Fungal Diversity
DOI 10.1007/s13225-015-0324-y
Fungal diversity notes 1–110: taxonomic and phylogenetic
contributions to fungal species
Jian Kui Liu 1,3 & Kevin D. Hyde 1,2,3 & E. B. Gareth Jones 4,16 & Hiran A. Ariyawansa 2,3 &
Darbhe J. Bhat 6,7 & Saranyaphat Boonmee 3 & Sajeewa S. N. Maharachchikumbura 1,2,3 &
Eric H. C. McKenzie 5 & Rungtiwa Phookamsak 2,3 & Chayanard Phukhamsakda 2,3 &
Belle Damodara Shenoy 25 & Mohamed A. Abdel-Wahab 16,17 & Bart Buyck 8 & Jie Chen 3 &
K. W. Thilini Chethana 3,10 & Chonticha Singtripop 2,3 & Dong Qin Dai 2,3 & Yu Cheng Dai 9 &
Dinushani A. Daranagama 2,3 & Asha J. Dissanayake 3,10 & Mingkwan Doilom 2,3 &
Melvina J. D’souza 2,3 & Xin Lei Fan 11 & Ishani D. Goonasekara 3 & Kazuyuki Hirayama 14 &
Sinang Hongsanan 2,3 & Subashini C. Jayasiri 3 & Ruvishika S. Jayawardena 2,3,10 &
Samantha C. Karunarathna 2,3 & Wen Jing Li 2,3 & Ausana Mapook 2,3 &
Chada Norphanphoun 3 & Ka Lai Pang 22 & Rekhani H. Perera 2,3 & Derek Peršoh 23 &
Umpava Pinruan 26 & Indunil C. Senanayake 2,3 & Sayanh Somrithipol 26 &
Satinee Suetrong 27 & Kazuaki Tanaka 20 & Kasun M. Thambugala 1,2,3 & Qing Tian 2,3 &
Saowaluck Tibpromma 3 & Danushka Udayanga 2,3 & Nalin N. Wijayawardene 2,3,29 &
Dhanuska Wanasinghe 2,3 & Komsit Wisitrassameewong 3 & Xiang Yu Zeng 3 &
Faten A. Abdel-Aziz 17 & Slavomir Adamčík 18 & Ali H. Bahkali 16 & Nattawut Boonyuen 27 &
Timur Bulgakov 35 & Philippe Callac 37 & Putarak Chomnunti 2,3 & Katrin Greiner 12 &
Akira Hashimoto 13,20 & Valerie Hofstetter 15 & Ji Chuan Kang 30 & David Lewis 19 &
Xing Hong Li 10 & Xing Zhong Liu 36 & Zuo Yi Liu 1 & Misato Matsumura 20 &
Peter E. Mortimer 2 & Gerhard Rambold 12 & Emile Randrianjohany 24 & Genki Sato 20 &
Veera Sri-Indrasutdhi 27 & Cheng Ming Tian 11 & Annemieke Verbeken 28 &
Wolfgang von Brackel 21 & Yong Wang 29 & Ting Chi Wen 30 & Jian Chu Xu 31 & Ji Ye Yan 10 &
Rui Lin Zhao 36 & Erio Camporesi 32,33,34
Received: 7 January 2015 / Accepted: 14 February 2015
# School of Science 2015
* Zuo Yi Liu
gzliuzuoyi@163.com
8
Muséum national d’histoire naturelle, Dépt. Systématique et
évolution, UMR 7205, 75005 Paris, France
9
Institute of Microbiology & Beijing Key Laboratory for Forest Pest
Control, Beijing Forestry University, PO Box 61, Beijing 100083,
China
1
Guizhou Key Laboratory of Agricultural Biotechnology, Guizhou
Academy of Agricultural Sciences, Guiyang 550006, Guizhou,
China
10
2
Key Laboratory for Plant Diversity and Biogeography of East Asia,
Kunming Institute of Botany, Chinese Academy of Science,
Kunming 650201, Yunnan, China
Institute of Plant and Environment Protection, Beijing Academy of
Agriculture and Forestry Sciences, No. 9 of Shu Guang Hua Yuan
Zhong Lu, Haidian District, Beijing 10097, China
11
3
Institute of Excellence in Fungal Research and School of Science,
Mae Fah Luang University, Chiang Rai 57100, Thailand
The Key Laboratory for Silviculture and Conservation of Ministry
of Education, Beijing Forestry University, Beijing 100083, China
12
4
Department of Botany and Microbiology, King Saudi University,
Riyadh, Saudi Arabia
Mycology, University of Bayreuth, Universitätsstraße 30,
95440 Bayreuth, Germany
13
5
Manaaki Whenua Landcare Research, Private Bag 92170,
Auckland, New Zealand
The United Graduate School of Agricultural Sciences, Iwate
University, 18-8 Ueda 3 chome, Morioka 020-8550, Japan
6
Formerly Department of Botany, Goa University, Goa, India
14
7
No. 128/1–J, Azad Housing Society, Curca, Goa Velha, India
Apple Experiment Station, Aomori Prefectural Agriculture and
Forestry Research Center, 24 Fukutami, Botandaira,
Kuroishi, Aomori 036-0332, Japan
Fungal Diversity
Abstract This paper is a compilation of notes on 110 fungal
taxa, including one new family, 10 new genera, and 76 new
species, representing a wide taxonomic and geographic range.
The new family, Paradictyoarthriniaceae is introduced based
on its distinct lineage in Dothideomycetes and its unique morphology. The family is sister to Biatriosporaceae and
Roussoellaceae. The new genera are Allophaeosphaeria
(Phaeosphaeriaceae), Amphibambusa (Amphisphaeriaceae),
Brunneomycosphaerella (Capnodiales genera incertae cedis),
Chaetocapnodium (Capnodiaceae), Flammeascoma
(Anteagloniaceae), Multiseptospora (Pleosporales genera
incertae cedis), Neogaeumannomyces (Magnaporthaceae),
Palmiascoma (Bambusicolaceae), Paralecia
(Squamarinaceae) and Sarimanas (Melanommataceae). The
newly described species are the Ascomycota Aliquandostipite
manochii, Allophaeosphaeria dactylidis, A. muriformia,
Alternaria cesenica, Amphibambusa bambusicola,
Amphisphaeria sorbi, Annulohypoxylon thailandicum,
Atrotorquata spartii, Brunneomycosphaerella laburni,
Byssosphaeria musae, Camarosporium aborescentis,
C. aureum, C. frutexensis, Chaetocapnodium siamensis,
Chaetothyrium agathis, Colletotrichum sedi, Conicomyces
pseudotransvaalensis, Cytospora berberidis, C. sibiraeae,
D i a p o r t h e t h u n b e rg i i c o l a , D i a t r y p e p a l m i c o l a ,
Dictyosporium aquaticum, D. meiosporum, D. thailandicum,
Didymella cirsii, Dinemasporium nelloi, Flammeascoma
bambusae, Kalmusia italica, K. spartii, Keissleriella
sparticola, Lauriomyces synnematicus, Leptosphaeria ebuli,
Lophiostoma pseudodictyosporium, L. ravennicum,
Lophiotrema eburnoides, Montagnula graminicola,
Multiseptospora thailandica, Myrothecium macrosporum,
Natantispora unipolaris, Neogaeumannomyces bambusicola,
Neosetophoma clematidis, N. italica, Oxydothis atypica,
Palmiascoma gregariascomum, Paraconiothyrium nelloi,
P. thysanolaenae, Paradictyoarthrinium tectonicola,
Paralecia pratorum, Paraphaeosphaeria spartii,
Pestalotiopsis digitalis, P. dracontomelon, P. italiana,
P h a e o i s a r i a p s e ud o c l e m a t i d i s, P h r a g m o c a p n i a s
p h i l i p p i n e n s i s , P s e u d o c a m a ro s p o r i u m c o t i n a e ,
Pseudocercospora tamarindi, Pseudotrichia rubriostiolata,
P. thailandica, Psiloglonium multiseptatum, Saagaromyces
mangrovei, Sarimanas pseudofluviatile, S. shirakamiense,
Tothia spartii, Trichomerium siamensis, Wojnowicia
dactylidicola, W. dactylidis and W. lonicerae. The
Basidiomycota Agaricus flavicentrus, A. hanthanaensis,
A. parvibicolor, A. sodalis, Cantharellus luteostipitatus,
Lactarius atrobrunneus, L. politus, Phylloporia dependens
and Russula cortinarioides are also introduce d.
15
Department of Plant Protection, Agroscope Changins-Wadenswil
Research Station ACW, Rte de Duiller, 1260 Nyon, Switzerland
27
16
Department of Botany and Microbiology, College of Science, King
Saud University, P.O. Box: 2455, Riyadh 1145, Saudi Arabia
Fungal Biodiversity Laboratory, BIOTEC, National Science and
Technology Development Agency (NSTDA), 113 Thailand Science
Park, Thanon Phahonyothin, Tombon Khlong Nueng, Amphoe
Khlong Luang, Pathum Thani 12120, Thailand
17
28
Department of Botany, Faculty of Science, Sohag University,
Sohag, Egypt
Research Group Mycology, Department of Biology,
Gent University, K.L. Ledeganckstraat 35, 9000 Gent, Belgium
29
18
Institute of Botany, Slovak Academy of Sciences, Dúbravská cesta
9, 84523 Bratislava, Slovakia
Department of Plant Pathology, Agriculture College, Guizhou
University, Guiyang 550025, People’s Republic of China
30
19
Hirosaki University, Bleakwood, 262 CR 3062,
Newton, Texas 75966, USA
20
Faculty of Agriculture and Life Sciences, Hirosaki University, 3
Bunkyo-cho, Hirosaki, Aomori 036-8561, Japan
The Engineering and Research Center of Southwest
Bio-Pharmaceutical Resource, Ministry of Education, Guizhou
University, Guiyang, Guizhou Province 550025, People’s Republic
of China
31
CAS Key Laboratory of Genome Sciences and Information, Beijing
Institute of Genomics, Chinese Academy of Sciences, No. 1
Beichen West Road, Chaoyang District, Beijing 100101, People’s
Republic of China
32
A.M.B. Gruppo Micologico Forlivese “Antonio Cicognani”, Via
Roma 18, Forlì, Italy
33
A.M.B. Circolo Micologico “Giovanni Carini”, C.P. 314,
Brescia, Italy
34
Società per gli Studi Naturalistici della Romagna, C.P. 144,
Bagnacavallo (RA), Italy
35
Academy of Biology and Biotechnology (Department of Biological
Sciences), Southern Federal University, Stachki str, 344090 Rostovna-Donu, Russia
36
The State Key Lab of Mycology, Institute of Microbiology, Chinese
Academic of Science, Beijing 100101, China
37
Mycology and Food Safety Research Unit, INRA,
Bordeaux, France
21
Institute for Vegetation Studies and Landscape Ecology, GeorgEger-Str. 1b, 91334 Hemhofen, Germany
22
Institute of Marine Biology and Center of Excellence for the
Oceans, National Taiwan Ocean University, 2 Pei-Ning Road,
Keelung 20224, Taiwan, Republic of China
23
Geobotany, Ruhr-Universität Bochum, Universitätsstraße 150,
44780 Bochum, Germany
24
Centre National de Recherche sur l’Environnement (CNRE), Lab.
de Microbiologie de l’Environnement (LME), BP 1739,
Antananarivo, Madagascar
25
CSIR-National Institute of Oceanography, Dona
Paula, Goa 403004, India
26
Microbe Interaction Laboratory, BIOTEC, National Science and
Technology Development Agency (NSTDA), 113 Thailand Science
Park, Thanon Phahonyothin, Tombon Khlong Nueng, Amphoe
Khlong Luang, Pathum Thani 12120, Thailand
Fungal Diversity
Epitypifications or reference specimens are designated for
Hapalocystis berkeleyi, Meliola tamarindi,
Pallidocercospora acaciigena, Phaeosphaeria musae,
Plenodomus agnitus, Psiloglonium colihuae, P. sasicola and
Zasmidium musae while notes and/or new sequence data are
provided for Annulohypoxylon leptascum, A. nitens,
A. stygium, Biscogniauxia marginata, Fasciatispora nypae,
Hypoxylon fendleri, H. monticulosum, Leptosphaeria
doliolum, Microsphaeropsis olivacea, Neomicrothyrium,
Paraleptosphaeria nitschkei, Phoma medicaginis and
Saccotheciaceae. A full description of each species is provided with light micrographs (or drawings). Molecular data is
provided for 90 taxa and used to generate phylogenetic trees
to establish a natural classification for species.
Keywords Ascomycota . Basidiomycota . Global Taxonomy
Initiative . Phylogeny . Taxonomy
Table of Contents
Amphisphaeriaceae sensu lato
1. Amphibambusa D.Q. Dai & K.D. Hyde, gen. nov.
2. Amphibambusa bambusicola D.Q. Dai & K.D. Hyde, sp.
nov.
3. Amphisphaeria sorbi Senanayake & K.D. Hyde, sp. nov.
4. Atrotorquata spartii Thambugala, Camporesi & K.D.
Hyde, sp. nov.
5. Oxydothis atypica Pinruan, sp. nov.
6. Pestalotiopsis digitalis Maharachch. & K.D. Hyde, sp.
nov.
7. Pestalotiopsis dracontomelon Maharachch. & K.D.
Hyde, sp. nov.
8. Pestalotiopsis italiana Maharachch, Camporesi & K.D.
Hyde, sp. nov.
Chaetosphaeriaceae
9. Conicomyces pseudotransvaalensis A. Hashim., G. Sato
& Kaz. Tanaka, sp. nov.
10. Dinemasporium nelloi W.J. Li, Camporesi & K.D.
Hyde, sp. nov.
Diaporthaceae
11. Diaporthe thunbergiicola Udayanga & K.D. Hyde, sp.
nov.
Diatrypaceae
12. Diatrype palmicola J.K. Liu & K.D. Hyde, sp. nov.
13. Phaeoisaria pseudoclematidis D.Q. Dai & K.D. Hyde,
sp. nov.
Glomerellaceae
14. Colletotrichum sedi Jayawardena, Bulgakov & K.D.
Hyde, sp. nov.
Halosphaeriaceae
15. Natantispora unipolaris K.L. Pang, S.Y. Guo & E.B.G.
Jones, sp. nov.
16. Saagaromyces mangrovei Abdel-Wahab, Bahkali &
E.B.G. Jones, sp. nov.
Hypocreales
17. Myrothecium macrosporum D.Q. Dai & K.D. Hyde, sp.
nov.
Magnaporthaceae
18. Neogaeumannomyces D.Q. Dai & K.D. Hyde, gen. nov.
19. Neogaeumannomyces bambusicola D.Q. Dai & K.D.
Hyde, sp. nov.
Meliolaceae
20. Meliola tamarindi Syd. & P. Syd.
Sydowiellaceae
21. Hapalocystis berkeleyi Auersw. ex Fuckel
Valsaceae
22. Cytospora berberidis C.M. Tian, X.L. Fan & K.D.
Hyde, sp. nov.
23. Cytospora sibiraeae C.M. Tian, X.L. Fan & K.D. Hyde,
sp. nov.
Xylariaceae
24.
25.
26.
27.
28.
29.
30.
31.
32.
Annulohypoxylon leptascum (Speg.) Y.M. Ju et al.
Annulohypoxylon nitens (Ces.) Y.M. Ju & J.D. Rogers
Annulohypoxylon stygium (Lév.) Y.M. Ju et al.
Annulohypoxylon thailandicum Daranagama & K.D.
Hyde, sp. nov.
Biscogniauxia marginata (Fr.) Pouzar
Fasciatispora nypae K.D. Hyde
Hypoxylon fendleri Berk. Ex Cooke
Hypoxylon lenormandii Berk. & M.A. Curtis apud Berk.
Hypoxylon monticulosum Mont.
Dothideomycetes
Anteagloniaceae
Fungal Diversity
33. Flammeascoma Phookamsak & K.D. Hyde, gen. nov.
34. Flammeascoma bambusae Phookamsak & K.D. Hyde,
sp. nov.
53. Paraphaeosphaeria spartii W.J. Li, Camporesi & K.D.
Hyde, sp. nov.
54. Pseudocamarosporium cotinae Norphanphoun,
Bulgakov & K.D. Hyde, sp. nov.
Bambusicolaceae
Hysteriaceae
35. Palmiascoma Phookamsak & K.D. Hyde, gen. nov.
36. Palmiascoma gregariascomum Phookamsak & K.D.
Hyde, sp. nov.
Capnodiaceae
37. Chaetocapnodium Hongsanan & K.D. Hyde, gen. nov.
38. Chaetocapnodium siamensis Hongsanan & KD Hyde,
sp. nov.
39. Phragmocapnias philippinensis Hongsanan & K.D.
Hyde, sp. nov.
Capnodiales genera incertae sedis
40. Brunneomycosphaerella Dissanayake J.K. Liu & K.D.
Hyde, gen. nov
41. Brunneomycosphaerella laburni Dissanayake J.K. Liu
& K.D. Hyde, sp. nov
Dictyosporaceae
42. Dictyosporium aquaticum Abdel-Aziz, sp. nov.
43. Dictyosporium meiosporum Boonmee & K.D. Hyde,
sp. nov.
44. Dictyosporium thailandicum D’ souza, D.J. Bhat &
K.D. Hyde sp. nov.
55. Psiloglonium colihuae (Lorenzo & Messuti) E. Boehm
56. Psiloglonium multiseptatum Phookamsak & K.D.
Hyde, sp. nov.
57. Psiloglonium sasicola (N. Amano) E. Boehm & C.L.
Schoch
Jahnulales
58. Aliquandostipite manochii Sri-indr., Boonyuen,
Suetrong, K.L. Pang & E.B.G. Jones, sp. nov.
Lentitheciaceae
59. Keissleriella sparticola Singtripop & K.D. Hyde, sp.
nov.
Leptosphaeriaceae
60. Leptosphaeria doliolum (Pers.) Ces. & De Not.
61. Leptosphaeria ebuli Jayasiri, Camporesi & K.D. Hyde,
sp. nov.
62. Paraleptosphaeria nitschkei (Rehm ex G. Winter)
De-Gruyter et al.
63. Plenodomus agnitus (Desm.) Gruyter et al.
Lophiostomataceae
Didymellaceae
45. Didymella cirsii Mapook, Camporesi & K.D. Hyde, sp.
nov.
46. Microsphaeropsis olivacea (Bonord.) Höhn.
47. Phoma medicaginis Malbr. & Roum.
64. Lophiostoma pseudodictyosporium Qing Tian,
Camporesi & K.D. Hyde, sp. nov.
65. Lophiostoma ravennicum Tibpromma, Camporesi &
K.D. Hyde, sp. nov.
Lophiotremataceae
Didymosphaeriaceae
48. Kalmusia italica Thambugala, Camporesi & K.D.
Hyde, sp. nov.
49. Kalmusia spartii Wanasinghe, Camporesi, E.B.G. Jones
& K.D. Hyde, sp. nov.
50. Montagnula graminicola Chethana, Thambugala,
Camporesi & K.D. Hyde, sp. nov.
51. Paraconiothyrium nelloi W.J. Li, Camporesi & K.D.
Hyde, sp. nov.
52. Paraconiothyrium thysanolaenae Phookamsak,
Chethana & K.D. Hyde, sp. nov.
66. Lophiotrema eburnoides Kaz. Tanaka, A. Hashim. & K.
Hiray., sp. nov.
Melanommataceae
67. Byssosphaeria musae Phookamsak & K.D. Hyde, sp.
nov.
68. Pseudotrichia rubriostiolata Phookamsak & K.D.
Hyde, sp. nov.
69. Pseudotrichia thailandica Phookamsak & K.D. Hyde,
sp. nov.
Fungal Diversity
70. Sarimanas Matsumura, K. Hiray. & Kaz. Tanaka, gen.
nov.
71. Sarimanas pseudofluviatile Matsumura, K. Hiray. &
Kaz. Tanaka, sp. nov.
72. Sarimanas shirakamiense Matsumura, K. Hiray. &
Kaz. Tanaka, sp. nov.
Microthyriales genus incertae sedis
73. Neomicrothyrium Boonmee, H.X. Wu & K.D. Hyde
Mycosphaerellaceae
74. Pallidocercospora acaciigena (Crous & M.J. Wingf.)
Crous & M.J. Wingf.
75. Pseudocercospora tamarindi Goon. & K.D. Hyde, sp.
nov.
76. Zasmidium musae (Arzanlou & Crous) Crous & U.
Braun
89. Alternaria cesenica Phukhamsakda, Qing Tian,
Camporesi & K.D. Hyde, sp. nov.
Pleosporales genera incertae sedis
90. Camarosporium aborescentis Phukhamsakda,
Bulgakov & K.D. Hyde, sp. nov.
91. Camarosporium aureum Norphanphoun, Bulgakov &
K.D. Hyde, sp. nov.
92. Camarosporium caraganicola Phukhamsakda,
Bulgakov & K.D. Hyde, sp. nov.
93. Multiseptospora Phookamsak & K.D. Hyde, gen. nov.
94. Multiseptospora thailandica Phookamsak & K.D.
Hyde, sp. nov.
Saccotheciaceae
95. Saccotheciaceae Bonord.
Venturiales
Paradictyoarthriniaceae
77. Paradictyoarthriniaceae Doilom, J.K. Liu & K.D.
Hyde, fam. nov.
78. Paradictyoarthrinium tectonicola Doilom & K.D.
Hyde, sp. nov.
96. Tothia spartii Qing Tian, Camporesi & K.D. Hyde, sp.
nov.
Eurotiomycetes
Chaetothyriales
Chaetothyriaceae
Phaeosphaeriaceae
79. Allophaeosphaeria Ariyawansa, Camporesi & K.D.
Hyde, gen. nov.
80. Allophaeosphaeria dactylidis Wanasinghe, Camporesi,
E.B.G. Jones & K.D. Hyde, sp. nov.
81. Allophaeosphaeria muriformia Ariyawansa, Camporesi
& K.D. Hyde, sp. nov.
82. Neosetophoma clematidis Wijayawardene, Camporesi
& K.D. Hyde, sp. nov.
83. Neosetophoma italica W.J. Li, Camporesi & K.D.
Hyde, sp. nov.
84. Phaeosphaeria musae Arzanlou & Crous
85. Wojnowicia dactylidicola Wijayawardene, Camporesi &
K.D. Hyde, sp. nov.
86. Wojnowicia dactylidis Wijayawardene, Camporesi and
K.D. Hyde, sp. nov.
87. Wojnowicia lonicerae Wijayawardene, Camporesi &
K.D. Hyde, sp. nov.
97. Chaetothyrium agathis Hongsanan & K.D. Hyde, sp.
nov.
Trichomeriaceae
98. Trichomerium siamensis Hongsanan & K.D. Hyde, sp.
nov.
Lecanorales
Squamarinaceae
99. Paralecia Brackel, Greiner, Peršoh & Rambold, gen.
nov.
100. Paralecia pratorum Brackel, Greiner, Peršoh &
Rambold, sp. nov.
Ascomycota genera, incertae sedis
101. Lauriomyces synnematicus Somrithipol, sp. nov.
Pleomassariaceae
Basidiomycota
88. Splanchnonema pupula (Fr.) Kuntze
Pleosporaceae
102. Agaricus flavicentrus Karunarathna & K.D. Hyde, sp.
nov.
Fungal Diversity
103. Agaricus hanthanaensis Karunarathna & K.D. Hyde,
sp. nov.
104. Agaricus parvibicolor L.J. Chen, R.L. Zhao & K.D.
Hyde, sp. nov.
105. Agaricus sodalis L.J. Chen, R.L. Zhao & K.D. Hyde,
sp. nov.
106. Cantharellus luteostipitatus Buyck, Randrianjohany
& V. Hofstetter, sp.nov.
107. Lactarius atrobrunneus Wisitrassameewong & K. D.
Hyde, sp. nov.
108. Lactarius politus Wisitrassameewong & K.D. Hyde,
sp. nov.
109. Phylloporia dependens Y.C. Dai, sp. nov.
110. Russula cortinarioides Buyck, Adamčík, Lewis & V.
Hofstetter, sp. nov.
Introduction
The estimated number of described fungi is 100,000
(including 17,500 lichens), but only approximately 1000
new species are described annually (Kirk et al. 2008).
The total number of fungi is however, estimated at between 700,000 to 1.5 million species (Hawksworth 1991,
2001; Schmit and Mueller 2007). The majority of the
undescribed fungal species are expected to be discovered
in poorly studied areas (Hyde et al. 2001), such as in
tropical forests or in under-explored habitats, for example
living on or in insects, plants or lichens (Hawksworth and
Rossman 1997; Fröhlich and Hyde 1999; Taylor et al.
2000a; Sipman and Aptroot 2001; Lawreya and Diederich
2003; Arnold and Lutzoni 2007). Phylogenetic studies are
revealing numerous new genera and species (Alves et al.
2004, 2006; Crous et al. 2006, 2009a, b; Phillips and Alves
2009; Phillips et al. 2008, 2013; de-Gruyter et al. 2010; Liu
et al. 2011, 2012; Maharachchikumbura et al. 2012). Before
molecular phylogenetics, most fungal species have been described based on their morphology and host associations and
thus the number of species is understandably underestimated.
Phylogenetic studies have distinctly indicated that numerous
morphologically similar taxa may be represented as distinct
lineages in different families (e.g. Botryosphaeriaceae, Liu
et al. 2012; Phaeosphaeriaceae, Phookamsak et al. 2014;
Didymosphaeriaceae, Ariyawansa et al. 2014b). Cryptic species of plant pathogens have been shown to comprise several
taxa (e.g. Colletotrichum Hyde et al. 2009, Sharma et al. 2014,
Pestalotiopsis, Maharachchikumbura et al. 2011, 2012, 2014,
Diaporthe, Udayanga et al. 2011, 2012).
Traditionally, systematic study papers and monographic
revisions have just one or a few authors, but the aforementioned collaborative initiatives have initiated a mind-shift with
a multi-authored, community-wide classification for the fungal kingdom (Hibbett et al. 2007). In addition, the
mycological community has several, regularly updated fundamental resources for species and higher level classification
and nomenclature available, such as Myconet (http://www.
fieldmuseum.org/myconet), Index Fungorum (http://www.
indexfungorum.org) and other CABI Bioscience databases,
and the Dictionary of the Fungi (Kirk et al. 2008).
Materials and methods
The phylogenetic analyses were performed based on up to
date ex-type, ex-epitype or otherwise authentic sequence data
available in GenBank as a concerted effort of multiple contributors listed in the authors section. New and reference species were sequenced based on the genomic DNA which was
extracted from the fresh mycelium except for lichenized and
lichenicolous fungi and fungi not readily cultivatable, specimens in this case were used for direct extraction of DNA.
Gene sequences and genetic markers used for each genus were
selected based on the current publications and have commonly
been used for each of the genera. The single gene sequence
alignments were initially aligned with ClustalX2 and improved in MAFFT V. 7.017 (Katoh et al. 2002) and BioEdit
7.0 (Hall 2004). Individual alignments were then concatenated
and used to construct the backbone trees of each group listed.
The phylogenetic analyses were performed for maximum parsimony in PAUP v. 4.0b10 (Swofford 2002), maximum likelihood in RAxML 7.4.2 Black Box or RAxMl GUI
(Stamatakis 2006; Stamatakis et al. 2008), PhyML 3.0
(Guindon et al. 2010) or Bayesian inference in MrBayes v.
3.1.2 (Huelsenbeck and Ronquist 2001) as specified in the
legend of each phylogenetic tree. The trees used to represent
each order, family and genus were analyzed by multiple contributors based on the selection of genes in given publications
under each description.
Results and discussion
The new species are described below in alphabetical order.
They represent a total of 67 genera in 33 families, 17 orders
and three classes in the Ascomycota. The individual taxon
entries are standardized as far as possible, but reflect in style
and content the diversity of participating authors and different
groups of fungi.
Contributions to Ascomycota
Sordariomycetes
A recent outline for the Class Sordariomycetes was provided
by Lumbsch and Huhndorf (2010), while there is presently no
Fungal Diversity
backbone tree and this is much needed. Below we introduce 20 species in the families Amphisphaeriaceae,
Chaetosphaeriaceae, Diaporthaceae, Diatrypaceae,
Glomerellaceae, Halosphaeriaceae, Magnaporthaceae,
Valsaceae and Xylariaceae.
Amphisphaeriaceae (sensu lato)
The family Amphisphaeriaceae is an important group of
ascomycetes within the order Xylariales. It was introduced by Winter (1887) to include Amphisphaeria and
related genera. Amphisphaeriaceae is characterised by
immersed ascomata in the host, and dark peridial walls
and ascal apices that are usually amyloid (Barr 1975).
The family mainly produces appendaged coelomycetous
asexual morphs. Amphisphaeriaceae is a relatively large
and heterogeneous family and is widely distributed
throughout tropical and temperate regions (Barr 1975; Kang
et al. 1999a, b). Several genera in the family are well known
for their ability to produce novel medicinal compounds (Xu
et al. 2010, 2014; Maharachchikumbura et al. 2012). Many
species of Amphisphaeriaceae cause a variety of disease in
plants and are often isolated as endophytes or saprobes (Nag
Raj 1993; Kang et al. 1998). In the present study, we introduce
a new genus and seven new species in the family
Amphisphaeriaceae based on molecular and morphological
characters. In this study we treat Amphisphaeriaceae in a wide
sense but expect it to be split into smaller family units. A
phylogenetic tree for amphisphaeriaceous genera (sensu lato)
is presented in Fig. 1.
1. Amphibambusa D.Q. Dai & K.D. Hyde, gen. nov.
Index Fungorum number: IF 550940, Facesoffungi
number: FoF00447
Etymology: In reference to a new genus in
Amphisphaeriaceae and its host Bambusa.
Type species: Amphibambusa bambusicola D.Q. Dai &
K.D. Hyde
Saprobic on decaying bamboo culms, forming black
circular spots on the host surface. Ascomata solitary,
scattered, immersed under host epidermis, globose to
subglobose, light brown, coriaceous, ostiolate at the
centre, surrounded by a small blackened clypeus and
ostiolar opening surrounded by white margin. Peridium
composed of thick-walled, brown to hyaline cells of
textura angularis. Hamathecium composed of filamentous, septate, paraphyses with hyaline, guttulate cells.
Asci 8-spored, unitunicate, cylindrical, short pedicellate,
with a cylindrical, J+, subapical ring. Ascospores 2-seriate, fusiform to broad fusiform, 1-septate, deeply constricted at the septum, hyaline, pointed at both ends,
with a longitudinally striated wall and surrounded by a
gelatinous sheath.
2. Amphibambusa bambusicola D.Q. Dai & K.D. Hyde,
sp. nov.
Index Fungorum number: IF550941, Facesoffungi
number: FoF00448, Fig. 2
Etymology: In reference to the host Bambusa and cola
meaning loving.
Holotype: MFLU 14–0825
Saprobic on decaying bamboo culms, forming a black circular spot on host surface. Sexual morph Ascomata 500–
800μm diam., 450–700μm high, solitary, scattered, immersed
under the host epidermis, globose to subglobose, light brown,
coriaceous, ostiole at the centre, surrounded by a small blackened clypeus and ostiolar opening surrounded by white margin. Peridium laterally 20–30μm thick, composed of thickwalled, brown to hyaline cells of textura angularis.
Hamathecium composed of long, septate, paraphyses, 6–
7μm wide at the base, 2.5–3μm wide at the apex, with hyaline, guttulate cells. Asci 150–200×17.5–20μm (x=180.6×
19.1μm, n=20), 8-spored, unitunicate, cylindrical, short pedicellate, with a cylindrical, J+, subapical ring, 1.5–3μm high,
2–3.5 μm diam. Ascospores 25–27 × 5.5–6 μm (x= 26.6 ×
5.7μm, n=20), 2-seriate, fusiform to broad-fusiform, 1-septate, deeply constricted at the septum, hyaline, pointed at both
ends, with a longitudinally striated wall and surrounded by
10 μm thick, gelatinous sheath. thick, gelatinous sheath.
Asexual morph Undetermined
Culture characters: Ascospores germinating on PDA within 36 h and germ tubes produced from upper cells. Colonies
growing slowly on PDA, reaching 5 mm in 2 weeks at 28 °C,
effuse, velvety to hairy, circular, irregular at the margin, white
from above, yellowish from below. Mycelium immersed in
the media, composed of branched, septate, smooth-walled,
hyaline, hyphae.
Material examined: THAILAND, Chiang Rai, Jiew
Santonkok, on dead culm of bamboo, 11 August 2011,
Dong-Qin Dai DDQ00104 (MFLU 14–0825, holotype),
(isotype in KUN, under the code HKAS 83940); ex-type living culture, MFLUCC 11–0617, IMCP. GenBank ITS:
KP744433; LSU: KP744474.
Notes: Kang et al. (1999a, b) included ten sexual
genera and their pestalotia-like asexual morphs in
Amphisphaeriaceae (sensu lato). Amphibambusa is clearly
different from other genera of Amphisphaeriaceae based on
molecular data and morphological characters. This monotypic
genus is introduced to accommodate taxa characterized by
immersed ascomata surrounded by a small blackened
clypeus and ostiolar opening surrounded by a white
margin, and cylindrical asci with fusiform ascospores
surrounded by wide gelatinous sheath. Amphibambusa
bambusicola is similar to Amphisphaeria coronata in
having immersed, black ascomata, cylindrical asci and
fusiform ascospores (Saccardo 1925). However,
Amphibambusa bambusicola is distinct in having wider
Fungal Diversity
Fungal Diversity
Phylogram generated from Maximum Likelihood analysis based
on LSU gene region of Amphisphaeraceae and related taxa in Xylariales.
Maximum likelihood bootstrap support values greater than 50 % are
indicated above or below the nodes. The ex-types (reference strains) are
in bold; the new isolates are in blue. The tree is rooted with Hypocrea
gelatinosa NBRC 104900
asci (17.5–20μm wide versus 7–8μm) (Saccardo 1913). Our
new taxon can also be compared with Amphisphaeria
bambusae which has ellipsoid, 14–17×8–9μm spores, while
the spores in Amphibambusa bambusicola are fusiform,
25–27×5.5–6μm.
Fig. 2 Amphibambusa bambusicola (holotype) a Ascomata immersed
in bamboo host b Section of ascoma c Peridium of ascoma d–f Asci g
Paraphyses j–m Ascospores surrounded by a wide gelatinous sheath n
Germinating ascospore h, i Culture on PDA o J+, subapical ring staining
by Melzer’s reagent. Scale bars: a=100 mm, b=100μm, c–g=10μm, h,
i=25 mm, j–o=5μm
R Fig. 1
Fungal Diversity
3. Amphisphaeria sorbi Senanayake & K.D. Hyde, sp.
nov.
Index Fungorum number: IF550904, Facesoffungi
number: FoF00414; Figs. 3 and 4
Etymology: Named after the host genus on which the fungus occurs.
Holotypus: MFLU 14–0797
Saprobic on branch of Sorbus aucuparia L. Sexual morph
Ascomata 350–380μm high×450–505μm diam. (x=370×
482μm, n=10), immersed to erumpent, visible as black spots
opening through the cracks of host surface, solitary, scattered,
globose to subglobose, short papillate, ostiole periphysate,
dark brown. Peridium 30–35μm (x=31μm, n=15) at the base,
65–70μm (x=68μm, n=15) at the neck, comprising 8–10
layers, inner layer of hyaline cells of textura angularis,
outer layer of brown cells of textura angularis.
Paraphyses 2–5μm wide (x=3μm, n=10), longer than
asci, filamentous, septate, embedded in gelatinous matrix. Asci 125–170×9–13μm (x=145×11μm, n=20), 8spored, unitunicate, cylindrical, short pedicellate, apically rounded, with a J-, apical apparatus. Ascospores 16–
24 × 6–8 μm (x= 19 × 6.5 μm, n = 20), uniseriate, rarely
Fig. 3 Amphisphaeria sorbi (holotype) a Ascomata on substrate b Cross section of ascoma c Peridium d–g Asci in water h. Asci in Melzer’s reagent i
Paraphyses j–l Ascospores m Sheath around spore n Germinating ascospore. Scale bars: a=1000μm, b=100μm, c=50μm, d-h=20μm, i–n=10μm
Fungal Diversity
Fig. 4 Amphisphaeria sorbi
(ex-type culture) Asexual morph
in culture a, b Conidiomata on
MEA c Peridium d–g
Conidiophore and conidiogenous
cells with attached conidia h
Conidia. Scale bars: a=500μm,
b=1000μm, c=100μm, d–h=
10μm
overlapping uniseriate, ellipsoidal, light brown, one median septate, slightly constricted at the septum, smoothwalled, surrounded by a thick mucilaginous sheath.
Asexual morph Coelomycete, Conidiomata 500–
900μm diam. (x=800μm, n=10), superficial on MEA,
solitary or aggregated, globose, dark brown. Peridium
consisting of thick walled, septate, brown mycelium.
Conidiophores 17–20μm long, 1.5–2.5μm (x=18×2μm,
n=20), arising from peridium, septate, branched, thick
walled, hyaline. Conidiogeneous cell elongated conical,
0.7–1μm wide at the apex, 2–2.5μm wide at the base
(x=1×2μm, n=20), thin-walled, septate, hyaline, annelidic.
Conidia 10–12×1–1.5μm (x=10×1μm, n=20), elongate-fusiform, hyaline, smooth-walled.
Culture characters: Colonies on MEA reaching 4 cm diam.
after 14 days at 18 °C, white, cottony, flat, low, dense, with
slightly wavy margin and few ariell mycelia.
Material examined: ITALY, Trento [TN], Dimaro,
Folgarida, on branch of Sorbus aucuparia L. (Rosaceae), 2
August 2013, E. Camporesi IT 1400 (MFLU 14–0797,
holotype); ex-type living cultures, MFLUCC 13–0721.
GenBank LSU: KP744475.
Notes: Amphisphaeria was introduced by Cesati and De
Notaris (1863) without designating a generic type (Wang
et al 2004). Petrak (1923) proposed A. umbrina as the lectotype of the genus. Different studies have listed more than 250
species in Amphisphaeria and Wang et al. (2004) accepted 12
species in the genus after examining more than 170 type
Fungal Diversity
specimens. Amphisphaeria sorbi shows more similarities
to A. vibratilis. Amphisphaeria sorbi however, differs
from A. vibratilis in having small perithecia, a peridium
with a cell arrangement of textura angularis, and wide,
non-flexuose paraphyses. The ascus apical apparatus is
discoid in Amphisphaeria sorbi and has shorter, smoothwalled ascospores without deeply pigmented septa.
Molecular analysis of the LSU gene region (Fig. 1)
confirms that Amphisphaeria sorbi clusters with
A. umbrina in Amphisphaeriaceae with 52 % bootstrap
support.
Fig. 5 Atrotorquata spartii
(holotype) a, b Appearance of
ascomata on host substrate as
minute ostiolar dots c
Section through ascoma d
Ostiole. e Peridium f
Pseudoparaphyses g Ascus in
Melzer’s reagent showing J+,
apical apparatus h–j Immature
and mature unitunicate
ascospores k–o Ascospores. Scale
bars: c=150μm, d, h–j=50μm,
e=20μm, f, k–o=10μm
4. Atrotorquata spartii Thambugala, Camporesi & K.D.
Hyde, sp. nov.
Index Fungorum number: IF550914, Facesoffungi
number: FoF00387, Fig. 5
Etymology: In reference to the host genus Spartium
Holotype: MFLU 14–0738
Saprobic on Spartium junceum. Sexual morph Ascomata
400–475μm high×280–375μm diam. (x=435×337μm, n=
5), solitary to scattered, or sometimes gregarious, immersed
beneath clypeus, dark brown to black, unilocular, globose to
subglobose with a central ostiole. Peridium 15–32μm wide,
Fungal Diversity
composed of thick-walled, dark brown to black cells of
textura angularis. Hamathecium comprising 3–4μm wide,
septate, guttulate, hyaline paraphyses, embedded in a gelatinous matrix. Asci 160–230×14–24μm (x=196×20μm, n=
10), 8-spored, unitunicate, cylindric-clavate, straight or
curved, short pedicellate, with an apical ring bluing in
Melzer’s reagent. Ascospores 19–23×8.5–9.5μm (x=21.2×
9μm, n=15), uniseriate, ellipsoid to fusiform, rounded at both
sides, hyaline when immature, becoming brown to yellowishbrown when mature, 1-septate, constricted at the septum,
guttulate, smooth-walled or striate, surrounded by a mucilaginous sheath. Asexual morph Undetermined.
Culture characters: Ascospores germinating on PDA within 24 h and germ tubes arising from both end cells. Colonies
growing slow on PDA, reaching a diam. of 20 mm after
30 days at 18 °C, circular, dense, initially white becoming pale
white, velvety, radiating towards the entire to slightly undulate
edge, non-pigmented.
Material examined: ITALY, Province of Rimini [RN],
Pennabilli, on dead branches of Spartium junceum L.
(Fabaceae), 9 October 2012, E. Camporesi, IT 799–1
(MFLU 14–0738 holotype), ex-type living cultures,
MFLUCC 13–0444. GenBank LSU: KP325443; ITALY,
Province of Forlì-Cesena [FC], Fiumicello, Premilcuore, on
dead branches of Spartium junceum (Fabaceae), 29 April
2013, E. Camporesi 799–2 (MFLU 14–0821), living cultures,
MFLUCC 13–0445.
Notes: Atrotorquata was introduced by Kohlmeyer and
Volkmann-Kohlmeyer (1993) as a monotypic genus in order
to accommodate Atrotorquata lineata. This genus is characterized by ascomata immersed under a clypeus, cylindrical
asci with a J+, apical ring and brown, two-celled, ascospores,
with 5–7 longitudinal striations at each apex, and surrounded
by a mucilaginous sheath (Kang et al. 1999a, b). Kang et al.
(1999a, b) placed Atrotorquata in Cainiaceae based on morphology. Unfortunately, the LSU sequence data for
Atrotorquata lineata are not publicly available, while the
available ITS sequence (AF009807) has not been included
in any publication. In our LSU phylogenetic tree,
Atrotorquata spartii is closely allied to Monographella nivalis
(UPSC 3273) and clustered outside of the Cainiaceae. Based
on a megablast search of NCBIs GenBank nucleotide database, the closest hit using the ITS sequence for our second
collection MFLU 14–0821/ MFLUCC 13–0445 is
Atrotorquata lineata (GenBank AF009807; Identities=405/
489(83 %). Atrotorquata lineata shares similar morphology
with A. spartii in having immersed, subglobose ascomata,
cylindrical asci with a J+, apical ring, and uniseriate, brown,
1-septate ascospores, but differs in having a hyaline clypeus, a
light brown to hyaline peridium and ellipsoidal, slightly
curved ascospores. Based on both morphology and molecular
phylogeny, A. spartii is identified as a new species of the
genus Atrotorquata. Multi-gene phylogenetic analysis of the
type species of Atrotorquata is required in order to confirm the
family placement in this genus.
5. Oxydothis atypica Pinruan, sp. nov.
Index Fungorum Number: IF551015, Facesoffungi
number: FoF00485, Fig. 6
Etymology: atypica refers to the differences from the typical ascospore characters.
Holotypus: Pinruan 135 in BBH.
Saprobic on dead leaves of Licuala longicalycata Furtado
in peat swamp. Ascomata 270–275μm, visible on the host
surface as a minute blackened dot with eccentric, periphysate
ostiole, immersed, lenticular to subglobose, brown, coriaceous, scattered. Peridium 10–12.5μm wide, comprising 3–6
layers of flattened, brown-walled cells of textura angularis,
fusing at the outside with the host tissues. Paraphyses 8.75μm
wide at the base, hypha-like, tapering to the apex, longer than
asci, not embedded in a gelatinous matrix. Asci 87.5–95×7.5–
8.75μm, 8-spored, cylindrical, unitunicate, thin-walled, pedicellate, apically rounded, with a J+, cylindrical, subapical ring,
5–5.25μm high × 2.2–2.5μm diam. Ascospores 42.5–45×5–
6.5μm, overlapping 2-seriate, hyaline, 1-septate, tapering
gradually to a point at the apex with a mucilaginous drop
and with a long drawn out spine at the base.
Holotypus: THAILAND, Narathiwat Province, Sirindhorn
Peat Swamp Forest, on dead leaves of Licuala longicalycata
(Arecaceae), 13 February 2002, U. Pinruan (Pinruan 135 in
BBH, holotype).
Notes: The characteristic features of Oxydothis atypica are
immersed ascomata with an eccentric neck, well-developed
paraphyses, cylindrical asci with a J+, subapical ring and
pointed ascospores (Hyde 1995; Hyde and Alias 2000). This
species differs from all species of Oxydothis as it has asymmetrical ascospores, with a short pointed apex and a long
spine at the base (Hyde 1995; Fröhlich et al. 2000).
Pestalotiopsis
Pestalotiopsis is an appendage-bearing conidial asexual
coelomycetous genus in the family Amphisphaeriaceae
(Barr 1975; Kang et al. 1998) that is common in tropical and
temperate ecosystems (Maharachchikumbura et al. 2011,
2012). Species of Pestalotiopsis occur commonly as plant
pathogens and are often isolated as endophytes or saprobes
(Maharachchikumbura et al. 2013). Pestalotiopsis represents a
fungal group known to produce a wide range of chemically
novel, diverse metabolites (Xu et al. 2010, 2014).
Maharachchikumbura et al. (2014) segregated two novel genera from Pestalotiopsis, namely Neopestalotiopsis and
Pseudopestalotiopsis. In this study, we introduce three
new species namely Pestalotiopsis dracontomelon,
P. italiana and P. digitalis. The phylogenetic tree is
presented in Fig. 7.
Fungal Diversity
Fig. 6 Oxydothis atypica (holotype) a Colonies on substratum b Section of ascoma c Asci with J+ ring d Asci e Paraphyses f–i Ascospores. Scale bars
a=1000μm; b=100μm; c-i=10μm
6. Pestalotiopsis digitalis Maharachch & K.D. Hyde, sp. nov.
Index Fungorum number: IF550945, Facesoffungi
number: FoF00459; Fig. 8
Etymology: named after host genus, where it was isolated.
Holotype: MFLU 14–0208
Pathogen on Digitalis purpurea. Sexual morph
Undetermined. Asexual morph Conidiomata pycnidial in
culture on PDA, globose, scattered or gregarious and confluent, semi-immersed, dark brown, up to 100 μm diam.
Conidiophores often reduced to conidiogenous cells.
Conidiogenous cells discrete ampulliform to lageniform,
smooth, thin-walled, hyaline, with 1–2 proliferations, sometimes remain vegetative. Conidia 18–22×7–9μm (x=20×
8.2μm, n=20), fusiform, straight to slightly curved, 4-septate,
basal cell conic to obconic, hyaline or slightly olivaceous,
thin- and verruculose, 2–3.5μm long (x=2.7μm), with three
median cells, doliform, concolourous, olivaceous, septa and
periclinal walls darker than the rest of the cell, together 11–
17μm long (x=15μm) second cell from base 3–4.5μm
(x=4.1μm); third cell 3–4.5μm (x=4.1μm); fourth cell 3–
4.5μm (x=4.1μm); apical cell hyaline, conic, 2–3.5μm long
(x=2.7μm); with 1–3 tubular apical appendages (mainly 2),
arising from the apex of the apical cell, 8–17μm long (x=
13μm); basal appendage 4–7μm long.
Culture characters: Colonies on PDA attaining 30–40 mm
diam. after 7 d at 25 °C, with smooth edge, pale honeycoloured, with dense aerial mycelium on the surface with
black, gregarious conidiomata; reverse similar in colour.
Material examined: NEW ZEALAND, on leaf spots of
Digitalis purpurea, 01 June 1972, J.M. Dingley 7270
(MFLU 14–0208, holotype); ex-type living cultures ICMP
5434. GenBank ITS: KP781879; TUB: KP781883.
Notes: Pestalotiopsis digitalis forms a sister clade to species including P. parva and P. rosea (Fig. 7). Pestalotiopsis
Fungal Diversity
Fig. 7 Phylogram generated from Maximum Likelihood analysis based
on combined ITS, β-tubulin and TEF gene regions of Pestalotiopsis.
Maximum likelihood bootstrap support values greater than 50 % are
indicated above or below the nodes. The ex-types (reference strains) are
in bold; the new isolates are in blue. The tree is rooted with
Neopestalotiopsis saprophyta MFLUCC 12-0282
rosea differs from P. dracontomelon in having distinctly narrow conidia. Furthermore, the reddish colony is unique to
P. rosea and this reddish colour can be seen even in
conidiogenous cells and some conidia. Furthermore, conidia
of P. digitalis are longer than those of P. parva.
7. Pestalotiopsis dracontomelon Maharachch. & K.D.
Hyde, sp. nov.
Index Fungorum number: IF550943, Facesoffungi
number: FoF00457; Fig. 9
Etymology: named after host genus, where it was isolated.
Holotype: MFLU 14–0207
Pathogen on Dracontomelon. Sexual morph
Undetermined. Asexual morph Conidiomata pycnidial in
culture on PDA, globose, aggregated or scattered, black, up
to 150 μm diam. Conidiophores 2–3-septate, sparsely
branched at the base, subcylindrical, up to 20 μm long.
Conidiogenous cells discrete or integrated, cylindrical,
percurrently proliferating 1–3 times. Conidia 18–23×5.5–
7.5μm (x=20×6.5μm, n=20), fusoid, ellipsoid, straight to
slightly curved, 4-septate; basal cell conic with a truncate base, hyaline, rugose and thin-walled, 4–5μm long; three median cells doliform, 13–17 μm long (x= 15 μm), wall
verruculose, concolourous, olivaceous, (second cell from the
base 3.5–4.5μm (x=4); third 3.5–4.5μm (x=4μm); fourth cell
3.5–4.5 μm (x= 4 μm); apical cell 3–4 μm long, hyaline,
subcylindrical, rugose and thin-walled; with 2–3 tubular apical appendages, arising from the apical crest, unbranched,
filiform, flexuous 11–20μm long (x=16μm); basal appendage
single, tubular, unbranched, centric, 2–7 μm long.
Culture characters: Colonies on PDA attaining 40–50 mm
diam. after 7 d at 25 °C, with smooth edge, whitish, with
sparse aerial mycelium on the surface with black, gregarious
conidiomata; reverse similar in colour.
Material examined: THAILAND, Chiang Rai, Nam Tak
Huey Mesak Forest Park, on disease leaves of Dracontomelon
dao Merr. & Rolfe (as D. mangiferum Blume), 10 February 2010,
S.S.N Maharachchikumbura, SAJ-0011 (MFLU 14–0207,
holotype); ex-type living culture, MFLUCC 10–0149.
GenBank ITS: KP781877; TEF: KP781880.
Notes: Pestalotiopsis dracontomelon is a pathogenic species collected from leaves of Dracontomelon mangifera from
Thailand. This species is a sister taxon to P. grevilleae and
P. knightiae (Fig. 7). It differs from P. grevilleae and
P. knightiae in having smaller conidia.
Fungal Diversity
Fig. 8 Pestalotiopsis digitalis (holotype) a Conidiomata on PDA b Conidia c Conidiogenous cells d–f Conidia. Scale bars: b–g=10μm
8. Pestalotiopsis italiana Maharachch., Camporesi & K.D.
Hyde, sp. nov.
Index Fungorum number: IF550944, Facesoffungi
number: FoF00458; Fig. 10
Etymology: named after the country, where it was collected.
Holotype: MFLU 14–0214
Saprobic on Cupressus glabra. Sexual morph
Undetermined. Asexual morph Conidiomata pycnidial
in culture on PDA, globose, scattered or gregarious
and confluent, semi-immersed, dark brown, up to
200 μm diam. Conidiophores septate near the base,
branched, subcylindrical. Conidiogenous cells discrete,
subcylindrical or ampulliform to lageniform, smooth-walled,
percurrently proliferating 1–3 times. Conidia 26–35× 8–
11 μm (x=30×9.6μm, n=20), ellipsoid, straight to slightly
curved, 4-septate, basal cell conic with obtuse end, hyaline,
thin-walled and verruculose, 5–7 μm long (x=6μm), with
three median cells, doliform to cylindrical, with thick
verruculose walls, constricted at the septa, concolourous,
olivaceous, septa and periclinal walls darker than the rest
of the cell, wall rugose, together 18–28μm long (x=23μm)
second cell from base 5.5–8.5 μm (x=6.7μm); third cell 6–9 μm
(x=7μm); fourth cell 6–9μm (x=7.7μm); apical cell hyaline,
conic to subcylindrical 4–6.5 μm long (x=4.9μm); with 2–5
tubular apical appendages (mostly 3–4), arising from the apex
of the apical cell (rarely 1 appendage arising from just above the
septum separating upper median and apical cell), 20–40μm long
(x=32μm); basal appendage present 6–10μm (x=7μm).
Culture characters: Colonies on PDA reaching 50–60 mm
diam. after 7 d at 25 °C, with an undulate edge, whitish to pale
grey-coloured, with dense aerial mycelium on surface, and
black, gregarious conidiomata; reverse similar in colour.
Material examined: ITALY, Province of Forlì-Cesena,
Camposonaldo, Santa Sofia, on dead twigs of Cupressus glabra,
24 November 2011, E. Camporesi IT051 (MFLU 14–0214,
holotype); ex-type living culture MFLUCC 12–0657.
GenBank ITS: KP781878; TEF: KP781881; TUB: KP781882.
Notes: Pestalotiopsis italiana was collected on dead twigs
of Cupressus glabra in Italy, and forms a sister clade to
P. brassicae (CBS 170.26), P. hollandica (CBS 265.33) and
P. verruculosa (MFLUCC 12–0274) which were isolated
from seeds of Brassica napus L. in New Zealand,
Sciadopitys verticillata (Thunb.) Siebold & Zucc. in the
Netherlands and Rhododendron sp. in China, respectively.
Fungal Diversity
Fig. 9 Pestalotiopsis dracontomelon (holotype) a Conidiomata on PDA b–d Conidiogenous cells e–h Conidia. Scale bars: b-h=10μm
Pestalotiopsis italiana overlaps morphologically with
P. verruculosa. However, in the phylogenetic analyses it
formed a distinct lineage apart from P. verruculosa (Fig. 7)
and geographically they are clearly distinct.
Fig. 10 Pestalotiopsis italiana (holotype) a Conidiomata on PDA b, c Conidiogenous cells d–g Conidia. Scale bars: b–g=10μm
Fungal Diversity
Chaetosphaeriaceae
The phylogenetic tree is presented in Fig. 11.
9. Conicomyces pseudotransvaalensis A. Hashim., G.
Sato & Kaz. Tanaka, sp. nov.
Index Fungorum number: IF551057, Facesoffungi
number: FoF00486; Fig. 12
Etymology: named after its morphological similarity to
Conicomyces transvaalensis R.C. Sinclair et al.
Holotype: HHUF 29956
Saprobic on Machilus japonica Siebold & Zucc. Sexual
morph Undetermined. Asexual morph Conidiomata
stromatic, synnematous, scattered, superficial, cone-shaped,
black to dark brown, setose, up to 780μm high, 260–360μm
wide at the base, cornuted with a head and a stipe; head slightly swollen, 65–100μm wide, bearing a concave conidial hymenium; stipe cylindrical or contorted, 30–100μm wide, composed of rectangular, thin-walled, brown, 1.5–2.5μm wide
cells of textura porrecta. Setae arising from stroma or stipe,
straight or curved, erect, septate, brown but pale at the apex,
thick-walled, smooth, unbranched, up to 450μm long, acute
and 2.5–4μm wide at the apex, 5–7μm wide at the base.
Conidiophores arising from inner elements of the stipe, hyaline to pale brown, unbranched or branched, up to 40μm long.
Conidiogenous cells phialidic, cylindrical, hyaline to pale
brown, smooth, 16–88×2.5–3μm. Conidia 105–170×7.5–
10μm (x=150.8×8.8μm, n=50), L/W 12–22.7 (x=17.4μm,
n=50), claviform, slightly obtuse at the apex, slightly truncate
at the base, 15–22-septate, hyaline, smooth-walled, guttulate,
bearing an unbranched appendage at the apex; appendage 40–
80μm long (x=59.8μm, n=50).
Culture characters: Conidia formed in culture are similar
to those on natural substrate.
Material examined: JAPAN, Kagoshima, Yakushima,
Yakusugi Land, dead twigs of Machilus japonica
(Lauraceae), 15 March 2007, K. Tanaka & H. Yonezawa,
GS 20 (HHUF 29956, holotype designated here); ex-type
living culture, MAFF 244767. GenBank ITS: LC001710;
LSU: LC001708.
Notes: The genus Conicomyces was established to accommodate C. transvaalensis having synnematous conidiomata
and apically appendaged conidia (Sinclair et al. 1983).
Conicomyces currently contains three described species
(Sinclair et al. 1983; Illman and White 1984; Seifert 1999),
but no molecular studies have been undertaken for the genus.
Morphologically C. pseudotransvaalensis is similar to
C. transvaalensis in having large conidia more than 100μm
long, but the latter has slightly long and slender conidia with
more septation (122–200×5.5–7.5μm, L/W 25, 19–29-septate; Nag Raj 1993). Based on a megablast search, the closest
hits to the 28S sequence of C. pseudotransvaalensis are
Chaetosphaeria fuegiana (GenBank EF063574; Identities=
729/754 (96.7 %), Gaps 5/754 (0.7 %)), Chaetosphaeria
hebetiseta (GenBank AF178549; Identities = 723/754
(95.9 %), Gaps 5/754 (0.7 %)) and Chaetosphaeria dilabens
(GenBank AF178557; Identities=720/751 (95.9 %), Gaps=5/
751 (0.7 %)). These results clearly indicate that the genus is a
member of Chaetosphaeriaceae (Sordariomycetes), as previously suggested by Hashimoto et al. (2015) based on
morphological grounds. In Fig. 11, C. pseudotransvaalensis
clusters in Chaetosphaeriaceae and is related to species of
Chaetosphaeria sensu lato.
10. Dinemasporium nelloi W.J. Li, Camporesi & K.D.
Hyde, sp. nov.
Index Fungorum number: IF550919, Facesoffungi
number: FoF00424, Fig. 13
Etymology: Named after Nello Camporesi, who collected
the sample from which the species was isolated.
Holotypus: MFLU 14–0811
Saprobic on dead stem of Dactylis glomerata L., forming
conspicuous, rounded to irregular, black, conidiomata. Sexual
morph Undetermined. Asexual morph coelomycetous.
Conidiomata 100–200μm high, 250–350μm diam., pycnidial, superficial, erumpent, cupulate when dry, solitary, scattered
or gregarious, black, with stroma cells of textura angularis at
the base and a lateral excipulum of textura porrecta or textura
intricata of pale to dark brown cells. Setae divergent, stiff,
brown, smooth-walled, septate, tapering towards an acute
apex. Conidiophores cylindrical, septate, smooth, thickwalled, hyaline, arising from basal and periclinal wall cells.
Conidiogenous cells 10–15 μm long × 1.5–3.5 μm wide,
phialidic, cylindrical, hyaline, smooth-walled, tapering toward
the apex. Conidia 10–20 × 2.5–3.5 μm (x= 15 × 3; n = 20),
blastic-phialidic, amerosporous, allantoid or lenticular,
guttulate, hyaline, smooth-walled, with a single unbranched
setula at each end; setula 5–15μm long×0.5–1.5μm wide.
Culture characters: Colonies on PDA slow growing, white
in the first few days, becoming yellowish, and reaching 20–
25 mm diam. after one week, with the middle area becoming
pink, whitened at the edge, becoming felt-like after two
weeks, dense, aerial, filamentous; reverse brown, pigments
produced.
Material examined: ITALY, Province of Forlì-Cesena [FC],
Castrocaro Terme, Converselle, on dead stem of Dactylis
glomerata L. (Poaceae), 1 December 2012, E. Camporesi
IT-934 (MFLU 14–0811, holotype), ex-type living culture,
MFLUCC 13–0482. GenBank ITS: KP711358; LSU:
KP711363; SSU: KP711368; ibid. (KUN! HKAS 83970,
isotype).
Notes: Dinemasporium was introduced by Léveillé (1846)
with D. graminum (Lib.) Lév. as the type species. This is a
comparatively large heterogeneous genus in which 85 taxa
have been recorded according to the Index Fungorum
(2015). The species of Dinemasporium are characterized by
Fungal Diversity
Fig. 11 Phylogram generated from Maximum likelihood (RAxML)
analysis based on LSU sequence data. Maximum Likelihood bootstrap
support values greater than 50 % are indicated above or below the nodes.
The ex-types (reference strains) are in bold. New isolates are in blue. The
tree is rooted with Camarops tubulina SMH 4614 and Camarops
ustulinoides SMH 1988
Fungal Diversity
Fig. 12 Conicomyces pseudotransvaalensis (holotype). a, b
Conidiomata on host surface c-e Conidiomata in culture f Conidioma in
longitudinal section g, h Conidiogenous cells i–n Conidia o Germinating
conidium a, b, f, h, k, l from HHUF 29956 (holotype); c-e, g, i, j, m–o
from MAFF 244767 (ex-type isolate). Scale bars: a, c=500μm, b, e=
100μm, d=250μm, f, i–o=20μm, g, h=5μm
superficial, cupulate conidiomata with setae; “phialidic”
conidiogenous cells; and hyaline, oblong to allantoid conidia
with single appendage at each end (Sutton 1980; Nag Raj
1993). Dinemasporium nelloi is morphologically similar with
Fungal Diversity
Fig. 13 Dinemasporium nelloi (holotype) a Specimen b, c Black
conidiomata on the host surface d Vertical section of conidioma e
Section of peridium f Seta g–i Conidiogenous cells and developing
conidia j, l–m Conidia k Germinated conidia n Culture on PDA. Scale
bars: b=500μm, c=100μm, d=50μm, e=10μm, f=20μm, g–m=5μm,
n=25 mm
D. americana, D. morbidum and D. polygonum. However,
D. nelloi produces conidia that are much longer than in these
species (i.e. (9–)12–13(–16) μm long in D. americana, 8–
13μm long in D. morbidum and (9–)10–12(–13) μm long in
Fungal Diversity
D. polygonum), although shorter than D. lanatum (13–27μm
long) (Nag Raj 1993; Crous et al. 2012). According to the
LSU sequence data, D. nelloi is distinct from any other species
within Dinemasporium. Combined with morphological data
and molecular data, we introduce the new species,
Dinemasporium nelloi.
Diaporthaceae
The family Diaporthaceae (order Diaporthales, class
Sordariomycetes) includes Diaporthe, as the species-richest
genus (Wehmeyer 1933; Ueker et al. 1988; Hyde et al.
2014; Udayanga et al. 2014). About 2000 names are listed
in Diaporthe and the asexual morph Phomopsis in the Index
Fungorum (2015). In addition, the genera Mazzantia,
Ophiodiaporthe and Pustulomyces are known within the family (Castlebury et al. 2002; Fu et al. 2013; Dai et al. 2014a, b).
The genus Mazzantia is poorly known with little sequence
data and few cultures available (Castlebury et al. 2002;
Rossman et al. 2007). Ophiodiaporthe, has been recently described from China as a monotypic genus comprising the homothallic species O. cyatheae Y.M. Ju et al. (Fu et al. 2013).
However, the phylogenetic distinctiveness of this genus is
uncertain as the analyses revealed a high similarity with
Diaporthe. A new coelomycetous genus, Pustulomyces, was
described from decaying bamboo in Thailand which is morphologically similar to Bambusicola (Bambusicolaceae) (Dai
et al. 2014a, b). Lamprecht et al. (2011) recognized
coelomycetous fungi with pigmented conidia, Stenocarpella
macrospora (Earle) B. Sutton, S. maydis (Berk.) B. Sutton,
and Phaeocytostroma ambiguum (Mont.) Petr, causing root
and crown rot of maize, to be placed within Diaporthaceae.
Herein, we introduce a new species, Diaporthe thunbergiicola
as a new species in the genus Diaporthe, contributing to the
diversity within the family. The phylogenetic tree is presented
in Fig. 14.
11. Diaporthe thunbergiicola Udayanga & K.D. Hyde, sp.
nov.
Index Fungorum Number: IF551072, Facesoffungi
number: FoF00472; Fig. 15
Etymology: Referring to the original host association with
tropical ornamental vine Thunbergia laurifolia Lindl.
Holotype: MFLU 14-0816
Pathogen on leaves of Thunbergia laurifolia. Sexual
morph Undetermined. Asexual morph Pycnidia on alfalfa
twigs on WA: subglobose to ovate, 100–200μm diam., embedded in tissue, erumpent at maturity, with an elongated,
black neck 100–200μm long, often with a yellowish, conidial
cirrus extruding through ostiole; walls parenchymatous,
consisting of 3–4 layers of medium brown textura angularis.
Conidiophores hyaline, smooth, unbranched or branched at
the basal cell, ampulliform, cylindrical to sub-cylindrical, with
larger basal cell 6–14×1–2μm (x±SD=11±2×1.5±0.2, n=
30). Conidiogenous cells phialidic, cylindrical, terminal,
slightly tapering towards apex, 0.5–1μm diam. Paraphyses
absent. Alpha conidia 5.7–7.5×2–3μm, abundant in culture
and on alfalfa twigs, aseptate, hyaline, smooth-walled, ovate
to ellipsoidal, biguttulate, base subtruncate (x±SD=6.6±0.4×
2.8±0.1 , n=30). Beta conidia undetermined.
Cultural characteristics: In dark at 25 °C for 1 wk, colonies
on PDA fast growing, 5±0.2 mm/day (n=8), white, raised,
aerial mycelium, reverse yellowish pigmentation developing
in centre; stroma not produced in 1 wk old culture.
Holotype: THAILAND, Chiang Mai Province, Doi Suthep
Pui herbal garden, on leaves of Thunbergia laurifolia
(Acanthaceae), 10 January 2012, D. Udayanga DPH 114
(MFLU 14–0816, holotype), ex-type culture, MFLUCC 12–
0033, ICMP. GenBank ITS: KP715097; TEF: KP715098.
Notes: This is the second species described from
Thunbergia laurifolia in Thailand. Udayanga et al. (2012)
described Diaporthe thunbergii Udayanga, X.Z. Liu &
K.D. Hyde from the same host. Both species were recovered from similar disease symptoms on leaves; however D. thunbergiicola has both abundant and distinct alpha
and beta conidia. The alpha conidial dimensions overlap with
other taxa in the genus, making morphological identification
impossible. The fungus was associated with typical leaf
spots and in latent phase of yellowing and leaf necrosis.
In the phylogenetic analysis, the species was placed
within D. sojae species complex (Udayanga et al.
2014), mostly consisting of crop and weed associated
species. The comparison of ITS and EF1-α sequences revealed that D. thunbergiicola is distinct from the sister species
D. ueckerae, originally reported from Cucumis melo L. in
Oklahoma, USA. The pair-wise comparison revealed high
similarities of the ITS (96 %) and EF1-α sequences (92 %)
between the two species. The combined phylogenetic tree
(Fig. 14) revealed the placement of D. thunbergii and
D. thunbergiicola compared to a selection of closely related
ex-type isolates.
Diatrypaceae
Species of Diatrypaceae (Xylariales) are widespread inhabitants of dead wood and bark of a broad variety of plants
worldwide (Trouillas et al. 2011). They can be recognized
by their perithecial ascomata embedded usually in a black
stroma, long stalked asci and allantoid ascospores (Glawe
and Rogers 1984; Rappaz 1987). Kirk et al (2008) listed
13 genera and more than 220 morphological species in
this family. The available phylogenetic studies are those
of Acero et al. (2004) and Trouillas et al. (2010, 2011).
In this study, we introduce a new species based on both
the morphology and phylogeny. The phylogenetic tree is
presented in Fig. 16.
Fungal Diversity
Fig. 14 Phylogram generated
from RAxML based on combined
alignment of ITS, TEF, β-tubulin
and CAL gene regions.
Maximum likelihood bootstrap
support values greater than 90 %
are indicated above or below the
nodes. The ex-types are in bold;
the newly isolate is in blue. The
tree is rooted with Diaporthella
corylina CBS 121124
12. Diatrype palmicola J.K. Liu & K.D. Hyde, sp. nov.
Index Fungorum number: IF551017; Facesoffungi
number: FoF00487, Fig. 17
Etymology: Named after the host on which the fungus was
collected.
Holotype: MFLU 15-0040
Saprobic on palm fronds. Stromata well-developed, or
comprising mostly fungal tissue, often delimited with a black
line perceptible in the wood below, bursting through bark or
wood and often surrounded by remaining adherent epidermis
or wood fragments, discoid to pulvinate or hemisphaerical,
generally strongly emerged from the host surface.
Entostroma poorly developed between ascomata, only present
at the level of the neck. Sexual morph Ascomata immersed in
the stromata, with the stromata surface flat or slightly convex.
Ostiole flattened and circular or subconical. Asci 70–110×7–
9μm, 8–spored, unitunicate, long pedicellate, clavate, apically
rounded to truncate. Ascospores (6−)7–8(−9)×1.5–2μm, 2–3
Fungal Diversity
Fig. 15 Diaporthe
thunbergiicola (holotype) a, b
Leaf sports on Thunbergia
laurifolia c Pycnidia on alfalfa
stem in WA d conidiophores e.
alpha conidia. Scale bars:
c=1000μm, d, e=8μm
seriate, allantoid, 1–celled, hyaline to subhyaline, rarely
pale olivaceous, slightly to moderately curved and
smooth-walled.
Material examined: THAILAND, Chiang Rai Province,
Muang District, Khun Korn Waterfall, on dead branch of
Caryota urens L. (Arecaceae), 14 March 2010. J.K. Lui,
JKA0031 (MFLU 15-0040, holotype) ex-type living culture=MFLUCC 11–0018, GenBank ITS: KP744438; LSU:
KP744481; SSU: KP753949; ibid. on dead branch of
Caryota urens, 6 September 2010, J.K. Liu, JKA0032
(MFLU 15-0041, paratype); living culture, MFLUCC 11–
0020, GenBank ITS: KP744439; LSU: KP744482; SSU:
KP753950.
Notes: Diatrype palmicola has morphological characteristics typical of taxa in the genus Diatrype. Four species, namely
D. euterpes, D. palmarum, D. palmarum var. rimosa and
D. urticaria are described from palms (Liu et al. 2014).
Analyses showed that D. palmicola ITS sequences differ from
all sequences of taxa of Diatrype in GenBank, including the
close species D. favacea and D. pulvinata. The reconstruction
generated from ITS sequences showed similar results as previous studies (Acero et al. 2004; Trouillas et al. 2011). This
suggests that the current taxonomic scheme for the
Diatrypaceae may not reflect the true phylogenetic relationship of these fungi. The number of spores per ascus (eight
spores versus more than eight spores) has been used traditionally to define genera of Diatrypaceae (Diatrype vs.
Diatrypella and Eutype vs. Eutypella). However the
polysporous ascus feature has been shown to be not significant in Diatrypaceae based on recent studies (Acero et al.
2004; Vasilyeva and Stephenson 2005; Trouillas et al. 2011;
Chacon et al. 2013) and the phylogenies showed that the
genera Cryptovalsa and Eutypella, as well as Diatrype and
Diatrypella have molecular affinities. Until more species are
collected, epitypified and sequenced, as well as an increased
sampling of taxa included, the understanding of Diatrypaceae
will remain fragmentary.
13. Phaeoisaria pseudoclematidis D.Q. Dai & K.D. Hyde,
sp. nov.
Index Fungorum number: IF550942, Facesoffungi
number: FoF00452, Figs. 18 and 19
Etymology: In reference to its similarity with Phaeoisaria
clematidis.
Holotype: MFLU 14–0824
Saprobic on bamboo culms, formed mostly at the nodal
region on host surface. Mycelium immersed on the substrate,
composed of septate, branched, brown hyphae. Sexual
morph Undetermined. Asexual morph Conidiophores 50–
500×2–3μm, macronematous, synnematous, brown to dark
brown, septate, branched, smooth, single conidiophores.
Synnemata erect, rigid, dark brown, velvety, smooth, composed of compactly and parallels adpressed conidiophores,
200–500μm long, 40–80μm wide at the base, 40–60μm wide
in the middle, 20–30 μm wide at the apex, with flared
conidiogenous cells in the above half. Conidiogenous cells
5–20×2–3μm (x=13.1×2.6μm, n=50), terminal, integrated
or discrete, short, recurved, ellipsoidal, brown to dark brown,
smooth, denticulate, polyblastic, sympodial, each with one to
several denticulate conidiogenous loci. Conidia 5–8.5×3–
4 μm (x= 6.7 × 3.3 μm, n = 50), cylindric-ovate, straight,
aseptate, hyaline, smooth-walled, guttulate.
Culture characters: Conidia germinating on PDA within
24 h and germ tubes produced from lower end. Colonies
Fungal Diversity
Fig. 16 Phylogram generated from parsimony analysis based on ITS
sequence data of Diatrypaceae. Parsimony bootstrap support values
greater than 50 % are indicated above the nodes, and branches with
Bayesian posterior probabilities greater than 0.95 are given in bold. The
ex-types (reference strains) are in bold; the new isolates are in blue. The
tree is rooted with Eutypella cryptovalsoidea CBS 128335
Fungal Diversity
Fig. 17 Diatrype palmicola
(holotype). a, b Appearance of
the stromata on host surface c–f
Asci with long pedicels g-i
Allantoid ascospores. Scale Bars:
a, b=1 mm. c–f=30μm, g–i=
10μm
growing slowly on PDA, reaching 3 mm in 2 week at 28 °C,
flat, circular, dark brown from forward and the reverse.
Mycelium immersed in media, composed of branched, septate, smooth, dark brown hyphae.
Material examined: THAILAND, Chiang Rai, Mae Sae
Village, on dead culm of bamboo (Bambusae), 14 May
2011, Dong-Qin Dai DDQ 0020 (MFLU 14–0824, holotype),
(isotype deposit in KUN, under the code of HKAS 83939),
living culture, MFLUCC 11–0393=ICMP. GenBank ITS:
KP744457; LSU: KP744501; SSU: KP753962.
Notes: Phaeoisaria was introduced by Höhnel (1909) for a
collection on Gigantochloa sp. (Bambusae) and is typified by
P. bambusae Höhn. This genus is characterized by black
synnemata with polyblastic, sympodial, denticulate,
conidiogenous cells which produce small, hyaline conidia.
Phaeoisaria pseudoclematidis is different from the type species, P. bambusae, in having shorter synnemata (200–500μm
versus 1–1.5mm), and wider conidia (3–4μm versus 1.5–
2 μm) (Sitzungsberichte et al. 1909, Seifert et al. 2011).
Phaeoisaria bambusae was originally collected from
Indonesia. Our new isolate is similar to P. clematidis in morphology. Synnemata of P. clematidis are however, more than
800μm long and conidia narrower than 3μm (Hughes 1958).
Phaeoisaria was placed in Pezizomycotina order incertae
sedis, Ascomycota (Höhnel 1909). Wijayawardene et al.
(2012) linked Phaeoisaria to Annulatascaceae
(Rhamphoria), Boliniaceae and Diatrypaceae (Eutypella).
There are a few new taxa published or recorded in
Phaeoisaria (Matsushima 1996; Castañeda Ruíz et al. 2002;
Mel’nik 2012). In this paper, based on our collection, we
Fungal Diversity
Fig. 18 Phylogram generated from Maximum Parsimony analysis based
on LSU sequence data. Parsimony bootstrap support values greater than
50 % are indicated above or below the nodes. The ex-type (reference
strains) are in bold, the newly new isolates are in blue. The tree is
rooted with Microascus longirostris CBS 267.49
provide the phylogenetic tree to clarify the natural
placement of Phaeoisaria (Fig. 18) and introduce a
new species.
RPB2 genes further elucidated the phylogenetic position of
Glomerellaceae. The phlogenetic tree is presented in Fig. 20.
Glomerellaceae
The family name Glomerellaceae was introduced by Locquin
(1984) and was validated by Seifert and W. Gams in Zhang
et al (2006). Glomerellaceae is a monotypic family characterized by the Glomerella sexual morph and the Colletotrichum
asexual morph. The first attempts to place Colletotrichum
within a molecular phylogenetic system using 18S rDNA
sequences were by Illingworth et al. (1991) and Berbee and
Taylor (1992). Réblová et al. (2011) using ITS, LSU, SSU and
14. Colletotrichum sedi Jayawardena, Bulgakov & K.D.
Hyde, sp. nov.
Index Fungorum number: IF550762, Facesoffungi
number: FoF00332; Fig. 21
Etymology: Based on the host genus Sedum.
Holotypus: MFLU 14–0623.
Saprobic on dead stalks of Sedum sp. Asexual morph
Conidiomata 262–410μm (x=329μm, n=10) diam., solitary,
acervulus, black, oval. Setae abundant, 50–125μm long,
pale to medium brown, smooth-walled, 1–4-septate, base
cylindrical, 2.8–6.8 μm diam., apex somewhat acute.
Fungal Diversity
Fig. 19 Phaeoisaria
pseudoclematidis (holotype) a
Fruiting bodies on bamboo host b
Apex of synnemata c Synnemata
d, f Conidiogenous cells e Base of
conidiophores g, h Hyaline
conidia i Germinating conidia j, k
Cultures on PDA. Scale bars: a=
1 mm, b=30μm, c, e=50μm, d,
f=10μm, g–i=5μm. j=1 mm, k=
3 mm
Conidiophores simple, to 37 μm long, hyaline, smooth–
walled. Conidiogenous cells (7–)10.5–19(–22)×(1–)2.39–
5.2(–6) μm (x=14×3.4μm, n=20), hyaline, smooth–walled,
cyllindrical to slighty inflated, opening 1.2–3.7μm diam.,
Collarette 0.5–1 μm long, periclinal thickening visible,
Conidia (2–)6.6–10.5(–14)×(1–)2–5(–6)μm (x=7.3×3.6μm,
n = 40) hyaline, smooth-walled or verruculose, aseptate,
curved, both sides gradually tapering towards the round to
slightly acute apex and truncate base, guttulate.
On PDA Conidiomata solitary or aggregated, acervulus,
submerged, black, circular to oval. Sporulation abundant.
Setae 50–120μm long, dark brown up to the tip, opaque, septa
difficult to distinguish, 2 to 4-septate, smooth-walled,
base cylindrical, straight or±bent, 3–7μm diam., apex
acute. Conidiophores to 30 μm long, simple, hyaline.
Conidiogenous cells (5–)8.5–16(–19)×(1–)2.5–5(–6)μm
(x=12×3.5μm, n=20), hyaline to pale brown, smooth–
walled, cyllindrical to slighty inflated, opening 1.2–
3.7 μm diam., Collarette 0.5–1 μm long, periclinal
thickening visible. Conidia (4–)6–10.3(–14) × (1–)2.3–
5.2(–6) μm (x= 6.9 × 3.6 μm, n = 40) hyaline, curved,
both sides gradually tapering towards the round to
slightly acute apex and truncate base, guttulate. Appressoria
(6–)8.5–16.4(–18)×(1–)2.5–4(–5) μm (x=12.5×2.8μm, n=
Fungal Diversity
Fig. 20 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-types
(reference strains) are in bold; the
new isolates are in blue. The tree
is rooted with Monilochaetes
infuscans CBS 869.96
10) solitary to aggregated, in small groups or short chains,
medium to dark brown, smooth-walled, round, oval or irregular. Sexual morph Undetermined.
Culture Characters: Colonies on PDA reaching 45 mm in
7 days at 28 °C, flat with entire margin, olivaceous grey aerial
mycelium becoming dull green towards the edge with olivaceous grey to iron grey acervuli from forward, reverse greyolivaceous to dull green, concentric.
Material examined: RUSSIA, Rostov region, Rostov-naDonu city, Botanical garden of Southern Federal University,
flowerbed, on Sedum sp. (Crassulaceae), 5 March 2014,
Timur Bulgakov (T94), (MFLU 14–0623, holotype); extype living cultures, MFLUCC 14–1002, CGMCC 3.17570.
GenBank ITS: KM974758; GADPH: KM974755; ACT:
KM974756; CHS: KM974754; β-tubulin: KM974757.
Notes: The genus Colletotrichum was introduced by Corda
(1831) and subsequently treated by Hyde et al. (2009) and Cai
et al. (2009). The most recent treatment is by Hyde et al.
(2014) which was based on multi-gene phylogeny. The
dematium complex contains the type species of the genus,
Colletotrichum lineola (Damm et al. 2009; Cannon et al.
2012). Colletotrichum dematium is mainly characertized by
its curved conidia whch is also a character of the species of the
truncatum complex but found to occupy distinct separate
clades (Damm et al. 2009; Cannon et al. 2012).
Colletotrichum sedi clusters in the subclade comprising
C. circinans and C. spinaciae within the dematium complex
(Fig. 20). Colletotrichum sedi forms a separate branch with
100 % bootstrap support and 1.00 Bayesian posterior probability. This species differs from C. circinans and C. spinaceae
in having longer setae (50–125 μm) with 1–4-septate, simple
conidiophores, solitary to aggregated appressoria, in small
groups or short chains and smaller conidia (6.6–10.5×2–5 μm).
Halosphaeriaceae
The family Halosphaeriaceae was detailed in Jones et al.
(2009) and treated is here accordingly. The phylogenetic tree
is presented in Fig. 22.
15. Natantispora unipolaris K.L. Pang, S.Y. Guo &
E.B.G. Jones, sp. nov.
Index Fungorum number: IF551074; Facesoffungi
number: FoF00564; Fig. 23
Etymology: In reference to the unipolar ascospore
appendage.
Holotype: F27870 (National Museum of Natural Science,
Taiwan)
Saprobic on dead stems. Sexual morph Ascomata
78–(124)–158 × 75–(119)–216 μm (n = 9), light to dark
coloured when mature, globose to subglobose, solitary or gregarious, immersed, coriaceous, ostiolate. Necks 30–(59)–
96μm long (n=8), periphysate. Peridium 5–33μm (n=18),
composed of one layer of elongated cells with large lumina,
cells of textura angularis. Catenophyses present, persistent.
Asci 59–(87)–108 × 15–(18)–20 μm (n = 8), 8-spored,
unitunicate, clavate, thin-walled, persistent, short pedicellate,
developing from inner wall of ascoma base, no apical apparatus or retraction of plasmalemma. Ascospores 21–(25)–28×
6–(8)–8μm (n=50), elongate-ellipsoidal, hyaline, 1-septate,
not constricted at the septum. Appendages unipolar, initially
adpressed to the ascospore wall and extended over the mid-
Fungal Diversity
Fig. 21 Colletotrichum sedi (holotype) a Specimen with conidiomata b
Black acervuli c Brown setae d Conidiophores with basal parts of setae e
Hyaline conidiogenous cells f Conidiomata on PDA g Hyaline conidia h
Germinating conidium i Appressoria j Reverse view of the colony k
Upper view of the colony. Scale bars: b=200μm, c=20μm, d=15μm,
e, f=10μm, g-i=5μm
Fungal Diversity
Fig. 22 Phylogram generated from Maximum likelihood (RAxML)
analysis based on combined SSU and LSU sequence data of the
Halosphaeriaceae. Maximum Likelihood bootstrap support values
greater than 50 % are indicated above or below the nodes, the branches
with Bayesian posterior probabilities greater than 0.95 are given in bold.
The ex-types (reference strains) are in bold, the new isolates are in blue.
The tree is rooted with Microascus trigonosporus AFTOL-ID 914 and
Petriella setifera AFTOL-ID 956
septum, unravelling immediately in sea water to form a long
thin filament. Asexual morph Undetermined.
Material examined: TAIWAN, Nankunshen, on a dead
stem of Phragmites australis (Cav.) Trin. ex Steud.
(Poaceae), 22 November 2010, Ka-Lai Pang, F27870
(holotype, National Museum of Natural Science, Taiwan);
ex-type living culture, BCRC FU 30316. GenBank LSU:
KM624522; SSU: KM624521.
Notes: Natantispora unipolaris forms in a monophyletic
clade with N. retorquens in the Halosphaeriaceae
(Microascales, Ascomycota) with high bootstrap support
(Fig. 22). Natantispora unipolaris is morphologically very
Fungal Diversity
Fig. 23 Natantispora unipolaris (holotype). a Immersed ascoma with neck protruding to the surface b One-layered peridium of elongated cells with
large lumina c Asci at various stages of development d–f Ascospores with an unfurling appendage at one end. Scale bars: a=30 μm, b–f=10 μm
similar to the type species N. retorquens, including globose to
subglobose ascomata, presence of catenophyses, hyaline,
elongate-ellipsoidal ascospores with one septum, a polar appendage pad extending over the mid-septum which unfurls to
form a long thread (Shearer and Crane 1980). Natantispora
unipolaris differs in having only one ascospore appendage
instead of two in N. retorquens; ascomata are smaller with
many small oil globules and one large one in each cell.
Tirispora unicaudata also has a single polar hamate appendage visible when mounted in sea water, but phylogenetically it
is distant from N. unipolaris.
16. Saagaromyces mangrovei Abdel-Wahab, Bahkali &
E.B.G. Jones, sp. nov.
Index Fungorum number: IF551073; Facesoffunginumber:
FoF 00488; Figs. 24 and 25
Etymology: In reference to the habitat where the fungus
was first found.
Holotype: CBS H-22126
Saprobic on submerged wood in mangroves. Ascomata
370–520μm high, 200–380μm wide, sub-globose, ovate,
obpyriform, ellipsoidal, horizontal, yellow-brown to brown,
immersed, membranous, ostiolate and papillate. Peridium
45–65μm thick, two-layered, forming textura angularis, outer
stratum 30–42μm thick, 8–15 layers of polygonal yellow
brown to brown melanized cells; inner stratum 15–25μm
thick, 4–7 layers of elongated polygonal to flattened, hyaline
cells, merging with the pseudoparenchyma of the venter.
Necks lateral bending upward, 260–550μm long, 90–130μm
wide, cylindrical, hyaline to yellow brown, ostiolar canal
periphysate, periphyses up to 20μm long. Catenophyses
consisting of chains of cells that are subglobose, ovate to
elongated, up to 28μm wide. Asci 120–202 (x=159μm, n=
20) μm long, 25–38 (x=32μm, n=20) μm wide, 8-spored,
clavate, with narrow long stalks, 40–65μm long, 4–9μm wide,
thin-walled, persistent, without an apical apparatus, developing
at the base of the ascoma venter on small-celled ascogenous
tissue. Ascospores 24–32 (x=29μm, n=50) μm long (excluding appendages), 10–16 (x=13μm, n=50) μm wide (excluding
appendages), broadly ellipsoidal, 1-septate, not constricted at
the septum, hyaline, thin-walled, with bipolar apical appendages, at first cap-like and stiff, 13–16μm long, 2.5–3μm wide,
rapidly unfurling in water into long, thin filaments.
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 H-22126 holotype); ex-type living culture, MF 1206. GenBank LSU: JX911896; SSU: JX911897.
Notes: Several genera with polar unfurling ascospore appendages were described from marine habitats including:
Aniptodera, Ascosacculus, Gesasha, Halosarpheia,
Magnisphaera, Natantispora, Oceanitis and Saagaromyces.
Among these, Saagaromyces is characterized by hyaline to
light brown ascomata; thin-walled, 1-septate, broadly ellipsoidal ascospores and thin-walled, persistent asci. However, the
single character that distinguishes the genus is the large
Fungal Diversity
Fig. 24 Saagaromyces
mangrovei (holotype). a Vertical
section of ascomata b Magnified
part of the vertical section of the
ascomata showing the peridium
structure and the periphysate neck
c Ascomatal squash showing
immature asci and catenophyses
(arrowed) d Ascomatal squash
showing mature asci. c, d Stained
with toluidine blue. Scale bars:
a=50μm, b-d=25μm
hamate polar appendages that are equal to or longer than half
of the ascospore length and width (Kohlmeyer 1984; Pang
et al. 2003). Saagaromyces mangrovei has smaller ascospores
and asci than the other described species in the genus.
Hypocreales
Hypocreales is a diverse order of Sordariomycetes, including
more than 2000 species, in about 115 genera which are related
to seven families (Lumbsch and Huhndorf 2010). Species of
Hypocreales are pathogens or saprobes, and are usually characterized by their brightly coloured (often yellow, orange or
red), perithecial fruiting bodies, or spore-producing structures.
The phylogenetic tree is presented in Fig. 26.
17. Myrothecium macrosporum D.Q. Dai & K.D. Hyde,
sp. nov.
Index Fungorum number: IF550939, Facesoffungi
number: FoF00451, Fig. 27
Etymology: In reference to the big conidia.
Holotype: MFLU 14–0823
Saprobic on decaying bamboo culms, forming dark green,
glistening, slimy drops on the centre of white mycelial
sporodochia. Mycelium partly immersed on the substrate, partly superficial, composed of septate, branched, hyaline, smooth
hyphae, 400–600 μm diam. Conidiophores sporodochial,
macronematous, cylindrical, brown to dark brown, septate,
branched, straight, smooth, single conidiophores 200–500×
4–5μm. Conidiogenous cells phialidic, integrated or discrete,
cylindrical, pale brown, smooth, straight. Conidia 25–35×2–
3μm (x=29.6×2.7μm, n=20), cylindrical, narrow at the apex,
truncate at the base, straight to slightly curved, flexuous, 0–3septate, hyaline, smooth-walled, with guttulate cells.
Culture characters: Conidia germinating on PDA within
24 h and germ tubes developing from both ends. Colonies
growing slowly on PDA, reaching 4 mm in 2 week at 28 °C,
circular, with irregular edge, hyaline from forward and reverse
view. Mycelium superficial to immersed in/on media, with
branched, septate, smooth hyphae.
Material examined: THAILAND, Chiang Rai, Mae Sae
Village, on dead culm of bamboo (Bambusae), 14 May
2011, Dong-Qin Dai, DDQ 00019 (MFLU 14–0823,
Fungal Diversity
Fig. 25 Saagaromyces mangrovei (holotype). a Mature ascus b–d Ascospores with different degrees of uncoiling for polar appendages. c, d Stained
with toluidine blue. Scale bars: a=10μm, b–d=5μm
holotype), isotype in KUN, under the code of HKAS 83938),
ex-type living culture at MFLUCC 11–0392. GenBank ITS:
KP744448; LSU: KP744491.
Notes: Tulloch (1972) monographed the genus
Myrothecium, redescribed eight species and added two new
species and three new combinations. The genus is typified by
M. inundatum Tode ex S.F. Gray (Preston 1948). Presently,
there are 79 records of Myrothecium in Index Fungorum
(2015), most of them are saprobes and some are pathogens
(Yamazaki et al. 2014). The species concept within the genus
Myrothecium is rather broad and sometimes controversial,
with species having a saprobic, pathogenic or hyperparasitic
habitat, sessile or stalked sporodochia to synnematous
conidiomata and smooth-walled or striated conidia
(Ellis 1971, 1976). The genus awaits revision. Our
new species differs from other species of Myrothecium
by the larger size of its conidia (more than 20μm long).
Myrothecium macrosporum is somewhat closely related
to M. prestonii and M. setiramosum in the phylogenetic
analysis (Fig. 26). However, M. prestonii was isolated
from soil and M. setiramosum from leaves of Eugenia
glabrata.
Magnaporthaceae
Magnaporthaceae was introduced by Cannon (1994), and is
typified with Magnaporthe. This family includes parasites,
saprobes and endophytes (Huhndorf et al. 2008). More than
100 species in 12 sexual and seven asexual genera currently
are placed in Magnaporthaceae (Luo and Zhang 2013).
Magnaporthaceae is characterized by solitary, immersed, globose to subglobose, perithecial ascomata, with a long, brown to
black, smooth neck, which break through the host surface (Luo
and Zhang 2013). The phylogenetic tree is presented in Fig. 28.
18. Neogaeumannomyces D.Q. Dai & K.D. Hyde, gen.
nov.
Index Fungorum number: IF550936, Facesoffungi
number: FoF00449
Etymology: In reference to a new genus similar with
Gaeumannomyces and neo meaning new.
Type species: Neogaeumannomyces bambusicola D.Q. Dai
& K.D. Hyde
Saprobic on decaying bamboo culms, forming dark circular erumpent spots on host surface with ascomatal ostiolar
Fungal Diversity
Fig. 26 Phylogram generated
from Bayesian analysis based on
ITS sequence data. Bayesian
posterior probabilities (PP)
greater than 0.90 are indicated
above or below the nodes. The extypes (reference strains) are in
bold, the new isolates are in blue.
The tree is rooted with
Melanospora pascuensis IMI
378527
neck on raised areas. Sexual morph Ascomata perithecial,
gregarious, immersed, solitary, globose to subglobose, black,
ostiolate. Ostiolar neck long, lined with periphyses. Peridium
comprising host and fungal tissues, with outer layer composed
of brown and thick-walled, cells; inner layer composed of
hyaline, thin-walled, large cells of textura angularis.
Hamathecium dense, with long, septate, paraphyses
intermixed with asci. Asci 8-spored, unitunicate, cylindrical,
with a short furcate pedicel, with an apical ring. Ascospores
filiform, curved to sigmoid, filiform to long fusiform, elongate, narrow and curved at the ends, 2–3-septate, distoseptate
when mature, guttulate, hyaline, smooth-walled.
19. Neogaeumannomyces bambusicola D.Q. Dai & K.D.
Hyde, sp. nov.
Index Fungorum number: IF550937, Facesoffungi
number: FoF00450, Fig. 29
Etymology: In reference to the host Bambusa and cola
meaning loving.
Holotype: MFLU 14–0822
Saprobic on decaying bamboo culms, forming dark circular erumpent spots on host surface with ascomatal ostiolar
neck with raised areas. Sexual morph Ascomata 500–
850μm diam., 350–500μm high, gregarious, solitary, immersed, globose to subglobose, black, ostiolate, soft.
Fungal Diversity
Fig. 27 Myrothecium macrosporum (holotype) a Fruiting bodies on bamboo b, d Conidia c Germinating conidium e–g Cultures on PDA. Scale bars:
a=500μm, b–d=20μm, e=25 mm, f, g=5 mm
Ostiolar neck 50–80 μm long, forming at the centre of
ascomata, raised from the host surface when mature, lined
with periphyses. Peridium comprising host and fungal tissues,
laterally 50–70μm thick, with outer layer composed of brown
and thick-walled, cells; inner layer composed of hyaline, thinwalled, large cells of textura angularis, cells 10–25×5–10μm.
Hamathecium dense, with long, 5–10μm wide, septate paraphyses, intermixed with asci. Asci 105–110×10–20μm
(x=107.1×12μm, n=20), 8-spored, unitunicate, cylindrical,
with a short furcate pedicel, with an apical ring. Ascospores
85–105×4–5.5μm (x=99.5×4.7μm, n=20), filiform, curved
to sigmoid, filiform to long fusiform, elongate, narrow
and curved at the ends, 2–3-septate when mature,
guttulate, hyaline, smooth-walled. Asexual morph
Undetermined.
Culture characters: Ascospores germinating on PDA
within 24 h with germ tubes produced from both ends.
Colonies growing slowly on PDA, reaching 45 mm in
2 weeks at 28 °C, effuse, velvety to hairy, fimbriate at
the margin, drift white from forward, dark brown from
the reverse. Mycelium superficial and immersed
branched, septate, smooth, hyaline and irregular.
Fungal Diversity
Fig. 28 Phylogram generated
from Bayesian analysis based on
combined alignment of ITS and
LSU gene regions. Bayesian
posterior probabilities (PP)
greater than 0.90 are indicated
above or below the nodes. The extypes (reference strains) are in
bold, the new isolates are in blue.
The tree is rooted with Sordaria
fimicola CBS 723.96
Material examined: THAILAND, Chiang Rai, Doi Tung,
on dead culm of bamboo (Bambusae), 4 May 2011, Dong-Qin
Dai DDQ 0004 (MFLU 14–0822, holotype), (isotype in
KUN, under the code of HKAS), ex-type living culture,
MFLUCC 11–0390. GenBank ITS: KP744449; LSU:
KP744492; SSU: KP753956.
Notes: The morphology and phylogeny of taxa in the
family Magnaporthaceae have been studied in detail
(Walker 1972; Hirata et al. 2007; Huhndorf et al.
2008; Thongkantha et al. 2009; Wong et al. 2012; Luo
and Zhang 2013; Murata et al. 2014). Our new species
can be separated from other genera in Magnaporthaceae
by morphology combined with the phylogenetic analysis
(Fig. 28). Neogaeumannomyces is similar to Gaeumannomyces
in having perithecial ascomata with long necks and cylindrical
asci producing filiform ascospores (Walker 1972). However,
Neogaeumannomyces differs in having a mycelial subiculum
growing on the host without hyphopodia, whereas
Gaeumannomyces species produce hyphopodiate mycelia
(Walker 1972).
Fungal Diversity
Fungal Diversity
Neogaeumannomyces bambusicola (holotype) a Bamboo
culm b–d Fruiting bodies on bamboo host e, f Sections of ascoma with
necks i Germinating ascospore j Paraphyses k–l Asci observed in water
m Asci observed in cotton blue n–r Ascospores s, t Cultures on PDA.
Scale bars: a=2 mm, b–d=500μm, e–f=500μm, g, h=50μm, i, j, o–r=
10μm, k–n=50μm, s, t=25 mm
Fig. 29
Meliolaceae
Species of Meliolaceae are a group of obligate parasitic fungi
with approximately 2000 species in 22 genera (Kirk et al.
2008), which are known as black mildews. They are characterized by branched, dark brown, superficial mycelium with
phialides and two-celled appressoria; dark brown, superficial
ascomata with clavate to globose, 2-4-spored asci and dark
brown, 3–4-septate ascospores. Previous studies of this family
are mostly based on morphology, as it is difficult to extract
DNA directly from the fruiting bodies. Technological progress
enabled, phylogenetic studies of the 28S nrDNA region of
several species from Meliolaceae, showing that Meliolales
represent a monophyletic order of the Sordariomycetes
(Pinho et al. 2012). A recent study carried out by Justavino
et al. (2014) indicates co-evolutionary relationships in the speciation of Meliolales on their hosts. The phylogenetic tree is
presented in Fig. 30.
20. Meliola tamarindi Syd. & P. Syd., Annls mycol. 10(1):
79 (1912)
Index Fungorum number: IF247233, Facesoffungi
number: FoF00489, Fig. 31
Epifoliar pathogen on living leaves. Sexual morph
Colonies epiphyllous, scattered, 1-2 mm diam. Hyphae superficial, substraight to crooked, branching opposite at acute to
wide angles, closely reticulate, with dark brown mycelial setae. Appressoria 20–25×9–12μm (x=22×10μm, n=10), opposite to alternate, straight to curved, two-celled, clavate to
Fig. 30 Phylogram generated from Bayesian analysis based on LSU
sequence data of Meliolaceae. Parsimony bootstrap support values
greater than 50 % are indicated above the nodes, and branches with
spathulate. Phialides 20–22(-25)×7–9μm (x=22×8μm, n=
5), few mixed with appressoria, alternate to opposite,
ampulliform. Ascomata 210–240μm wide (x=222μm, n=
10) and 170–220μm high (x=195μm, n=10), subdense, superficial on the mycelium, composed of hyaline inner cell and
dark brown outer wall with textura angularis, globose to
subglobose, dark brown, with ostiolate and dark brown setae.
Asci 65–85 × 28–40 μm (x= 73 × 34 μm, n = 5), 2-spored,
unitunicate, ellipsoid to ovoid, evanescent. Ascospores 43–48×
19–22μm (x=45×20μm, n=20), ellipsoid to fusiform, hyaline
with 2 septa when young state, becoming dark brown with 4
septa when mature, constricted at septa, middle cell slightly
longer than others. Asexual morph Undetermined.
Material examined: THAILAND, Chiang Rai province,
Bandu; on the living leaves of Tamarindus indica L.
(Fabaceae), 10 January 2014, Xiang-Yu Zeng (MFLU 14–
0282, reference specimen designated here). GenBank
LSU: KP744489.
Notes: In Fig. 30, Meliola tamarindi clusters with a putatively named strain of Asteridiella obesa and four other
Meliola species. Asteridiella sp., Endomeliola dingleyae and
Appendiculella sp. also cluster in Meliolaceae (Fig. 30). Our
collection of Meliola tamarindi has larger spores than in the
protologue (43–48 × 19–22 μm, versus 36–44 × 13–17 μm
(Sydow and Sydow 1912). The addition of reference specimen sequence data for M. tamarindi is important for this phylogenetically poorly understood group.
Sydowiellaceae
Sydowiellaceae (Diaporthales) is based on Sydowiella and its
generic type S. fenestrans (Duby) Petr. Several fungal genera
and their species are accommodated in this family, however
they do not have any clear features in common (Rossman et al.
2007). Genera in this family include Chapeckia, Hapalocystis,
Rossmania, Stegophora and Sillia. These taxa are plant
Bayesian posterior probabilities greater than 0.95 are given in bold. The
ex-types (ex-epitypes, reference strains) are in bold, the new isolates are
in blue. The tree is rooted with Chaetomium globosum CBS 147.51
Fungal Diversity
Fig. 31 Meliola tamarindi (MFLU 14–0282). a Host b Host leaves c
Colony on leaf d Ascomata e Seta f Apex of seta g Mycelium with
phialides and appressoria h Cross section of ascoma I Peridium, j–o
Immature and mature asci, p–s Immature and mature ascospores. Scale
bars: c-d=200μm, e=100μm, h-i=50μm, f-g, j-s=20μm.
pathogens and saprobes on herbaceous, dicotyledonous plants
and hardwood trees. The phylogenetic tree is presented in
Fig. 32.
245μm, n=15), immersed, solitary or aggregated, scattered,
visible through dark cracks on the host surface, subglobose to
globose, ostiolate. Ostiole 130–195 μm high× 90–113 μm
wide (x=145×105μm, n=10), short papillate, black, cylindrical, periphysate. Periphyses hyaline. Peridium 4–16μm wide
(x=10μm, n=15) black, thick-walled, 3–5-layered of cells of
textura angularis. Paraphyses 3–7μm wide, few, septate, hyaline, attached to the base, longer than asci. Asci 112–127×
46–80μm (x=121×66μm, n=20) 4 or 8-spored, unitunicate,
clavate to fusoid, with a short pedicel, with an inconspicuous
flat, refractive ring at the lower end of the thickened apical
21. Hapalocystis berkeleyi Auersw. ex Fuckel, Fungi
rhenani exsic., fasc. 6: no. 585 (1863)
Index Fungorum number: IF357284, Facesoffunginumber:
FoF00413; Figs. 33 and 34
Reference specimen: MFLU14–0798
Saprobic on bark of Platanus hybrid. Sexual morph
Ascomata 90–185 μm high × 206–302 μm wide (x= 150 ×
Fungal Diversity
Fig. 32 Phylogram generated from Maximum Parsimony analysis based
on LSU sequence data of Sydowiellaceae. Parsimony 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
in bold. The ex-types (ex-epitypes and reference strains) are in bold; the
new isolates are in blue. The tree is rooted with Diaporthe padi AR 3419
and D. pustulata AR 3430
wall, apex narrow and blunted. Ascospores 34–42×12–18μm
(x=37×17μm) inequilaterally ellipsoidal, with broadly rounded ends, initially hyaline, dark brown at maturity, (1–)2(–3)septate, with cells of equal length, thick and smooth-walled,
with short, hyaline strap-like appendages situated at both
rounded ends with same width as the ascospore. Asexual
morph Conidiomata on MEA 165–195 μm diam. (x=
182μm, n=15), pycnidia, superficial, aggregated, 3–5 in one
group, globose, orange to brown, Conidiomatal wall 7–13μm
wide (x=10μm, n=10) small, thick-walled, orange, 5–10 cells
layers of textura angularis. Conidiophores 1.5–2μm high×1–
1.5μm wide (x=2×1.5μm, n=10) branched, hyaline, short,
few conidiogenous cells arising from one conidiophore, attached to conidiomatal wall. Conidiogenous cells 10–14μm
long, 5–7μm wide, cylindrical, hyaline, bottle-shaped, septate, ends pointed, phialidic. Conidia 0.5–1.5 diam. (x=
1μm, n=20) ellipsoid, one-celled, hyaline, smooth-walled.
Culture characters: Ascospores germinating on MEA after
2 days at 16 °C forming hyaline, branched germ tubes from end
cells. Colonies on MEA reaching 2 cm after 14 days at 25 °C in
light, greenishash, effuse, circular, margin entire, surface rough.
Material examined: ITALY, Province of Forlì-Cesena [FC],
Modigliana, Montebello, on branch of Platanus hybrid, 14
April 2013, E. Camporesi It 1187 (MFLU 14-0798, reference
specimen of Hapalocystis berkeleyi designated here); extype living cultures, MFLUCC 13-0662. GenBank LSU:
KP744486.
Notes: Hapalocystis (Sydowiellaceae, Diaporthales) is typified by H. berkeleyi. LSU gene analysis of diaporthalean
fungi showed Hapalocystis as a distinct genus in
Sydowiellaceae (Castlebury et al. 2002). Jaklitsch and
Voglmayr (2004) evaluated Hapalocystis introducing a new
species. Hapalocystis comprises H. occidentalis, H. ulmi,
H. bicaudata, H. corticalis and H. berkeleyi (H. berkeleyi
Auersw. ex Fuckel var. berkeleyi and H. berkeleyi var. kickxii
(Westend.) M.E. Barr (Jaklitsch and Voglmayr 2004). The
asexual morph of Hapalocystis is given as Stilbospora,
Hendersonia or Dothiorella (Wehmeyer 1941), or as
stilbospora-like (Barr 1978). Glawe (1985) reported a Phomalike asexual morph for H. berkeleyi in culture. In this study we
observed a phoma-like asexual morph on MEA and we made
fresh collections of Hapalocystis berkeleyi which is identical to
the morphology of type specimen. The type specimen is in
good condition (at herbarium K) and hence, we designate this
as a reference specimen (Ariyawansa et al. 2014a) and provide
molecular data; the asexual morph was also produced in culture. LSU gene analysis of taxa in family Sydowiellaceae
showed our reference strain of H. berkeleyi cluster with other
H. berkeleyi strains with 100 % bootstrap support (Fig. 32).
Valsaceae
The family Valsaceae (Diaporthales, Ascomycota) was
established by Tulasne and Tulanse (1861) and later recognized as family of Diaporthales by Barr (1978). It includes
important phytopathogens that cause dieback and canker disease on a wide range of plants, causing severe commercial and
ecological damage and significant losses worldwide (Adams
et al. 2005; Fan et al. 2014). The taxa are characterized by
aggregated ascomata surrounded by well-developed
entostromata, emerging beaks arranged centrally through a
white to black disc, having allantoids ascospores (Castlebury
et al. 2002; Rossman et al. 2007). Previously, the Valsaceae
was restricted to five genera, i.e. Cytospora and its segregated
sexual morph, i.e. Leucostoma, Valsa, Valsella, and
Valseutypella (Fries 1823; Saccardo 1884; Spielman 1985;
Adams et al. 2002; Castlebury et al. 2002). However, all the
genera were recently combined under Valsa as subgenera or
species (Adams et al. 2005). Cytospora (1818) is an older
name than Valsa (1849) and the asexual morph is more common in nature; therefore, we chose to adopt Cytospora and treat
Valsa as synonym of Cytospora. Cytospora is characterised by
a single or labyrinthine of locules (or diaporthalean-like perithecia), filamentous conidiophores (or clavate to elongate obovoid asci) and allantoid hyaline conidia (or ascospores)
(Spielman 1985; Adams et al. 2005). In most conditions, the
conidia emerge and spread from fruiting bodies in yellow to red
gelatinous tendrils (Adams et al. 2005, 2006). Cytospora contains 110 species in Kirk et al. (2008). Ex-type sequence data,
are however, available for only a very few species rendering
identification to species level difficult. Therefore, research into
cryptic species and a backbone tree for Cytospora is needed as
Fungal Diversity
Fungal Diversity
for other pathogenic genera (Hyde et al. 2014). The phylogenetic trees are presented in Figs. 35 and 36.
22. Cytospora berberidis C.M. Tian, X.L. Fan & K.D.
Hyde, sp. nov.
MycoBank MB811223, Facesoffungi number: FOF00491;
Fig. 37
Etymology: berberis (Lat.), referring to Berberis
dasystachya, on which the fungus was collected.
Holotype: BJFC-S681.
Fig. 34 “Phoma” like asexual morph of H. berkeleyi (MFLUCC 130662)a Germinating ascospore b Culture on upper surface, growing on
MEA c-f Conidiomata g Cross section of conidiomata h Peridium i–p
Conidiophores, conidiogenous cells and conidia q Conidia. Scale bars:
a=50μm, c=1000μm, d-f=500μm, g=100μm, h=20μm, i-p=10μm,
q=5μm
Hapalocystis berkeleyi (MFLU 14-0798) a Ascomata on
substrate b Cross section of ascomata c Peridium d–k Asci l–r
Ascospores. Scale bars: a=1000μm, b=100μm, c=20μm, d-k=50μm,
l-r=20μm
Fig. 33
Fungal Diversity
Fig. 35 Phylogram generated
from parsimony analysis based on
the combined genes of ITS,
nrLSU, RPB2 and ACT gene
regions. Bootstrap support values
above the branches indicate
maximum parsimony bootstrap
(MPBP≥50 %) and maximum
likelihood bootstrap (MLBP≥
70 %), and branches with
Bayesian posterior probabilities
greater than 0.95 are given in
bold. The ex-types (ex-epitypes)
are in bold, the new isolates are in
blue. The tree is rooted with
Phomopsis vaccinii ATCC 18451
Pathogen on twigs and branches of Berberis dasystachya.
Sexual morph Undetermined. Asexual morph Stromata immersed in bark. Conidiomata, erumpent through the surface of
bark, discoid, nearly flat, with a solitary undivided locule.
Conceptacle absent. Disc white to light brown, circular to
ovoid, (0.32–)0.36–0.41(–0.47) mm (x=0.38 mm, n=20),
with one ostiole per disc. Ostiole in the centre of the disc, at
the same level as the disc surface, (12.1–)13.9–18.5(–21.7)
mm (x= 16.3 mm, n = 20) diam. Locule uniloculate,
(0.63–)0.69–0.87(–0.91) mm (x= 0.78 μm, n = 20) diam.
Conidiophores hyaline, unbranched or occasionally branched
at the bases, (16.4–)17.5–25.5(–26.2) μm (x=21.7μm, n=20).
Conidia hyaline, elongate-allantoid, aguttulate, aseptate,
(5.8–)6–6.9(–7.1)×(1.8–)1.9–2.1(–2.2) μm (x in 6.5×2μm,
n=50).
Culture characters: Colony white, flat, felty, texture uniform, conidiomata sparse, irregularly distributed.
Material examined: CHINA, Qinghai Province, Huzhu,
Jiading, 36°56′12.62″ N, 102°30′11.85″ E, 2324 m asl., on
twigs and branches of Berberis dasystachya Maximowicz
(Berberidaceae), 15 August 2012, X.L. Fan (BJFC-S681,
holotype); ex-type living culture, CFCC 89927. GenBank
ITS: KP340985; LSU: KP340989; ACT: KP340997; RPB2:
KP340993; ibid., living culture, CFCC 89933. GenBank ITS:
KP340986; LSU: KP340990; ACT: KP340998; RPB2:
KP340994.
Notes: Cytospora berberidis with a single locule, differs
from most Cytospora species with multi-locules, including
type species C. chrysosperma. The new species can be distinguished from C. pruinosa and C. atrocirrhata with
single locules by the absence of conceptacles and size
of conidia. Molecular analysis also confirms the novelty
of the species which forms a single clade with high
support (Figs. 35 and 36). This novel species represents
the first record of Cytospora isolated from Berberis
dasystachya.
23. Cytospora sibiraeae C.M. Tian, X.L. Fan & K.D.
Hyde, sp. nov.
MycoBank MB811222, Facesoffungi number: FOF00490,
Fig. 38
Etymology: sibiraeae (Lat.), referring to Sibiraea
angustata, on which the fungus was collected.
Holotype: BJFC-S783.
Pathogen on twigs and branches of Sibiraea angustata.
Sexual morph Stromata immersed in bark. Ascostromata
erumpent through the surface of bark, circular to ovoid, extending to a large circular area, (0.98–)1.15–1.53(–1.71) mm
(x=1.32 mm, n=20) diam., 4–10 locules arranged at one
depth in the bark, conceptacles dark. Disc deep grey to black,
nearly flat, circular to ovoid, (0.28–)0.32–0.38(–0.41) mm
(x=0.35 mm, n=20) diam. Ostioles numerous, dark brown
to black, at the same level as the disc, occasionally area below
disc a lighter entostroma, (67.2–)71.3–107.1(–126.3) μm
(x=88.2μm, n=20) diam. Locules dark brown, arranged
circularly, flask-shaped to sphaerical, (0.22–)0.28–0.35(–
0.39) mm (x=0.34 mm, n=20) diam. Asci (36.3–) 41.5–
49.9 (–50.1)×(5.2–)6.4–7.8(–9.7) μm (x in 45.7×7.4μm,
n = 20), 8-spored, free, clavate to elongate-obovoid.
Ascospores (7.8–)8.1–13.3 (–13.6)× (1.9–)2.1–2.5(–3.1)
Fungal Diversity
Fig. 36 Phylogram generated from parsimony analysis based on the ITS
gene region. Bootstrap support values above the branches indicate
maximum parsimony bootstrap (MPBP ≥ 50 %) and maximum
likelihood bootstrap (MLBP ≥ 70 %), and branches with Bayesian
posterior probabilities greater than 0.95 are given in bold. The ex-types
(ex-epitypes) are in bold; the new isolates are in blue. The tree is rooted
with Phomopsis vaccinii ATCC 18451
μm (x in 11.7 × 2.3 μm, n = 50), biseriate, elongateallantoid, thin-walled, hyaline, lacking guttules, aseptate.
Asexual morph Undetermined.
Culture characters: Colony white, flat, felty, texture uniform, conidiomata sparse, irregularly distributed.
Material examined: CHINA, Gansu Province, Gannan,
Lintan, 34°31′50.10″ N, 103°08′32.47″ E, 2750 m asl., on
twigs and branches of Sibiraea angustata (Rehder) Hand.-
Mazz. (Rosaceae), 8 August 2012, X.L. Fan (BJFC-S783,
holotype); ex-type living culture, CFCC 50045. GenBank
ITS: KP340987; LSU: KP340991; RPB2: KP340995; ibid.,
living culture, CFCC 50046. GenBank ITS: KP340988; LSU:
KP340992; RPB2: KP340996.
Notes: Cytospora species with black conceptacles were
previously placed in the genus Leucostoma. Adams et al.
(2005) redefined Cytospora using ITS phylogeny and
Fungal Diversity
Fig. 37 Cytospora berberidis
(holotype) a Habit of
conidiomata on a twig b
Transverse sections through
conidiomata c Longitudinal
sections through conidiomata d
Conidiophores e Conidia. Scale
bars: a-c=0.5 mm; d–e=10μm
combined Leucostoma and other sexual morphs (Valsa,
Valsella and Valseutypella) under Valsa, as subgenera or species with no additional infrageneric rank. Cytospora sibiraeae
possesses obviously black conceptacles, which are similar to
those of C. nivea and C. atrocirrhata. However the taxon
clusters in an individual clade in ITS and multi-gene
phylograms (Figs. 38 and 39). Few pathogenic fungi have
been reported from Sibiraea angustata. This novel species
represents the first record of a Cytospora isolated from
Sibiraea angustata.
Fig. 38 Cytospora sibiraeae (holotype) a Habit of ascomata on a twig b Multi-ostioles on disc c Transverse sections through ascomata d Longitudinal
sections through ascomata e ascus f ascospores. Scale bars: a–d=1 mm; e–f=10μm
Fungal Diversity
Fig. 39 Phylogram generated from Maximum likelihood (RAxML)
analysis based on combined ITS, LSU, RPB2 and β-tubulin sequenced
data of Xylariaceae. Maximum Likelihood bootstrap support values
greater than 50 % are indicated above or below the nodes. The ex-types
(ex-epitypes) are in bold. New sequences are in blue. The tree is rooted
with Sordaria fimicola CBS 723.96
Xylariaceae
Isotype: BPI 738677
Saprobic on corticated and decorticated wood. Sexual
morph Ascostromata 0.5–8×0.5–1.2×0.06–0.13 cm (x=4×
0.8×0.08 cm), pulvinate to effuse-pulvinate, with inconspicuous perithecial mounds, surface sepia (63), dark brick (60),
or fuscous (103); blackish granules immediately beneath surface and between perithecia, with KOH-extractable pigments
greenish-olivaceous (90), dull green (70), or dark green (21);
the tissue below the perithecial layer inconspicuous, perithecia
0.3–0.5 diam. × 0.3–0.7 mm high (x=0.4×0.5 mm), immersed, obovoid to tubular, ostioles conical-papillate,
encircled with a convex truncatum-type disc 0.2–0.3 mm
(x=0.3 mm) diam., paraphyses not seen. Asci (120–)125–
160(–167)×(3.8–)4–5.5(–5.8) μm (x=147×4.5μm, n=20),
8-spored, unitunicate, cylindrical, pedicellate, apical ring bluing in Melzer’s reagent, discoid, 0.5×1.5–2μm (x=0.5×
1.8μm, n=20). Ascospores (7.2–)7.5–13.8(–14)×(3.6–)4–
6(–6.2) μm (x=9.7×5.5μm, n=30), uniseriate, one-celled, ellipsoid, inequilateral, with narrowly rounded ends, pale
brown, with straight germ slit much less than spore-length
and originating from one end, perispore indehiscent in 10 %
KOH, epispore smooth. Asexual morph Reported by Ju and
Rogers (1996), found on the surface of young stromata on
natural substrates, Conidiogenous structures periconiella-like,
dark brown, coarsely roughened. Conidiogenous cells pale
Phylogenetic analysis of combined gene sequence data (ITS,
LSU, RPB2 and β-tubulin) resolve Xylariaceae as two major
monophyletic groups representing the subfamilies Xylarioideae
(93 % bootstrap support) and Hypoxyloideae (79 % bootstrap
support). The Xylarioideae clade contains 14 resolved
s u b c l a d e s n a m e d A m p h i ro s e l l i n i a , A s t ro c y s t i s ,
Brunneiperidium, Collodiscula, Discoxylaria, Entoleuca,
Euepixylon, Lunatiannulus, Kretzschmaria, Nemania,
Podosodaria, Poronia, Rosellinia and Vamsapriya. The
Hypoxyloideae contains ten well-resolved subclades named
A n n u l o h y p o x y l o n, A n t h o c a n a l i s , A n t h o s t o m e l l a ,
Biscogniauxia, Daldinia, Hypoxylon, Rhopalostroma,
Rostrohypoxylon, Ruwenzoria and Thamnomyces.
Fasciatispora nypae K.D. Hyde is placed as a basal group of
Xylariaceae (70 % bootstrap support) parallel to Creosphaeria
and Pyriformiascoma, but as a singleton. Annulohypoxylon
thailandicum is a new species introduced here and clustered
with other Annulohypoxylon species (80 % bootstrap support).
The phylogenetic tree is presented in Fig. 39.
25. Annulohypoxylon leptascum (Speg.) Y.M. Ju., et al.,
Mycologia 97(4): 859 (2005).
Facesoffunginumber: FoF00296, Fig. 40
Fungal Diversity
Fungal Diversity
Annulohypoxylon leptascum (MFLU 13–0118). a Stromatal
habit in wood b Ostioles seen from above c Stroma inside view d Cross
section of the stroma showing perithecia e Pigment formation in KOH f, g
Mature ascus in water h, i Ascospore in water j Ascospore showing germ
slit k Dehiscent perispore in KOH. Scale bars: a-c=5 mm, d=1 mm, f–
k=10μm
Fig. 40
brown, smooth to finely roughened. Conidia hyaline, smoothwalled, ellipsoid.
Culture characters: Colonies on OA at 25–28 °C reaching
the edge of 6 cm in 7 days, whitish, velvety to felty, azonate,
with diffuse margins, reverse at first yellow green (71) and
turning greenish-olivaceous (90) in the centre after 5–7 days.
Material examined: BRAZIL, Apiahy, on wood, 1888,
Puiggari (BPI 738677, isotype); THAILAND, Chiang Mai,
Doi Inthanon, on decaying wood, 3 December 2012, D.A.
Daranagama and K.D. Hyde AXL 036 (MFLU 13–0118),
living cultures, MFLUCC 12–0829, ICMP).
Notes: Annulohypoxylon leptascum and A. truncatum are
the potential taxa that could be compared with this specimen.
Both these taxa share common stromatal characters with KOH
extractable pigments. The inconspicuous tissue below the
perithecial layer in the specimen is a feature similar to
A. leptascum. The perithecia are obovoid as in A. leptascum.
Even with slight deviations in sizes, the asci and ascospores
are quite similar to that of A. leptascum. The ascus apical ring
was however not observed as mentioned in the description of
A. leptascum (Ju and Rogers 1996). Annulohypoxylon
elevatidiscus reported in Taiwan and described in Ju et al.
(2004) is also a possible match with our specimen. It has
similar stromatal characters except for the conspicuous tissue
layer beneath perithecia. However, A. elevatidiscus has large,
sphaerical perithecia compared with this specimen. In addition, the dehiscent perispore in KOH was not observed in this
specimen. Annulohypoxylon leptascum has so far been reported only from temperate countries and hence recorded for the
first time from Thailand with the new molecular data.
26. Annulohypoxylon nitens (Ces.) Y.M. Ju & J. D.
Rogers, A Revision of the Genus Hypoxylon, p. 220. 1996.
Facesoffungi number: FoF00029, Figs. 41 and 42
Isotype: IMI 339738
Saprobic on corticated and decorticated wood. Sexual
morph Ascostromata 0.5–7×0.25–5×0.07–0.2 cm thick (x=
4.5 ×3.5 ×0.12 cm), glomerate, hemisphaerical to effusepulvinate, with perithecial mounds exposing up to 1/4–1/2;
when young surface dark brown vinaceous (84), becoming
blackish at maturity, with reddish-brown tone, finally shiny
black; blackish granules immediately beneath surface, with
KOH-extractable pigments, greenish-olivaceous (90), woody
tissue below the perithecial layer, blackish, perithecia 0.5–
1 mm diam. (x=0.7 mm), sphaerical, ostioles conical-papillate, encircled with a flattened bovei-type disc, 0.2–0.5 mm
diam. (x=0.4 mm), paraphyses, not seen. Asci (105–)110–
140(–145)×(3.5–)4–5(–5.2) μm (x=125×4.2μm, n=20), 8spored, unitunicate, cylindrical, pedicellate, apical ring bluing
in Melzer’s reagent, discoid, 0.5×1–1.5μm (x=0.5×1.2μm).
Ascospores (6.2–)6.5–10(–10.3)×(2.8–)3–4.5(4.9) μm (x=
8.4 × 3.8 μm, n = 30), uniseriate, one-celled, ellipsoidinequilateral, with narrowly rounded ends, light brown to
brown, with straight germ slit the entire spore-length,
perispore dehiscent in 10 % KOH, epispore smooth.
Asexual morph Sporulating regions scattered over entire surface of colony, honey (64) to fawn (87). Conidiogenous
structure: nodulisporium-like, brown, thickened and dense.
Conidiogenous cells 10–24×2–3μm (x=20×1.6μm, hyaline,
smooth-walled. Conidia 4–5×2.5–3μm (x=4.3×2.7μm), hyaline, smooth to finely roughened, ellipsoid.
Culture characters: Colonies on OA at 25–28 °C reaching
7 cm in 7 days, averse whitish with black pigments, azonate
with diffuse margins, reverse honey (64) or dull green (70),
azonate.
Material examined: THAILAND, Chiang Mai, Doi Pui
mountain, on decaying wood, 3 November 2012, D.A.
Daranagama and K.D. Hyde AXL 030 (MFLU 13–0109),
living cultures, MFLUCC 12–0823, ICMP. GenBank ITS:
KJ934991; LSU: KJ934992; RPB2: KJ934994.
Notes: Annulohypoxylon nitens (Ces.) Y.M. Ju et al. and
Hypoxylon truncatum (Schwein.) Y.M. Ju et al. differ in their
stromatal characters in that the former has a bovei-type ostiolar
disc as well as more effuse stromata. Unlike many other related taxa, A. nitens possesses stromata which are thinly covered
by fragments of the outermost stromatal layer exposing free
perithecial mounds (Ju and Rogers 1996). In identification,
these stromatal characters are quite useful. Another striking
feature is that A. nitens has a dull reddish diffusible colour in
the stromata though at maturity; it is black and shiny with
woody layer beneath the perithecia (Ju and Rogers 1996).
The asexual morph of this species is nodulisporium-like and
the conidiophores are likely brown bearing hyaline
conidiogenous cells and conidia. Many records of A. nitens
in GenBank have only ITS and β-tubulin data from authentic
strains. We hereby contribute sequences in addition from LSU
and RPB2 for molecular analyses.
27. Annulohypoxylon stygium (Lév.) Y.M. Ju et al.
Mycologia 97(4): 861 (2005)
Facesoffunginumber: FoF000298, Fig. 43
Isotype: K 42247
Saprobic on decorticated wood. Sexual morph
Ascostromata 1–10 × 0.5–3 × 0.04–0.1 cm (x= 6 × 1.8 ×
0.08 cm), pulvinate to effuse-pulvinate, with conspicuous
perithecial mounds, surface shiny blackish, with reddishbrown tone, dull reddish brown granules immediately beneath
surface and between perithecia, with KOH-extractable pigments greenish-olivaceous (90) or dull green (70), the tissue
below the perithecial layer inconspicuous, perithecia obovoid,
Fungal Diversity
Fungal Diversity
Annulohypoxylon nitens (MFLU 13–0109) a Stromatal habit in
wood b Ostioles in stroma c Stromata from side view d Alignment of
perithecia in across section e Perithecia from above f Formation of
coloured pigments in KOH g, h Mature ascus in water i Ascus in
Melzer’s reagent showing inconspicuous apical apparatus j–l
Ascospores in water m Ascospore showing the germ slit n Indehiscent
perispore in KOH. Scale bars: a-c=5 mm, d, e=1 mm, g- n=10μm
Fig. 41
0.2–0.3×0.3–0.5 mm (x=0.3×0.4 mm), ostioles papillate,
encircled with a convex truncatum-type disc 0.1–0.2 mm
diam. (x=0.15 mm). Asci (48–)55–90(–97)×(2.8–)3–4(–4.7)
μm (x=73×3.7μm, n=20), 8-spored, unitunicate, cylindrical,
pedicellate, apical ring bluing in Melzer’s reagent, discoid,
0.5 × 1 μm (x= 0.3 × 0.4 μm, n = 2 0). Ascospores (5.3–)6–
10.5(–10.9)×(2.4–)3–5.5(–5.8) μm (x=8.3×4.4μm, n=30),
uniseriate, one-celled, ellipsoid-inequilateral, with narrowly
rounded ends, light brown, with straight germ slit sporelength on flattened side, perispore dehiscent in 10 % KOH,
smooth, epispore smooth. Asexual morph Undetermined.
Fig. 42 Annulohypoxylon
nitens on OA after 2 weeks a
From above white with black
pigments b From below honey
(64) or dull green (70) c
Conidiogenous structure d
Conidiogenous cells e Conidia.
Scale bars: c–d=10μm
Culture characters: Colonies on OA at 25–28 °C reaching
5 cm in 7 days, from above, at first whitish developing hazel
(88) shade in the middle after 5–7 days with the time the hazel
shade spreads over the entire plate, azonate with diffuse margins, reverse at first citrine greenish-olivaceous (90) and developing black patches in the centre after 5 days, azonate.
Material examined: SRI LANKA, 1915, T. Petch 4465 (K
42247, isotype), THAILAND, Chiang Mai, Doi Inthanon
National Park on decaying wood, 3 December 2012, D.A.
Daranagama and K.D. Hyde, AXL 034 (MFLU 13–0109),
living cultures, MFLUCC 12–0826, ICMP. GenBank ITS:
KJ940870; LSU: KJ940869; RPB2: KJ940868.
Notes: Annulohypoxylon stygium, A. nitens and
A. truncatum were possible species to accommodate this specimen regarding the stromatal characters. In the latter two species, the asci possess longer stipes, whereas in this specimen,
the stipe is short. Annulohypoxylon nitens has KOH extractable greenish-olivaceous pigments, whereas in the studied
Fungal Diversity
Fungal Diversity
Annulohypoxylon stygium (MFLU 13–0109) a Stromatal habit
in wood b, c Ostioles seen from above d Stroma in side view e Cross
section of the stroma showing perithecia f Pigment formation in KOH g
Perithecia from above h-j Mature ascus in water k Ascus in Melzer’s
reagent showing weakly stained inconspicuous amyloid ascal apical
apparatus l Ascospore in water m Dehiscent perispore in KOH
n Ascospore showing germ slit. Scale bars: a = 1 cm, b-g =
0.5 mm, h–n=10μm
Fig. 43
specimen, the pigment is more towards green series. In this
specimen, there are more conspicuous, glomerate, perithecial
mounds, in contrast to the description given by Ju and Rogers
(1996). The majority of the sequences deposited in the
GenBank are ITS and a few reliable sequences for β-tubulin.
In this study, we provide new LSU and RPB2 sequences from
an authentic strain.
27. Annulohypoxylon thailandicum Daranagama & K.D.
Hyde, sp. nov.
Index Fungorum number: IF550799, Facesoffunginumber:
FoF00367, Fig. 44
Etymology: refers to the country, Thailand where the new
species was collected.
Holotype: MFLU 13–0441
Saprobic on corticated and decorticated wood, Sexual
morph Ascostromata 0.5–0.7×0.8–1.7×0.8 mm (x=0.7×
1.5×0.8 mm), effuse–pulvinate, with conspicuous perithecial
mounds exposing almost completely, sphaericalhemisphaerical, surface black, carbonaceous, blackish granules
immediately beneath surface and between perithecia, with KOH
extractable pigments greenish-olivaceous (90), perithecia 0.35–
0.5×0.3–0.7 mm (x=0.4×0.5 mm), immersed, encased in carbonaceous tissue, sphaerical, ostioles coarsely papillate,
encircled with a small truncatum–type disk, 0.3–0.45 mm
diam., paraphyses not seen. Asci (76.7–)89–100.8(–
107) × (4.1–)4.5–6.5(–6.9) μm (x= 97 × 5.6 μm), 8-spored,
unitunicate, cylindrical, short-pedicellate, with apical ring faintly
bluing at base or not bluing in Melzer’s reagent, globose, 1.5×
2μm. Ascospores (5.8–)6–11.5(12)×(3.2–)4–6.5(6.8) μm (x=
12.5×5.6μm), uniseriate, one-celled, ellipsoidal inequilaterally
with narrowly rounded ends, brown, with a straight germ slit
running full length of the spore, epispore smooth, perispore
dehiscent in 10 % KOH. Asexual morph Undetermined.
Culture characters: Colonies on OA at 25–28 °C reaching
5 cm in 7 days, whitish colonies, azonate with diffuse margins,
reverse at first whitish and turning light brown after 3–4 days,
developing black pigments after 8–10 days.
Material examined: THAILAND, Chiang Mai, Doi
Suthep, on decaying wood, 10 November 2012, D.A.
Daranagama and K.D. Hyde AXL 107 (MFLU 13–0441,
holotype); ex-type living culture, MFLUCC 13–0118,
ICMP. GenBank ITS: KP744434; LSU: KP744476.
Notes: Annulohypoxylon thailandicum, with characters
such as brown ascospores with a germ slit originating from
one end more or less up to the full length of the spore, greenish
olivaceous/dull green KOH extractable pigments, dehiscent
perispore in 10 % KOH and ostioles encircled with a small
disk, can be compared with A. archeri or A. microcarpum.
Both A. archeri and A. microcarpum have ostiolar disks of
around 0.2 mm diam., with conspicuous perithecial mounds
with reddish tones and sphaerical perithecia. A. microcarpum
has rather small perithecia (0.15–0.2 mm diam) and small
ascospores (7–8×3–4μm), whereas Annulohypoxylon archei
has perithecia of 0.1 mm diam. and ascospores of 9–10.5μm×
4–5μm (Ju and Rogers 1996). Therefore, A. archeri could be
morphologically the most simialr taxon. However, the new
species has a black surface, larger perithecia and wider ostiolar
disks as well as smaller ascospores, as compared to A. archeri.
28. Biscogniauxia marginata (Fr.: Fr.) Pouzar, Ceska
Mykology. 33: 216. 1979.
Facesoffunginumber: FoF000299, Figs. 45 and 46
Holotype: UPS: BOT:F-175466
Saprobic on wood. Sexual morph Ascostromata 3.5–7×
2–3 mm (x=5.6×2.8 mm), raised–discoid, globose, with concave surface, distinct raised margins, outer dehiscing layer,
surface black, woody layer immediately beneath the stromatal
surface and between perithecia with carbonaceous tissue
encasing each ostiole, tissue beneath perithecia comprising
with host tissue, carbonaceous, perithecia obovoid, 0.2–0.5×
0.5–1 mm (x=0.4×0.8 mm), thick-walled, light brown inner
cell layers, outer cell layers carbonaceous, ostioles slightly
umbilicate with punctuate openings, with white residues. paraphyses numerous, between asci, filamentous, septate, 3–5×
50 μm or longer (x= 4.4 × 50 μm). Asci (145–)160–200(–
204)×(8.3)9–12(12.2) μm (x=176×13μm, n=20), 8-spored,
unitunicate, cylindrical, pedicellate, with apical ring bluing in
Melzer’s reagent, discoid, 0.5–0.7×1.5–2μm (x=0.6×1.7μm,
n–20). Ascospores (12.4–)13–16.5(–16.8)×10–13.5μm (x=
15 × 11.7 μm, n = 30 ), uniseriate, one-celled, globose to
subglobose, with broadly rounded ends, dark brown to black,
with sigmoid germ slit the entire spore-length, smooth-walled.
Asexual morph Sporulating regions scattered over entire central part of the colony, brown-vinaceous (84) after 4 weeks.
Conidiogenous structure nodulisporium-like, arising as
roughened masses of hyphae, hyaline. Conidiogenous cells
50–60×4–5μm (x=56×4.4μm), hyaline, finely roughened.
Conidia 6–8×3.5–4μm (x=7.2×3.8μm), hyaline, smooth to
finely roughened, ellipsoid.
Culture characters: Colonies on OA at 25–28 °C reaching
the edge of 6 cm Petri-dish in 14 days, whitish, velvety to felty,
azonate, with diffuse margins, reverse at first straw (46), later
developing into dark brown colonies after 3–4 weeks.
Material examined: FRANCE, on wood, 16 April 2012,
Erio Gardiennet AXL 001 (MFLU 13–0099), living cultures,
MFLUCC 12–0740, ICMP. GenBank ITS: KJ958407; LSU:
KJ958408; RPB2: KJ958409.
Fungal Diversity
Fungal Diversity
Annulohypoxylon thailandicum (holotype) a Stromatal habit
on wood b Perforations seen from above c Ostioles with ostiolar disks d
Stromatal surface in side view e Perithecia from above f Cross section of
the stroma showing perithecia g Formation of pigments in KOH h Mature
ascus in water i Ascus in Melzer’s reagent showing inconspicuous apical
apparatus j, k Ascospores in water l Ascospore with germ slit m
Dehiscent perispore in 10 % KOH. Scale bars: a–d=5 mm, e, f=1 mm,
h–m=10μm
Fig. 44
Notes: Biscogniauxia marginata and B. baileyi are similar
in their stromatal characters. Both have subglobose ascospores
which make them different from many of the other species in
the genus. However, B. marginata has globose ascospores
with a sigmoidal germ slit, while B. baileyi has straight germ
slit. Biscogniauxia marginata has a nodulosporium-like asexual morph as observed here. Callan and Rogers (1986), Petrini
and Müller (1986) and Ju and Rogers (1996) mentioned
the asexual morph as nodulosporium–like, while Whalley
and Edwards (1985) recognised the asexual morph as
geniculosporium-like. In GenBank there is only one ITS sequence of belonging to B. marginata and we add ITS, LSU,
RPB2 and β-tubulin gene data from an authentic specimen.
29. Fasciatispora nypae K.D. Hyde., Trans. Mycol. Soc.
Japan 32(2): 267 (1991); Fig. 47
Material examined: THAILAND, Tambon Ngao, near
Ngao Mangrove Forest Research Centre, Ranong Province,
on frond of Nypa fruticans Wurmb. (Arecaceae), 11 January
2011; Jian-Kui Liu, JKA 0066 (MFLU 15-0042, reference
specimen designated here), living culture MFLUCC 110382. GenBank ITS: JN846716; LSU: KP744484.
Notes: This fungus was collected from Nypa fruticans, in
Ranong mangroves, Thailand. Due to the paucity of ascomata
on the specimen, we were only able to isolate a few ascospores. Thus we provide a reference sequence for the first time
for this genus in this study.
30. Hypoxylon fendleri Berk. ex Cooke, Grevillea 11(no.
60): 132 (1883).
Facesoffunginumber: FoF00301, Figs. 48 and 49
Holotype: K 120974
Saprobic on wood, Sexual morph Ascostromata 3–7×
0.05–0.08 cm (x=5.3×0.07 cm), effuse-pulvinate, with inconspicuous to conspicuous perithecial mounds, surface brown
vinaceous (84), dark vinaceous (82), dark brick (60), sepia
(63) or, less frequently, bay (6); orange red granules immediately beneath surface and between perithecia, with KOHextractable pigments orange (7), the tissue below the perithecial layer inconspicuous, perithecia obovoid, 0.2–0.4×0.3–
0.6 mm (x=0.3×0.5 mm), paraphyses not seen, ostioles lower
than the stromatal surface. Asci (98–)100–156(–
160) × (6.1–)6.5–10(–10.2) μm (x= 143 × 8 μm, n=20), 8spored, unitunicate, cylindrical, pedicellate, with discoid apical ring bluing in Melzer’s reagent, 0.5–1.2×1.8–2.5μm (x=
0.9×2μm, n=20). Ascospores (9.5–)10–15(15.3)×(4.6–)5–
7(–7.4) μm (x=14×6μm, n=30), uniseriate, one-celled, ellipsoid-inequilateral, with narrowly rounded ends, brown to dark
brown, with sigmoid germ slit spore-length, perispore dehiscent in 10 % KOH, smooth or with inconspicuous coil-like
ornamentation, epispore smooth. Asexual morph
Conidiogenous structure nodulisporium-like, arising from
brownish, roughened masses. Conidiogenous cells hyaline,
finely roughened, 10–20×2–3μm (x=15×2.4μm). Conidia
hyaline, smooth to finely roughened-walled, ellipsoid, 4–6×
2.5–4μm (x=5×3.5μm).
Culture characters: Colonies on OA at 25–28 °C covering
9 cm diam. Petri-dish in 3 wk, at first white, becoming hazel
(88) towards the centre, velvety, azonate, with diffuse margins, usually with vinaceous-buff (86) to isabelline (65) pigments diffusing beyond colonies; reverse amber (47) to grayolivaceous (107). Sporulating regions scattered over entire
central of colony, brown-vinaceous (84).
Material examined: VENEZUELA, corticated wood, 1855,
A. Fendler (K 120974, isotype) THAILAND, Chiang Mai, Doi
Suthep, on decaying wood, 10 November 2012, D.A.
Daranagama and K.D. Hyde AXL 039 (MFLU 12–0816), living culture, MFLUCC 12–0832. GenBank ITS: KM017563;
LSU: KM017565; RPB2: KM017566; Chiang Rai, Doi Mae
Saloung, on dead bamboo clumps, 12 December 2012, D.A.
Daranagama and K.D. Hyde AXL 055 (MFLU 12–0823), living cultures, MFLUCC 13–0104, ICMP.
Notes: Hypoxylon fendleri, H. trugodes and
H. crocopeplum are taxa that share similar stromatal characters. These taxa have a wide distribution in the tropics and
subtropics on a variety of host substrates. Hypoxylon fendleri
is different in having unique vinaceous tone in stromatal extractable pigment colour. Hypoxylon fendleri is similar to
H. trugodes, but differs in its orange extractable stromatal
pigments (Ju and Rogers 1996). Hypoxylon fendleri is differentiated from H. crocopeplum by its vinaceous stromatal surfaces. However, some collections of H. fendleri lack vinaceous tones and hence were categorized under
H. crocopeplum (Ju and Rogers 1996). The asexual morphs
of these two taxa are quite different. While the former produces a nodulisporium-like conidial morph, the latter has a
virgariella-like asexual morph (Ju and Rogers 1996).
Hypoxylon subcrocopeplum can also be compared with
H. fendleri. The sexual morph is similar to H. crocopeplum
in having almost similar stromatal characters, but the
perispore is indehiscent in the former species. Hypoxylon
subcrocopeplum possesses a nodulisporium-like asexual
morph, which is similar to H. fendleri. Both H. fendleri and
H. subcrocopeplum have been recorded from New Zealand
(Ju and Rogers 1996). Many of sequences from H. fendleri
deposited in GenBank are from unpublished and unverified
strains. In this study, we introduce new sequences including
RPB2 from an authentic specimen.
Fungal Diversity
Fungal Diversity
Biscogniauxia marginata (MFLU 13–0099). a Habit b, c
Concave stromatal surface in with carbonaceous tissue d Cross section
of stromata showing the perithecium and ostiole e J+ apical apparatus in
Melzer’s reagent f Mature ascus in Lactophenol Cotton Blue g Ascus in
Lactophenol Cotton Blue showing sigmoid germ slit h. Paraphyses in
water i Ascospores in water j Germinating ascospore. Scale bars: b, c=
5 mm, d=0.2 mm, e–h=20μm, i, j=15μm
Fig. 45
31. Hypoxylon lenormandii Berk. & M.A. Curtis apud
Berk., J. Linn. Soc., Bot. 10: 385. 1869.
Facesoffunginumber: FoF 00300, Fig. 50
Holotype: K 42245
Fig. 46 Biscogniauxia
marginata in MEA after 2 weeks
a From above – white colony b
From below – straw (46) c
Conidiogenous structure d
Conidiogenous cells e, f Conidia.
Scale bars: c, d=30μm, e, f=5μm
Saprobic on decorticated wood. Sexual morph
Ascostromata 0.1–1×0.04–0.1 cm diam. (x=0.6×0.08 cm),
glomerate and effuse-pulvinate, stromata almost single and
rosellinoid, connected by very thin stromatal tissue at the base,
with very conspicuous perithecial mounds, half or sometimes
entire perithecial mounds exposed, surface greyish-sepia
(106), fuscous (103), or brown-vinaceous (84), dull orange
brown to dark brown granules immediately beneath surface
and between perithecia, with KOH extractable pigments hazel
(88), fulvous (43), umber (9) or ochreous (44), the tissue below the perithecial layer inconspicuous, perithecia 0.3–
0.5 mm diam. (x=0.4 mm), immersed, sphaerical, ostioles
Fungal Diversity
Fig. 47 Fasciatispora nypae
(MFLU 15-0042) a Acus. b
Ascospores. c Germinating spore.
Scale bars: a=30μm, b=10μm
slightly papillate, without apparent disk formation. Paraphyses
not seen. Asci (120–)122–150(–153)×(5.3–)5.5–8.5(–8.7) μm
(x =134×7.1μm), 8-spored, unitunicate, cylindrical, pedicellate,
with apical ring not bluing in Melzer’s reagent, discoid, 0.2–
0.5×2–3μm broad (x=0.3×2.7μm). Ascospores (10.3–)10.5–
15(–15.4)×6–8.5μm (x=13.8×7.6μm), uniseriate, one-celled,
ellipsoid-inequilateral, with narrowly rounded ends, brown to
dark brown, with straight, to slightly sigmoid germ slit more or
less entire spore-length, perispore dehiscent in 10 % KOH,
epispore smooth. Asexual morph Conidiogenous structures
nodulisporium-like, hyaline. Conidiogenous cells 12–17×3–
3.5μm (x=15×3.3μm), hyaline, smooth. Conidia 5.5–7×2.5–
3μm (x=6.2×2.7μm), hyaline, ellipsoid.
Culture characters: Colonies on OA at 25–28 °C reaching
the edge of 4cm in 7 days, at first white, becoming hazel (88)
to greyish-sepia (106), azonate with diffuse margins, reverse
at first fuscous (103) and turning black after 10–12 days.
Sporulating regions scattered over the surface of colony particularly at the centre, hazel (88).
Material examined: CUBA, on bark, C. Wright 485 (K
42245, holotype); THAILAND, Chiang Mai, Doi Suthep,
on decaying wood, 10 November 2012, D.A. Daranagama
and K.D. Hyde AXL 104 (MFLU 12–0831), living cultures,
MFLUCC 13–0311, ICMP; Chiang Rai, Doi Mae Saloung, on
dead bamboo clumps, 12 December 2012, D.A. Daranagama
and K.D. Hyde AXL 112 (MFLU 12–0843), living culture,
MFLUCC 13–0304; Chiang Rai, Mae Fah Luang University,
on decaying wood, 20 December 2012, D.A. Daranagama and
K.D. Hyde AXL 115 (MFLU 13–0847), living cultures,
MFLUCC 13–0120. GenBank ITS: KM039135; LSU:
KM039136; RPB2: KM039137.
Notes: Hypoxylon lenormandii, with its glomerate stromata
can easily be distinguished from other species, such as
H. undulatum. Both taxa have very conspicuous stromata,
but only H. lenormandii has characteristic glomerate stromata,
more similar to the Rosellinoid type, which in fact led mycologists to name this fungus several times under Rosellinia.
Hypoxylon undulatum is characterized by the lack of KOH
extractable pigments (Ju and Rogers 1996). Therefore, our
specimen cannot be placed under H. undulatum. Hypoxylon
sublenormandii described by Suwannasai et al. (2005) from
Thailand is similar to H. lenormandii. However, the stromatal
surface of the new taxon is strongly reddish brown, while
H. lenormandii is greyish sepia, fuscous or brownvinaceous.
In addition, asci (95–110 × 3.8–5 μm) and ascospores of
H. lenormandii are longer than H. sublenormandii (Ju and
Rogers 1996, Suwannasai et al. 2005). Hypoxylon
sublenormandii is so far known only from bamboo, whereas
Fungal Diversity
Fig. 48 Hypoxylon fendleri (MFLU 12–0816) a Stromata in wood b
Surface of stromata with ostioles c KOH extractable pigments d Side
view of stromata e Cross section of stroma f Perithecia g Mature asci
and immature asci in cluster h Mature asci in water i Asci in Melzer’s
reagent with J+, discoid, apical apparatus j Mature ascospore with
dehiscent perispore in KOH H Mature ascospores in water i Mature
ascospore with straight germ slit. Scale bars: a=1 cm, b, c=1 mm, d,
e=100μm, g-k=10μm
Fungal Diversity
Fig. 49 Hypoxylon fendleri on
OA after 2 weeks a From above b
From below c, d Conidiogenous
structures e Conidia. Scale bars: c,
d=30μm, e=5μm
this specimen is from angiosperm wood. According to Ju and
Rogers (1996), H. lenormandii can occur both on dicotyledonous and monocotyledonous substrates, indicating a fairly
wide host range. According to Ju and Rogers (1996) the details in the description of H. lenormandii are compatible with
this specimen and its nodulisporium-like asexual morph. We
provide new ITS, LSU, RPB2 and β-tubulin gene sequence
data from an authenticate strain of H. lenormandii which can
be used in the future molecular analysis.
32. Hypoxylon monticulosum Mont., Syll. Gen. Sp.
Crypt., p. 214. 1856.
Facesoffunginumber: FoF 000302, Fig. 51
Holotype: BPI 589948
Saprobic on decorticated wood. Sexual morph
Ascostromata 0.3–5 × 0.5–2.5 × 0.5–1 cm (x= 3.2 × 1.8 ×
0.7 cm), pulvinate to effuse-pulvinate, with conspicuous perithecial mounds, surface rust (39) to sepia (63), blackish, shiny
when mature; with blackish, woody to carbonaceous tissue
immediately beneath surface and between perithecia, without
apparent KOH-extractable pigments; the tissue below the perithecial layer 0.2–0.3 mm thick (x= 0.2 mm), perithecia
sphaerical to obovoid, 0.2–0.5 × 0.3–0.5 mm (x= 0.3 ×
0.4 mm), immersed in carbonaceous tissue, arranged as
multi-layers, ostioles papillate, without apparent disk formation, paraphyses not seen. Asci (86–)90–130(–132)×(4.6–)5–
7(7.3) μm (x=117×6.8μm, n=20), 8-spored, unitunicate, cylindrical, pedicellate, with discoid apical ring inconspicuously
bluing in Melzer’s reagent, 0.5–1.2 × 1.5–2.5 μm (x= 1 ×
2.1μm, n=20). Ascospores (7.7–)8–12.5(–12.9)×(3.2–)3.5–
5(–5.3) μm (x=10.4×4.8μm), uniseriate, one-celled, ellipsoid-inequilateral, with narrowly rounded ends, brown to dark
brown, with sigmoid germ slit the entire spore length,
perispore dehiscent in 10 % KOH, smooth or with inconspicuous coil-like ornamentation, epispore smooth. Asexual
morph Undetermined.
Fungal Diversity
Fig. 50 Hypoxylon lenormandii (MFLU 12–0831) a Stromatal habit on
wood b Perforations and ostioles seen from above c Stroma in side view d
Cross section of the stroma showing perithecia e Perithecia from above f
Formation of pigments in KOH g Mature ascus in water i Ascus in
Melzer’s reagent showing non amyloid ascal apical apparatus j
Ascospore in water k Dehiscent perispore in KOH l Ascospore
showing germ slit. Scale bars: a-c=5 mm, d, f=30μm, g-l=10μm
Culture characters: Colonies on OA at 25–28 °C covering
Petri-dish in 3 weeks, at first white, becoming hazel (88) to
pale mouse gray (117), cinnamon (62) towards the centre,
velvety, azonate, with diffuse margins, usually with vinaceous buff (86) to isabelline (65) pigments diffusing
beyond colonies; reverse gray olivaceous (107) or
greyish-sepia (106).
Material examined: THAILAND, Chiang Mai, Doi Pui
mountain, on decaying wood, 2 December 2012, D.A.
Daranagama and K.D. Hyde AXL 111 (MFLU 12–0827),
living cultures, MFLUCC 13–0133, ICMP. Chiang Mai, Doi
Suthep mountain, on decaying wood, 12 December 2012,
D.A. Daranagama and K.D. Hyde AXL 111 (MFLU 12–
0848), living cultures, MFLUCC 12-0818. GenBank ITS:
KM052716; LSU: KM052717; RPB2: KM052719.
Notes: Hypoxylon monticulosum, H. submonticulosum and
H. investiens are quite similar in their morphological characters which resulted in some old collections of H. investiens
being named as H. monticulosum (Ju and Rogers 1996).
Besides, the variable stromatal characters in H. investiens
make it rather difficult to identify. For example,
H. investiens possesses varying ostiolar characters.
H. monticulosum and H. submonticulosum differ from
H. investiens in the lack of apparent KOH-extractable
Fungal Diversity
Fig. 51 Hypoxylon monticulosum (MFLU 12–0827) a Stromatal habit
in wood b Ostioles seen from above c Stromata in KOH (note–lack of
apparent colour formation) d Stroma in side view e Cross section of the
stroma showing perithecia f Mature and immature ascus in water g
Mature ascus in water h Ascospores in ascus showing germ slit i Ascus
in Melzer’s reagent showing inconspicuous ascal apical apparatus j
Dehiscent perispore in KOH k, l, m Ascospore in water. Scale bars: a=
2 cm, b, d, e=1 mm, f-m=10μm
pigments, while the latter has green–olivaceous KOH extracta b l e p i g m e n t s ( J u a n d R o g e r s 1 9 9 6 ) . H o w e v e r,
H. monticulosum has been reported to produce purple
stromatal pigments when young but at the maturity, the pigments are not observable. Hypoxylon monticulosum has a
blackish surface on mature stromata and even in the young
stromata which sometimes make it different from others,
coupled with Virgariella-like conidiogenous structures (Ju
and Rogers 1996). Hypoxylon monticulosum has darker,
inequilateral ascospores with dehiscent perispore in KOH as
compared to the temperate H. submonticulosum. In GenBank
there are sequences of H. monticulosum but many are from
unverified strains. We introduce new ITS, RPB2, LSU and βtubulin gene sequences from an authentic strain.
Fungal Diversity
Dothideomycetes
Recent outlines for the Dothideomycetes were provided by
Hyde et al. (2013) and Wijayawardene et al (2014). Below
we introduce 44 new species in the families Anteagloniaceae,
Bambusicolaceae, Capnodiaceae, Didymellaceae,
Didymosphaeriaceae, Hysteriaceae, Lentitheciaceae,
Leptosphaeriaceae, Lophiostomataceae,
Lophiostremataceae, Melanommataceae,
M y c o s p h a e re l l a c e a e , P a r a d i c t y o a r t h r i n i a c e a e ,
Phaeosphaeriaceae and Pleosporaceae.
Anteagloniaceae K.D. Hyde & A. Mapook., Fungal
Diversity 63: 33 (2013)
Anteagloniaceae was introduced by Hyde et al. (2013) to
accommodate a monotypic genus Anteaglonium, which had
previously been placed in Hysteriaceae because of its
hysterothecial ascomata (Boehm 2009; Mugambi and
Huhndorf 2009; Hyde et al. 2013). Phylogenetic evidence distinguishes Anteaglonium from taxa of Hysteriales (Mugambi
and Huhndorf 2009; Schoch et al. 2009; Zhang et al. 2012a;
Hyde et al. 2013). The phylogenetic tree is presented in Fig. 52.
33. Flammeascoma Phookamsak & K.D. Hyde, gen. nov.
Index Fungorum number: IF550925, Facesoffungi
number: FoF 00426
Etymology: The generic epithet Flammeascoma refers to
the reddish-orange pigmented produced at the ascomata apex.
Type species: Flammeascoma bambusae Phookamsak &
K.D. Hyde
Saprobic on submerged bamboo. Sexual morph
Ascostromata solitary or gregarious, semi-immersed to superficial, dark at the basal region and orange brown at the apex of
the ascoma, uni- to bi-loculate, conical to lenticular, with a
Fig. 52 Phylogram generated
from Maximum likelihood
(RAxML) analysis based on
combined LSU, SSU and TEF1
sequence data of Anteagloniaceae
and Lophiotremataceae.
Bootstrap support values for
maximum likelihood (ML) and
maximum parsimony (MP)
greater than 50 % are indicated
above and below the nodes, and
branches with Bayesian posterior
probabilities greater than 0.95 are
given in bold. The ex-types (exepitypes, reference strains) are in
bold; the new isolates are in blue.
The tree is rooted withSalsuginea
ramicola KT 2597.1
flattened base, coriaceous, ostiole central, with pore-like opening. Peridium thick-walled, of unequal thickness, poorly developed at the base, composed of several layers of brown to
dark brown cells, with host cells plus fungal tissue, organised
in a textura epidermoidea. Hamathecium comprising 1–
1.8μm wide, dense, narrow, cellular pseudoparaphyses, embedded in a hyaline gelatinous matrix. Asci 8-spored,
bitunicate, fissitunicate, clavate to cylindric-clavate, short
pedicellate with foot-like pedicel, apically rounded with
well-developed ocular chamber. Ascospores overlapping 1–
2-seriate, didymosporous, fusiform, slightly curved, hyaline,
becoming brown when released from asci and geminating, 1septate, constricted at the septum, surrounded by mucilaginous sheath. Asexual morph Undetermined.
Notes: The combined analysis of LSU, SSU and TEF1
genes by maximum likelihood analysis (ML) and maximum
parsimony (MP) of 16 taxa from Anteagloniaceae and
Lophiotremaceae (Pleosporales), show three clades which
are represented by the species in Anteagloniaceae,
Lophiotremataceae and a new genus Flammeascoma
(Fig. 52). Flammeascoma forms a robust clade basal to
Anteagloniaceae, while the species in Anteagloniaceae and
Lophiotremataceae form a well-resolved clade.
Flammeascoma is monotypic genus introduced to accommodate taxa associated with submerged bamboo and characteristically has orange-brown pigments in the ascostromata.
The genus is similar to Fissuroma in having coriaceous
ascomata, trabeculate pseudoparaphyses and fusiform ascospores. However, Flammeascoma differs from Fissuroma as
the former has a pore-like, orangish, pigmented ostiole,
whereas Fissuroma has a dark, slit-like opening (Tanaka and
Harada 2005; Liu et al. 2011). Based on the phylogenetic
evidence, Flammeascoma forms a single clade basal to
Fungal Diversity
Anteagloniaceae. Flammeascoma differs from Anteaglonium
which has hysterothecial ascomata and small ascospores,
while Flammeascoma has pseudothecial ascomata with larger
ascospores (Mugambi and Huhndorf 2009; Hyde et al. 2013).
We place the Flammeascoma in Anteagloniaceae.
34. Flammeascoma bambusae Phookamsak & K.D.
Hyde, sp. nov.
Fig. 53 Flammeascoma bambusae (holotype) a Ascostromata visible
as black raised structures, orangish at the apex, raised on host surface b
Section through an ascoma c Section through peridium d Asci with
Index Fungorum number: IF550924, Facesoffungi
number: FoF 00427, Fig. 53.
Etymology: The specific epithet bambusae refers to the
host bamboo in reference to the host from which the fungus
was isolated.
Holotypus: MFLU 11–0143
Saprobic on submerged bamboo. Sexual morph
Ascostromata 400–630 μm, high, 1000–1400 μm diam.,
narrow pseudoparaphyses e–g Asci h–l Ascospores m Spore
germination. Scale bars: b=200μm, c=50μm, d–g, m=20μm, h–l=5μm
Fungal Diversity
solitary, gregarious, semi-immersed to superficial, dark at the
basal region and orange brown at the apex of the ascoma, unito bi-loculate, conical to lenticular, with a flattened base, coriaceous, ostiole central, with pore-like opening. Peridium 70–
110μm wide, thick-walled, of unequal thickness, poorly developed at the base, composed of several layers of brown to
dark brown cells, with host cells plus fungal tissue, organised
in a textura epidermoidea. Hamathecium comprising 1–
1.8μm wide, dense, narrow pseudoparaphyses, embedded in
a hyaline gelatinous matrix. Asci (125–)130–160(–
175)×(14.5–)15–17(–20) μm (x=147.7×16.6μm, n=20), 8spored, bitunicate, fissitunicate, clavate to cylindric-clavate,
short pedicellate with foot-like pedicel, apically rounded with
well-developed ocular chamber. Ascospores (35–)40–43(–
48)×7–9(–10.5) μm (x=41.9×8.7μm, n=25), overlapping
1–2-seriate, didymosporous, fusiform, slightly curved, hyaline, becoming brown when released from asci and geminating, 1-septate, constricted at the septa, swollen near the septa,
with small and large guttules, wall rough, surrounded by mucilaginous sheath. Asexual morph Undetermined.
Culture characters: Colonies on PDA slow growing, 10–
20 mm diam. after 4 weeks at 25–30 °C, dark-greenish to
black at the margin, grey to dark grey in the centre; reverse
dark-greyish to black; medium dense to dense, irregular,
slightly raised to low convex or umbonate, dull with undulate
edge, hairly to woolly, slightly wavy.
Material examined: THAILAND, Chiang Mai Province,
Chom Tong District, Doi Inthanon, on dead stem of submerged bamboo (Bambusae), 5 September 2009, R.
Phookamsak RP0013 (MFLU 11–0143, holotype); ex-type
living culture, MFLUCC 10–0551. GenBank ITS:
KP744440; SSU: KP753952.
Bambusicolaceae
Bambusicolaceae was introduced by Hyde et al. (2013) to
accommodate a monotypic genus Bambusicola D.Q. Dai &
K.D. Hyde and is typified by Bambusicola massarinia D.Q.
Dai & K.D. Hyde (Dai et al. 2012; Hyde et al. 2013). Four
new species were first introduced in this genus which their
morphological characters similar to species in Massarina
sensu lato (Hirayama et al. 2010; Dai et al. 2012). However,
Bambusicola differs from Massarina sensu lato in its asexual
morph. Bambusicola formed a distinct clade from
Massarinaceae and is closely related to Lentitheciaceae in
Pleosporales (Dai et al. 2012; Hyde et al. 2013). The phylogenetic tree is presented in Fig. 54.
35. Palmiascoma Phookamsak & K.D. Hyde, gen. nov.
Index Fungorum number: IF550926, Facesoffungi
number: FoF 00428
Etymology: The generic epithet Palmiascoma refers to the
host from which the fungus was isolated.
Type species: Palmiascoma gregariascomum Phookamsak
& K.D. Hyde
Saprobic on palms. Sexual morph Ascomata gregarious,
scattered to clusterred, semi-immersed to erumpent, visible as
raised, dark spots on the host surface, uni-loculate, conical to
subglobose or irregular, glabrous, ostiole central, with minute
papilla. Peridium composed of several layers, of brown to
dark brown, pseudoparenchymatous cells, organized in a
textura angularis. Hamathecium comprising 1–2μm wide,
dense, broad cellular, pseudoparaphyses, with distinct septa,
but constricted at the the septa, anastomosing at the apex,
embedded in a hyaline gelatinous matrix. Asci 8-spored,
bitunicate, fissitunicate, clavate, short pedicellate with rounded to obtuse pedicel, apically rounded, with a well-developed
ocular chamber. Ascospores overlapping 1–2-seriate,
didymosporous, clavate to ellipsoidal, slightly curved, pale
yellowish when young, becoming brown to dark brown at
maturity, 1-septate, echinulate, surrounded by a mucilaginous
sheath. Asexual morph Conidiomata pycnidial, solitary, immersed in agar to superficial, visible as black dots covered by
vegetative hyphae, 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, outer layers comprising 3–5 cell layers of thick-walled, dark brown to black,
organized in a textura angularis to textura prismatica, inner
layers comprising 2–3 layers of thin-walled, hyaline, organized in a textura angularis. Conidiophores arising from basal
cavity of conidiomata, mostly reduced to conidiogenous cells.
Conidiogenous cells holoblastic, phialidic, discrete,
ampulliform to cylindrical, hyaline, aseptate, smooth-walled.
Conidia solitary, one-celled, oblong to ellipsoidal, with rounded or obtuse ends, initially hyaline, becoming brown at maturity, aseptate, smooth-walled.
Notes: Palmiascoma is a monotypic genus associated with
palms and produces echinulate, didymospores. The genus is
similar to Didymosphaeria Fuckel and Verruculina Kohlm. &
Volkm.-Kohlm. due to its didymosporous, brown and
echinulate ascospores (Zhang et al. 2012a). However, they are
distinguished based on molecular data (Zhang et al. 2012a;
Hyde et al. 2013; Ariyawansa et al. 2014b). Palmiascoma differs from Bambusicola species, as Palmiascoma produces ellipsoidal, dark brown, echinulate ascospores with broad cellular
pseudoparaphyses. Bambusicola species have fusiform, hyaline, smooth-walled ascospores with narrow cellular
pseudoparaphyses (Dai et al. 2012; Hyde et al. 2013).
36. Palmiascoma gregariascomum Phookamsak & K.D.
Hyde, sp. nov.
Index Fungorum number: IF550927, Facesoffungi
number: FoF00429, Fig. 55
Etymology: The specific epithet “gregariascomum” refers
to the gregarious ascomata on the host.
Fungal Diversity
Fig. 54 Phylogram generated
from Maximum likelihood
(RAxML) analysis based on
combined LSU, SSU and RPB2
sequence data of respective
families in Pleosporales.
Maximum likelihood 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-types (ex-epitypes)
are in bold; the new isolates are in
blue. The tree is rooted with
Melanomma pulvis-pyrius CBS
124080
Holotypus: MFLU 11–0211
Saprobic on palms. Sexual morph Ascomata 130–180μm
high, 130–250μm diam., gregarious, scattered to clusterred,
semi-immersed to erumpent, visible as raised, dark spots on
the host surface, uni-loculate, conical to subglobose or irregular, glabrous, ostiole central, with minute papilla. Peridium
10–50μm wide, thin to thick walled, of unequal thickness,
slightly thick at sides, composed of several layers, of brown
to dark brown, pseudoparenchymatous cells, organized in a
textura angularis. Hamathecium comprising 1–2μm wide,
dense, broad cellular, pseudoparaphyses, with distinct septa,
but constricted at the the septa, anastomosing at the apex,
embedded in a hyaline gelatinous matrix. Asci (45–)50–60(–
67)×7–10μm (x=53×8.2μm, n=25), 8-spored, bitunicate,
fissitunicate, clavate, short pedicellate with rounded to obtuse
pedicel, apically rounded, with a well-developed ocular chamber. Ascospores 11–13×3–4μm (x=11.9×3.9μm, n= 30),
overlapping 1–2-seriate, didymosporous, clavate to ellipsoidal, slightly curved, pale yellowish when young, becoming
brown to dark brown at maturity, 1-septate, not constricted
at the septa, wall rough, echinulate, surrounded by a mucilaginous sheath. Asexual morph Coelomycetes forming on bamboo pieces on WA or immersed in agar after 12 weeks.
Conidiomata 210–325μm high, 240–420μm diam., pycnidial, solitary, immersed in agar to superficial, visible as black
dots covered by vegetative hyphae, uni- to multi-loculate, globose to subglobose, glabrous, ostiole central, with minute papilla. Conidiomata walls 28–60μm wide, thick-walled, of
equal thickness, composed of several layers of hyaline to dark
brown, pseudoparenchymatous cells, outer layers comprising
3–5 cell layers of thick-walled, dark brown to black, organized
in a textura angularis to textura prismatica, inner layers comprising 2–3 layers of thin-walled, hyaline, organized in a
textura angularis. Conidiophores arising from basal cavity
of conidiomata mostly reduced to conidiogenous cells.
Conidiogenous cells 5–12×2–4μm (x=7.9×3.1 μm, n=30),
holoblastic, phialidic, discrete, ampulliform to cylindrical, hyaline, aseptate, smooth-walled. Conidia 4–6×2–3μm (x=
4.7×2.5μm, n=30), solitary, one-celled, oblong to ellipsoidal,
with rounded or obtuse ends, initially hyaline, becoming
brown at maturity, aseptate, smooth-walled.
Culture characters: Colonies on PDA fast growing, 70–
80 mm diam. after 4 weeks at 25–30 °C, greenish-grey to grey,
forming white tufts on surface, slightly radiating; reverse
brown to dark brown at the margin, dark brown to black in
the centre; medium dense, circular, flattened to slightly raised,
Fungal Diversity
Fig. 55 Palmiascoma gregariascomum (holotype) a Appearance of
ascomata on host surface b Section through an ascoma c
Section through peridium d Pseudoparaphyses e–g Asci h–l Ascospores
m Ascospore stained in Indian ink n Conidiomata produced on bamboo
pieces on WA o Section through conidiomata p Section through pycnidial
wall q–s Conidiophores t–x Conidia. Scale bars: o=100μm, b, p=50μm,
c, d=20μm, e–g, q=10μm, h–m, r–t=5μm, u–x=2μm
Fungal Diversity
dull to rough with entire edge, fairy fluffy to velvety, slightly
radially furrowed.
Material examined: THAILAND: Chiang Rai, Muang
District, Khun Korn Waterfall, on dead frond of palm, 17
December 2010, R. Phookamsak RP0091 (MFLU 11–0211,
holotype), ex-type living culture, MFLUCC 11–0175.
GenBank ITS: KP744452; LSU: KP744495; SSU:
KP753958.
Capnodiaceae
Capnodiaceae was introduced by von Höhnel (1910); the generic type is Capnodium Mont. The family Capnodiaceae are
sooty moulds in the order Capnodiales (Batista and Ciferri
1963; Hughes 1972; Crous et al. 2009a, b; Chomnunti et al.
2011; Hyde et al. 2013). Phylogenetic trees for Capnodiaceae
have been provided by Crous et al. (2009a, b) and Chomnunti
et al. (2011). In this study, one new genus and two new species
of Capnodiaceae, Capnodium siamensis and
Fig. 56 Phylogram generated
from Maximum likelihood
(RAxML) analysis based on LSU
gene region of Capnodiales.
Maximum likelihood 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-types (ex-epitypes
and reference strains) are in bold;
the new isolates are in blue. The
tree is rooted with Elsinoe
verbenae CPC 18561
Phragmocapnias philippinensis are presented and
phylogeneticially analyzed, using a dataset as used in previous
studies. The phylogenetic tree is presented in Fig. 56
37. Chaetocapnodium Hongsanan & K.D. Hyde, gen. nov.
Index Fungorum number: IF550888, Facesoffungi
number: FoF00399
Etymology: From Greek Chaeto (chaítē) meaning hair,
capnodium is from Greek kapnōdēs meaning smoky or dark.
Type species: Chaetocapnodium siamensis Hongsanan &
K.D. Hyde
Saprobic on the surface of leaves. Superficial hyphae with
septa, constricted and dark at the septum. Sexual morph
Ascomata superficial, solitary, subglobose, dark brown to
black, with setae. Setae short, aseptate, brown. Peridium comprising two layers of cells of textura angularis, inner layer
hyaline, outer layer brown, thickened at the base.
Hamathecium lacking psuedoparaphyses. Asci 8-spored,
bitunicate, fissitunicate, cylindrical to clavate, short
Fungal Diversity
pedicellate or apedicellate. Ascospores bi to tri-seriate, oblong
to ellipsoid, 3-septate, constricted and dark at the septum,
hyaline when immature and brown at maturity, smoothwalled.
Notes: Molecular analyses (LSU gene) of
Chaetocapnodium siamensis indicate that it is separate from
other known genera in Capnodiaceae with 86 % ML and
100 % PP support. The morphology of C. siamensis is typical
of Capnodiaceae based on its saprobic habitat as sooty
moulds, subglobose to globose, ascomata forming from thalli,
with a central ostiole, and bitunicate asci. Chaetocapnodium
siamensis is most similar to Capnodium tiliae based on ascospores shape and septation. However, it is distinct from other
genera in Capnodiaceae by having short setae at upper part of
ascomata and cylindrical to clavate asci. Therefore,
Chaetocapnodium should be a new genus in Capnodiaceae,
typified by C. siamensis.
38. Chaetocapnodium siamensis Hongsanan & K.D.
Hyde, sp. nov.
Index Fungorum: IF550889, Facesoffungi number: FoF
00397; Fig. 57
Etymology: siamensis is siam + ensis from Latin meaning
“of Siam” or “Thailand”.
Holotypus: MFLU 14–0747
Saprobic on the upper surface of leaves. Superficial hyphae
4–5μm wide, septate, constricted and dark at the septum.
Sexual morph Ascomata 110–120μm diam. (x=116μm, n=
5), superficial, mostly solitary, subglobose, broad at the centre,
dark brown, with apical setae. Setae 12–17×2–4μm (x=15×
3μm, n=5), short, aseptate, brown. Peridium 20–25μm (x=
22 μm, n = 5), comprising two layers of cells of textura
angularis, inner layer hyaline, outer layer brown, thickened
at the base. Hamathecium lacking pseudoparaphyses. Asci
34–41×11–14μm (x=38×13μm, n=5), 8-spored, bitunicate,
fissitunicate, cylindrical to clavate, short pedicellate or
apedicellate. Ascospores 9–15×4–6μm (x=12×5μm, n=10),
bi to tri-seriate, oblong to ellipsoid, 3-septate, constricted and
dark at the septum, hyaline when immature and brown at
maturity, smooth-walled. Asexual morph Undetermined.
Material examined: THAILAND, Chiang Rai, Tasud, on
leaves of unidentified plant, 17 September 2013, S.
Hongsanan SOY3 (MFLU 14–0747, holotype); ex-type living culture, MFLUCC 13–0778. GenBank LSU: KP744479.
39. Phragmocapnias philippinensis Hongsanan & K.D.
Hyde, sp. nov.
Index Fungorum number: IF550890, Facesoffungi
number: FoF00398; Fig. 58
Etymology: From Latin philippinensis meaning
Philippines, because the sample was collected from
Philippines.
Holotypus: MFLU 14–0748
Saprobic on the upper surface of living leaves of
Arecaceae. Sexual morph Superficial hyphae 2–4μm wide
(x=3μm, n=10), septate, slightly constricted and dark at the
septum, pale brown to brown. Ascomata 97–104μm diam.
(x=102μm, n=10), superficial on surface of substrate, solitary
or clustered, globose to subglobose, rounded above, attached
to the basal hyphae, ostiole present at maturity, thick-walled,
brown to greenish, dark brown at the base. Setae 5–8μm diam.
(x=6μm, n=10), aseptate, developing 3–5 setae at the upper
part of ascomata, brown to dark brown, pale brown at the
apex. Peridium 13–18μm (x=16μm, n=10) comprising cell
layers of textura angularis, inner layer hyaline, outer layer
dark brown or greenish. Hamathecium lacking
pseudoparaphyses. Asci 39–48×12–18μm (x=41×15μm,
n=10), 8-spored, bitunicate, fissitunicate, cylindrical when
immature, broadly clavate at maturity, short pedicellate or
sometimes apedicellate, with small ocular chamber.
Ascospores 23–27×4–6μm (x=26×5μm, n=10), bi to tri-seriate, cylindrical to clavate, 3–4-septate, constricted at the septum, basal cell narrow and slightly long, hyaline, wall rough,
occasionally smooth-walled. Asexual morph Undetermined.
Material examined: PHILIPPINES, Laguna, Mount
Makiling, on leaves of palm (Arecaceae), February 2012,
K.D. Hyde HSA14/1 (MFLU 14–0748, holotype); ex-type
living culture, MFLUCC 12–0110, CPC 20474). GenBank
LSU: KP744503.
Notes: Phragmocapnias philippinensis is most similar to
P. betle, but differs as ascospores have long and narrow basal
cells and lack a mucilaginous sheath in P. philippinensis,
whereas ascospores are slightly rounded at the base and have
a mucilaginous sheath in P. betle (Chomnunti et al. 2011).
Molecular analyses also indicate that P. philippinensis is a
new species.
Capnodiales genera incertae sedis
40. Brunneomycosphaerella Dissanayake., J.K. Liu & K.D.
Hyde, gen. nov
Index Fungorum number: IF551058, Facesoffungi
number: FoF00565
Etymology: Brunneomycosphaerella, in reference to the
brown spores of a mycospherella-like species.
Type species: Brunneomycosphaerella laburni
Dissanayake., J. K. Liu & K. D. Hyde
Saprobic on dead branches. Sexual morph Ascostromata
solitary, scattered or gregarious, immersed to erumpent, dark
brown to black, uni- to bi-loculate, globose to sub-globose.
Peridium comprising brown to black-walled cells of textura
angularis. Hamathecium comprising septate, hyaline
pseudoparaphyses. Asci 8-spored, bitunicate, fissitunicate,
elongate, cylindrical to slightly clavate at the base, pedicellate,
rounded at apex with a canal-like structure, with an ocular
chamber. Ascospores multi-seriate, dark yellow to brown, 1-
Fungal Diversity
Fig. 57 Chaetocapnodium siamensis (holotype). a, b Specimens c
Ascoma on surface of leaves d Section through ascoma e Peridium f
Thallus under ascoma g Setae on ascoma h Upper wall of ascoma i
Ascus when immature j Ascus at maturity k, l Ascospores when
immature m–o Ascospores at maturity p Conidia of possible associated
taxa. Scale bars: c, d=100μm, f, p=20μm, e, h, i, j=10μm, g, k, l, m, n,
o=5μm
septate, deeply constricted at the septum, ellipsoidal with
broadly rounded ends, with small and large guttules,
smooth-walled, lacking a sheath. Asexual morph
Undetermined.
Notes: Phylogenetic analysis of combined SSU and LSU
gene sequence data (Fig. 59) place Brunneomycosphaerella in
Fig. 58 Phragmocapnias philippinensis (holotype). a Ascomata and
hyphae on surface of leaves b Section through ascoma c Hyphae d
Ascoma with setae when viewed in squash mount e, g Asci f, h Asci in
Melzer’s reagent i Asci in cotton blue reagent j 3-septate ascospore k 4septate ascospore in Melzer’s reagent l 3-septate ascospore in cotton blue
reagent. Scale bars: b, d=50μm, c, h, i=20μm, e, f, j, k, l=10μm, g=
100μm
Fungal Diversity
Fungal Diversity
Fig. 59 Phylogram generated
from Maximum likelihood
(RAxML) analysis based on
combined LSU and SSU gene
region sequence data of related
taxa in Capnodiales. Maximum
likelihood 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-types (ex-epitypes
and reference strains) are in bold;
the new isolates are in blue. The
tree is rooted with Botryosphaeria
dothidea AFTOL ID-946
Clade A in an unresolved group in Capnodiales with rock
inhabiting fungi (Schoch et al. 2009; Hyde et al. 2013). We
therefore introduce a new genus and place it in Capnodiales,
family incertae sedis.
Dictyosporaceae
41. Brunneomycosphaerella laburni Dissanayake., J.K.
Liu & K.D. Hyde, sp. nov.
Index Fungorum number: IF551059; Facesoffunginumber:
FoF00566; Fig. 60
Etymology: in reference to its occurrence on Laburnum.
Holotypus: MFLU 14-0808
Saprobic on dead branches of Laburnum anagyroides
Medik. Sexual morph Ascomata 372–480 μm high ×
325–554 μm diam. (x=435× 490 μm), solitary, scattered
or gregarious, immersed to erumpent through the epidermis, dark brown to black, uni- to bi-loculate, globose to
sub-globose. Peridium 10–15μm wide, comprising several layers of brown to black-walled cells of textura
angularis. Hamathecium comprising septate, hyaline
pseudoparaphyses. Asci 70–100 × 45–60 μm (x= 82 ×
50μm, n =20), 8-spored, bitunicate, fissitunicate, elongate, cylindrical to slightly clavate at the base, pedicellate, rounded at the apex with a canal-like structure,
with an ocular chamber. Ascospores 18–25 × 4–5 μm
(x= 22 ×4 μm), multi-seriate, dark yellow to brown, 1septate, deeply constricted at the septum, ellipsoidal
with broadly rounded ends, with small and large
guttules, smooth-walled, lacking a sheath. Asexual
morph Undetermined.
Material examined: ITALY, Province of Forlì-Cesena,
Bagno di Romagna, Ridracoli, Santa Sofia, on dead branches
of Laburnum anagyroides IT 1761 (Fabaceae), 10 March
2014, Erio Camporesi (MFLU 14-808, holotype). GenBank
LSU: KP703171; SSU: KP703172.
Dictyosporium
The family Dictyosporaceae (Pleosporales) will be introduced
in a publication in prep. (Boonmee et al. 2015).
The genus Dictyosporium was established by Corda (1836) to
include species with sporodochia and cheiroid conidia produced on micronematous conidiophores (Corda 1836). The
concept of this genus is clear and the species are known to
be distributed worldwide (Goh et al. 1999). The genus has
been revised by Goh et al. (1999) and a dichotomous key
was provided for species (Goh et al. 1999; Cai et al. 2003a;
Crous et al. 2009a, b; Whitton et al. 2012). Currently 43 species are accepted in Dictyosporium (Whitton et al. 2012).
Molecular phylogenetic analyses place Dictyosporium within
the Pleosporales (Tsui et al. 2006; Crous et al. 2009a, b) and a
new family, Dictyosporaceae, will be introduced by Boonmee
et al. (in prep) to accommodate this group. The phylogenetic
tree is presented in Fig. 61.
42. Dictyosporium aquaticum Abdel-Aziz, sp. nov.
Index Fungorum number: IF551060, Facesoffungi
number: FoF00552; Figs. 62 and 63
Etymology: From the Latin adjective aquaticus, in reference to the freshwater habitat of the fungus.
Holotype: CBS H-21460
Saprobic on submerged wood. Sexual mor ph
Undetermined. Asexual morph Mycelium superficial and immersed in the substrate, sub-hyaline to yellow brown, 2–3μm
thick. Sporodochia on natural substratum superficial, circular,
subglobose, elongated to irregular, covering most of the wood
surface, dark brown to black, ranging diam. Conidia born
Fungal Diversity
Fig. 60 Brunneomycosphaerella laburni (holotype). a, b Ascostromata on host substrate. c Cross section of ascostromata. d, e ascus. f, g Brown
ascospores. Scale bars: c=200μm, d–f=40μm, g=20μm
directly on mycelium or on short lateral, hyaline to yellow
brown, one-celled hyphae. Conidia 60–85 μm long, 20–
30μm wide, excluding appendage (x=69×25μm, n=50),
cheiroid when young, become cylindrical or obclavate in frontal view and cylindrical in side view, medium to dark brown,
smooth-walled, euseptate, one cell layer thick, cell rows originating from a central basal cell, rows attached along their
length, cell rows increase in length from outside to inside,
conidial cells are of the same colour and consist of 66 to 82
cells arranged in 5 or 6 rows. Conidial appendages 5 to 6
globose to sub-globose, hyaline appendages that are
surrounded by thick gelatinous sheath that spreads in water,
each appendage is connected to a row.
Culture characters: Colonies on potato dextrose agar at
25 °C 15–20 mm radius after 15 days, white mycelium with
dense yellow reverse, and yellow droplets on mycelium and
Fungal Diversity
Fig. 61 Phylogram generated
from Maximum Parsimony
analysis based on combined ITS,
SSU and LSU sequence data of
cheirosporous and dictyosporious
taxa. Parsimony bootstrap support
values greater than 70 % are
indicated above and below the
nodes, and branches with
Bayesian posterior probabilities
greater than 0.95 are given in
bold. The ex-types (reference
strains) are in bold; the new
isolates are in blue. The tree is
rooted with Letendraea
helminthicola CBS 884.85
sporodochia, sporulating after about 3 weeks. Sporodochial
and conidial dimensions are similar to those recorded on natural wood.
Material examined: EGYPT, Sohag, River Nile, on submerged decaying date palm, 14 August 2014, F.A. AbdelAziz (CBS H-21460, holotype); ex-type living culture,
A136. GenBank ITS: KM610236.
Notes: Dictyosporium aquaticum is distinguished from the
other species described under the genus by the rounded hyaline appendages attached to the apical cell of each conidial cell
row. Based on the key provided by Cai et al. (2003b) and
Crous et al. (2009a, b), D. aquatica is morphologically similar
to D. bulbosum and D. strelitziae in that the three species have
rounded apical appendages. The latter two species, however, have smaller conidia and two apical appendages (Goh
et al. 1999; Crous et al. 2009a; b). Phylogenetically,
D. aquatica is closest to D. stellatum (conidia 95–140×
30–40μm; appendages absent), but has smaller conidia than
the latter species.
43. Dictyosporium meiosporum S. Boonmee & K.D.
Hyde, sp. nov.
Index Fungorum number: IF550896, Facesoffungi
number: FoF00409, Figs. 64 and 65
Etymology: meiosporum, meaning the sexual spore reproduction of this species
Holotypus: MFLU 10–0064
Saprobic on decaying wood. Sexual morph: Ascomata
281–346μm high×287.5–388μm diam., uniloculate, superficial, solitary or scattered, globose to subglobose, dark brown
to black,with ostiole, collapsing when dry. Peridium 20–
28μm thick, membracenous, composed of cells of textura
angularis, with compressed, dark brown inner layers, and
black outer layers. Hamathecium comprising numerous cellular, hyaline pseudoparaphyses. Asci 83–135.5×13–17μm (x=
101.5×14μm, n=20), 8-spored, bitunicate, fissitunicate, clavate to cylindrical or saccate, pedicel 7–19μm long, apically
rounded. Ascospores 31–39×6–8.5μm (x=35×7μm, n=20),
2–seriate, fusiform, elongated-ellipsoid, clavate, 1-septate,
Fungal Diversity
Fig. 62 Dictyosporium
aquaticum (holotype) a
Sporodochia on wood b–e
Variously shaped conidia with
apical hyaline appendages Scale
bars: a=150μm, b=20μm, c, d=
10μm, e=5μm
slightly curved, constricted at the septa, hyaline, surrounded
by a thin mucilaginous sheath, smooth-walled. Asexual
morph: hyphomycetous species. Conidiophores 14–30μm
long, 2–3 μm wide (x= 20×2.5 μm), septate, light brown,
smooth-walled. Conidiogenous cells holoblastic, terminal, integrated, dark brown. Conidia 17–27.5×6–8.5μm (x=22×
7μm, n=20), dark brown, solitary, cheirosporous, with 4 contiguous rows of cells often apically incurved at maturity, each
row 1–5-transverse septate, constricted at the septa, smoothwalled.
Culture characters: Ascospores germinating on MEA
within 36 h. Colonies on MEA, slow growing, attaining ca
Fig. 63 Dictyosporium aquaticum (ex-type culture) a Young conidia at different stages of development b, c variously shaped conidia with apical
hyaline appendages produced in single spored pure culture. Scale bars: a-c=8μm
Fungal Diversity
Fig. 64 Sexual morph of Dictyosporium meiosporum (holotype) a, b Ascomata occurring superficial on substrate c Section through of ascoma d
Peridium e. Pseudoparaphyses f–h Asci i–l Ascospores. Scale bars: a=500μm, b, c=200μm, d=20μm, e, i-l=10μm, f-h=50μm
4 mm diam. in 7 days at 28 °C. Mycelium superficial, primary
white to yellow, later becomes dark grey to brown due to
development of hyphae and conidiophores.
Material examined: THAILAND, Chiang Rai, Khun Korn
waterfall, Chiang Rai, N19°51–54′ E 99°35.39′, 671 msl., on
decaying wood of unidentified trees, 13 November 2009, S.
Boonmee KK07 (MFLU 10–0064, holotype); ex-type living
culture, MFLUCC 10-0131, BCC 52296, IFRD 2188.
GenBank ITS: KP710944; LSU: KP710945; SSU: KP710946.
Notes: Dictyosporium meiosporum is introduced as a new
species based on its sexual characteristics and hand-like conidia. Based on sexual morphology, we considered this fungus
might be related to Lophiostomataceae or Melanommataceae
in Pleosporales (Mugambi and Huhndorf 2009; Zhang et al.
2009a, b, c, 2012a, b; Hyde et al. 2013). Dictyosporium
meiosporum is characterized by superficial black, ascomata,
with an apical ostiole; bitunicate, cylindric-clavate asci and
fusiform, elongate-ellipsoid, 1-septate, hyaline ascospores,
Fungal Diversity
Fig. 65 Asexual morph of Dictyosporium meiosporum (ex-type
culture) a Ascospore germination b Colonies culture on MEA c
Vegetative hyphae d–i Conidiophores j–o Conidia with branches apical,
dark brown, and several-septate. Note up to 4-conidium on
conidiogenous locus. Scale bars: a, f, g-o = 10 μm, b = 10 mm, c =
500μm, d, e=5μm
Fungal Diversity
surrounded by a thin mucilaginous sheath. Additionally, the
isolates from this taxon produced an asexual morph in culture
(Fig. 65). The new species, forms asexual cheirosporous conidia which differ from those of D. elegans Corda and other
Dictyosporium species (Goh et al. 1999; Tsui et al. 2006;
Crous et al. 2009a, b; Kirschner et al. 2013), and the two
new species introduced in this paper. Molecular data
(Fig. 61) confirms that this fungus belongs to Dictyosporium
but as a distinct species.
45. Dictyosporium thailandicum D’ souza, D.J. Bhat &
K.D. Hyde, sp. nov.
Index Fungorum number: IF551063, Facesoffungi
number: FoF 00394, Fig. 66
Etymology: The specific epithet thailandicum refers to
Thailand the country of origin. Holotypus: MFLU 14–0240
Saprobic on submerged wood in flowing stream.
Mycelium partly superficial, partly immersed, composed of
smooth, septate, branched, light brown 1.5–3.2μm wide, hyphae. Asexual morph Sporodochia 350–500μm wide, black,
s c a t t e r e d . C o n i d i o p h o re s 6 . 8 – 11 × 4 . 3 – 1 0 . 6 μ m ,
micronematous, mononematous, cylindrical, light brown,
smooth-walled. Conidiogenous cells 4.2–5×5–5.7μm terminal, integrated. Conidia 15.4–34.5×14.5–20.6μm (x=30.6×
19μm, n=50), acrogenous, solitary, dictyoseptate, cheiroid,
smooth-walled, complanate, light brown when young, turning
dark brown to black at maturity, with 28–32 cells per conidium and 6–9 cells per row, consisting of 5 rows of cells; outer
rows shorter than inner rows, with 6–8 cells, with two hyaline,
smooth, tubular, elongated appendages in the above half,
20.6–26×5.1–6.1μm.Sexual morph Undetermined.
Culture characters: Conidia germinate on water agar (WA)
within 24 h. Colonies on PDA, slow growing, covering 9 cm
Petri-dish in 2 months at 24–28 °C, yellow to pale brown,
sporulating in 45 days.
Material examined: THAILAND. Chiang Rai Province,
Mae Fah Luang University stream, on submerged wood, 15
September 2013, M. D’souza, MJD-2 (MFLU 14–0240,
holotype); ex-type living culture, MFLUCC 13-0773.
GenBank ITS: KP716706; LSU: KP716707.
Notes: Morphologically D. thailandicum is comparable to
D. alatum Emden, D. bulbosum Tzean & J.L. Chen,
D. elegans Corda and D. strelitziae Crous & A.R. Wood
(Table 1). Dictyosporium thailandicum is very similar to
D. alatum in having the same-sized conidia, appendages,
shape and rows of cells, but differs in the number of cells
per row. Dictyosporium alatum has 4–6 cells in a row, whereas
D. thailandicum has 6–9. Further, the rows of cells in
D. al atum m a y va ry fr om 4– 7 a n d , i n c o n t r a s t ,
D. thailandicum has consistently 5 rows of cells except when
in culture; D. thailandicum may occasionally possess 3 or 4
rows of cells. In our phylogenetic analysis, D. thailandicum
was nested in a clade together with D. alatum, D. bulbosum,
D. elegans and D. strelitziae with a strong bootstrap support.
Dictyosporium thailandicum is phylogenetically close to
D. strelitziae, but morphologically this species differs from
D. thailandicum in having longer conidia, and a larger number
of cells in the conidium.
Fig. 66 Dictyosporium thailandicum (holotype) a, b Condiomata on the substrate c-f conidia. Scale bars: a=500μm, b=100μm, c-f=20μm
Fungal Diversity
Table 1
Synopsis of Dictyosporium species discussed in the paper
Species
Conidial size In μm
Conidial
shape
Rows of
cells
Total cells in
conidium
Cells per
row
Special features
D. strelitziae
D. bulbosum
D. elegans
D. alatum
30–55×20–25
27–46×11–30
50–80×24–31
(22)-26–32×15–21
Complanate
Complanate
Complanate
Complanate
(4)-5–(6)
(2)-5–(6)
5–6
(4)-5–(7)
45–56
12–48
51–96
26–37
6–11
6–10
9–13
4–6
Apical cells of outside rows with appendages
Swollen apical appendages
2 apical elongate appendages
D. thailandicum
28–38×15–21
Complanate
5
28–32
6–9
2 hyaline elongated appendages
Didymellaceae
The family Didymellaceae was introduced by de Gruyter et al.
(2009) in order to accommodate the type species Didymella
exigua, including Phoma or phoma-like genera. Zhang et al.
(2009) included Didymellaceae in the order Pleosporales
within the suborder Pleosporineae. In the most recent study
of Hyde et al (2013) included Ascochyta, Boeremia,
Chaetasbolisia, Dactuliochaeta, Didymella, Epicoccum,
Leptosphaerulina, Macroventuria, Microsphaeropsis,
Monascostroma, Phoma, Piggotia and Pithomyces in this
family based on morphology and phylogenetic analysis.
Ascochyta and Phoma are well known pathogens that are significantly important in quarantine (Kaiser et al. 2008;
Aveskamp et al. 2010). A backbone tree is presented in
Fig. 67.
45. Didymella cirsii Mapook, Camporesi & K.D. Hyde, sp.
nov.
Index Fungorum number: IF550935, Facesoffungi
number: FoF00441; Fig. 68
Etymology: Name reflects the host genus Cirsium, from
which this species was isolated.
Holotype: MFLU 14–0023
Saprobic on a dead stem of Cirsium sp., noticeable as black
dots on host surface. Sexual morph Ascomata (118–)165–
209(–243)×(121–)200–245μm (x=193×209μm, n=10), immersed or erumpent, solitary or scattered, globose to
subglobose, brown to dark brown, without subiculum covering host, Ostiole central. Peridium (15–)30–34μm wide,
thick, dark brown, comprising dark brown cells of textura
angularis. Hamathecium comprising 1–1.5μm wide, cylindrical to filiform, pseudoparaphyses. Asci (60–)67–88(–109)×
13–16 μm (x= 85 × 15 μm, n = 10), 8-spored, bitunicate,
fissitunicate, cylindric-clavate, slightly curved, smoothwalled, apically rounded, short pedicellate, with an ocular
chamber (ca. 4–6μm wide). Ascospores 20–26×6–7μm (x=
23×7μm, n=20), 1–2-seriate, overlapping in the ascus, ellipsoid to obovoid, hyaline, 1-septate, constricted at septa, widest
at the centre and tapering toward narrow ends, straight or
slightly curved, thick and smooth-walled. Asexual morph
Undetermined.
Culture characters: Ascospores germinating on MEA
within 24 hr and germ tubes produced from both ends.
Colonies growing on MEA, reaching 20 mm diam. after 5 days
at 16–18 °C, flat, margin crenate, white to olivaceous grey,
fine mycelium, medium dense to dense, septate, hyaline.
Material examined: ITALY, province of Forlì-Cesena [FC],
Fiumicello - Premilcuore, on dead stems of Cirsium sp., 5
May 2013, E. Camporesi IT1218 (MFLU 14–0023,
holotype), ex-type living culture, MFLUCC 13–0687.
GenBank LSU: KP744483; SSU: KP753951.
Notes: Didymella cirsii resembles species of Didymella
(Didymellaceae) based on its immersed, erumpent ascomata
with one-celled ascospores. It differs from Didymella
applanata in the size of ascomata, asci and ascospores.
Phylogenetic analysis of combined ITS and LSU genes indicated that Didymella cirsii belongs in Didymellaceae with
high bootstrap support (Fig. 67), but as a distinct lineage; thus
a new species is proposed.
46. Microsphaeropsis olivacea (Bonord.) Höhn.,
Hedwigia 59: 267 (1917)
Index Fungorum number: IF 438686, Facesoffungi
number: FoF00400; Figs. 69 and 70
Saprobic on Sarothamnus scoparius twig. Sexual morph
Ascomata 390–400×398–405μm (x=395× 400μm, n=5), superficial on the wood, densely crowded, in roundish to oval
clusters, breaking through the bark, pyroid to globose, with
flattened base, ostiole central and short, basally with some
brown, thick-walled. Peridium 40–45μm (n=5), black, distinctly rough, thick, multi-layered, composed of cells of
textura globulosa to angularis with red brown and smooth
walls. Hamathecium comprising numerous, filiform, thinwalled, septate pseudoparaphyses, longer than the asci. Asci
105–142×12–14μm (x=130×13μm, n=5), 8-spored, cylindrical, bitunicate, thick-walled, apically rounded. Ascospores
25–29×10–12μm (x=26×10μm, n=5), muriform, ellipsoid,
straight, both part of the spore +/-equal in size, end cells conical, immature spores hyaline, mature spores golden-brownish, septa constricted only in the centre, all stages without a
gelatinous sheath. Asexual morph Coelomycetes species.
Pycnidia 144 – 160μm (n=2), globose, white when immature,
pigmented gradually with age, brown when mature, with
Fungal Diversity
Fig. 67 Phylogram generated from Maximum likelihood (RAxML)
analysis based on combined LSU and ITS sequence data of
Didymellaceae. Maximum likelihood 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-types (reference strains) are in bold; the new isolates are in blue. The
tree is rooted with Ascochytahordei var. Hordei CBS 544.74
indistinct apical ostioles: Peridium 24–34μm (n=2), yellowish brown, of cells of textura angularis. Conidia 5.8–7(4) μm
globose, smooth-walled, hyaline initially.
Culture characters: Ascospores germinating on PDA
within 30 h and germ tubes produced from sides of
the ascospores. Colonies growing on MEA, reaching
5 mm in 7 days at 15 °C, mycelium partly superficial,
partly immersed, slightly effuse.
Material examined: GERMANY, on attached corticated
Sarothamnus scoparius twig, 21 December 2013, R.K.
Schumacher (MFLU 14–0819), living cultures, MFLUCC
14–0507. GenBank ITS: KR025859; LSU: KR025863.
Fungal Diversity
Fig. 68 Didymella cirsii (holotype) a Ascomata b Section through of ascoma c Ostiole d Peridium e Pseudoparaphyses f-h Immature and mature asci i-l
Ascospores M Germinating ascospores. Scale bars: a=200μm, b, f-h=50μm, c-e=20μm, i-l, m=10μm
Notes: The genus is similar to Coniothyrium but is differs
rentiated by the mode of conidiogenous which is phialidic in
Microsphaeropsis and holoblastic in Coniothyrium (Sutton
1980). Microsphaeropsis species produce ornamented conidia and are distinguishable from one another by conidial
size and morphology. In the phylogenetic tree our sexual
morph collection clustered with Microsphaeropsis olivacea
and thus this is the first report of the sexual morph of this
species.
47. Phoma medicaginis Malbr. & Roum., Fungi Selecti
Galliaei Exs. 37: no. 3675 (1886)
Fungal Diversity
Fig. 69 Microsphaeriopsis
olivacea (MFLU 14–0819) a, b
Ascomata c Section through of
ascoma d Peridium e Setae f
Pseudoparaphyses g–j Immature
and mature asci. k–o Immature
and mature ascospores p
Germinating ascospore. Scale
bars: a=500μm, b=200μm, c=
100μm, d=50μm, e-j=30μm, k–
o=10μm, p=20μm
Index Fungorum number: IF 169294, Facesoffungi
number: FoF00423; Fig. 71
Saprobic on Scabiosa sp., forming numerous, conspicuous,
rounded to oval, dark brown, conidomata. Sexual morph
Undetermined. Asexual morph coelomycetous.
Conidiomata 100–150μm high, 150–250μm diam., pycnidial, solitary, separate, scattered or gregarious, globose, dark
brown, immersed to semi-immersed, unilocular, thin-walled,
with a single, papillate, centrally located ostiole. Peridium
composed of 3–4 wall-layers, 15–30μm wide, with outer 1–
Fungal Diversity
Fig. 70 Microsphaeriopsis olivaceae - asexual morph a. Germinating
ascospore b. Culture from above c. Culture from below d. Induced the
growth of asexual morph e. Conidiomata on toothpick f. Close up of
conidiomata g. Section through conidioma h. Ostiole with immature
conidiospores i Conidioma j Peridium k, l Arrangement of hyphae m, n
Conidia. Scale bars: a=20μm, g=50μm, h-l=20μm, m=5μm, n=10μm
2-layers dark brown, inner 1–2-layers hyaline, with thinwalled cells textura angularis. Conidiophores reduced to
conidiogenous cells arising from the innermost wall-layer
cells of the conidiomata. Conidiogenous cells 4–7μm long×
Fungal Diversity
Fig. 71 Phoma medicaginis (MFLU 14–0812) a Specimen. b, c Black
conidiomata on the host surface. d Vertical section of conidioma. e
Ostiole. f, g Section of peridium. h–l. Conidiogenous cells and
developing conidia. m, n Germinated spore. o, p Conidia. q, r Culture
on PDA (note r; reverse). Scale bars: b=500μm, c=100μm, d=50μm,
e–g=20μm, h–k=5μm, m–n=10μm, o–p=5μm, q–r=25 mm
Fungal Diversity
5–8μm wide, enteroblastic, phialidic, determinate, doliiform
and hyaline. Conidia 10–20×3.5–5.5μm wide (x=15.5×5,
n = 20), ellipsoidal to ovoid, hyaline, straight or slightly
curved, 1-septate, constricted at the septum, obtuse at both
ends, thin-walled, smooth-walled, guttulate.
Culture characters: Colonies on PDA slow growing,
reaching 25 mm diam. after one month at 25–30 °C, circular,
grey to black, white at the edge, flattened with dense, filamentous, aerial, fluffy hyphae; reverse black in the middle, white
at the edge, without any diffusible pigments.
Material examined: ITALY, Province of Ravenna [RA],
Zattaglia, on dead twig of Scabiosa sp. (Caprifoliaceae), 30
December 2012, E. Camporesi IT-988 (MFLU 14–0812), living culture, MFLUCC 13–0485, ICMP 20794. GenBank ITS:
KP711359; LSU: KP711364; SSU: KP711369; ibid. (KUN!
HKAS 83971).
Notes: Phoma medicaginis was introduced by Malbranche
& Roumeguère (1886). This species is cosmopolitan with a
worldwide distribution and occurs on various hosts such as
Brassica oleracea L., Medicago sativa Lam., Cicer arietinum
L., Glycine soja sensu auct., Lathyrus odoratus L. and
Medicago sativa L. (Sutton 1980; Aveskamp et al. 2010).
Phoma medicaginis is an economically important pathogen,
and most likely a species complex comprising morphologically indistinguishable, but genetically and biologically isolated
species (Ellwood et al. 2006; Aveskamp et al. 2008, 2010). In
addition, it has been described as an endophyte in the roots of
Taxus globosa (Rivera-Orduña et al. 2011). The species is
characterized by globose, large, pycnidia, with straight or
slightly irregular, cylindrical conidia, often becoming 1septate and variably guttulate (Sutton 1980). Our isolate
(MFLUCC 13–0485) is morphologically similar with Ph.
medicaginis, and the only distinguish morphological character
is the dimension of conidia, but it should be noted that this
character has proven not be very reliable in coelomycetes
(Verkley et al. 2014). Phylogenetic analyses based on the
combination of LSU and ITS sequence data coupled with
morphological characters showed that the strain (MFLUCC
13–0485) is conspecific with Phoma medicaginis. Hence,
we provided here a description of this species for further
reference.
Didymosphaeriaceae
Munk (1953) introduced the family Didymosphaeriaceae typified by the genus Didymosphaeria. T he fam ily
Didymosphaeriaceae includes saprobes, endophytes and
pathogens associated with a wide variety of substrates worldwide. This family was restudied by Ariyawansa et al. (2014)
and Montagnulaceae was treated as a synonym. Ariyawansa
et al. (2014) included Alloconiothyrium, Bimuria,
Deniquelata, Didymocrea, Didymosphaeria, Kalmusia,
Karstenula, Letendraea, Montagnula, Neokalmusia,
Paraconiothyrium, Paraphaeosphaeria, Phaeodothis and
Tremateia in Didymosphaeriaceae based on both morphology
and phylogeny. A backbone tree for the family was provided
and this is followed here. The phylogenetic tree is presented in
Fig. 72.
48. Kalmusia italica Thambugala, Camporesi & K.D.
Hyde, sp. nov.
Index Fungorum number: IF550915, Facesoffungi
number: FoF00388, Figs. 73 and 74
Etymology: in reference to its occurrence in Italy.
Holotype: MFLU 14–0620
Saprobic on branches of Spartium junceum. Sexual morph
Ascomata (140–)200–300μm high×150–280μm diam. (x=
214×198μm, n=10), gregarious or grouped, immersed to
semi-immersed, dark brown to black, uniloculate, globose to
subglobose, medium to large sized and coriaceous, fusing
with the host tissue, with a central ostiole. Peridium 10–
20μm wide, comprising a few layers of dark brown to lightly
pigmented cells of textura angularis. Hamathecium comprising 1–2 μm wide, dense, cellular, hyaline, aseptate,
pseudoparaphyses, anastomosing above the asci, embedded
in a gelatinous matrix. Asci 90–135×14–20μm (x=107.5×
17.7 μm, n = 15), 4–8-spored, bitunicate, fissitunicate,
cylindro-clavate to clavate, pedicellate, apically rounded with
a small ocular chamber. Ascospores 21–27×6–8.6μm (x=
23 × 7.2 μm, n = 25), overlapping 1–2-seriate, fusoid to
fusoid-ellipsoidal, with broadly to narrowly rounded ends,
pale brown to dark yellowish-brown, 3-septate, strongly constricted at the septum, with verruculose ascospore wall
surrounded by mucilaginous sheath. Asexual morph
Pycnidia solitary, gregarious or grouped, superficial on
PDA, ostiolate. Pycnidial wall up to 50–90μm wide, comprising several layers of pseudoparenchymatous, cells of textura
angularis and textura prismatica, the outer layer composed of
thick-walled, dark brown cells, lighter towards the inner layers
of hyaline cells. Conidiogenous cells 3.4–6.2×2–3.5μm (x=
4.8×3.1μm, n=20), short cylindric, conidiogenous holoblastic, hyaline, smooth. Conidia 1–1.5 × 2–3.4 μm (x= 1.2 ×
2.9μm, n=25), oblong, ellipsoid-cylindric, aseptate, hyaline
to lightly pigmented, smooth-walled.
Culture characters: Ascospores germinating on PDA within 18 h and producing germ tubes from both ends. Colonies
growing slowly on PDA MEA, reaching a diam. of 30 mm
after 9 d at 25 °C, flat, surface smooth to velvety, with entire
edge, white, fairly dense, filamentous; reverse white to pale
white, non-pigmented.
Material examined: ITALY, province of Forlì-Cesena [FC],
Montebello, Modigliana, on dead branch of Spartium
junceum L. (Fabaceae), 14 October 2012, E. Camporesi, IT
827 (MFLU 14–0620, holotype), ex-type living cultures,
MFLUCC 13–0066. GenBank ITS: KP325440; LSU:
KP325441; SSU: KP325442; ibid. (PDD, isotype).
Fungal Diversity
Fungal Diversity
Phylogram generated from Maximum likelihood analysis based
on combined SSU, LSU, ß- tubulin and ITS sequence data of
Didymosphaeriaceae. Maximum likelihood bootstrap support values
greater than 50 % are indicated above and below the nodes, and
branches with Bayesian posterior probabilities greater than 0.95 are given
in bold. The ex-types (reference strains) are in bold; the new isolates are in
blue. The tree is rooted with Massarina eburnea CBS 473.64
Fig. 72
Notes: Kalmusia was introduced by Niessl (1872) based on
K. ebuli Niessl and is characterized by immersed to erumpent
ascomata, filiform, delicate, septate pseudoparaphyses,
bitunicate, clavate asci with narrowly ovoid to clavate, pale
brown, 3-septate ascospores. Kalmusia has been reported to
have coniothyrium-like, Cytoplea, Microsphaeropsis and
Fig. 73 Kalmusia italica
(holotype) a, b Ascomata on host
surface c Section through
ascomata d Peridium e Immature
ascus f, g Mature bitunicate asci
(note the pseudoparaphyses in g)
h–j Ascospores k Germinating
ascospore l Ascospore stained
with Indian ink. Scale bars: c=
200μm, d=50μm, e-g=25μm, hl=10μm
Paraconiothyrium asexual morphs (Zhang et al. 2012a;
Ariyawansa et al. 2014b; Zhang et al. 2014). Kalmusia italica
is distinct from K. ebuli in having aseptate pseudoparaphyses,
asci with short pedicel and a mucilaginous sheath.
49. Kalmusia spartii Wanasinghe, Camporesi, E.B.G.
Jones & K.D. Hyde, sp. nov.
Index Fungorum Number: IF550895, Facesoffungi
number: FoF00385; Figs. 75 and 76
Etymology: Named after the host genus from which it was
collected, Spartium
Holotype: MFLU 14–0751
Saprobic on dead herbaceous branches. Sexual morph
Ascomata 350–450μm high×250–400μm diam. (x=395.3×
Fungal Diversity
Fig. 74 Kalmusia italica Asexual morph (ex-type culture). a, b Pycnidia on MEA c Section of pycnidia d Pycnidial wall e, f=Conidiogenous cells and
developing conidia g Conidia. Scale bars: c=200μm, d=100μm, e-f=15μm, g=10μm
323.7μm, n=10), solitary, scattered, immersed to erumpent,
globose or subglobose, coriaceous, wall black, with or without
papilla, ostiolate. Ostiole 150–200μm high, 90–130μm diam.
(x=174.3×109μm, n=10), blackish-brown, smooth, ostiolar
canal filled with hyaline cells. Peridium 15–30μm wide at the
base, 25–50μm wide at the sides, thick, 1-layered, composed
of small heavily pigmented thick-walled cells of textura
angularis. Hamathecium comprising numerous, 1.9μm (n=
30) wide, filamentous, branched septate, pseudoparaphyses.
Asci 110–130 × 10–16 μm (x= 119.6 × 13.9 μm, n = 40), 8spored, bitunicate, fissitunicate, clavate, with a long, narrow,
furcate pedicel, up to 30–35 μm long, with a minute ocular
chamber. Ascospores 19–22×5.5–9μm (x=20.2×7.6μm, n=
50), overlapping 1–2-seriate, narrowly ovoid to clavate, 3distoseptate, constricted at the septa, initially hyaline, becoming yellowish-brown at maturity, narrowly rounded at both
ends, smooth-walled, not surrounded by a mucilaginous
sheath. Asexual morph Conidiomata 300–550μm diam.,
200–250μm high, superficial or immersed in the agar, dark
brown to black, clothed with white hyphal projections, simple
cavities. Conidiomatal wall composed of a 40–50μm wide
outer layer and 10–20μm wide inner layer of cells of textura
angularis. Conidiogenous cells 5–8×2–4μm discrete, assembled into protruding masses, or integrated in very compact
conidiophores. Conidia 2.5–4×1.5–2.5μm (x=3.2×1.9μm,
n=20) narrowly ellipsoidal or short-cylindrical, straight or
slightly curved, rounded at both ends, one-celled, with one
or two small, polar guttules, initially hyaline, becoming light
brown, smooth-walled.
Culture characters: Colonies on PDA reaching a diam. of
30–35 mm in 21 d, flat, with undulate to lobate margin, hyaline, covered by thin, felty, white, aerial mycelium, honeyyellow in reverse, sporulation after 8 weeks.
Material examined: ITALY, Forlì-Cesena Province:
Castello di Corniolo, Santa Sofia, dead and hanging branches
of Spartium junceum L. (Fabaceae), 15 March 2013, E.
Camporesi (MFLU 14–0751, holotype); ex-type living culture, MFLUCC 14–0560. GenBank ITS: KP744441; LSU:
KP744487; SSU: KP753953.
Notes: Multi-gene phylogenetic analyses indicated that
Kalmusia spartii belongs to Didymosphaeriaceae and
grouped together with other Kalmusia strains. Kalmusia
spartii has a similar morphology to K. ebuli, but in our phylogenetic tree, K. ebuli and K. spartii were well resolved
(Fig. 72).
51. Montagnula graminicola Chethana, Thambugala,
Camporesi & K.D. Hyde sp. nov.
Index Fungorum number: IF550763, Facesoffungi
number: FoF00379; Fig. 77
Etymology: The specific epithet graminicola was given
after the host family Graminae in which the fungus was
collected.
Holotypus: MFLU 14–0622
Fungal Diversity
Fig. 75 Kalmusia spartii (holotype) a Ascomata on host substrate b Section of ascoma c Close up of ostiole d Peridium e Pseudoparaphyses f–i Asci j–
n Ascospores. Scale bars: b=100μm, c=50μm, d–i=20μm, j–n=10μm
Fungal Diversity
Fig. 76 Asexual morph of Kalmusia spartii (ex-type culture) a, b Colonies on PDA (b from below) c conidiomata d Longitudinal sections of
conidiomata e Conidiomatal wall f Conidia. Scale bars: c=100μm, d=50μm, e=20μm, f=5μm
Saprobic on dead stem of grass. Sexual morph Ascomata
37–117.22μm diam. (x= 56.67 μm, n = 20), pseudothecia,
mostly solitary, semi-immersed to erumpent, black, globose
to subglobose, coriaceous, with a minutely papillate ostiole.
Peridium 14.9μm thick at side walls, up to 16μm thick near
the apex and 9μm thick at the base, consisting of a 3–4 layers
of cells, outer 3–4 layers of thick-walled, dark brown cells of
textura angularis and 2–3 layers of hyaline to pale brown
inner cells. Hamathecium comprising 2–3μm wide, filiform,
hyaline, septate, pseudoparaphyses, anastomosing above the
asci, embedded in a gelatinous matrix. Asci (45–)50–132(–
137) × (6–)8–13(–15) μm (x= 81.3 × 10.10 μm, n = 10), 8spored, bitunicate, fissitunicate, cylindrical to clavate, long
pedicellate and apically rounded with a small ocular chamber
distinct at immature asci. Ascospores (7.8–)9.8–13(–
15)×(2.8–)3.8–5.5(–6.5) μm (x=11.3×4.9μm, n=25), partially overlapping, biseriate, ellipsoidal, tapering towards the
ends, brown, two-celled, septate median, slightly constricted
at the septum, wall verruculose, surrounded by a mucilaginous
sheath. Asexual morph Undetermined.
Culture characters: Ascospores germinated on water agar
within 48 h with 3–4μm diam. germ tubes. Colonies slow
growing on PDA, attaining 6 mm diam. after 14 days at
28 °C, entire edged, white to pale white, dense, cottony mycelium on the surface and reverse pale white colour mycelium.
Material examined: ITALY, Province of Forlì-Cesena [FC],
Montebello, Modigliana, on dead stem of grass (Graminae),
27 May 2013, E. Camporesi (MFLU 14–0622, holotype); extype living cultures, MFLUCC 13–0352. GenBank ITS:
KM658314; LSU: KM658315; SSU: KM658316.
Notes: Montagnula was established by Berlese (1896),
with M. infernalis and M. gigantea based on the presence of
hyphal stromatic tissue over the ascomata and long pedicellate
asci. Montagnula infernalis was selected as the lectotype species and the genus was characterized by immersed to
erumpent, globose to subglobose ascomata, bitunicate,
cylindro-clavate to clavate, pedicellate asci and reddishbrown to dark yellowish-brown, muriform or
phragmosporous ascospores (Ariyawansa et al. 2014b).
Leuchtmann (1984) and Aptroot (1995) included some
phragmosporous and didymosporous species, making it heterogenous (Hyde et al. 2013; Ariyawansa et al. 2014). Recent
phylogenetic analyses have shown the robust clustering of
M. opulenta with Bimuria, Didymocrea, Letendrea,
Paraphaeosphaeria, Didymosphaeria,
Fig. 77 Montagnula graminicola (holotype) a–c Ascomata submersed
in the host tissue d Section of the ascoma e Section of the peridium cells f
Pseudoparaphyses g Immature asci h–i Mature asci j Fissitunicate nature
of the ascus k–p Brown ascospores with clear sheath. Scale bars: a=
200μm, b, c, d=100μm, e=5μm, f=15μm, g–j=20μm, k–p=5μm
Fungal Diversity
Fungal Diversity
Pseudocamarosporium, Paracamarosporium,
Paraconiothyrium, Tremateia, Deniquelata, Neokalmusia,
Phaeodothis, Alloconiothyrium and Kalmusia forming the
family clade (Zhang et al. 2009a; Hyde et al. 2013;
Ariyawansa et al. 2014b; Wijayawardene et al. 2014).
Ariyawansa et al. (2014) synonymized Montagnulaceae under the older family name Didymosphaeriaceae.
In the current study, a new species, Montagnula
graminicola is introduced in the genus Montagnula based on
the morphological and phylogenetic evidence. Our new species Montagnula graminicola is distinct from M. infernalis in
having smaller ascomata, and smaller, ellipsoidal, one-celled
ascospores with a sheath. Montagnula graminicola resembles M. opulenta, but differs in having smaller
ascomata, and spores with verruculose wall. The phylogenetic analysis provides strong evidence that
M. graminicola belongs in the genus Montagnula
(Fig. 72), where it forms a sister clade to Montagnula
o p u l e n t a ( C B S 1 6 8 . 3 4 , A F TO L - I D 1 7 3 4 ) a n d
Montagnula aloes (CPC 19671) with high bootstrap
support; thus a new species is proposed.
51. Paraconiothyrium nelloi W.J. Li, Camporesi & K.D.
Hyde, sp. nov.
Index Fungorum number: IF550918, Facesoffungi
number: FoF00422; Fig. 78
Etymology: Named after Camporesi Nello, who collected
the sample from which the species was isolated.
Holotypus: MFLU 14–0813
Saprobic on dead stem of Spartium junceum L. Sexual
morph Undetermined. Asexual morph coelomycetous.
Conidiomata 250–350μm high, 200–300μm diam., pycnidial, solitary, immersed, globose to obpyriform, unilocular, centrally ostiolate, thick-walled. Peridium 15–25μm wide, 4–5layered, composed of outer 3–4-layers brown and inner 1–2layers hyaline, thin-walled cells of textura angularis.
Conidiophores reduced to conidiogenous cells, arising from
the base and sides of the conidioma. Conidiogenous cells 3.5–
10μm long×5–10μm wide, enteroblastic, phialidic, determinate, ampulliform, lining the inner wall layer of the pycnidium, hyaline, smooth. Conidia 6.5–8.5×5–6μm (x=7.5×5.5;
n=10), globose to obovate, thick-walled, smooth-walled, onecelled, hyaline when young, becoming dark brown at
maturity.
Culture characters: Culture on PDA slow growing,
reaching 10–15 mm diam. after one week, circular, yellowish
in the centre, with whitened edge, after one month, sparse,
aerial, filamentous, no pigments produced.
Material examined: ITALY, Province of Forlì-Cesena [FC],
Premilcuore, Fiumicello, on dead twig of Spartium junceum
L. (Fabaceae), 13 January 2013, E. Camporesi IT-1008
(MFLU 14–0813, holotype), ex-type living culture,
MFLUCC 13–0487, ICMP 20796. GenBank ITS:
KP711360; LSU: KP711365; SSU: KP711370; ibid. (KUN!
HKAS 83972, isotype).
Notes: Paraconiothyrium was introduced by Verkley et al.
(2004) to accommodate four species, namely Parac.
estuarinum (type specie), Parac. Brasiliense, Parac.
cyclothyrioides and Parac. fungicola. Subsequently, the genus
was expanded to include four more species, viz. Parac.
africanum, Parac. babiogorense, Parac. hawaiiense and
Parac. variabile (Damm et al. 2008; Budziszewska et al.
2011). Presently, Paraconiothyrium comprises 21 species
(de-Gruyter et al. 2013; Ariyawansa et al. 2014b; Verkley et
al. 2014), including Parac. nelloi and the sexual species
Parac. thysanolaenicola described in this paper. The morphological characters of Paraconiothyrium are variable. The
conidiomata can be eustromatic to pycnidial, the
conidiogenous cells are phialidic or annelidic, and the conidia
smooth-walled or minutely warted and hyaline to brown at
later stages of development (Verkley et al. 2004; Gruyter et
al. 2013). The description of Parac. nelloi fits well with this
generic concept, and Parac. nelloi shares similarities with
Parac. fuckelii in having pycnidial conidiomata with a single
ostiole, and subglobose to ellipsoid or obovoid conidia
(Gruyter et al. 2013; Verkley et al. 2014). Parac. nelloi differs
from Parac. fuckelii by the conidiogenous cells, which are
phialidic in Parac. nelloi and annelidic in Parac. fuckelii.
Combined phylogenetic analyses of ITS, SSU and LSU sequence data (Fig. 72) show that Parac. nelloi is distinct from
any other species of Paraconiothyrium. Based on the morphological characters together with molecular sequence data,
Parac. nelloi is introduced as a new species.
52. Paraconiothyrium thysanolaenae Phookamsak,
Chethana & K.D. Hyde, sp. nov.
Index Fungorum number: IF550930, Facesoffungi
number: FoF00432; Fig. 79
Etymology: The specific epithet thysanolaenicola refers to
the host from which the fungus was isolated.
Holotypus: MFLU 11–0142
Saprobic on stems of Thysanolaena maxima. Sexual
morph Ascostromata 115–250μm high, 410–840μm diam.,
scattered or clustered, gregarious, immersed to semi-immersed, visible as raised, black structures, longitudinal axis
vertical to the host surface. Locules 130–200μm high, 170–
300μm diam., clustered, immersed under pseudoclypeus, unito multi-loculate, subglobose to quadrilateral, dark brown to
black, ostioles central in each locule, with black, minute papilla. Peridium 10–30μm wide, thin-walled, of equal thickness, composed of 3–5 layers, of brown to dark brown,
pseudoparanchymatous cells, arranged in a textura angularis.
Hamathecium of dense, 1–2μm wide, filamentous, broad cellular, pseudoparaphysese, branched, anastomosing above the
asci, embedded in a gelatinous matrix. Asci (52–)55–70(–75×
6–7(–7.5) μm (x= 65 ×7μm, n=25), 8-spored, bitunicate,
Fungal Diversity
Fig. 78 Paraconiothyrium nelloi (holotype) a Specimen b Black
conidiomata on the host surface c Section of peridium d Vertical section
of conidiomata e, f Conidiogenous cells and developing conidia g Ostiole
h Germinated spore i–k Conidia l Culture on PDA. Scale bars: b=
200μm, c=20μm, e–f=5μm, g=50μm, i–k=5μm, l=25 mm
Fungal Diversity
Fig. 79 Paraconiothyrium thysanolaenae (holotype) a Black, raised,
longitudinal ascostromata on host surface b Section of an ascoma c
Peridium d Ocular chamber e Ascospore stained in Indian ink f
fissitunicate, cylindrical, with short acute or knob-like pedicel,
apically rounded with well-developed ocular chamber.
Ascospores 10.5–12(–13)×3–4μm (x=11.7×3.9μm, n=30),
overlapping 1–2-seriate, phragmosporous, fusiform to ellipsoidal, brown to reddish-brown, 3-septate, constricted at the
septum. Asexual morph Undetermined.
Pseudoparaphyses g-h Asci i–m Ascospores n–o Culture characters
Scale bars: b=200μm, c, g, h=20μm, f=10μm, d, e, i, j, k, l, m=5μm
Fig. 80 Paraphaeosphaeria spartii (holotype). a Specimen b–c Black
conidiomata on host surface d, e Vertical section of conidioma f
Section of peridium g Ostiole h–k Conidiogenous cells and developing
conidia l Germinated spore m–n Conidia o. p Culture on PDA. Scale
bars: b=500μm, c=200μm, d=100μm, e=5μm, f=5μm, g=10μm, h–i=
5μm, j–n=5μm
Fungal Diversity
Fungal Diversity
Culture characters: Colonies on MEA 78–84 mm diam.
after 4 weeks at 25–30 °C, cream to orangish-white at the
margins, white grey in the centre; reverse white to
cream or yellowish-white at the magins, light brown to
yellowish-brown at the cantre; medium dense, circular,
flattened, smooth, with entire edge, fairly fluffy to
flocose, slightly radiating in the lower part, not producing pigments.
Material examined: THAILAND, Chiang Mai, Mae Taeng
District, Huai Nam Dung, 8 September 2009, on dead stem of
Thysanolaena maxima Kuntze (Poaceae), R. Phookamsak
RP0007 (MFLU 11–0142, holotype), ex-type living culture,
MFLUCC 10–0550, GenBank ITS: KP744453; LSU:
KP744496; SSU: KP753959.
Notes: In this study, we introduce a new sexual species in
the genus Paraconiothyrium, Parac. thysanolaenae, based on
the phylogenetic evidence. Paraconiothyrium thysanolaenae
formed a robust clade with Parac. estuarinum and Parac.
cyclothyrioides in our phylogenetic analyses. However, the
species did not produce an asexual morph; thus the asexual
morphology of these species is not comparable. Ariyawansa et
al. (2014) reported the sexual species of Parac. fuckelii and
Parac. magnolia which have clearly distinguish morphological characters as compared to Parac. thysanolaenae.
Paraconiothyrium thysanolaenae differs from P. fuckelii
and Parac. magnolia by its ascostromata and mucilaginous
sheath surrounding the ascospores. Paraconiothyrium
thysanolaenae forms pseudostromata with uni- to multilocules and ascospores lack a mucilaginous sheath, while
Parac. fuckelii and Parac. magnolia have uniloculate
ascomata.
53. Paraphaeosphaeria spartii W.J. Li, Camporesi & K.D.
Hyde, sp. nov.
Index Fungorum number: IF550917, Facesoffungi
number: FoF00421; Fig. 80
Etymology: Referring to the host, Spartium, on which the
fungus was found.
Holotypus: MFLU 14–0810
Saprobic on dead stem of Spartium junceum L. Sexual
morph Undetermined. Asexual morph coelomycetous.
Conidiomata 200–300 (x=250) μm diam., 250–300 (x=260)
μm high, pycnidial, epidermal to subepidermal, scattered, globose to subglobose, unilocular, pale brown, ostiolate,
appearing pruinose, thick-walled. Ostiole circular, papillate.
Peridium 10–20μm wide, composed of 3–4-layers, with outer
1–3-layers brown and inner 1–2-layers hyaline, thin-walled
cells of textura angularis. Conidiophores reduced to
conidiogenous cells, arising from basal layers of conidioma.
C o n i d i o g e n o u s c e l l s 2 – 6 × 3 – 6 . 5 ( x= 4 × 4 . 5) μ m ,
enteroblastic, phialidic with an inconspicuous periclinal thickening, cylindrical to subcylindrical, or subcylindrical to
ampulliform, integrated, hyaline, smooth-walled. Conidia 4–
7.5×3.5–5μm (x=6×4, n=50), subglobose to ellipsoid or obovoid, hyaline when young, becoming pigmented to pale
brown at maturity, smooth-walled, guttulate, aseptate, thinwalled.
Culture characters: Colonies on PDA slow growing,
reaching 10–20 diam. after 3 weeks, glabrous and with
colourless margin; mycelium immersed, initially
colourless, later becoming yellowish aerial mycelium,
non sporulating.
Material examined: ITALY, Province of Forlì-Cesena [FC],
Santa Sofia, Collina di Pondo. on dead stem of Spartium
junceum L. (Fabaceae), 16 October 2012, E. Camporesi IT816 (MFLU 14–0810, holotype); ex-type living culture,
MFLUCC 13–0398, ICMP 20789. GenBank ITS:
KP711357; LSU: KP711362; SSU: KP711367; ibid. (KUN!
HKAS 83969, isotype).
Notes: Morphologically, Paraph. spartii shares similarities
with Paraph. sporulosa in having subglobose to ellipsoid or
obovoid, aseptate conidia with one large and often also 1–2
additional smaller oil-droplets. Parac. spartii can be distinguished from Paraph. sporulosa by its conidiogenous cells.
Parap. spartii has subcylindrical to ampulliform, integrated,
phialidic conidiogenous cells, with an inconspicuous
periclinal thickening and collarette. Paraph. sporulosa has
globose to ampulliform, hyaline, discrete, conidiogenous cells
with 1–2 percurrent proliferations. Phylogenetically, Paraph.
spartii is also distinct from any other species within
Paraphaeosphaeria (Fig. 72)
54. Pseudocamarosporium cotinae Norphanphoun,
Bulgakov & K.D. Hyde, sp. nov.
Index Fungorum number: IF550922, Facesoffungi
uumber: FoF00492; Fig. 81
Etymology: Named after the generic name of host, Cotinus
Holotype: MFLU 14–0774
Saprobic on dead herbaceous branches of Cotinus
coggygria. Sexual morph Undetermined. Asexual morph
Conidiomata pycnidial, 400–600μm diam. 370–620μm high,
solitary to gregarious, black, sub-immersed, unilocular, with a
papillate ostiole. Pycnidial wall 50–60μm, multi–layered,
with 3–5 outer layers of brown–walled cells of textura
angularis, with inner most layer thin, hyaline.
C o n i d i o p h o re s r e d u c e d t o c o n i d i o g e n o u s c e l l s .
Conidiogenous cells blastic, phialidic, hyaline, smooth,
formed from the inner most layer of pycnidium wall.
Conidia 12–15×6–9μm (x=13.5×7.5μm, n=30) oblong to
ellipsoidal, mostly straight, infrequently slightly curved,
muriform, with 1–3 transverse septa, with 1–2 longitudinal
septa, constricted at the septa, initially hyaline, brown to dark
brown at maturity, narrowly rounded at both ends, smooth–
walled.
Culture characters: Conidia germinating on MEA within
24 h. Colonies on PDA slow growing, reaching 1 cm diam.
Fungal Diversity
Fig. 81 Pseudocamarosporium cotinae (holotype) a Habit in wood b
Conidiomata on substrate c Single conidioma d Cross section conidioma
e Peridium f Apex of conidioma g, h Conidiogenous cells with attached
conidia i Immature conidia j, k Mature conidia l Germinating conidium.
Scale bars: d=200μm, e, f=50μm, g, h=20μm, i-k=10μm , l=50μm
after 1 week at 25 °C, later producing dense mycelium, circular with rough margin, white at first, greenish-yellow after
7 days, flat or effuse on the surface, without aerial mycelium.
Hyphae septate branched, hyaline, thin.
Fungal Diversity
Material examined: RUSSIA, Rostov region, Shakhty city,
near Grushevsky pond, shelterbelt artificial forest, on dead
branches of Cotinus coggygria Scop. (Anacardiaceae), 18
March 2014, T. Bulgakov (MFLU 14 – 0774, holotype); living culture, MFLUCC 14–0624; GenBank ITS: KP744460;
LSU: KP744505; SSU: KP753964.
Notes: Wijayawardene et al. (2014) introduced
Pseudocamarosporium, with the type species P. propinquum,
from Salix vitellina, and four other species.
Pseudocamarosporium propinquum has oblong conidia. Our
collection differs from the type in having oblong to ellipsoidal
conidia which are narrowly rounded at both ends. The new
species also has larger conidiomata than other species of
Pseudocamarosporium, and also differs in sequence data
(Fig. 72). Pseudocamarosporium cotinae clusters with P.
corni (Wijayawardene et al. 2014) which provides good evidence that P. cotinae belongs in Pseudocamarosporium,
where it forms a well-resolved clade. Hence we introduce it
as a new species.
Hysteriaceae
The family Hysteriaceae was introduced by Chevallier (1826)
to accommodate Dothideomycete species producing
hysterothecial ascomata in Hysteriales (Boehm 2009; Boehm
et al. 2009a, b; Hyde et al. 2013). The family has been
monographed by Zogg (1962) and included six genera viz.
Hysterium, Hysterographium, Gloniopsis, Gloniella, Glonium,
Farlowiella and Hysterocarina with Hysterium as the type genus (Boehm et al. 2009b, c; Eriksson 2006). Actidiographium
was added in Hysteriaceae (Boehm et al. 2009a; Hyde et al.
2013). Based on the phylogenetic analyses, Boehm et al.
(2009a) removed three genera viz. Glonium, Hysterographium
and Farlowiella from Hysteriaceae (Hyde et al. 2013). Some
species previously accommodated in Hysterographium have
been transferred to the new genera Hysterobrevium and
Oedohysterium in Hysteriaceae, whereas the genus
Farlowiella was tentatively placed in the Dothideomycetes,
genera incertae sedis (Hyde et al. 2013). The asexual morphs
of Hysteriaceae are coelomycetes and hyphomycetes (Lohman
1931, 1933a, b, 1934; Boehm et al. 2009b). Hysteriaceae presently composes 12 sexual genera viz. Actidiographium,
Coniosporium, Gloniella, Gloniopsis, Hysterobrevium,
Hysterocarina, Sphaeronaema, Oedohysterium, Ostreichnion,
Psiloglonium, Rhytidhysteron and Sphaeronaema (Hyde et al.
2013). The phylogenetic tree is presented in Fig. 82.
55. Psiloglonium colihuae (Lorenzo & Messuti) E.
Boehm., Stud. Mycol. 64: 71 (2009)
≡ Glonium colihuae Lorenzo & Messuti, Mycol. Res.
102(9): 1104 (1998)
Index Fungorum number: IF 444903, Facesoffungi
number: FoF00435; Fig. 83
Saprobic on stems of Thysanolaena maxima. Sexual
morph Ascomata 135–195 μm high, 590–970 μm diam.,
hysterothecial, gregarious, superficial, dark, elongate, shieldshaped, or irregular, rarely furcate, joined at ridges, with central, slit-like opening. Peridium 6–20μm wide, thin-walled, of
unequal thickness, poorly developed at the base, carbonaceous, opaque dark and glossy, composed of several cell
layers of small, opaque, dark, dull cells arranged in textura
angularis to textura globulosa. Hamathecium composed of
0.8–1.5 μm wide, dense, trabeculate, anastomosing,
pseudoparaphyses, embedded in a hyaline to brown gelatinous matrix, slightly swollen at the apex. Asci (76–)80–95(–
120)×(15–)17–18(20) μm (x=94.5×18μm, n=25), 8-spored,
bitunicate, fissitunicate, clavate, short pedicellate with obtuse
ends, apically rounded with well-developed ocular chamber.
Ascospores 30–35×4–6μm (x=32.9×5.7μm, n=30), overlapping 1–3-seriate, fusiform with acute ends, hyaline, 1-septate,
constricted at the septa, slightly curved, smooth-walled,
surrounded by distinct mucilaginous sheath. Asexual morph
Undetermined.
Culture characters: Colonies on PDA slow growing, 20–
24 mm diam. after 4 weeks at 25–30 °C, grey at the magins,
dark grey at the centre; reverse dark grey to black; dense,
irregular, raised to umbonate, dull, undulate edge with entire
magin, velvety, slightly wrinkled, producing reddish pigments
in agar.
Material examined: THAILAND, Chiang Mai, Chom
Tong District, Doi Inthanon, on dead stem of Thysanolaena
maxima Kuntze (Poaceae), 16 November 2010, R.
Phookamsak RP0094 (MFLU 11–0214, reference specimen
designated here), living culture, MFLUCC 11–0178,
GenBank ITS: KP744466; LSU: KP744511.
Notes: Psiloglonium colihuae was combined in the genus
Psiloglonium by Boehm et al. (2009a, b). The species is poorly
known and lacks sequence data in GenBank. Psiloglonium
colihuae is similar to P. sasicola in the size of the ascospores,
but they differ in having acute ends in P. colihuae. P. colihuae
has slightly larger ascospores (30–43 ×4–9.8μm) than P.
sasicola (N. Amano) E. Boehm & C.L. Schoch (25–32×5–
8μm). Our isolate has the same ascospore size range and acute
ends as the type of P. colihuae. Therefore we treat our isolate
as P. colihuae and designate our collection as a reference specimen (sensu Ariyawansa et al. 2014a). Based on multi-gene
phylogenetic analyses, P. colihuae clusters with P. araucanum
and a new species P. multiseptatum Phookamsak & K.D.
Hyde, and is phylogenetically distinct from P. sasicola.
56. Psiloglonium multiseptatum Phookamsak & K.D.
Hyde, sp. nov.
Fungal Diversity
Fig. 82 Phylogram generated
from Maximum likelihood
(RAxML) analysis based on
combined LSU, TEF and RPB2
sequence data. Maximum
likelihood 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-types (reference
strains) are in bold; the new
isolates are in blue. The tree is
rooted with Pleospora herbarum
CBS 191.86
Index Fungorum number: IF550931, Facesoffungi
number: FoF00434; Fig. 84
Etymology: The specific epithet multiseptatum refers to the
septation of the ascospores.
Holotypus: MFLU 11–0200
Saprobic on stem of Thysanolaena maxima. Sexual morph
Ascomata 130–265 μm high, 450–1000 μm diam.,
hysterothecial, solitary to gregarious, scattered, superficial,
elongate, cupulate to lenticular, or irregular, joined at the
ridges, slightly straight, with central slit-like opening.
Peridium 4–20μm wide, thin-walled, of unequal thickness,
thick at the base and sides, carbonaceous, composed of several
layers of small, opaque, dark cells, arranged in textura
angularis to textura globulosa. Hamathecium 0.5–2μm wide,
composed of dense, trabeculate, anastomosing,
pseudoparaphyses, embedded in a hyaline to brown gelatinous
matrix. Asci (86–)90–110(–118)×(19–)22–23(–27) μm (x=
104.5×22.6μm, n=25), 8-spored, bitunicate, fissitunicate, cy-
Fungal Diversity
Fig. 83 Psiloglonium colihuae (MFLU 11–0214) a Hysterothecia
black, irregular, elongate, lying on host surface. b Vertical section
through hysterothecia. c, d Section throu gh peridium. e
Pseudoparaphyses stained in congo red reagent. f Asci embedded in
pseudoparaphyses. g–j Asci. k–p Ascospores. Scale bars: b=200μm, d,
f–j=20μm, c, e, k–p=10μm
lindric-clavate to clavate, subsessile to short pedicellate with
furcate pedicel, apically rounded, with indistinct ocular chamber, easily broken in external layer. Ascospores (38.5–)46–
51(–53) × (6–)7–8(–9) μm (x= 47.6 × 7.6 μm, n = 30),
overlapping 2–4-seriate, elongated fusiform, with acute ends,
hyaline, 5–7-septate, constricted at the septum, easily separated
in to part ascospores, smooth-walled, surrounded by distinct
mucilaginous sheath. Asexual morph Undetermined.
Fungal Diversity
Fig. 84 Psiloglonium multiseptatum (holotype) a Hysterothecia on host surface b Vertical section through a hysterothecium c Peridium d Asci
embedded in pseudoparaphyses e–h Asci. i–m Ascospores Scale bars: b=100μm, c–h=20μm, i–m=10μm
Fungal Diversity
Culture characters: Colonies on PDA slow growing,
10–15 mm diam. after 4 weeks at 25–30 °C, dark grey
to dark greenish at the magins, yellowish in centre; reverse dark grey to black; dense, circular to slightly irregular, raised to low umbonate, dull, undulate edge with
entire margin, felty, with hard small granules, standing or
tufts of yellowish hyphae, producing dark brown pigments
in agar.
Material examined: THAILAND, Chiang Mai Province,
Chom Tong District, Doi Inthanon, on dead stem of
Thysanolaena maxima Kuntze (Poaceae), 6 October 2010,
R. Phookamsak RP0080 (MFLU 11–0200, holotype), ex-
Fig. 85 Psiloglonium sasicola (MFLU 11–0159) a Black elongated
ascomata lying on host surface b Section through ascoma c Asci
embedded in pseudoparaphyses d Pseudoparaphyses stained by cotton
blue e, f Asci g–j Ascospores k Ascospores stained in Indian ink l–m
Culture characteristic. Scale bars: b=100μm, c, e, f, k=20μm, d, g-j=
10μm
Fungal Diversity
type living culture, MFLUCC 11–0164; GenBank LSU:
KP744512; SSU: KP753969.
Notes: Psiloglonium multiseptatum is introduced to accommodate a gloniella-like species which is similar to Gloniella
bambusae Zogg and G. normandina Rehm in having transseptate ascospores, but differs in the size range. Psiloglonium
multiseptatum has larger ascospores (46–51×7–8μm) than
Gloniella bambusae H. Zogg (30–35 × 5–8 μm) and G.
normandina Rehm (22–25×3–3.5μm) (Rehm 1912; Boehm
2009). Based on multi-gene phylogenetic analyses,
Psiloglonium multiseptatum groups in a clade with P.
araucanum and is placed in Psiloglonium and not Gloniella.
57. Psiloglonium sasicola (N. Amano) E. Boehm & C.L.
Schoch
≡ Glonium sasicola N. Amano, Trans. Mycol. Soc. Japan
24(3): 287 (1983)
Index Fungorum number: IF515322, Facesoffungi
number: FoF00433; Fig. 85
Saprobic on bamboo. Sexual morph Ascomata 130–
180μm high, 165–210μm wide, hysterothecial, solitary to
gregarious, scattered, superficial, dark, short to elongate, or
irregular, rarely bifurcate, with central, slit-like opening.
Peridium 20–40μm wide, thick-walled, of unequal thickness,
poorly developed at the base, slightly thick at sides, carbonaceous, opaque dark and glossy. Hamathecium 0.5–1.2μm
wide, composed of dense, trabeculate pseudoparaphyses,
anastomosing, embedded in a hyaline to pale brown gelatinous matrix, slightly swollen at the apex. Asci (74–)80–105(–
115) × 13–15 μm (x= 92 × 14.3 μm, n = 25), 8-spored,
bitunicate, fissitunicate, cylindrical to cylindric-clavate, short
pedicellate with knob-like pedicel, apically rounded with
well-developed ocular chamber, easily broken in external layer. Ascospores (25–)27–30×6–7(–8) μm (x=28.3×6.8μm,
n=30), overlapping 1–2-seriate, fusiform, with slightly roung
ends, hyaline, 1-septate, constricted at the septa, individaully
upper cell larger than lower cell, smooth-walled, with small
and large guttules, surrounded by wide mucilaginous sheath.
Asexual morph Undetermined.
Culture characters: Colonies on MEA slow growing,
reaching 30–35 mm diam. after 4 weeks at 25–30 °C, white
to cream, or pale yellowish at the magins, pale yellowish at the
centre, distinguished from the margin by white
embossed hyphae with grey tufts in the centre; slightly
radiating; dense, irregular, slightly raised to umbonate,
dull with undulate edge, velvety to floccose, zonate in
the centre at the lower part, reverse white cream at the
margin, yellowish at the centre.
Material examined: THAILAND, Chiang Rai Province,
Phan District, Poo Kaeng Waterfall, on dead branch of bamboo (Bambusae), 23 May 2010, R. Phookamsak RP0039
(MFLU 11–0159, reference specimen designated here), living culture, MFLUCC 10–0565; GenBank: ITS: KP744467;
LSU: KP744513.
Notes: Psiloglonium sasicola was transferred to
Psiloglonium by Boehm et al. (2009a, b) due to its
didymosporous, fusiform hyaline, 1-septate ascospores with
rounded ends, which are surrounded by gelatinous sheath
(Boehm 2009; Boehm et al. 2009a, b). Psiloglonium sasicola
is similar to P. uspallatense (Speg.) E. Boehm & C.L. Schoch
and P. ephedrae (Henn.) E. Boehm & C.L. Schoch, but differs
in the size of ascospores (Boehm 2009). Our collection is
similar to P. sasicola and is from the same host bamboo.
This isolate is treated as P. sasicola and is designated as a
reference specimen (sensu Ariyawansa et al. 2014a). P.
sasicola forms a robust clade in the phylogenetic tree with P.
simulans (Fig. 82)
Jahnulales
58. Aliquandostipite manochii Sri-indr., Boonyuen,
Suetrong, K.L. Pang & E.B.G. Jones, sp. nov.
Index Fungorum number: IF550752, Facesoffungi
number: FoF00493; Fig. 86
Etymology: “manochii” – in honour to Prof. Dr. Leka
Manoch as her family name, a Thai distinguished mycologist
and a co-chair, IMC10 steering committee, The 10th
International Mycological Congress (IMC 2014).
Holotype: BBH 38816
Saprobic on dead wood. Sexual morph Ascomata 250–
310 high×290–320μm wide, globose to subglobose, gregarious, immersed, superficial with stalk (up to 500μm long and
25–40μm wide) or erumpent, shedding wood particles and
becoming superficial with base remaining immersed, dark
brown to black, coriaceous, ostiolate. Stalk septate 2.5–
7.5μm wide. Peridium Undetermined. Hamathecium comprising ca 2.5–7.5μm diam., hypha-like, filamentous, hyaline,
septate, pseudoparaphyses, unbranched between the asci,
branching and anastomosing above. Asci 125–200 × 20–
55μm, 8–spored, obclavate, short pedicellate, bitunicate,
fissitunicate, with an ocular chamber and a faint ring.
Ascospores 50–80×10–17.5μm, brown, biseriate, 1-septate,
light brown, guttulate, ellipsoid–fusiform, minutely
verruculose, constricted at the septum, surrounded by a wide
mucilaginous sheath. Asexual morph Undetermined.
Culture characters: Ascospores did not germinate on
media
Material examined: THAILAND, Nakhon Ratchasima,
Khao Yai National Park, Wang Champi Waterfall (Latitude:
14.7117, Longitude: 101.4117, 757 meters above sea level),
on submerged wood of an unidentified tree, 3 April 2003, V.
Sri-indrasutdhi, N. Boonyuen & S. Sivichai, BBH 38816
(holotype).
Fungal Diversity
Fungal Diversity
Aliquandostipite manochii (holotype). a-c Immersed and
superficial ascomata with septate stalk marked by arrow (S) in the
substratum d-f Asci which are 8–spored, obclavate, pedicellate,
bitunicate, fissitunicate, with an ocular chamber and a faint ring with
pseudoparaphyses marked by arrow (P) g-m Ascospores with a broad
mucilaginous sheath marked by arrow (M), brown, biseriate, 1-septate
and ellipsoid-fusiform. Scale bars a=200μm, b-c=100μm, d=50μm, ef=20μm, g=15μm, h-m=30μm
Fig. 86
Notes: A genus in Aliquandostipitaceae (Jahnulales,
Dothideomycetes) based on morphological observations as
the species did not germinate on standard media. Our new
species is morphologically similar to species of Jahnula; however, morphological features of A. manochii agree with species of Aliquandostipite as proposed by Inderbitzin et al.
(2001) and Suetrong et al. (2011). It is characterized by the
dimorphic ascomata, with a superficial stalk, and the arrangement of spores in the asci, and ascospores with a mucilaginous
sheath or pad. Morphologically A. manochii is most similar to
A. khaoyaiensis in having ascospores with a broad gelatinous
sheath, but differs in having a branched and anastomosing
hamathecium above of the asci, lacking arcicular crystals
within the spores, and in ascospore size (Raja et al. 2005;
Raja and Shearer 2007).
Lentitheciaceae
Lentitheciaceae was introduced by Zhang et al. (2012a, b)
to accommodate massarina-like species in the suborder
Massarineae (Zhang et al. 2009a, b, c, 2012a, b; Hyde et al.
2013). Lentitheciaceae recently included Katumotoa,
Keissleriella Tingoldiago, Setoseptoria and Lentithecium
and is typified by Lentithecium fluviatile (Zhang et al.
2012a, b; Hyde et al. 2013; Quaedvlieg et al 2013;
Fig. 87 Phylogram generated
from Maximum likelihood
(RAxML) analysis based on
combined LSU and SSU
sequence data of Lentitheciaceae.
Maximum likelihood 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-types (reference
strains) are in bold; the new
isolates are in blue. The tree is
rooted with Massarina eburnea
CBS 473.64
Wijayawardene et al. 2014). Lentitheciaceae was reported as
being saprobic on herbaceous and woody plants and producing coelomycetous asexual morphs. Setoseptoria was introduced for stagonospora-like or dendrophoma-like taxa
(Quaedvlieg et al 2013). The phylogenetic tree is presented
in Fig. 87.
60. Keissleriella sparticola Singtripop & K.D. Hyde, sp.
nov.
Index Fungorum number: IF551011, Facesoffungi
number: FoF00481, Fig. 88
Etymology: The species epithet refers to “Spartium sp.” the
host genus.
Holotypus: MFLU 14–0116
Saprobic on a dead branch of Spartium junceum. Sexual
morph Ascomata 134–259μm high × 130–260μm diam. (x=
180×182μm, n=5), immersed in the host tissue, globose to
subglobose, dark brown to black, round, dimidiate, ostiole
central. Ostiole 47–84μm high×69–88μm diam. (x=60×
81μm, n=5), papillate, dark brown, smooth, with long, dark
brown setae, with about 28–79μm. Peridium 27–44μm wide,
thick, comprising 5–6 layers, outer layer comprising heavily
pigmented, thick-walled, brown to dark brown cells of textura
angularis, inner layer composed of hyaline, thin-walled cells
of textura angularis. Hamathecium comprising numerous,
1.7–3.2μm wide, filamentous, occasionally branched, septate,
hyaline, pseudoparaphyses. Asci 104–123×17–23μm (x=
116×19μm, n=7), 8-spored, bitunicate, broadly cylindrical,
smooth-walled, with an ocular chamber. Ascospores 24–
29μm high×9–11μm diam. (x=27×10μm, n=10), 2–4-overlapping seriate, oval to ellipsoid, with 2 layers, upper part
Fungal Diversity
Fig. 88 Keissleriella sparticola (holotype) a Appearance of ascomata on
host surface b Section of ascoma on host c Peridium d Vertical section
through ascoma e Close up of periphyses f Pseudoparaphyses g Immature
ascus h–j Mature asci k–n Mature ascospores o Germinating ascospore.
Scale bars: a=1000μm, b, d, e=200μm, c=100μm, g–j=50μm, f, o=
20μm, k–n=10μm
Globose to subglobose, immersed, Brown to dark brown Cylindrical 104–123×17–23μm Oval to ellipsoid, hyaline to pale This study
134–259×130–260μm
setae 28–79μm long
yellow, 4–5-septate, with both
cross and longitudinal septa,
24–29×9–11μm
K. cladophila On dry twigs and branches of Aesculus Globose to sub-globose,
Dark brown setae
Cylindrical to clavate
Ellipsoid, hyaline, 1–3-septate, Corbaz 1957
spp., Berberis spp., Genista spp.,
immersed 300–450 diam.
30–45×3–4.5μm
70–125×9–12μm
9–16×4–6μm
Salix spp., Sarcothamnus spp.,
Smilax spp.,
Hyaline to brown, 1-septate
Müll & von
K. genistae
On dry twig of Genista spp.,
No data
No data
18–21×5–6μm
Arx 1962
Sarcothamnus spp. (Leguminosae)
Brown to dark brown Cylindrical 117–124×14–19μm Ellipsoid to fusiform, hyaline,
Dennis 1964
K. linearis
On dead stems of Phragmites
Globose to subglobose,
setae 34–48μm long
1–3-septate, 32–42×8–10μm
australis (Poaceae)
immersed or erumpent,
584–762×369–495μm
Lentithecium Submerged wood of Fraxinus
Immersed, 130-160×240–320 μm Absent
Cylindrical
Broadly fusiform, hyaline, 1–3 Ying Zhang
aquaticum
excelsior (Oleaceae)
130-190 μm×17-23 μm
septate, 25–30×8–12 μm
et al. 2009
On dead branch of Spartium junceum
Keissleriella
sparticola
Ascomata
Fungus name Host
Table 2
Synopsis comparing Keissleriella sparticola and some related species
Setae
Asci
Ascospore
Original
reference
Fungal Diversity
wider than the lower part, 4–5-septate, with 1–3 vertical septa,
hyaline, becoming yellowish at maturity. Asexual morph
Undetermined.
Culture characters: Colonies on PDA slow growing,
reaching 0.5–1 cm diam. after 1 week at 18 °C, later with
dense mycelium, circular, margin smooth, white at first, pinkish-ash after 4 weeks, Hyphae thin, smooth-walled, branched
septate, hyaline. Reverse similar.
Material examined: ITALY, Ravenna Province, Monticino,
Brisighella, on dead branch of Spartium junceum L.
(Fabaceae), 18 September 2013, E. Camporesi IT 990
(MFLU 14–0116, holotype). ex-type culture, MFLUCC 14–
0196; GenBank LSU: KP639571.
Notes: Keissleriella was introduced by Höhnel (1919) and
is typified by K. aesculi Sacc (≡ Pyrenochaeta aesculi Höhn.).
The genus Keissleriella is characterized by black setae in the
ostiolar canal and hyaline 1 to multi-septate ascospores (Barr
1990). Based on the phylogenetic tree (Fig. 87) the new species of Keissleriella sparticola has a close relationship with
Keissleriella cladophila, Keissleriella genistae, Lentithecium
lineare and Lentithecium aquaticum. However our new species differ from these species in host, ascomata, periphyses,
asci and/or ascospore characters (Table 2).
Leptosphaeriaceae
The family Leptosphaeriaceae which belongs in the order
Pleosporales was introduced by Barr (1987) and is typified
by Leptosphaeria Ces. & De Not. (1863), while the familial
status of the Leptosphaeriaceae is also supported by molecular phylogenetic studies (Schoch et al. 2009; Zhang et al.
2009a, b, c; Hyde et al. 2013). Zhang et al. (2012a, b) included
only Leptosphaeria and Neophaeosphaeria in family
Leptosphaeriaceae, while Hyde et al. (2013) accepted
Heterospora, Leptosphaeria, Neophaeosphaeria,
Paraleptosphaeria, Plenodomus and Subplenodomus based
on combined analysis of LSU, SSU, RBP2 and TEF1 gene
data. Members of this family are saprobes, hemibiotrophs or
pathogens on plants, occurring in terrestrial and aquatic
habitats (Hyde et al. 2013). Camarosporium, Heterospora,
Plenodomus and Subplenodomus are accepted asexual
morphs of Leptosphaeriaceae (de Gruyter et al. 2009;
Hyde et al. 2013). The phylogenetic tree is presented in
Fig. 89.
61. Leptosphaeria doliolum (Pers.) Ces. & De Not.
Comm. Soc. crittog. Ital. 1(4): 234 (1863)
Facesoffunginumber: FoF 00391; Fig. 90
Saprobic on decaying stem of Urtica dioica L. Sexual
morph Ascomata 154–211μm high×160–225μm diam. (x=
177×201μm, n=6), superficial, semi-immersed at the base,
solitary, scattered, globose to subglobose, apically conical,
wider and flattened at the base, carbonaceous, glabrous, black,
Fungal Diversity
Fig. 89 Phylogram generated
from Maximum likelihood
(RAxML) analysis based on
combined LSU and ITS sequence
data of Leptosphaeriaceae.
Maximum likelihood 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-types (reference
strains) are in bold; the new
isolates are in blue. The tree is
rooted with Phoma herbarum
CBS 615.75
ostiolate, with a shiny papilla. Peridium 24–39μm, thickwalled, multi-layered, comprising cells of textura angularis;
cells thick-walled at external layers, inner layer of thin-walled
cells, surface heavily pigmented. Hamathecium comprising
numerous, 1–2μm wide, long cylindrical, cellular, branched,
septate, hyaline, pseudoparaphyses, anastomosing above the
Fungal Diversity
Fig. 90 Leptosphaeria doliolum (MFLU 14–0565). a, b Ascomata c Cross section of ascoma d Section the peridium e Pseudoparaphyses f, g Asci h-k
Ascospores l Germinating spore m, n Culture. Scale bars: a=200μm, b–c=20μm, d=5μm, e=2μm, f–g=10μm
asci. Asci 52–71×4–5μm (x=63×4μm, n=15), 8-spored,
bitunicate, fissitunicate, narrowly cylindrical, short pedicellate
or apedicellate, rounded at the apex, with a small ocular
chamber. Ascospores 15–19×2–3μm (x=18×3μm, n=15),
overlapping uni-seriate, 3-septate, straight or slightly curved,
constricted at the septa, yellowish brown to brown, guttulate,
Fungal Diversity
lacking a mucilaginous sheath, smooth-walled. Asexual
morph Undetermined.
Culture characters: Ascospores germinate on MEA at
18 °C within 24 h and germ tubes produced from both ends
of ascospores. Colonies on MEA fast growing, white-gray.
Mycelium septate, branched, hyaline to light brown, and
smooth.
Material examined: ITALY, Province of Forlì-Cesena [FC],
Monte Fumaiolo, on stems of Urtica dioica L. (Urticaceae), 3
July 2013, E. Camporesi IT661 (MFLU 14–0565), living culture, MFLUCC 13–0740. GenBank ITS: KP729444; LSU:
KP729445.
Notes: Leptosphaeria was introduced by Cesati and De
Notaris (1863) and included 26 species, with L. doliolum
(Pers.:Fr.) Ces. & De Not. selected as the lectotype species
for the genus (Shearer et al. 1990). Confusion surrounding the
type of L. doliolum has been discussed in Hyde et al. (2013).
Due to the uncertainty of the placement of this genus, several
authors have included it under different families, such as
Leptosphaeriaceae (Barr 1987; Eriksson and Hawksworth
1991) or Phaeosphaeriaceae (Eriksson and Hawksworth
1986). Recent studies based on multi-gene analysis showed
that Leptosphaeria clustered within the order Pleosporales, in
the family Leptosphaeriaceae. Species of Leptosphaeria (including the type of Leptosphaeriaceae) and
Neophaeosphaeria form a paraphyletic clade with moderate
bootstrap support (Schoch et al. 2009; Zhang et al. 2012a, b).
In our study we signify an additional collection for the type of
L. doliolum to stabilise the placement of the genus in
Leptosphaeriaceae which is morphologically and phylogenetically identical to the epitypified strain of L. doliolum (CBS
541.66).
61. Leptosphaeria ebuli Jayasiri, Camporesi & K.D. Hyde,
sp. nov.
Index Fungorum number: IF550958, Facesoffunginumber:
FoF: 00454; Fig. 91
Etymology: In reference to its occurrence on Sambucus
ebulus
Holotype: MFLU 14–0820
Saprobic on dead stem of Sambucus ebulus. Sexual
morph Ascomata 226–396×241–251μm (=300×241μm,
n=10), solitary or scattered, initially immersed, becoming
erumpent to near superficial, globose to subglobose, broadly
or narrowly conical, coriaceous, smooth-walled, ostiolate.
Ostiole usually broadly papillate, darkened at the base,
Peridium 24–26μm (=45μm, n=20) wide, comprising two
types of cells, outer cells of 1–2 layers of heavily pigmented
cells of textura angularis, inner layer composed of small,
light brown to hyaline cells of textura angularis.
Hamathecium of septate, long, hyaline, cellular
pseudoparaphyses, embedded in gelatinous matrix between
and above the asci. Asci 80–109×8–9μm (=100×9μm, n=
20), 8-spored, bitunicate, fissitunicate, clavate to sub-cylindrical, with a short, broad pedicel, thickened and rounded at
apex with a distinct ocular chamber surrounded by a large,
distinct, apical ring. Ascospores 23–28×4–5μm (x=25×
5.8μm, n=40), biseriate to overlapping tri-seriate, ellipsoidal
with broadly rounded ends, hyaline to light brown when
immature, becoming brown to chestnut brown when mature,
3-septate, smooth-walled, lacking a sheath. Asexual morph
Undetermined.
Culture characters: Ascospores germinated on MEA at
18 °C within 24 h and germ tubes were produced from both
ends of ascospores. Colonies on MEA were growing fast
(within one week 2 cm diam. colony), white-gray.
Material examined: ITALY, Province of Monte Fumaiolo
(FC), on stems of Sambucus ebulus L. (Adoxaceae), 23 July
2014, E. Camporesi (MFLU 14–0820, holotype); ex-type living cultures, MFLUCC 14–0828; GenBank ITS: KP744446;
LSU: KP744488; SSU: KP753954.
Notes: Notes: Multi-gene phylogenetic analysis (Fig. 89)
indicated that Leptosphaeria ebuli forms a distinct clade sister
to Leptosphaeria sydowii clade (BS support 96 %) whichis
known as an asexual morph (de Gruyter et al. 2012).The morphological characters of L. ebuli is fit with the generic concept
of Leptosphaeria in having superficial ascomata with a shiny
papilla, multi-layered peridium comprising cells of textura
angularis, cellular pseudoparaphyses and fissitunicate, narrowly cylindrical asci bearing multi septate, yellowish-brown
to brown ascospores but differ from the generic type of L.
doliolum in the shape of the ascomata (globose to subglobose
versus conical), width of the ostiole (narrow versus broad)
ostiole pore and the size of the ascospores (25× 5.8μm versus
18 × 3 μm). Hence we introduced Leptosphaeria ebuli in
Leptosphaeria as new species based on both molecular data
coupled with morphology.
62. Paraleptosphaeria nitschkei (Rehm ex G. Winter) DeGruyter et al. Stud. Mycol.75: 20 (2012)
Facesoffunginumber: FoF00442; Figs. 92, 93
Saprobic on dead stems of Petasites sp. Sexual morph
Ascomata (296–)328–362 × (406–)420–423 μm (x= 329 ×
416μm, n=5), superficial, solitary or scattered, subglobose
to obpyriform, dark brown, without subiculum covering host.
Ostiolar neck protruding. Peridium (38–)45–56(–64) μm
wide, brown to dark brown, comprising dark brown cells of
textura angularis. Hamathecium comprising 0.5–1μm wide,
cylindrical to filiform, pseudoparaphyses. Asci (80–)97–102(–
107)×9–10μm (x=98×9μm, n=10), 8-spored, bitunicate, cylindric-clavate, slightly curved, smooth-walled, apically
rounded, short pedicellate, with an ocular chamber (ca. 1–
1.5μm wide). Ascospores 27–29(–34)×4.5–6μm (x=29×
5μm, n=10), 1–2-seriate, overlapping in the ascus, fusiform
to inequilateral, hyaline, 3-euseptate, constricted at medium
septum, widest at the middle and tapering towards narrow
Fungal Diversity
Fig. 91 Leptosphaeria ebuli (holotype). a, b Ascomata c Section through the ascoma d Peridium e Pseudoparaphyses f Immature ascus g, h Mature asci
j-l Ascospores. Scale bars: a=500μm, b=100μm, c=100μm, d, f–h=30μm, e=10μm, i–l=10μm
Fungal Diversity
Fungal Diversity
Paraleptosphaeria nitschkei (MFLU 14–0021) a, b Ascomata c
Cross section of ascoma d Peridium e Ostiole f Pseudoparaphyses g, h
Asci i–l Ascospores. Scale bars: a=500μm, b–c=100μm, d–e=20μm, f=
2μm, g–h=20μm, i–l=5μm
Fig. 92
ends, straight or slightly curved, thick and smooth-walled.
Asexual morph Coelomycete. Conidiomata 435–455(–
475)×(276–)300–330μm (x=455×305μm, n=5), pycnidial,
globose to subglobose, solitary, peridium 30–35(–38)
μm, thick-walled, scleroplectenchymatous.
Conidiogenous cells 3–4 × 5–6 μm, phialidic. Conidia
1.5–2× (–12)15–20 μm, hyaline, aseptate, cylindrical to
filiform. Undetermined.
Cultural characteristics: Ascospores germinating on MEA
within 48 hours at 18 °C and germ tubes produced from both
ends. Colonies growing on MEA, reaching 20 mm diam. after
7 days at room temperature, flat, margin crenated, olivaceous
brown.
Fig. 93 Paraleptosphaeria nitschkei - Asexual morph (MFLUCC 13–
0688) a, b Colonies on MEA media. c Asexual test on WA media d, e
Conidiomata forming on substrate f Section through of conidiomata. g
Material examined: ITALY, Province of Forlì-Cesena [FC],
Passo la Calla, Santa Sofia, on stems of Petasites sp.
(Asteraceae), 14 July 2012, E. Camporesi IT563 (MFLU
14–0021, reference specimen designated here), living culture, MFLUCC 13-0688; GenBank ITS: KR025860; LSU:
KR025864. ITALY, Province of Forlì-Cesena (FC), Monte
Falco, on decaying grass stems of Petasites sp., 21 May
2013, E. Camporesi IT648 (MFLU 13–0644), DNA was extracted from the fruiting body), GenBank ITS: KP729446;
LSU: KP729447.
Notes: Paraleptosphaeria is typified by Paraleptosphaeria
nitschkei (Rehm ex G. Winter) De-Gruyter et al. Munk
(1957) recognized Leptosphaeria section ParaLeptosphaeria, an invalid taxon, as a heterogeneous group.
The section was differentiated from Eu-Leptosphaeria,
which included the generic type species Leptosphaeria
doliolum. Leptosphaeria nitschkei was considered a typical
representative of section Eu-Leptosphaeria (Müller and von
Arx 1950). Gruyter et al. (2013) introduced
Peridium wall. h-i Conidiogenous cell and conidia. Scale bars: a-c=2 cm,
d-e=200μm, f=100μm, g=10μm, h-i=5μm
Fungal Diversity
Paraleptosphaeria to accommodate Leptosphaeria nitschkei
and its relatives. The putative strain of Paraleptosphaeria
nitschkei (holotype) clustered with our newly collected
strains from Italy. Paraleptosphaeria nitschkei was originally
described from Austria, whereas our specimen is from grass
stems of Petasites sp. in Italy. Even though the morphology
and the molecular data of our Italy collection show 100 %
identity to the putatively named Paraleptosphaeria nitschkei,
it would be unwise to epitypify this species with the Italy
collection as they are from different continents. We therefore
designate our collection as a reference specimen of P. nitschkei
so that further work on these taxa can be carried out.
63. Plenodomus agnitus (Desm.) Gruyter et al., Stud.
Mycol. 75: 21 (2012)
Facesoffungi number: FoF00411; Fig. 94
Basionym: Sphaeria agnita Desm., Annls Sci. Nat., Bot.,
sér. 2 15: 18 (1841)
≡Leptosphaeria agnita (Desm.) Ces. & De Not., Comm.
Soc. crittog. Ital. 1(4): 236 (1863)
Saprobic on dead stems of Eupatorium cannabinum.
Sexual morph Ascomata 137–250μm high, 235–453μm
diam., solitary, scattered, or in small groups of 1–3, semi-immersed, subglobose to ovoid, with a flattened base, smooth,
easily removed from host substrate, black. Ostiole papillate,
black. Peridium 37–107μm (x=71μm, n=10) wide, composed of three layers, externally with a 5–14μm thick layer
of dark brown to black, opaque cells, central layer composed
of hyaline, small, thick-walled cells of textura angularis, inner
layer composed of pale brown, textura prismatica.
Hamathecium comprising 1–3 μm wide, dense, hyaline,
broad, septate, cellular pseudoparaphyses, branching above
asci and embedded in gelatinous matrix. Asci 66–115×7–
10μm (x=94×9μm, n=20), 8-spored, bitunicate, cylindricsubclavate, with a short, furcate pedicel, rounded at the apex.
Ascospores 26–32×3–6μm (x=29×4μm, n=40), overlapping
uniseriate, narrowly fusoid with sharp to narrowly rounded
ends, 6–8-septate, constricted at third or fourth septum, upper
third or fourth cell enlarged, pale yellowish-ash, smoothwalled. Asexual morph Undetermined.
Material examined: GERMANY, Senckenberganlage 25,
D-60325 Frankfurt (Main), on dead stem of Eupatorium
cannabinum L. (Asteraceae), 13 November 2013, H.A.
Ariyawansa (MFLU 15-0039, reference specimen designated here). The DNA was extracted from fruiting body.
GenBank ITS: KP744459; LSU: KP744504.
Notes: Leptosphaeria agnita was transferred to Plenodomus
agnitus by de Gruyter et al. (2013) based on the sequence data.
The putative strain of Plenodomus agnitus (CBS 121.89) clustered with our new collection from Germany, on dead stem of
Eupatorium cannabinum. Saccardo (1883) originally described
Fig. 94 Plenodomus agnitus (MFLU 15-0039) a, b Ascomata on host
substrate c Vertical section of ascoma d Close up of the peridium e
Cellular pseudoparaphyses f, g Asci h Ascus in Melzer’s reagent i
Ascospores in the ascus j–l Ascospores. Scale bars: a = 1 cm, b =
500μm, c=100μm, d=50μm, e–h=20μm, i–l=10μm
Plenodomus agnitus as Leptosphaeria agnita and mentioned
the different hosts including dry stems of Eupatorium
cannabinum in France, Germany, Britain, Italy and
Netherlands. The ascomata, size of asci and ascospores
are typical of P. agnitus (Saccardo 1883) and the molecular data is identical to the putatively named CBS
121.89 strain. We therefore designate our collection as
a reference specimen (sensu Ariyawansa et al. 2014a) of
P. agnitus to stabilize the application of this name.
Lophiostomataceae
The family Lophiostomataceae was established by
Nitschke (1869) and is characterized by slit-like ostiolar
opening on a laterally compressed papilla, mostly clavate asci, 1- to several-septate and hyaline to dark
brown ascospores with terminal appendages or mucilaginous sheaths. (Chesters and Bell 1970; Holm and Holm
1988; Zhang et al. 2012a, b). The genera Lophiostoma,
Misturatosphaeria and Tumularia are presently accepted
in Lophiostomataceae (Hirayama and Tanaka 2011;
Zhang et al. 2009a, b, c, 2012a, b; Hyde et al. 2013).
The phylogenetic tree is presented in Fig. 95.
64. Lophiostoma pseudodictyosporium Qing Tian,
Camporesi & K.D. Hyde, sp. nov.
Index Fungorum number: IF550887, Facesoffungi
number: FoF: 00403; Fig. 96
Etymology: In reference to the dictyosporous conidia similarity of the taxon to Dictyosporium.
Holotypus: MFLU 14–0737
Saprobic under periderm or semi-immersed in woody plant
substrates Colonies growing on dead branch of Spartium
junceum. Conidiomata pycnidial, on the upper surface of
stem, (130–)155–180 μm high × (145–)160–195 μm diam.
(x=165×180μm, n=10), solitary, scattered, superficial, globose to subglobose, black. Peridium thin at the apex, base and
sides 12–15μm (x=13.5μm, n=7), a single layer, composed
of brown to black thin-walled cells of textura angularis.
Conidiogenous cells (2–)3–4.5μm broad (x=3.5μm, n=
10), holoblastic, integrated, smooth, brown, producing a
branched conidium at the tip, cup-shaped or doliiform.
Conidia (8–)10–15μm diam. × 12–26(–32) μm high (x=
12.5 × 23.5 μm, n = 10), uniformly medium brown,
complanate, dictyosporous conidia, regularly consisting
of 6–8 rows of cells, each rows comprising 5–7 cells,
3–3.5μm wide.
Fungal Diversity
Fungal Diversity
Fig. 95 Phylogram generated
from Maximum likelihood
(RAxML) analysis based on LSU
sequence data 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 0.95 are given in
bold. The ex-types (reference
strains) are in bold; the new
isolates are in blue. The tree is
rooted with Sporormiella minima
CBS 524.50
Culture characters: Conidia germinating on PDA within
24 h and germ tubes arising from both ends. Colonies growing
slowly on PDA, reaching a diam. of 1 cm after 30 days at
16 °C, velvety, radiating towards the edge. Mycelium initially
hyaline and light pink at the margin, after 30 days, dense,
filamentous, reverse black, pigments produced.
Material examined: ITALY, Province of Forlì-Cesena [FC],
Fiumicello, Premilcuore, on stem of Spartium junceum
(Fabaceae), 6 March 2013, E. Camporesi (MFLU 14–0737,
holotype); ex-type living culture, MFLUCC 13–0451.
GenBank ITS: KR025858; LSU: KR025862.
Notes: Lophiostoma is a morphologically well-studied genus (Barr 1990; Chesters and Bell 1970; Holm and Holm
1988; Hyde et al. 2013; Mugambi and Huhndorf 2009; Yuan
and Zhao 1994; Zhang et al. 2012a, b). Lophiostoma was
introduced by Cesati and De Notaris (1863), while Tanaka
and Harada (2003a) typified L. macrostomum which was characterized by immersed to erumpent ascomata, with a
cylindrical or crest-like papilla and slit-like ostiole; this has
been regarded as a prominent morphological character of
Lophiostoma macrostomum (Chesters and Bell 1970; Holm
and Holm 1988). Combined gene analysis of LSU and SSU
sequence data indicated Lophiostoma to be polyphyletic,
grouping into two monophyletic clades (Zhang et al. 2009a,
b, c; Suetrong et al. 2009). The asexual morph of Lophiostoma
has rarely been reported. In this study, we introduce an asexual
morph taxon, Lophiostoma pseudodictyosporium, which is
shown to be related to Lophiostoma based on analysis of
LSU gene sequence data. Lophistoma dictyosporium is morphologically similar with the species of Dictyosporium, both
having brown hand-like conidia. In Dictyosporium the conidia
form directly on mycelia, while in Lophiostoma
pseudodictyosporium the conidia form in pycnidia. In the phylogenetic analysis they clustered into different families, therefore, we place the new species, Lophiostoma
pseudodictyosporium in Lophiostoma, Lophiostomataceae.
Fungal Diversity
Fig. 96 Lophiostoma pseudodictyosporium (holotype) a Specimen
material b Black conidiomata on the host c–e Vertical section of
conidioma f–g Conidiogenous cells and developing conidia h
Germinating ascospore i–k Conidia l Colony on PDA from above.
Scale bars: a=500μm, b=100μm, c–d=50μm, e, h=20μm, f–g, i–k=
10μm
65. Lophiostoma ravennicum Tibpromma, Camporesi &
K.D. Hyde, sp. nov.
Index Fungorum number: IF550884, Facesoffungi
number: FoF00389; Fig. 97
Fungal Diversity
Fig. 97 Lophiostoma ravennicum (holotype) a, b Ascomata c Cross section of ascoma d Ostiole e Peridium f Pseudoparaphyses g–i Asci. j–l
Ascospores m Germinating ascospore. Scale bars: a = 500 μm, b = 100 μm, c = 50 μm, d-e = 10 μm, f = 2 μm, g-i = 10 μm, j-m = 5 μm
Etymology: Refers to the name of the province in Italy
where the fungus was collected
Holotype: MFLU 14–0692
Saprobic on decaying grass stems of Juncus sp. Sexual
morph Ascomata 211–282μm high×121–187μm diam. (x=
244×158μm, n=5), superficial, solitary, scattered, black, globose to subglobose, not easy to remove from host, neck long,
black. Peridium 10–27μm, 1-layered, composed of small,
light brown to dark brown, thin-walled cells of textura
angularis. Hamathecium comprising numerous, 1.1–1.9μm
wide, long cellular, septate, hyaline, pseudoparaphyses,
branching above the asci. Asci 55–70×9–11μm (x=64×
10μm, n=10), 8-spored, bitunicate, fissitunicate, narrowly cylindrical, short pedicellate or apedicellate, apically rounded,
Fungal Diversity
with an ocular chamber. Ascospores 18–21×4–6μm (x=19×
5μm, n=15), 1–2-seriate, brown, ellipsoidal-fusiform, narrowly fusoid with rounded ends, or fusiform, usually 6-septate, the cells above central septum often broader than the
lower ones, with mucilaginous sheath, smooth-walled.
Asexual morph Undetermined.
Material examined: ITALY, Ravenna Province, Marina
Romea, on stems of Juncus sp. (Juncaceae), 28 November
2013, E. Camporesi IT 1544 (MFLU 14–0692, holotype),
ex-type living culture, MFLUCC 14–0005; GenBank ITS:
KP698413; LSU: KP698414; SSU: KP698415.
Notes: The molecular data confirms that our strain groups
within the genus Lophiostoma (Fig. 95). Lophiostoma
ravennicum however, differs in having ellipsoidal-fusiform,
usually 6-septate ascospores, with thick mucilaginous sheath
and forms an individual lineage in the phylogenetic tree
(Fig. 95).
Lophiotremataceae
The family Lophiotremataceae was established by Hirayama
and Tanaka (2011) with Lophiotrema as the type for
lophiostomatoid taxa distinguished from Lophiostoma, it
forms a well-supported monophyletic group on the dendrogram (Zhang et al. 2009b; Hirayama and Tanaka 2011; Hyde
et al. 2013).
66. Lophiotrema eburnoides Kaz. Tanaka, A. Hashim. &
K. Hiray., sp. nov.
Index Fungorum number: IF551065, Facesoffunginumber:
FoF 00495; Fig. 98
Etymology: In reference to the ascospore similarity of the
taxon to Massarina eburnea.
Holotype: HHUF 30079
Saprobic on Vitis coignetiae. Sexual morph Ascomata
240–300μm high, 150–300μm diam., scattered, immersed,
erumpent at the beak, subglobose in section. Neck 40–50μm
high, composed of carbonaceous, black, thick-walled cells,
without clypeus, with a slit-like ostiole and periphyses.
Peridium 5–10μm thick, composed of 2–4 layers of polygonal
to elongate, thin-walled cells (3.5–5×2–3μm). Hamathecium
numerous, trabeculate, 1–2μm wide, septate, branched and
anastomosed. Asci (125–)138–175 × 15–20 μm (x= 150 ×
17.9μm, n=27), 8-spored, numerous, bitunicate, fissitunicate,
cylindrical, with a short stipe (8–18μm long, x=14.1μm, n=
19), apically rounded with an ocular chamber. Ascospores 30–
37×(6–)7–9μm (x=32.9×8.3μm, n=42), L/W 3.5–4.5 (x=4,
n=42), broadly fusiform with rounded ends, mostly straight to
slightly curved, 1(–3)-septate, with a primary septum nearly
median (0.49–0.54, x=0.51, n=42), hyaline, smooth-walled,
guttulate when young, with an entire gelatinous sheath (3–
15μm wide at sides). Asexual morph Undetermined.
Culture characters: Colonies on PDA, pale mouse grey
with an entire olivaceous margin; on MEA, olivaceous with
white margin, with hyaline exudates on the surface; on CMA,
grey olivaceous in the centre. On rice straw agar (RSA:
Tanaka and Harada 2003a), numerous ascomata are produced
on the surface of rice straw. Ascospores are slightly wider than
those on natural specimen, 31–36×9–11μm (x=33.2×9.7μm,
n=20), L/W 2.9–3.8 (x=3.4, n=20), with a submedian primary septum (0.51–0.56; x=0.53, n=20).
Material examined: JAPAN, Hokkaido, Notsuke, Bekkai,
Keneyaushubetsu-river, near Mannen-bashi, on vines of Vitis
coignetiae (Vitaceae), 8 September 2003, K. Tanaka & S.
Hatakeyama, KT 1424 (HHUF 30079, holotype designated
here); ex-type living cultures, JCM 17826, MAFF 242970.
GenBank ITS: LC001709; LSU: LC001707; SSU: LC001706.
Notes: Morphological features of this species, such as compressed beak of ascomata with a slit ostiole, cylindrical asci
with a short stipe (mostly less than 15μm long), and 1(–3)septate ascospores with an entire sheath, agree with the generic concept of the genus Lophiotrema recently circumscribed
(Hirayama and Tanaka 2011). This species can be distinguished from other taxa in Lophiotrema (Tanaka and Harada
2003b; Zhang et al. 2009b) by its relatively large ascospores.
A BLAST search of the GenBank database with the 28S and ITS
sequences of L. eburnoides indicated that it is most close to L.
vagabundum (GenBank AF383954; Identities = 487/491
(99.1 %), Gaps=2/491 (0.4 %)), but the latter species has smaller
ascospores (20–26(–29)×4–5.5μm; Tanaka and Harada 2003b).
Melanommataceae
Melanommataceae is characterized by globose or depressed
ascomata, bitunicate and fissitunicate asci, pigmented
phragmosporous ascospores as well as the trabeculate
pseudoparaphyses (Barr 1990; Sivanesan 1984).
Phylogenetic studies of this family have shown it to be wellsupported (Liew et al. 2000; Kodsueb et al. 2006a; Kruys et al.
2006; Wang et al. 2007; Hyde et al. 2013; Wijayawardene et
al. 2014). The asexual morphs of Melanommataceae are
mostly coelomycetous and rarely hyphomycetous. The phylogenetic tree is presented in Fig. 99.
67. Byssosphaeria musae Phookamsak & K.D. Hyde, sp.
nov.
Index Fungorum number: IF550932, Facesoffungi
number: FoF00436; Fig. 100
Etymology: The specific epithet musae refers to the host.
Holotypus: MFLU 11–0182
Fungal Diversity
Fig. 98 Lophiotrema eburnoides (holotype). a Ascomata on host
surface b Ascoma in longitudinal section c Peridium d, e Asci f Ascus
apex g Pseudoparaphyses h Ascospores in India ink i–l Ascospores m
Colonies on PDA (upper), MEA (left), and CMA (right) after 30 d at
20 °C in the dark. a-g, i, j from HHUF 30079 (holotype); h, k–m from
JCM 17826=MAFF 242970 (ex-holotype isolate). Scale bars: a=500μm,
b=50μm, c-g, i-l=10μm, h=50μm, m=1 cm
Saprobic on leaf sheath of Musa sp. Sexual morph
Ascomata 430–540μm high, 450–630μm diam., gregarious,
scattered, superficial on subiculum, visible as dark spots on
host, orange to yellow around pore, uni-loculate, globose to
subglobose, setose, apex rounded, ostiole central, with pore-like
opening. Peridium 35–80μm wide, thick-walled, of equal
thickness, composed of several layers of dark brown to black
cells, arranged in textura angularis to textura prismatica.
Hamathecium 0.5–1.7μm wide, composed of dense, trabeculate, distinctly septate, anastomosing, pseudoparaphyses, embedded in a hyaline gelatinous matrix. Asci (120–)125–135(–
145)×(11.5–)12–14(–17) μm (x=132.8×13.7μm, n=25), 8-
spored, bitunicate, fissitunicate, clavate, long pedicellate (40–
50μm long) with knob-like pedicel, apically rounded, with
well-developed ocular chamber. Ascospores (27–)30–33(–
36)×(4–)5–6μm (x=32.3×5.9μm, n=30), overlapping 1–2-seriate, fusiform, with acute ends, hyaline to pale brown when
young, becoming light brown at maturity, 1(–3)-septate,
not constricted at the septa, slightly curved, smoothwalled, bearing delicate hyaline appendages over ends
with wing-like appendages near the central septum.
Asexual morph Undetermined.
Culture characters: Colonies on PDA fast growing,
80–90 mm diam. after 2 weeks at 25–30 °C, white to
Fungal Diversity
Fig. 99 Phylogram generated
from Maximum likelihood
(RAxML) analysis based on
combined LSU, SSU and TEF1
sequence data of
Melannomataceae. Maximum
likelihood 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-types (reference
strains) are in bold; the new
isolates are in blue. The tree is
rooted with Biatriospora marina
strain CY 1228
cream or pale yellowish, intermixed with yellowish to
orangish hyphae; reverse cream to white yellowish at
the margin, yellowish-brown at the centre, medium
dense to dense, irregular, flattened to slightly raised,
dull with entire edge, fluffy to feathery, effuse,
zonate.
Material examined: THAILAND: Chiang Rai, Muang
District, Khun Korn Waterfall, on leaf sheath of Musa sp.
(Musaceae), 5 September 2010, R. Phookamsak RP0062
(MFLU 11–0182, holotype); ex-type living culture,
MFLUCC 11–0146. GenBank ITS: KP744435; LSU:
KP744477; SSU: KP753947.
Notes: Byssosphaeria musae is similar to B. salebrosa
(Cooke & Peck) M.E. Barr and B. schiedermayeriana
(Fuckel) M.E. Barr (1984; Mugambi and Huhndorf 2009),
with ascomata, asci and ascospores having a similar size
range, but being smallest in B. musae which has smaller
ascomata, asci and ascospores than these species.
Byssosphaeria musae differs from B. schiedermayeriana in
ascospore appendages, which are present in B. musae, but
lacking in B. schiedermayeriana. Phylogenetic analyses of
combined genes (Fig. 99) showed that Byssosphaeria musae
forms a separate robust clade (77 % MP and 67 % ML) close
to B. schiedermayeriana and B. salebrosa.
Fungal Diversity
Fig. 100 Byssosphaeria musae (holotype) a Ascomata on host surface b Section through ascoma c Peridium d Pseudoparaphyses e–i Asci j–n
Ascospores o Ascospore stained in Indian ink. p Germinating spore. Scale bars: b=100μm, c-i, p=20μm, j-o=10μm
Fungal Diversity
68. Pseudotrichia rubriostiolata Phookamsak & K.D.
Hyde, sp. nov.
Index Fungorum number: IF550933, Facesoffungi
number: FoF00437; Fig. 101
Etymology: The specific epithet rubriostiolata refers to the
ascomata ostioles with red pigments.
Holotypus: MFLU 11–0174
Saprobic on Thysanolaena maxima. Sexual morph
Ascomata 140–220μm high (excluding neck), 150–190μm
diam., solitary, scattered, immersed to semi-immersed or
erumpent with neck, visible as dark spots, with reddish pigment surrounding the ostiole, uni-loculate, globose to
subglobose, glabrous, ostiole central, with short papilla (70–
95×70–90μm). Peridium 5–20μm wide, thin-walled, of equal
thickness, composed of 2–3 layers, brown to dark brown,
flattened, pseudoparenchymatous cells, arranged in a textura
angularis. Hamathecium 1–1.5(–2) μm wide, composed of
dense, narrow cellular pseudoparaphyses, wall rough, with
small guttules, distinctly septate, anastomosing above the asci,
embedded in a hyaline gelatinous matrix. Asci (55–)70–80(–
88) × (13–)15–16(–17) μm (x=75.5 × 15.4 μm, n = 25), 8spored, bitunicate, fissitunicate, clavate, short pedicellate with
foot-like pedicel, apically rounded with well-developed ocular
chamber. Ascospores (23.5–)25–27(–29) × 4–5(–6) μm
(x=26.5×4.5μm, n=30), overlapping 1–3-seriate, fusiform,
slightly curved, hyaline, becoming light brown when geminated, 1-septate, constricted at the septa, swollen near the septa, smooth-walled, surrounded by distinct mucilaginous
sheath. Asexual morph forming stromatic fruiting bodies on
bamboo pieces on water agar (WA) after 12 weeks.
Conidiomata 80–205μm high, 80–210μm diam., pycnidial,
scattered or clustered, gregarious, semi-immersed to superficial on bamboo pieces, visible as black spots on surface, globose to subglobose or irregular in shape, glabrous, surrounded
by vegetative hyphae, ostiole undetectable, becoming reddish
to reddish-purple. Conidiomata walls 7.5–15.5μm wide, composed of several layers of dark brown to black, pseudoparenchymatous cells, initially arranged in textura intricata, becoming textura angularis. Conidiophores simple, rarely branched,
aseptate, hyaline, mostly reduced to conidiogenous cells.
Conidiogenous cells 4–16×1.5–2(–3) μm (x=7×1.8μm, n=
40), enteroblastic, phialidic, single, discrete, determinate,
doliiform to cylindrical or ampulliform, hyaline, arising from
basal stratum. Conidia 2–3.5×1–2μm (x=3.1×1.6μm, n=
40), one-celled, oblong or rod-shaped to obovoid, with rounded to obtuse ends, hyaline, smooth-walled.
Culture characters: Colonies on MEA slow growing,
reaching 45–50 mm diam. after 4 weeks at 25–30 °C, cream
to pale yellowish or pale reddish at the margin, white to cream
or pale yellowish at the centre with small reddish droplets
and the periphery of raised; reverse cream to white, yellowish or pale reddish at the margin, reddish to blond yellowish
at the centre, slightly radiating; medium dense to dense,
Fig. 101 Pseudotrichia rubriostiolata (holotype) a Ascomata on host
surface visible as black raised areas with reddish pigments at the apex b
Section through an ascoma c Section through peridium d Asci with
pseudoparaphyses stained in Congo red reagent e–g Asci h–l
Ascospores m Germinating spore n Pycnidia on bamboo pieces on WA
o Section through stromatic pycnidia p Section though pycnidial walls q–
t Conidiophores u–ac Conidia. Scale bars: b=50μm, c-g, m=20μm, h-l,
p-u=10μm, r-t=2μm, v-ac=1μm, o=100μm
circular, flattened to slightly raised, dull with entire edge,
initially mucoid, becoming velvety to felty, seperating from
agar, produced reddish-brown to light brown pigments diffusing in the agar and conidiomata immersed in agar after
8 weeks.
Material examined: THAILAND, Chiang Rai Province,
Muang District, Mae Fah Luang University campus grounds,
on dead stem of Thysanolaena maxima (Poaceae), 13 August
2010, R. Phookamsak RP0054 (MFLU 11–0174, holotype),
ex-type living culture, MFLUCC 11–0138, GenBank ITS:
KP744463; SSU: KP744508; ibid., on dead stem of
Thysanolaena maxima, 13 August2010, R. Phookamsak
RP0053 (MFLU 11–0173), living culture, MFLUCC 11–
0137. GenBank ITS: KP744462; LSU: KP744507; SSU:
KP753966; Phrae Province, Rongkwang District, Maejo
University Phrae Campus, on dead stem of Thysanolaena
maxima, 20 August 2010, R. Phookamsak RP0072 (MFLU
11–0192), living culture, MFLUCC 11–0156. GenBank ITS:
KP744464; LSU: KP744509; SSU: KP753967.
Notes: Pseudotrichia rubriostiolata clusters together with
P. guatopoensis in the phylogenetic tree. Morphologically it
differs from other Pseudotrichia species in the smaller size of
its ascomata, asci and ascospores. P. rubriostiolata is similar
to P. thailandica, but in P. rubriostiolata the ascomata, asci
and ascospores are smaller and the ascospores are slightly
constricted at the septum. In P. thailandica ascospores are
larger and deeply constricted at the septum, and easily separate into two parts. In the phylogenetic analyses, P.
rubriostiolata forms a clade with P. thailandica and P.
guatopoensis (strain SMH 4535).
69. Pseudotrichia thailandica Phookamsak & K.D. Hyde,
sp. nov.
Index Fungorum number: IF550934, Facesoffungi
number: FoF 00438; Fig. 102
Etymology: The specific epithet thailandica refer to the
country in which the fungus was first collected.
Holotypus: MFLU 11–0191
Saprobic on dead stem of Thysanolaena maxima. Sexual
morph Ascomata 140–180μm high (including neck), 165–
310μm diam., solitary to gregarious, scattered, semi-immersed, visible on host surface as raised, dark spots, with
reddish pigment surrounding the ostiole, uni-loculate, rarely
bi-loculate, subglobose, glabrous, ostiole oblique, with short
Fungal Diversity
Fungal Diversity
Fig. 102 Pseudotrichia thailandica (holotype) a Ascomata on host surface b Section through an ascoma c Peridium d Asci with pseudoparaphyses e–h
Asci i Ascospores in Indian ink. j–n Ascospores. Scale bars: b=50μm, c–i=20μm, j-n=10μm, o=100μm
papilla, carbonaceous. Peridium 7–22μm wide, thin-walled,
of unequal thickness, slightly thin at the base of ascoma, composed of several layers of brown to dark brown, pseudoparenchymatous cells, outer layer comprising 2–3 layers, flattened,
arranged in a textura prismatica, inner layer comprising 2–3
layers, arranged in textura angularis. Hamathecium composed of dense, 0.8–2.5μm wide, dense, narrow, cellular
pseudoparaphyses, wall rough with small guttules, distinctly
septate, anastomosing above asci, embedded in a hyaline gelatinous matrix. Asci (77.5–)80–100(–115.5)×(14–)16–18(–
Fungal Diversity
20) μm (x= 95.1× 17.3 μm, n = 25), 8-spored, bitunicate,
fissitunicate, clavate, short pedicellate with foot-like or
knob-like pedicel, apically rounded, with well-developed ocular chamber. Ascospores (27–)28–32(–34.5)×4–5(–6) μm
(x=30×4.7μm, n=30), overlapping 1–3-seriate, fusiform,
slightly curved, hyaline to subhyaline, 1(–5)-septate, constricted at the septa, easily seperated into two part spores,
swollen near the septa in the upper cell, smooth-walled,
surrounded by a distinct mucilaginous sheath. Asexual
morph Undetermined.
Culture characters: Colonies on PDA slow growing,
reaching 30–35 mm diam. after 4 weeks at 25–30 °C, cream
to pale yellowish at the margin, grey to dark grey at the centre,
with small black droplets; reverse cream to yellowish-white at
the margin, dark greenish at the centre, slighty radiating,
forming sectors; medium dense to dense, circular, flattened
to slightly raised, dull with undulate edge, mucoid, fairly
fluffy at the magins, seperating from agar, felty in the centre
with sparse hyphae at the margin.
Material examined: THAILAND: Phrae Province,
Rongkwang District, Maejo University Phrae Campus, on
dead stem of Thysanolaena maxima (Poaceae), 20 August
2010, R. Phookamsak RP0071 (MFLU 11–0191, holotype),
ex-type living culture, MFLUCC 11–0155. GenBank ITS:
KP744465; LSU: KP744510; SSU: KP753968.
Notes: Pseudotrichia thailandica is similar to P.
rubriostiolata in having a red pigment in the ostiole.
However, Pseudotrichia thailandica has larger ascomata, asci
and ascospores than P. rubriostiolata. These two species form
a clade together with P. guatopoensis (strain SMH 4535) in
the phylogenetic tree (Fig. 99).
Fig. 103 Sarimanas pseudofluviatile (holotype) a Ascoma in
longitudinal section b Peridium c, d Asci e Ascus apex f
Pseudoparaphyses g–j Ascospores k Colonies on PDA (upper), MEA
(left), and CMA (right) after 30 d at 20 °C in the dark. a–j from HHUF
27552 (holotype); k from MAFF 239465 (ex-type). Scale bars: a=
100μm, b, c–j=10μm, k=1 cm
70. Sarimanas Matsumura, K. Hiray. & Kaz. Tanaka, gen.
nov.
Index Fungorum number: IF551052, Facesoffungi
number: FoF00497
Etymology: An anagram of Massarina.
Type species: Sarimanas shirakamiense.
Ascomata immersed, scattered or 2–3 gregarious, globose to subglobose, ostiolate. Neck short papillate, without clypeus. Ascomatal wall composed of polygonal
thin-walled cells. Hamathecium cellular, numerous, septate. Asci fissitunicate, cylindrical to ovoid, with a short
stipe. Ascospores broadly fusiform with rounded ends,
1-septate, hyaline, smooth-walled, with an entire gelatinous sheath.
71. Sarimanas pseudofluviatile Matsumura, K. Hiray. &
Kaz. Tanaka, sp. nov.
Index Fungorum number: IF551053, Facesoffungi
number: FoF00498; Fig. 103
Fungal Diversity
Etymology: In reference to the ascospore similarity of the
taxon to Massarina fluviatilis (= Lentithecium fluviatile).
Holotype: HHUF 27552
Saprobic on dead wood. Sexual morph Ascomata 140–
220μm high, 180–320μm diam., globose to subglobose, covered with sparse hyphae, ostiolate. Neck 37–50μm long, 50–
75μm wide. Peridium in surface view textura prismatica, in
longitudinal section 20–35μm thick at sides, composed of 4–6
layers of polygonal, hyaline to brown, 5–13×5–8μm cells.
Hamathecium 1.5–2.5μm wide, septate. Asci 90–110×18–
23.5μm (x=99×20.2μm, n=30), fissitunicate, cylindrical,
short pedicellate (5–15μm long; x=8.8μm, n=19), apically
rounded with a shallow ocular chamber. Ascospores
(27–)28.5–33(–36)×(7–)8–10μm (x=30.6×8.7μm, n=100),
L/W (3–)3.2–3.9(–4.1) (x=3.5, n=100), broadly fusiform to
ellipsoidal with rounded ends, with a septum submedian
(0.51–0.55, x=0.53, n=85), hyaline, smooth-walled, with an
entire sheath of 2–5μm wide. Asexual morph Undetermined.
Culture characters: On RSA, numerous ascomata are
formed on the surface of rice straw and the ascospores are
similar to those found in nature, measuring 28.5–33×8.5–
10 μm (x=30.7 × 9.3 μm, n = 30). No asexual morph is
formed.
Materials examined: JAPAN, Hokkaido, Sapporo
Toyohira-river, riverbank, on dead twigs of woody plant, 2
September 2001, K. Tanaka, KT 760 (= HHUF 27552,
holotype designated here); ex-type living culture=MAFF
239465. GenBank ITS: LC001717; LSU: LC001714; SSU:
LC001711; ibid., KT 759 (= HHUF 27551, paratype).
Notes: This species was reported as Massarina fluviatilis
by Tanaka and Harada (2003c), but this was obviously a misidentification. We examined the holotype of M. fluviatilis (Van
Ryckegem 509 in GENT; Van Ryckegem and Aptroot 2001)
and found that M. fluviatilis on Phragmites australis
(Poaceae) has ascomata composed of thick-walled, slightly
larger cells (5–30×2.5–11μm) when compared to those of S.
pseudofluviatile on woody plants. The ascospores of M.
fluviatilis are 1–3-septate in sequence 2:1:3, but those are consistently 1-septate in S. pseudofluviatile. Zhang et al. (2009b)
established a new genus Lentithecium using M. fluviatilis as
the type species, and later placed the genus in Lentitheciaceae
(Zhang et al. 2009a). In a phylogenetic analysis, S.
pseudofluviatile deviated from Lentitheciaceae and clustered
in the clade of Melanommataceae together with S.
shirakamiense (Fig. 99).
72. Sarimanas shirakamiense Matsumura, K. Hiray. &
Kaz. Tanaka, sp. nov.
Index Fungorum number: IF551054, Facesoffunginumber:
FoF00497; Fig. 104
Etymology: In reference to the location where the specimen
was collected.
Holotype: HHUF 30454
Fig. 104 Sarimanas shirakamiense (holotype). a, b Ascomata on host
surface c, d Ascomata in longitudinal section e Peridium of surface view f
Peridium in longitudinal section g, h Asci i Ascus apex j
Pseudoparaphyses k–n Ascospores o Ascospores with an expanding
gelatinous sheath p Ascomata on rice straw in culture q Colonies on
PDA (upper), MEA (left), and CMA (right) after 30 d at 20 °C in the
dark. a-g, i, m from HHUF 30081 (paratype); n-q from JCM 17825=
MAFF 242969 (ex-paratype); h, j, k from HHUF 30454 (holotype); l
from MAFF 244768 (ex-type). Scale bars: a, b=1 mm, c, d=100μm, e–
o=10μm, p, q=1 cm
Saprobic on Swida controversa. Sexual morph Ascomata
220–300μm high, 200–320μm diam., mostly scattered, sometimes 2–3 gregarious, immersed, globose in section. Neck 30–
45μm high, 70–80μm wide, short papillate, without clypeus.
Peridium uniformly 25–30μm thick, composed of 4–6 layers
o f p o l y g o n a l , 1 0– 20 × 4 –8 μ m , th i n - w a l l e d c e l l s .
Hamathecium numerous, cellular, 2–3μm wide, hyaline, septate, branched and anastomosed. Asci 95–158(–175)×26.5–
52.5μm (x=133.2×39.6μm, n=30), 8-spored, not numerous,
bitunicate, fissitunicate, cylindrical to ovoid, with a short pedicel (8.5–20μm) long, apically rounded with an ocular chamber. Ascospores (30–)32–51×10–17(–18.5) μm (x=40.3×
13.9μm, n=60), L/W 2.2–3.8 (x=2.9, n=60), broadly fusiform with rounded ends, mostly straight, thick-walled, with
a septum mostly submedian (0.49–0.62; x=0.53, n=60), hyaline, smooth-walled, guttulate when young, with an entire
sheath; sheath when fresh condition diffuse, gelatinous, up
to 8μm wide, later becoming sharply delimited firm sheath
of 2–4μm thick. Asexual morph Undetermined.
Culture characters: Colonies on PDA, white, buff pigment
produced; on MEA, white to olivaceous grey; on CMA, white
to lavender grey. On RSA, numerous ascomata are produced
on the surface of rice straw. Ascomata ca. 400μm high, ca.
250μm diam., immersed, globose, with papilla. Asci and ascospores are considerably larger than those on the natural
specimen; asci 162–225×34–46.5μm (x=192.8×38.5μm,
n = 20); ascospores (42.5–)45–51(–53) × (10–)15–17.5 μm
(x=47.5×15.4μm, n=50), L/W (2.6–) 2.8–3.3(–3.6) (x=3.1,
n=50), with a septum mostly submedian (0.48–0.54; x=0.51,
n=50).
Material examined: JAPAN, Aomori, Nishimeya,
Shirakami Natural Science Park of Hirosaki Univ., on twigs
of Swida controversa (Cornaceae), 3 June 2012, K. Tanaka,
KT 3000 (HHUF 30454, holotype designated here); ex-type
living culture, MAFF 244768. GenBank ITS: LC001718;
LSU: LC001715; SSU: LC001712; ibid., Aomori,
Nishimeya, Seisyu-trail, on dead twigs of woody plant, 29
May 2007, K. Hirayama et al., KH 13 (HHUF 30081,
paratype); ex-paratype living cultures, JCM 17825, MAFF
242969. GenBank ITS: LC001719; LSU: LC001716; SSU:
LC001713.
Notes: A new genus, Sarimanas, is established for S.
shirakamiense and S. pseudofluviatile based on their shared
Fungal Diversity
Fungal Diversity
morphological characters, such as globose to subglobose
ascomata composed of polygonal thin-walled cells, cylindrical to ovoid asci with a short pedicel, and broadly fusiform, 1septate, hyaline ascospores with an entire sheath. These morphological features of Sarimanas are somewhat similar to
those of Massarina, but the latter genus has clypeate ascomata
and belongs to Massarinaceae (Hyde et al. 1999; Zhang et al.
2012). Ascospore morphology of Sarimanas superficially resembles to that of Wettsteinina in Pleomassariaceae (Kodsueb
et al. 2006) or Lentitheciaceae (Schoch et al. 2009), but the
latter genus has ascomata lacking a hamathecium when mature (Zhang et al. 2012a, b). The phylogenetic analysis based
on LSU (Fig. 99) suggested that this genus has close affinity to
genera in Melanommataceae, such as Aposphaeria and
Herpotrichia, but within in this family no genus phenotypically similar to Sarimanas is known. Sarimanas
shirakamiense is similar to S. pseudofluviatile, but the asci
and ascospores of S. shirakamiense are considerably larger
than those of the latter (asci 99×20.2μm, ascospores 30.6×
8.7μm in average). Sequence differences between these two
taxa were found at eight positions in the ITS region, suggesting that they are not conspecific.
Microthyriaceae
73. Neomicrothyrium Boonmee, H.X. Wu & K.D. Hyde.,
Fungal Diversity 51(1): 217 (2011)
Index Fungorum number: IF 563368.
Type species: Neomicrothyrium siamense Boonmee, H.X.
Wu & K.D. Hyde
Neomicrothyrium siamense Boonmee, H.X. Wu & K.D.
Hyde., Fungal Diversity 51(1): 217 (2011)
Index Fungorum number: IF 563369
Notes: Wu et al. (2011) proposed a new genus
Neomicrothyrium including a new species Neomicrothyrium
siamense under Dothideomycetes genera incertae sedis. Since
the publication, Index Fungorum (2015) has indicated that this
species is a Nom. Inval., Art. 40.6 and the genus name is also
Nom. Inval., Art. 35.1. According to the Melbourne
International Code of Nomenclature (ICN) in McNeill et al.
(2012), these articles are reported in the errors of typus not
assigned. Here we designate and validly publish the type species of Neomicrothyrium, Neomicrothyrium siamense
Boonmee, H.X. Wu & K.D. Hyde.
Mycosphaerellaceae
The family Mycosphaerellaceae was introduced by Lindau
(1896) with Mycosphaerella as the type genus. It was initially
placed in the order Dothideales (Hawksworth et al. 1995).
Kirk et al. (2001) introduced a separate order –
Mycosphaerellales for the family, but later again it was placed
in the order Capnodiales by Kirk et al. (2008). This placement
of Mycosphaerellaceae in Capnodiales has been phylogenetically confirmed in studies by Crous et al. (2007; 2009a, b).
The family includes 14 sexual morph-typified genera, namely Achorodothis, Brunneosphaerella, Cymadothea,
Euryachora, Gillotia, Melanodothis, Mycosphaerella,
P l a co c re a , P olysporella, Pseudostigmidium,
Sphaerellothecium, Sphaerulina, Stigmidium and
Wernerella (Lumbsch and Huhndorf 2010). More than 30
asexual genera linked to the type genus Mycosphaerella
are also included in the family (Hyde et al. 2013), while
further genera have subsequently been added in recent studies (Crous et al. 2013; Quaedvlieg et al. 2013). These asexual genera consist of some of the most common and destructive plant pathogens that affect a wide variety of host plants,
especially economically important fruit and vegetable crops,
and cereals (Farr et al. 1995; Crous and Braun 2003). The
phylogenetic tree is presented in Fig. 105.
74. Pallidocercospora acaciigena (Crous & M.J. Wingf.)
Crous & M.J. Wingf. Stud. Mycol. 75: 74 (2012)
≡ Mycosphaerella acaciigena Crous & M.J. Wingf., in
Crous, Groenewald, Pongpanich, Himaman, Arzanlou &
Wingfield, Stud. Mycol. 50(2): 463 (2004)
Index Fungorum number: IF564821,
Facesoffunginumber: FoF 00439; Fig. 106
Saprobic on palms. Sexual morph Ascomata 50–70μm
high., scattered, solitary to gregarious, immersed to semi-immersed, with protruding papilla, visible as black spots on host
surface, uniloculate, globose to subglobose, glabrous, ostiole
central, with minute papilla, obtuse at the apex. Peridium 4–
10μm wide, thin-walled, of equal thickness, comprising 2–6
layers of brown pseudoparenchymatous cells, arranged in
textura angularis. Hamathecium lacking pseudoparaphyses.
Asci (27–)30–40(–53)×(5–)6–8(–9) μm (x=36.6×7.2μm,
n=25), 8-spored, bitunicate, fissitunicate, cylindric-clavate to
ampulliform, or obclavate, often ventricose, apedicellate, apically rounded, with an ocular chamber (0.5–1μm wide).
Ascospores 9–12×2–3μm (x=10.5×2.9μm, n=30), overlapping 2–3-seriate, oblong to clavate, hyaline, 1-septum, not
contricted to slightly constricted at the septa, with smooth or
rough walls, with small guttules, or slightly echinulate, mostly
upper cell wider than lower cell. Asexual morph
Undetermined.
Culture characters: Colonies on MEA slow growing,
reaching 18–20 mm diam. after 3 weeks at 25–30 °C, dark
greenish at the margin, pale greenish at the centre, becoming pale
greenish-grey to grey at the centre; reverse dark greenish, becoming paler in the centre; dense, irregular, raised to umbonate, with
entire edge, velvety, slightly radiating, curved, radially furrowed.
Material examined: THAILAND: Chiang Rai, Muang
District, Khun Korn Waterfall, on dead leaves of palm, 12
Fungal Diversity
Fig. 105 Phylogram generated
from Maximum likelihood
(RAxML) analysis based on
combined LSU and ITS sequence
data of Capnodiales. Maximum
likelihood 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-types (reference
strains) are in bold; the new
isolates are in blue. The tree is
rooted with Lecanosticta acicola
CBS 871.95
May 2010, R. Phookamsak RP0032 (MFLU 11–0153), living
culture, MFLUCC 10–0561. GenBank ITS: KP744451; LSU:
KP744494.
Notes: Pallidocercospora was introduced by Crous et al.
(2013) to accommodate taxa that are not congeneric with
Mycosphaerella (Crous et al. 2012). The genus was considered to be distinguished from Mycosphaerella based on its
asexual morph. Mycosphaerella is represented by Ramularia
asexual morphs (Quaedvlieg et al. 2013), whereas,
Pallidocercospora has pseudocercospora-like asexual morphs
(Crous et al. 2004, 2012). Pallidocercospora acaciigena is
morphologically similar to Mycosphaerella thailandica
Crous et al., but differs in its asexual morph and ascomata
(Crous et al. 2004). Pallidocercospora acaciigena has longer
pseudocercospora-like conidia than M. thailandica and the
ascomata of P. acaciigena are also dense, superficial and clustered (Crous et al. 2004). Pallidocercospora acaciigena has
been reported as pathogen on Acacia in Venezuela (Crous et
al. 2004), while our isolate was found as a saprobe on palms in
Thailand, representing a new host and continent. Based on a
megablast nucleotide search in GenBank of ITS gene, our
isolate is related to P. acaciigena (99 % similarity), P.
crystallina (99 %), and Mycosphaerella heimii (99 %).
However, the ITS pairwise comparison has shown that our
isolate is most closely related to P. acaciigena. Phylogenetic
analyses of LSU genes shows that our isolate forms a strongly
Fungal Diversity
Fig. 106 Pallidocercospora acaciigena (MFLU 11–0153) a Ascomata visible as black spots on host surface b Vertical section through an ascoma c
Peridium d, e, f Asci g, i, j, k, l Ascospores h Germinating ascospore m, n Culture characters. Scale bars: b=20μm, c–f=10μm, g, h, = 5μm, i–l=2μm
supported clade with P. acaciigena.
75. Pseudocercospora tamarindi Goon. & K.D. Hyde, sp.
nov.
Index Fungorum number: IF550996, Facesoffungi
number: FoF00473; Fig. 107
Etymology: in reference to the host Tamarindus indica.
Holotype: MFLU 14–0805
Pathogen on living leaves of Tamarindus indica. Sexual
morph Undetermined. Asexual morph Leaf spots
epiphyllous, indistinct, subcircular but mostly irregular, 0.5–
1cm diam., at most 2cm diam. when confluent, pale brown to
golden brown, delimited by leaf mid rib. Caespituli
amphigenous but predominantly epiphyllous, abundant on
leaf spots, dense, dark, punctiform. Stromata superficial, globular, pale to dark brown 11–25μm diam., cells rounded,
Fungal Diversity
Fig. 107 Pseudocercospora tamarindi (holotype) a Symptoms on
leaves (upper surface) b, c Close up of leaf spot d, e Stroma with
attached conidiophores and conidia f Stromatal cells g-p Conidia of
different sizes and maturity q Colony on PDA (3 weeks). Scale bars:
b=500μm, c=50μm, d–f, h–o=10μm, g, p=20μm
oblong or irregular, 2–3μm diam., wall thickness less than
0.3μm. Conidiophores fasciculate, arising from the stromata,
brown becoming pale brown towards the apex, smooth, 0–2septate, often short or one-cells, straight to slightly curved,
apex rounded to truncate, unbranched, 3–12×1–2μm (x=
7.37 × 1.81 μm, n = 20), wall thickness less than 0.25 μm.
Conidiogenous cells 5–7×0.8–2μm (x=5.66×1.59, n=10),
integrated, terminal or conidiophores reduced to
conidiogenous cells, occasionally branched, smooth, hyaline
to pale brown, integrated, proliferating percurrently,
conidiogenous loci inconspicuous. Conidia (13–)17–25(–
30)×(1.8–)2–3(–3.5) μm (x=21×2.38μm, n=30), solitary,
subhyaline to brown, cylindrical to obclavate, some showing
constrictions at septa, some guttulate, subacute to obtuse apex,
obconically subtruncate base, 2–14–septate, hila neither thickened or darkened, 0.7–2μm diam.
Culture characters: Colonies on PDA show slow growth,
attaining 10–12mm diam. after 3 weeks at 25 °C, dark-grey to
black at the margin, grey to pale grey in the centre; reverse
dark-green to black; medium dense with dense center, irregular, convex surface, smooth lobate margin, hairy, nonpigmented.
Material examined: THAILAND, Chiang Rai Province,
Mueang Chiang Rai District, on living leaves of Tamarindus
indica L. (Fabaceae), 12 February 2014, K.D. Hyde IG018
(MFLU 14–0805, holotype); ex-type living culture,
MFLUCC 14–0805. GenBank ITS: KP744461; LSU:
KP744506; SSU: KP753965.
Notes: The genus Pseudocercospora was introduced by
Spegazzini (1910) with Pseudocercospora vitis as the type
species. The genus comprises asexual morphs or species with
Mycosphaerella-like sexual morphs, which includes many
Fungal Diversity
important plant pathogens (Hyde et al. 2013). They are characterized morphologically by their conidiophores and conidia
being pigmented and by the conidiogenous loci being inconspicuous or at least unthickened and not darkened (Crous and
Braun 2003; Crous et al. 2013). This new species is the first
cercosporoid to be described from the host Tamarindus indica
in Thailand. Previously, a species Cercospora tamarindi had
been recorded on leaves of Tamarindus indica in Uttar
Pradesh, India (Khan et al. 1988; Index Fungorum 2015).
Pseudocercospora tamarindi differs from this species by having conidiophores that lack distinct spore scars with much less
thick wall (>0.25μm) and darker coloured conidia. Although
they both have a similar range for the number of septa (-14),
the spores of P. tamarindi are much smaller (13–)17–25(–
30)×(1.8–)2–3(–3.5) μm compared to those of C. tamarindi
(38.4–93×3.8μm according to Khan et al. 1988). A molecular
analysis based on a combined gene analysis of ITS and LSU
shows that Pseudocercospora tamarindi clusters within the
major clade that includes the type species Pseudocercospora
vitis with a bootstrap value of 84 % but is distant from the type
s p e c i e s . Ye t i t i s c l e a r l y s e p a r a t e d f r o m o t h e r
Pseudocercospora species. Pseudocercospora tamarindi is
related with Pseudocercospora punctata (Wakef.) B. Sutton
(= Cercostigmina punctata Wakef.) which has verruculose
conidia (which the former lacks). It is also morphologically
similar, but different in conidial size and the number of septa
to Pseudocercospora ocimicola which has 1–8 septate conidia
of (15–)25–75(–85)×2–4μm (Crous et al. 2013).
76. Zasmidium musae (Arzanlou & Crous) Crous & U.
Braun., Schlechtendalia 20: 102 (2010)
≡ Stenella musae Arzanlou & Crous, Persoonia 20: 31
(2008)
Index Fungorum number: IF516587, Facesoffunginumber:
FoF 00440; Fig. 108
Biotrophic or hemibiotrophic causing necrotic leaf spots on
palms. Lesions initially small, globose to subglobose, becoming irregularly broader lesions, dark brown to reddish-brown
at the margin, dry, pale brown at the centre. Sexual morph
Ascomata 50–60μm high, 60–75μm diam., scattered, solitary
to gregarious, immersed to semi-immersed, with protruding
papilla, visible as black spots on host surface, uniloculate,
globose to subglobose, glabrous, ostiole central, with minute
papilla, rounded at the apex. Peridium 5–11μm wide, thinwalled, of equal thickness, comprising 3–7 cell layers of
brown to dark brown, pseudoparenchymatous cells, arrenged
in a textura angularis. Hamathecium lacking
pseudoparaphyses. Asci (26.5–)27–33(–36)×9–12μm (x=
31.9×10.9μm, n=25), 8-spored, bitunicate, fissitunicate,
ampulliform to ovoid, with subsessile to knob-like pedicel,
apically rounded, with well-developed ocular chamber,
thick-walled at the apex. Ascospores 8–10×2–3μm (x=9.5×
2.7μm, n=30), irregular overlapping 1–4-seriate, oblong to
clavate, hyaline, 1-septate, not contricted at the septa, wall
smooth to rough, thick-walled, mostly upper cell wider than
lower cell. Asexual morph Undetermined.
Culture characters: Colonies on MEA slow growth, 24–
25 mm diam. after 3 weeks at 25–30 °C, dark greenish to dark
grey at the margin, white to pale grey at the centre; reverse
dark greenish to black; dense, irregular, raised, rough with
entire edge, glabrous to velvety, radially furrowed.
Material examined: THAILAND: Chiang Mai, San Sai
District, Maejo University, on living leaves of palm, 18 July
2010, R. Phookamsak RP0048 (MFLU11–0168), living culture, MFLUCC 11–0132. GenBank ITS: KP744472; LSU:
KP744514; SSU: KP753970.
Notes: Zasmidium musae (Arzanlou & Crous) Crous & U.
Braun was described as Stenella musae Arzanlou & Crous
based on its asexual morph and lack of known sexual morph
(Arzanlou et al. 2008). Based on a megablast nucleotide
search in GenBank of the ITS sequences, our isolate is identified as Stenella musae and the pariwise comparison of ITS
shows that our isolate and Z. musae differ only in one base
pair. Based on phylogenetic analysis of LSU gene data, our
isolate forms a clade clustered with Zasmidium scaevolicola.
Therefore, we report the sexual morph of Zasmidium musae.
77. Paradictyoarthriniaceae Doilom, J.K. Liu & K.D.
Hyde, fam. nov.
Index Fungorum number: IF550921; Facesoffungi
number: FoF00499.
Saprobic on dead wood. Sexual morph Undetermined.
Asexual morph Colonies on natural substrate, superficial,
scattered, black, powdery. Conidiophores macronematous,
erect to slightly curved, constricted at septa. Conidiogenous
cells blastic, integrated, terminal, determinate. Conidia
muriform, subglobose to ellipsoidal, brown to black, verrucose, solitary or developing in branched chains with short 1–
3 chains, very variable in size and shape, circular to irregular
with a protruding basal cell; rounded to truncate at the base.
Notes: The family Paradictyoarthriniaceae is established
to accommodate the genus Paradictyoarthrinium in the order
Pleosporales, Dothideomycetes based on its unique morphology and distinct lineage in the phylogenetic analysis. The
combined phylogeny of LSU, SSU, TEF1 and RPB2 gene
data (Fig. 109) shows that Paradictyoarthrinium species
groups close to Roussoellaceae (Liu et al. 2014) and
Biatriosporaceae (Hyde et al. 2013), but is a distinct clade
with MLBS (64 %) and PP (0.99) support within the order
Pleosporales (Fig. 109). The morphology differs from the
asexual morph of Roussoellaceae in many characters (see
Liu et al. 2014). The family presently comprises two species,
namely Paradictyoarthrinium diffractum Matsush and P.
tectonicola sp. nov.
Type genus: Paradictyoarthrinium Matsush., Matsush.
Mycol. Mem. 9:18 (1996)
Fungal Diversity
Fig. 108 Zasmidium musae (MFLU11–0168) a-c Ascomata visible as black spots on leaf lesions. d Vertical section through an ascoma. e Peridium. f–j
Asci. k–n Ascospores. o, p Culture characteristic. Scale bars: d=20μm, e–j=10μm, k–n=2μm
Type species: Paradictyoarthrinium diffractum Matsush.,
Matsush. Mycol. Mem. 9:18 (1996).
Notes: Paradictyoarthrinium was introduced as a monotypic genus by Matsushima (1996) with
Paradictyoarthrinium diffractum as the type species. It was
described from a dead decaying spathe of Cocos nucifera
from a rivulet at Rustenburg in South Africa. Prabhugaonkar
and Bhat (2011) stated that based on ML analysis of ITS gene
data, the genus Paradictyoarthrinium is related with members
of the order Pleosporales, Dothideomycetes. However, multigene phylogenetic analysis with an extended dataset and the
relevant type strains were thought to be required to resolve
their suitable placement within the families of Pleosporales
(Prabhugaonkar and Bhat 2011).
In the current study we resolve the placement of
Paradictyoarthrinium. The MP ITS analysis showed two
isolates in this study (MFLUCC 12-0557 and MFLUCC
13-0466) grouping with P. diffractum (ex-type GUFCC
15514) with strong bootstrap support (91 % MPBS, tree
not shown). We established isolates MFLUCC 12–0557
and MFLUCC 13–0466 as P. diffractum based on identical morphology and phylogenetic analysis as compared to
the type. The combined dataset of LSU, SSU, TEF1 and
RPB2 were then used for family placement (see note for
family). Our new sequence data of Paradictyoarthrinium
diffractum is provided here. GenBank: (MFLUCC
12-0557, ITS: KP744454; LSU: KP744497); (MFLUCC
13-0466, ITS: KP744455; LSU: KP744498; SSU:
KP753960).
78. Paradictyoarthrinium tectonicola Doilom & K.D.
Hyde, sp. nov.
Fungal Diversity
Fungal Diversity
Phylogram generated from Maximum likelihood (RAxML)
analysis based on combined LSU, SSU, TEF1 and RPB2 sequence data
of Pleosporales. Maximum likelihood 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-types (reference strains) are in bold; the new isolates are in blue. The
tree is rooted with Dothidea insculpta CBS 189.58
Fig. 109
IndexFungorum no: IF550900, Facesoffungi number:
FoF00315; Fig. 110
Etymology: In reference to the host Tectona, and cola
meaning loving.
Holotypus: MFLU 14–0629.
Saprobic on dead stump of Tectona grandis L.f. Sexual
morph Undetermined. Asexual morph Colonies on natural substrate, superficial, gregarious, scattered, black, powdery. Conidiophores up to 33μm long, 1.5–3μm diam.,
macronematous, erect to slightly curved, constrict at the
septa, arising from hyphae. Conidiogenous cells blastic,
integrated, terminal, determinate. Conidia (8–) 17–21 (–
26) high × (8–) 16–19 (–22) μm (x= 18 × 16 μm, n = 30),
muriform, subglobose to ellipsoidal, brown to black, verrucose, solitary or developing in branched chains, with 1–
3 short chains, very variable in size and shape; circular to
irregular with a protruding basal cell; rounded to truncate
at the base.
Culture characters: Ascospores germinating on PDA within 24 h. Germ tubes produced around conidia. Colonies on
MEA reaching 18–28 and 26–35 mm diam. after 7 and 10 d in
the dark at 25 °C respectively (av=20 mm (7 d), 29 mm (10 d)
n=10), circular shape, superficial, flattened to effuse, velvety,
dense, grey (26E1), entire edge. Mycelium 1.5–3.5μm broad,
partly superficial, partly immersed, pale brown to dark brown,
septate, branched, verruculose. Chlamydospores reddishbrown. Conidiophores up to 30μm long, 3–4 diam., arising
from hyphae. Conidia (9–) 15–23 (–37) high×(9–) 14–18 (–
35) μm (x=19×17μm, n=30), muriform, reddish-brown to
dark brown.
Fig. 110 Paradictyoarthrinium tectonicola (holotype) a Colonies on host b Conidia with basal cells c–e Conidia with conidiogenous cells on host f, g
Conidia with conidial chains i Germinating conidia j Culture on MEA after 1 weekk-n Conidia with conidiophores on MEA. Scale bars: a=500μm, b, d,
h=10μm, c, e-f, i, k-n=20μm
Fungal Diversity
Material examined: THAILAND, Chiang Rai Province,
Mae Chan District, Kiu Tap Yang garden, on dead stem of
Tectona grandis (Lamiaceae), 3 March 2013, M. Doilom
(MFLU 14–0629, holotype), ex-type living culture,
MFLUCC 13–0465, ICMP 20686. GenBank ITS:
KP744456; LSU: KP744500; SSU: KP753961; ibid., MFLU
14-0630, living culture MFLUCC 12-0556. GenBank LSU:
KP744499.
Notes: Paradictyoarthrinium tectonicola is introduced as
second species of Paradictyoarthrinium. Two species can be
distinguished as colonies of P. tectonicola are slower growing
on MEA for at least 10 days than P. diffractum
(Prabhugaonkar and Bhat 2011). Paradictyoarthrinium
tectonicola and P. diffractum are very variable in size and
shape of conidia, thus having no obvious different conidia
dimensions. Nevertheless, phylogenetic analyses based on
ITS sequence data can separate them at the species level.
Isolates of P. tectonicola (MFLUCC 12–0556 and ex-type
MFLUCC 13–0465) grouped separately from P. diffractum
(MFLUCC 12–0557 and MFLUCC 13–0466) with strong
MLBS (100 %) and PP (1.00) support in the combined
LSU, SSU, TEF1 and RPB2 phylogeny (Fig. 109).
Phaeosphaeriaceae
A phylogenetic analysis and monograph of genera of
Phaeosphaeriaceae was published by Phookamsak et al.
(2014) and this is followed here. The phylogenetic tree is
presented in Fig. 111.
79. Allophaeosphaeria Ariyawansa, Camporesi, J.K. Liu
& K.D. Hyde, gen. nov.
Index Fungorum number: IF550997; Facesoffunginumber:
FoF 00494
Etymology: Named after its morphological resemblance to
Phaeosphaeria in contrast to the phylogenetic distance between both genera.
Type species: Allophaeosphaeria muriformia Ariyawansa,
Camporesi & K.D. Hyde
Saprobic on dead wood. Sexual morph Ascomata superficial, globose to subglobose, ostiolate. Ostiole papillate, without
periphyses Peridium comprising 2-layers, outer layer composed of heavily pigmented thick-walled, innermost layer of
broad, hyaline compressed rows of cells of textura
angularis. Hamathecium lacking pseudoparaphyses. Asci
bitunicate, fissitunicate, elongate cylindrical to slightly clavate with an ocular chamber. Ascospores oblong to narrowly
oblong, multi-septate, muriform, constricted at each septa,
hyaline, pale brown when mature, smooth-walled. Asexual
morph Undetermined.
80. Allophaeosphaeria dactylidis Wanasinghe, Camporesi,
E.B.G. Jones & K.D. Hyde, sp. nov.
Index Fungorum Number: IF550894, Facesoffungi
Number: FoF 00384; Fig. 112
Etymology: Named after the host genus from which it was
collected, Dactylis.
Holotype: MFLU 14–0754
Saprobic on dead herbaceous branches. Sexual morph
Ascomata 100–150μm high 100–175μm diam. (x=142.2×
131.2μm, n=10), solitary, scattered, immersed to erumpent,
globose or subglobose, dark brown to black, coriaceous,
ostiolate. Ostiole 30–70μm high 20–30μm diam. (x=48.7×
23.8μm, n=10), blackish-brown, smooth, ostiolar canal filled
with dark brown cells. Peridium 10–15μm wide at the base,
8–12μm wide at the sides, composed with reddish to dark
brown cells of textura angularis. Hamathecium comprising
numerous, 1.5–2.5μm (n=30) wide, filamentous, branched
septate, pseudoparaphyses. Asci 60–90×10–20μm (x=69×
14.6μm, n=40), 8-spored, bitunicate, fissitunicate, cylindricclavate to clavate, pedicellate, thick-walled at the apex, with
minute ocular chamber. Ascospores 15–20 × 5–8 μm (x=
18.2×6.3μm, n=50), overlapping 1–2-seriate, ellipsoidal to
subfusiform, upper part wider than the lower part, muriform,
3–5 transversely septate, with 1–3 vertical septa, constricted at
the central septum, initially hyaline, becoming yellowishbrown at maturity, ends remaining lighter and cone-shaped,
with rounded ends, without a mucilaginous sheath. Asexual
morph Undetermined.
Material examined: ITALY, Province of Forlì-Cesena [FC],
Corniolo, dead upright stems of Dactylis glomerata
(Poaceae), 12 July 2013, E. Camporesi (MFLU 14–0754,
holotype); ex-type living culture, MFLUCC 13–0618.
GenBank ITS: KP744432; LSU: KP744473; SSU:
KP753946.
Notes: Allophaeosphaeria dactylidis is the second species
of Allophaeosphaeria and it has muriform ascospores similar
to those characterized in P. vagans (Niessl) O.E. Erikss., P
phragmiticola Leuchtm and P. phragmitis (Hollós) Leuchtm
(Shoemaker and Babcock 1989). Multi-gene phylogenetic
analyses (ITS, LSU and SSU sequences) indicated that
Allophaeosphaeria dactylidis belongs to
Phaeosphaeriaceae, but is distinct from Phaeosphaeria sensu
stricto.
81. Allophaeosphaeria muriformia Ariyawansa,
Camporesi & K.D. Hyde, sp. nov.
Index Fungorum number: IF550998, Facesoffunginumber:
FoF00410; Fig. 113
Etymology: The specific epithet muriformia is based on the
ascospore septation.
Holotype: MFLU 15-0067
Saprobic on dead wood. Sexual morph Ascomata 275–
340μm high×300–380μm diam. (x=290×340μm), solitary,
scattered, superficial, globose to subglobose, ostiolate. Ostiole
papillate, black, smooth, with neck and without periphyses
Peridium 45–70μm wide, comprising 2-layers, outer layer
Fungal Diversity
Fig. 111 Phylogram generated from Maximum likelihood (RAxML)
analysis based on combined LSU, SSU and ITS sequence data of
Phaeosphaeriaceae. Maximum likelihood 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-types (reference strains) are in bold; the new isolates are in blue.
The tree is rooted with Didymella exigua CBS 183.55
composed of heavily pigmented thick-walled cells, innermost
layer of broad, hyaline compressed rows of cells of textura
angularis. Hamathecium lacking pseudoparaphyses. Asci
210–237 × 50–70 μm (x= 235 × 62 μm, n = 20), 8-spored,
bitunicate, fissitunicate, elongate cylindrical to slightly clavate, with a minute pedicel, thick-walled and rounded at apex,
Fungal Diversity
Fungal Diversity
Allophaeosphaeria dactylidis (holotype). a Ascomata on host
substrate b Section of ascoma c Close up of ostiole d Peridium e
Pseudoparaphyses f–i Asci j–l Ascospores m Ascospores stained in
Indian ink. Scale bars: a=200μm, b=50μm, c, d=20μm, e–i=10μm, j–
m=5μm
Fig. 112
with an ocular chamber. Ascospores 40–60×20–30μm (x=
56×26μm, n=40), overlapping 2–3-seriate, oblong to narrowly oblong, straight to slightly curved, muriform, multi-septate,
constricted at each septa, hyaline, pale brown when mature,
smooth-walled. Asexual morph Undetermined.
Material examined: ITALY, Province of Forlì-Cesena,
Montevescovo, on dead stem, 4 February 2013, E.
Camporesi (MFLU 14–1122, holotype), ex-type living culture, MFLUCC 13–0349. GenBank ITS: KP765680; LSU:
KP765681; SSU: KP765682.
Notes: Allophaeosphaeria resembles many species of
Phaeosphaeria in having a peridium comprising 2–3 layers
of brown to dark brown cells of textura angularis and multiseptate ascospores with a gelatinous sheath, but differs in
Fig. 113 Allophaeosphaeria muriformia (holotype) a, b Ascomata on
the host surface c Section of an ascoma d Close up of the peridium e–h
Cylindrical asci with a minute pedicel i–k Hyaline to pale brown
having elongate-cylindrical to slightly clavate asci with a clear
ocular chamber and lacks pseudoparaphyses. The phylogenetic analysis of combined ITS, LSU and SSU sequences provided strong evidence that the type species of Allophaeosphaeria,
A. muriformia belongs in Phaeosphaeriaceae and clustered
together with putatively named strains of Phaeosphaeria
phragmiticola (CBS 459.84) and Phaeosphaeria vagans
(CBS 604.86).
82. Neosetophoma clematidis Wijayawardene, Camporesi
& K.D. Hyde, sp. nov.
Index Fungorum number: IF 505909, Faces of Fungi number: FoF 00419; Fig. 114
Etymology: Named after host on which it was first collected
Holotypus: MFLU 14–0746
Saprobic on branch of Clematis vitalba. Sexual morph
Undetermined. Asexual morph Conidiomata 425–475μm
diam., 220–270μm high, pycnidial, superficial, black, gregarious, unilocular, but occasionally multi-locular, with a papillate
ostiole. Pycnidial wall 30–55μm wide, thick at outer layer,
ascospores k Ascospores mounted in Indian ink. Scale bars: c=50μm,
d=5μm, e-g=30μm, h-j=5μm, k=10μm
Fungal Diversity
Fig. 114 Neosetophoma clematidis (holotype) a, b Conidiomata on the host plant c Cross section of conidioma d, e Pycnidium wall f-i Different stages
of conidiogeny j-m Conidia n Germinating conidia. Scale bars: a=500μm, b=200μm, c=100μm, d-e=50μm, f=5μm, g-m=10μm
brown to dark brown, with 4–10 cell layers, inner layer hyaline,
3–5 cell layers, cells of textura angularis. Conidiophores reduced to conidiogenous cells. Conidiogenous cells 3–5×1–
3 μm, simple, smooth, hyaline, integrated, enteroblastic,
phialidic. Conidia 11–15×2–4μm (x=12.71×2.95μm, n=20),
slightly brown, straight to curve, continuous, with truncate base, with obtuse apex, 3-euseptate, often slightly narrower in the
middle, smooth-walled, guttulate.
Culture characters: on PDA white from above and greyish
white from reverse, cottony, circular, zonate, slow growing,
attaining a diam. of 1.5 cm in 7 days at 18 °C.
Material examined: Italy, Province of Forlì-Cesena,
Pietrapazza, Bagno di Romagna, on dead branch of Clematis
vitalba L. (Ranunculaceae), 20 January 2013, E. Camporesi,
IT1027 (MFLU 14–0746, holotype); ibid., GUHC 7204,
isotype), ex-type living culture, MFLUCC 13–0734, GUCC
Fungal Diversity
20. GenBank ITS: KP744450; LSU: KP684153; SSU:
KP684154.
Notes: The phylogenetic tree showed that this species belongs to Neosetophoma and is a different species to N. italica
and N. samarorum (Fig. 111).
83. Neosetophoma italica W.J. Li, Camporesi & K.D.
Hyde, sp. nov.
Index Fungorum number: IF550920, Facesoffunginumber:
FoF 00425; Fig. 115
Etymology: Named after the country where it was collected, Italy.
Holotypus: MFLU 14–0809
Saprobic on dead leaves of Iris germanica L., forming
conspicuous, dark brown conidiomata. Sexual morph
Undetermined. Asexual morph coelomycetous.
Conidiomata 50–60μm high, 40–75μm diam., pycnidial, separate, dark brown, globose, subepidermal, unilocular, thinwalled, papillate. Peridium 5–10μm wide, composed of 3–4
layers, with outer 1–2 layers comprising brown and inner 1–2
layers of pale brown to hyaline cells of textura angularis.
C o n i d i o p h o re s r e d u c e d t o c o n i d i o g e n o u s c e l l s .
Conidiogenous cells 2 –5 μ m l o n g × 3–6 μ m w i d e ,
enteroblastic, phialidic, doliiform to ampulliform, determinate, hyaline, smooth-walled. Conidia 6–11×3–4μm (x=9×
3; n=30), subcylindrical, fusiform, or ellipsoid to fusiform,
pale brown, smooth-walled, 1–2-euseptate, constricted at the
septa, apex and base obtuse, or sometimes with truncate base,
smooth-walled.
Culture characters: Colonies on PDA, reaching 30–35 mm
diam. after one month, slow growing, white to yellowish in
the first week, with central area becoming grey after three
weeks, circular, dense, aerial, reverse dark brown, filamentous. Colonies on MEA, pale grey to almost white, reverse
dark brown, without any pigments produced.
Material examined: ITALY, Province of Forlì-Cesena [FC],
Castrocaro Terme, on dead leaf of Iris germanica L.
(Iridaceae), 20 September 2012, E. Camporesi IT–733
(MFLU 14–0809, holotype), ex-type living cultures,
MFLUCC 13–0388, ICMP. GenBank ITS: KP711356; LSU:
KP711361; SSU: KP711366; ibid. (KUN! HKAS 83968,
isotype).
Notes: Neosetophoma, is typified by N. samararum
(Desm.) Gruyter et al. and is monotypic (Gruyter et al.
2010). The species has been reported as a pathogen causing
leaf spots of various hosts (Phookamsak et al. 2014). The
genus is characterized by globose to irregular conidiomata,
with papillate ostioles, and with yellowish conidia that are
attenuate at one end (Gruyter et al. 2010). The morphological
characters of N. italica are in accordance with the generic
concepts of Neosetophoma, and the only difference between
N. italica and N. samararum is the colour of the conidia (pale
brown in N. italica, versus yellowish in N. samararum).
Phylogenetic analyses based on multi-genes (Fig. 111) show
that N. italica groups with two strains of N. samararum with
high bootstrap support (93 %). However, N. italica is distinct
from N. samararum strains, forming a distinct branch.
Therefore Neosetophoma italica is introduced as novel species.
84. Phaeosphaeria musae Arzanlou & Crous., Fungal
Planet 9: (2006)
Facesoffungi number: FoF00597; Fig. 116
Ascomata immersed, mostly erumpent, raising the host tissue, visible as black pustules on surface view, subglobose,
brown to black, solitary, clustered or scattered; in section
120–240 μm high ×150–310 μm diam., rounded at base.
Ostiole central, brown to black, papillate, aperiphysate layers.
Peridium thin, comprising several layers of brown walled angular pseudoparenchymatous cells. Asci 60–80×9–12 μm, 8spored, bitunicate, fissitunicate, cylindrical-clavate, short pedicellate, apically rounded, with an ocular chamber. Ascospores
20–25×4–6 μm, biseriate, ellipsoidal, fusiform to narrowly fusiform with narrowly rounded ends, 5-celled, euseptate, lightly
constricted at the septum, guttulate at each cell, hyaline when
young, turning light brown and concolourous while in the
ascus, wall smooth, surrounded by a mucilaginous sheath.
Material examined: THAILAND: Chiang Mai Province,
Doi Inthanon National Park, junction of Highway 1009 and
road to Mae Chen, N19°31.58′ E 98°29.64′, elev. 1700 m, on a
dead leaf of palms. 21, Sep. 2009. Jian-Kui Liu, JKA-0021
(MFLU 15-0043), living culture MFLUCC 11-0015.
GenBank ITS: KP744458; LSU: KP744502; SSU:
KP753963.
Notes: Phaeosphaeria musae (MFLUCC 11-0015) clustered in the Phaeosphaeria sensu stricto clade and could not
be well-resolved. Phookamsak et al (2014) showed that this
group is a complex section in Phaeosphaeriaceae, and protein
coding genes, such as TEF1 and RPB2 are useful for resolving
the genus complex, or even the species complex. In this study
the taxon was isolated from leaves of palms as the sexual
morph, whereas it was first described from Musa sp. based
on its asexual morph.
85. Wojnowicia dactylidicola Wijayawardene, Camporesi
& K.D. Hyde, sp. nov.
Index Fungorum number: IF550906, Facesoffungi
number: FoF00415; Fig. 117
Etymology: Named after the host Dactylis.
Holotypus: MFLU 14–0743
Saprobic on branch of Dactylis sp. Sexual morph
Undetermined. Asexual morph Conidiomata 70–100μm
diam., 100–130μm high, pycnidial, superficial, black, gregarious to solitary, unilocular, with an eccentric papillate ostiole.
Pycnidial wall 18–20μm wide, with thick outer layer, dark
brown, 5–6 cell layered, with inner most layer hyaline, 2–3
Fungal Diversity
Fig. 115 Neosetophoma italica (holotype) a Specimen b, c Brown
conidiomata on the host surface d Vertical section of conidioma e
Section of peridium f–i Conidiogenous cells and developing conidia j
Germinated spore k–o Conidia p Culture on PDA. Scale bars: c=
200μm, d=25μm, e=5μm, f–i=5μm, j–o=5μm, p=25 mm
Fungal Diversity
Fig. 116 Phaeosphaeria musae.
a Appearance of ascomata on the
host surface. b Section of ascoma.
c Peridium. d, e-i Asci e-g, j.
Ascospores a-i In water. j In India
ink. Scale bars: a=500μm. b, c,
d=30μm. e-g=10μm. h, i=
20μm. j=10μm
cell layered, with cells of textura angularis. Conidiophores
reduced to conidiogenous cells. Conidiogenous cells 1–2.5×
1.5–3 μm, simple, smooth, hyaline, short, integrated, holoblastic to phialidic. Conidia 25–35×3.5–6.5μm (x=28.38×
4.87μm, n=20), pale brown, fusiform to cylindrical, straight
to curved, with obtuse apex and base, 3–5-septate, discrete,
smooth-walled, guttulate.
Culture characters: on PDA olive brown from above and
brown from reverse, with thin mycelium, zonate, slow growing, attaining a diam. of 2.5 cm in 7 days at 18 °C.
Material examined: Italy, Province of Forlì-Cesena,
Fiumicello, Premilcuore, on dead branch of Dactylis glomerata
L. (Poaceae), 15 March 2013, E. Camporesi, NNW IT1259
(MFLU 14–0743, holotype); ibid., (GUHC (Guizhou
University Herbarium Collection) 7201, isotype), ex-type
living culture, MFLUCC 13–0738, GUCC 11. GenBank ITS:
KP744469; LSU: KP684147; SSU: KP684148.
Notes: Saccardo (1892) established Wojnowicia, with W.
hirta as the type species. Currently only four species are accepted (Sutton 1980; Farr and Bills 1995; Wijayawardene et
al. 2013). In conidial dimension, our collection is similar to
Wojnowicia colluvium D.F. Farr & Bills (19–36×3–4μm).
However, Wojnowicia colluvium only has 3 septa, while our
collection has 3–5 septa. Multi-gene analysis (Fig. 111) shows
that the new strain is distinct from other Wojnowicia species,
thus we introduce a new species, W. dactylidicola, to accommodate our collection.
86. Wojnowicia dactylidis Wijayawardene, Camporesi and
K.D. Hyde, sp. nov.
Fungal Diversity
Fig. 117 Wojnowicia dactylidicola (holotype). a, b Conidiomata on host material c Cross sections of the pycnidium d Pycnidial wall e-g Conidia
attached to conidiogenous cell h-l Conidia. Scale bars: b=200μm, c=50μm, c=50μm, e-g=10μm, h-l=15μm
Index Fungorum number: IF550910, Faces of Fungi
number: FoF00417; Fig. 118
Etymology: Named after the generic name of host it occurs
Holotypus: MFLU14–0745
Saprobic on grass. Sexual morph Undetermined. Asexual
morph Conidiomata 200–250μm diam., 120–150μm high,
pycnidial, superficial, black, solitary to gregarious, unilocular,
with a central, papillate ostiole. Pycnidial wall 10–20μm, with
thick outer layer, brown to dark brown, with 5–7 cell layers,
inner layer hyaline, 2–3 cell layered, with cells of textura
angularis. Conidiophores reduced to conidiogenous cells.
Conidiogenous cells 2–4×2–3μm, simple, smooth, hyaline,
enteroblastic, phialidic. Conidia 35–40 × 4–5.5 μm (x=
37.05×5.45 μm, n=20), pale brown, fusiform to cylindrical,
straight to curved, with obtuse apex and base, 7–11-septate,
smooth-walled.
Culture characters: on PDA olive brown from above and
dark brown from reverse, cottony, circular, zonate, slow growing, attaining a diam. of 2 cm in 7 days at 18 °C.
Material examined: Italy, Province of Forlì-Cesena,
Teodorano, Meldola, on dead branch of Dactylis glomerata
L. (Poaceae), 7 March 2013, E. Camporesi, NNW IT1101
(MFLU 14–0745, holotype); ibid., GUHC 7203, isotype),
ex-type living culture, MFLUCC 13–0735, GUCC 9.
GenBank ITS: KP744470; LSU: KP684149; SSU:
KP684150.
Notes: Wojnowicia dactylidicola was also reported from
Dactylis glomerata. The species has smaller conidia and only
3–5 transverse septa, while our new collection has larger conidia (35–40×4–5.5μm) and 7–11 septa. Phylogenetic analysis showed Wojnowicia dactylidicola and W. dactylidis to be
distantly related (Fig. 111). Thus, based on both morphological and molecular characters our new collection is introduced
as a new species, W. dactylidis.
87. Wojnowicia lonicerae Wijayawardene, Camporesi &
K.D. Hyde, sp. nov.
Index Fungorum number: IF550907, Facesoffungi
number: FoF00416, Fig. 119
Fungal Diversity
Fig. 118 Wojnowicia dactylidis (holotype) a Host material b Conidiomata on host surface c, d Cross section of pycnidia e-h Developing conidia
attached to conidiogenous cells i-n Conidia. Scale bars: b=200μm, c=100μm, d-e=50μm, f-n=20μm
Etymology: Named after the generic name of host it occurs
Holotypus: MFLU 14–0744
Saprobic on branch of Lonicera sp. Sexual morph
Undetermined. Asexual morph Conidiomata 150–175μm
diam., 120–130μm high, pycnidial, superficial, black, solitary,
unilocular, with a central, papillate ostiole. Pycnidial wall 15–
30μm wide, with thick outer layer, dark brown, with 5–8 cell
layers, inner layer hyaline, 2–3 layered, with cells of textura
angularis. Conidiophores reduced to conidiogenous cells.
Conidiogenous cells 1–3×1.5–2μm, simple, smooth, hyaline,
enteroblastic, phialidic. Conidia 38–49×5–6μm (x=42.34×
5.45μm, n=20), pale brown, fusiform to cylindrical, straight
to curved, with obtuse apex and base, 8–11-septate, constricted at the septa.
Culture characters: on PDA olive brown from above and
greyish-brown from reverse, cottony, circular, zonate, slow
growing, attaining a diam. of 3 cm in 7 days at 18 °C.
Material examined: Italy, Arezzo Province, Pieve Santo
Stefano, on dead branch of Lonicera sp. L. (Caprifoliaceae),
15 March 2013, E. Camporesi, IT1246 (MFLU 14–0744,
holotype); GUHC 7202, isotype), ex-type living culture,
MFLUCC 13–0737, GUCC 10. GenBank ITS: KP744471;
LSU: KP684151; SSU: KP684152.
Notes: In conidial length, Wojnowicia lonicerae is similar to
W. hirta (31–44×3–4μm), but the former has narrower conidia.
Wojnowicia lonicerae has 8–11 transverse septate; whereas W.
hirta has only 7–8 transverse septa. Multi-gene analyses shows
Wojnowicia lonicerae grouping with W. dactylidis, but with low
Fungal Diversity
Fig. 119 Wojnowicia lonicerae (holotype). a Host material b Conidiomata on the host c Cross section of pycnidium d Pycnidium wall e-g Conidia
attach to conidiogenous cell h-l Conidia. Scale bars: b=200μm, c=100μm, d=50μm, e-g=10μm, h-l=20μm
bootstrap support. However, conidial morphology of our new
strain and W. dactylidis is distinct. Hence we introduce a new
species, W. lonicerae, to accommodate our collection.
Pleomassariaceae
The family Pleomassariaceae comprises saprobes or pathogens on wood or lichens in terrestrial environments. The
family was introduced by Barr (1979a) and included the generic type Pleomassaria, plus Asteromassaria and
Splanchnonema. Pleomassaria differs from the other two genera in producing muriform and somewhat asymmetrical ascospores with a submedian primary septum. Eopyrenula
(Aptroot 1991), Kirschsteiniothelia, Macrovalsaria (Barr
1993a) and Lichenopyrenis (Calatayud et al. 2001) have also
been added. Currently the family comprises Lichenopyrenis,
Fungal Diversity
Splanchnonema, Peridiothelia and Pleomassaria (Zhang et
al. 2012a). Pleomassaria was treated as a synonym of
Splanchnonema (Barr 1993) based on their similar characteristics, but this was not followed by many researchers.
The asexual morphs of the Pleomassariaceae are mostly
coelomycetous. Pleomassaria siparia, the type of this family
has been linked with Prosthemium betulinum. The asexual
morphs of Pleomassaria are Prosthemium and Shearia (Barr
1982; Sivanesan 1984; Tanaka et al. 2010). Pleomassariaceae
was treated as a synonym of Melanommataceae based on
molecular studies (Zhang et al. 2009a). In the study of
Zhang et al. (2012a) Pleomassaria siparia and four
Prosthemium species formed a well-supported monophyletic
clade, thus proving Pleomassariaceae as a separate family in
the order Pleosporales. The phylogenetic tree based on the
LSU is presented in Fig. 120.
88. Splanchnonema pupula (Fr.) Kuntze, Revis. gen. pl.
(Leipzig) 3(2): 531 (1898)
Stigmatomassaria pupula (Fr.) Munk, Dansk bot. Ark.
15(no. 2): 127 (1953)
Facesoffungi number: FoF00475; Fig. 121
Saprobic on dead branches of Acer pseudoplatanus.
Sexual morph Ascomata 437–521μm high×672–882μm
diam. (x= 482 × 717 μm, n = 10), solitary, immersed, dark
brown to black, slightly depressed, uniloculate, globose to
subglobose, ostiole central, papilla projecting through the
bark. Peridium 35–47μm wide, composed of 4–5 layers of
brown to black-walled cells of textura angularis.
Hamathecium comprising 1.5–2.5μm wide, hyaline, septate,
branched pseudoparaphyses, embedded in a gelatinous matrix. Asci 118–152×31–36μm (x=134×32μm, n=20), 8spored, bitunicate, fissitunicate, cylindrical or fusoid, pedicellate, rounded at apex, with a distinct ocular chamber.
Ascospores 37–51×11–18μm (x=44×14μm, n=20), partially
overlapping to biseriate, hyaline to light brown, always brown
at maturity, 1-septate in early states, constricted at the septum,
muriform, becoming 3-euseptate, somewhat asymmetrical, ellipsoidal with broadly rounded ends, thick-walled, smoothwalled, surrounded by mucilaginous sheath. Asexual morph
Undetermined.
Material examined: ITALY, Province of Monte Secchieta,
P r a t o m a g n o , A r e z z o , o n d e a d b r a nc h e s o f A c e r
pseudoplatanus, (Sapindaceae), 5 June 2012, E. Camporesi
(MFLU 14–0807, holotype). The DNA was extracted from
the fungal fruiting body. GenBank ITS: KP659196; LSU:
KP659197.
Notes: This collection is identified as Splanchnonema
pupula; however, there is no sequence data in GenBank for
the type of this genus, S. pustulatum. Splanchnonema platani
is a synonym of Macrodiplodiopsis desmazieresii and belongs
near Bambusicolaceae and is not Lophiostomataceae (sensu
Wijayawardene et al. 2014). Other species of Splanchnonema
are placed in Massarinaceae and Pleosporaceae (Fig. 120),
and thus the genus is polyphyletic. As there is no type sequence data it is not clear which family Splanchnonema sensu
stricto belongs in. Stigmatomassaria pupula is the type species of Stigmatomassaria and this may be a good generic
name for this species in Pleomassariaceae, however until
the type species of Splanchnonema is made available we name
it as Splanchnonema pupula.
Pleosporaceae
Pleosporaceae is a family in the order Pleosporales, with
Pleospora herbarum (Pers.) Rabenh. as the generic type.
The family was revised by Ariyawansa et al. (2015) and 18
genera accepted. Phylogenetic analysis of 45 strains
representing 24 sections from the Alternaria species complex,
including our new sexual strains, indicated it was a new species. The novelty of Alternaria cesenica as new species receives strong support in the phylogenetic tree (Fig. 122).
89. Alternaria cesenica Phukhamsakda, Qing Tian,
Camporesi & K.D. Hyde, sp. nov.
Index Fungorum number: IF550899, Facesoffungi
number: FoF 00408; Fig. 123.
Etymology: The specific epithet Cesenica refers to the
place in which the fungus was first collected.
Holotypus: MFLU 14–0795
Saprobic on dead branch of Bellevalia romana (L.) Sweet.
Sexual morph Ascomata 100–200μm high, 84–225μm diam.
(x=170×160μm, n=10), on the surface of the host, solitary,
scattered, or sometimes gregarious, globose, superficial,
black, smooth. Peridium 22–33.5μm (x=28.5μm, n=6), at
the outside composed of irregular, thick-walled, brown to
black cells of textura angularis, inner layer composed of
slightly, smaller cells of textura angularis. Asci 68–104×
11–14μm (x=85×12μm, n=10), 6–8-spored, bitunicate, clavate to subellipsoid, short pedicellate, apically rounded, with
an ocular chamber. Hamathecium comprising 1–2μm broad,
septate, long, branched or simple, pseudoparaphyses.
Ascospores 16–23×8–10μm (x=19×9μm, n=10), uniseriate
or overlapping uniseriate, ellipsoid to fusiform, muriform,
dark brown to blackish-brown, constricted at the center, lower cell narrow and longer, smooth-walled. Asexual morph
Undetermined.
Culture characters: Colonies on PDA, reaching 2 cm diam.
after 2 weeks at 16 °C, later with dense mycelium, circular,
margin rough, white, surface flat, without aerial mycelium.
Hyphae septate branched, hyaline, thin.
Material examined: ITALY, Forlì-Cesena Province:
Fiumana, Predappio, dead and hanging branches of
Bellevalia romana (Asparagaceae), 7 February 2013, E.
Camporesi IT1052 (MFLU 14–0795, holotype); ex-type living culture, MFLUCC 13–0450. GenBank ITS: KP711383;
LSU: KP711384; SSU: KP711385; TEF: KP711386.
Fungal Diversity
Fig. 120 Phylogram generated from Maximum likelihood (RAxML)
analysis based on LSU sequence data of Pleosporales. Maximum
likelihood 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-types
(reference strains) are in bold; the new isolates are in blue. The tree is
rooted with Aigialus grandis BCC 18419
Fungal Diversity
Fig. 121 Splanchnonema
pupula (MFLU 14–0807). a
Ascomata on host substrate b
Transverse section of ascomata c
Immature asci d Immature ascus e
Immature ascospores showing
primary septum f, g Mature ascus
h, i Hyaline ascospores j Mature
brown ascospore. Scale bars: b=
400μm, c=200μm, d–g=80μm,
h, j=30μm
Notes: Alternaria is a genus of pathogenic and saprobic
taxa (Woudenberg et al. 2013). The genus was originally introduced by Nees (1816) as Alternaria tenuis, which is currently treated as a synonym of Alternaria alternata
(Woudenberg et al. 2013). Comoclathris has been reported
as a sexual morph of Alternaria and has muriform, yellowish
to dark brown ascospores. Comoclathris clusters within
Alternaria in phylogenetic analyses (Woudenberg et al.
2013). Molecular analysis provides evidence that Alternaria
cesenica is a new species grouping in section infectoriae
Fungal Diversity
Fig. 122 Phylogram generated
from parsimony analysis based on
LSU, SSU, ITS and TEF1
sequence data of Pleosporaceae.
Parsimony bootstrap support
values greater than 50 % are
indicated above the nodes, and
branches with Bayesian posterior
probabilities greater than 0.95 are
given in bold. The ex-types
(reference strains) are in bold; the
new isolates are in blue. The tree
is rooted with Pleospora
herbarum CBS 191.86
(relatively high bootstrap support of 88 % from RAxML and
the Bayesian PP of 1.00, Fig. 122). The sexual morph is rarely
found in Alternaria.
Pleosporales genera incertae sedis
Camarosporium
Camarosporium was introduced by Schulzer (1870) with
Camarosporium quaternatum (Hazsl.) Schulz. as the
type species. The phylogenetic tree is presented in
Fig. 124.
90. Camarosporium aborescentis Phukhamsakda,
Bulgakov & K.D. Hyde, sp. nov.
Index Fungorum number: IF550911, Facesoffungi
number: FoF00406; Fig. 125
Etymology: The specific epithet arborescentis refers to the
host species Colutea arborescens and latin entis means residing that reference to the host from which the fungus was first
collected.
Fungal Diversity
Fig. 123 Alternaria cesenica (holotype) a, b Ascomata on host C Section of ascomata d Peridium e Pseudoparaphyses f Immature ascus. g, h Asci with
ascospores i Fissitunicate ascus j-l. Ascospores. Scale bars: b=500μm, c=200μm, d=100μm, e-i=10μm, j-l=5μm
Holotypus: MFLU 14–0793
Saprobic on dead branch of Colutea arborescens L. Sexual
morph Undetermined. Asexual morph Conidiomata 490–
770 μm diam., 350–600 μm high, pycnidial, immersed,
erumpent at maturity, solitary to gregarious, scattered, unilocular, dark brown, papillate ostiole. Pycnidial wall 32–54μm (–
70μm at apex), multi-layered, with 4–6 layers, comprising of
brown walled cells of textura angularis. Conidiophores reduced to conidiogenous cells. Conidiogenous cells
enteroblastic with percurrent annelidic, discrete, doliiform,
hyaline, brown when mature, smooth-walled, formed from
the inner layer of pycnidium wall. Conidia 12–25×5–13μm
(x=20.3×9μm, n=50), oblong, ellipsoid, straight or rarely
slightly curved, narrowly rounded at both ends, muriform,
with 1–3 transverse septa, and 1–4 longitudinal septa, initially
hyaline, pale brown to brown at maturity, smooth-walled.
Culture characters: Colonies on PDA reaching 25–33 mm
diam. after 1 week at 16 °C, later with dense mycelium, circular, fimbriate, margin rough, white at first, olive-green after
5 days in the centre of the colony, convex with papillate on the
surface, without aerial mycelium. Hyphae septate, branched,
hyaline.
Fungal Diversity
Fig. 124 Phylogram generated
from Maximum likelihood
(RAxML) analysis based on
combined LSU, SSU and ITS
sequence data of Pleosporales.
Maximum likelihood 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-types (reference
strains) are in bold; the new
isolates are in blue. The tree is
rooted with Montagnula
anthostomoides CBS 615.86
Material examined: RUSSIA, Rostov Province, Rostovna-Donu, Badia Tega, Botanical Garden of Southern Federal
University, Systematic Arboretum, on dead twigs of Colutea
arborescens L. (Fabaceae), 8 May 2014, T. Bulgakov T44
(MFLU 14–0793, holotype); ex-type living culture,
MFLUCC 14–0604. GenBank ITS: KP711377; LSU:
KP711378; SSU: KP711379.
Notes: Our collection is morphologically similar to
Camarosporium feurichii which has erumpent, black
conidiomata and brown, smooth-walled, oblong, 3
transverse septa conidia, usually with one longitudinal septum (Eille and Eille 1985). Molecular data analyses of combined SSU, LSU and ITS sequence data (Fig. 124) show
our strain cluster with Camarosporium sensu stricto which
is a distinct lineage in Pleosporinae, Pleosporales
(Wijayawardene et al. 2014).
91. Camarosporium aureum Norphanphoun, Bulgakov &
K.D. Hyde, sp. nov.
Index Fungorum number: IF550905, Facesoffungi
Number: FoF 00551; Fig. 126
Fungal Diversity
Fig. 125 Camarosporium aborescentis (holotype) a, b Conidiomata on
surface of twigs c Side view of conidiomata erumpent on surface of host d
Vertical section of multi-locular conidiomata e Vertical section of
unilocular conidiomata f Part of pycnidial wall g-j Conidiogenous cells
and developing conidia k-l Immature conidia m-o Conidia at maturity.
Scale bars: b-d=500μm, e=200μm, f=50μm, g–o=10μm
Etymology: From the Latin, aureum meaning gold, in reference to the colour nature of the conidia.
Holotype: MFLU 14–0770
Saprobic on dead herbaceous branches of Cotinus
coggygria (Scop.). Sexual morph Undetermined. Asexual
morph Conidiomata 400–550μm high, 350–550μm diam.
pycnidial, solitary, immersed, unilocular, dark brown, with a
central ostiole. Pycnidial wall 25–30μm multi-layered, with
4–5 layers of brown, cells of textura angularis, with inner
most layer thin, hyaline. Conidiophores reduced to
conidiogenous cells. Conidiogenous cells enteroblastic,
phialidic, hyaline, smooth, formed from the inner most layer
Fungal Diversity
Fig. 126 Camarosporium aureum (holotype), a Stromatal habit in
wood b, c Fruiting bodies on substrate d Cross section of the stroma
showing perithecia e Peridium f Apex of perithecia g-h Conidiogenous
cell with attached conidia i Immature conidia. j–k Mature conidia. l
Germinating conidia. Scale bars: d=200μm, e=50μm, f=100μm, g–h=
20μm, i–k=10μm, l=30μm
of pycnidium wall. Conidia (19–)–19.5–22.5(–24)×8–9(–10)
μm (x=21×9.5μm, n=30) oblong, mostly straight, infrequently slightly curved, muriform, with 4–5 transverse septa,
with 2–3 longitudinal septa, not constricted at the septa, initially hyaline, pale gold to brown at maturity, narrowly rounded at both ends, smooth-walled.
Culture characters: Colonies on PDA slow growing,
reaching 2 cm diam. after 11 days at 25 °C, later producing dense mycelium, circular, margin rough, white at first,
greenish-yellow after 5 days, flat or effuse on surface,
without aerial mycelium. Hyphae septate branched, hyaline, thin.
Fungal Diversity
Material examined: RUSSIA, Rostov region, Shakhty city,
near Grushevsky pond, shelterbelt artificial forest, on dead
branches of Cotinus coggygria Scop. (Anacardiaceae) 18
March.2014, T. Bulgakov (MFLU 14 – 0770, holotype); ibid.
(PDD isotype); ex-type culture, MFLUCC 14–0620.
GenBank ITS: KP744436; LSU: KP744478; SSU:
KP753948.
Notes: Sutton (1980) reported the type species of
Camarosporium, C. quaternatum from Lycium halimifolium,
which was characterized by fusiform conidia, with 3–4 transverse septa, and several secondary longitudinal or oblique
septa. Our new species has oblong, larger conidia and 4–5
transverse septa, with 2–3 longitudinal septa and is golden
brown at maturity. Molecular phylogenetic analysis of combined ITS and LSU gene data places C. aureum close to C.
spartii Trail from Cytisus sp. (Fig. 124). However, C. spartii
has shorter and narrower conidia (13–16×6–7μm).
92. Camarosporium caraganicola Phukhamsakda,
Bulgakov & K.D. Hyde, sp. nov.
Index Fungorum number: IF550912, Facesoffungi
number: FoF00407; Fig. 127
Etymology: The specific epithet caraganicolarefers to the
origin of host species Caragana frutex and latin cola meaning
originating, in reference to the host from which the fungus was
first isolated.
Holotypus: MFLU 14–0794
Saprobic on dead branch of Caragana frutex L. Sexual
morph Undetermined. Asexual morph Conidiomata 280–
780μm diam., 413–604μm high, pycnidial, erumpent at maturity, confluent, gregarious, dark brown to black, papillate
ostiole, lacking periphyses. Pycnidial wall 15–45 μm (–
50μm at apex), multi-layered, with 5–6 layers of brown cells
o f te x tur a angular is. Coni diophore s r e d u c e d t o
conidiogenous cells. Conidiogenous cells enteroblastic with
percurrent annelidic, doliiform, discrete, solitary, hyaline,
brown when mature, smooth-walled, and formed from the
inner layer of pycnidium wall. Conidia 13–26×6–13μm (x=
19.5×9.8μm, n=50), oblong, ellipsoid, straight or occasionally curved at the apex, muriform, with 1–4 transverse septa,
with 1–4 longitudinal septa, initially hyaline, pale brown to
brown at maturity, narrowly rounded at both ends, smoothwalled.
Culture characters: Colonies on PDA reaching 3 cm diam.
after 1 week at 16 °C, later with dense mycelium, circular,
margin rough, white at first, dark green after 5 days in the
centre of the colony, surrounded by white mycelium, flat on
the surface, without aerial mycelium. Hyphae septate
branched, hyaline.
Material examined: RUSSIA, Rostov Province, Rostovna-Donu, BadiaTega, Botanical Garden of Southern Federal
University, Systematic Arboretum, on dead twigs of
Caraganafrutex (L). (Fabaceae), 26 April 2014, T.
Bulgakov T46 (MFLU 14–0794, holotype); ex-type living
culture, MFLUCC 14–0605. GenBank ITS: KP711380;
LSU: KP711381; SSU: KP7113824.
Notes: Camarosporium caraganae is recorded from
Caragana spp. and has brown conidia with 3–5 transverse
septa and 1(−2) longitudinal septa (Karst 1885). Our collection Camarosporium caraganicola was discovered on C.
frutex shows unique conidia morphology with 1–4 transverse,
and 1–4 longitudinal septa. The molecular analysis of combined SSU, LSU and ITS data (Fig. 124) shows a close relationship with C. aborescentis. Morphologically, C.
caraganicola and C. aborescentis are similar but colonize
different hosts.
93. Multiseptospora Phookamsak & K.D. Hyde, gen. nov.
Index Fungorum number: IF550928, Facesoffungi
number: FoF00430
Etymology: The generic epithet Multiseptospora refers to
the phragmosporous ascospores.
Type species: Multiseptospora thailandica Phookamsak &
K.D. Hyde
Saprobic on Poaceae. Sexual morph Ascomata solitary to
gregarious, immersed, uni-loculate, globose to subglobose,
setose, ostiole central, with pore-like opening. Peridium
thick-walled, of unequal thickness, thick at the base, composed of black, pseudoparenchymatous cells, textura
angularis to textura prismatica. Hamathecium composed of
dense, narrow, cellular pseudoparaphyses. Asci 8-spored,
bitunicate, fissitunicate, broadly cylindrical, subsessile, apically rounded with indistinct ocular chamber. Ascospores
overlapping 2–3-seriate, fusiform to vermiform, with acute
ends, hyaline to pale brown when young, becoming brown
when mature, 10–11-septate, slightly curved, slightly constricted at the septum, smooth-walled, surrounded by distinct
mucilaginous sheath. Asexual morph Undetermined.
Notes: Multiseptospora is a monotypic genus introduced to
accommodate the fungus associated with Thysanolaena
maxima. The genus is similar to Falciformispora K.D.
Hyde in its ascospores, however, they differ with regard
to their ascomata and host (Zhang et al. 2012a, b).
Multiseptospora forms setose ascomata with pore-like
openings, immersed in the host, while Falciformispora
forms epapillate, glabrous coriaceous, erumpent to superficial, ascomata, (Zhang et al. 2012a, b). Based on multigene phylogenetic analyses, Multiseptospora forms a sister clade basal to Trematosphaeriaceae (Fig. 54).
Therefore, we introduce a new genus to accommodate a
single species and placed it in Pleosporales, genera
incertae sedis.
94. Multiseptospora thailandica Phookamsak & K.D.
Hyde, sp. nov.
Index Fungorum number: IF550929, Facesoffungi
number: FoF00431; Fig. 128
Fungal Diversity
Fig. 127 Camarosporium caraganicola (holotype) a, b Conidiomata
on host surface c Gregarious conidiomata on host surface d Vertical
section of multi-loculate conidiomata e Ostioles f Part of pycnidium
wall g-i Conidiogenous cells and developing conidia j-l Conidia when
immature m-o Mature conidia. Scale bars: b=500μm, c-e=200μm, f=
100μm, g-h=20μm, i–o=10μm
Etymology: The specific epithet “thailandica” refers to the country where the taxon was first
collected.
Holotypus: MFLU 11–0219
Saprobic on Thysanolaena maxima. Sexual morph
Ascomata 180–260μm high, 210–320μm diam., solitary to
gregarious, scaterred, immersed, visible as dark spots with
sparse, with tufts of hyphae on host surface, uni-loculate, globose to subglobose, setose, ostiole central, with pore-like
opening, obtuse at the apex. Peridium 10–60μm wide, with
thin to thick walls, of unequal thickness, slightly thick at the
base and sides, composed of several layers of dark brown to
black cells, oganized in a textura angularis to textura
prismatica. Hamathecium composed of dense, 0.8–1.5μm
Fungal Diversity
Fig. 128 Multiseptospora thailandica (holotype) a Ascomata on host surface b Section through an ascoma c Peridium d Pseudoparaphyses e–i Asci j–
n Ascospores o Ascospore at maturity. Scale bars: b=100μm, e=50μm, c-i=20μm, j-o=10μm
wide, narrow cellular, distinctly septate pseudoparaphyses not
constricted at the septa, anastomosing above the asci, embedded in a hyaline gelatinous matrix. Asci (77–)80–95(–
110) × (18–)19–22.5 μm (x= 85 × 20 μm, n = 25), 8-spored,
bitunicate, fissitunicate, broadly cylindrical, short pedicellate,
apically rounded with indistinct ocular chamber. Ascospores
(35–)37–43(–45)×6–8μm (x=40.6×7.3μm, n=30), overlapping 2–3-seriate, fusiform to vermiform, with acute ends,
slighty curved, initially hyaline to pale brown, becoming
brown to dark brown at maturity, 10– to 11-septate, slightly
Fungal Diversity
constricted at the septa, smooth-walled, surrounded by thin,
distinct, mucilaginous sheath. Asexual morph
Undetermined.
Culture characters: Colonies on PDA fast growing, 70–
80 mm diam. after 4 weeks at 25–30 °C, brown to orangishbrown at the margin, dark brown to dark grey in the centre;
reverse brown to orangish-brown at the margin, dark grey in
the middle, black at the centre; dense, circular, slightly raised
to umbonate, smooth to dull with entire edge, concave at the
centre, fluffy to floccose, aspect smooth, producing brown
pigments in agar.
Material examined: THAILAND: Chiang Mai, Doi
Suthep-Pui, on dead stem of Thysanolaena maxima Kuntze
(Poaceae), 13 Febuary 2011, R. Phookamsak RP0099
(MFLU 11–0219, holotype), ex-type living culture,
MFLUCC 11–0183. GenBank ITS: KP744447; LSU:
KP744490; SSU: KP753955.
95. 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)
Type species: Saccothecium sepincola (Fr.) Fr. [as
‘saepincola’], Summa veg. Scand., Section Post.
(Stockholm): 398 (1849)
≡ Sphaeria sepincola Fr. [as ‘saepincola’], Observ.
mycol. (Havniae) 1: 181 (1815)
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 (2014) synonymized Dothioraceae under
Dothideaceae based on morphology and molecular phylogeny, and also 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). Nevertheless, Aureobasidiaceae should be
synonymized under Saccotheciaceae because the latter is the
oldest available name for the family that contains
Aureobasidium and Saccothecium.
Venturiales Y. Zhang et al., Fungal Diversity 51(1): 251
(2011)
Venturiales was excluded from the Pleosporales and introduced as a new order by Zhang et al. (2011a). It presently
includes Sympoventuriaceae and Venturiaceae, families which
are supported by morphological characteristics and strong support in multi-gene phylogenetic analyses (Hyde et al. 2013).
Venturiaceae E. Müll. & Arx ex M.E. Barr, Mycologia 71(5):
947 (1979)
The family Venturiaceae was introduced Müller and von
Arx (1950) and validated by Barr (1979). Genera in
Venturiaceae are usually characterized by yellowish, greenishbrown to brown, two-celled ascospores and obclavate asci
(Barr 1968, 1989; Wu et al. 2011). Members of Venturiaceae
are distinct and distantly related to other members of
Pleosporales and have a unique set of morphological and ecological characters (Kruys et al. 2006; Zhang et al. 2012a, 2012a,
b), as it is confirmed that Venturiaceae was excluded from the
Pleosporales by Zhang et al. (2011a) and placed in
Venturiales based on strong support of multi-gene phylogenetic
analyses.
Tothia Bat., Annls hist.-nat. Mus. natn. hung. 52: 105 (1960).
Index Fungorum number: IF5506, Facesoffungi number:
FoF 00401
Notes: Wu et al. (2011) epitypified the generic type
T. fuscella which clustered outside the family
Microthyriaceae and was related to Venturiales in phylogenetic analysis of LSU and ITS. A New species is
introduced in this genus and the phylogenetic tree is
presented in Fig. 129.
96. Tothia spartii Qing Tian, Camporesi & K.D. Hyde, sp. nov.
Index Fungorum number: IF550898, Facesoffungi
number: FoF00401; Fig. 130
Etymology: The specific epithet spartii refers to the host
genus on which the fungus occurs.
Holotypus: MFLU 14–0739
Saprobic on a dead or living branches of Spartium
junceum. Sexual morph Superficial mycelium absent.
Ascomata (170–)185–250(–270)×(150–)180–(280–) 300μm
(x=232×210.5μm, n=10), superficial on branches, solitary,
scattered, sometimes gregarious, thyriothecial , dome-shaped
or flat-conical, black, membranaceous, opening by a short,
papillate ostiole. Peridium thick at the apex,
18(–25)–(20–)23μm (x=21.5μm, n=5), upper wall comprising irregular-lobed cells, tightly fitting together like a jigsaw
puzzle, one-layered, lower wall poorly developed.
Hamathecium comprising 1–3 μm broad long, hyaline,
pseudoparaphyses, with transverse septa, branched, longer
than asci. Asci 30(–35)–50(–55) × 7(–9)–12(–13) μm
(x=42×10.5μm, n=10), 8-spored, bitunicate, fissitunicate,
obclavate, with a knob-like short-pedicellate, thin-walled.
Ascospores (10–)11–12(–13)×(3–)4–4(–5) μm (x=10.5×
4.2μm, n=10), biseriate or multi-seriate, ellipsoid to fusiform,
1-septate, normally unequal, upper cell slightly wider than the
lower, slightly constricted at the septum, rounded at both
Fungal Diversity
Fig. 129 Phylogram generated
from Maximum Likelihood
(RAxML) analysis based on
combined LSU, SSU, TEF and
RPB2 sequenced data of
Venturiaceae. Maximum
likelihood bootstrap support
values greater than 50 % are
indicated above and below the
nodes, and branches with
Bayesian posterior probabilities
greater than 0.95 are given in
bold. The ex-types (reference
strains) are in bold; the new
isolates are in blue. The tree is
rooted with Schismatomma
decolorans DUKE 0047570
sides, hyaline and becoming light brown when mature, with
four guttules, smooth-walled. Asexual morph Undetermined.
Culture characters: Colonies on PDA erumpent, slow
growing, reaching 1 cm diam. after 20 d, velvety, mycelium black, dense, globose to irregular, in the centre
raised, with water droplets on the surface, reverse dark,
sunken. No asexual morph was produced on PDA after
60 d incubation.
Material examined: ITALY, Province of Forlì-Cesena [FC],
Fiumicello, Premilcuore, on living and dead branches of
Spartium junceum (Fabaceae), 17 March 2012, E. Camporesi
(MFLU 14–0739, holotype); ex-type living culture, MFLUCC
14–0615. GenBank ITS: KR025861; LSU: KR025865.
Notes: Tothia is a monotypic genus which was introduced
to accommodate type species Tothia fuscella (Sacc.) Bat. It is
a poorly known genus forming thyriothecia and classified in
the Microthyriaceae by Lumbsch and Huhndorf (2010).
Tothia shares morphological characters with typical representatives of Microthyriaceae, such as two-celled, guttulate, and
slightly asymmetrical ascospores which resemble
Microthyrium, but differ in being brown. Thyriothecial
ascomata are unusual for the Venturiaceae, however, most
genera in Venturiaceae usually have yellowish, greenish
brown to brown, two-celled ascospores and obclavate asci
which match Tothia well (Barr 1968, 1989). In recent studies,
Tothia was found to be most closely related to the
Venturiaceae based on analysis of LSU and ITS rRNA genes
(Hyde et al. 2013; Wu et al. 2011). In addition, our study
confirmed the placement of Tothia in Venturiaceae and indicated the new collection to be a new species based on analysis
of LSU, SSU, TEF1 and RPB2 genes. Ascospores of T.
fuscella are fusiform or oblong-ellipsoid, tapering at both
sides, while ascospores are ellipsoid to fusiform and rounded
at both sides in T. sparticola.
Eurotiomycetes
Chaetothyriales
The order Chaetothyriales was introduced by Batista and
Ciferri (1962) and validated by Barr (1987); and the family
type is Chaetothyriaceae (Batista and Ciferri 1962). Schoch et
al. (2006) confirmed the placement of Chaetothyriales placement in Eurotiomycetes based on phylogenetic analysis.
Chaetothyriales comprised two families, Chaetothyriaceae
and Herpotrichiellaceae (Geiser et al. 2006; Kirk et al.
2008). Chomnunti et al. (2012a, b) introduced
Trichomeriaceae under Chaetothyriales based on morphology and phylogeny. Réblová et al. (2013) introduced the family
Cyphellophoraceae in Chaetothyriales with only asexual
morphs, which was separated from the Herpotrichiellaceae;
they also transferred Trichomeriaceae to Chaetothyriaceae
based on their phylogenetic tree. Hui et al. (2014) and
C h o m n u n t i e t a l . (2014) , h o w e v e r, a c c e p t e d
Trichomeriaceae as distinct from Chaetothyriaceae because
of its morphological and phylogenetic differences.
Chaetothyriaceae
The family Chaetothyriaceae was introduced by Hansford
(1946), with the generic type Chaetothyrium. Species in
Chaetothyriaceae are referred to as sooty moulds because of
the similarity of appearance and ecology with other sooty
moulds in Capnodiaceae (Chomnunti et al. 2012a; b, 2014).
However, these two families are clear distinct in morphology
and phylogeny. Phylogenetic analyses of Chaetothyriaceae
confirmed that this family should be placed in the order
Chaetothyriales, class Eurotiomycetes with strong support
(Schoch et al. 2006; Chomnunti et al. 2012a; b; Hui et al.
Fungal Diversity
Fig. 130 Tothia spartii (holotype) a, b Ascomata on the host c, f Squash mount of ascoma d, e Vertical section of ascoma g-j Asci with ascospores k
Pseudoparaphyses l Germinating ascospore m–p Ascospores. Scale bars: a=1000μm, b=200μm, c-k=20μm, l=50μm, m-p=10μm
2014). A new species of Chaetothyrium is introduced here.
The phylogenetic tree is presented in Fig. 131.
97. Chaetothyrium agathis Hongsanan & K.D. Hyde, sp.
nov.
Index Fungorum number: IF550893, Facesoffungi
number: FoF00396; Fig. 132
Etymology: From the Latin agathis, meaning precious
thing which is the genetic name of the host plant.
Holotypus: MFLU 14–0749
Saprobic on surface of leaves of Agathis. Sexual morph
Ascomata 80×124μm diam. (x=94μm, n=5), rather small,
solitary or clustered (usually solitary), superficial on surface
Fungal Diversity
Fig. 131 Phylogram generated
from Maximum likelihood
(RAxML) analysis based on
combined LSU and ITS sequence
data of Chaetothyriales.
Maximum likelihood 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-types (reference
strains) are in bold; the new
isolates are in blue. The tree is
rooted with Capronia munkii
AFTOL-ID 656
of leaves, subglobose to globose, rounded above, wall composed of pale brown cells, lacking an ostiole. Setae 62–107
long×1.8–2.6 wide μm (x=74×2μm, n=10), dark brown,
rounded at the base and wider than apex, stellate when viewed
under stereomicroscope because of setae arrangement, dark
brown to black. Peridium 21–34μm (x=26μm, n=5) comprising two cell layers of textura angularis, inner layer hyaline,
outer layer brown. Hamathecium lacking paraphyses. Asci
52–43 × 16–20 μm (x= 48 × 19 μm, n = 10), 8-spored,
bitunicate, fissitunicate, broadly ovoid or oblong, usually
short pedicellate, with an ocular chamber. Ascospores 21–
29×4–5μm (x=26×5μm, n=10), 2–3-seriate, oblong to ellipsoidal, with 4–7 trans-septa, small and narrow at both end
cells, hyaline, slightly rough-walled and constricted at septum
in water, rough-walled and not constricted at the septum when
in Melzer’s reagent, smooth-walled and not constricted at the
septum when in cotton blue reagent. Asexual morph
Undetermined.
Material examined: PHILIPPINES, Laguna Province,
Mount Makiling, on leaves of Agathis sp. (Araucariaceae),
February 2012, K.D. Hyde HSA15 (MFLU 14–0749,
holotype); ex-type living culture, MFLUCC 12–0113, CPC
20477. GenBank ITS: KP744437; LSU: KP744480.
Notes: Chaetothyrium agathis is most similar to C.
brischoficola (see Chomnunti et al. 2012a; b) in having setae.
The setae in C. brischoficola are however, short, while those
in C. agathis are long. Ascospores of C. brischoficola are
obovoid, 3–4-septate or muriform, while in C. agathis they
are cylindrical with 3–7-septate.
Trichomeriaceae
The family Trichomeriaceae was introduced by Chomnunti &
K.D. Hyde (2013) based on generic type Trichomerium.
Trichomeriaceae are sooty moulds under the order
Chaetothyriales and similar to Capnodiaceae and
Chaetothyriaceae, but differ in having an apical ascal ring,
ascospores with or without sheath and septation (Chomnunti
et al. 2012a, b; 2014). Réblová et al. (2013) transferred species
of Trichomeriaceae to Chaetothyriaceae based on their phylogenetic tree. In other studies, phylogenetic analysis and morphology demonstrated that the family Trichomeriaceae clusters separately from Capnodiaceae and Chaetothyriaceae
(Chomnunti et al. 2012a, b, 2014; Hui et al. 2014). The family
Trichomeriaceae comprises a single genus, Trichomerium
(Chomnunti et al. 2012a; b; 2014), which contains 27 species
in Index Fungorum (2015). In this paper a new species of
Trichomerium is introduced. The phylogenetic tree is presented in Fig. 133.
98. Trichomerium siamensis Hongsanan & K.D. Hyde,
sp. nov.
Index Fungorum number: IF550885, Facesoffunginumber:
FoF 00395; Fig. 134
Etymology: siamensis is from Latin derived from siam+
ensis which means “of Siam” or “Thailand”.
Holotypus: MFLU 14–0750.
Saprobic on the lower surface of leaves. Superficial
hyphae 3–4μm wide, septate, constricted at the septum,
pale brown to brown. Sexual morph Ascomata 112–
130μm diam. (x=126μm, n=5), superficial, mostly solitary, subglobose, rounded above, light brown, ostiolate,
held to the leaf surface by basal mycelium, with apical
setae. Setae 38–56×4–5μm (x=48×4μm, n=10) at upper part of ascomata, larger and rounded at the base,
curved, dark brown to black. Peridium 9–14 μm (x=
10μm, n=5), comprising two-layers of cells of textura
angularis, outer layer composed of pigmented, thickwalled cells of textura angularis, inner layer composed
Fungal Diversity
Fungal Diversity
Chaetothyrium agathis (holotype). a, b Ascomata on host
surface c, d Ascomata viewed in squash mount e Ascoma with long
setae around the centre. f Section through ascoma g Base of setae h
Asci when immature i Asci in Melzer’s reagent with ocular chamber j,
k Multi-septate, verrucose ascospores l Multi-septate ascospore in
Melzer’s reagent m Ascospore in cotton blue reagent n Germinating
ascospore. Scale Bars: c, d=20μm, e, f=50μm, h, i=10μm, j, k, l, m=
5μm
Fig. 132
while setae are aseptate in T. criniporum. Molecular
analysis (ITS) supports the fact that T. siamensis is the
new species in Trichomerium with 91 % ML and 100 %
BP support.
Lecanorales
Squamarinaceae
of pale, pigmented and thin-walled cells of textura
angularis. Hamathecium lacking paraphyses. Asci 48–
64×14–21μm (x=62×17μm, n=5), 8-spored, bitunicate,
broadly cylindrical or oblong, usually with short pedicel. Ascospores 20–23×6–7μm, (x=21×6μm, n=10), bi
to tri-seriate, oblong to ellipsoid, 3-septate, not constricted at the septum, end cells narrow and smaller than
central cells, hyaline, wall slightly roughened. Asexual
morph Undetermined.
Material examined: THAILAND, Chiang Rai, Tasud,
on leaves of Tecoma sp. (Bignoniaceae), November
2 0 11 , S . H o n gs a n an H S A 09 ( M F L U 1 4 – 0 75 0 ,
holotype); ex-type living culture, MFLUCC 12–0097,
CPC 20469. GenBank ITS: KP744468.
Notes: This new taxon is similar to other species of
Trichomerium (Trichomeriaceae) based on its superficial
ascomata on the surface of leaves, and dark setae on the
upper part of ascomata (Chomnunti et al. 2014).
Trichomerium siamensis is most similar to T. ornatum
based on ascospore shape and septation, but differs in
having short setae at the upper part of ascomata in T.
siamensis, while ascomata have long setae in T.
ornatum. Moreover, T. siamensis is also similar to T.
criniporum due to the ascomata with setae and shape
of ascospores, but differs in having setae with septa,
Fig. 133 Phylogram generated
from Maximum likelihood
(RAxML) analysis based on ITS
sequence data of
Trichomeriaceae. Maximum
likelihood 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-types (reference
strains) are in bold; the new
isolates are in blue. The tree is
rooted with Phaeococcomyces
catenatus UAMH 4357
Among the ascomycetes, the subclass of Lecanoromycetidae
is characterized by discocarp ascomata, often with an I+ hymenium and typically by thick-walled asci with amyloid apical structures (Kirk et al 2008). The fungi are mainly lichenforming with mutualistic relationship to green algal and
cyanobacterial photobionts. While many groups of the subclass Ostropomycetidae exhibit associations with
trentepohlioid algae, the remaining three subclasses
Acarosporomycetidae, Umbilicariomycetidae and
Lecanoromycetidae mostly form thalli with, aside from
cyanobacterial associations, chlorococcoid and trebouxioid
algal species (Peršoh et al. 2004; Miadlikowska et al. 2006;
Miadlikowska et al. 2014). Lecanoromycetidae represent
about half of all lichenized species (Rambold et al. 2001 onwards), and form the majority of rock-inhabiting communities
with apothecial structures. About two percent of those are
lichenicolous and some 40 species exhibit endolichenic
growth of the vegetative thallus (about half of that number
within Lecanorales Nannf.), at least during their juvenile
stages. Based on morphological traits, the lichen family
Squamarinaceae was established by Hafellner (1984), but,
based on molecular phylogenetic data, has been transferred
into the synonymy of Stereocaulaceae Chevall (Lumbsch
and Huhndorf 2010). The LSU sequence of P. pratorum, its
closest relatives according to GenBank BLAST query, and
Fungal Diversity
Fig. 134 Trichomerium siamensis (holotype) a, b Ascomata on lower
surface of leaf c Dark setae clustered at the apex of ascomata d
Section through ascomata showing asci arrangement e Peridium and
ascus arrangement f Mycelium on surface of leaves g Asci embedded in
gelatinous matrix h, i Asci j–l Ascospore with 3-septa. Scale bars: c, d, h,
i=20μm, e, g=50μm, f, j, k, l=10μm
further representatives of the Lecanoromycetidae were aligned
using MUSCLE, as implemented in Geneious v. 6
(Biomatters, Auckland, New Zealand), the phylogenetic tree
generated for Lecanoromycetidae is presented in Fig. 135.
99. Paralecia Brackel, Greiner, Peršoh & Rambold, gen.
nov.
Index Fungorum number: IF551012, Facesoffunginumber:
FoF 00563
Etymology: The genus name ‘Paralecia’ refers to ‘para-’
(along with, beside) and to ‘lecia’ (refers to the lecideine
apothecia).
Type species: Paralecia pratorum Brackel, Greiner, Peršoh
& Rambold
Notes: This new species is characterized by its
lichenicolous, endothallic growth, lecideine apothecia, asci
with I+ dark blue tube-like apical structure, hyaline, and
Fungal Diversity
Fig. 135 Phylogram generated
from Maximum likelihood
(PhyxML v. 2.20) analysis based
on LSU sequence data. Maximum
likelihood bootstrap support
values greater than 70 % are
indicated above the nodes.
Species names are followed by
the voucher IDs and the new
species is in blue
simple ascospores. Among families with lecanoralean asci
(i.e. Lecanoromycetidae), this combination of traits matches
with species of the families (sensu Triebel, D. & Bensch, K.
2005 onwards) Aphanopsidaceae, Cladoniaceae,
Lecanoraceae, Lecideaceae, Malmidiaceae, Micareaceae,
Pannariaceae, Pilocarpaceae, Psoraceae, Ramalinaceae,
Stereocaulaceae, Vezdaceae, and the not yet family-assigned
genus Myochroidea (Rambold et al. 2001 onwards:
lichenicolous growth ignored; thallus growth habit set to crustose endosubstratal, episubstratal, (sub-)crustose, OR
(sub-)squamulose, primary photobiont set to
chlorophytaceous). However, Myochroidea differs by its
well-developed thallus and reddish brown apothecia (black
in Paralecia).
According to molecular data, all families, but
Stereocaulaceae (incl. Squamarinaceae Hafellner) and
Psoraceae (incl. Protomicarea Hafellner), may be discarded
as taxa to be considered. The LSU (GenBank Acc. No.:
KP224503) and ITS (KP224502) rRNA gene sequences were
compared with data in GenBank, using the BLAST function
of NCBI (Zhang et al. 2000). The next similar LSU sequence
originated from Squamarina cartilaginea (With.) P. James
Fungal Diversity
(DQ986763, grade 98.4 %, coverage 100 %, similarity
96.9 %). The ITS sequence of Paralecia pratorum, however,
did not reasonably match (similarity<80 %) any sequences in
the international sequence databases (INSDC; www.insdc.
org). According to LSU phylogeny topology (Fig. 135),
which is largely not conflicting with published phylogenies
(e.g., Andersen and Ekman 2005; Bendiksby and Timdal
2013; Wedin et al. 2009), P. pratorum is phylogenetically
close to the lichenized genus Squamarina (with S.
cartilaginea, DQ986763, and S. lentigera (Weber) Poelt,
AY756363) and is tentatively considered a member of the
Squamarinaceae Hafellner (Hafellner 1984). Within this
family, it is unique by its lichenicolous growth, lecideine
apothecia, and a more pronounced amyloid tube-structure of
the tholus. The ascus type (with a tube-like amyloid structure)
is in accordance with that known from Squamarina gypsacea
(Sm.) Poelt (Hertel and Rambold 1988). Next related, but with
only 91 % similarity (722 out of 795 bp are identical) and
without bootstrap support, appears to be Protomicarea
limosa (Ach.) Hafellner (AY756332), exhibiting a rather
similar combination of apothecial traits (excipulum,
pigmentation, ascus, ascospores).
Hitherto, only one record of an undescribed taxon,
‘Lecidea aff. insidiosa’, growing on P. muralis, exists (Nimis
and Poelt 1987; Rambold and Triebel 1992). Lecidea
Fig. 136 Paralecia pratorum
(holotype). a Habitius: ascomata
of P. pratorum on young lobes of
Protoparmeliopsis muralis b
apothecium in longitudinal
section c asci at different
ontogenetical stages and
paraphyses in Lugol’s solution
after pretreatment with KOH.
Scale bars: a=1 mm, b=50μm,
c=20μm
insidiosa Th. Fr. (KJ766650), exhibiting a more or less typical
Lecanora-type ascus, has meanwhile been combined to
Ramboldia insidiosa (Th. Fr.) Hafellner, but clusters in the
present LSU tree with Miriquidica complanata (Körb.)
Hertel & Rambold (KF562179; generic type of Miriquidica)
(Fig. 135). Phylogenetic analysis results further show a closer
relationship of the Squamarinaceae Hafellner to Lobariaceae
and Pannariaceae, but not to the Stereocaulaceae, which supports the suggestion of Miadlikowska et al. (2014) to resurrect
and recircumscribe this family.
100. Paralecia pratorum Brackel, Greiner, Peršoh &
Rambold, sp. nov.
Index Fungorum number: IF550913, Facesoffunginumber:
FoF 00553; Fig. 136
Etymology: The specific epithet ‘pratorum’ (lat., genitive
plural of ‘pratum’) refers to the name of the type locality ‘Prati
[= meadows] di Logarghena’.
Holotype: M-0045925
Lichenicolous on Protoparmeliopsis muralis. Thallus externally not visible, endothallic (Fig. 136a). Sexual morph
Apothecia breaking through the cortex of the host thallus, in
loose groups, sessile, constricted at the base, 200–450μm
wide; disc plane to convex at maturity, roundish, black,
epruinose; proper margin developed, mostly paler than the
disc and somewhat translucent, disappearing at maturity;
Fungal Diversity
excipulum c. 70μm thick, hyaline to slightly brownish on
upper side, filled with numerous crystals, composed of
thick-walled cells, 6–12×2–4μm; epihymenium pale brown,
K+ greyish brown, 10–20μm thick, filled with a multitude of
small crystals dissolving in K; hymenium hyaline, slightly
brownish on upper side, 60–85μm thick; hypothecium hyaline, 120–150μm thick (Fig. 136b). Paraphyses hyaline,
slightly brownish at the ends, with a brown cap, simple or
sparingly branched, 1.5–3μm wide; apices swollen up to
5μm. Asci (4–)8-spored, cylindrical to narrowly clavate, 55–
80×10–18μm, with a thickened tholus; tholus I+ blue with a
I+ dark blue tube-like structure, outer gelatinous layer I+ blue
(Fig. 136c). Ascospores hyaline, simple, ellipsoid, with a
rounded upper and a slightly attenuated lower end, or sometimes with both ends being slightly attenuated, (11–)11.8–
13.7(–15)×(4.5–)4.7–5.6(–6.5) μm, l/b=(1.7–)2.2–2.8(–3.1)
[n=40]. Asexual morph Undetermined.
Material examined: ITALY, Toscana, Prov. di MassaCarrara, Prati di Logarghena above the city of Pontremonli,
N44° 22.848′, E9° 56.573′, elev. 845 msl., growing on
Protoparmeliopsis muralis (Schreb.) M. Choisy, on schistose
rock outcrops in a meadow, 7 October 2013, W. v. Brackel
(M-0045925, holotype; GenBank ITS: KP224502; LSU:
KP224503; hb. Brackel 6996, isotype). ITALY, Toscana,
Prov. di Massa-Carrara, Valdantena, above Molinello, N44°
25.165′, E9° 55.883′, 460 msl., on P. muralis on a shady old
wall at the roadside, 5 October 2013, W. v. Brackel (hb.
Brackel 6914); below Passo Cirone, N44° 25.317′, E9°
54.988′, c. 1000 msl, on P. muralis on sandstone outcrops in
beech forest, 5 October 2013, W. v. Brackel (hb. Brackel
6941).
Notes: The fungus causes no recognisable harm to the host
thallus, but induces the formation of new lobes, also originating from the apothecial margin. The production of host
apothecia is suppressed to some degree. One has been
analysed to confirm the presence of Lecanora-type asci.
Ascomycota genera, incertae sedis
101. Lauriomyces synnematicus Somrithipol, sp. nov.
Index Fungorum number: IF551016, Facesoffungi number: FoF 00554; Figs. 137, 138
Etymology: synnematicum, refers to the synnematal
structure
Holotype: SFC 2156 in BBH
S a p ro b i c o n d e c a y i n g l e a v e s . S e x u a l m o r p h
Undetermined. Asexual morph Colonies scattered, white conidial sporulation on scattered synnemata. Synnemata solitary
or in groups, scattered, black, 50–225μm high, 10–40μm
diam. with 5–20 filaments closely adpressed along most of
their length. Filaments septate, thick and smooth-walled,
Fig. 137 Lauriomyces synnematicus (holotype). a A synnema with conidial mass b Synnematous filaments bearing conidiogenous cells and branched
chains of conidia and c Cylindrical conidia. Scale bars: a=50μm, b, c=10μm
Fungal Diversity
Hsieh 1990), Melanographium selenioides (Ellis 1963),
Memnoniella stilboidea (Ellis 1976) and Dictyoarthrinium
synnematicum (Somrithipol 2007). Conidia of L.
synnematicum are rather narrower than those of the other species in the genus with cylindrical conidia. Somrithipol and
Jones (2007) illustrated the gelatinization process during the
conidial formation of L. cylindricus, resulting in deposition of
mucilage between the conidia to join them into a persistent
chain. This conidial chain is the only character distinguishing
the genus Lauriomyces from Haplographium. In L.
synnematicum, the mucilage was less evident but some persistent chains could be observed.
Contributions to Basidiomycota
Fig. 138 Lauriomyces synnematicus (holotype). Line drawing of a
synnema with filaments bearing branched chains of conidia and
cylindrical conidia. Scale bars: = 10μm
brown to dark brown, 5–6μm wide, with terminal branches.
Branches cylindrical, thin and smooth-walled, hyaline to
subhyaline. Primary branches in clusters of 3–5 at the apex
of the filament, 6–7.5 μm long (x= 7 ± 0.68 μm, n =5), 1–
2.5μm wide (x=2±0.28 μm, n=5). Subsequent branches in
clusters of 3–5, 6–7.5μm long (x=6.75±0.68μm, n=5), 1–
2 μm wide (x=1.4±0.28μm, n=5). Ramoconidia and conidia
holoblastic, cylindrical, one-celled, hyaline to subhyaline, thinand smooth-walled, in acropetal branched chains, 4.5–7.5μm
long (x= 5.4 ± 0.61 μm, n = 50), 0.7–1.3 μm wide (x= 1 ±
0.12μm, n=50).
Material examined: THAILAND, Nakhon Rachasima, on
decaying leaves, 23 August 2008, B. Thongnuch (SFC 2156
in BBH, holotype).
Notes: Lauriomyces synnematicus differs from other species of the genus in possessing synnemata. The synnema is an
important characteristic to distinguish several hyphomycete
species, for example: Janetia synnematosa (Sivanesan and
Agaricus L.: Fr.
Agaricus (= Psalliota Fr.), the large and well-known edible
mushroom genus which also includes a small number of species that are toxic if eaten (Kerrigan et al. 2006; Zhao et al.
2012), comprises numerous species. According to Bas (1991),
the number of Agaricus described worldwide is probably
close to 400; Zhao et al. (2011) recognized 386 species in
the genus. Taxa from temperate regions are grouped into eight
commonly recognized sections based on morphological
and organoleptic traits, as well as macrochemical reactions (Cappelli 1984; Parra 2008, 2013). Compared with
temperate areas, knowledge of species diversity is lessdeveloped in tropical regions. Phylogenetic analyses of
Zhao et al. (2011) revealed seven (TR I to TR VII)
strongly supported tropical clades in addition to the
clades of the eight classical sections. In the past decade
about 36 Agaricus species have been described and 15
out of 36 were from tropical areas. The phylogenetic
tree is presented in Fig. 139.
102. Agaricus flavicentrus Karunarathna & K.D. Hyde,
sp. nov.
Index Fungorum number: IF550891, Facesoffungi
number: FoF 00392; Figs. 140, 141
Etymology: flavicentrus, in reference to the “yellow centre”, which is the colour of the pileus centre of this new
species.
Holotypus: MFLU 12–0146
Pileus 30–70 mm diam., circular from above,
hemisphaerical when young, convex when old; pileus splits
when matured, surface dry, smooth, with small yellow flakes
on the centre, yellowish white (2A2) at the centre white (5A1)
towards the margin when young and old; no colour changes
on pileus and stipe surfaces. Context in pileus 3–5 mm thick at
the disc, fragile, white, soft, and colour unchangeable.
Lamellae free, crowded with 4 tiers of lamellulae, 2–3 mm
wide with homomorph edge, light brown (5D4) at first, then
dark brown to chocolate brown (6F) at maturity, with
concolourous even edge. Annulus is simple, single, fragile,
Fungal Diversity
Fig. 139 Phylogram generated from Maximum Likelihood (PhyML ver
3.0) analysis based on ITS sequences of Agaricus from the sections
Agaricus, Sanguinolenti, Spissicaules, Arvenses and Minores (Lebel
and Syme 2012; Zhao et al. 2011). Maximum Likelihood bootstrap
support values greater than 50 % are indicated above the nodes, and
branches with Maximum parsimony bootstrap support values greater
than 75 % are given in bold. The type species are given in bold; the
new specimens are in blue. The tree is rooted with Barcheria willisiana
no floccose pieces, 10 mm diam., pendent, stretched,
white, cottony, white to slight yellowish white (3A2).
Stipe 40–70×10 mm at the base, at the middle, and at
the top, cylindrical, centrally attached to pileus, buried,
fistulose; context white, cottony; surface smooth, slight
yellowish white (2A2) at the top and yellowish white
Fig. 140 Agaricus flavicentrus a
fruiting bodies in the field (MFLU
12–0146, holotype). b fruiting
bodies in the laboratory (MFLU
12–0148). c fruiting bodies in the
field (MFLU 12–0147). d fruiting
bodies in the field (MFLU 12–
0149). Scale bars: a, b, d=10cm,
c=5cm
Fungal Diversity
Fig. 141 Agaricus flavicentrus
(MFLU12–0146, holotype) a
Basidia. b Basidiospores. c
Cheilocystidia. d Hyphae of
stipitipellis. e Hyphae of
pileipellis. Scale bars: a–e=20μm
(3A2) at base; rhizomorphs absent at base. No
discolouration observed in the pileus or stipe context
on touching or cutting. Odour bitter almond-like.
Macrochemical reaction: no reaction with 5 % KOH on
pileus surface and stipe surface (negative). Negative results
obtained for the Schaeffer reaction on fresh context of pileus
and stipe base.
Basidiospores 10–11 × 6–7 μm (x=10.50 × 6.50 μm,
Q=1.75–2.12, Qm =1.97, n=40), mostly oblong but rarely
ellipsoid, without germ pore, smooth, chocolate-brown in
mass, thick-walled. Basidia 18–25×4–6.5 (–7) μm, clavate,
4-spored. Cheilocystidia 12–18×8–10μm, broadly clavate,
single, hyaline, smooth. Pleurocystidia absent. Pileipellis a
cutis; hyphae 2.5–3μm diam., inflated, hyaline, unbranched.
Stipitipellis hyphae similar to pileipellis hyphae 1.5–2μm
diam. Clamp connections absent.
Habitat: as a group on grassland with humus-rich, organic
matter rich soil. Fruiting at the beginning of the rainy season,
found in open grassland with lot of organic litter mixed soil.
Material examined: THAILAND, Chiang Rai Province,
Mueang Chiang Rai District, Mae Fah Luang University park,
20° 2′44.39″N 99°53′37.38″E, 258 m, 21 March 2011, S.C.
Karunarathna (MFLU12–0146, holotype); GenBank ITS:
KR025857. THAILAND, Chiang Mai Province, Mueang
Chiang Mai District, Chiang Mai University park, 18°48′22″
N 98°57′10″E, 345 m, 27 July 2011, S.C. Karunarathna
(MFLU12–0147); GenBank ITS: KR025854. THAILAND,
Chiang Rai Province, Mueang Chiang Rai District, Mae Fah
Luang University park, 20° 2′44.39″N 99°53′37.38″E, 258 m,
23 March 2011, 15 May 2013, S.C. Karunarathna (MFLU12–
0148; GenBank ITS: KR025855; MFLU12–0149; GenBank
ITS: KR025856).
Notes: This new species is distinguished by its relatively
small to moderate basidiomes with a soft, hemisphaerical and
yellowish centered pileus; 10–11×6–7μm sized mostly oblong, thick-walled basidiospores, well visible single, nonfloccose, fragile, annulus; 12–18×8–10μm, broadly clavate
cheilocystidia and pleasant bitter almond odour.
103. Agaricus hanthanaensis Karunarathna & K.D. Hyde,
sp. nov.
Index Fungorum number: IF550892, Facesoffungi
number: FoF00393; Figs. 142, 143
Etymology: refers to the type collecting site “Hanthana”.
Holotypus: MFLU11–0017
Fungal Diversity
Fig. 142 Agaricus
hanthanaensis (MFLU 11–0017,
holotype) a–b Basidiome in the
field. c–d Basidiome in the
laboratory
Pileus 190–200 mm diam., ovoid, plano-convex,
applanate, at centre slightly depressed; surface squarrose or
densly squamose, dark to light brown (6D5); squamules on
the pileus reddish-dark brown (6D1), silky-fibrous, woolly,
soft; colour readily changing to red on bruising, touching or
cutting. Context in pileus 20–25 mm thick at the disc, fragile,
white soon after cut, white to light yellow with age. Lamellae
free, crowded with 8 tiers of lamellulae, 10 mm wide, ventricose,
light brown (6D4) at first, then dark brown (6E4) to chocolate
brown (6F8) at maturity, with concolourous even edge. Annulus
is not well visible but the remnants of the universal veil are
visible on the stipe which forms the annulus, brown, floccose.
Stipe 95×30 mm, at base 35 mm diam., cylindrical and
subbulbous at base, without rhizomorphs, buried, solid; surface
scabrous, reddish dark brown (6D1). Immediate discolouration
on touching and cutting, context at top of stipe pale red, at bottom
of the stipe pale yellow. Odour strong and peculiar.
Macrochemical reaction: 5 % KOH reaction on pileus surface and stipe surface yellow (positive). Schaeffer reaction on
dried tissue of the flesh of the cap and stipe base positive.
Basidiospores 4.2–6.5×2–3.5μm (x=5.1×2.8μm, Q=1.6–
2.2, Qm =1.7, n=40), ellipsoid, without germ pore, smooth,
reddish-brown in mass, thick-walled. Basidia 20–27×6–8 (–
9) μm, clavate, 4-spored. Cheilocystidia 30–50 (–56)×5–
11μm, clavate, single, hyaline, smooth. Pleurocystidia absent.
Pileipellis a cutis; hyphae 7–24μm diam., inflated, hyaline or
light brown, with intracellular light brown pigments in dots.
Stipitipellis hyphae similar to pileipellis hyphae 12–20μm
diam. Clamp connections absent.
Habitat: solitary in humus-rich soil under heavily rotted
litter in a highland forest. Fruiting during the middle of the
rainy season, found under Dipterocarpus spp. and Pinus spp.
Material examined: SRI LANKA, Central Province, Kandy
District, Hanthana Mountain Range, Peradeniya, rainforest
with Dipterocarpus spp. and Pinus spp., 7°15′35.03″N 80°36′
4.07″E, elev. 590 m, 5 July 2009, S.C. Karunarathna (MFLU
11–0017, holotype). GenBank ITS: JQ246405.
Notes: The morphological characters (yellow KOH reaction, positive Schaeffer’s reaction, peculiar odour, well-developed pileus squamules, and clavate cheilocystidia) support the
molecular data in that the species belongs to the section
Spissicaules (Nauta et al. 1999). The basal part of the stipe
context stains pale yellow, whereas the upper part stains pale
red on cutting or bruising which are also characteristic features
of the section Spissicaules (Nauta et al. 1999) where we place
A. hanthanaensis. The dark brown pileus squamules on a dark
to light brown background, the very thick fleshy (20–25 mm)
large pileus (190–200 mm) and the universal veil remnants
which forms the ring on the stipe of A. hanthanaensis easily
distinguishes it from the other species of section Spissicaules.
Agaricus hanthanaensis is remarkable amongst tropical
Agaricus species because of its relatively large-sized solitary
basidiocarps, as most tropical species of Agaricus have solitary, but small-sized basidiocarps (Zhao et al. 2011b). The
surface of the basidiome changes from brownish to red on
bruising or cutting. This new species bears superficial resemblance to A. bresadolanus but the latter species is distinct in
having 36–85 mm pilei with 5.5–7.5 × 4–5 μm, ellipsoid
Fungal Diversity
Fig. 143 Agaricus
hanthanaensis (MFLU 11-0017,
holotype) a Basidia. b
Cheilocystidia. c Basidiospores. d
Hyphae on Stipipellis. e Hyphae
on Pileipellis. Scale bars: a, b, d,
e=20μm, c=10μm
basidiospores and 13–25×7–11μm, clavate cheilocystidia
(Nauta et al. 1999). In some aspects Agaricus hanthanaensis
is similar to Agaricus simulans Berk. which was first described from Sri Lanka as a fleshy species typical of the subgenus Lanagaricus with a large basidiome (~140 mm), but it
differs in having ovoid spores, a floccose annulus formed by
universal veil, sticky scales on the pileus and stipe and smaller
cheilocystidia (28–32×6.5–7.5μm) (Pegler 1986). Our new
species also shares some similarities with A. parasilvaticus
Heinem., but the latter is distinct in having a 6 mm thick
context, 25–60 mm wide pilei, a superior annulus and 13–
37.5×7–15μm cheilocystidia (Heinemann 1962a).
104. Agaricus parvibicolor L.J. Chen, R.L. Zhao & K.D.
Hyde, sp. nov.
Index Fungorum number: IF550999, Facesoffungi
number: FoF00467; Figs. 144, 145
Etymology: Refers to the size of sporocarps and the white
and violaceous pileus.
Holotype: MFLU 12–0953
Pileus 15–40 mm diam., 1 mm thick, first parabolic, then
hemisphaerical to convex, and finally applanate; surface dry,
with reddish brown to violet brown fibrils, densely arranged at
disc, very few or none towards the margin on a white background; occasionally, with pileus expansion, starting from the
centre, the surface disrupted into triangular fibrillose
squamules; margin crenulate, at times appendiculate by
remnants from of the annulus, finely striate when sporocarps become mature; Lamellae free, crowded,
lamellulae with more than 5 series, 3–4 mm broad, at
first white, then pinkish, later greyish-brown and finally
dark brown. Stipe 32–57×2.5–5 mm, fistulose, cylindrical to slightly bulbous at base, with numerous
Fungal Diversity
Fig. 144 a–d Agaricus
parvibicolor (MFLU 12–0953,
holotype) a Sporocarps in situ. b
Pileus characters. c Finely striate
on the margin. d Membranous
annulus. e–h Agaricus sodalis
(MFLU 12–0987, holotype) e
Sporocarps in situ. f Sporocarp. g
Lamellae and annulus. h Staining
yellow when cut. Scale bars: a–
h=10 mm
rhizomorphs, surface white, staining orange by handing
or bruising, smooth above annulus, fibrillose below.
Annulus membranous, single, white, fragile. Context,
firm, white. Odour strong of almonds.
Macrochemical reactions: KOH reaction positive yellow.
Schäffer’s reaction positive reddish orange.
Basidiospores 4.7–5.5 (–5.7)×3–3.5 (–3.7) μm, (x=5.2×
3.3μm, Q=1.39–1.77, Qm =1.55, n=20), ellipsoid to oblong,
smooth, brown, thick–walled. Basidia 10–17×5.5–8μm, clavate to broadly clavate, hyaline, smooth, 4-spored.
Cheilocystidia 18–38×7.5–13μm, abundant, simple, broadly
clavate, pyriform or sphaeropedunculate, hyaline, smooth.
Pleurocystidia absent. Pileipellis a cutis composed of hyphae
of 5–9μm diam., cylindrical, light brown, smooth, at times
slightly constricted at the septa.
Habit: gregarious in small groups or solitary in soil of
grassland.
Material examined: THAILAND, Chiang Rai Province,
Mae Fah Luang University, 21 July 2012, collector Jie
Chen, LD2012116 (MFLU 12–0953, holotype). GenBank
ITS: KP715162; Chiang Rai Province, Mae fah Luang
University, 13 July 2012, collector Jie Chen, LD201288
(MFLU 12–0926), GenBank ITS: KP715165; LD201295
(MFLU 12–0933), GenBank ITS: KP715164; Mae Fah
Luang University, 21 July 2012, collector Jie Chen,
LD2012118 (MFLU 12–0954), GenBank ITS: KP715163;
Fungal Diversity
Fig. 145 Agaricus parvibicolor
(holotype) a Cheilocystidia b
Basidia c Basidiospores d
Pileipellis. Scale bars: a–c=5μm,
d=10μm
Mae Souy Dist., Pamae Lao National Park., 2 August 2006,
collected by Ruilin Zhao, ZRL3091 (SFSU).
Notes: The positive KOH and Schäffer’s reactions and almond-like odour suggest this species belongs to Agaricus section Minores. Agaricus parvibicolor is characterized by its
slender sporocarps, with reddish-brown to violet brown fibrils
at the centre of the pileus surface, and whitish towards the
margin, the simple cheilocystidia, frequently broadly clavate
or sphaeropedunculate in shape, the spore with an average size
of 5.2×3.3μm. Morphologically, the most similar species is A.
purpurellus (F.H. Møller) F.H. Møller, which also shares a
slender sporocarp and purplish fibrillose pileus. However, A.
purpurellus shows an entirely purplish pileus, slightly wider
in spore size (5.16×4μm) and a distinctive habitat of conifer
woods (Parra 2013). Another species with reddish-purple fibrils on the disc, but white elsewhere, is A.
dulcidulus Schulzer, but it can be easily distinguished
by its much smaller spore size which average is to
4.31×3μm (Parra 2013). In the phylogenetic analysis,
A. parvibicolor forms a single clade in Agaricus section
Minores.
105. Agaricus sodalis L.J. Chen, R.L. Zhao & K.D. Hyde,
sp. nov.
Index Fungorum number: IF551000, Facesoffungi
number: FoF00477; Figs. 144, 146
Etymology: Refers to the Latin adjective sodalis meaning
“the Agaricus of friends”
Holotype: MFLU 12–0987
Pileus 42–90 mm diam., 3.5 mm thick, plano-convex with
truncate centre; surface dry, with violet brown fibrils, densely
arranged at disc, rarely distributed or none towards the margin
on whitish background; margin finely crenulate; Lamellae
free, crowded, lamellulae with 3 series, 2.5–4 mm broad, at
first white, then pinkish, later greyish-brown and finally dark
brown; lamellae not exceeding the margin. Stipe 40–80×5–9
(10–15 at base) mm, fistulose, cylindrical with a bulbous base,
surface white, becoming yellow by handing, smooth above
annulus, fibrillose below. Annulus membranous, single, fragile
white, turning yellowish on bruising. Context firm, first white,
becoming conspicuously yellow when cut. Odour like almond.
Macrochemical reactions: KOH reaction positive yellow.
Schäffer’s reaction positive reddish orange.
Basidiospores 5–5.6 (–6.2)×(3.2–) 3.4–3.8μm, (x=5.4×
3.6μm, Q=1.39–1.63, Qm =1.50, n=20), ellipsoid, smooth,
brown, thick-walled. Basidia 12.5–25×7–8.5μm, clavate, hyaline, smooth, 4-spored; Cheilocystidia 21–38×6.5–14μm, abundant, simple, broadly clavate, pyriform or sphaeropedunculate,
with yellowish pigment, smooth. Pleurocystidia absent.
Pileipellis a cutis composed of hyphae of 5–12μm diam., cylindrical, rarely branched, hyaline, smooth, constricted at the septa.
Fungal Diversity
Fig. 146 Agaricus sodalis
(holotype) a Cheilocystidia b
Basidia c Basidiospores d
Pileipellis. Scale bars: a–c 5=μm,
d=10μm
Habit: gregarious or solitary in open area of forest.
Material examined: THAILAND, Chiang Rai Province,
Doi Pui site 2, 31 July 2012, collector Jie Chen, LD2012159
(MFLU 12–0987, holotype), GenBank ITS: KP715161;
Chiang Rai Province, Doi Pui site 1, 1 September 2011, collector Benjarong Thongbai, LD2011029 (MFLU 11–1311),
GenBank ITS: KP715160.
Notes: The positive KOH and Schäffer’s reactions, the almond-like odour and the simple cheilocystidia, place A.
sodalis in Agaricus section Minores. It is characterized by its
robust sporocarps, violet brown fibrils on the disc of the pileus
surface, but few or none elsewhere on a whitish background,
the simple cheilocystidia containing yellowish pigments, generally broadly clavate or sphaeropedunculate in shape and the
spores with an average size of 5.4×3.6μm. Within the section,
there are few species having robust sporocarps, exceptions are
A. brunneolus (J.E. Lange) Pilát, A. megalosporus J. Chen et
al. and A. pseudolutosus (G. Moreno et al.) G. Moreno et al.
Firstly, A. brunneolus exhibits much more coloured fibrils on
the entire pileus surface while the colour can be very variable,
from dark reddish purple to white and all the intermediate
colour shades, and there is no yellowish pigments been described from its cheilocystidia. The two other species can be
differentiated by their larger sized spores (A. megalosporus
6×3.5μm; A. pseudolutosus 6.37×4.78μm ); additionally A.
megalosporus shows a well coloured fibrillose squamules pileus and a variable shape of cheilocystidia has been observed
in A. pseudolutosus (Chen et al. 2012; Parra 2013). In the phylogenetic analysis, A. sodalis forms a clade with A. huijsmanii
and three undescribed species labelled as A. sp. ZRL3101, A. sp.
NT62 and A. sp. NTT72, but without significant support.
Cantharellus Adans.:Fr.
The economically very important edible genus
Cantharellus (Cantharellales) has been a taxonomical headache for several centuries because of the poor differentiation
of its microscopical features, an aspect that is probably related
to the fact that it represents one of the oldest clades of
mushroom-forming basidiomycetes. The molecularly oriented approach of the past decade has led to an explosion of newly described species, in particular from the
tropics, and revealed important differences between
northern and southern hemisphere species groups
(Buyck et al. 2014; Buyck et al. 2015a, b). The genus
presently counts slightly over one hundred species, and
all of them are ectomycorrhizal symbionts. The phylogenetic tree is presented in Fig. 147.
106. Cantharellus luteostipitatus Buyck, Randrianjohany
& V. Hofstetter, sp. nov.
MycoBank: MB 810671, Facesoffungi number: FoF
00555; Figs. 148, 149
Etymology: refers to the partly or entirely bright yellow
colouration of the stipe (luteo- meaning yellow and stipus
meaning stipe.
Holotype: PC0085575
Fungal Diversity
Fig. 147 Phylogram generated
from Maximum Parsimony
analysis based on TEF1 sequence
data of Cantharellus. Sequences
used in this study have been
sampled from Buyck et al. (2014)
or newly generated for
C. luteostipitatus sp. nov.
Branches indicated in bold
received significant (≥70 %)
bootstrap support. Species for
which obtained sequences are
based on type material have
names in bold
Pileus mostly 30–50(–60) mm diam., regular or with uneven surface when young and with sunken center from the
beginning, the margin strongly inrolled when young and remaining so for a long time, only in older specimens becoming
uplifted or revolute at the very extreme border only, sometimes slightly striate over a distance of max. 10 mm,
hygrophanous, when wet smooth and with velvety or even
distinctly greasy aspect, under a hand lens having almost a
frozen aspect, on drying becoming cottony-tomentose toward
the center, more rarely fragmenting in larger, appressed
squamules, colours varying between reddish pink and
yellowish orange tints, often unevenly distributed on the surface and fading rapidly on drying. Hymenophore decurrent, of
well-developed gill folds, 2–4(–6) mm high, very variable in
colour, starting out as very pale cream or nearly whitish, but
sometimes rapidly developing yellowish orange tints, particularly toward the cap margin, gills predominantly unequal,
forkings abundant only close to the very cap margin, varying
from completely smooth to heavily anastomosed in between
gills. Stipe mostly shorter than the cap diam., 8–17 mm diam.,
subcylindrical or slightly narrowing upwards, tinged with
bright chrome yellow to a diluted yellowish orange over part
Fungal Diversity
Fig. 148 Cantharellus luteostipitatus (PC0085576). General habit
(photo B. Buyck)
or all of its surface, except for the extreme base which remains
generally pure white, solid. Flesh firm and thick beneath the
cap center but then rapidly becoming very thin toward the
margin, white but with orange-reddish tinges immediately below the cap surface, sometimes showing distinct bluish tinges
Fig. 149 Cantharellus luteostipitatus (holotype) a Spores. b Basidia,
basidiola and subhymenial cells. c Hyphal extremities of the pileipellis.
Scale bars=10μm, but only 5μm for spores. Drawings B. Buyck
when cut, on the surface of stipe and gills yellowing when
handled. Taste mild. Smell of apricot. Spore print pale
yellowish.
Spores ellipsoid, (7.1)7.5–8.04–8.5(8.8)×(4.8)5.2–5.77–
6.3(6.7) μm, Q=(1.25)1.31–1.40–1.49(1.56), smooth, filled
with numerous oily inclusions. Basidia 50–60 × 7–9 μm,
subcylindrical to weakly clavulate, (2)4(–5)-spored and with
5–8 μm long sterigmata (longer for 2–3 spored basidia);
basidiola not particularly irregular in form. Subhymenium
pseudoparenchymatic, of large voluminous cells. Cystidia
none. Pileipellis a loose (tricho)cutis of ramifying, thin-walled
hyphal extremities, measuring mostly 5–10μm diam., terminal cells subcylindrical and undifferentiated or sometimes remarkably short and occasionally more inflated to claviform;
incrusting pigments most distinct on the narrowest hyphae of
subpellis and trama. Clamp connections absent from all
tissues.
Material examined: MADAGASCAR, Province of
Toamasina, East Coast, 15 km south of Brickaville, on a private property near Ambila Lemaitso, growing in groups of 3
to 6 individuals under Uapaca littoralis in the deep sandy soils
of the littoral forest, 27 June 2011, Buyck & V. Hofstetter
11.042 (PC0085573, paratypus), GenBank TEF: KP033509;
Buyck & V. Hofstetter 11.044 (PC0085575, holotypus),
GenBank TEF: KP033510; Buyck & V. Hofstetter 11.045
(PC0085576, paratypus), GenBank TEF: KP033511.
MADAGASCAR Central Plateau, Ibity, in Uapaca bojeri
woodland, 1500–1700 m alt., 27 January 2008, Buyck & V.
Hofstetter 08.210 (PC 0085130).
Notes: Madagascar is extremely rich in chanterelles (Buyck
2012, 2014; Buyck et al. 2014; Buyck and Randrianjohany
2013) with many species showing very close relationships to
mainland African taxa. The newly described chanterelle in this
paper strongly resembles the type variety of the mainland
African Cantharellus platyphyllus Heinemann in the field because of the pinkish red tinges in its cap colour and palecoloured hymenophore of distinct gill folds. The Malagasy
C. platyphyllus ssp. bojerienis Eyssart. & Buyck, which is
common and abundant in the Uapaca bojeri woodlands on
the Central Plateau, has more grayish-yellowish-greenish-orange tints on the cap. Our species differs from both essentially
in the bright yellow colouration of its stipe. Nevertheless,
there is quite some variation in the general habit and
colouration of these taxa. The only feature so far that allowed
to separate all the forms of C. platyphyllus from the mainland
African C. symoensii Heinemann, a red-capped species that
typically develops a deep, egg-yolk yellow hymenophore at
maturity, was the form of the spores: shortly ellipsoid to ellipsoid (mean length-width ratio ca 1.4) for C. platyphylllus and
its various infraspecific taxa versus narrowly ellipsoid to elongate spores for C. symoensii (mean length-width ratio of ca 1.9
– see Eyssartier and Buyck 1999). Our new species, although
genetically closer to C. symoensii, has the same spores as C.
Fungal Diversity
Fig. 150 Phylogram generated
from Maximum likelihood
(RAxML) analysis based on ITS
sequence data. Maximum
likelihood bootstrap support
values greater than 50 % are
indicated above or below the
nodes, new species are in blue and
species for which obtained
sequences are based on type
material have names in bold.The
tree is rooted with Multifurca
furcatus RH804, Multifurca
ochricompacta BB02-107 and
Multifurca zonaria DED7442
platyphyllus and therefore makes the identification of the
African taxa in subgen. Afrocantharellus Eyssart. & Buyck
even more delicate.
All specimens of C. luteostipitatus were collected in the
deep sandy soils of the littoral forest, but are genetically close
to one collection (Fig. 147, as C. luteostipitatus cf) made on
the rocky, lateritic soils of the seasonally much colder woodlands of the Central highlands near Ibity at 1500–1700 m
altitude. The latter collection consists of a single very immature specimen that lacks the yellow stipe and, therefore, looks
even more like typical mainland C. platyphyllus. As these
ecologically and climatically very different sites harbor mutually exclusive host tree species, it would seem surprising that
their associated fungi were less affected. Additional collecting
will be needed to solve the status of this Central Highland’s
collection.
Cantharellus luteostipitatus is the second species of subgenus Afrocantharellus described from Madagascar. Whereas
this subgenus was for many years exclusively known from
Africa, its distribution has very recently been extended to
Malaysia (Buyck et al. 2014) and China (Shao et al. 2014).
Lactarius Pers.
Fungal Diversity
Fig. 151 Lactarius atrobrunneus (holotype). Basidiocarps in the field
(photo by K. Wisitrassameewong)
Fig. 152 Lactarius
atrobrunneus (holotype) a
Basidiospore b Basidia c
Marginal cells d Pseudocystidia e
pileipellis. Scale bar=10μm.
Drawings by K.
Wisitrassameewong
Lactarius is a dominant genus of ectomycorrhizal fungi
represented worldwide which exudes latex when the fruiting
body is bruised. According to the critical revision of the genus
(Buyck et al. 2010), three subgenera are conserved: L. subg.
Lactarius, L. subg. Russularia and L. subg. Plinthogalus. In
this contribution two new species presented from Thailand are
introduced. Both species belong to subg. Russularia. The phylogenetic tree is presented in Fig. 150.
107. Lactarius atrobrunneus Wisitrassameewong & K. D.
Hyde, sp. nov.
Index Fungorum number: IF551005 Facesoffungi number:
FoF 00483, Figs. 151, 152
Etymology: atrobrunneus refers to dark brown colour.
Holotypus: MFLU 12–1136
Fungal Diversity
Basidiocarps small sized. Pileus 6–31 mm diam., broadly
convex in young specimens, with or without a small papilla,
becoming plane to infundibuliform at maturity; surface dry,
radially rugulose, particularly in inner half of pileus, strongly
and irregularly wrinkled in the outer half of pileus, with
grooves that show the lamellae in transparency; margin striate,
occasionally fissured, incurved; surface blackish to dark
brown when young, turning less unicolourous and between
dark brown (9 F8), reddish-brown (9E8) and blackish when
older. Lamellae subdecurrent to decurrent, 1–2 mm broad,
distant, with 1–2 series of lamellulae, beige to greyish beige
(4C2–4C3) to fuliginous. Stipe 14–37×1–4 mm, fragile, cylindrical, more or less concolourous to pileus, with darker
brown tints when old. Context thin-fleshed, 0.5–1 mm broad
in pileus, pinkish to clay buff, darker in cortex, unchanging
when cut, but changing yellow with 10 % KOH within seconds; smell like L. quietus, like Pentatomidae bugs; taste mild.
Latex rather abundant, pure watery, unchanging when exposure, unchanging with 10 % KOH, unchanging on white tissue
or paper; taste faintly bitter. Basidiospores subglobose to ellipsoid; 6.9–8.1–8.2–9.6 (−10.7)×6–7–7.2–8.3(−8.5) μm,
Q=1.06–1.15–1.16–1.29 (n=40); ornamentation amyloid,
subreticulate, composed of ridges up to 1.5μm high; ridges
often spiny and irregular, rather broad, interconnected by finer
lines, forming an incomplete to subcomplete reticulum; isolated warts infrequent; plage distally amyloid. Basidia 55–
72×18–20μm, mostly 4–spored, with small or large guttulate
contents. Pleuromacrocystidia absent. Pseudocystidia 4–6μm
diam., cylindrical, often curved, slightly emergent to emergent. Lamellae edge heterogeneous, with some basidia present; marginal cells 11–44 × 5–12 μm, subcylindrical to
subclavate; cheilocystidia absent. Lamellar trama mixed,
composed of small and large globose cells, septate hyphae
and lactiferous hyphae. Pileipellis an epithelium, about 40–
70μm thick, composed of globose cells of 5–25μm diam.,
with some subclavate elements on top; underlying layer composed of smaller globose cells and cylindrical hyphae.
Habitat: gregarious on naked soil, in montane tropical forest with Fagaceae trees.
Material examined: THAILAND, Chiang Mai Province,
Mae Taeng District, Pa Pae sub-district, Mushroom
Research Centre, N19°17.12 E98°44.00, elev. ca 900 msl.,
KW347, 9 July 2012 (MFLU12–1136, holotype, isotype in
GENT. GenBank ITS: KP744442); ibid., KW270, 7 June
2012 (MFLU12–1079, paratypes; isoparatypes in GENT
GenBank ITS: KP744443).
Notes: Lactarius atrobrunneus is rather unique and wellrecognizable in the subgenus by its small size and striking
blackish-brown colour. Furthermore the strongly rugose cap,
the pure transparent latex and distant gills are striking field
characters. Microscopically, the rather spiny and heavy spore
ornamentation and the well-developed epithelium as a
pileipellis structure are striking. Other species of the subgenus
with remarkably distant lamellae are L. laccarioides
Wisitrassameewong & Verbeken, L. sublaccarioides
W i s i t r a s s a m e e w o n g & Ve r b e k e n , L . s t u b b e i
Wisitrassameewong & Verbeken and L. pasohensis
Wisitrassameewong & Stubbei (Wisitrassameewong et al.
2014). All of them are larger and have distinctly orange brown
colours and white latex.
108. Lactarius politus Wisitrassameewong & K.D. Hyde,
sp. nov.
Index Fungorum number: IF551004 Facesoffungi number:
FoF00484; Figs. 153, 154
Etymology: politus refers to polished cap surface
Holotypus: MFLU 12–0822
Basidiocarp medium sized. Pileus 25–66 mm diam., at first
broadly convex, later turning infundibuliform, inconspicuously mucronate in centre; surface dry and polished, somewhat
smooth when young, becoming minutely wrinkled; margin
incurved, translucently striate up to almost halfway the radius,
crenulate when mature; surface reddish brown or dark brown
(8F8), with darker zonation at least in inner half of pileus,
palest at pileus margin, in outer half ranging from dark brown
(7F8–8F8) to reddish brown (7E8) to brownish orange (5C4–
C5) to brown (5E8) to olive brown (4E8) to greyish orange
(5B4). Lamellae decurrent, 3 mm broad, medium crowded,
sometimes forked, with 1–2 series of lamellulae, light yellow
(4A4) to greyish yellow (4B4) to olive brown (4D5–E3), with
greyish brown (5E3) tinge. Stipe 30–62×5–8 mm, cylindrical;
surface dry, longitudinally wrinkled, brown (5E5) to greyish
orange (6B3) to dark brown (9F8). Context 2 mm broad in
pileus, medium firm, hollow in stipe, greyish pink; smell
strong, sweetish and spicy, maggi-like in dried specimens;
unchanging with 10 % KOH and unchanging with FeSO4.
Latex transparent, unchanging on exposure, unchanging with
10 % KOH, unchanging on white tissue or paper; taste mild.
Basidiospores globose to subglobose, sometimes ellipsoid;
Fig. 153 Lactarius politus (MFLU 12–0822, holotype). Basidiocarp in
the field (photo by K. Wisitrassameewong)
Fungal Diversity
Fig 154 Lactarius politus
(MFLU 12–0822, holotype) a
Basidiospore b
Pleuromacrocystidia c
Cheilocystidia d Basidia e
Pseudocystidia f Marginal cells g
Pileipellis. Scale bar=10μm.
Drawings by K.
Wisitrassameewong
7.7–8.6–9–10.2(−10.6)×6.7–7.6–7.8–8.7μm; Q=1.03–1.11–
1.17–1.33 (n = 80); ornamentation amyloid, reticulate,
ornamented with very heavy, dense and high ridges up to
2.5μm; ridges irregular and thick interconnected with finer
lines, forming a dense reticulum, sometimes with a slight tendency of zebroid ornamentation; plage sometimes distally amyloid. Basidia 55–84×18–23μm, 4–spored, subclavate, with
guttulate contents. Pleuromacrocystidia very scarce, 55–77×
16–20 μm, not emergent, subfusiform to subclavate,
mucronate to rounded, with small granules and guttulate contents. Pseudocystidia 3–6μm diam., not emergent, tortuous
cylindrical, often curved, sometimes ramified. Lamellae edge
heterogeneous, with some basidia present; marginal cells
subcylindrical to subclavate to obovoid to capitate, 25–43×
11–22μm; cheilocystidia scarce, 30–55×8–14μm, not emergent, subcylindrical to subclavate, with mucronate apex.
Lamellar trama consisting of small and large globose cells,
septate hyphae and lactiferous hyphae. Pileipellis a
Fungal Diversity
hyphoepithelium; upper layer 20–30μm thick, consisting of
repent cylindrical hyphae; subpellis consisting of globose cells
of 10–40μm diam.
Habitat: solitary or scattered on soil in montane tropical
forest near Lithocarpus sootepensis stand.
Material examined: THAILAND, Chiang Rai Province,
Mae Fah Luang district, Doi Mae Salong Nok sub-district,
Doi Mae Salong, elev. ca 1269 msl., N20°16.90 E99°62.30,
13 July 2012, KW351 (MFLU 12–0822, holotype, isotype in
GENT; GenBank ITS: KF433019), ibid., 22 August 2012,
KW396 (MFLU 12–0845, paratype, isoparatype in GENT;
GenBank ITS: KF433018), ibid., 10 September 2013,
KW468 (MFLU13–0589, paratype, isoparatype in GENT;
GenBank ITS: KP744445); Chiang Mai province, Mae
Taeng district, Pa Pae sub-district, Mushroom Research
Center, elev. ca 900 msl., N19°07.20 E98°44.04, 3 July
2012, KW331(MFLU 12–0812, paratype, isoparatype in
GENT; GenBank ITS: KF433020), Chiang Mai province,
Bahn Mae Sae village, on Highway no.1095 near 50 km
marker, elev. ca 962 msl., N19°14.59 E098°39.45, 4 July
2011, KW107 (MFLU 11–1214, paratype, isoparatype in
GENT; GenBank ITS: KP744444).
Notes: Lactarius politus is recognized by its large, ellipsoid
spores and firm ornamentation forming a complete reticulum.
The other important characters are the watery latex and the
strong smell. The species is rather variable in pileus colour
which could be brown zonation or rather even dark brown in
inner half of pileus and with paler margin. The pileipellis
structure is hyphoepithelium which is common feature in subgenus Russularia.
Phylloporia
Phylloporia Murrill, typified by P. parasitica Murrill, is one of
the most economically important wood decaying genera within the Hymenochaetaceae, Hymenochaetales, because some
species in this genus are considered to be forest pathogens.
Phylloporia is evidenced from nLSU sequences to be a monophyletic genus, but its species are highly diverse in morphology. A species with distinct morphological characters is newly
described from China below. The phylogenetic tree is presented in Fig. 155.
109. Phylloporia dependens Y.C. Dai, sp. nov.
Index Fungorum number: IF551014 Facesoffungi number:
FoF00556; Figs. 156, 157
Etymology: referring to the hanging down growing habit.
Holotype: BJFC013379
Basidiocarps perennial, pileate to pendent, corky and without odour or taste when fresh, becoming woody hard up on
drying. Pilei projecting up to 5 cm, 4 cm wide and 5 cm thick
at base. Pileal surface vinaceous brown to black when fresh
and dry, narrowly sulcate, glabrous; margin obtuse, vinaceous
brown to brown. Pore surface cream-brown to pale brown
when fresh, buff-yellow to cinnamon-buff when dry, more
or less glancing; margin buff-yellow, narrow to almost lacking; pores circular or angular, 7–9 per mm; dissepiments thin,
entire. Context yellowish-brown to cinnamon, corky, up to
1 mm thick. Tubes cinnamon, slightly darker than pore surface, up to 49 mm long. Hyphal system monomitic; generative
hyphae with simple septa; neither amyloid nor dextrinoid,
acyanophilous; tissue darkening but otherwise unchanged in
KOH. Contextual hyphae pale yellowish brown to yellowish
brown, thick-walled with a wide to narrow lumen, occasionally collapsed, rarely branched, regularly arranged, 2.4–
3.1μm diam. Tramal hyphae pale yellowish brown to yellowish brown, thick-walled with a wide to narrow lumen, occasionally branched, straight, subparallel along the tubes, 2–
3μm diam. Setae absent; cystidia absent; cystidioles present,
fusoid, hyaline, thin-walled, 9–17×3–4.5μm; basidia more or
less barrel-shaped, with four sterigmata and a simple septum
at the base, 9–12×4–5μm; basidioles mostly pear-shaped,
slightly smaller than basidia. Crystals polyhedric, frequently
present among trama and subhymenium. Basidiospores
broadly ellipsoid, yellowish, thick-walled, usually collapsed
when mature, neither amyloid nor dextrinoid, moderately
cyanophilous, 3–3.4 × 2.7–3 (–3.1) μm, L = 3.16 μm, W =
2.9μm, Q=1.09 (n=30/1).
Material examined: CHINA, Yunnan Province, Ruili, Moli
Tropical Rainforest, on rotten angiosperm stump, 1 November
2012, Dai 13167 (BJFC013379, holotype; GenBank LSU:
KP698746; IFP 019122, isotype).
Notes: Phylloporia verae-crucis (Berk. ex Sacc.) Ryvarden
has similar pores (7–9 per mm) to P. dependens, but has larger
basidiospores (4–4.5×3–3.5μm, Wagner and Fischer 2002).
In addition, it lives on ground over buried wood, and occurs in
South America (Wagner and Fischer 2002). Phylloporia was
well studied in China, and 13 species were recorded in the
county besides the new species (Dai 2010, 2012; Zhou and
Dai 2012; Zhou 2013). Phylloporia dependens differs from all
the existed species in the genus by its hanging down growing
habit. In addition, phylogenetic analysis based on nLSU region indicates that P. dependens belonged to Phylloporia, and
occupied a distinct lineage with strong support (96 % BS and
1 BPP, Fig. 155). So the morphological and molecular evidence confirmed the placement of the new species.
Russula Pers.
Russula Pers. is one of the most diverse genera of
ectomycorrhizal basidiomycetes with an estimated diversity
of several thousands of species ranging from completely hypogeous to stipitate epigeous or pleurotoid forms. It is also
very well represented in the tropics. The phylogenetic tree is
presented in Fig. 158.
110. Russula cortinarioides Buyck, Adamčík, Lewis & V.
Hofstetter, sp. nov.
MycoBank MB 810670; Facesoffungi number: FoF 00557;
Figs. 159, 160, 161
Fungal Diversity
Fig. 155 Phylogram generated
from Maximum Likelihood
(RAxML) analysis based on LSU
sequence data. Maximum
likelihood bootstrap support
values greater than 50 % and
Bayesian posterior probabilities
greater than 0.90 are indicated
above the nodes, new species are
in blue and species for which
obtained sequences are based on
type material have names in bold.
The tree is rooted with Inonotus
hispidus
Etymology: referring to the gradually developing reddish
brown colour of the gills, giving this species almost a
Cortinarius-like aspect.
Holotypus: PC 0142175.
Basidiocarps growing individually or in small groups. Cap
36–50(65) mm diam., regular, smooth, hardly pealing, glabrous, slightly depressed and not discolouring in the centre,
unevenly tinged in colours that vary from grayish brown, yellowish brown to reddish brown, sometimes also predominantly dirty cream to whitish, particularly closer to the cap margin,
greasy-viscose when wet, shiny even when dry, separable up
to mid-radius, not striate at margin. Gills adnate to
subdecurrent, brittle, unequal and irregularly alternating with
lamellulae of one, two to sometimes three lengths, not widely
spaced (8–15 l+L/cm at cap margin), 3–5 mm high, sometimes forked or anastomosing particularly close to the cap
margin, whitish when very young, then rapidly cream to pale
yellowish brown and finally turning vinaceous to deep brownish red when old; edges concolourous, even. Stipe central,
shorter or more rarely longer than cap diam., 31–33(55)×7–
Fungal Diversity
Fig. 156 Phylloporia dependens basidiocarps (holotype)
11 mm, cylindrical, with the base often irregularly deformedwrinkled and sometimes narrowing (as in R. adusta), surface
smooth, and although not pruinose, often more or less silveryglistening and whitish, rapidly dirty grayish when handled or
rubbed gently as the initial whitish covering disappears and
revealing a distinct longitudinal striation, relatively firm,
spongy inside under a hard outer cortex which is grayish
brown in section, not really chambered but irregularly
hollowing in the central medulla. Context firm and fleshy in
the cap center, only 2 mm thick at cap mid radius, whitish,
turning brown with age, weakly reddening to becoming vinaceous when cut. Smell distinct and nauseous, somewhat alkaline (as in R. pseudolateriticola). Taste unpleasant and faintly
astringent. Spore print white.
Macrochemical reactions: stipe surface reacting strongly
with guaiac, insensitive to FeSO4.
Spores ellipsoid, (7.5–)7.8–8.1–8.4(–8.7)×(5.6–)5.8–6.1–
6.4(–6.6) μm, Q = (1.25–)1.28–1.32–1.36(–1.4), with
subreticulate to reticulate ornamentation of low, distinctly amyloid, obtuse warts [6–8 warts in a 3μm diam. circle on the
spore surface], 0.2–0.3μm high, connected by fine line connections [2–5 lines in the circle] or fused in pairs or short
chains [(1–)2–4 fusions in the circle], with a smooth, relatively
large, nonamyloid suprahilar spot. Basidia (39–)46–50–57×
8–9–10μm, 4-spored, narrowly clavate to subcylindrical;
basidiola first cylindrical, then narrowly clavate.
Subhymenium composed of densely septate, narrow cells.
Lamellar trama with large sphaerocytes only present near
the cap trama. Hymenial cystidia on gill sides abundant,
[3000–4000 per mm2], (62–)67–80–93(–115)×7–8–9μm, also numerous but much shorter near the gill edge, measuring
(26–)40–47–56×6–8(–10) μm, thin–walled, subulate, narrowly lageniform to subcylindrical, rarely clavate or lanceolate, sometimes mucronate with a short, 1–3(–5) μm long
appendage, the interior partly filled with granular or slightly
crystalline contents, turning reddish grey in sulfovanilin.
Marginal cells not differentiated. Pileipellis 110–140μm
deep, gelatinized, orthochromatic in Cresyl blue, ill-delimited
from the underlying context. Incrustations absent. Hyphal extremities ascending, long and slender, becoming gradually
denser downwards, uniformly cylindrical, sparsely branching,
with obtuse terminal cells measuring (19–)30–41–60(–
85)×(3–)4–5(–6) μm near cap margin, but narrower, 2.5–
3.5–4(–4.5) μm diam. in the cap center. Pileocystidia numerous, usually very long (frequently>100μm) and originating
deep in subpellis or cap trama where they continue as abundant cystidioid hyphae, one–celled, subcylindrical, apically
attenuated and slightly mucronate, without or with one or
two appendages (2–8μm long), often shorter, obtuse–rounded
and more clavate in cap center, thin–walled, (3.5–)4.5–5–6μm
diam., filled with granular to yellowish-oily contents that do
not react in sulfovanilin. Clamp connections absent in all
parts.
Specimens examined: UNITED STATES, Texas, Newton
Co., Canyon Rim, near Mayflower, along State Highway 87,
in Beech-Magnolia-Loblolly Pine woods, 28 July 2007,
Buyck 07.133 (PC 0142175, holotypus); Ibid., 5 July 2002,
Buyck 02.115 (PC0142176), 02.116bis (PC0142183), 29 Jul
2005, Buyck 05.084 (PC0124614), 28 July 2007, Buyck
07.131 (PC0142177 paratypus); Bleakwood, on Core‘s residence, in open Oak-Pine vegetation, N 30° 42.068′, W 93°
49.770′, Lewis 9185 (PC0142182). Montgomery Co.,
Conroe, in Oak-Pine woods, 27 July 2007, Buyck 07.103
(PC0124672), 07.104 (PC 0142178 paratypus), 07.105 (PC
0142179), 07.111 (PC 0142180 paratypus); Hardin Co., near
Saratoga, Lance Rosier Unit, Big Thicket National Preserve,
along Teel Cemetery road near cypress swamp among
Cantharellus texensis in Oak-Loblolly Pine woods, N 30°
15.901′, W 94° 30.759′, ca 50 m alt., 16 June 2013, DP
Lewis 10797 (PC 0142181), ibidem, 23 June 2014, Buyck,
Hofstetter & Lewis leg., Buyck 14.024–14.027 (PC 0142213–
PC 014226); Turkey Creek unit, Big Thicket National
Preserve, along Kirby nature trail, N 30.47181-W 94.34888,
2 July 2014, Buyck 14.118 (PC 0142307)
Notes: The combined phylogenetic analysis of ITS, LSU &
RPB2 sequences (Fig. 158) shows that the new species,
Russula cortinarioides, is the most related to R. polyphylla
Peck, the type species of subsect. Polyphyllinae Singer 1951
nom. inval. (Art. 39.1). This subsection was originally defined
by the presence of numerous “gloeo-vessels or
macrocystidioid oleiferous hyphae” in the upper layer of the
cap cuticle, to distinguish them from other members of sect.
Rigidae Fr. (= subgenus Heterophyllidia Romagn.). Singer
(from 1951 onwards) recognized four species in this subsection: R. polyphylla (Peck 1898), R. magnifica (Peck 1903), R.
polycystis (Singer 1939) and R. viridella (Peck 1906).
Even before introducing this new subsection, Singer (1943)
had already suggested the co-identity of the former two species, a synonymy he maintained up to the last edition of his
Agaricales (Singer 1986), and in which he was followed by
Fungal Diversity
Fig. 157 Phylloporia dependens
(holotype) microscopic structures
a Basidiospores. b Basidia and
basidioles. c Cystidioles. d
Hyphae in trama. e Hyphae in
context
most mycologists. It is therefore difficult to understand why
Singer placed Polyphyllinae in sect. Rigidae near subsect.
Virescentinae (Schaeff.) Fr. because R. magnifica was described (Peck 1903) as a whitish species having unequal gills,
with suggested affinities to a completely different group:
subsect. Lactarioideae in subgen. Compacta Fr. (see Buyck
and Adamčík 2013). Even Singer (as early as 1926) agreed
with such a systematic placement for R. magnifica until he
changed opinion in 1951 without any clear argumentation.
The two other species, R. polycystis and R. viridella, have
been placed in Virescentinae, until Singer (1951) transferred
them to Polyphyllinae. R. polycystis was originally described
as a new species of Virescentinae (Singer 1939) and this position has been maintained, although with some hesitation,
when Bills (1984) reported on recollecting it for the first time
since its original description. R. viridella has also been associated with Virescentinae (Singer 1932 and thereafter) not
with standing its acrid taste. It is more than likely that both
these species are unrelated to either of these subsections.
It was Buyck (in Buyck et al. 2005) who, for the first time,
pointed out the strong microscopical similarities between R.
polyphylla and R. eccentrica Peck, a relationship that is fully
Fungal Diversity
Fig. 158 Phylogram generated
from Maximum Parsimony
analysis based on combined
sequence data of RPB2, LSU, ITS
for 31 Russula and six outgroup
sequences (2 Multifurca spp., 2
Lactarius spp. and 2 Lactifluus
spp.). Sequences used in this
study have been sampled from a
previous study (Buyck et al 2008)
or newly generated for
R. cortinarioides, R. polyphylla,
R. eccentrica, R. lateriticola and
R. fistulosa var. grata [see
GenBank accession numbers
KP033498–KP033508 (RPB2),
KP033487–KP033497
(nucLSU), KP033476–
KP033486 (ITS). Branches
indicated in bold received
significant (≥70 %) bootstrap
support
supported (BS = 100 %) by our phylogenetic analysis
(Fig. 158). The molecular data indicate that Polyphyllinae
are neither related to Virescentinae, nor to Lactarioideae, but
are closely related to subsect. Nigricantes Fr., a species group
typically characterized by their, often first reddening, then
blackening context. The closest, native American taxon to
our new species seems to be R. densifolia var. paxilloides
Peck, which differs from our species by the differentiation of
marginal cells and in the less reticulate and less dense, slightly
higher spore ornamentation (see Adamčík and Buyck 2014).
Polyphyllinae differ from Nigricantes by the fact that their
context is only reddening but never blackening, a feature most
easily observed on the ageing hymenophore which turns pinkish to reddish brown at maturity. As morphology in our diagnosis, R. cortinarioides is most similar to R. eccentrica in the
field but has denser gills, a feature that is even much more
Fungal Diversity
Fig. 159 Russula cortinarioides (holotype). Habit (Buyck 07.133).
Photo B. Buyck
Fig. 160 Russula cortinarioides
(holotype). Microscopic features
of the pileipellis. a Pileocystidia
with indication of contents as
observed in Congo Red for two
elements, schematically in all
other elements. b Hyphal
extremities. Scale bars=10μm.
Drawings B. Buyck
pronounced in the larger, nearly whitish R. polyphylla, its
sister-species, which has crowded gills.
In the field, R. cortinarioides is also very easily confused
with R. compacta Frost because of the unequal gills, brownish
colours, unpleasant smell, white spore print and overall similar habit. The latter differs from our new species in the hymenium bruising brownish upon handling but not slowly turning
reddish to reddish brown as in R. cortinarioides; both species
are also easily distinguished by the very different microscopic
features of their pileipellis and our phylogeny distinctly shows
both species to be quite unrelated.
As a result of our molecular analysis, we therefore restrict
Polyphyllinae in North America presently to R. polyphylla, R.
eccentrica and the here newly described R. cortinarioides.
Fungal Diversity
Fig. 161 Russula cortinarioides.
Microscopic features of the
hymenium. a Basidia. b
Basidiola. c Spores. d
Gloeopleurocystidia. e
Gloeocheilocystidia (no marginal
cells differentiated).
Gloeocystidia with indication of
contents as observed in Congo
Red for two elements,
schematically in all other
elements. Scale bars=10μm, but
only 5μm for spores. Drawings B.
Buyck
Polyphyllinae is also known from Costa Rica (see Buyck and
Halling 2004; Buyck et al. 2005) and the subsection is well
represented in most other tropical and subtropical areas of the
world (Buyck unpubl.). All species of this subsection are recognizable by their unequal gills becoming pinkish to reddish
brown at maturity, a slowly, mostly weakly reddening, but not
blackening context; they have gloeoplerous elements in all
parts of the carpophore and possess very similar spores.
Acknowledgments The research was supported by 1) Chinese Academy of Sciences (Project No. KSCX2–EW-Z-9). K. Tanaka would like to
thank the Japan Society for the Promotion of Science (JSPS, 25440199)
for financial support. This project was supported by the Distinguished
Scientist Fellowship Program (DSFP), King Saud University, Saudi Arabia. This project was also supported by NSTIP strategic technologies
program, number (12–BIO2840–02) in the Kingdom of Saudi Arabia.
B. Buyck and V. Hofstetter acknowledge the ATM “Emergences” (Dirs.
P. Janvier & S. Peigné) from the National museum of natural history,
Paris, for financial support related to sequencing as well as travel expenses in Madagascar. Overseas travel of B. Buyck was funded by the
ATM research program “Etat et structure phylogénétique de la
biodiversité actuelle et fossile” of the Paris’ Museum. (director Ph.
Janvier); S. Adamčík acknowledges funding by his national Slovak Project VEGA 02/0075/14. B.D. Shenoy acknowledges funding to visit Mae
Fah Luang University, Chiang Rai under Indo-Thailand bilateral programme (THAI-1205). The Lewis family is warmly thanked for their
generous hospitality, offering excellent and very agreeable working and
Fungal Diversity
field condition. K. Wisitrassameewong and A. Verbeken were financially
supported by the joint doctorate program of the “Bijzonder
Onderzoeksfonds Gent University” (BOF), Gent University. This research was also financial supported by The Biodiversity and Training
Program (BRT R 149009) and Interaction Laboratory (BMIT), National
Center for Genetic Engineering and Biotechnology (BIOTEC) Khlong
Luang, Pathum Thani 12120, Thailand. Thanks are extended to the Thailand Research Fund grant (BRG 5580009) under the research grant entitled “Taxonomy, Phylogeny and Biochemistry of Thai Basidiomycetes”
for financial support. J. Chen and R. L. Zhao would like to thank the
National Natural Science Foundation of China (Project ID: 31000013,
31360014 and 31470152). Uwe Braun, Walter Jaklitsch, Tom May, Marc
Stadler and Jo Taylor, are thanks for commenting on some parts of the
manuscript. Shaun Pennycook is thanked for checking and suggesting
corrections to most of the Latin names.
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