Pleurotus with particular
reference to Malaysian species. Nova Hedwig Beih 69:1–169
8553
8554
8555
8556
8557
8558
8559
8560
8561
8562
8563
8564
8565
8566
8567
8568
8569
8570
8571
8572
8573
8574
8575
8576
8577
8578
8579
8580
8581
8582
8583
8584
8585
8586
8587
8588
8589
8590
8591
8592
8593
8594
8595
Crous PW, Palm ME (1999) Reassessment of the anamorph genera Botryodiplodia,
Dothiorella and Fusicoccum. Sydowia 51:167–175
Crous PW, Verkley GJM, Christensen M, Castañeda-Ruiz RF, Groenewald JZ (2012)
How important are conidial appendages? Persoonia 28:126–137
Crous PW, Wingfield MJ, Schumacher RK, Summerell BA, Giraldo A, Gené J,
Guarro J, Wanasinghe DN, Hyde KD, Camporesi E, Gareth Jones EB,
Thambugala KM, Malysheva EF, Malysheva VF, Acharya K, Álvarez J,
Alvarado P, Assefa A, Barnes CW, Bartlett JS, Blanchette RA, Burgess TI,
Carlavilla JR, Coetzee MP, Damm U, Decock CA, den Breeÿen A, de Vries B,
Dutta AK, Holdom DG, Rooney-Latham S, Manjón JL, Marincowitz S,
Mirabolfathy M, Moreno G, Nakashima C, Papizadeh M, Shahzadeh Fazeli SA,
Amoozegar MA, Romberg MK, Shivas RG, Stalpers JA, Stielow B, Stukely MJ,
Swart WJ, Tan YP, van der Bank M, Wood AR, Zhang Y, Groenewald JZ
(2014a) Fungal Planet Description Sheets: 281–319. Persoonia 33: 212−289
Crous PW, Shivas RG, Quaedvlieg W, van der Bank M, Zhang Y, Summerell BA,
Guarro J, Wingfield MJ, Wood AR, Alfenas AC, Braun U, Cano-Lira JF, García
D, Marin-Felix Y, Alvarado P, Andrade JP, Armengol J, Assefa A, den Breeÿen
A, Camele I, Cheewangkoon R, De Souza JT, Duong TA, Esteve-Raventós F,
Fournier J, Frisullo S, García-Jiménez J, Gardiennet A, Gené J,
Hernández-Restrepo M, Hirooka Y, Hospenthal DR, King A, Lechat C, Lombard
L1, Mang SM, Marbach PA, Marincowitz S, Marin-Felix Y, Montaño-Mata NJ,
Moreno G, Perez CA, Pérez Sierra AM, Robertson JL, Roux J, Rubio E,
Schumacher RK, Stchigel AM, Sutton DA, Tan YP, Thompson EH, van der
Linde E, Walker AK, Walker DM, Wickes BL, Wong PT, Groenewald JZ
(2014b) Fungal Planet description sheets: 214–280. Persoonia 32:184–306
Crous PW, Schumacher RK, Wingfield MJ, Lombard L, Giraldo A, Christensen M,
Gardiennet A, Nakashima C, Pereira OL, Smith AJ, Groenewald JZ (2015a)
Fungal Systematics and Evolution: FUSE 1. Sydowia 67:81–118
Crous PW, Müller MM, Sánchez RM, Giordano L, Bianchinotti MV, Anderson FE,
Groenewald JZ (2015b) Resolving Tiarosporella spp. allied to
Botryosphaeriaceae and Phacidiaceae. Phytotaxa 202:073–093
Crous PW, Wingfield MJ, Guarro J, Hernández-Restrepo M, Sutton DA, Acharya K,
Barber PA, Boekhout T, Dimitrov RA, Dueñas M, Dutta AK, Gené J,
Gouliamova DE, Groenewald M, Lombard L, Morozova OV, Sarkar J, Smith
MTh, Stchigel AM, Wiederhold NP, Alexandrova AV, Antelmi I, Armengol J,
Barnes I, Cano-Lira JF, Castañeda Ruiz RF, Contu M, Courtecuisse PrR, da
Silveira AL, Decock CA, de Goes A, Edathodu J, Ercole E, Firmino AC, Fourie
A, Fournier J, Furtado EL, Geering ADW, Gershenzon J, Giraldo A, Gramaje D,
Hammerbacher A, He X-L, Haryadi D, Khemmuk W, Kovalenko AE,
Krawczynski R, Laich F, Lechat C, Lopes UP, Madrid H, Malysheva EF,
Marin-Felix Y, Martin MP, Mostert L, Nigro F, Pereira OL, Picillo B, Pinho DB,
Popov ES, Rodas Peláez CA, Rooney-Latham S, Sandoval-Denis M, Shivas RG,
Silva V, Stoilova-Disheva MM, Telleria MT, Ullah C, Unsicker SB, van der
8596
8597
8598
8599
8600
8601
8602
8603
8604
8605
8606
8607
8608
8609
8610
8611
8612
8613
8614
8615
8616
8617
8618
8619
8620
8621
8622
8623
8624
8625
8626
8627
8628
8629
8630
8631
8632
8633
8634
8635
8636
8637
8638
8639
8640
Merwe NA, Vizzini A, Wagner H-G, Wong PTW, Wood AR, Groenewald JZ
(2015c) Fungal planet Description Sheets: 320–370. Persoonia 34:167–266
Crous PW, Carris LM, Giraldo A, Groenewald JZ, Hawksworth DL,
Hernández-Restrepo M, Jaklitsch WM, Lebrun M-H, Schumacher RK, Stielow B,
van der Linde EJ, Vilcāne J, Voglmayr H, Wood AR (2015d) The Genera of
Fungi - fixing the application of the type species of generic names–G 2:
Allantophomopsis, Latorua, Macrodiplodiopsis, Macrohilum, Milospium,
Protostegia, Pyricularia, Robillarda, Rotula, Septoriella, Torula, and
Wojnowicia. IMA Fungus 6(1):163–198
Crous PW, Hawksworth DL, Wingfield MJ (2015e) Identifying and naming
plant-pathogenic fungi: past, present, and future. Annual Review of
Phytopathology 53: 247–267
Cui BK, Dai YC, He SH, Zhou LW, Yuan HS (2015) A novel Phellinidium sp. causes
laminated root rot on Qilian juniper (Sabina przewalskii) in Northwest China.
Plant Dis 99:39–43
Cunningham G H (1921) The genus Cordyceps in New Zealand. Trans Proc NZ Inst
53:372–382
Dai YC (2010) Hymenochaetaceae (Basidiomycota) in China. Fungal Divers
45:131–343
Dai, YC, Cui BK, Decock C (2008) A new species of Fomitiporia
(Hymenochaetaceae, Basidiomycota) from China based on morphological and
molecular characters. Mycol Res 112:375-380
Dai YC, Cui BK, Liu XY (2010) Bondarzewia podocarpi, a new and remarkable
polypore from tropical China. Mycologia 102:881–886
Dai YC, Cui BK, Yuan HS, Li BD (2007) Pathogenic wood-decaying fungi in China.
Forest Pathol 37:105–120
Dai YC, Yang ZL, Cui BK, Yu CJ, Zhou LW (2009) Species diversity and utilization
of medicinal mushrooms and fungi in China (Review). Int J Med Mushrooms
11:287–302
Damm U, Woudenberg JHC, Cannon PF, Crous PW (2009) Colletotrichum species
with curved conidia from herbaceous hosts. Fungal Divers 39:45–87
Das K, Chakraborty D (2014) Three interesting species of wild mushrooms from
Sikkim (India). Indian Journal of Plant Sciences 3(1): 101–108
Das K, Miller SL, Sharma JR (2005) Russula in Himalaya 1: A new species of
subgenus Amoenula. Mycotaxon 94:85–88
Das K, Miller SL, Sharma JR (2006a) Russula in Himalaya 2: Four new taxa.
Mycotaxon 95:205–215
Das K, Sharma JR, Atri NS (2006b) Russula in Himalaya 3: a new species of
subgenus Ingratula. Mycotaxon 95:271–275
Das K, Putte VDP, Buyck B (2010) New or interesting Russula from Sikkim
Himalaya (India). Cryptog Mycol 31:373–387
Das K, Atri NS, Buyck B (2013) Three new Russula (Russulales) from India.
Mycosphere 4:722–732
Das K, Dowie NJ, Li GJ, Miller SL (2014) Two new species of Russula (Russulales)
from India. Mycosphere 5:612–622
8641
8642
8643
8644
8645
8646
8647
8648
8649
8650
8651
8652
8653
8654
8655
8656
8657
8658
8659
8660
8661
8662
8663
8664
8665
8666
8667
8668
8669
8670
8671
8672
8673
8674
8675
8676
8677
8678
8679
8680
8681
8682
8683
de Almeida DAC, Gusmão LFP, Miller AN (2014) A new genus and three new
species of hysteriaceous ascomycetes from the semiarid region of Brazil.
Phytotaxa 176(1):298–308
De Bertoldi M, Lepidi AA, Nuti MP (1972) Classification of the genus Humicola
Traaen I. preliminary reports and investigations. Mycopathologia 46:289–304
De Crop E, Nuytinck J, Van de Putte K, Lecomte M, Eberhardt U, Verbeken A (2014)
Lactifluus piperatus (Russulales, Basidiomycota) and allied species in Western
Europe and a preliminary overview of the group worldwide. Mycol Prog 13
(3):493–511.
De Crop E, Nuytinck J, Van de Putte K, Wisitrassameewong K, Hackel J, Stubbe D,
Hyde KD, Roy M, Halling RE, Wang XH, Moreau PA, Eberhardt U, Verbeken
A (subm.) A multi-gene phylogeny of Lactifluus (Basidiomycota, Russulales)
translated into a new infrageneric classification of the genus. Mycol Prog (In
press)
De Gruyter JD, Woudenberg JHC, Aveskamp AA, Verkley GJM, Groenewald JZ,
Crous PW (2012) Redisposition ofPhoma-like anamorphs in Pleosporales. Stud
Mycol 75:1–36
Decock C, Figueroa SH, Robledo G, Castillo G (2007) Fomitiporia punctata
(Basidiomycota, Hymenochaetales) and its presumed taxonomic synonyms in
America: taxonomy and phylogeny of some species from tropical/subtropical
areas. Mycologia 99(5):733–752
Dexter Y, Cooke RC (1984) Fatty acids, sterols and catotenoids of the psychrophile
Mucor strictus and some mesophilic Mucor species. Trans Brit Mycol Soc
83:455–461
Doweld AB (2012) (2068–2070) Proposals to conserve Pertusariaceae against
Variolariaceae, Chrysothrichaceae against Pulverariaceae and Dothioraceae
against Saccotheciaceae (Fungi) with revision of dates of publication given in
Taxonomic Literature II. Taxon, 61(3):681–682
Du Y, Shi P, Huang H, Zhang X, Luo H, Wang Y, Yao B (2013) Characterization of
three novel thermophilic xylanase from Humicola insolens Y1 with application
potentials in the brewing industry. Bioresource Technology 130:161–167
Duhem B, Michel H (2009) Une espèce nouvelle de Corticium s.st. Études dans les
genres Dendrocorticium et Dentocorticium (Basidiomycotina). Cryptogam Mycol
30(2):161–179
Eberhardt U (2002) Molecular kinship analyses of the agaricoid Russulaceae:
correspondence with mycorrhizal anatomy and sporocarp features in the genus
Russula. Mycol Prog 1:201–223
Ellis MB (1971) Dematiaceous hyphomycetes. Commonwealth Mycological Institute,
Kew
Ertz D, Diederich P, Lawrev JD, Berger F, Freebury CE, Coppins B, Gardiennet A,
Hafellner J (2015) Phylogenetic insights resolve Dacampiaceae (Pleosporales)
as polyphyletic: Didymocyrtis (Pleosporales, Phaeosphaeriaceae) with
Phoma-like anamorphs resurrected and segregated from Polycoccum
8684
8685
8686
8687
8688
8689
8690
8691
8692
8693
8694
8695
8696
8697
8698
8699
8700
8701
8702
8703
8704
8705
8706
8707
8708
8709
8710
8711
8712
8713
8714
8715
8716
8717
8718
8719
8720
8721
8722
8723
8724
8725
8726
8727
(Trypetheliales,
Polycoccaceae
fam.
nov.)
Fungal
Divers
DOI
10.1007/s13225-015-0345-6
Fabre JH (1879). Essai sur les Sphériacées du Départment de Vaucluse. Annales des
Sciences Naturelles Botanique. 9:66–118
Fan XL, Liang YM, Ma R, Tian CM (2014a) Morphological and phylogenetic studies
of Cytospora (Valsaceae, Diaporthales) isolates from Chinese scholar tree, with
description of a new species. Mycoscience 55:252–259
Fan XL, Tian CM, Yang Q, Liang YM, You CJ, Zhang YB (2014b) Cytospora from
Salix in northern China. Mycotaxon 129:303–315
Fan XL, Hyde KD, Liu M, Liang YM, Tian CM (2015a) Cytospora species associated
with walnut canker disease in China, with description of a new species C.
gigalocus. Fungal Biol 119:310–319
Fan XL, Hyde KD, Yang Q, Liang YM, Ma R, Tian CM (2015b) Cytospora species
associated with canker disease of three antidesertification plants in northwestern
China. Phytotaxa 197:227–244
Farr DF, Rossman AY (2015) Fungal Databases, Systematic Mycology and
Microbiology Laboratory, ARS, USDA. Retrieved January 11, 2015, from
http://nt.ars−grin.gov/fungaldatabases/
Fell JW, Master IM (1975). Phycomycetes (Phytophthora spp. nov. and Pythium sp.
nov.) associated with degrading mangrove (Rhizophora mangle) leaves. Can J
Bot 53:2908–2922
Fernández FA, Huhndorf SM (2005) New species of Chaetosphaeria,
Melanopsammella and Tainosphaeria gen. nov. from the Americas. Fungal
Divers 18:15–57
Fernández FA, Miller AN, Huhndorf SM, Lutzoni FM, Zoller S (2006) Systematics of
the genus Chaetosphaeria and its allied genera: morphological and phylogenetic
diversity in north temperate and neotropical taxa. Mycologia 98:121–130
Ferreira JHS (1987) Dieback of grapevines in South Africa. PhD Thesis, University of
the Western Cape, South Africa.
Fidalgo O (1968) Phellinuspachyphloeus and its allies. Mem NY Bot Gard
17:109–147
Fliegerová K, Hodrová B, Voigt K (2004) Classical and molecular approaches as a
powerful tool for the characterization of rumen polycentric fungi. Folia
Microbiol (Praha) 49:157–164
Fliegerová K, Mrázek J, Hoffmann K, Zábranská J, Voigt K (2010) Diversity of
anaerobic fungi within cow manure determined by ITS1 analysis. Folia
Microbiol (Praha) 55:319–325
Fotouhifar KB, Hedjaroude GA, Leuchtmann A (2010) ITS rDNA phylogeny of
Iranian strains of Cytospora and associated teleomorphs. Mycologia
102:1369–1382
Fries EM (1823) Systema mycologicum vol 2. Greifswald, Germany. (in Latin)
Fries GE (1874) Hymenomycetes. Eur. Ed. Uppsala: Berling.
Frye CB, Reinhardt DJ (1993) Characterization of groups of the zygomycete genus
Rhizopus. Mycopathologia 124:139–147
8728
8729
8730
8731
8732
8733
8734
8735
8736
8737
8738
8739
8740
8741
8742
8743
8744
8745
8746
8747
8748
8749
8750
8751
8752
8753
8754
8755
8756
8757
8758
8759
8760
8761
8762
8763
8764
8765
8766
8767
8768
8769
8770
Fuckel L (1870) Symbolae mycologicae. Beiträge zur Kenntniss der Rheinischen
Pilze. Jb. nassau. Ver. Naturk. 23–24:212
Fujimura R, Nishimura A, Ohshima S, Sato Y, Nishizawa T, Oshima K, Hattori M,
Narisawa K, Ohta H (2014) Draft genome sequence of the
betaproteobacterialendosymbiont associated with the fungus Mortierella
elongata FMR23-6. Genome Announcements 2(6):e01272–14
Fungal Names (2016) http://124.16.146.175:8080/checklist/checklist.html. Accessed
on March 2016
Gaillard-Martinie B, Breton A, Dusser M, Julliand V (1995) Piromyces citronii sp.
nov., a strictly anaerobic fungus from the equine caecum: a morphological,
metabolic, and ultrastructural study. FEMS Microbiol Lett 130:321–326
Gams W (1976) Some new or noteworthy species of Mortierella. Persoonia
9(1):111–140
Gams W (1977) A key to the species of Mortierella. Persoonia 9(3):381–391
Garnica S, Weiss M, Walther G, Oberwinkler F (2007) Reconstructing the evolution
of agarics from nuclear gene sequences and basidiospore ultrastructure. Mycol
Res 111:1019–1029
Garnica S, Spahn P, Oertel B, Ammirati J, Oberwinkler F (2011) Tracking the
evolutionary history of Cortinarius species in section Calochroi, with
transoceanic disjunct distributions. BMC Evol Biol 11:213–231
Gelardi M, Vizzini A, Ercole E, Voyron S, Sun JZ, Liu XZ (2013) Boletus
sinopulverulentus, a new species from Shaanxi Province (central China) and
notes on Boletus and Xerocomus. Sydowia 65(1):45–57
Ghobad-Nejhad M, Nilsson RH, Hallenberg N (2010) Phylogeny and taxonomy of the
genus Vuilleminia (Basidiomycota) based on molecular and morphological
evidence, with new insights into Corticiales. Taxon 59(5):1519–1534
Gold JJ, Heath IB, Bauchop T (1988) Ultrastructural description of a new chytrid
genus of caecum anaerobe, Caecomyces equi gen. nov., sp. nov., assigned to the
Neocallimasticaceae. Biosystems 21:403–415
Gorjón SP, Saitta A (2014) Leptocorticium gloeocystidiatum sp. nov. (Corticiales,
Basidiomycota), a new corticioid fungus from Sicily, Italy. Mycosphere
5(3):406–409
Grove WB (1937) British stem-and leaf-fungi (Coelomycetes). Vol. II.
Gruninger RJ, Puniya AK, Callaghan TM, Edwards JE, Youssef N, Dagar SS,
Fliegerova K, Griffith GW, Forster R, Tsang A, McAllister T, Elshahed MS
(2014) Anaerobic Fungi (Phylum Neocallimastigomycota): Advances in
understanding of their taxonomy, life cycle, ecology, role, and biotechnological
potential. FEMS Microbiol Ecol 90:1–17
Guindon S, Gascuel O (2003) A Simple, fast, and accurate algorithm to estimate large
phylogenies by Maximum Likelihood. SystBiol 52:679–704
Guindon S, Dufayard JF, Lefort V, Anisimova M, Hordijk W, Gascuel O (2010) New
algorithms and methods to estimate maximumlikelihood phylogenies: assessing
the performance of PhyML 3.0. Syst Biol 59:307–321
8771
8772
8773
8774
8775
8776
8777
8778
8779
8780
8781
8782
8783
8784
8785
8786
8787
8788
8789
8790
8791
8792
8793
8794
8795
8796
8797
8798
8799
8800
8801
8802
8803
8804
8805
8806
8807
8808
8809
8810
8811
8812
8813
8814
Gvritishvili MN (1982) The fungal genus Cytospora in the USSR. (Izdatelstve
Sabchota Sakarstvelo: Tbilisi)
Hall T (2004) BioEdit. Ibis Therapeutics, Carlsbad, CA, 92008,
USA.(http://www.mbio.ncsu.edu/BioEdit/bioedit.html) (18 Mar 2005)
Hallenberg N (1980) New taxa of Corticiaceae from N. Iran (Basidiomycetes).
Mycotaxon 11(2):447–475
Hallenberg N (2012) Leptocorticium tenellum (Agaricomycetes) found in Chile.
Kurtziana 37(1):109–112
Hansen EM, Reeser PW, Sutton W (2012). Phytophthora borealis and Phytophthora
riparia, new species in Phytophthora ITS Clade 6.Mycologia 104:1133–1142
Hao YJ, Qin J, Yang ZL (2014) Cibaomyces, a new genus of Physalacriaceae from
East Asia. Phytotaxa 162:198–210
Hardham AR (2009). The asexual life cycle. In: Lamour K, Kamoun S (eds.)
Oomycete Genetics and Genomics: Diversity, Interactions, and Research Tools.
John Wiley & Sons, Hoboken, pp 93–119
Harkness HW (1884) Bulletin of the California Academy of Sciences. S.l.: The
Academy 1 (no. 1):page 45
Harrower E, Ammirati JF, Cappuccino AA, Ceska O, Kranabetter JM, Kroeger P,
Lim S, Taylor T, Berbee ML (2011) Cortinarius species diversity in British
Columbia and molecular phylogenetic comparison with European specimen
sequences. Botany 89(11):799–810
Hashimoto A, Sato G, Matsuda T, Hirayama K, Hatakeyama S, Harada Y, Shirouzu T,
Tanaka K (2015a) Molecular taxonomy of Dinemasporium and its allied genera.
Mycoscience 56:86–101
Hashimoto A, Sato G, Matsuda T, Matsumura M, Hatakeyama S, Harada Y, Tanaka
K (2015) Taxonomic revision of Pseudolachnea and Pseudolachnella and
establishment of Neopseudolachnella and Pseudodinemasporium gen. nov.
Mycologia 107:383–408
Hawksworth DL (1982) A new species of Caryospora from Eugenia in East Africa.
Trans Br Mycol Soc 79:69–74
Hawksworth DL, Eriksson O (1988). (895) – (906) Proposal to conserve 11 family
names in the Ascomycotina (Fungi). Taxon 37:190–193
He MQ, Zhao RL. (2015) A new species of Agaricus section Minores from China.
Mycology 6:182–186
Heath IB, Bauchop T, Skipp RA (1983) Assignment of the rumen anaerobe
Neocallimastix frontalis to the Spizellomycetales (Chytridiomycetes) on the
basis of its polyflagellate zoospore ultrastructure. Can J Bot 61:295–307
Heinemann P. (1978). Essai d'une clé de détermination des genres Agaricus et
Micropsalliota. Sydowia 30(1–6):6–37
Hermet A, Méheust D, Monunier J, Barbier G, Jany JL (2012) Molecular systematics
in the genus Mucor with special regards to species encountered in cheese. Fungal
Biol 116:692–705
Hesseltine CW, Ellis JJ (1961) Notes on Mucorales, especially Absidia. Mycologia
53:406–426
8815
8816
8817
8818
8819
8820
8821
8822
8823
8824
8825
8826
8827
8828
8829
8830
8831
8832
8833
8834
8835
8836
8837
8838
8839
8840
8841
8842
8843
8844
8845
8846
8847
8848
8849
8850
8851
8852
8853
8854
8855
8856
8857
8858
Hesseltine CW, Ellis JJ (1964) The genus Absidia: Gongronella and cylindrical
spored species of Absidia. Mycologia 56:568–601
Hesseltine CW, Ellis JJ (1966) Species of Absidia with ovoid sporangiospores.
Mycologia 58:761–785
Hibbett DS, Binder M, Bischoff JF, Blackwell M, Cannon PF, Eriksson OE,
Huhndorf S, James T, Kirk PM, Lucking R, Thorsten Lumbsch H, Lutzoni F,
Matheny PB, McLaughlin DJ, Powell MJ, Redhead S, Schoch CL, Spatafora JW,
Stalpers JA, Vilgalys R, Aime MC, Aptroot A, Bauer R, Begerow D, Benny GL,
Castlebury LA, Crous PW, Day Y-C, Gams W, Geiser DM, Griffith GW,
Gueidan C, Hawksworth DL, Hestmark G, Hosaka K, Humber RA, Hyde KD,
Ironside IE, Koljalg U, Kurtzman CP, Larsson K-H, Lichtwardt R, Longcore J,
Miądlikowska J, Miller A, Moncalvo J-M, Mozley-Standtridge S, Oberwinkler F,
Parmasto E, Reeb W, Rogers J-D, Roux C, Ryvarden L, Sampaio JP, Schussler
A, Sugiyama J, Thorn RG, Tibell L, Untereiner WA, Walker C, Wang Z, Weir A,
Weiss M, White MM, Winka K, Yao Y-J, Zhang N (2007) A higher- level
phylogenetic classification of the Fungi. Mycol Res 111:509–547
Hino I, Katumoto K (1958) On Chaetopatella, a new genus of the Excipulaceae. J Jpn
Bot 33:233–239
Hirayama K, Tanaka K, Raja HA, Miller AN, Shearer CA (2010) A molecular
phylogenetic assessment of Massarina ingoldianasensu lato. Mycologia
102:729–746
Ho HM, Chen ZC (1990) Morphological study of Gongronellabutleri (Mucorales)
from Taiwan. Taiwania 35:259–263
Ho HM, Chuang SC, Chen SJ (2004) Notes on Zygomycetes of Taiwan (IV): Three
Absidia species (Mucoraceae). Fung Sci 19:125–131
Ho YW, Barr DJS, Abdullah N, Jalaluldin S, Kudo H (1993a) A new species of
Piromyces from the rumen of deer in Malaysia. Mycotaxon 47:285-293
Ho YW, Barr DJS, Abdullah N, Jalaluldin S, Kudo H (1993b) Piromyces spiralis, a
new species of anaerobic fungus from the rumen of goat. Mycotaxon 48:59–68
Hodhod MS, Abdel-Wahab MA, Bahkali AH, Hyde KD (2012) Amergraphium
solium sp. nov. from Yanbu mangroves in the Kingdom of Saudi Arabia.
Cryptogam Mycol 33:285–294
Hoffmann K, Voigt K (2009) Absidia parricida plays a dominant role in biotrophic
fusion parasitism among mucoralean fungi (Zygomycetes): Lentamyces, a new
genus for A. parricida and A. zychae. Plant Biology 4:537–554
Hoffmann K, Discher S, Voigt K (2007) Revision of the genus Absidia (Mucorales,
Zygomycetes) based on physiological, phylogenetic, and morphological
characters; thermotolerant Absidia spp. form a coherent group, Mycocladiaceae
fam. nov. Mycol Res 111:1169–1183
Hoffmann K, Voigt K, Kirk P (2011) Mortierellomycotina subphyl. nov., based on
multi-gene genealogies. Mycotaxon 115:353–363
Hoffmann K, Pawlowska J, Walther G, Wrzosek M, de Hoog GS, Benny GL, Kirk
PM, Voigt K (2013) The family structure of the Mucorales: a synoptic revision
based on comprehensive multigene-genealogies. Persoonia 30:57–76
8859
8860
8861
8862
8863
8864
8865
8866
8867
8868
8869
8870
8871
8872
8873
8874
8875
8876
8877
8878
8879
8880
8881
8882
8883
8884
8885
8886
8887
8888
8889
8890
8891
8892
8893
8894
8895
8896
8897
8898
8899
8900
8901
8902
Hofstetter V, Clémençon H, Vilgalys R, Moncalvo J-M (2002) Phylogenetic analyses
of the Lyophylleae (Agaricales, Basidiomycota) based on nuclear and
mitochondrial rDNA sequences. Mycol Res 106:104–1059
Höhnel F. von (1918) Mycologische Fragmente, 272. Über die Hysteriaceen. Ann
Mycol 16:145–154
Höhnel F (1919) Fünfte vorläufige Mitteilung mykologische Ergebnisse (Nr.
300–500). Berichte der Deutschen Botanischen Gesellschaft 37:153–161
Holm L (1957) Nomenclatural notes on Pyrenomycetes. Taxon 24:475–488
Holm L, Holm K (1988) Studies in the Lophiostomataceae with emphasis on the
Swedish species. Symb Bot Upsal 28:1–50
Hong CX, Gallegly ME, Richardson PA, Kong P, Moorman GW, Lea-Cox JD, Ross
DS (2010). Phytophthora hydropathica, a new pathogen identified from
irrigation water, Rhododendron catawbiense and Kalmia latifolia. Plant Pathol
59:913–921
Hong CX, Richardson PA, Hao W, Ghimire SR, Kong P, Moorman GW, Lea-Cox JD,
Ross DS (2012). Phytophthora aquimorbida sp. nov. and Phytophthora taxon
‘aquatilis’ recovered from irrigation reservoirs and a stream in Virginia, USA.
Mycologia 104:1097–1108
Hongo T (1982) The Amanitas of Japan. Acta Phytotaxon Geobot 33:116–126
Hosen MI, Ge ZW. (2011). Clarkeinda trachodes (Agaricales, Basidiomycetes), first
record from Bangladesh. Mycotaxon 118: 331–336
Hongsanan S, Tian Q, Peršoh D, Zeng XY, Hyde KD, Chomnunti P, Boonmee S,
Bahkali AH, Wen TC (2015) Meliolales. Fungal Diversity 74(1):91−141
Hsieh HM, Ju YM, Rogers JD (2005) Molecular phylogeny of Hypoxylon and closely
related genera. Mycologia 97:844–865
Hundorf SM, Miller AN, Fernández FA (2004) Molecular systematics of the
Coronophorales and new species of Bertia, Lasiobertia and Nitschkia. Mycol
Res 108:1384–1398
Hyde KD (1990) A new marine ascomycete from Brunei. Aniptodera longispora sp.
nov. from intertidal mangrove wood. Bot Mar 33:335–338
Hyde KD (2002) Aniptodera triseptata sp. nov.(Halosphaeriales) from submerged
wood in freshwater. Cryptogam Mycol 23:5–7
Hyde KD, Ho WH, Tsui CKM (1999) The genera Aniptodera, Halosarpheia, Nais
and Phaeonectriella from freshwater habitats. Mycoscience 40:165183
Hyde KD, Vrijmoed LLP, Chinnaraj S, Jones EBG (1992) Massarina armatispora sp.
nov., a new intertidal ascomycete from mangroves. Bot Mar 35:325–328
Hyde KD, Cai L, McKenzie EHC, Yang YL, Zhang JZ, Prihastuti H (2009)
Colletotrichum: a catalogue of confusion. Fungal Divers 39:1–17
Hyde KD, Abd-Elsalam K, Cai L (2010) Morphology: still essential in a molecular
world. Mycotaxon 114:439–451
Hyde KD, Jones EBG, Liu JK, Ariyawansa H, Boehm E, Boonmee S, Braun U,
Chomnunti P, Crous PW, Dai DQ, Diederich P, Dissanayake A, Doilom M,
Doveri F, Hongsanan S, Jayawardena R, Lawrey JD, Li YM, Liu Y-X, Lucking
R, Monkai J, Muggia L, Nelsen MP, Pang KL, Phookamsak R, Senanayake IC,
8903
8904
8905
8906
8907
8908
8909
8910
8911
8912
8913
8914
8915
8916
8917
8918
8919
8920
8921
8922
8923
8924
8925
8926
8927
8928
8929
8930
8931
8932
8933
8934
8935
8936
8937
8938
8939
8940
8941
8942
8943
8944
Shearer CA, Suetrong S, Tanaka K, Thambugala KM, Wijayawardene NN,
Wikee S, Wu HX, Zhang Y, Aguirre-Hudson B, Alias SA, Aptroot A, Bahkali
AH, Bezerra JL, Bhat DJ, Camporesi E, Chukeatirote E, Gueidan C,
Hawksworth DL, Hirayama K, De Hoog S, Kang JC, Knudsen K, Li WJ, Li XH,
Liu ZY, Mapook A, McKenzie EHC, Miller AN, Mortimer PE, Phillips AJL,
Raja HA, Scheuer C, Schumm F, Taylor JE, Tian Q, Tibpromma S, Wanasinghe
DN, Wang Y, Xu JC, Yacharoen S, Yan JY, Zhang M (2013) Families of
Dothideomycetes. Fungal Divers 63 (1):1–313
Hyde KD, Nilsson RH, Alias SA, Ariyawansa HA, Blair JE, Cai L, de Cock AWAM,
Dissanayake AJ, Glockling SL, Goonasekara ID, Gorczak M, Hahn M,
Jayawardena RS, van Kan JAL, Laurence MH, Lévesque CA, Li XH, Liu JK,
Maharachchikumbura SSN, Manamgoda DS, Martin FN, McKenzie EHC,
McTaggart AR, Mortimer PE, Nair PVR, Pawłowska J, Rintoul TL, Shivas RG,
Spies CFJ, Summerell BA, Taylor PWJ, Terhem RB, Udayanga D, Vaghefi N,
Walther G, Wilk M, Wrzosek M, Xu JC, Yan JY, Zhou N (2014) One stop
shop:backbones trees for important phytopathogenic genera: I. Fungal Divers
67:21–125
Index Fungorum (2016) http://www.indexfungorum.org/Names/Names.asp. Accessed
on March 2016
Jacobs K, Botha A (2008) Mucor renisporus sp. nov., a new coprophilous species
from Southern Africa. Fungal Divers 29:27–35
Jaklitsch W, Baral HO, Lücking R, Lumbsch HT (2016) Engler's Syllabus of Plant
Families (Editor Frey W), 13th ed., Part 1/2: Ascomycota. Borntraeger, Stuttgart,
Germany
Jami F, Slippers B, Wingfield MJ, Gryzenhout M (2012) Five new species of the
Botryosphaeriaceae from Acacia karroo in South Africa. Cryptogam Mycol
33:245–266
Jayasiri SC, Hyde KD, Abd-Elsalam KA, Abdel-Wahab MA,Ariyawansa HA, Bhat J,
Buyck B, Dai YC, Ertz D, Hidayat I, Jeewon R, Jones EBG, Karunarathna SC,
Kirk P, Lei C, Liu JK, Maharachchikumbura SSN, McKenzie E, Ghobad-Nejhad
M, Nilsson H, Pang KL, Phookamsak R, Rollins AW, Romero AI,Stephenson S,
Suetrong S, Tsui CKM, Vizzini A, Wen TC, De Silva NI, Promputtha I, Kang JC
(2015) The Facesoffungi database: fungal names linked with morphology,
molecular and human attributes. Fungal Divers 74(1):3–18
Jeewon R, Yeung SYQ, Hyde KD (2009) A novel phylogenetic group within
Thozetella (Chaetosphaeriaceae): a new taxon based on morphology and DNA
sequence analyses. Can J Microbiol 55:680–687
Jones EBG (1995) Ultrastructure and taxonomy of the aquatic ascomycetous order
Halosphaeriales. Can J Bot 73(Suppl 1):S790–S801
Jones EBG, Sakayaroj J, Suetrong S, Somrithipol S, Pang KL (2009) Classification of
marine Ascomycota, anamorphic taxa and Basidiomycota. Fungal Divers
35:1–203
8945
8946
8947
8948
8949
8950
8951
8952
8953
8954
8955
8956
8957
8958
8959
8960
8961
8962
8963
8964
8965
8966
8967
8968
8969
8970
8971
8972
8973
8974
8975
8976
8977
8978
8979
8980
8981
8982
8983
8984
8985
8986
8987
8988
Jones EBG, Suetrong S, Sakayaroj J,Bahkali AH, Abdel-Wahab MA, Boekhout T,
Pang KL (2015) Classification of marine Ascomycota, Basidiomycota,
Blastocladiomycota and Chytridiomycota. Fungal Divers 73:1–72
Ju YM, Rogers JD (1996) A revision of the genus Hypoxylon. Mycologia Memoir no.
20. APS Press, St. Paul, 365
Ju YM, Hsieh HM, Ho MC, Szu DH, Fang MJ (2007) Theissenia rogersii sp. nov.
and phylogenetic position of Theissenia. Mycologia 66(4):612–621
Judelson HS (2009). Sexual reproduction in Oomycetes: biology, diversity, and
contributions to fitness. In: Lamour K, Kamoun S (eds.) Oomycete Genetics and
Genomics: Diversity, Interactions, and Research Tools. John Wiley & Sons,
Hoboken, pp 121–138
Kamoun S (2003). Molecular Genetics of Pathogenic Oomycetes. Eukaryot Cell
2:191–199
Karadžić D (2003) Tiarosporella species: Distribution and significance. Glasnik
Sumarskog Fakulteta 87:9–23
Karsten PA (1881) Enumeralio boletinearum et polypore arum fennicarum, systemate
novo dispositarum. Rev. Mycol. 3, 16–19
Karunarathna SC, Yang ZL, Zhao R, Vellinga EC, Bahkali AH, Chukeatirote E, Hyde
KD (2011) Three new species of Lentinus from northern Thailand. Mycol Prog
10:389–398
Kepler RM, Sung GH, Harada Y, Tanaka K, Tanaka E, Hosoya T, Bischoff JF,
Spatafora JW (2012) Host jumping onto close relatives and across Kingdoms by
Tyrannicordyceps
(Clavicipitaceae)
gen.
nov.
and
Ustilaginoidea
(Clavicipitaceae). Am J Bot 99(3):552–561
Kepler R M, Sung GH, Ban S, Nakagiri A, Chen MJ, Huang B, Spatafora JW (2012)
Newteleomorph combinations in the entomopathogenic genus Metacordyceps.
Mycologia 104(1):182–197
Kepler RM, Humber RA, Bischoff, JF, Rehner SA (2014) Clarification of generic and
species boundaries for Metarhizium and related fungi through multi-gene
phylogenetics. Mycologia 106(4):811–29
Kirisits T (2007) Die Petrakia-Blattbräune des Bergahorns. Forstschutz Aktuell
(Wien) 40:28–31
Knapp DG, Kovács GM, Zajta E, Groenewald JZ, Crous PW (2015) Dark septate
endophytic pleosporalean genera from semiarid areas. Persoonia 35:87–100
Kobayashi Y (1981) “The genus Cordyceps and its allies from Taiwan (Formosa)”
Bull Nat Sci Mus Ser B (Tokyo), 7(4):113–122
Kraichak, E., P.K. Divakar, A. Crespo, S.D. Leavitt, M.P. Nelsen, R. Lücking & H.T.
Lumbsch (2015) A tale of two hyper-diversities: diversification dynamics of the
two largest families of lichenized fungi. Sci. Rep. 5:10028
Kränzlin F (2005) Fungi of Switzerland, 6, Russulaceae: Lactarius, Russula. Verlag
Mykologia, Luzern
Kroon LPNM, Bakker FT, Van den Bosch GBM, Bonants PJM, Flier WG
(2004).Phylogenetic analysis of Phytophthora species based on mitochondrial
and nuclear DNA sequences. Fungal Genet Biol 41:766–782
8989
8990
8991
8992
8993
8994
8995
8996
8997
8998
8999
9000
9001
9002
9003
9004
9005
9006
9007
9008
9009
9010
9011
9012
9013
9014
9015
9016
9017
9018
9019
9020
9021
9022
9023
9024
9025
9026
9027
9028
9029
9030
9031
9032
Kroon LPNM, Brouwer H, de Cock AWAM, Govers F (2012). The genus
Phytophthora anno 2012. Phytopathol 102:348–364
Kirk PM, Cannon PF, Minter DW, Stalpers JA (2008) Ainsworth & Bisby’s
dictionary of the fungi, 10th edn. CABI, Wallingford
Kirk P M, Stalpers J A, Braun E, Crous P W, Hansen K (2013) A without-prejudice
list of generic names of fungi for protection under the International Code of
Nomenclature for algae, fungi, and plants. IMA Fungus 4:381–443
Ko WH, Yang CH, Lin MJ, Chen CY, Tsou YJ (2011) Humicola phialophoroides sp.
nov. from soil with potential for biological control of plant disease. Bot Stud
52:197–202
Kohlmeyer J, Kohlmeyer E (1979) Marine mycology. The higher fungi. Academic
Press, New York
Kohlmeyer J, Volkman-Kohlmeyer B (1987) Marine fungi of Aldabra, the Galapago,
and other tropical islands. Can J Bot 65:571–582
Kornerup A, Wanscher JH (1978) Methuen handbook of colour, 3rd edn. Methuen,
London
Kornerup A, Wanscher JH (1981) Taschenlexikon der Farben. 3. Aufl.
Muster-Schmidt Verlag, Göttingen
Kuhnert E, Fournier J, Peršoh D, Luangsa-ard JJD, Stadler M (2014) New Hypoxylon
species from Martinique and new evidence on the molecular phylogeny of
Hypoxylon based on ITS rDNA and β-tubulin data. Fungal Divers 64: 181-203.
Kuhnert E, Surup F, Sir EB, Lambert C, Hyde KD, Hladki AI, Romero AI, Stadler M
(2016) Lenormandins A – G, new azaphilones from Hypoxylon lenormandii and
Hypoxylon jaklitschii sp. nov., recognised by chemotaxonomic data. Fungal
Divers 71:165-184.
Kundsen H, Borgen T (1982) Russlaceae in Greenland. In: Artic and Alpine
Mycology:216–236
Kuntze O. (1891) Revisio generum plantarum (in Latin) 2. Leipzig, Germany: A.
Felix. 848pp
Kwasna H, Ward E, Bateman GL (2006) Phylogenetic relationships among
Zygomycetes from soil based on ITS1/2 rDNA sequences. Mycol Res
110:501–510
Larkin MA, Blackshields G, Brown NP, Chenna R, McGettigan PA, McWilliam H,
Valentin F, Wallace IM, Wilm A, Lopez R, Thompson JD, Gibson TJ, Higgins
DG. (2007) Clustal W and Clustal X version 2.0. Bioinformatics 23:2947–2948
Larsson KH (2007) Re-thinking the classification of corticioid fungi. Mycol Res
111(9):1040–1063
Larsson E, Larsson KH (2003) Phylogenetic relationships of russuloid basidiomycetes
with emphasis on aphyllophoralean taxa. Mycologia 95:1037–1065
Læssøe T, Spooner BM (1994) Rosellinia & Astrocystis (Xylariaceae): new species
and generic concepts. Kew Bull. 49(1):1–70
Læssøe T, Srikitikulchai P, Luangsa-ard JJ, Stadler M (2013) Theissenia reconsidered,
including molecular phylogeny of the type species T. pyrenocrata and a new
genus Durotheca (Xylariaceae, Ascomycota). IMA FUNGUS 4(1):57–69
9033
9034
9035
9036
9037
9038
9039
9040
9041
9042
9043
9044
9045
9046
9047
9048
9049
9050
9051
9052
9053
9054
9055
9056
9057
9058
9059
9060
9061
9062
9063
9064
9065
9066
9067
9068
9069
9070
9071
9072
9073
9074
9075
Le HT, Verbeken A, Nuytinck J, Lumyong S, Desjardin DE (2007) Lactarius in
Northern Thailand: 3. Lactarius subgenus Lactoriopsis. Mycotaxon 102:281–291
Lebel T, Tonkin JE (2007) Australasian species of Macowanites are sequestrate
species of Russula. Aust Syst Bot 20:355–381
Lebel T. (2013) Two new species of sequestrate Agaricus (sectionMinores) from
Australia. Mycol Prog. 12:699–707
Lechat C (2010) Ochronectria courtecuissei sp. nov. Bull. Soc. Mycol. Fr.
126(2):97–101
Leelavathy KM, Zachariah S, Sankaran KV (1981) Clarkeinda rhacodes a new record
from India. Mycologia 73(1):204–207
Li GJ (2014) Taxonomy of Russula from China. Ph.D. dissertation. Institute of
Microbiology, Chinese Academy of Sciences & University of Chinese Academy
of Sciences (In Chinese)
Li GJ (2015a) Recent research progress of Russula (Russulales, Agaricomycetes): a
review. Mycosystema 34:821–848 (In Chinese)
Li GJ, Wen HA (2009) Research of prospects on taxonomy of the Russula in China.
Mycosystema 28:303–309 (In Chinese)
Li GJ, Li SF, Wen HA (2011a) Russula zhejiangensis sp. nov. from East China.
Cryptogam Mycol 32:127–133
Li GJ, Li SF, Liu XZ, Wen HA (2012) Russula jilinensis sp. nov. (Russulaceae) from
northeast China. Mycotaxon 120:49–58
Li GJ, Zhao Q, Zhao D, Yue SF, Li SF, Wen HA, Liu XZ (2013a) Russula
atroaeruginea and R. sichuanensis spp. nov. from southwest China. Mycotaxon
124:137–188
Li GJ, Zhao D, Li SF, Yang HJ, Wen HA, Liu XZ (2013b) Russula changbaiensis sp.
nov. from northeast China. Mycotaxon 124:269–278
Li GJ, Zhao D, Li SF, Wen HA (2015b) Russula chiui and R. pseudopectinatoides,
two new species from southwestern China supported by morphological and
molecular evidence. Mycol Prog 14:33
Li J, Heath IB, Bauchop T (1990) Piromyces mae and Piromyces dumbonica, two
new species of uniflagellate anaerobic chytridiomycete fungi from the hindgut of
the horse and elephant. Can J Bot 68:1021–1033
Li J, Zhang J, Chen H, Chen X, Lan J. (2013c) Complete Mitochondrial Genome of
the Medicinal Mushroom Ganoderma lucidum. PLoS ONE 8(8):72038
Li WJ, Liu JK, Bhat DJ, Camporesi E, Xu J, Hyde KD (2014) Introducing the novel
species, Dothiorella symphoricarposicola, from snowberry in Italy. Cryptogam
Mycol 35:257–270
Li WJ, Bhat DJ, Camporesi E, Tian Q, Wijayawardene NN, Dai DQ, Phookamsak R,
Chomnunti P, Bahkali AH, Hyde KD (2015c). New asexual morph taxa in
Phaeosphaeriaceae. Mycosphere 6(6):681–708
Li YC, Yang ZL, Tolgor B (2009) Phylogenetic and biogeographic relationships of
Chroogomphus species as inferred from molecular and morphological data.
Fungal Divers 38:85–104
9076
9077
9078
9079
9080
9081
9082
9083
9084
9085
9086
9087
9088
9089
9090
9091
9092
9093
9094
9095
9096
9097
9098
9099
9100
9101
9102
9103
9104
9105
9106
9107
9108
9109
9110
9111
9112
9113
9114
9115
9116
9117
9118
Li YC, Feng B, Yang ZL (2011b) Zangia, a new genus of Boletaceae supported by
molecular and morphological evidence. Fungal Divers 49:125–143
Li YC, Ortiz-Santana B, Zeng NK, Feng B, Yang ZL (2014). Molecular phylogeny
and taxonomy of the genus Veloporphyrellus. Mycologia 106:291–306
Li YK, Zhang X, Yuan Y, Cao Z, Liang JF (2015b) Morphological and molecular
evidence for a new species of Russula (Russulaceae) from southern China.
Phytotaxa 202(2):94–102
Liang ZQ (2007) Cordyceps. Flora Fungorum Sinicorum. Science Press, Beijing.
32:1–190
Liebetanz E (1910) Die parasitischen Protozoen des Wiederkäuermagens. Arch. Für
Protistenkd 19:19–80
Liew ECY, Aptroot A, Hyde KD (2002) An evaluation of the monophyly of
Massarina based on ribosomal DNA sequences. Mycologia 94:803–813
Liimatainen K (2015) Nomenclatural novelties, Cortinarius. Index Fungorum no. 241
Linaldeddu BT, Deidda A, Scanu B, Franceschini A, Serra S, Berraf-Tebbal A,
Zouaoui Boutiti M, Jamâa MLB, Phillips AJL (2015) Diversity of
Botryosphaeriaceae species associated with grapevine and other woody hosts in
Italy, Algeria and Tunisia, with descriptions of Lasiodiplodia exigua and
Lasiodiplodia mediterranea sp. nov. Fungal Divers 71:201–204
Lindau G (1897) Pyrenomycetineae, Laboulbeniineae. In Engler, A.; Prantl, K. [eds],
Die Natürlichen Pflanzenfamilien. Teil. 1 1 (1):321–505. Leipzig; Verlag von
Wilhelm Engelmann.
Liu JK, Phookamsak R, Doilom M, Wikee S, Li YM, Ariyawansha H, Boonmee S,
Chomnunti P, Dai DQ, Bhat JD, Romero AI, Zhuang WY, Monkai J, Jones EBG,
Chukeatirote E, KoKo TW, Zhao YC, Wang Y, Hyde KD (2012) Towards a
natural classification of Botryosphaeriales. Fungal Divers 57:149–210
Liu JK, Hyde KD, Jones EBG, Ariyawansa HA. Bhat JD, Boonmee S,
Maharachchikumbura SSN., Mckenzie EHC, Phookamsak R, Phukhamsakda C,
Shenoy BD, Abdel-Wahab MA, Buyck B, Chen J, Chethana KWT, Singtripop C,
Dai DQ, Dai YC, Daranagama DA, Dissanayake AJ, Doilom M, D'souza MJ,
Fan XL, Goonasekara ID, Hirayama K, Hongsanan S, Jayasiri SC, Jayawardena
RS, Karunarathana SC, Li WJ, Mapook A, Norphanphoun C, Pang KL, Perera
RH, Peršoh D, Pinruan U, Senanayake IC, Somrithipol S, suetrong S, Tanaka K,
Thambugala KM, Tian Q, Tibpromma S, udayanga D, Wuayawardene NN,
Wanasinghe D, Wisitrassameewong K, Zeng XY, Abdel-Aziz FA, Adamčík S,
Bahkali AH, Boonyuen N, Bulgakov T, Callac P, Chomnunti P, Greiner K,
Hashimoto A, Hofstetter V, Kang JC, Lewis D, Li XL, Liu XX, Liu ZY,
Matsumura M, Mortimer PE, Rambold G, Randrianjohany E, Sato G,
Sriindrasutdhi V, Tian CM, Verbeken A, Von Brackel W, Wang Y, Wen TC, Xu
JC, Yan JY, Zhao RL, Camporesi E (2015) Fungal Diversity Notes 1-110:
Taxonomic and phylogenetic contributions to fungal species. Fungal Divers
72(1):1–197
Locquin M (1984) Mycologie Générale et Structurale. Masson, Paris
9119
9120
9121
9122
9123
9124
9125
9126
9127
9128
9129
9130
9131
9132
9133
9134
9135
9136
9137
9138
9139
9140
9141
9142
9143
9144
9145
9146
9147
9148
9149
9150
9151
9152
9153
9154
9155
9156
9157
9158
9159
9160
9161
9162
Lodge DJ, Padamsee M, Matheny PB, Aime MC, Cantrell SA, Boertmann D,
Kovalenko A, Vizzini A, Dentinger BTM, Kirk PM, Ainsworth AM, Moncalvo
J-M, Vilgalys R, Larsson E, Lücking R, Griffith GW, Smith ME, Norvell LL,
Desjardin DE, Redhead SA, Ovrebo CL, Lickey EB, Ercole E, Hughes KW,
Courtecuisse R, Young A, Binder M, Minnis AM, Lindner DL, Ortiz-Santana B,
Haight J, Læssøe T, Baroni TJ, Geml J, Hattori T (2014) Molecular phylogeny,
morphology, pigment chemistry and ecology in Hygrophoraceae (Agaricales).
Fungal Divers 64:1–99
Luangsa-ard JJ, Tasanathai K, Mongkolsamrit S, Hywel-Jones NL. (2007) Atlas of
invertebrate-pathogenic fungi of Thailand. vol. 1. BIOTEC, National Science and
Tecnology Development Agency, Thailand
Lücking R (2014) A key to species of the Ocellularia papillata, perforata and
terebrata morphodemes (Ascomycota: Graphidaceae). Glalia 6(3):1–35
Lücking R (2015) Thelotremoid Graphidaceae from the NYBG herbarium: New
species, range extensions, and a forgotten lichen. Opusc Philolich 14:1–57
Lücking R, Pérez-Ortega S (2015) Four new species of Ocellularia (lichenized
Ascomycota: Graphidaceae) from Cuba, with a revised taxonomy of the O.
bahiana complex and a key to thelotremoid taxa with small, brown, (sub-)
muriform ascospores. Lichenologist 47:305–322
Lücking R, Johnston MK, Aptroot A, Kraichak E, Lendemer JC, Boonpragob K,
Cáceres MES, Ertz D, Ferraro LI, Jia ZF, Kalb K, Mangold A, Manoch L,
Mercado-Díaz JA, Moncada B, Mongkolsuk P, Papong K, Parnmen S, Peláez R,
Poengsoengnoen V, Rivas Plata E, W. Saipunkaew, Sipman HJM,
Suttjaritturakan J, Van den Broeck D, von Konrat M, Weerakoon G, Lumbsch
HT. (2014) One hundred and sventy five new species of Graphidaceae: closing
the gap or a drop in the bucket? Phytotaxa 189:7–38
Lücking R, Johnston MK, Aptroot A, Kraichak E, Lendemer JC, Boonpragob K,
Cáceres MES, Ertz D, Ferraro LI, Jia ZF, Kalb K, Mangold A, Manoch L,
Mercado-Díaz JA, Moncada B, Mongkolsuk P, Papong K, Parnmen S, Peláez R,
Poengsoengnoen V, Rivas Plata E, Saipunkaew W, Sipman HJM,
Suttjaritturakan J, van den Broeck D, von Konrat M, Weerakoon G, Lumbsch HT
(2014) One hundred and sventy five new species of Graphidaceae: closing the
gap or a drop in the bucket? Phytotaxa 189:7–38
Lumbsch HT, Huhndorf SM (2010) Outline of Ascomycota – 2009. Fieldiana Life and
Earth Sciences 1:1–60
Luque J, Sierra D, Torres E, Garcia F (2006) Cryptovalsa ampelina on grapevines in
N.E. Spain: identification and pathogenicity. Phytopathologia Mediterranea
45:S101–S109
Maerz A, Paul MR (1950) A dictionary of color. USA: McGraw-Hill book Company.
Madden AA, Stchigel AM, Guarro J, Sutton D, Starks PT (2012). Mucor nidicola sp.
nov., a fungal species isolated from an invasive paper wasp nest. Int J Syst Evol
Microbiol 62:1710–1714
Maharachchikumbura SSN, Hyde KD, Jones EBG, McKenzie EHC, Huang S-K,
Abdel-Wahab MA, Daranagama DA, Dayarathne M, D’souza MJ, Goonasekara
9163
9164
9165
9166
9167
9168
9169
9170
9171
9172
9173
9174
9175
9176
9177
9178
9179
9180
9181
9182
9183
9184
9185
9186
9187
9188
9189
9190
9191
9192
9193
9194
9195
9196
9197
9198
9199
9200
9201
9202
9203
9204
9205
ID, Hongsanan S, Jayawardena RS, Kirk PM, Konta S, Liu J-K, Liu Z-Y,
Norphanphoun C, Shenoy BD, Xiao Y, Bahkali AH, Kang J, Somrothipol S,
Suetrong S, Wen T, Xu J (2015) Towards a natural classification and backbone
tree for Sordariomycetes. Fungal Divers 72:199–301
Maharachchikumbura SSN, Hyde KD, Jones EBG, McKenzie EHC, Huang S-K,
Abdel-Wahab MA, Daranagama DA, Dayarathne M, D’souza MJ, Goonasekara
ID, Hongsanan S, Jayawardena RS, Kirk PM, Konta S, Liu J-K, Liu Z-Y,
Norphanphoun C, Shenoy BD, Xiao Y, Bahkali AH, Kang J, Somrothipol S,
Suetrong S, Wen T, Xu J (2016) Families of Sordariomycetes. Fungal Divers
72:199–301
Mains EB (1959a) North American species of Aschersonia parasitic on Aleyrodidae.
Journal of Insect Pathology 1:43–47
Mains EB (1959b) Species of Aschersonia (Sphaeropsidales). Lloydia 22:215–221
Marano AV, Pires-Zottarelli CLA, de Souza JI, Glockling SL, Leaño E, Gachon
CMM, Strittmatter M, Gleason FH (2012). Hyphochytriomycota, Oomycota and
Perkinsozoa (SupergroupChromalveolata). In: Jones EBG, Pang KL (orgs.)
Marine Fungi and fungal-like organisms. De Gruyter, Berlin, pp 167–213.
Marano AV, Jesus AL, Pires-Zottarelli CLA, James TY, Gleason FH, de Souza JI
(2014). Phylogenetic relationships of Pythiales and Peronosporales (Oomycetes,
Straminipila) within the “peronosporalean galaxy”. In: Jones EBG, Hyde KD,
Pang KL (orgs.) Freshwater Fungi and fungal-like organisms. De Gruyter,
Berlin, pp 177–199.
Marano AV, Jesus AL, de Souza JI, Jerônimo GH, Gonçalves DR, Boro MC, Rocha
SCO, Pires-Zottarelli CLA (2016). Ecological roles of saprotrophic
Peronosporales (Oomycetes, Straminipila) in natural environments. Fungal Ecol
19:77–88
Matheny PB, Curtis JM, Hofstetter V, Aime MC, Moncalvo JM, Ge ZW, Slot JC,
Ammirati JF, Baroni TJ, Bougher NL, Hughes KW, Lodge DJ, Kerrigan RW,
Seidl MT, Aanen DK, DeNitis M, Daniele GM, Desjardin DE, Kropp BR,
Norvell LL, Parker A, Vellinga EC, Vilgalys R, Hibbett DS (2006) Major clades
of Agaricales: a multi-locus phylogenetic overview. Mycologia 98:982–995
Martín MT, Martín L, de-Francisco MT, Cobos R. (2009) First report of Lasidiplodia
theobromae and Cryptovalsa ampelina associated with grapevine decline from
Castillay Leon, Spain. Plant Disease 93(5):545
McNeill J, Barrie FR, Burdet HM, Demoulin V, Hawksworth DL, Marhold K,
Nicolson DH, Prado J, Silva PC, Skog JE, Wiersema JH, Turland NJ (eds.)
(2006). International Code of Botanical Nomenclature (Vienna Code). A. R. G.
Gantner Verlag, Ruggell, Liechtenstein.
McNeill J, Barrie FR, Buck WR, et al. (2012) International code of nomenclature for
algae, fungi, and plants (Melbourne Code). Regnum Veg 154
Mehrabi M, Mohammadi Goltapeh E, Fotouhifar KB (2011) Studies on Cytospora
canker disease of apple trees in Semirom region of Iran. Journal of Agricultural
Technology 7(4):967-982
9206
9207
9208
9209
9210
9211
9212
9213
9214
9215
9216
9217
9218
9219
9220
9221
9222
9223
9224
9225
9226
9227
9228
9229
9230
9231
9232
9233
9234
9235
9236
9237
9238
9239
9240
9241
9242
9243
9244
9245
9246
9247
9248
Mehrabi M, Hemmati R, Vasilyeva LN, Trouillas FP (2015) A new species and a new
record of Diatrypaceae from Iran. Mycosphere 6(1):60–68
Mehrotra BS, Mehrotra BM. (1978/1979). Another azygosporic species of Mucor
from India. Sydowia 31:94–96
Mostert L, Halleen F, Creaser ML, Crous PW (2004) Cryptovalsa ampelina, a
forgotten shoot and cane pathogen of grapevines. Austral Pl Pathol 33:295–299
Miller SL, Buyck B (2002) Molecular phylogeny of the genus Russula in Europe with
a comparison of modern infrageneric classifications. Mycol Res 106(3):259–276
Miller SL, McClean TM, Walker JF, Buyck B (2001) A molecular phylogeny of the
Russulales including agaricoid, gasteroid and pleurotoid taxa. Mycologia
93(2):344–354
Miller SL, Larsson E, Larsson KH, Verbeken A, Nuytinck J (2006) Perspectives in
the new Russulales. Mycologia 98(6):960–970
Miller SL, Aime MC, Henkel TW (2012) Russulaceae of Pakaraima Mountain of
Guyana 2. New species of Russula and Lactifluus. Mycotaxon 121:233–253
Mirarab S, Nguyen N, Warnow T (2014) PASTA: Ultra-Large Multiple Sequence
Alignment. In R. Sharan (Ed.), Research in Computational Molecular Biology
(RECOMB)
Mirza JH, Khan SM, Begum S, Shagufta S (1979) Mucorales of Pakistan. Faisalabad,
Pakistan: University of Agriculture
Moncalvo JM, Ryvarden L (1997) A nomenclatural study of the Ganodermataceae
Donk. Fungi Flora 10:1–114
Moncalvo JM, Lutzoni FM, Rehner SA, Johnson J, Vilgalys R (2000) Phylogenetic
relationships of agaric fungi based on nuclear large subunit ribosomal DNA
sequences. Syst Biol 49 (2):278–305
Moncalvo JM, Vilgalys R, Redhead SA, Johnson JE, James TY, Aime MC, Hofstetter
V, Verduin, SJW, Larsson E, Baroni TJ, Thorn GR, Jacobsson S, Clémençon H,
Miller OK (2002) One hundred and seventeen clades of euagarics. Mole Phyl &
Evol 23:357–400
Müller E, von Arx JA (1962) Die Gattungen der didymosporen Pyrenomyceten.
Beiträge. Kryptogamenflora Schweiz 11(2):1–922
Musumeci E (2014) Fungi Non Delineati raro vel haud perspecte et explorate
descripti aut definite picti. Pars LXVII–LXIX. Contributo alla conoscenza della
Micoflora europea: Specie nuove endemiche, funghi rari con microclima
localizzato. Candusso Edizioni, I-Alassio-(SV)
Muszewska A, Pawlowska J, Krzys’sciak P (2014) Biology, systematics, and clinical
manifestations of Zygomycota infections. Eur J Clin Microbiol Infect Dis
33:1273–1287
MycoBank (2016) http://www.mycobank.org/Biolomics.aspx?Table=Mycobank.
Accessed on March 2016
Naher M, Motohash K, Watanabe H, Chikuo Y, Senda M, Suga H, Brasier C,
Kageyama K (2011). Phytophthora chrysanthemi sp. nov., a new species causing
root rot of chrysanthemum in Japan. Mycol Prog 10:21–31
9249
9250
9251
9252
9253
9254
9255
9256
9257
9258
9259
9260
9261
9262
9263
9264
9265
9266
9267
9268
9269
9270
9271
9272
9273
9274
9275
9276
9277
9278
9279
9280
9281
9282
9283
9284
9285
9286
9287
9288
9289
9290
9291
Nag Raj TR (1993) Coelomycetous anamorphs with appendage-bearing conidia.
Waterloo, Canada: Mycologue Publications
Nakagiri A (2002). Diversity and phylogeny of Halophytophthora (Oomycetes). In:
Book of abstracts of the IMC7 – The 7th International Mycological Congress.
Oslo, 11–17 Aug 2002. No 55 Book of Abstracts p 19
Nakagiri A, Izumi O (2005). Phylogeny, taxonomy and ecology of
Halophytophthoraspinosa (marine Oomycetes). Inoculum 56:43
Niemelä T, Wagner T, Fischer M, Dai YC (2001) Phellopilus gen. nov. and its
affinities within Phellinus s. lato and Inonotus s. lato (Basidiomycetes). Ann Bot
Fenn 38:51–62
Nitschke TRJ (1867) Pyrenomycetes Germanici. Die Kernpilze Deutschlands
Bearbeitet von Dr. Th. Nitschke. Erster Band. Erste Lieferung i-ii, 156 Germany,
Bresalau; Verlag von Eduard Trewendt
Norphanphoun C, Maharachchikumbura SSN, Daranagama A, BulgakovTS, Bhat DJ,
Bahkali AH, Hyde KD (2015) Towards a backbone tree for Seimatosporium,
with S. physocarpi sp. nov. Mycosphere 6(3):385–400
O’Donnell K, Cigelnik E, Benny GL (1998) Phylogenetic relationships among the
Harpellales and Kickxellales. Mycologia 90:624–639
O’Donnell K, Lutzoni FM, Ward TJ, Benny GL (2001) Evolutionary relationships
among mucoralean fungi (Zygomycota): Evidence for family polyphyly on a
large scale. Mycologia 93:286–296
Ogawa J, Sakurdani E, Kishino S, Ando A, Yokozeki K, Shimizu S (2012)
Polyunsaturated fatty acids production and transformation by Mortierellaalpina
and anaerobic bacteria. European Journal of Lipid Science and Technology
114:1107–1113
Okane I, Nakagiri A. Ito T (1996) Discostroma tricellulare, a new endophytic
ascomycete with a Seimatosporium anamorph isolated from Rhododendron. Can
J Bot 74(8):1338−1344
Omvik A (1955) Two new species of Chaetomium and one new Humicola species.
Mycologia 47:748–757
Orpin CG (1975) Studies on the rumen flagellate Neocallimastix frontalis. J. Gen.
Microbiol. 91:249–262
Orpin CG (1977a) The occurrence of chitin in the cell walls of the rumen organisms
Neocallimastix frontalis, Piromonas communis and Sphaeromonas communis. J.
Gen. Microbiol 99:215–218
Orpin CG (1977b) The rumen flagellate Piromonas communis: its life-history and
invasion of plant material in the rumen. J Gen Microbiol 99:107–117
Orpin CG, Munn EA (1986) Neocallimastix patriciarum sp. nov., a new member of
the Neocallimasticaceae inhabiting the rumen of sheep. Trans. Br. Mycol. Soc.
86:178–181
Ott M, Zola J, Aluru S, Stamatakis A (2007) Large-scale maximum likelihood-based
phylogenetic analysis on the IBM BlueGene/L. Proceedings of ACM/IEEE
Supercomputing conference. Article No. 4
9292
9293
9294
9295
9296
9297
9298
9299
9300
9301
9302
9303
9304
9305
9306
9307
9308
9309
9310
9311
9312
9313
9314
9315
9316
9317
9318
9319
9320
9321
9322
9323
9324
9325
9326
9327
9328
9329
9330
9331
9332
9333
9334
9335
9336
Pang KL, Vrijmoed LLP, Kong RYC, Jones EBG (2003) Lignincola and Nais,
polyphyletic genera of the Halosphaeriales (Ascomycota). Mycol Prog 2:29–36
Pang KL, Lin HJ, Lin HY, Huang YF, Chen YM (2015). Production of arachidonic
and eicosapentaenoic acids by the marine oomycete Halophytophthora. Mar
Biotechnol 17:121–129
Parra LA. 2008. Agaricus L Allopsalliota Nauta & Bas. Italy:Pars 1Edizioni Candusso
Alassio
Parra LA. 2013. Fungi Europaei, Volume 1A, Agaricus L.:Allopsalliota, Nauta & Bas
(Parte II). Candusso Edizioni
Pegler DN (1983) The Genus Lentinus: A World Monograph. HMSO, London
Pegler DN (1985) The genus Clarkeinda (Basidiomycotina: Agaricaceae). Bot J Linn
Soc 91(1-2):245–252
Pegler DN (1986) Agaric flora of Sri Lanka. Kew Bull. Additional Series 6, Her
Majesty’s Stationary Office, London
Pei KQ (2000) A new variety of Mucor variosporus and the validation of M. luteus
Linnemann and M. variosporus Schipper. Mycosystema 19:10–12
Peintner U, Bougher NL, Castellano MA, Moncalvo JM, Moser MM, Trappe JM.,
Vilgalys R. (2001) Multiple origins of sequestrate fungi related to Cortinarius
(Cortinariaceae). Am. J. Bot. 88(12):2168–2179
Peintner U, Horak E, Moser M., Vilgalys R. (2002) Phylogeny of Rozites, Cuphocybe
and Rapacea inferred from ITS and LSU rDNA sequences. Mycologia
94(4):620–629
Petch T (1921) Studies in entomogenous fungi. II. The genera of Hypocrella and
Aschersonia. Ann Roy Bot Gard Perad 7:167–278
Petch T (1931) Notes on entomogenous fungi. Trans Brit Mycol Soc 16(1):55–75
Petch T (1932) A list of the entomogenous fungi of Great Britain. Trans Brit Mycol Soc
17(3):170–178
Petch T, Bisby GR (1950) The Fungi of Ceylon. Peradeniya Manual 6:111pp
Petrak F (1923a). Mykologische Notizen VI. No. 226. Über die Gattung Glonium
Muhl. Ann Mycol 21:225–227
Petrak F (1923b). Mykologische Notizen VI, Nr. 284. Psiloglonium finkii nov. sp.
Annales Mycologici 21:308–309
Petrak F. (1934) Mykologische Notizen. XII. Annales Mycologici. 32(5-6):317-447
Petrak F (1941) Mykologische Notizen. XIV. AnnalesMycologici. 39 (4–6):251–349
Paterson RRM (2006) Ganoderma – a therapeutic fungal bio factory. Photochemistry
67:1985–2001
Petrini L.E. (1992) Rosellinia species of the temperate zones. Sydowia 44:169–281
Petrini L.E. (2013) Rosellinia - a world monograph. Bibl Mycol 205:1–410
Peyronel B, Dal Vesco G (1955) Ricerche sulla microflora di un terreno agrario
presso Torino. Allionia 2:357–417
Pici G (1955) Qualche osservazione sopra due Mucoraceae. Atti Inst Bot Univ
Pavia 13:38–44
Pilotti CA, Sanderson FR, Aitken AB, Armstrong W (2004) Morphological variation
and host range of two Ganoderma species from Papua New Guinea
Mycopathologia 158:251–265
9337
9338
9339
9340
9341
9342
9343
9344
9345
9346
9347
9348
9349
9350
9351
9352
9353
9354
9355
9356
9357
9358
9359
9360
9361
9362
9363
9364
9365
9366
9367
9368
9369
9370
9371
9372
9373
9374
9375
9376
9377
9378
9379
9380
Pinnoi A, Phongpaichit P, Jeewon R, Tang AMC, Hyde KD, Jones EBG. (2010)
Phylogenetic relationships of Astrocystis eleiodoxae sp. nov. (Xylariaceae).
Mycosphere I:1–9
Phillips A, Alves A, Correia A, Luque J (2005) Two new species of Botryosphaeria
with brown, 1-septate ascospores and Dothiorella anamorphs. Mycologia
97:513–529
Phillips AJL, Alves A, Abdollahzadeh J, Slippers B, Wingfield MJ, Groenewald JZ,
Crous PW (2013) The Botryosphaeriaceae: genera and species known from
culture. Stud Mycol 76:51–167
Phookamsak R, Liu JK, McKenzie EHC, Manamgoda DS, Ariyawansa HA,
Thambugala KM, Dai DQ, Camporesi E, Chukeatirote E, Wijayawardene NN,
Bahkali AH, Mortimer PE Xu JC, Hyde KD (2014) Revision of
Phaeosphaeriaceae. Fungal Divers 68:159–238
Phookamsak R, Norphanphoun C, Tanaka K, Dai DQ, Luo ZL, Liu JK,Su HY, Bhat
DJ, Bahkali AH, Mortimer PE, Xu JC, Hyde KD (2015) Towards a natural
classidication of Astrosphaeriella-like species; introducing Astrosphaeriellaceae
and Pseudoastrosphaeriellaceae fam. nov and Astrosphaeriellaceaeopsis, gen.
nov. Fungal Divers 74:143–197
Pitt WM, Trouillas FP, Gubler WD, Savocchia S, Sosnowski MR (2013a)
Pathogenicity of diatrypaceous fungi on grapevines in Australia. Plant Dis.
97:749–756
Pitt WM, Úrbez-Torres JR, Trouillas FP (2013b) Dothiorella vidmadera, a novel
species from grapevines in Australia and notes on Spencermartinsia. Fungal
Divers 61:209–219
Pitt WM, Úrbez-Torres JR, Trouillas FP (2015) Dothiorella and Spencermartinsia,
new species and records from grapevines in Australia. Aust Pl Pathol 44:43–56
Quaedvlieg W, Kema GHJ, Groenewald JZ, Verkley GJM, Seifbarghi S, Razavi M,
Mirzadi Gohari A, Mehrabi R, Crous PW (2011). Zymoseptoria gen. nov.: a new
genus to accommodate Septoria- like speciesoccurring on graminicolous hosts.
Persoonia 26:57–69
Quaedvlieg W, Verkley GJM, Shin H-D, Barretto RW, Alfenas AC, Swart WJ,
Groenewald JZ, Crous PW (2013) Sizing up Septoria. Stud Mycol 75:307–390
Quandt C A, Kepler R M, Gams W, Araújo J P, Ban S, Evans H C, Luangsa-Ard J J
(2014) Phylogenetic-based nomenclatural proposals for Ophiocordycipitaceae
(Hypocreales) with new combinations in Tolypocladium. IMA fungus 5(1):121
Raja HA, Shearer CA (2008) Freshwater ascomycetes: new and noteworthy species
from aquatic habitats in Florida. Mycologia 100:467–489
Raja HA, Tanaka K, Hirayama K, Miller AH, Shearer CA (2011) Freshwater
ascomycetes: two new species of Lindgomyces (Lindgomycetaceae, Pleosporales,
Dothideomycetes) from Japan and USA. Mycologia 103:1421–1432
Raja HA, Oberlies NH, El-Elimat T, Miller AN, Zelski SE, Shearer CA (2013)
Lindgomyces angustiascus (Lindgomycetaceae, Pleosporales, Dothideomycetes),
a new lignicolous species from freshwater habitats in the USA. Mycoscience
54:353–361
9381
9382
9383
9384
9385
9386
9387
9388
9389
9390
9391
9392
9393
9394
9395
9396
9397
9398
9399
9400
9401
9402
9403
9404
9405
9406
9407
9408
9409
9410
9411
9412
9413
9414
9415
9416
9417
9418
9419
9420
9421
9422
9423
9424
Rajchenberg M, Pildain MB, Bianchinotti MV, Barroetaveña C (2015) The
phylogenetic position of poroid Hymenochaetaceae (Hymenochaetales,
Basidiomycota) from Patagonia, Argentina. Mycologia 107:754–767
Rea AJ, Burgess TI, Hardy GESJ, Stukely MJC, Jung T (2011). Two novel and
potentially endemic species of Phytophthora associated with episodic dieback of
Kwongan vegetation in the south-west of Western Australia. Plant Pathol
60:1055–1068
Réblová M, Winka K (2000) Phylogeny of Chaetosphaeria and its anamorphs based
on morphological and molecular data. Mycologia 92:939–954
Réblová M, Barr ME, Samuels GJ (1999) Chaetosphaeriaceae, a new family
forChaetosphaeria and its relatives. Sydowia 51:49–70
Réblová M, Gams W, Seifert KA (2011) Monilochaetes and allied genera of the
Glomerellales, and a reconsideration of families in the Microascales. Stud Mycol
68:163–191
Reeser PW, Sutton W, Hansen EM, Remigi P, Adams GC (2011).Phytophthora
species in forest streams in Oregon and Alaska. Mycologia 103:22–35
Ribaldi MS (1952) Sopra un interessante Zigomicete terricolo: Gongronella
urceolifera n. gen. et n. sp. Rivista di Biologia 49:157–166
Ribes JA, Vanover-Sams CL, Baker DJ (2000) Zygomycetes in human disease. Clin
Microbiol Rev 13:236–301
Ridgway R (1912) Colour standards and colour nomenclature. Robert Ridgway,
Washington
Rivas Plata E, Lücking R, Lumbsch HT (2008) When family matters: an analysis of
Thelotremataceae (Lichenized Ascomycota: Ostropales) as bioindicators of
ecological continuity in tropical forests. Biodiv Cons 17:1319–1351
Rivas Plata E, Lumbsch HT, Lücking R (2012a) A new classification for the lichen
family Graphidaceae s.lat. (Ascomycota: Lecanoromycetes: Ostropales). Fung
Divers 52:107–121
Rivas Plata E, Lücking R, Lumbsch HT (2012b) Molecular phylogeny and
systematics of the Ocellularia-clade (Ascomycota: Ostropales: Graphidaceae).
Taxon 61:1161–1179
Rivas Plata E., Parnmen S, Staiger B, Mangold A, Frisch A, Weerakoon G,
Hernández JE, Cáceres MES, Kalb K, Sipman HJM, Common RS, Nelsen MP,
Lücking R, Lumbsch HT (2013) A molecular phylogeny of Graphidaceae
(Ascomycota: Lecanoromycetes: Ostropales) including 428 species. MycoKeys
6:55–94
Roa Engel CA, Straathof AJJ, Zijlmans TW, van Gulik WM, van der Wielen LAM
(2008) Fumaric acid production by fermentation. App Microbiol Biotech
78:379–389
Robideau GP, de Cock AWAM, Coffey MD, Voglmayr H, Brouwer H, Bala K,
Chitty DW, Désaulniers N,Eggertson QA, Gachon CMM, Hu CH, Küpper FC,
Rintoul TL, Sarhan E, Verstappen ECP, Zhang Y, Bonants PJM, Ristaino JB,
Lévesque AC (2011). DNA barcoding of oomycetes with cytochrome c oxidase
subunit I and internal transcribed spacer. Mol Ecol Resour11:1002–1011
9425
9426
9427
9428
9429
9430
9431
9432
9433
9434
9435
9436
9437
9438
9439
9440
9441
9442
9443
9444
9445
9446
9447
9448
9449
9450
9451
9452
9453
9454
9455
9456
9457
9458
9459
9460
9461
9462
9463
9464
9465
9466
9467
9468
Rogers JD (2000) Thoughts and musings about tropical Xylariaceae. Mycol Res
104:1412–1420
Romagnesi H (1967) Russules d’Europe et d’Afique du Nord. Bordas, Paris
Romagnesi H (1985) Les Russules d’ Europe et d’ Afrique du Nord. Reprint with
supplement. J. Cramer, Lehre
Romagnesi H (1987) Status et noms nouveaux pour les taxa infragénériques dans le
genere Russula. Documentation Mycologique 18:39–40
Ronquist F, Teslenko M, Van Der Mark P, Ayres DL, Darling A, Hohna S, Larget B,
Liu L, Suchard MA, Huelsenbeck JP (2012). MrBayes 3.2: efficient Bayesian
phylogenetic inference and model choice across a large model space. Sist Biol
61:1–4
Rossman AY, Palm ME (2006). Why are Phytophthora and other Oomycota not true
Fungi? Outlook Pest Manag 17:217–219
Rossman, AY, Samuels GJ, Rogerson CT, Lowen R (1999) Genera of Bionectriaceae,
Hypocreaceae and Nectriaceae (Hypocreales, Ascomycetes). Stud Mycol
42:1–248
Rossman AY, McKemy JM, Pardo-Schultheiss RA, Schroers HJ (2001) Molecular
studies of the Bionectriaceae using large subunit rDNA sequences. Mycologia
93(1):100–110
Rossman AY, Adams GC, Cannon PF, Castlebury LA, Crous PW, Gryzenhout M,
Jaklitsch WM, Mejia LC, Stoykov D, Udayanga D, Voglmayr H, Walker DM
(2015) Recommendations of generic names in Diaporthales competing for
protection or use. IMA Fungus 6:145–154
Ryvarden L (1991) Genera of polypores, nomenclature and taxonomy, Fungiflora,
Oslo
Ryvarden L (2005) The genus Inonotus, a synopsis. Synopsis Fungorum 21:1–149
Saccardo, P. A. (1880). Conspectus generum fungorum italiae inferiorum nempe ad
Sphaeropsideas, Melanconieas et Hyphomyceteas pertinentium, systemate
sporologico dispositu. Michelia 2:1–38
Saccardo PA (1882) Sylloge Fungorum 1: i-xviii, 187–188. Italy, Padua
Saccardo PA (1884) Sylloge Fungorum vol 3. Typis Seminarii, Italy. (in Latin).
Saccardo PA (1925–1928) Sylloge Fungorum XXIV:1040
Sakayaroj J, Pang KL, Jones EBG (2011) Multi-gene phylogeny of the
Halosphaeriaceae: its ordinal status, relationships between genera and
morphological character evolution. Fungal Divers 46:87109
Samuels GJ, Candoussau F, Magni JF (1997) Fungicolous pyrenomycetes 1.
Helminthosphaeria and the new family Helminthosphaeriaceae. Mycologia
89:141–155
Sánchez-García M, Matheny PB, Palfner G, Lodge DJ (2014) Deconstructing the
Tricholomataceae (Agaricales) and introduction of the new genera Albomagister,
Corneriella, Pogonoloma and Pseudotricholoma. Taxon 63:993–1007
Santiago ALCM de A, Santos PJP, Trufem SFB, Malosso E, Cavalcanti MAQ (2011)
Zygomycetes from herbivore dung in the Ecological Reserve of Dois Irmãos,
Northeast Brazil. Braz J Microbiol 42:89–95
9469
9470
9471
9472
9473
9474
9475
9476
9477
9478
9479
9480
9481
9482
9483
9484
9485
9486
9487
9488
9489
9490
9491
9492
9493
9494
9495
9496
9497
9498
9499
9500
9501
9502
9503
9504
9505
9506
9507
9508
9509
9510
9511
9512
9513
Santiago ALCM de A, Santos, PJP, Maia, LC (2013) Mucorales from the semiarid of
Pernambuco, Brazil. Braz J Microbiol 44 (1):299–305
Santos MJS, de Oliveira PC, Trufem SFB (2003) Morphological observations on
Absidia corymbifera and Absidia blakesleeana strains perserved under mineral
oil. Mycoses 46:402–406
Sarnari M (1998) Monografia illustrate de genere Russula in Europa. Tomo Primo.
AMB, Centro Studi Micologici, Trento
Sarnari M (2005) Monografia illustrate de genere Russula in Europa. Tomo Secondo.
AMB, Centro Studi Micologici, Trento
Sato G, Tanaka K, Hosoya T (2008) Bambusicolous fungi in Japan (8): a new species
of Pseudolachnella from Yakushima Island, southern Japan. Mycoscience
49:392–394
Schardl C L, Young C A, Moore N, Krom N, Dupont P-Y, Pan J, Florea S, Webb J S,
Jaromczyk J, Jaromczyk JW, Cox M, Farman ML (2014) Chapter Ten –
Genomes of Plant-Associated Clavicipitaceae. In: Martin, F. M., Advances in
Botanical Research. (pp. 291-327). INRA Centre de Nancy, Centre de Nancy,
Champenoux,: Academic Press Inc
Schipper MAA (1973) A study on variability in Mucor hiemalis and related species.
Stud Mycol 4:1–40
Schipper MAA (1975) Mucor mucedo, Mucor flavus and related species. Stud Mycol
10:1–33
Schipper MAA (1976) On Mucor circinelloides, Mucor racemosus and related species.
Stud Mycol 12:1–40
Schipper MAA (1978) On certain species of Mucor with a key to all accepted species.
Stud Mycol 17:1–52
Schipper MAA (1984) A revision of the genus Rhizopus I. The Rh. solonifer-group
and Rh. oyzae. Stud Mycol 25:1–19
Schipper MAA, Stalpers JA (1984) A revision of the genus Rhizopus II. The Rh.
microsporus-group. Stud Mycol 25:20–34
Schipper MAA (1990) Note on Mucorales-I. Observations on Absidia. Persoonia
14:133–149
Schipper MAA, Samson RA (1978). On certain species of Mucor with a key to all
accepted species. Stud Mycol 17:1–52
Schipper MAA, Samson, RA (1994) Miscellaneous notes on Mucoraceae. Mycotaxon
50:475–491
Slippers B, Boissin E, Phillips AJL, Groenewald JZ, Lombard L, Wingfield MJ,
Postma A, Burgess T, Crous PW (2013) Phylogenetic lineages in the
Botryosphaeriales: a systematic and evolutionary framework. Stud Mycol
76:31–49
Slippers B, Roux J, Wingfield MJ, van der Walt, FJJ, Jami F, Mehl JWM, Marais GJ
(2014) Confronting the constraints of morphological taxonomy in the
Botryosphaeriales. Persoonia 33:55–168
Schoenlein-Crusius IH, Milanez AI, Trufem SFB, Pires-Zottarelli CLA, Grandi RAP,
Santos ML, Giustra KC (2006) Microscopic fungi in the Atlantic Rainforest in
Cubatão, São Paulo, Brazil. Braz J Microbiol 37:267–275
9514
9515
9516
9517
9518
9519
9520
9521
9522
9523
9524
9525
9526
9527
9528
9529
9530
9531
9532
9533
9534
9535
9536
9537
9538
9539
9540
9541
9542
9543
9544
9545
9546
9547
9548
9549
9550
9551
9552
9553
9554
9555
9556
Seguy E (1936) Code universel des couleurs. P. Lechevalier, Paris
Senanayake IC, Maharachchikumbura SSN, Hyde KD, Bhat DJ, Jones EBG,
McKenzie EHC, Dai DQ, Daranagama DA, Dayarathne MC, Goonasekara ID,
Konta S, Li WJ, Shang QJ, Stadler M, Wijayawardene NN, Xiao YP,
Norphanphoun C, Li QR, Liu XZ, Bahkali AH, Kang JC, Wang Y, Wen TC,
Wendt L, Xu JC, Camporesi E. (2015) Towards unraveling relationships in
Xylariomycetidae (Sordariomycetes). Fungal Divers 73:73–144
Sharma JR (1995) Hymenochaetaceae of India. Botanical Survey of India, Calcutta
Sharma S, Chadha BS, Kaur M, Ghatora SK, Saini HS (2008) Molecular
characterization of multiple xylanase producing thermophilic/thermotolerant
fungi isolated from composting materials. Let Appl Microbiol 46:526–535
Shearer CA (1989) Aniptodera (Halosphaeriaceae) from wood in freshwater habitats.
Mycologia 81:139146
Shearer CA, Miller M. (1977) Fungi of the Chesapeake Bay and its tributaries V.
Aniptodera chesapeakensis gen. et sp. nov. Mycologia 69:887–897
Shoemaker RA (1964) Seimatosporium (= Cryptostictis) parasites of Rosa, Vitis, and
Cornus. Can J Bot 42:411−417
Shoemaker RA, Muller E (1964) Generic correlations and concepts: Clathridium (=
Griphosphaeria) and Seimatosporium (= Sporocadus). Can J Bot 42:403−407
Silvestro D, Michalak I (2011) RaxmlGUI: A graphical front-end for RAxML. Org
Divers Evol 12:335–337
Simmons D R, Kepler R M, Rehner S A, Groden E (2015) Phylogeny of Hirsutella
species (Ophiocordycipitaceae) from the USA: remedying the paucity of
Hirsutella sequence data. IMA Fungus 6(2):345-356
Singer R (1940) Notes sur quelques Basidiomycètes. Rev Mycol 5:3–13
Singer R (1947) The Boletoideae of Florida. The Boletineae of Florida with notes on
extralimital species III. Am Midl Nat 37:1–135
Singer R (1986) The Agaricales in modern taxonomy. 4th ed. Koeltz Scientific Books:
Koenigstein
Singh SK, Doshi A, Pancholy A, Pathak R (2013) Biodiversity in wood–decay
macro–fungi associated with declining arid zone trees of India as revealed by
nuclear rDNA analysis. Eur J Plant Pathol. 136(2):373-382
Singtripopa C, Camporesi E, Ariyawansa HA, Wanasinghe DN, Bahkali AH,
Chomnunti P, Boonmee S, Hyde KD (2015) Keissleriella dactylidis, sp. nov.,
from Dactylis glomerata and its phylogenetic placement. Science Asia
41:295–304
Sir EB, Kuhnert E, Lambert C, Hladki AI, Romero AI, Stadler M (2016) New species
and reports of Hypoxylon from Argentina recognized by a polyphasic approach.
Mycol Prog, in press (DOI: 10.1007/s11557-016-1182-z).
Sivanesan A (1984) The bitunicate ascomycetes and their anamorphs. J. Cramer,
Vaduz
Slippers B, Boissin E, Phillips AJL, Groenewald JZ, Lombard L, Wingfield MJ,
Postma A, Burgess T, Crous PW (2013) Phylogenetic lineages in the
9557
9558
9559
9560
9561
9562
9563
9564
9565
9566
9567
9568
9569
9570
9571
9572
9573
9574
9575
9576
9577
9578
9579
9580
9581
9582
9583
9584
9585
9586
9587
9588
9589
9590
9591
9592
9593
9594
9595
9596
9597
9598
9599
Botryosphaeriales: a systematic and evolutionary framework. Stud Mycol
76:31–49
Slippers B, Roux J, Wingfield MJ, van der Walt, FJJ, Jami F, Mehl JWM, Marais GJ
(2014) Confronting the constraints of morphological taxonomy in the
Botryosphaeriales. Persoonia 33:55–168
Smith AH, Thiers HD (1971) The Boletes of Michigan. University of Michigan Press,
Ann Arbor, USA
Smith JDG, Liew ECY, Hyde KD (2001) The Xylariales: a monophyletic order
containing 7 families. Fungal Divers 13:175–208
Smith ME, Gryganskyi A, Bonito G, Nouhra E, Moreno-Arroyo B, Benny G (2013)
Phylogenetic analysis of the genus Modicella reveals an independent
evolutionary origin of sporocarp-forming fungi in the Mortierellales. Fung Gen
Biol 61:61–68
Sogonov MV, Castlebury LA, Rossman AY, Mejía LC, White JF, Jr (2008).
Leaf-inhabiting genera of the Gnomoniaceae, Diaporthales. Stud Mycol 62:1–79
Somrithipol S, Sakayaroj J, Rungjindamai N, Plaingam N, Jones EBG (2008)
Phylogenetic relationship of the coelomycete genus Infundibulomyces based on
nuclear rDNA data. Mycologia 100:735–741
Song J, Chen Y, Cui B, Liu H, Wang Y (2014) Morphological and molecular
evidence for two new species of Laetiporus (Basidiomycota, Polyporales) from
southwestern China. Mycologia 106:1039–1050
Spatafora JW, Quandt CA, Kepler RM, Sung GH, Shrestha B, Hywel-Jones NL,
Luangsa-ard J J (2015) New 1F1N species Combinations in Ophiocordycipitaceae
(Hypocreales). IMA Fungus 6(2):357–362
Spatafora JW, Sung GH, Sung JM, Hywel-Jones NL, White JF (2007) Phylogenetic
evidence for an animal pathogen origin of ergot and the grass endophytes. Mol
Ecol 16:1701–1711
Spatafora J W, Quandt C A, Kepler R M, Sung G H, Shrestha B, Hywel-Jones N L,
Luangsa-ard J J (2015) New 1F1N species Combinations in
Ophiocordycipitaceae (Hypocreales). Phytopathology 6(2):357–362
Spielman LJ (1983) Taxonomy and Biology of Valsa Species on Hardwoods in North
America, with Special Reference to Species on Maples. Cornell University, NY,
USA
Spielman LJ (1985) A monograph of Valsa on hardwoods in North America. Can J
Bot 63:1355–1378
Spooner BM. (1981). New records and species of British microfungi. Trans Brit
Mycol Soc 76:265–301
Stadler M (2011) Importance of secondary metabolites in the Xylariaceae as
parameters for assessment of their taxonomy, phylogeny, and functional
biodiversity. Curr Res Environ Appl Mycol 1:75–133
Stadler M, Kuhnert E, Peršoh D, Fournier J (2013) The Xylariaceae as model
example for a unified nomenclature following the “One Fungus-One Name”
(1F1N) concept. Mycology 4(1):5–21
9600
9601
9602
9603
9604
9605
9606
9607
9608
9609
9610
9611
9612
9613
9614
9615
9616
9617
9618
9619
9620
9621
9622
9623
9624
9625
9626
9627
9628
9629
9630
9631
9632
9633
9634
9635
9636
9637
9638
9639
9640
9641
9642
9643
Stamatakis A (2006) RAxML-VI-HPC: maximum likelihood-based phylogenetic
analyses with thousands of taxa and mixed models. Bioinformatics
22:2688–2690
Stamatakis A (2014) RAxML version 8: a tool for phylogenetic analysis and
post-analysis of large phylogenies. Bioinformatics 30 (9):1312–1313
Stamatakis A, Hoover P, Rougemont J (2008) A rapid bootstrap algorithm for the
RAxML web servers. Syst Biol 57:758–777
Stubbe D, Nuytinck J, Verbeken A (2010) Critical assessment of the Lactarius
gerardii species complex (Russulales). Fungal Biol 114 (2-3):271–283
Su, HY, Hyde KD, Maharachchikumbura SSN, Ariyawansa HA, Luo ZL, Promputtha
I, Tian Q, Lin CG, Shang QJ, Zhao YC, Chai HM, Liu XY, Bahkali AH, Bhat
JD, McKenzie EHC, Zhou DQ (2016) The families Distoseptisporaceae fam.
nov., Kirschsteiniotheliaceae, Sporormiaceae and Torulaceae, with new species
from freshwater in Yunnan Province, China. Fingal Divers: DOI
10.1007/s13225-016-0362-0
Subrahamanyam A (1983) Studies on themomycology. Mucor thermo-hyalospora sp.
nov. Bibl Mycol 91:421–423
Suetrong S, Schoch CL, Spatafora JW, Kohlmeyer J, Volkmann-Kohlmeyer B,
Sakayaroj J, Phongpaichit S, Tanaka K, Hirayama K, Jones EBG (2009)
Molecular systematics of the marine Dothideomycetes. Stud Mycol 64:155–173
Sung GH, Hywel-Jones NL, Sung JM, Luangsa-ard JJ, Shrestha B, Spatafora JW
(2007) Phylogenetic classification of Cordyceps and the clavicipitaceous fungi.
Stud Mycol 57:5–59
Šutara J (2008) Xerocomus s.l. in the light of the present state of knowledge. Czech
Mycol 60:29–62
Sutton BC (1980) The Coelomycetes. Fungi imperfecti with pycnidia, acervuli and
stromata. Commonwealth Mycological Institute, Kew.
Swofford DL (2002) PAUP: phylogenetic analysis using parsimony, version 4.0b10.
Illinois Natural History Survey, Champion, III
Tadych M, Chaverri P, Bergen M, White JF (2009) Moelleriella zhongdongii: stroma
development and identification of hirsutella-like and Aschersonia synanamorphs.
Mycol Res 113:611–615
Tanabe Y, Saikawa M, Watanabe MM, Sugiyama J (2003) Molecular phylogeny of
Zygomycota based on EF-1alpha and RPB1 sequences: limitations and utility of
alternative markers to rDNA. Mol Phylogenet Evol 30:438–449
Tanaka K, Harada Y (2003). Pleosporales in Japan (2): the genus Lophiotrema.
Mycoscience 44:115–121
Tanaka K, Hirayama K, Yonezawa H, Hatakeyama S, Harada Y, Sano T, Shirouzu T,
Hosoya T (2009) Molecular taxonomy of bambusicolous fungi:
Tetraplosphaeriaceae, a new pleosporalean family with Tetraploa-like
anamorphs. Stud Mycol 64:175-209
Tanaka K, Endo M, Hirayama K, Okane I, Hosoya T, Sato T (2011) Phylogeny of
Discosia and Seimatosporium, and introduction of Adisciso and Immersidiscosia
genera nova. Persoonia 26:85−98
9644
9645
9646
9647
9648
9649
9650
9651
9652
9653
9654
9655
9656
9657
9658
9659
9660
9661
9662
9663
9664
9665
9666
9667
9668
9669
9670
9671
9672
9673
9674
9675
9676
9677
9678
9679
9680
9681
9682
9683
9684
9685
Tanaka K, Hirayama K, Yonezawa H, Sato G, Toriyabe A, Kudo H, Hashimoto A,
Matsumura M, Harada Y, Kurihara Y, Shirouzu T, Hosoya T, (2015) Revision of
the Massarineae (Pleosporales, Dothideomycetes). Stud Mycol 82:75–136
Tang LP, Hao YJ, Cai Q, Tolgor B, Yang ZL (2014) Morphological and molecular
evidence for a new species of Rhodotus from tropical and subtropical Yunnan,
China. Mycol Progr 13:45–53
Teng SC (1936) Additional fungi from China IV. Sinensia 7:752–823
Teunissen MJ, Op den Camp HJ, Orpin CG, Huis in ’t Veld JH, Vogels GD (1991)
Comparison of growth characteristics of anaerobic fungi isolated from ruminant
and non-ruminant herbivores during cultivation in a defined medium. J Gen
Microbiol 137:1401–1408
Thambugala KM, daranagama DA, Camporesi E, Singtripop C, Liu ZY, Hyde KD
(2014a) Multi-locus phylogeny reveals the sexual state of Tiarosporella in
Botryosphaeriaceae. Cryptogam Mycol 35:359–367
Thambugala KM, Ariyawansa HA, Li YM, Boonmee S, Hongsanan S, Tian Q,
Singtripop C, Bhat DJ, Camporesi E, Jayawardena R, Liu ZY, Xu JC,
Chukeatirote E, Hyde KD (2014b) Dothideales. Fungal Diversity 68(1):105–158
Thambugala KM, Ariyawansa HA, Liu ZY, Chukeatirote E, Hyde KD (2014c)
Towards a natural classification of Dothideomycetes 6: the genera Dolabra,
Placostromella, Pleosphaerellula, Polysporidiella and Pseudotrichia
(Dothideomycetes Incertae sedis). Phytotaxa 176(1):55–67
Thambugala KM, Hyde KD, Eungwanichayapant PD, Romero AI, Liu ZY (2016)
Additions to the genus Rhytidhysteron in Hysteriaceae. Cryptog Mycolog (in
press)
Thambugala KM, Hyde KD, Tanaka K, Tian Q, Wanasinghe DN, Ariyawansa, HA,
Jayasiri SC, Boonmee S, Camporesi E, Hashimoto A, Hirayama K, Schumacher
RK, Promputtha I, Liu ZY (2015a) Towards a natural classification and
backbone tree for Lophiostomataceae, Floricolaceae, and Amorosiaceae fam.
nov. Fungal Divers 74:199–266
Thambugala KM, Chunfang Y, Camporesi E, Bahkali AH, Liu ZY, Hyde KD (2015b).
Pseudodidymosphaeria gen. nov. in Massarinaceae. Phytotaxa 231(3):271–282
Theissen F, Sydow H (1917) Synoptische Tafeln. Ann Mycol 15(6):389–491
Tian Q, Hyde KD, Liu JK, Wanasinghe DN, Boonmee S., Senanayak IC, Luo ZL,
Ariyawansa H, Li WJ, Thambugala KM, Jones EBG, Bhat DJ, Bahkali AH,
Chomnunti P, Mortimer PE, Xu JC, Campesori E (2015) Phylogenetic
relationships and morphological reappraisal of Melanommataceae. Fungal
Divers 74(1):267–324
Traaen AE (1914) Untersuchungen über Bodenpilze aus Norwegen. Nytt Magazin for
Naturvidenskapene 52:1–121
Trakunyingcharoen T, Lombard L, Groenewald JZ, Cheewangkoon R, Toanun C,
Alfenas AC, Crous PW (2014) Mycoparasitic species of Sphaerellopsis, and
allied lichenicolous and other genera. IMA Fungus 5(2):391–414
9686
9687
9688
9689
9690
9691
9692
9693
9694
9695
9696
9697
9698
9699
9700
9701
9702
9703
9704
9705
9706
9707
9708
9709
9710
9711
9712
9713
9714
9715
9716
9717
9718
9719
9720
9721
9722
9723
9724
9725
9726
9727
9728
Trouillas FP, Úrbez-Torres JR, Gubler WD (2010) Diversity of diatrypaceous fungi
associated with grapevine canker diseases in California. Mycologia
102(2):319–336
Trouillas FP, Pitt WM, Sosnowski MR, Huang R, Peduto F, Loschiavo A, Savocchia
S, Scott ES, Gubler WD (2011) Taxonomy and DNA phylogeny of Diatrypaceae
associated with Vitis vinifera and other woody plants in Australia. Fungal Divers
49(1):203–223
Tuckwell DS, Nicholson MJ, McSweeney CS, Theodorou MK, Brookman JL (2005)
The rapid assignment of ruminal fungi to presumptive genera using ITS1 and
ITS2 RNA secondary structures to produce group-specific fingerprints.
Microbiol 151:1557–1567
Tulasne LR, Tulasne C. (1861) Selecta Fungorum Carpologia
Tulloss, R.E., Yang Z.L. (2016) Amanitaceae studies (http://www.amanitaceae.org)
Upadhyay HP (1969) Soil fungi from North-East and North Brazil–VII. The genus
Gongronella. Nova Hedwigia 17:65–73
Van de Putte K, Nuytinck J, Stubbe D, Huyen TL, Verbeken A (2010) Lactarius
volemus sensu lato (Russulales) from northern Thailand: morphological and
phylogenetic species concepts explored. Fungal Divers 45 (1):99–130
van Tieghem P (1876) Sur le développement du fruit des Ascodesmis, genre nouveau
de l'ordre des Ascomycètes. Bull Soc Bot Fr 23:271–279
Vasilyeva LN, Stephenson SL (2005) Pyrenomycetes of the Great Smoky Mountains
National Park. II. Cryptovalsa Ces. et De Not. and Diatrypella (Ces. et De Not.)
Nitschke (Diatrypaceae). Fungal Divers 19:189–200
Vasilyeva LN, Mel'nik VA (2006) New pyrenomycetous species from Korea and South
Africa. Mikol Fitopatol 40(2):89–92
Vavra J, Joyon L (1966) Étude sur la morphologie, le cycle évolutif et la position
systematique de Callimastix cyclopsis Weissenberg 1912. Protistologica 2:15–16
Vizzini A (2014) Nomenclatural novelties. Index Fungorum 176:1.
Vizzini A, Contu M, Ercole E (2011) Musumecia gen. nov. in the Tricholomatoid
clade (Basidiomycota, Agaricales) related to Pseudoclitocybe. Nord J Bot
29:734–740
Vizzini A, Consiglio G, Ercole E, Setti L (2016) Pseudoporpoloma, a new genus for
Agaricus pes-caprae (Agaricales, Tricholomataceae). Phytotaxa 243:271–280
Vlasák J, Kout J (2011) Pileate Fomitiporia species in the USA. New combinations
Fomitiporia calkinsii and F. bakeri. Mycol Prog 10:445–452
Vlasák J, Li HJ, Zhou LW, Dai YC (2013) A further study on Inonotus linteus
complex (Hymenochaetales, Basidiomycota) in tropical America. Phytotaxa 124
(1):25–36
Wagner T, Fischer M (2002) Proceedings towards a natural classification of
theworldwide taxa Phellinus s.l. and Inonotus s.l., and phylogenetic relationships
of allied genera. Mycologia 94:998–1016
Wagner L, Stielow B, Hoffmann K, Petkovits T, Papp T, Vágvölgyi C, de Hoog GS,
Verkley G, Voigt K (2013) A comprehensive molecular phylogeny of the
9729
9730
9731
9732
9733
9734
9735
9736
9737
9738
9739
9740
9741
9742
9743
9744
9745
9746
9747
9748
9749
9750
9751
9752
9753
9754
9755
9756
9757
9758
9759
9760
9761
9762
9763
9764
9765
9766
9767
9768
9769
9770
9771
9772
Mortierellales (Mortierellomycotina) based on nuclear ribosomal DNA.
Persoonia 30:77–93
Walther G, Pawłowska J, Alastruey-Izquierdo A, Wrzosek M, Rodriguez-Tudela JL,
Dolatabadi S, Chakrabarti A, de Hoog GS (2013) DNA barcoding in Mucorales:
an inventory of biodiversity. Persoonia 30:11–47
Wanasinghe DN, Jones EBG, Camporesi E, Boonmee S, Ariyawansa HA,
Wijayawardene NN, Hyde KD (2014) An Exciting Novel Member of
Lentitheciaceae in Italy from Clematis Vitalba. Cryptogam Mycol
35(4):323–337
Wanasinghe DN, Jones EBG, Camporesi E, Dissanayake AJ, Kamolhan S, Mortimer
PE, Xu JC, Abd-Elsalam K, Hyde KD (2016) Taxonomy and phylogeny of
Laburnicola gen. nov. (Didymosphaeriaceae, Massarinae). Fungal Biol (in
press)
Wang DM, Wu SH, Li TH. (2009a) Two records of Ganoderma new to mainland
China. Mycotaxon 108:35–40
Wang J, Zhou W, Yuan H, Wang Y (2008) Characterization of a novel fungal
chitosanase Csn2 from Gongronella sp. JG. Carbohydr Res 343:2583–2588
Wang XH, Yang ZL, Li YC, Knudsen H, Liu PG (2009b) Russula griseocarnosa sp.
nov. (Russulaceae, Russulales), a commercially important edible mushroom in
tropical China: mycorrhiza, phylogenetic position, and taxonomy. Nova Hedwig
88:269–282
Wang YW, Hong TW, Tai YL, Wang YJ, Tsai SH, Lien PTK, Chou TH, Lai JY, Chu R,
Ding ST, Irie K, Li TK, Tzean SS, Shen TL (2015a) Evaluation of an epitypified
Ophiocordyceps formosana (Cordycepss.l.) for its pharmacological potential.
Evid Based Complement Alternat Med 2015:13
Wang ZR, Parra L, Callac P, Zhou JL, Fu WJ, Dui SH, Hyde KD, Zhao RL (2015b)
Edible species of Agaricus (Agaricaceae) from Xinjiang Province (Western
China). Phytotaxa 202:185–197
Watanabe T (1994) Two new species of homothallic Mucor in Japan. Mycologia
86:691–695
Webb J, Theodorou MK (1991) Neocallimastix hurleyensis sp. nov., an anaerobic
fungus from the ovine rumen. Can J Bot 69:1220–1224
Weerakoon G (2015) Fascinating Lichens of Sri Lanka. Dilmah Conservation,
Colombo, Sri Lanka.
Weerakoon G, Aptroot A (2013) Some new lichen species from Sri Lanka, with a key
to the genus Heterodermia in Sri Lanka. Cryptogam Mycol 34:321–328
Weerakoon G, Aptroot A (2014) Over 200 new lichen records from Sri Lanka, with
three new species to science. Cryptogamie Mycol 35:51–62
Weerakoon G, Aptroot A, Lumbsch HT, Wolseley PA, Wijeyaratne SC, Gueidan C
(2012a). New molecular data on Pyrenulaceae from Sri Lanka reveal two
well-supported groups within this family. Lichenologist 44:639–647
Weerakoon G, Rivas Plata E, Lumbsch HT, Lücking R (2012b) Three new species of
Chapsa (lichenized Ascomycota: Ostropales: Graphidaceae) from tropical Asia.
Lichenologist 44:373–379.
9773
9774
9775
9776
9777
9778
9779
9780
9781
9782
9783
9784
9785
9786
9787
9788
9789
9790
9791
9792
9793
9794
9795
9796
9797
9798
9799
9800
9801
9802
9803
9804
9805
9806
9807
9808
9809
9810
9811
9812
9813
9814
9815
Weerakoon G, Wijeyaratne SC, Wolseley PA, Rivas Plata E, Lücking R, Lumbsch
HT (2012c) Six new species of Graphidaceae from Sri Lanka. Bryologist
115:74–83
Weerakoon G, Lücking R, Lumbsch HT (2014) Thirteen new species of
Graphidaceae (lichenized Ascomycota: Ostropales) from Sri Lanka. Phytotaxa
189:331–347
Weerakoon G, Jayalal U, Wijesundara S, Karunaratne V, Lücking R (2015) Six new
Graphidaceae (lichenized Ascomycota: Ostropales) from Horton Plains National
Park, Sri Lanka. Nova Hedwigia 101:77–88
Wehmeyer LE (1957) The genus Saccothecium, Pringsheimia, Pleosphaerulina and
Pseudoplea. Mucologia 49:83–94
Wei F, Hong Y, Liu J, Yuan J, Fang W, Peng H, Xiao Y (2010) Gongronella sp.
induces overproduction of laccase in Panus rudis. J Basic Microbiol50:98–103
Wen TC, Zhu RC, Kang JC, Huang MH, Tan DB, Ariyawansha H, Liu H (2013)
Ophiocordyceps xuefengensis sp. nov. from larvae of Phassusnodus (Hepialidae)
in Hunan Province, southern China. Phytotaxa 123(1):41–50
Westendorp GD (1857) Sur quelques hypoxylées inédites ou nouvelles pour la flore
de la Belgique. Bulletin de l’Académie Royale de Belgique Classe de Sciences,
Série 22:554–579
Wicklow DT, Joshi BK, Gamble WR, Gloer JB, Dowd PF (1998) Antifungal
metabolites (monorden, monocillin IV, and cerebrosides) from Humicola
fuscoatra Traaen NRRL 22980, a mycoparasite of Aspergillus flavus sclerotia.
Appl Environ Microbiol 64:4482–4484
Wijayawardene DNN, McKenzie EHC, Hyde KD (2012) Towards incorporating
anamorphic fungi in a natural classification—checklist and notes for 2011.
Mycosphere 3:157–228
Wijayawardene NN, Hyde KD, Bhat DJ, Camporesi E, Schumacher RK, Chethana
KT, Wikee S, Bahkali AH, Wang Y (2014a) Camarosporium-like species are
polyphyletic in Pleosporales; introducing Paracamarosporium and
Pseudocamarosporium gen. nov. in Montagnulaceae. Cryptogam Mycol
35(2):177–198
Wijayawardene NN, Crous PW, Kirk PM, Hawksworth DL, Boonmee S, Braun U,
Chomnunti P, Dai DQ, D’Souza MJ, Diederich P, Dissanayake A, Doilom M,
Hongsanan S, Jones EBG, Groenewald JZ, ayawardena R, Lawrey JD, Liu JK,
Lucking R, Madrid H, Manamgoda DS, Muggia L, Nelsen MP, Phookamsak R,
Suetrong S, Tanaka K,Thambugala KM, Wikee S, Zhang Y, Aptroot A,
Ariyawansa HA, Bahkali AH, Bhat JD, Gueidan C, DeHoog GS, Knudsen K,
Mckenzie EHC, Miller AN, Mortimer PE, Wanasinghe DN, Phillips AJL, Raja
HA, Slippers B, Shivas RS, Taylor JE, Wang Y, Woudenberg JHC, Piatek M,
Cai L, Jaklitsch WM, Hyde KD. (2014b) Naming and outline of
Dothideomycetes–2014. Fungal Divers 69:1–55
Wilson AW, Desjardin DE (2015) Phylogenetic relationships in the gymnopoid and
marasmioid fungi (Basidiomycetes, euagarics clade). Mycologia 97:667–679
9816
9817
9818
9819
9820
9821
9822
9823
9824
9825
9826
9827
9828
9829
9830
9831
9832
9833
9834
9835
9836
9837
9838
9839
9840
9841
9842
9843
9844
9845
9846
9847
9848
9849
9850
9851
9852
9853
9854
9855
9856
9857
9858
9859
Wingfield MJ, De Beer ZW, Slippers B, Wingfield BD, Groenewald JZ, Lombard L,
Crous PW (2012) One fungus, one name promotes progressive plant pathology.
Mol Plant Pathol 13:604–613
Winter G (1885) Pilze - Ascomyceten. In GL Rabenhorst's Kryptogamen-Flora von
Deutschland, Oesterreich und der Schweiz 1:65–528
Winter G (1886) Rabenhorst's Kryptogamen-Flora, Pilze - Ascomyceten 1(2):570pp
Whalley AJS, Jones EBG, Hyde KD, Læssøe T. (1999). Halorosellinia gen. nov. to
accommodate Hypoxylon oceanicum,a common mangrove species. Mycol. Res.
104 (3):368–374
White WL, Downing MH (1953) Humicola grisea, a soil-inhabiting, cellulolytic
hyphomycete. Mycologia 45:951–963
White MM, James TY, O'Donnell K, Cafaro MJ, Tanabe Y, Sugiyama J (2006)
Phylogeny of the Zygomycota based on nuclear ribosomal sequence data.
Mycologia 98:872–884
Wu G, Feng B, Xu J, Zhu XT, Li YC, Zeng NK, Hosen MI, Yang ZL (2014)
Molecular phylogenetic analyses redefine seven major clades and reveal 22 new
generic clades in the fungal family Boletaceae. Fungal Divers 69(1):93–115
Wu SH, Dai YC, Hattori T, Yu TW, Wang DM, Parmasto E, Chang HY, Shih SY
(2012) Species clarification for the medicinally valuable ‘sanghuang’ mushroom.
Bot Stud 53:135–149
Yan JY, Xie Y, Zhang W, Wang Y, Liu JK, Hyde KD, Seem RC, Zhang GC, Wang
ZY, Yao SW, Bai XJ, Dissanayake AJ, Peng YL, Li XH (2013) Species of
Botryosphaeriaceae involved in grapevine dieback in China. Fungal Divers
61:221–236
Yang Q, Fan ZL, Crous PW, Liang YM, Tian CM (2015) Cytospora from Ulmus
pumila in Northern China. Mycol Prog 14:74–85
Yang X, Copes WE, Hong C (2014a). Two novel species representing a new clade
and cluster of Phytophthora. Fungal Biol 118:72–82
Yang X, Gallegly ME, Hong CX (2014b). A high-temperature tolerant species in
clade 9 of the genus Phytophthora: P. hydrogena sp. nov. Mycologia 106:57–65
Yang X, Hong CX (2013). Phytophthora virginiana sp. nov., a high-temperature
tolerant species from irrigation water in Virginia. Mycotaxon 126:167–176
Yang ZL. (1991). Clarkeinda trachodes, an agaric new to China. Acta Botanica
Yunnanica 13:279–282
Yang ZL (1997) Die Amanita-Arten von Suedwestchina. Bibl Mycol 170:1–240
Yang ZL, Doi Y (1999) A contribution to the knowledge of Amanita (Amanitaceae,
Agaricales) in Japan. Bull. Natl. Sci. Mus. Tokyo 25(3):108-130
Yang ZL, Li TH, Wu XL (2001) Revision of Amanita collections made from Hainan,
Southern China. Fungal Diversity 6:149-165
Yang ZL, Weiß M, Oberwinkler F (2004) New species of Amanita from the eastern
Himalaya and adjacent regions. Mycologia 96(3):636-646
Yang ZL, Li YC, Tang LP, Shi GQ, Zeng G (2012) Trogia venenata (Agaricales), a
novel poisonous species which has caused hundreds of deaths in southwestern
China. Mycol Prog 11:937–945
9860
9861
9862
9863
9864
9865
9866
9867
9868
9869
9870
9871
9872
9873
9874
9875
9876
9877
9878
9879
9880
9881
9882
9883
9884
9885
9886
9887
9888
9889
9890
9891
9892
9893
9894
9895
9896
9897
9898
9899
9900
9901
9902
Yang ZL, Feng B, Hao YJ (2013) Pseudoarmillariella bacillaris, a new species with
bacilliform basidiospores in Asia. Mycosystema 32:132–132
Yang ZL, Qin J, Xia CF, Hu Q, Li QQ (2015) Ophiocordyceps highlandensis, a new
entomopathogenic fungus from Yunnan, China. Phytotaxa 204:287–295
Zalar P, Hennebert GL, Gunde-Cimerman N, Cimerman A (1997) Mucor troglophilus,
a new species from cave crickets. Mycotaxon 65:507–516
Zang M, Li TH, Petersen RH (2001) Five new species of Boletaceae from China.
Mycotaxon 80:481–487
Zeng NK, Tang LP, Li YC, Tolgor B, Zhu XT, Zhao Q, Yang ZL (2013) The genus
Phylloporus (Boletaceae, Boletales) from China: morphological and multi-locus
DNA sequence analyses. Fungal Divers 58:73–101
Zeng NK, Wu G, Li YC, Liang ZQ, Yang ZL (2014) Crocinoboletus, a new genus of
Boletaceae (Boletales) with unusual boletocrocin polyene pigments. Phytotaxa
175:133–140
Zhang LF, Yang JB, Yang ZL (2004) Molecular phylogeny of eastern Asian species
of Amanita (Agaricales, Basidiomycota): taxonomic and biogeographic
implications. Fungal Divers 17:219–238
Zhang N, Castlebury LA, Miller AN, Huhndorf SM, Schoch CL, Seifert KA,
Rossman AY, Rogers JD, Kohlmeyer J, Volkmann-Kohlmeyer B, Sung GH
(2006) An overview of the systematics of the Sordariomycetes based on
four-gene phylogeny. Mycologia 98:1077–1088
Zhang P, Chen ZH, Xiao B, Tolgor B, Bao HY, Yang ZL (2010) Lethal amanitas of
East Asia characterized by morphological and molecular data. Fungal Divery
42:119–133
Zhang P, Yang ZL, Ge ZW (2005) Two new species of Ramaria from southwestern
China. Mycotaxcon 94:235–240
Zhang Y, Schoch CL, Fournier J, Crous PW, De Gruyter J, Woudenberg JHC,
Hirayama K, Tanaka K, Pointing SB, Spatafora JW, Hyde KD (2009a)
Multi-locus phylogeny of the Pleosporales: a taxonomic, ecological and
evolutionary re-evaluation. Stud Mycol 64:85–102
Zhang Y, Crous PW, Schoch CL, Hyde KD (2012) Pleosporales. Fungal Divers
53:1–221
Zhang Y, Zhang X, Fournier J, Chen J, Hyde KD (2014) Lindgomyces griseosporus, a
new aquatic ascomycete from Europe including new records. Mycoscience
55:43–48
Zhao GC, Zhao RL (2012) The Higher Microfungi from forests of Yunnan Province.
Yunnan Science and Technology Press, Kunming.
Zhao JD, Zhang XQ (2000) Flora Fungorum Sinicorum. Vol. 8. Ganodermataceae.
Science Press, Beijing
Zhao K, Wu G, Yang ZL (2014) A new genus, Rubroboletus, to accommodate
Boletus sinicus and its allies. Phytotaxa 188:61–77
Zhao RL, Yang YM, Zhao GC (2004) Pseudolachnella vermospora sp. nov. from
Yushania vigens in China. Fungal Divers 15:255–260
9903
9904
9905
9906
9907
9908
9909
9910
9911
9912
9913
9914
9915
9916
9917
9918
9919
9920
9921
9922
9923
9924
9925
9926
9927
9928
9929
9930
9931
Zhao RL, Karunarathna SC, Raspé O, Parra LA, Guinberteau J, Moinard M, Kesel
AD, Barroso G, Courtecuisse R, Hyde KD, Guelly AK, Desjardin DE, Callac P.
(2011) Major clades in tropical Agaricus. Fungal Divers 51:279–296
Zhao RL, Zhou JL, Chen J, Margaritescu S, Sánchez-Ramírez S, Hyde KD, Callac P,
Parra LA, Li GJ, Moncalvo J-M (2016) Divergence times and the reconstruction
of a standardized taxonomic system based on multi-gene sequences: case from
the genus Agaricus. Fungal Diversity. (accepted).
Zhou LW (2015a) Cylindrosporus flavidus gen. et comb. nov. (Hymenochaetales,
Basidiomycota) segregated from Onnia. Phytotaxa 219:276–282
Zhou LW (2015b) Four new species of Phylloporia (Hymenochaetales,
Basidiomycota) from tropical China with a key to Phylloporia species worldwide.
Mycologia 107:1184–1192
Zhou LW, Dai YC (2012) Phylogeny and taxonomy of Phylloporia
(Hymenochaetales): new species and a worldwide key to the genus. Mycologia
104:211–222
Zhou LW, Vlasák J, Decock C, Assefa A, Stenlid J, Abate D, Wu SH, Dai YC
(2016a) Global diversity and taxonomy of the Inonotus linteus complex
(Hymenochaetales, Basidiomycota): Sanghuangporus gen. nov., Tropicoporus
excentrodendri and T. guanacastensis gen. et spp. nov., and 17 new
combinations. Fungal Divers http://dx.doi.org/10.1007/s13225-015-0335-8
Zhou LW, Vlasák J, Qin WM, Dai YC (2016b) Global diversity and phylogeny of the
Phellinus igniarius complex (Hymenochaetales, Basidiomycota) with the
description of five new species. Mycologia 108:192–204
Zhou W, Yuan H, Wang J, Yao J (2008) Production, purification and characterization
of chitosanase produced by Gongronella sp. JG. Lett Appl Microbiol 46:49–54
Zogg H (1962) Die Hysteriaceaes. str. und Lophiaceae, unter besonderer
Berücksichtigung der mitteleuropäischen Formen. Beitr Kryptogamenflora
Schweiz Band 11:1–190