Fungal Diversity (2020) 102:1–203
https://doi.org/10.1007/s13225-020-00448-4
Microfungi associated with Clematis (Ranunculaceae)
with an integrated approach to delimiting species boundaries
Chayanard Phukhamsakda1,2,3 · Eric H. C. McKenzie4 · Alan J. L. Phillips5 · E. B. Gareth Jones9,10 · D. Jayarama Bhat14 ·
Stadler Marc16 · Chitrabhanu S. Bhunjun3 · Dhanushka N. Wanasinghe2,3,8,15 · Benjarong Thongbai16 ·
Erio Camporesi11,12,13 · Damien Ertz17,20 · Ruvishika S. Jayawardena3 · Rekhani Hansika Perera3,18 ·
Anusha H. Ekanayake3 · Saowaluck Tibpromma2,8,19 · Mingkwan Doilom2,8,19 · Jianchu Xu2,19,8 · Kevin D. Hyde1,2,3,6,7,8
Received: 5 March 2020 / Accepted: 6 May 2020 / Published online: 7 July 2020
© The Author(s) 2020
Abstract
The cosmopolitan plant genus Clematis contains many climbing species that can be found worldwide. The genus occurs in
the wild and is grown commercially for horticulture. Microfungi on Clematis were collected from Belgium, China, Italy,
Thailand and the UK. They are characterized by morphology and analyses of gene sequence data using an integrated species
concept to validate identifications. The study revealed two new families, 12 new genera, 50 new species, 26 new host records
with one dimorphic character report, and ten species are transferred to other genera. The new families revealed by multigene
phylogeny are Longiostiolaceae and Pseudomassarinaceae in Pleosporales (Dothideomycetes). New genera are Anthodidymella (Didymellaceae), Anthosulcatispora and Parasulcatispora (Sulcatisporaceae), Fusiformispora (Amniculicolaceae),
Longispora (Phaeosphaeriaceae), Neobyssosphaeria (Melanommataceae), Neoleptosporella (Chaetosphaeriales, genera
incertae sedis), Neostictis (Stictidaceae), Pseudohelminthosporium (Neomassarinaceae), Pseudomassarina (Pseudomassarinaceae), Sclerenchymomyces (Leptosphaeriaceae) and Xenoplectosphaerella (Plectosphaerellaceae). The newly described
species are Alloleptosphaeria clematidis, Anthodidymella ranunculacearum, Anthosulcatispora subglobosa, Aquadictyospora
clematidis, Brunneofusispora clematidis, Chaetosphaeronema clematidicola, C. clematidis, Chromolaenicola clematidis,
Diaporthe clematidina, Dictyocheirospora clematidis, Distoseptispora clematidis, Floricola clematidis, Fusiformispora
clematidis, Hermatomyces clematidis, Leptospora clematidis, Longispora clematidis, Massariosphaeria clematidis, Melomastia clematidis, M. fulvicomae, Neobyssosphaeria clematidis, Neoleptosporella clematidis, Neoroussoella clematidis,
N. fulvicomae, Neostictis nigricans, Neovaginatispora clematidis, Parasulcatispora clematidis, Parathyridaria clematidis,
P. serratifoliae, P. virginianae, Periconia verrucose, Phomatospora uniseriata, Pleopunctum clematidis, Pseudocapulatispora clematidis, Pseudocoleophoma clematidis, Pseudohelminthosporium clematidis, Pseudolophiostoma chiangraiense, P.
clematidis, Pseudomassarina clematidis, Ramusculicola clematidis, Sarocladium clematidis, Sclerenchymomyces clematidis,
Sigarispora clematidicola, S. clematidis, S. montanae, Sordaria clematidis, Stemphylium clematidis, Wojnowiciella clematidis, Xenodidymella clematidis, Xenomassariosphaeria clematidis and Xenoplectosphaerella clematidis. The following
fungi are recorded on Clematis species for the first time: Angustimassarina rosarum, Dendryphion europaeum, Dermatiopleospora mariae, Diaporthe ravennica, D. rudis, Dichotomopilus ramosissimum, Dictyocheirospora xishuangbannaensis,
Didymosphaeria rubi-ulmifolii, Fitzroyomyces cyperacearum, Fusarium celtidicola, Leptospora thailandica, Memnoniella
oblongispora, Neodidymelliopsis longicolla, Neoeutypella baoshanensis, Neoroussoella heveae, Nigrograna chromolaenae,
N. obliqua, Pestalotiopsis verruculosa, Pseudoberkleasmium chiangmaiense, Pseudoophiobolus rosae, Pseudoroussoella
chromolaenae, P. elaeicola, Ramusculicola thailandica, Stemphylium vesicarium and Torula chromolaenae. The new combinations are Anthodidymella clematidis (≡ Didymella clematidis), A. vitalbina (≡ Didymella vitalbina), Anthosulcatispora
brunnea (≡ Neobambusicola brunnea), Fuscohypha kunmingensis (≡ Plectosphaerella kunmingensis), Magnibotryascoma
rubriostiolata (≡ Teichospora rubriostiolata), Pararoussoella mangrovei (≡ Roussoella mangrovei), Pseudoneoconiothyrium
euonymi (≡ Roussoella euonymi), Sclerenchymomyces jonesii (≡ Neoleptosphaeria jonesii), Stemphylium rosae (≡ Pleospora
rosae), and S. rosae-caninae (≡ Pleospora rosae-caninae). The microfungi on Clematis is distributed in several classes of
Extended author information available on the last page of the article
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Fungal Diversity (2020) 102:1–203
Ascomycota. The analyses are based on morphological examination of specimens, coupled with phylogenetic sequence data.
To the best of our knowledge, the consolidated species concept approach is recommended in validating species.
Keywords 73 new taxa · Ascomycota · Belgium · China · Dothideomycetes · Incertae sedis · Italy · Lecanoromycetes ·
Phylogeny · Sordariomycetes · Taxonomy · Thailand · UK
Table of Contents
Phylum Ascomycota R.H. Whittaker
Subphylum Pezizomycotina Erikss. & K. Winka
Class Dothideomycetes Erikss. & K. Winka
Subclass Pleosporomycetidae Schoch et al.
Pleosporales Luttr. ex M.E. Barr
Amniculicolaceae Zhang et al.
1. Fusiformispora Phukhams. & K.D. Hyde, gen.
nov.
2. Fusiformispora clematidis Phukhams., M.V. de
Bult & K.D. Hyde, sp. nov.
Amorosiaceae Thambug. & K.D. Hyde
3. Angustimassarina rosarum Tibpromma, Camporesi & K.D. Hyde, new host record
Cyclothyriellaceae Jaklitsch & H. Voglmayr
4. Massariosphaeria clematidis Phukhams., Wanas.,
Camporesi & K.D. Hyde, sp. nov.
Dictyosporiaceae Boonmee & K.D. Hyde
5. Aquadictyospora clematidis Phukhams., Bhat &
K.D. Hyde, sp. nov.
6. Dictyocheirospora clematidis Phukhams., Bhat &
K.D. Hyde, sp. nov.
7. Dictyocheirospora xishuangbannaensis Tibpromma & K.D. Hyde, new host record
8. Pseudocoleophoma clematidis Phukhams. & K.D.
Hyde, sp. nov.
Didymellaceae Gruyter, Aveskamp & Verkley
9. Anthodidymella Phukhams., Camporesi & K.D.
Hyde, gen. nov.
10. Anthodidymella clematidis (Woudenb., Spiers &
Gruyter) Phukhams. & K.D. Hyde, comb. nov.
11. Anthodidymella ranunculacearum Phukhams.,
Camporesi & K.D. Hyde, sp. nov.
12. Anthodidymella vitalbina (Petr.) Phukhams. &
K.D. Hyde, comb. nov.
13. Neodidymelliopsis longicolla Hou, Crous & L. Cai,
new host record
14. Xenodidymella clematidis Phukhams., Camporesi
& K.D. Hyde, sp. nov.
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Didymosphaeriaceae Munk
15. Chromolaenicola clematidis Phukhams. & K.D.
Hyde, sp. nov.
16. Didymosphaeria rubi-ulmifolii Ariyaw., Camporesi & K.D. Hyde, new host record
Hermatomycetaceae Locq. ex A. Hashim. & K. Tanaka
17. Hermatomyces clematidis Phukhams., Bhat &
K.D. Hyde, sp. nov.
Leptosphaeriaceae Barr
18. Alloleptosphaeria clematidis Phukhams. & K.D.
Hyde, sp. nov.
19. Sclerenchymomyces Phukhams. & K.D. Hyde,
gen. nov.
20. Sclerenchymomyces clematidis Phukhams. & K.D.
Hyde, sp. nov.
21. Sclerenchymomyces jonesii (Wanasinghe, Camporesi & K.D. Hyde) Phukhams. & K.D. Hyde, comb.
nov.
Longiostiolaceae Phukhams., Doilom & K.D. Hyde
22. Longiostiolaceae Phukhams., Doilom & K.D.
Hyde, fam. nov.
Lophiostomataceae Luerss.
23. Neovaginatispora clematidis Phukhams., Ertz,
Gerstmans & K.D. Hyde, sp. nov.
24. Pseudocapulatispora clematidis Phukhams. & K.D.
Hyde, sp. nov.
25. Pseudolophiostoma chiangraiense Phukhams. &
K.D. Hyde, sp. nov.
26. Pseudolophiostoma clematidis Phukhams. & K.D.
Hyde, sp. nov.
27. Sigarispora clematidis Phukhams. & K.D. Hyde,
sp. nov.
28. Sigarispora clematidicola Phukhams., Camporesi
& K.D. Hyde, sp. nov.
29. Sigarispora montanae Phukhams., Sue & K.D.
Hyde, sp. nov.
Melanommataceae Winter
30. Neobyssosphaeria Wanas., Jones & K.D. Hyde,
gen. nov.
31. Neobyssosphaeria clematidis Wanas., Phukhams.,
Jones & K.D. Hyde, sp. nov.
Fungal Diversity (2020) 102:1–203
Neomassarinaceae Mapook & K.D. Hyde
32. Pseudohelminthosporium Phukhams. & K.D.
Hyde, gen. nov.
33. Pseudohelminthosporium clematidis Phukhams.
& K.D. Hyde, sp. nov.
Nigrogranaceae Jaklitsch & H. Voglmayr
34. Nigrograna chromolaenae Mapook & K.D. Hyde,
new host record
35. Nigrograna obliqua Jaklitsch & H. Voglmayr, new
host record
Occultibambusaceae Dai & K.D. Hyde
36. Brunneofusispora clematidis Phukhams. & K.D.
Hyde, sp. nov.
Paradictyoarthriniaceae Doilom, Liu & K.D. Hyde
37. Xenomassariosphaeria clematidis Wanas.,
Phukhams., Camporesi & K.D. Hyde, sp. nov.
Periconiaceae Nann.
38. Periconia verrucosa Phukhams, Ertz, Gerstmans
& K.D. Hyde, sp. nov.
Phaeoseptaceae Boonmee, Thambug. & K.D. Hyde
39. Pleopunctum clematidis Phukhams., Bhat & K.D.
Hyde, sp. nov.
Phaeosphaeriaceae Barr
40. Chaetosphaeronema clematidicola Phukhams,
Ertz, Gerstmans & K.D. Hyde, sp. nov.
41. Chaetosphaeronema clematidis Phukhams, Ertz,
Gerstmans & K.D. Hyde, sp. nov.
42. Dermatiopleospora mariae Wanas., Camporesi,
Jones & K.D. Hyde, new host record
43. Leptospora clematidis Phukhams., Ertz, Gerstmans, & K.D. Hyde, sp. nov.
44. Leptospora thailandica Phukhams. & K.D. Hyde,
new host record
45. Longispora Phukhams. & K.D. Hyde, gen. nov.
46. Longispora clematidis Phukhams. & K.D. Hyde,
sp. nov.
47. Pseudoophiobolus rosae Phookamsak, Wanas.,
Phukhams., Camporesi & K.D. Hyde, new host
record
48. Wojnowiciella clematidis Phukhams., Ertz, Gerstmans & K.D. Hyde, sp. nov.
Pleosporaceae Nitschke
49. Stemphylium clematidis Wanas., Camporesi &
K.D. Hyde, sp. nov.
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50. Stemphylium rosae (Wanas. et al.) Phukhams. &
K.D. Hyde, comb. nov.
51. Stemphylium rosae-caninae (Wanas. et al.)
Phukhams. & K.D. Hyde, comb. nov.
52. Stemphylium vesicarium (Wallr.) E.G. Simmons,
new host record
Pseudoberkleasmiaceae Phukhams. & K.D. Hyde
53. Pseudoberkleasmium chiangmaiense Lu & K.D.
Hyde, new host record
Pseudomassarinaceae Phukhams. & K.D. Hyde
54. Pseudomassarinaceae Phukhams. & K.D. Hyde,
fam. nov.
55. Pseudomassarina Phukhams. & K.D. Hyde, gen.
nov.
56. Pseudomassarina clematidis Phukhams, Camporesi & K.D. Hyde, sp. nov.
Pseudolophiotremataceae Hyde & S. Hongsanan
57. Clematidis italica Tibpromma, Camporesi & K.D.
Hyde
Roussoellaceae Liu, Phookamsak, Dai & K.D. Hyde
58. Neoroussoella clematidis Phukhams. & K.D.
Hyde, sp. nov.
59. Neoroussoella fulvicomae Phukhams. & K.D.
Hyde, sp. nov.
60. Neoroussoella heveae Senwanna, Phookamsak &
K.D. Hyde, new host record
61. Pararoussoella mangrovei (Phukhams. & K.D.
Hyde) Phukhams. & K.D. Hyde, comb. nov.
62. Pseudoneoconiothyrium euonymi (Crous & Akulov) Phukhams. & K.D. Hyde, comb. nov.
63. Pseudoroussoella chromolaenae Mapook & K.D.
Hyde, new host record
64. Pseudoroussoella elaeicola (Konta & K.D. Hyde)
Mapook & K.D. Hyde, new host record
Sulcatisporaceae Tanaka & K. Hirayama
65. Anthosulcatispora Phukhams. & K.D. Hyde, gen.
nov.
66. Anthosulcatispora brunnea (Chen & C. Norphanphoun) Phukhams. & K.D. Hyde, comb. nov.
67. Anthosulcatispora subglobosa Phukhams. & K.D.
Hyde, sp. nov.
68. Parasulcatispora Phukhams. & K.D. Hyde, gen.
nov.
69. Parasulcatispora clematidis Phukhams. & K.D.
Hyde, sp. nov.
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Teichosporaceae Barr
70. Floricola clematidis Phukhams., Camporesi &
K.D. Hyde, sp. nov.
71. Magnibotryascoma rubriostiolata (Jaklitsch &
Voglmayr) Phukhams., Jones & K.D. Hyde, comb.
nov. and new host record
72. Ramusculicola clematidis Phukhams. & K.D.
Hyde, sp. nov.
73. Ramusculicola thailandica Thambug. & K.D.
Hyde, new host record
Thyridariaceae Tian & K.D. Hyde
74. Parathyridaria clematidis Phukhams., Camporesi
& K.D. Hyde, sp. nov.
75. Parathyridaria serratifoliae Phukhams., Ertz, Gerstmans & K.D. Hyde, sp. nov.
76. Parathyridaria virginianae Phukhams., Ertz, Gerstmans & K.D. Hyde, sp. nov.
Torulaceae Corda
77. Dendryphion europaeum Crous & R.K. Schumacher, new host record
78. Torula chromolaenae Li, Phookamsak, Mapook &
K.D. Hyde, new host record
Dothideomycetes, family incertae sedis
Dyfrolomycetales Pang, Hyde & E.B.G. Jones
Pleurotremataceae Watson
79. Melomastia clematidis Phukhams., & K.D. Hyde,
sp. nov.
80. Melomastia fulvicomae Phukhams., & K.D. Hyde,
sp. nov.
Class Lecanoromycetes Erikss. & K. Winka
Subclass Ostropomycetidae Reeb, Lutzoni & Cl. Roux
Ostropales Nannf.
Stictidaceae Fr.
81. Fitzroyomyces cyperacearum Crous, new host
record
82. Neostictis Ekanayaka, Camporesi & K.D. Hyde,
gen. nov.
83. Neostictis nigricans Ekanayaka, Phukhams.,
Camporesi & K.D. Hyde, sp. nov.
Class Sordariomycetes Erikss. & K. Winka
Subclass: Sordariomycetidae Erikss. & K. Winka
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Chaetosphaeriales Huhndorf, Mill. & F.A. Fernández
Chaetosphaeriales, genera incertae sedis
84. Neoleptosporella Phukhams. & K.D. Hyde, gen.
nov.
85. Neoleptosporella clematidis Phukhams., Konta &
K.D. Hyde, sp. nov.
Sordariales Chadef. ex Hawksw. & O.E. Erikss.
Chaetomiaceae G. Winter
86. Dichotomopilus ramosissimum (X. Wei Wang &
L. Cai) X. Wei Wang & Samson, new host record
Sordariaceae Winter
87. Sordaria clematidis Phukhams. & K.D. Hyde, sp.
nov.
Subclass Diaporthomycetidae Senan., Maharachch. &
K.D. Hyde
Diaporthales Nannf.
Diaporthaceae Hohn. ex Wehm.
88. Diaporthe clematidina Phukhams., M.V. de Bult
& K.D. Hyde, sp. nov.
89. Diaporthe ravennica Thambug., Camporesi &
K.D. Hyde, new host record
90. Diaporthe rudis (Fr.) Nitschke, new host record
Phomatosporales Senan., Maharachch. & K.D. Hyde
Phomatosporaceae Senan. & K.D. Hyde
91. Phomatospora uniseriata Phukhams., M.V. de
Bult & K.D. Hyde, sp. nov.
Diaporthomycetidae, family incertae sedis
Distoseptisporaceae Hyde & E. McKenzie
92. Distoseptispora clematidis Phukhams., M.V. de
Bult & K.D. Hyde, sp. nov.
Subclass Xylariomycetidae Erikss. & W. Winka
Amphisphaeriales Hawksw. & O.E. Erikss
Sporocadaceae Corda
93. Pestalotiopsis verruculosa Maharachch. & K.D.
Hyde, new host record
Xylariales Nannf.
Diatrypaceae Nitschke
94. Neoeutypella baoshanensis Raza, Shang,
Phookamsak & L. Cai, new host record
Subclass Hypocreomycetidae Erikss. & K. Winka
Glomerellales Chadef. ex Re´blova´ et al.
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Plectosphaerellaceae Gams, Summerb. & R. Zare
95. Fuscohypha kunmingensis (Phookamsak, J.F. Li
& K.D. Hyde) Jayaward., Phukhams. & K.D. Hyde,
comb. nov.
96. Xenoplectosphaerella Jayaward., Phukhams. &
K.D. Hyde, gen. nov.
97. Xenoplectosphaerella clematidis Jayaward.,
Phukhams. & K.D. Hyde, sp. nov.
Hypocreales Lindau
Nectriaceae Tul & C. Tul
98. Fusarium celtidicola Shang, Camporesi & K.D.
Hyde, new host record
Sarocladiaceae Lombard
99. Sarocladium clematidis Phukhams., Ertz, Gerstmans & K.D. Hyde, sp. nov.
Stachybotryaceae Lombard & P. Crous
100. Memnoniella oblongispora Lin, McKenzie, Wang
& K.D. Hyde, new host record
Introduction
Clematis (Ranunculaceae) is a flowering climber which
has become a popular plant in horticulture (Linnaeus 1753;
Yuan et al. 2010). The genus contains between 250 and 350
species and hybrids (Grey-Wilson 2000; Lehtonen et al.
2016; He et al. 2019). Clematis is widespread in warm-temperate or montane ecosystems and is native to most areas
of China, Europe, Korea, and Russia (Tamura 1956; Ziman
and Keener 1989; Yuan and Yang 2020). Clematis vitalba
(old man’s beard), the type species of Clematis is an invasive weed broadly distributed in Europe, and also expanding to New Zealand and South America (Ogle et al. 2000;
Leuschner and Ellenberg 2017; Redmond and Stout 2018).
Clematis vitalba can influence the biodiversity dynamics of native plants (Ogle et al. 2000; Ashton and Lerdau
2008). In Thailand, Clematis species are mostly found in
the northern provinces of Chiang Mai, Chiang Rai and Nan,
which are mountainous areas with a tropical savanna climate
(Tamura 1997, 2000). Many Clematis species are grown as
ornamental plants. Clematis species are also used in Traditional Chinese Medicine and some secondary metabolites
isolated from Clematis have been tested in vitro, but there
are no reports of successful clinical tests or if it is safe to
consume the plant parts (Ding et al. 2009; Fu et al. 2010;
Feng et al. 2011; Hawaze et al. 2012; Lu et al. 2014; Zhao
et al. 2016). Clematis species are herbaceous vines, with
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opposite compound, bipinnate to tripinnate leaves, and
leather-like flowers with feather achenes (Johnson 2001)
(Fig. 1). Section-level phylogenetic classification of Clematis by Lehtonen et al. (2016) includes specific characteristics
and geographic distribution for each section. The estimated
divergence time of the available sequences for Clematis have
shown that the stem age was in the Oligocene (25.99 million
years ago; Lehtonen et al. 2016).
Fungal species associated with Clematis have been documented since the late eighteenth century (Lamarck 1805;
Saccardo 1892; Farr and Rossman 2020; Index Fungorum
2020). Index Fungorum and the U.S. National Fungus Collections Fungal Database lists over 500 records, mainly as
saprobes, or pathogens that can cause leaf lesions and wilt
in Clematis species (Baylis 1954; Braun 1992; Ahn and
Shearer 1998; Wanasinghe et al. 2014; Chen et al. 2015;
Crous et al. 2019).
In this study, Clematis samples were collected in Belgium, China, Italy, Thailand, and the UK to establish the
microfungi associated with this host and to analyze their
host-specificity. In addition, fungal isolates were evaluated
for their antagonistic activity against selected microorganism (Phukhamsakda et al. 2018; Hyde et al. 2019b; Macabeo
et al. 2020). The delineation of new species introduced in
this study relies on a polyphasic approach based on morphological traits (MSC), molecular data (PSC), and application of Genealogical Concordance Phylogenetic Species
Recognition (GCPSR) (Taylor et al. 2000). GCPSR model
relies on performing a pairwise homoplasy index coupled
with phylogenetic relatedness in a multi-locus dataset and
the interpretation of nucleotide differences (Turner et al.
2013; Quaedvlieg et al. 2014; Jeewon and Hyde 2016). We
also compared the morphology of our new collections with
documented fungal taxa recorded in public databases and
discuss their ecological species concepts.
Materials and methods
Sample collection, morphological study
and isolation
Fresh Clematis specimens were collected or received from
Belgium, China, Italy, Thailand and England (the UK).
Some specimens have single collection because this study
mainly focused on the diversity of fungi associated with
Clematis. One collection is defined as a sample of fungus
that can be identified with a single collecting trip which was
used to cast the number of species. Thus, the plant materials
were mainly collected and received from a single trip of the
aforementioned countries. The specimens were maintained
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Fig. 1 a–c Habitats of Clematis species. d Opposite compound with
bi-pinnate to tri-pinnate leaves. e Woody climbing stem. f Inflorescence of C. pitcheri. g Inflorescence of C. “Crystal Fountain”.
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h Inflorescence of C. vitalba. i Achenes of C. vitalba. j Enlarged
achenes of C. subumbellata with plumose style
Fungal Diversity (2020) 102:1–203
in paper bags for transport to the laboratory. The specimens
were examined using a Motic SMZ 168 Series stereo-microscope. Thereafter, vertical free-hand sections were made by
a razor blade and placed on a droplet of sterilized water on
a glass slide. A Nikon ECLIPSE 80i compound microscope
was used to examine the samples and a Canon 600D digital camera fitted to the microscope was used to photograph
the samples. Tarosoft (R) Image Frame Work program was
used for measurements and photo-plate were made by using
Adobe Photoshop CS6 Extended version 10.0 software
(Adobe Systems, United States).
Pure cultures were obtained from single ascospores isolation on malt extract agar (MEA: 33.6 g/L, malt extract
Difco™) or potato dextrose agar (PDA: 39 g/L, potato
dextrose media Difco™) as described by Chomnunti et al.
(2014) which were incubated at 16–25 °C with the standard
light cycles, 12 h in the light followed by 12 h in the dark
for about four up to eight weeks. Asexual reproduction was
induced by placing agar squares with mycelia on water agar
or MEA placed with additional substances such as sterile
pine needles or rice straws. Authentic type specimens are
deposited in Mae Fah Luang University (MFLU) herbarium and ex-type living cultures are deposited at the Mae
Fah Luang Culture Collection (MFLUCC). Faces of fungi
numbers (Jayasiri et al. 2015) and Index Fungorum numbers
(2020) are provided.
DNA extraction, amplification and sequencing
The Biospin Fungus Genomic DNA Extraction Kit (BioFlux®) (Hangzhou, P. R. China) and gene extraction kit
(Bio Basic Inc., Canada) were used for DNA extraction from
mycelium. The fruiting bodies DNA was extracted by using
Forensic DNA Kit–D3591-01 (OMEGA bio-tek) following
the manufacturer’s instructions. Polymerase chain reaction
(PCR) was used to amplify partial gene regions with primer
pairs as described in Tibpromma et al. (2018). The PCR
amplifications were performed in a total volume of 25 µL
solution containing 10–20 ng of DNA template, Easy Taq
PCR Super Mix (mixture of Easy Taq TM DNA Polymerase,
dNTPs, and optimized buffer) and 10 picomolar forward and
reverse primers. Amplification reactions were performed following Phukhamsakda et al. (2016) and Tibpromma et al.
(2018). Genomic DNA and PCR amplification products
were checked on 1% agarose gel. PCR products were purified as described in the manufacturer’s instructions (EZ-10
PCR Products Purification Kit, Bio basic Canada INC.).
Sequences were generated by Shanghai Sangon Biological
Engineering Technology & Services Co. (Shanghai, P.R.
China) and the sequencing service at Helmholtz Centre For
Infection Research (HZI, Braunschweig, Germany).
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Sequence alignment and phylogenetic analysis
Consensus sequences were assembled using SeqMan v.
7.0.0 (DNASTAR, Madison, WI). Sequences of closely
related strains were retrieved using BLAST searches against
GenBank (http://www.ncbi.nlm.nih.gov). Sequences were
aligned with MAFTT version 7 (Katoh et al. 2019) (http://
mafft.cbrc.jp/alignment/server), with minimal adjustment
of the ambiguous nucleotides by visual examination and
manually corrected in AliView program (Larsson 2014).
Leading or trailing gaps exceeding the primer binding site
were trimmed from the alignments prior to tree building and
alignment gaps were treated as missing data. The concatenation of the multigene datasets was created in Sequence
Matrix (Vaidya et al. 2011).
Phylogenetic analyses of the single gene and combined
gene were based on maximum parsimony (MP), maximum
likelihood (ML) and Bayesian inference posterior probabilities (BYPP). PAUP program was used for MP bootstrap
analyses, with 1000 bootstrap replicates using 10 rounds
of the heuristic search replicates to estimate the homoplasy
yield. The random addition of sequences and subsequent
TBR branch swapping during each bootstrap replicate, with
each replicate was limited to 1000 rearrangements. Gaps
were treated as missing data; all characters were unordered
and given equal weight. The statistics for parsimony were
described under the phylogenetic legend with the values of
Tree Length (TL), Consistency Index (CI), Retention Index
(RI), Relative Consistency Index (RC) and Homoplasy Index
(HI) calculated for trees generated under different optimality
criteria. The best fitting substitution model for each single
gene partition and the concatenated data set was determined
in MrModeltest 2.3 (Nylander 2004) for Bayesian inference
posterior probabilities and ML. Maximum likelihood analyses, including 1000 bootstrap replicates, were performed
using the RAxML-HPC2 on XSEDE (8.2.12) in the CIPRES
Science Gateway (Stamatakis 2014; Miller et al. 2017). The
general time reversible (GTR) model was used for nucleotide
substitution with a discrete gamma distribution plus invariant site (GTR + I + G). The bootstrap replicates were summarized onto the best scoring tree (Miller et al. 2017). The
Bayesian inference posterior probabilities (PP) distribution
(Zhaxybayeva and Gogarten 2002) was estimated by Markov
Chain Monte Carlo sampling (MCMC) in MrBayes 3.2.2 on
XSEDE (Ronquist and Huelsenbeck 2003). Six simultaneous Markov chains were run for 1,000,000 to 10,000,000
generations, depending on individual settings for the fungal
group. The resulted trees were sampled at one tree every
100th or 1000th generation. The first 10–25% of burn-in
phase of the analyses were discarded based on suitable burnin phases determined by using Tracer version 1.7 (Rambaut
et al. 2018). The remaining trees were used to calculate
13
8
Fungal Diversity (2020) 102:1–203
posterior probabilities in the majority-rule consensus (MRC)
trees (50%) with critical value for the topological convergence diagnostic set to 0.01.
FigTree v. 1.4 (Rambaut 2014) was used to visualize phylogenetic trees and data files and the phylogram was edited
using Adobe Illustrator CS v. 6 (Adobe Systems, USA). All
sequences generated in this study were submitted to GenBank. All entries are represented using phylogenetic tree
and relevant description.
multi-septate or muriform ascospores (Hyde et al. 2013).
The family comprises Amniculicola, Murispora, Neomassariosphaeria, Pseudomassariosphaeria and Vargamyces
(Zhang et al. 2009; Hyde et al. 2013; Ariyawansa et al.
2015a; Hernández-Restrepo et al. 2017). We introduce a
novel saprobic genus, Fusiformispora from Clematis collections in Thailand. Maximum likelihood and Bayesian
analyses of the combined dataset (LSU, SSU, ITS, tef1 and
rpb2) is shown in Fig. 2.
Genealogical concordance phylogenetic species
recognition analysis
Fusiformispora Phukhams. & K.D. Hyde, gen. nov.
Index Fungorum number: IF557106; Facesoffungi number: FoF 07242, Fig. 3.
Etymology: Genus name reflects the fusiform shape of
its ascospores.
Saprobic on decaying wood or herbaceous plant material
in terrestrial habitats. Sexual morph: Ascomata on surface
of the host, covered by a pseudoclypeus, visible as black
spots, solitary, scattered, uniloculate, obpyriform to compressed globose, coriaceous, brown to dark brown, ostiolate.
Ostioles central, brown to dark brown, papillate. Peridium
multilayered, cells of textura angularis, inner layers comprising thin, hyaline cells. Hamathecium composed of dense,
filiform, branched, transverly septate, trabecular pseudoparaphyses anastomosing above asci. Asci 8-spored, bitunicate, fissitunicate, thick-walled, cylindric-clavate, apically
rounded, short, with a furcate pedicel, with ocular chamber.
Ascospores biseriate, partially overlapping, broad fusiform,
tapering towards the acute ends, hyaline, with guttules in
each cell, constricted at the septa, smooth-walled, with a
thin mucilaginous sheath. Asexual morph: Undetermined.
Type species: Fusiformispora clematidis Phukhams.,
M.V. de Bult & K.D. Hyde
The closely related strains that resulted from morphology
and phylogeny evidence of recombination were prospectively analyzed using the genetic distances by performing
a pairwise homoplasy index test (Φw) (Taylor et al. 2000;
Bruen et al. 2006). A pairwise homoplasy index (PHI) test
was performed in SplitsTree (version 4.1.4.4) using the
Kimura’s two parameter (K2P) models for low genetic distance datasets. LogDet transformation were applied for the
average of nucleotide frequencies and splits decomposition
graph options (Gu and Li 1996a, b; Taylor et al. 2000; Bruen
et al. 2006; Huson and Bryant 2006; Gioan and Paul 2012;
Nishimaki and Sato 2019). The standard deviation of split
frequencies PHI test results (Φw) < 0.05 indicate significant
recombination within the dataset.
Taxonomy
Phylum Ascomycota R.H. Whittaker
The taxa are arranged as in the Outline of Fungi and
fungus-like organisms (Wijayawardene et al. 2016, 2020).
Subphylum Pezizomycotina Erikss. & K. Winka
Class Dothideomycetes sensu O.E. Erikss & Winka
For the classification of Dothideomycetes we follow Hyde
et al. (2013), Liu et al. (2017) and Hongsanan et al. (2020).
Subclass Pleosporomycetidae C.L. Schoch et al.
Pleosporales Luttrell ex M.E. Barr
Pleosporales is the largest and most diverse order in Dothideomycetes with over 75 families (Hongsanan et al. 2020).
Amniculicolaceae Y. Zhang, C.L. Schoch, J. Fourn., Crous
& K.D. Hyde
Amniculicolaceae was introduced for freshwater-associated ascomycetes. This family is characterized by solitary
ascomata with a rough black surface. The members usually stain the surface of the substrate purple and have short
pedicellate asci, with hyaline or pale brown or brown, 1- to
13
Fig. 2 The Bayesian 50% majority-rule consensus phylogram based ▸
on combined LSU, SSU, ITS, tef1 and rpb2 sequence data of related
families in Pleosporales. The topology and clade stability of the combined gene analyses was compared to the single gene analyses. The
tree is rooted with species of Hysteriales. One hundred and fiftythree strains were included in the DNA analyses which comprised
4394 characters (848 characters for LSU, 1044 characters for SSU,
556 characters for ITS, 910 characters for tef1, and 1036 characters
for rpb2, including gap regions). The tree from the maximum likelihood analysis had similar topology to the Bayesian analyses. The best
scoring RAxML tree had a final likelihood value of − 73089.933914.
The matrix had 2676 distinct alignment patterns, with 40.81% undetermined characters and gaps. Estimated base frequencies were as
follows; A = 0.246412, C = 0.245743, G = 0.272077, T = 0.235768;
substitution rates AC = 1.617324, AG = 3.695355, AT = 1.662826,
CG = 1.183453, CT = 8.283938, GT = 1.000000; gamma distribution
shape parameter α = 0.635095. The GTR + I + G model was selected
for every partition in Bayesian analysis. Bootstrap values (BS) greater
than 50% BS (ML, left) and Bayesian posterior probabilities (BYPP,
right) greater than 0.90 are given at the nodes. Hyphens (-) represent
support values less than 50% BS/0.90 BYPP. The type sequences are
in bold and the species determined in this study are indicated in blue
Fungal Diversity (2020) 102:1–203
9
Crassiperidium octosporum KT 3605
Crassiperidium octosporum MM 9
Crassiperidium
octosporum KT 3008
64/1.00
Crassiperidium octosporum KT 2894
64/0.99
Crassiperidium octosporum KT 3604
Crassiperidium octosporum KT 3046
Crassiperidium
octosporum KT 2144
94/0.99 1
Crassiperidium octosporum KT 3029
74/0.99
Crassiperidium octosporum KT 3188
98/1.00
Crassiperidium octosporum KT 3468
98/1.00
Crassiperidium quadrisporum KT 2798-1
100/1.00 Crassiperidium quadrisporum KT 27982
Longiostiolum tectonae MFLUCC 12-0562
Cyclothyriella rubronotata CBS 141486
85/1.00
Cyclothyriella rubronotata TR9
100/1.00
Cyclothyriella rubronotata TR
Cyclothyriella rubronotata CBS 419.85
Cyclothyriella rubronotata CBS 385.39
86/1.00
Massariosphaeria clematidis MFLU 16-0174
100/1.00
Massariosphaeria phaeospora CBS 611.86
100/1.00 Acrocalymma medicaginis CPC 24345
-100/0.99
Acrocalymma medicaginis CPC 24340
64/1.00
Acrocalymma ficus CBS 317.76
Acrocalymma aquatica MFLUCC 11-0208
--/0.99
100/1.00 Ascocylindrica marina MD6012
Ascocylindrica marina MD6011
79/0.98 100/1.00
Leptosphaeria doliolum CBS 505.75
Leptosphaeriaceae
Stemphylium vesicarium CBS 191.86
100/1.00
Pleosporaceae
69/1.00
70/0.99 Helminthosporium velutinum MFLUCC 13-0243
100/1.00 Helminthosporium velutinum MFLUCC 15-0423
66/1.00
Helminthosporium velutinum CBS 139923
Helminthosporium aquaticum MFLUCC 15-0357
100/1.00
--/0.93
Massarina eburnea CBS 473.64
Stagonospora pseudocaricis CBS 135132
57/0.93
Brevicollum hyalosporum MAFF 243400
Neohendersonia kickxii CBS 112403
100/1.00
Paradictyoarthrinium tectonicola MFLUCC 12-0556
98/0.99
61/0.94
Paradictyoarthrinium tectonicola MFLUCC 13-0465
100/1.00
Sirodesmium olivaceum CBS 395.59
60/0.99
Paradictyoarthrinium diffractum MFLUCC 12-0557
86/0.99 Paradictyoarthrinium diffractum MFLUCC 13-0466
Xenomassariosphaeria rosae CBS 612.86
77/1.00
100/1.00
100/1.00
Xenomassariosphaeria rosae MFLUCC 15-0179
Xenomassariosphaeria clematidis MFLUCC 14-0923
100/1.00 Ohleria modesta OM
--/0.99
Ohleria
modesta
MGC
54/0.94
Hobus wogradensis TI
Occultibambusa pustula MFLUCC 11-0502
100/1.00
99/1.00
Occultibambusa bambusae MFLUCC 11-0394
Versicolorisporium triseptatum HHUF 28815
100/1.00
Seriascoma didymospora MFLUCC 11-0179
Preussia lignicola CBS 264.69
100/1.00
99/1.00
Preussia lignicola CBS 363.69
86/1.00
Sporormiella minima CBS 524.50
Westerdykella ornata CBS 379.55
100/1.00
--/0.93
Westerdykella ornata CBS 297.56
Sporormia fimetaria UPS:Dissing Gr.81.194
--/0.92
Neomassarina pandanicola MFLUCC 16-0270
100/1.00
Neomassarina thailandica MFLU 11-0144
100/1.00
Neomassarina thailandica MFLUCC 17-1432
Neomassarina chromolaenae MFLUCC 17-1480
--/0.93
Pseudohelminthosporium clematidis MFLUCC 17–2086
--/0.92
Halotthia
posidoniae BBH 22481
94/1.00
93/1.00
Pontoporeia biturbinata BBH 23338
94/1.00
Mauritiana rhizophorae BCC 28866
Sulcosporium
thailandica
MFLUCC
12-0004
95/1.00
--/0.99
-Brunneoclavispora bambusae MFLUCC 11-0177
Neolophiostoma pigmenatum MFLUCC 10-0129
100/1.00 Berkleasmium nigroapicale BCC 8220
51/0.92
Berkleasmium micronesicum BCC 8141
100/1.00
Bahusandhika indica GUFCC 18001
-Lignosphaeria thailandica MFLUCC 11-0376
100/1.00
Lignosphaeria fusispora MFLUCC 11-0377
-100/1.00
Thyridaria macrostomoides GKM 1033
100/1.00
Thyridaria macrostomoides GKM 1159
Pleopunctum clematidis MFLUCC 17–2091
98/1.00
Phaeoseptum
aquaticum CBS 123113
92/-100/1.00
100/1.00
Phaeoseptum terricola MFLUCC 10-0102
Phaeoseptum malorum MFLUCC 17-2108
--/0.99
Decaisnella formosa BCC 25616
100/1.00
Decaisnella formosa BCC 25617
Angustimassarina populi MFLUCC 13-0034
51/-100/1.00
100/1.00
Angustimassarina quercicola MFLUCC 14-0506
Amorosia littoralis NN 6654
Lophiostoma arundinis CBS 621.86
--/0.94
100/1.00
Lophiostoma crenatum CBS 629.86
Pseudomassarina clematidis MFLU 16-0493
Pseudomassarinaceae
Longiostiolaceae
Cyclothyriellaceae
Acrocalymmaceae
Ascocylindricaceae
Massarinaceae
Neohendersoniaceae
Paradictyoarthriniaceae
Ohleriaceae
Occultibambusaceae
Sporormiaceae
Neomassarinaceae
Halotthiaceae
Lentimurisporaceae
Phaeoseptaceae
Amorosiaceae
Lophiostomataceae
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Fungal Diversity (2020) 102:1–203
Dendryphion europaeum CPC 22943
Dendryphion europaeum CPC 23231
Dendryphion nanum MFLUCC 16-0975
Neotorula aquatica MFLUCC 15-0342
100/1.00
69/0.99
Neotorula submersa KUMCC 15-0280
-100/1.00
Torula gaodangensis MFLUCC 17-0234
99/1.00
Torula herbarum CPC 24114
100/1.00
78/1.00
Rostriconidium aquaticum KUMCC 15-0297
Rostriconidium aquaticum MFLUCC 16-1113
100/1.00 Sporidesmioides thailandica KUMCC 16-0012
Sporidesmioides thailandica MFLUCC 13-0840
Lophiotrema nucula CBS 627.86
99/1.00
96/1.00
Lophiotrema eburnoides JCM 17826
Cryptoclypeus ryukyuensis MAFF 245615
97/1.00
Cryptocoryneum longicondensatum MAFF 245374
99/1.00
Cryptocoryneum japonicum MAFF 245370
99/1.00
82/1.00
Cryptocoryneum akitaense MAFF 245365
100/1.00 Aquasubmersa japonica MAFF 245219
100/1.00
Aquasubmersa japonica MAFF 245220
Aquasubmersa mircensis MFLUCC 11-0401
--/1.00
100/1.00 Pseudoberkleasmium chiangmaiense MFLUCC 17–2088
100/1.00
Pseudoberkleasmium chiangmaiense MFLUCC 17-1809 Pseudoberkleasmiaceae
Pseudoberkleasmium pandanicola MFLUCC 17-2264
--/1.00
Hermatomyces tectonae MFLUCC 14-1140
100/1.00
Hermatomyces indicus MFLUCC 14-1143
52/-61/1.00
Hermatomyces iriomotensis KT 2016
--/1.00 100/1.00
Antealophiotrema brunneosporum CBS 123095
Anteaglonium abbreviatum ANM 925.1
-98/1.00
Anteaglonium parvulum MFLUCC 14-0821
Anteaglonium globosum ANM 925.2
--/1.00
Clematidis italica MFLUCC 15-0084
100/1.00
Pseudolophiotrema elymicola JCM 13090
73/0.96
Pseudotetraploa
curviappendiculata JCM 12852
94/1.00
55/-Tetraploa sasicola JCM 13167
Triplosphaeria maxima JCM 13172
58/1.00
Byssolophis sphaerioides IFRDCC 2053
-Quadricrura septentrionalis CBS 125429
Polyplosphaeria fusca JCM 13175
--/0.96
Pseudoastrosphaeriella bambusae MFLUCC 11-0205
--/0.95
100/1.00
Pseudoastrosphaeriella longicolla MFLUCC 11-0171
Pseudoastrosphaeriellaceae
64/1.00
Pseudoastrosphaeriella thailandensis MFLUCC 10-0553
Aigialus grandis BCC 20000
100/1.00
Aigialus parvus BCC 18403
-Berkleasmium crunisia BCC 17023
100/1.00
Berkleasmium typhae BCC 12536
-100/1.00
Lindgomyces ingoldianus ATCC 200398
Aquamassariosphaeria typhicola CBS 609.86
100/1.00
60/0.94
Salsuginea ramicola KT 2597.1
Salsuginea ramicola CBS 125781
55/1.00
69/1.00 Vargamyces aquaticus HKUCC 10830
69/0.96
Vargamyces aquaticus CBS 639.63
71/1.00
Amniculicola immersa CBS 123083
88/1.00
Amniculicola parva CBS 123092
Murispora rubicunda CBS 291.56
98/1.00
Pseudomassariosphaeria bromicola MFLUCC 15-0031
100/1.00
100/1.00
Amniculicola lignicola CBS 123094
100/1.00
Fusiformispora clematidis MFLUCC 17-2077
Fusculina eucalyptorum CBS 145083
100/1.00
100/1.00
Fusculina eucalypti CBS 120083
Gordonomyces mucovaginatus CBS 127273
--/0.99
Neomassaria fabacearum MFLUCC 16-1875
Neomassaria formosana NTUCC 17-007
96/1.00
Delitschia didyma UME 31411
Delitschia winteri AFTOL-ID 1599
94/0.99
Hysterium angustatum MFLUCC 16-0623
Hysterium
angustatum
CBS 236.34
100/1.00
100/1.00
99/1.00
--/0.99
50/0.99
Torulaceae
Lophiotremataceae
Aquasubmersaceae
Hermatomycetaceae
Anteagloniaceae
Pseudolophiotremataceae
Tetraplosphaeriaceae
Aigialaceae
Lindgomycetaceae
Salsugineaceae
Amniculicolaceae
Fusculinaceae
Neomassariaceae
Delitschiaceae
Hysteriales
0.04
Fig. 2 (continued)
Notes: Fusiformispora is established as a monotypic
genus. In the multi-gene phylogenetic analyses, the isolate MFLUCC 17–2077 formed a basal lineage with other
genera in Amniculicolaceae (Fig. 2) with strong support
(100% ML/1.00 BYPP). The genus is compatible with the
13
concept of Amniculicolaceae in having compressed globose,
coriaceous, brown to dark brown ascomata, ostiolate, with
trabeculate, anastomosed pseudoparaphyses (sensu Liew
et al. 2000), and fusiform ascospores that are hyaline and
septate with mucilaginous appendages (Zhang et al. 2009).
Fungal Diversity (2020) 102:1–203
Fig. 3 Fusiformispora clematidis (MFLU 17–1485, holotype). a
Appearance of ascoma on host surface. b Close up of ascoma on host
substrate. c Vertical section of ascoma. d Ostiolar canal. e Section
11
of peridium. f Pseudoparaphyses. g–i Asci. j–m Ascospores. n Culture characteristics on MEA. Scale bars: b = 200 µm, c = 100 µm, d,
j–m = 20 µm, e–i = 50 µm
13
12
Fusiformispora is similar to Amniculicola Zhang & K.D.
Hyde, however, the genus differs by having thinner peridium
walls with sub-carbonaceous ascomatal type. Amniculicola
is an aquatic genus and its species have cylindrical asci and
uniseriate arrangement of ascospores, while Fusiformispora has cylindric-clavate asci and biseriate arrangement
of ascospores and is from a terrestrial habitat. We therefore,
introduce a new genus based on morphological and phylogenetic evidence for a fungal collection on Clematis fulvicoma.
Fusiformispora clematidis Phukhams., M.V. de Bult & K.D.
Hyde, sp. nov.
Index Fungorum number: IF557107; Facesoffungi number: FoF 07243, Fig. 3.
Etymology: Epithet reflects the host Clematis.
Holotype: MFLU 17–1485.
Saprobic on dead stems of Clematis fulvicoma. Sexual
morph: Ascomata 165–190 × 200–275 μm ( x̄ = 175 × 225 μm,
n = 5), on surface of host, covered by a pseudoclypeus, visible
as black spots, immersed to superficial, solitary, scattered,
uniloculate, obpyriform to compressed globose, base flattened, brown to dark brown, partially carbonaceous, roughwalled, with apical ostioles. Ostioles central, 55 × 35 μm,
brown to dark brown, papillate, with easily opening by a
pore, filled with periphyses. Peridium 10–18 μm wide, multilayered, comprising 4–5 layers of brown to dark brown
cells of textura angularis, inner layers comprising thin, hyaline cells. Hamathecium composed of dense, 0.5–1.5 μm
wide ( x̄ = 1.3 μm, n = 50), filiform, branched, trabeculate
pseudoparaphyses, anastomosing above the asci, reaching
the ostiole, transversely septate. Asci 86–127 × 18–24 μm
( x̄ = 100 × 25 μm, n = 40), 8-spored, bitunicate, fissitunicate,
thick-walled, cylindric-clavate, apically rounded, short, with
furcate pedicel, ocular chamber clearly visible when immature. Ascospores 24–36 × 5–10 μm ( x̄ = 30 × 8 μm, n = 50),
biseriate, partially overlapping, broad fusiform, tapering
towards the ends, acute at both ends, hyaline, with (1–)3–4
transverse septa, with large guttules in each cell, constricted
at the septa, deeply constricted at the median septum, cell
above median septum slightly wider than below, smoothwalled, with 4–12 μm wide mucilaginous sheath. Asexual
morph: Undetermined.
Culture characters: Colonies on MEA reaching 50 mm
diam. after 4 weeks at 25 °C. Cultures from above, greybrown, with reddish brown mixed in the mycelium, dense,
colonies circular, flat, umbonate, raised from the agar in the
centre, dull, covered with aerial mycelium, white mycelium
at the edge; reverse dark brown, dense, circular, with irregular, fimbriate margin, pinkish mycelium radiating outwardly.
Material examined: Thailand, Chiang Rai Province, on
dead stems of Clematis fulvicoma Rehder & E.H. Wilson, 20
March 2017, C. Phukhamsakda, CMTH22 (MFLU 17–1485,
holotype); ex-type living culture, MFLUCC 17–2077.
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Fungal Diversity (2020) 102:1–203
Host: Clematis fulvicoma—(This study).
Distribution: Thailand—(This study).
GenBank accession numbers: LSU: MT214542; SSU:
MT226661; ITS: MT310589; tef1: MT394725; rpb2:
MT394677.
Notes: In a BLASTn search of GenBank, the LSU
sequence of Fusiformispora clematidis MFLUCC 17–2077
showed 96% similarity to Lindgomyces pseudomadisonensis
KT 2742 (LC149916), while the ITS sequence had 91% similarity to Vargamyces aquaticus CBS 636.91 (NR_154471).
Fusiformispora clematidis is phylogenetically distinct, therefore we introduce the collection as a new species.
Amorosiaceae Thambug. & K.D. Hyde
Amorosiaceae was introduced for Amorosia Mantle &
D. Hawksw. and Angustimassarina Thambugala, Tanaka
& K.D. Hyde (Thambugala et al. 2015). Amorosiaceae is
characterized by immersed or semi-immersed ascomata
with a short crest-like papilla, and hyaline ascospores with
a mucilaginous sheath. Asexual morphs of this family
were described as sporodochia (Mantle et al. 2006; Thambugala et al. 2015). Jayasiri et al. (2019) reported Amorocoelophoma from a decaying pod of Cassia species. Phylogeny combined with morphological observations confirm
the placement of Amorocoelophoma as the first coelomycetous species in Amorosiaceae (Fig. 4).
Angustimassarina Thambug., Kaz. Tanaka & K.D. Hyde
Angustimassarina was described for fungal species that
have ascospores resembling Massarina while being narrowly
fusiform in shape. The genus has immersed to semi-immersed
ascomata, coriaceous, dark brown to black, globose to subglobose, ostiolate, cylindrical to cylindric-clavate asci and
fusiform to cylindrical or ellipsoidal-fusiform ascospores surrounded by a mucilaginous sheath. The asexual morph of this
genus is hyphomycetous. Twelve species are listed in Index
Fungorum for Angustimassarina (Thambugala et al. 2015;
Wanasinghe et al. 2018; Hyde et al. 2019a). In this study,
Angustimassarina rosarum was isolated from Clematis viticella and identification was based on multigene phylogenetic
analysis of LSU, SSU, ITS, and tef1 sequence data (Fig. 4)
and its compatible morphology (Fig. 5).
Angustimassarina rosarum Tibpromma, Camporesi & K.D.
Hyde, Fungal Diversity 89: [21] (2018), new host record.
Index Fungorum number: IF553939; Facesoffungi number: FoF 03964, Fig. 5.
Saprobic on dead stems of Clematis viticella. Sexual
morph: Ascomata 221–306 × 267–400 µm ( x̄ = 265 × 340 µm,
n = 5), scattered, gregarious, immersed, coriaceous, dark
globose to subglobose, brown to black, ostiolate. Ostioles
57 × 134 µm, central, rounded, papillate, with opening by a
pore. Peridium 14–40(–60) µm ( x̄ = 20 µm, n = 10), thick at
Fungal Diversity (2020) 102:1–203
Angustimassarina alni MFLUCC 15-0184
53/--
Angustimassarina italica MFLUCC 15-0082
ML/BYPP
Angustimassarina quercicola MFLUCC 15-0079
95/1.00
Angustimassarina quercicola MFLUCC 14-0506
Angustimassarina acerina MFLUCC 14-0505
82/1.00
Amorosiaceae
Fig. 4 The best scoring RAxML
tree with a final likelihood
value of − 10209.184183 based
on combined LSU, SSU, ITS
and tef1 sequence data for
Amorosiaceae. The topology and clade stability of the
combined gene analyses was
compared to the single gene
analyses. The tree is rooted
with species of Sporormiaceae.
The tree from the maximum
likelihood analysis had similar
topology to the Bayesian 50%
majority-rule consensus
phylogram. The matrix had
809 distinct alignment patterns
with 21.55% undetermined
characters and gaps. Estimated base frequencies were
as follows; A = 0.246053,
C = 0.243449, G = 0.269921,
T = 0.240577; substitution rates
AC = 1.221779, AG = 2.152972,
AT = 1.558241, CG = 0.998233,
CT = 7.007203, GT = 1.000000;
gamma distribution shape
parameter α = 2.180328. The
species determined in this study
is indicated in blue. Bootstrap
values (BS) greater than 50%
BS (ML, left) and Bayesian
posterior probabilities (BYPP,
right) greater than 0.90 are
given at the nodes. Hyphens (-)
represent support values less
than (BS ≥ 50%/BYPP ≥ 0.90)
13
Angustimassarina coryli MFLUCC 14-0981
Angustimassarina rosarum MFLUCC 17-2155
--/0.96
Angustimassarina rosarum MFLUCC 15-0080
Massarina corticola CBS 154.93
Angustimassarina populi MFLUCC 17-1069
53/--
Angustimassarina arezzoensis MFLUCC 13-0578
85/--
Angustimassarina populi MFLUCC 13-0034
Angustimassarina sylvatica MFLUCC 18–0550
100/1.00
Angustimassarina lonicerae MFLUCC 15-0088
Exosporium stylobatum CBS 160.30
77/1.00
Angustimassarina premilcurensis MFLUCC 15-0074
91/1.00
Amorocoelophoma cassiae MFLUCC 17-2283
Amorosia littoralis NN 6654
100/1.00
Lophiostoma macrostomum HHUF 27290
100/1.00
Floricolaceae
Guttulispora crataegi MFLUCC 13-0442
77/0.99
Floricola striata JK 5678I
100/1.00
Lophiostomataceae
Teichospora rubriostiolata TR7
Westerdykella ornata CBS 379.55
100/1.00
Preussia funiculata CBS 659.74
Sporormiaceae
(Out group)
0.01
the sides, broad at the apex and thinner at the base, comprising brown to dark brown cells of textura angularis,
fusing at the outside with the host tissues. Hamathecium
composed of dense, 1.5–2.5 µm wide, septate, long, cellular
pseudoparaphyses, embedded in a gelatinous matrix. Asci
77–85 × 10–16 µm ( x̄ = 78 × 15 µm, n = 30), 8-spored, bitunicate, fissitunicate, broad cylindrical to cylindrical-clavate,
with bulbous pedicel, rounded at the apex, with a minute
ocular chamber. Ascospores 17–23 × 4–4.5 µm ( x̄ = 20 × 4 µm,
n = 15), biseriate, partially overlapping, broad fusiform, hyaline, 1(–3) septate, deeply constricted at the primary septum,
widest at the centre and tapering towards the ends, straight,
smooth-walled, 1(–3)-guttulate, surrounded by a 5–10 µm
wide mucilaginous sheath. Asexual morph: Undetermined.
Culture characters: Colonies on MEA reaching 30 mm
diam. after 4 weeks at 25 °C. Culture black in the middle,
radiating white, dense, circular, umbonate, entries edge,
shiny, dull, undulate, radially furrowed, reverse black, radiating outwardly, white.
Material examined: Belgium, Flemish Brabant, Meise
Botanic Garden, Bouchout Domain, dead stems of Clematis
viticella L., 13 June 2017, D. Ertz & C. Gerstmans, BRCV1
(MFLU 17–1513); living culture, MFLUCC 17–2155.
Hosts: Clematis viticella, Rosa canina—(Wanasinghe
et al. 2018; this study).
Distribution: Belgium, Italy—(Wanasinghe et al. 2018,
this study).
GenBank accession numbers: LSU: MT214543; SSU:
MT226662; ITS: MT310590; tef1: MT394726; rpb2:
MT394678.
Notes: Angustimassarina rosarum (MFLUCC 15–0080)
was described from Rosa canina in Italy (Wanasinghe et al.
2018). We compared our collection with A. rosarum and
both have similar morphology in terms of ascomata, asci,
and ascospores. Angustimassarina rosarum (MFLUCC
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Fungal Diversity (2020) 102:1–203
Fig. 5 Angustimassarina rosarum (MFLU 17–1513). a Appearance
of ascomata on host surface. b Close up of ascoma on host substrate.
c Vertical section through ascoma. d Ostiolar canal. e Section of
peridium. f Pseudoparaphyses. g–h Asci. i–m Ascospores. Scale bars:
b = 500 µm, c = 200 µm, d–f = 50 µm, g, h = 20 µm, i–m = 10 µm
15–0080) has globose to subglobose, cylindric-clavate asci
(124 × 143 µm), and club-shaped pedicel (70 × 10 µm). It
has a minute ocular chamber, with fusiform to ellipsoidal
ascospores (19 × 5 µm) and a hyaline, 1 septate at the centre,
with two large guttules in each cell. Our collection has larger
ascomata (265 × 340 µm), but is similar in overall morphology (Fig. 5). In molecular analysis, the new strain forms a
close relationship with A. rosarum (MFLUCC 15–0080).
Comparison of the ITS sequence data reveals no significant
difference (one base pair difference) between our new collection and A. rosarum (MFLUCC 15–0080). However, the
tef1 sequence is not available for A. rosarum (MFLUCC
15–0080) for comparison. Therefore, we introduce a new
host record of A. rosarum on Clematis species herein.
Cyclothyriellaceae Jaklitsch & Voglmayr
Cyclothyriellaceae was introduced for Cyclothyriella
rubronotata (= Thyridaria rubronotata) and Massariosphaeria phaeospora as revealed by molecular phylogeny
(Jaklitsch and Voglmayer 2016). Cyclothyriellaceae is
characterized by scattered, immersed-erumpent ascomata,
with occasional purple stain on plant tissue, and narrow,
anastomosing, trabeculate pseudoparaphyses (sensu Liew
et al. 2000). Asci are cylindrical to clavate, bitunicate and
8-spored. Ascospores are brown, ellipsoid to fusoid, with
several eusepta. The asexual morph is pycnidial with hyaline
to brown conidia (Zhang et al. 2012; Jaklitsch and Voglmayer 2016). We describe a novel species of Massariosphaeria recorded on Clematis vitalba from Italy (Fig. 2).
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Massariosphaeria (E. Müll.) Crivelli
Massariosphaeria was introduced for species with reddish brown to brown, multi-septate, phragmosporous to
dictyosporous and usually with colouration on the surface
of the substrate (Crivelli 1983; Leuchtmann 1984; Zhang
et al. 2012). Several studies have proved the polyphyletic
placement of Massariosphaeria, classifying them into distinct genera (Zhang et al. 2012; Ariyawansa et al. 2015a;
Phukhamsakda et al. 2016). Massariosphaeria has comparable peridium and phragmosporous characters with Chaetomastia (Teichosporaceae) (Barr 1989). However, the characteristic of ascomata position and the ascospores characters of
Chaetomastia and Massariosphaeria are distinct. The type
species Massariosphaeria phaeospora (CBS 611.86) is currently placed in Cyclothyriellaceae (Fig. 2). Twenty-one epithets are listed under Massariosphaeria in Index Fungorum
(2020). Based on phylogenetic analysis including Cyclothyriellaceae, our collection (MFLU 16–0174) clustered with
M. phaeospora (CBS 611.86) with strong support (100%
MLBS/1.00 BYPP). A new Massariosphaeria species on
Clematis vitalba is introduced herein (Fig. 6).
Massariosphaeria clematidis Phukhams., Wanas., Camporesi & K.D. Hyde, sp. nov.
Index Fungorum number: IF557108; Facesoffungi number: FoF 07248, Fig. 6.
Etymology: Epithet reflects the host Clematis.
Holotype: MFLU 16–0174.
Saprobic on dead stems of Clematis vitalba. Sexual morph: Ascomata 340–430 × 215–300 μm
( x̄ = 280 × 255 μm, n = 5), with only black shiny ostioles
present on the surface of host, solitary, scattered, immersed,
globose to compressed globose, sub-carbonaceous to coriaceous, dark brown to black, rough-walled, with short hyphae
projecting from peridium, ostiolate. Ostioles centrally
located, 125–175 × 110–130 μm ( x̄ = 140 × 120 μm, n = 10),
carbonaceous, papillate, periphysoids. Peridium 22–33 μm
wide, composed of 6–8(–12 at apex) layers of dark brown
to black cells of textura angularis, inner layer composed
of hyaline gelatinous cells. Hamathecium composed of
numerous, dense, long, 1.6–3.5 μm wide ( x̄ = 2.5 μm,
n = 50), filiform, transversely septate, branched, anastomosing, cellular pseudoparaphyses. Asci 150–225 × 20–30 μm
( x̄ = 180 × 25 μm, n = 20), 8-spored, bitunicate, fissitunicate,
cylindric-clavate to broad cylindrical, with furcate pedicel,
with ocular chamber visible when immature. Ascospores
35–45 × 10–15 μm ( x̄ = 40 × 12 μm, n = 50), biseriate or
overlapping, broad fusiform, narrow towards the apex, initially hyaline, becoming yellowish to brown at maturity,
6–8-transversely euseptate, constricted at the septa, third
cell from apex usually enlarged, smooth-walled, guttulate
and indentations present, surrounded by a 8–20 μm wide,
mucilaginous sheath. Asexual morph: Undetermined.
15
Culture characters: Colonies on MEA reaching 20 mm
diam. after 4 weeks at 18 °C. Culture from above brown,
yellowish towards the edge, dense, circular, flat, dull, fimbriate, radially furrowed, and slightly covered with white aerial
mycelium; reverse black with radiating cream mycelium.
Material examined: Italy, Forlì-Cesena Province, Strada
San Zeno—Galeata, dead aerial stems of Clematis vitalba
L., 7 November 2013, E. Camporesi, IT1509 (MFLU
16–0174, holotype).
Host: Clematis vitalba—(This study).
Distribution: Italy—(This study).
GenBank accession numbers: LSU: MT214544; SSU:
MT226663; ITS: MT310591.
Notes: Our new collection MFLU 16–0174 from Italy
formed a close relationship with the type species of Massariosphaeria, M. phaeospora (CBS 611.86) with strong
support (100% ML/1.00 BYPP). The strain is compatible
with the concept of Massariosphaeria in having scattered,
immersed, papillate ascomata with thick ostiolar wall, dense
pseudoparaphyses, cylindro-clavate asci and broad fusiform,
reddish brown to brown, multi-septate ascospores with a
mucilaginous sheath (Müller 1950; Zhang et al. 2012).
MFLU 16–0174 is distinguishable from M. phaeospora by
its partial carbonaceous ostioles, narrower, 6–8-transversely
euseptate ascospores which are swollen at the third cell.
Massariosphaeria vitalbae (≡ Leptosphaeria vitalbae) was
described from Clematis vitalba in Switzerland (Ahn and
Shearer 1998). Characters of MFLU 16–0174 include a long
neck ( x̄ = 140 × 120 μm), with sub-carbonaceous to coriaceous peridium types and ascospores enlarged at the third
cell. Massariosphaeria vitalbae has sub-parenchymatous
cells type with short ostiolar necks and 9–10-septate and
ascospores enlarged at the third cell (Müller 1950). Massariosphaeria vitalbae closely resembles Paramassariosphaeria clematidicola, however, fresh collections are required to
clarify its taxonomic placement (Wanasinghe et al. 2016b).
In a BLASTn search of GenBank, the LSU sequence of
Massariosphaeria clematidis (MFLU 16–0174) was found
to be 96% similar to M. phaeospora strain CBS 611.86
(FJ795503). We introduce a novel species of Massariosphaeria, M. clematidis based on the morphological (Fig. 6) and
phylogenetic evidence (Fig. 2).
Dictyosporiaceae Boonmee & K.D. Hyde
Dictyosporiaceae was introduced with Dictyosporium as
the type genus in Dothideomycetes (Boonmee et al. 2016).
The sexual morph of Dictyosporiaceae has immersed to
erumpent or superficial, globose to subglobose and dark
brown to black ascomata including bitunicate, cylindricclavate asci with septate and hyaline ascospores with a thick
mucilaginous sheath (Tanaka et al. 2015; Boonmee et al.
2016). Most members of Dictyosporiaceae have a hyphomycetous asexual morph with cheirosporous or dictyosporous
13
16
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Fungal Diversity (2020) 102:1–203
Fungal Diversity (2020) 102:1–203
◂Fig. 6 Massariosphaeria clematidis (MFLU 16–0174, holotype).
a Appearance of ascomata on host surface. b Vertical section of
ascoma. c Ostiolar canal. d Section of peridium. e Pseudoparaphyses.
f–i Asci. j–m Ascospores. n Ascospore in 10% Indian ink. Scale bars:
a = 500 µm, b = 200 µm, c = 100 µm, d–i = 50 µm, j–n = 20 µm
conidia, pycnidia with coleophoma-like characters and phialidic conidiogenesis cells. Fourteen genera are listed under
Dictyosporiaceae (Iturrieta-González et al. 2018; Wijayawardene et al. 2018; Crous et al. 2019). We provide an
updated phylogenetic tree of Dictyosporiaceae and propose a
new species and a new host record of Aquadictyospora, Dictyocheirospora and Pseudocoleophoma on Clematis (Fig. 7).
Aquadictyospora Luo, Hyde & H.Y. Su
Aquadictyospora was introduced to accommodate a dictyosporous taxon on submersed decaying wood, and typified
with A. lignicola Z.L. Luo et al. The genus is characterized
by sporodochia, superficial, circular or subglobose conidiomata, micronematous conidiophores with monoblastic conidiogenesis cells, and uniformly medium brown dictyosporous conidia with a subglobose, hyaline cell at the basal end
(Li et al. 2016). We introduce a second species of Aquadictyospora based on morphology (Fig. 8) and phylogenetic
analyses (Fig. 7).
Aquadictyospora clematidis Phukhams., Bhat & K.D. Hyde,
sp. nov.
Index Fungorum number: IF557125; Facesoffungi number: FoF 07250, Fig. 8.
Etymology: Name refers to the host plant, Clematis.
Holotype: MFLU 17–1488.
Saprobic on dead stems of Clematis sikkimensis. Sexual
morph: Undetermined. Asexual morph: Hyphomycetous.
Colonies 53–97 × 121–213 μm ( x̄ = 72 × 153 μm, n = 10),
on natural substrate forming sporodochial conidiomata,
superficial, compact, scattered, subglobose to oval, dark
brown to reddish brown, velvety. Mycelium 2–3 μm wide,
immersed, consisting of branched septate hyphae. Conidiophores 6–11 × 2–4 μm ( x̄ = 8 × 3 μm, n = 10), micronematous, hyaline to pale brown, smooth. Conidiogenous cells
5–8 × 3–5 μm, holoblastic, monoblastic, solitary, discrete,
determinate. Conidia 17–38 × 15–24 μm ( x̄ = 32 × 20 μm,
n = 50), dictyosporous, compactly depressed, obovoid,
appearing broadly rounded in upper half, heavily pigmented at the upper half, smooth, entirely reddish brown,
with hyaline mammiform basal cell, smooth basal cell,
5–11 × 4–11 μm ( x̄ = 8 × 8 μm, n = 40), not complanate,
secession involves splitting of the basal, without appendages.
Culture characters: Colonies on MEA reaching 50 mm
diam. after 4 weeks at 25 °C. Cultures from above brownish
beige at the centre, grey radiating outwardly, dense, raised
with concave edge, circular, entire edge, umbonate, papillate
17
with fairly fluffy, wrinkled folded, covered with white aerial
mycelium; reverse dark brown at the centre, faintly zonate,
white mycelium at the edge.
Material examined: Thailand, Nan Province, on dead stem
of Clematis sikkimensis (Hook. f. & Thomson) Drumm. ex
Burkill, 2 May 2017, C. Phukhamsakda, CMTH26 (MFLU
17–1488, holotype); ex-type living culture, MFLUCC
17–2080.
Host: Clematis sikkimensis—(This study).
Distribution: Thailand—(This study).
GenBank accession numbers: LSU: MT214545; SSU:
MT226664; ITS: MT310592; tef1: MT394727, rpb2:
MT394679.
Notes: The species is assigned to Aquadictyospora based
on its compatible morphological features such as superficial sporodochia with subglobose to oval conidiomata,
micronematous conidiophores with monoblastic conidiogenous cell, and dictyosporous conidia with hyaline, mammiform basal cells (Li et al. 2016). Aquadictyospora clematidis has smaller non-cheiroid conidia (32 × 20 μm) than A.
lignicola which has larger cheiroid conidia (50 × 24 μm).
Aquadictyospora clematidis was found in a terrestrial habitat
while A. lignicola was from submerged substrates (Fig. 8).
In a BLASTn search of GenBank, the ITS sequence had
95% similarity while the tef1 sequence had 95% similarity
to the type species, A. lignicola. The new strain is introduced
as a new species of Aquadictyospora based on polyphasic
evidence.
Dictyocheirospora D’souza, Boonmee & K.D. Hyde
Boonmee et al. (2016) introduced Dictyocheirospora
(typified by D. rotunda) for an aero-aquatic sporodochial
fungus with cheiroid dictyospores. Nineteen species are
listed in Index Fungorum (2020) and interestingly, no sexual
morph has been described for this genus (Wang et al. 2016;
Tibpromma et al. 2018; Hyde et al. 2017, 2019a). We introduce Dictyocheirospora clematidis as a novel species and
D. xishuangbannaensis as a new host record from Clematis
based on compatible morphological and phylogenetic analysis (Fig. 9).
Dictyocheirospora clematidis Phukhams., D.J. Bhat & K.D.
Hyde, sp. nov.
Index Fungorum number: IF557126; Facesoffungi number: FoF 07251, Fig. 9.
Etymology: Name refers to the host plant, Clematis.
Holotype: MFLU 17–1497.
Saprobic on dead stem of Clematis sikkimensis. Sexual
morph: Undetermined. Asexual morph: Hyphomycetous.
Colonies 200–340 μm wide ( x̄ = 265 μm, n = 20), on natural substrate forming sporodochial conidiomata, superficial,
gregarious, scattered, punctiform, blackish brown, velvety,
glistening, orbicular, with abundant sporulation, conidia
13
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Fungal Diversity (2020) 102:1–203
100/1.00
79/0.99
100/1.00
Dictyosporium subramanianii BCC 3503
Dictyocheirospora indica MFLUCC 15–0056
Dictyocheirospora xishuangbannaensis MFLUCC 17–2267
Dictyocheirospora xishuangbannaensis MFLUCC 17–2087 DT3
Dictyocheirospora lithocarpi MFLUCC 17–2537
-Dictyocheirospora heptaspora CBS 396.59
Dictyocheirospora aquadulcis MFLUCC 17–2571
Dictyocheirospora bannica KH 332
69/1.00
Dictyocheirospora vinaya MFLUCC 14–0294
--/0.94
80/0.98
Dictyocheirospora pandanicola MFLUCC 16–0365
99/1.00
Dictyocheirospora pseudomusae KH 412
84/1.00
Dictyocheirospora pseudomusae yone 234
90/1.00
Dictyocheirospora gigantica BCC 11346
100/1.00 Dictyocheirospora metroxylonis MFLUCC 15–0028b
79/1.00
Dictyocheirospora metroxylonis MFLUCC 15–0028
89/1.00
Dictyocheirospora clematidis MFLUCC 17–2089 DT6
100/1.00
Dictyocheirospora taiwanense MFLUCC 17–2654
100/1.00 Dictyocheirospora rotunda KUMCC 18–0014
Dictyocheirospora rotunda MFLUCC 14–0293
99/1.00
95/1.00
Dictyocheirospora cheirospora KUMCC 17–0035
88/1.00 Dictyocheirospora garethjonesii MFLUCC 16–0909
100/1.00
89/1.00
Dictyocheirospora garethjonesii KUMCC 15–0396
Dictyocheirospora nabanheensis MFLUCC 17–0562
100/1.00
Dictyocheirospora aquatica KUMCC 15–0305
Digitodesmium chiangmaiense KUN–HKAS 102163
66/0.99
Digitodesmium bambusicola CBS 110279
Aquaticheirospora lignicola RK 2006a
66/0.97
99/0.99 Jalapriya pulchra LQXM47
Jalapriya pulchra MFLUCC 15–0348
96/1.00
Jalapriya inflata NTOU 3855
94/1.00
--/0.96
68/0.94 Dictyosporium sp. 19VA07
85/1.00
100/1.00
Dictyocheirospora
Digitodesmium
Aquaticheirospora
Jalapriya
Jalapriya toruloides FMR11942
Jalapriya toruloides CBS 209.65
Vikalpa
Vikalpa australiensis HKUCC 8797
Dictyosporium alatum ATCC 34953
Dictyosporium elegans NBRC 32502
76/1.00
Dictyosporium hughesii KT 1847
52/1.00
Dictyosporium
Dictyosporium aquaticum MF 1318
60/0.94
Dictyosporium tubulatum MFLUCC 15–0631
99/1.00
Dictyosporium sexualis MFLUCC 10–0127
Dictyosporium meiosporum MFLUCC 10–0131
96/1.00
Aquadictyospora clematidis MFLUCC 17–2080 CMTH26
Aquadictyospora
80/0.99
Aquadictyospora lignicola MFLUCC 17–1318
65/1.00 Pseudodictyosporium wauense KRP 88-6
87/0.97 Pseudodictyosporium wauense NBRC 30078
76/-100/1.00
Pseudodictyosporium
Pseudodictyosporium indicum CBS 471.95
100/1.00
Pseudodictyosporium elegans CBS 688.93
Pseudodictyosporium thailandica MFLUCC 16–0029
Cheirosporium triseriale HMAS 180703 Cheirosporium
Pseudocoleophoma bauhiniae MFLUCC 17–2280
98/1.00
---/0.97
Pseudocoleophoma bauhiniae MFLUCC 17–2586
52/-Pseudocoleophoma polygonicola KT 731
Pseudocoleophoma calamagrostidis KT 3284
Pseudocoleophoma
100/1.00
Pseudocoleophoma clematidis MFLUCC 17–2177
100/-Pseudocoleophoma clematidis MFLUCC 17–2177B
Pseudocoleophoma typhicola MFLUCC 16–0123
Neodendryphiella tarraconensis FMR 16234
100/1.00
100/1.00
Neodendryphiella mali CBS 139.95 Neodendryphiella
100/1.00
Dendryphiella fasciculata MFLUCC 17–1074
Dendryphiella
Dendryphiella paravinosa CBS 141286
96/1.00
Gregarithecium curvisporum KT 922
Gregarithecium
Gregarithecium sp. MFLUCC 13–0853
100/1.00
Murilentithecium clematidis MFLUCC 14–0562
Lentitheciaceae
Murilentithecium clematidis MFLUCC 14–0561
100/1.00
98/1.00
(Out group)
0.04
13
Dictyosporiaceae
ML/BYPP
Fungal Diversity (2020) 102:1–203
◂Fig. 7 The best scoring RAxML tree with a final likelihood value
of −15336.133569 based on LSU, ITS and tef1 sequence data for
Dictyosporiaceae species. The topology and clade stability of the
combined DNA analyses was compared to the single gene analyses.
The tree is rooted with sequences of Murilentithecium clematidis
(MFLUCC 14–0561, MFLUCC 14–0562) in Lentitheciaceae. The
tree from the maximum likelihood analysis had similar topology to
the Bayesian analyses. The matrix had 1009 distinct alignment patterns, with 37.31% undetermined characters and gaps. Estimated
base frequencies were as follows; A = 0.234307, C = 0.254705,
G = 0.271061, T = 0.239927; substitution rates AC = 1.414609,
AG = 2.411925, AT = 2.055489, CG = 0.612591, CT = 6.466528,
GT = 1.000000; gamma distribution shape parameter α = 0.761212.
The species determined in this study are indicated in blue. Bootstrap
values (BS) greater than 50% BS (ML, left) and Bayesian posterior
probabilities (BYPP, right) greater than 0.90 are given at the nodes.
Hyphens (-) represent support values less than 50% BS/0.90 BYPP.
Thick branches represent significant support values from all analyses
(BS ≥ 70%/BYPP ≥ 0.95)
readily liberated when agitated. Mycelium immersed, composed of brown, smooth, thin-walled, septate hyphae. Conidiophores 15 × 3 μm, micronematous, pale brown, smooth,
thin-walled. Conidiogenous cells 5–6 × 3–4 μm, holoblastic, monoblastic, integrated, terminal, determinate, hyaline,
smooth-walled. Conidia 42–60 × 15–30 μm ( x̄ = 50 × 23 μm,
n = 40), solitary, acrogenous, cheiroid, with a basal connecting cell, discharges after mounted in water, cognac brown,
consisting of 6–7 rows of cells, individual rows discoid.
Conidial arm 34–60 × 7–10 μm ( x̄ = 52 × 9 μm, n = 30), digitate, cylindrical, inwardly curved at the tip, arising from a
basal cell, 10–12 distosepta, slightly constricted at the septa,
with large guttule in each cell.
Culture characters: Colonies on MEA reaching 40 mm
diam. after 4 weeks at 25 °C. Cultures from above, black,
dense, circular, entire edge, umbonate, papillate with aerial
mycelium, wrinkled and folded, narrow fringe of submerged
mycelium, covered with grey aerial mycelium; reverse black,
white mycelium present at the edge.
Material examined: Thailand, Chiang Rai Province, Doi
Tung, on dead stem of Clematis sikkimensis, 2 May 2017,
C. Phukhamsakda & M.V. de Bult, CMTHDT06 (MFLU
17–1497, holotype); ex-type living culture, MFLUCC
17–2089.
Host: Clematis sikkimensis—(This study).
Distribution: Thailand—(This study).
GenBank accession numbers: LSU: MT214546; SSU:
MT226665; ITS: MT310593; tef1: MT394728, rpb2:
MT394680.
Notes: Dictyocheirospora species are highly diverse
especially in tropical regions (Boonmee et al. 2016; Tibpromma et al. 2018; Yang et al. 2018a; Hyde et al. 2019a).
Dictyocheirospora clematidis is similar to D. metroxylonis based on characters (Table 1), but the ITS sequence
shows 97% similarity (2.8% nucleotide differences) while
the tef1 sequence has 96% similarity (4.6% nucleotide
19
differences). Dictyocheirospora clematidis clustered in the
same clade as D. taiwanense but it has distinct characters.
Dictyocheirospora taiwanense usually has 5 rows of cells in
the conidia and they are longer than those of D. clematidis
(78 × 18 vs 50 × 23 μm). In the phylogenetic analysis, D.
clematidis (MFLUCC 17–2089) clustered with D. metroxylonis (MFLUCC 15–0282) and D. taiwanense (MFLUCC
17–2654) but formed a well-separated clade with good support of 79% in ML and 1.00 in BYPP (Fig. 7).
Dictyocheirospora xishuangbannaensis Tibpromma &
K.D. Hyde, Fungal Divers 93:14 (2018), new host record
Index Fungorum number: IF554476; Facesoffungi number: FoF 04485, Fig. 10
Saprobic on dead stem of Clematis sikkimensis. Sexual
morph: Undetermined. Asexual morph: Hyphomycetous. Colonies 235–423 μm wide ( x̄ = 326 μm, n = 20),
on natural substrate forming sporodochial conidiomata,
superficial, gregarious, scattered, punctiform, blackish
brown, velvety, glistening, orbicular, with abundant sporulation, conidia readily liberated when agitated. Mycelium 2 μm wide, immersed, composed of hyaline to pale
brown, smooth, thin-walled, septate hyphae. Conidiophores
10–20 × 3–6 μm ( x̄ = 12 × 4 μm, n = 10), micronematous,
hyaline to light brown, smooth, thin-walled. Conidiogenous cells 3–7 × 2–6 μm ( x̄ = 5 × 4 μm, n = 10), holoblastic,
integrated, terminal, determinate, hyaline, smooth-walled.
Conidia 32–53 × 16–27 μm ( x̄ = 46 × 20 μm, n = 50), solitary,
acrogenous, cheiroid, oblong or subglobose, with a basal
connecting cell, discharges after mounted in water, cognac
brown, heavily pigmented upper part, consisting of 6–7
rows of cells, rows digitate. Conidial arm 40–55 × 6–9 μm
( x̄ = 46 × 6 μm, n = 30), cylindrical, curved at both ends, arising from a basal cell, reddish brown, heavily pigmented at
the upper part, 7–11 distoseptate, slightly constricted at the
septa, with large guttule in each cell.
Culture characters: Colonies on MEA reaching 40 mm
diam. after 4 weeks at 25 °C. Cultures from above, dark
brown, dense, circular, edge entire, umbonate, papillate with
white aerial mycelium, wrinkled and folded, narrow fringe
of submerged mycelium; reverse dark brown, erose.
Material examined: Thailand, Chiang Rai Province,
on dead stem of Clematis sikkimensis, 2 May 2017, C.
Phukhamsakda & M.V. de Bult, CMTHDT03 (MFLU
17–1495); living culture, MFLUCC 17–2087.
Hosts: Clematis sikkimensis, Pandanus sp.—(Tibpromma
et al. 2018; this study).
Distribution: China, Thailand—(Tibpromma et al. 2018;
this study).
GenBank accession numbers: LSU: MT214547; SSU:
MT226666; ITS: MT310594; tef1: MT394729.
Notes: Dictyocheirospora xishuangbannaensis was
recorded from Pandanus by Tibpromma et al. (2018) and our
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◂Fig. 8 Aquadictyospora clematidis (MFLU 17–1488, holotype). a,
b Sporodochia on natural substrate. c Vertical section through sporodochium. d–e Conidia attached to conidiophores. f–k Mature conidia.
l Germinated conidia. m, n Culture characteristics on MEA. Scale
bars: b = 100 μm, c = 100 μm, d–l = 20 μm
collection resembles this species morphologically (Fig. 10).
The first record of D. xishuangbannaensis was from southern China (Xishuangbanna, Yunnan Province), while our
collection was found in the northern part of Thailand (Chiang Rai Province). Northern Thailand and Xishuangbanna
both have a tropical climate and share similar weather in
both the wet and dry seasons (Cao et al. 2006). It can be
hypothesized that D. xishuangbannaensis generally occurs
in tropical climates (Boonmee et al. 2016; Tibpromma
et al. 2018; Yang et al. 2018a). This is the first record of D.
xishuangbannaensis on Clematis species. We also provide
sequence data and phylogenetic analyses in Fig. 7.
Pseudocoleophoma Tanaka & K. Hiray.
Pseudocoleophoma is characterized by its immersed to
erumpent, ostiolate ascomata, cylindrical to clavate and
short pedicellate asci and fusiform, 1-septate, sheathed
ascospores. The genus produces coleophoma-like asexual
morph with phialidic, doliiform to lageniform conidiogenous
cells and cylindrical, hyaline conidia. We introduce a new
species in Pseudocoleophoma based on molecular and morphology (Figs. 7, 11).
Pseudocoleophoma clematidis Phukhams. & K.D. Hyde,
sp. nov.
Index Fungorum number: IF557127; Facesoffungi number: FoF 07252, Fig. 11.
Etymology: Name refers to the host plant, Clematis from
which the fungus was isolated.
Holotype: MFLU 16–0280.
Saprobic on dead stems of Clematis vitalba. Sexual
morph: Undetermined. Asexual morph: Conidiomata
130–150 × 100–130 μm ( x̄ = 140 × 110 μm, n = 5), pycnidial, solitary, aggregated, uniloculate, immersed, with black
shiny ostioles visible, globose to subglobose, coriaceous,
subcoriaceous at the outer layers, thick-walled, black to
dark brown, ostioles. Ostioles 30 × 50 μm, central, papillate, ovoid. Conidiomatal wall 20–30 μm wide, of equal
thickness, multilayered, outer layer composed of 8–10 layers of light brown to brown cells of textura angularis, lined
with a thick hyaline layer bearing conidiogenous cells. Conidiophores reduced to conidiogenous cells. Conidiogenous
cells 2–4 × 1.5–4 μm ( x̄ = 2.5 × 3 μm, n = 30), holoblastic,
phialidic, determinate, discrete, cylindrical to subcylindrical,
smooth-walled, hyaline, arising from inner layers of conidioma. Conidia 5–8 × 2–4 μm ( x̄ = 6 × 4 μm, n = 50), oval,
21
slightly curved towards the ends, aseptate, with 1(–2) guttules in each cell, hyaline when immature, yellowish brown
at maturity, smooth-walled.
Culture characters: Colonies on MEA reaching 20 mm
diam. after 2 weeks at 16 °C. Cultures from above, cream,
dense, circular, umbonate, papillate with fluffy, covered with
white aerial mycelium; reverse dark brown at the centre,
cream radiating outwardly.
Material examined: Italy, Arezzo Province, Badia Tega—
Ortignano Raggiolo, on dead aerial branch of Clematis
vitalba, 9 March 2013, E. Camporesi, IT 1110 (MFLU
16–0280, holotype); ex-type living culture, MFLUCC
17–2177.
Host: Clematis vitalba—(This study).
Distribution: Italy—(This study).
GenBank accession numbers: LSU: MT214548,
MT214549; SSU: MT226667; ITS: MT310595, MT310596;
tef1: MT394730.
Notes: Based on the multi-gene phylogenetic analyses
(Fig. 7), Pseudocoleophoma clematidis strain MFLUCC
17–2177 (Fig. 11) clusters between P. calamagrostidis
(KT 3284) and P. typhicola (MFLUCC 16–0123). Pseudocoleophoma clematidis is different from other Pseudocoleophoma species by having pycnidial walls which are
flat at the base and yellowish brown conidia (Tanaka et al.
2015; Hyde et al. 2016; Jayasiri et al. 2019, Fig. 11). This
study confirmed its placement in Pseudocoleophoma. In
a BLASTn search of GenBank, the closest match of the
LSU sequence of MFLUCC 17–2177 is P. calamagrostidis
(HHUF 30450) with 97% similarity, while the closest match
of the ITS sequence is P. polygonicola (HHUF 27558) with
92% similarity.
Didymellaceae Gruyter, Aveskamp & Verkley
Valenzuela-Lopez et al. (2018) accepted 26 genera in
Didymellaceae. Didymellaceous taxa frequently occur
as plant pathogens, causing drooping and wilting of plant
leaves or gummy stem blight leading to death of the plant
(Sudisha et al. 2004; Vaghefi et al. 2012; Ahmadpour et al.
2017; Wijayawardene et al. 2017). The combined dataset of
LSU, ITS and rpb2 sequences for a multilocus analysis tree
revealed distinct lineages in Didymellaceae (Fig. 12). In the
present study, a cluster of fungi associated with Clematis
species formed a distinct lineage, Anthodidymella, a novel
genus in Didymellaceae, a novel species in Xenodidymella,
and a new host record in Neodidymelliopsis.
Anthodidymella Phukhams., Camporesi & K.D. Hyde, gen.
nov.
Index Fungorum number: IF557128; Facesoffungi number: FoF 07255, Fig. 13
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Fig. 9 Dictyocheirospora clematidis (MFLU 17–1497, holotype). a, b Sporodochia on natural substrate. c Sporodochium mounted in water. d–i
Mature conidia. j Culture characteristics on MEA. Scale bars: b = 500 μm, c = 100 μm, d–g = 20 μm, h, i = 10 μm
Table 1 Synopsis of related Dictyocheirospora species and the new species from this study
Species
Conidia
References
Average size (µm) Shape
D. clematidis
50 × 23
D. metroxylonis 60 × 20
D. taiwanense 78 × 18
13
Cheiroid, acrogenous, cylindrical, cognac brown, consisting of 6–7 rows of
cells
Cheiroid dictyospores, cylindrical, pale brown, consisting of 4–6 rows of cells
Cheiroid dictyospores, brown, ellipsoid to cylindrical, consisting of 5 rows of
cells
This study
Phookamsak et al. (2019)
Hyde et al. (2019a)
Fungal Diversity (2020) 102:1–203
23
Fig. 10 Dictyocheirospora xishuangbannaensis (MFLU 17–1495).
a, b Sporodochia on natural substrate. c Sporodochium mounted in
water. d–h Mature conidia. i–j Separated rows of conidium. k Cul-
ture characteristics on MEA. Scale bars: b = 200 μm, c = 100 μm,
d–e = 50 μm, f–g = 20 μm. h–j = 10 μm
Etymology: Anthos-meaning flower, Anthodidymella refer
to species of Didymella that frequently occur on flowering
plants.
Saprobic or necrotic on leaf and dead stems of herbaceous
plants Sexual morph: Ascomata superficial, solitary or clustered, globose or subglobose to pyriform, with elongated
ostioles. Perithecial wall consisting of textura globulosa.
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Fig. 11 Pseudocoleophoma clematidis (MFLU 16–0280, holotype).
a Appearance of conidiomata on Clematis vitalba. b Vertical section
through conidioma. c Ostiolar canal. d Section of conidioma wall.
e–g Conidiogenous cells and conidia. h Conidia. i Culture characteristics on MEA. Scale bars: b = 200 µm, c = 50 µm, d = 20 µm, e–
h = 5 µm
Hamathecium composed of numerous, dense, pseudoparaphyses. Asci 8-spored, bitunicate, cylindrical club-shaped
pedicel. Ascospores uniseriate or partially overlapping,
ovate to obpyriform, 1-septate, hyaline (Woudenberg et al.
2009). Asexual morph: Conidiomata pycnidial, uniloculate,
immersed under host epidermis, subglobose to depressed,
coriaceous, thin-walled, dark brown to brown, with papillate
ostioles. Conidiomatal wall thin layers, pseudoparenchymatous, with brown cells of textura globulosa, lined with a hyaline cell-layer bearing conidiogenous cells. Conidiophores
reduced to conidiogenous cells. Conidiogenous cells phialidic, determinate, discrete, ampulliform, cylindrical to subcylindrical, hyaline. Conidia oblong or oval, rounded ends,
hyaline, aseptate or septate, smooth-walled. Chlamydospores
absent.
Type species: Anthodidymella ranunculacearum
Phukhams., Camporesi & K.D. Hyde
Notes: Anthodidymella is introduced for a strongly supported clade (93% ML/1.00 BYPP, Fig. 12) of Didymella
species unit that is associated with Clematis (Aveskamp
et al. 2010). Anthodidymella clematidis was described as
Phoma clematidina as it clustered with other Phoma clematidina isolates (Woudenberg et al. 2009; Golzar et al. 2011).
Phoma clematidina not only causes symptoms on Clematis
species, but also is a saprobe on other hosts. It has been
used as a control agent of Clematis vitalba in New Zealand
(Gourlay et al. 2000). An updated study classified Phoma
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25
Stagonosporopsis dorenboschii CBS 426.90
Stagonosporopsis dorenboschii CBS 320.90
Stagonosporopsis pini MFLUCC 18-1549 Stagonosporopsis
Stagonosporopsis loticola CBS 562.81
81/1.00
Stagonosporopsis actaeae CBS 106.96
69/---/0.99
Stagonosporopsis ligulicola var ligulicola CBS 137.96
89/1.00
Stagonosporopsis helianthi CBS 200.87
Stagonosporopsis hortensis CBS 104.42
100/1.00 Allophoma nicaraguensis CBS 506.91
70/1.00
99/1.00
Allophoma tropica CBS 436.75
90/1.00
Allophoma oligotrophica CGMCC 3.18114 Allophoma
Allophoma labilis CBS 124.93
100/1.00
96/1.00
Allophoma zantedeschiae CBS 131.93
-Allophoma cylindrispora CBS 142453
--/1.00
Heterophoma nobilis CBS 507.91
99/1.00
Heterophoma
Heterophoma verbascicola CGMCC:3.18364
90/0.98 Ectophoma multirostrata CBS 110.79
100/1.00 Ectophoma multirostrata CBS 368.65
100/1.00
Ectophoma
Ectophoma multirostrata CBS 274.60
--/0.95
Ectophoma pomi CBS 267.92
Anthodidymella vitalbina CBS 454.64
Anthodidymella vitalbina CBS 123707 ET
95/1.00
Anthodidymella vitalbina CBS 123706
Anthodidymella
Anthodidymella vitalbina CBS 911.87
92/1.00
Anthodidymella ranunculacearum MFLUCC 17–2184 HT
93/1.00 100/1.00 Anthodidymella ranunculacearum MFLUCC 17–2209
Anthodidymella clematidis CBS 123705 HT
100/1.00
69/0.99
84/1.00
Didymellaceae
ML/BYPP
Remotididymella destructiva CBS 162.78
Remotididymella destructiva CBS 378.73
Remotididymella
Remotididymella anthropophila CBS 142462
87/1.00
--/0.96
100/1.00
Boeremia lycopersici CBS 378.67
Boeremia
Boeremia exigua CBS 118.38
--/-Epicoccum sorghinum CBS 179.80
95/0.98
Epicoccum
latusicollum
CGMCC:3.18346
74/1.00
Epicoccum viticis CGMCC:3.18344
--/0.99
100/1.00
Epicoccum
Epicoccum ovisporum CBS 180.80
100/1.00
Epicoccum nigrum CBS 125.82
Epicoccum
nigrum
CBS
173.73
76/0.99
Similiphoma crystallifera CBS 193.82
Similiphoma
--/0.99
Nothophoma quercina CBS 633.92
Nothophoma infossa CBS 123395
Nothophoma
95/1.00
Nothophoma anigozanthi CBS 381.91
Didymella coffeae-arabicae CBS 123380
86/1.00
--/0.93
0.998
Didymella eucalyptica CBS 377.91
93/1.00
Didymella pinodes CBS 525.77
-Didymella aurea CBS 269.93
98/1.00
Didymella
heteroderae CBS 109.92
-Didymella maydis CBS 588.69
Didymella pteridis CBS 379.96
-Didymella
Didymella exigua CBS 183.55
97/1.00
Didymella rumicicola CBS 683.79
--/0.99
84/0.99
Didymella microchlamydospora CBS 105.95
71/1.00
Didymella bellidis CBS 714.85
87/1.00
Didymella molleriana CBS 229.79
97/1.00
Paraboeremia litseae CGMCC3.18109
100/1.00
Paraboeremia camelliae CGMCC3.18106
Paraboeremia
Paraboeremia adianticola CBS 187.83
79/0.99
Cumuliphoma indica CBS 991.95
100/1.00
100/1.00
Cumuliphoma indica CBS 654.77
Cumuliphoma
62/0.93
Cumuliphoma omnivirens CBS 341.86
Macroventuria anomochaeta CBS 525.71
100/1.00
Macroventuria
Macroventuria wentii CBS 526.71
--/0.98
Vacuiphoma bulgarica CBS 357.84
100/1.00
Vacuiphoma
Vacuiphoma oculihominis UTHSC:DI16-308
--/0.90
99/1.00
Juxtiphoma eupyrena CBS 374.91
Juxtiphoma
Juxtiphoma eupyrena CBS 527.66
99/1.00
100/1.00
Fig. 12 Phylogram generated from maximum likelihood analysis
based on combined LSU, ITS, and rpb2 sequence data representing
Didymellaceae species. Related sequences were taken from Chen
et al. (2017) and Valenzuela-Lopez et al. (2018). One hundred and
nine strains were included in the combined DNA analyses which
comprised 2094 characters (964 characters for LSU, 531 characters
for ITS, 599 characters for RPB2, including gap regions). Leptosphaeria conoidea (CBS 616.75) and L. doliolum (CBS 505.75) in Leptosphaeriaceae (Pleosporales) were used as out-group taxa. The topology and clade stability of the combined gene analyses was compared
to the single gene analyses. The tree from the maximum likelihood
analysis had similar topology to the Bayesian analyses. The best sorting RaxML tree with a final likelihood value of − 18910.278845 is
presented. The matrix had 602 distinct alignment patterns with 6.40%
undetermined characters or gaps proportions. Estimated base frequencies were as follows: A = 0.247270, C = 0.227052, G = 0.279182,
T = 0.246495; substitution rates AC = 2.142238, AG = 8.227445,
AT = 2.335028, CG = 1.021789, CT = 16.735281, GT = 1.000000;
gamma distribution shape parameter α = 0.490873. The GTR + I + G
model was used for each partition in Bayesian analysis. The species determined in this study are indicated in blue. Bootstrap values (BS) greater than 50% BS (ML, left) and Bayesian posterior
probabilities (BYPP, right) greater than 0.90 are given at the nodes.
Hyphens (-) represent support values less than 50% BS/0.90 BYPP.
Thick branches represent significant support values from all analyses
(BS ≥ 70%/BYPP ≥ 0.95)
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Phomatodes aubrietiae CBS 627.97
Phomatodes nebulosa CBS 740.96
Phoma herbarum CBS 377.92
Phoma herbarum UTHSC:DI16-319
-Briansuttonomyces eucalypti CBS 114879
--/0.93
Neomicrosphaeriopsis italica MFLUCC 15-0485
100/1.00
Neomicrosphaeriopsis italica MFLUCC 15-0484
99/1.00
Ascochyta pisi CBS 126.54
--/0.95
Ascochyta rabiei CBS 206.30
99/1.00
Pseudoascochyta novae-zelandiea CBS 141689
Pseudoascochyta pratensis CBS 141688
Calophoma clematidis-rectae PD 99.2069
99/0.98
0.9505
Calophoma clematidis-rectae CBS 507.63
--/1.00
Calophoma
clematidis-rectae PD 95.1958
--/0.99
92/0.95
Calophoma aquilegiicola CBS 108.96
Calophoma rosae CGMCC 3.18347
100/1.00
Calophoma clematidina CBS 201.49
-100/1.00 Calophoma clematidina CBS 195.64
Calophoma clematidina CBS 108.79
Calophoma clematidina CBS 102.66
100/1.00
Leptosphaerulina australis CBS 317.83
Leptosphaerulina americana CBS 213.55
100/1.00 Xenodidymella applanata CBS 195.36
--/0.99
--/0.99
Xenodidymella applanata CBS 205.63
--/1.00
Xenodidymella clematidis MFLUCC 16–1365
--/1.00
Xenodidymella
catariae CBS 102635
--/1.00
Xenodidymella asphodeli CBS 375.62
Xenodidymella humicola CBS 220.85
Xenodidymella saxea CBS 419.92
95/0.99 Neodidymelliopsis xanthina CBS 383.68
99/1.00
Neodidymelliopsis achlydis CBS 256.77
--/0.95
Neodidymelliopsis polemonii CBS 109181
Neodidymelliopsis cannabis CBS 234.37
92/1.00
100/1.00
Neodidymelliopsis longicolla MFLUCC 17–2167
97/-Neodidymelliopsis longicolla UTHSC:DI16-322
Neodidymelliopsis longicolla CBS 382.96
97/1.00
Neoascochyta triticicola CBS 544.74
67/0.94
Neoascochyta ardicrescens CBS 689.97
90/1.00
Neoascochyta europaea CBS 820.84
99/1.00
Neoascochyta desmazieri CBS 297.69
Neoascochyta desmazieri UTHSC:DI16-207
Leptosphaeria doliolum CBS 505.75
Leptosphaeria conoidea CBS 616.75
100/1.00
--/0.95
--/--
--
99/1.00
Didymellaceae
ML/BYPP
100/1.00
100/1.00
Phomatodes
Phoma
Briansuttonomyces
Neomicrosphaeriopsis
Ascochyta
Pseudoascochyta
Calophoma
Leptosphaerulina
Xenodidymella
Neodidymelliopsis
Neoascochyta
Leptosphaeriaceae
0.03
Fig. 12 (continued)
collections isolated from necrotic leaf tissues of Clematis species into Anthodidymella, Calophoma, and Phoma
(Woudenberg et al. 2009; Valenzuela-Lopez et al. 2018, this
study, Fig. 12).
Anthodidymella has similar morphology to Didymella
in its solitary or clustered, globose ascomata, thin-walled
cells of textura globulosa, with ovate to obpyriform, 1 septum, hyaline ascospores without a mucilaginous sheath. The
asexual morph is pycnidial with phialidic, determinate, and
discrete conidiogenous cells (Chen et al. 2017). However,
Anthodidymella have broad-cylindrical asci, obpyriform
ascospores while Didymella has oblong asci and broadfusiform ascospores (Woudenberg et al. 2009; Aveskamp
et al. 2010). The asexual morph of Anthodidymella has
globose or flask-shaped, phialidic conidiogenous cells with
oblong or elongated-oval conidia. The combined dataset
13
of the LSU, ITS, and rpb2 sequences for Didymellaceae
revealed a lineage including Anthodidymella clematidis, A.
ranunculacearum (type species) and A. vitalbina (Fig. 12).
Anthodidymella clematidis (Woudenb., Spiers & Gruyter)
Phukhams. & K.D. Hyde, comb. nov.
Index Fungorum number: IF557129; Facesoffungi number: FoF 07256
Basionym: Didymella clematidis Woudenb., Spiers &
Gruyter in Woudenberg et al., Persoonia 22:60 (2009)
Synonym: Phoma clematidina (Thüm.) Boerema, Versl.
Medsd. Plziektenk. Dienst Wageningen 153:17 (1979)
Notes: Since the fungus has been introduced before one
fungus = one name (Taylor 2011), the isolate CBS 123705
bears two names for its pleomorphic life-cycles The strain
was originally described as Phoma clematidina from a
Fungal Diversity (2020) 102:1–203
27
Fig. 13 Anthodidymella ranunculacearum (MFLU 17–1468, holotype). a Appearance of conidiomata on Clematis vitalba. b Close
up of conidioma on host substrate. c Vertical section through conidioma. d Ostiolar canal. e Section of partial conidioma wall. f–h Con-
idiogenous cells and conidia (h conidiogenous cells in cotton blue). i
Conidia. Scale bars: b = 200 µm, c = 100 µm, d = 20 µm, e = 10 µm,
f–i = 5 µm
necrotic leaf spot of Clematis ligusticifolia and developed
both sexual morph and asexual morph characters in pure culture (Woudenberg et al. 2009). Anthodidymella clematidis
was initially described as Didymella clematidis for its sexual
morph epithet (= Phoma clematidina as asexual name) as it
clustered with other Phoma clematidina isolates (Woudenberg et al. 2009; Golzar et al. 2011). The new combination,
Anthodidymella clematidis is proposed for Didymella clematidis (CBS 123705).
Host: Clematis ligusticifolia—(Woudenberg et al. 2009).
Distribution: USA—(Woudenberg et al. 2009).
Anthodidymella ranunculacearum Phukhams., Camporesi
& K.D. Hyde, sp. nov.
Index Fungorum number: IF557130; Facesoffungi number: FoF 07257, Fig. 13.
Etymology: The specific epithet reflects the host family,
Ranunculaceae.
Holotype: MFLU 17–1468.
Saprobic on dead stems of Clematis vitalba. Sexual
morph: Undetermined. Asexual morph: Conidiomata
99–214 × 130–246 μm ( x̄ = 142 × 169 μm, n = 5), pycnidial,
solitary, sometimes aggregated, uniloculate, immersed
under epidermal layer, subglobose to depressed, coriaceous,
13
28
thin-walled, brown to dark brown, with ostiolate. Ostioles
25 × 42 μm, central, papillate, with pore. Conidiomatal
wall 10–28(–36) μm wide, of 2–5 layers, each cell-layer
10 μm wide, light brown to brown cells of textura globulosa, heavily pigmented in the outer layers, lined with a
hyaline innermost layer bearing conidiogenous cells. Conidiophores reduced to conidiogenous cells. Conidiogenous
cells 2.5–5 × 1.5–3.5 μm ( x̄ = 3.5 × 2 μm, n = 30), phialidic,
determinate, discrete, ampulliform, cylindrical to sub-cylindrical, smooth-walled, hyaline, arising from the inner layer
of conidioma. Conidia 6–10 × 2–5 μm ( x̄ = 6 × 4 μm, n = 50),
oblong or oval, slightly curved toward the ends, rounded
ends, with 1(–2) guttules in each cell, hyaline, aseptate,
smooth-walled.
Culture characters: Colonies on MEA reaching 20 mm
diam. after 2 weeks at 25 °C. Cultures; above: greyish brown
or dark green, dense, circular, umbonate, papillate, fluffy,
covered with aerial mycelium, reverse dark brown.
Material examined: Italy, Forlì-Cesena Province, Valdinoce—Meldola, on dead aerial branch of Clematis
vitalba, 3 February 2015, E. Camporesi, IT 2364 (MFLU
17–1536, holotype); ex-type living culture, MFLUCC
17–2184 = MFLUCC 17–2209.
Host: Clematis vitalba—(This study).
Distribution: Italy—(This study).
GenBank accession numbers: MFLUCC 17–2184;
LSU: MT214550; SSU: MT226668; ITS: MT310597; tef1:
MT394731; rpb2: MT394681; act: MT394620. MFLUCC
17–2209; LSU: MT214551; SSU: MT226669; ITS:
MT310598; tef1: MT394732; act: MT394621.
Notes: Anthodidymella ranunculacearum (MFLUCC
17–2184) is similar to A. vitalbina (CBS 123707, ex-epitype), a strain recorded from the same host (Woudenberg
et al. 2009). However, A. ranunculacearum differs from
A. vitalbina in its thicker conidiomatal wall (10–36 vs
5.5–9.5 μm, Fig. 13). In a BLASTn search of GenBank,
the ITS sequence had 99.5% similarity (2.25% nucleotide
differences), while the act sequence had 91% similarity (77
nucleotide differences in 297 nucleotides). Thus, the new
strain is introduced as a new species of Anthodidymella
based on guidelines of Jeewon and Hyde (2016). Additionally, A. ranunculacearum is designated as the type species of
Anthodidymella based on available material and an ex-type
culture.
Anthodidymella vitalbina (Petr.) Phukhams. & K.D. Hyde,
comb. nov.
Index Fungorum number: IF557131; Facesoffungi number: FoF 07258.
Basionym: Ascochyta vitalbae Briard & Har. apud Briard,
Rev. Mycol. (Toulouse) 13: 17. (1891).
≡ Diplodina vitalbae (Briard & Har.) Allesch., Rabenh.
Krypt.-Fl., ed. 2. Pilze 6 (Lief. 69): 683. 1900 (1901).
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Synonym: Diplodina clematidina Fautrey & Roum. apud
Roum., Rev. Mycol. (Toulouse) 14: 105 (1892).
= Didymella vitalbina Petr, Annls mycol. 38(2/4): 348
(1940).
= Phoma clematidina (Thüm.) Boerema, Versl. Medsd.
Plziektenk. Dienst Wageningen, 1978 153: 17 (1979).
Notes: Anthodidymella vitalbina was introduced for Didymella vitalbina (= Phoma clematidina as asexual name)
which was reported from a necrotic leaf spot of Clematis species (Woudenberg et al. 2009). Phoma clematidina
(strain CBS 123707) was isolated from Clematis vitalba and
developed sexual and asexual morphs in culture. The sexual
morph is named as Didymella vitalbina Petr. and was chosen
as an epitype of D. vitalbina by Woudenberg et al. (2009). In
the analyses of combined LSU, ITS, and rpb2 sequence data
of Didymellaceae, the ex-epitype strain (CBS 123707) and
the related strains clustered with Anthodidymella clematidis.
Therefore, we transfer Didymella vitalbina (= Phoma clematidina) to Anthodidymella vitalbina based on phylogenetic
relationship of compatible morphology of both morphs. The
nomenclature changes are also based on one fungus = one
name protocol.
Host: Clematis vitalba—(Woudenberg et al. 2009).
Distribution: Austria, France, Switzerland—(Woudenberg et al. 2009).
Neodidymelliopsis Qian & L. Cai
Neodidymelliopsis was introduced for one section of
phoma-like species that reside within Didymellaceae and
is typified by N. cannabis (Chen et al. 2017). The genus
is characterized by immersed or erumpent subglobose to
pyriform, ostiolate ascomata, cylindrical to clavate asci and
subovoid to ellipsoidal, hyaline, septate ascospores. The
asexual morph is phoma-like with pycnidial conidiomata,
a 2–7-layered pseudoparenchymatous pycnidial wall, phialidic conidiogenous cells, and aseptate or occasionally 1-septate conidia (Chen et al. 2015, 2017). We introduce a new
host record of N. longicolla from Clematis vitalba in Italy
(Fig. 14).
Neodidymelliopsis longicolla Hou, Crous & L. Cai, Stud.
Mycol. 87: 153 (2017), new host record
Index Fungorum number: IF820006; Facesoffungi number: FoF 07259, Fig. 14.
Saprobic on dead stems of Clematis vitalba. Sexual
morph: Undetermined. Asexual morph: Conidiomata
70–95 × 124–134 μm ( x̄ = 84 × 130 μm, n = 5), pycnidial,
aggregated, uniloculate, superficial or covered by host epidermal layer, subglobose to depressed, cupulate when dried
coriaceous, thick-walled, light brown to brown, with papillate ostioles. Ostioles 80 × 38 μm, central, papillate, opening by a pore. Conidiomatal wall 10–17(–27) μm wide,
composed of 5–7 layers of light brown to brown cells of
Fungal Diversity (2020) 102:1–203
29
textura angularis, heavily pigmented at the outer layers,
lined with a hyaline layer bearing conidiogenous cells. Conidiophores reduced to conidiogenous cells. Conidiogenous
cells 2.5–7 × 2–4.5 μm ( x̄ = 4.5 × 3.5 μm, n = 30), phialidic,
annellidic, determinate, discrete, ampulliform, cylindrical
to sub-cylindrical, smooth-walled, hyaline, arising from the
inner layers of conidiomata. Conidia 6.5–10 × 2–4.5 μm
( x̄ = 8 × 4 μm, n = 50), oblong-elliptical, oval, slightly curved
towards the ends, rounded ends, with 1(–2) guttules in each
cell, initially aseptate and hyaline, becoming pale brown
and 1-septate at maturity, constricted at the septum, wall
verrucose.
Culture characters: Colonies on MEA reaching 20 mm
diam. after 2 weeks at 25 °C. Cultures from above, cream
with white at the centre, medium dense, circular, umbonate,
papillate, fluffy, covered with white aerial mycelium; reverse
brown white cream at the edge.
Material examined: Italy, Forlì-Cesena Province, Castrocaro Terme, on dead aerial branch of Clematis vitalba, 19
Fig. 14 Neodidymelliopsis longicolla (MFLU 20–0421). a Appearance of conidiomata on Clematis vitalba. b Vertical section through
conidioma. c Section of conidioma wall. d–f Conidiogenous cells and
conidia. g–j Conidia. k Culture characteristics on MEA. Scale bars:
b = 100 µm, c = 20 µm, d–f = 10 µm, g–j = 10 µm
13
30
September 2012, E. Camporesi, IT739 (MFLU 20–0421);
living culture, MFLUCC 17–2167.
Hosts: Soil in desert, Clematis vitalba—(Chen et al.
2017; this study).
Distribution: Israel, Italy—(Chen et al. 2017; this study).
GenBank accession numbers: LSU: MT214552; SSU:
MT226670; ITS: MT310599; tef1: MT394733; rpb2:
MT394682.
Notes: A new isolate of Neodidymelliopsis longicolla
(MFLUCC 17–2167) was collected from Clematis vitalba
in Italy. The new isolate formed a close relationship with the
type (100% ML/1.00 BYPP). Neodidymelliopsis longicolla
(CBS 382.96) was originally reported from a soil sample in
Israel. The characters of the type strain were obtained from
a culture on OA medium. The new collection differs slightly
from the type material in having a shorter ostioles (Fig. 14).
Xenodidymella Chen & L. Cai
Xenodidymella typified by X. applanata (≡ Didymosphaeria applanata) has Phoma argillacea as the asexual morph
(Corlett 1981; Gruyter et al. 2013; Chen et al. 2015). Xenodidymella is distinct from other genera in Didymellaceae in
having a thick peridium and ellipsoidal, allantoid or subcylindrical, unicellular conidia. Xenodidymella is reported
from Europe and USA and consists of five species (Farr
and Rossman 2020; Index Fungorum 2020). We introduce
a novel species of Xenodidymella clematidis from Clematis
vitalba in Italy and provide phylogenetic and morphological
comparisons (Figs. 12, 15).
Xenodidymella clematidis Phukhams., Camporesi & K.D.
Hyde, sp. nov.
Index Fungorum number: IF557132; Facesoffungi number: FoF 07260, Fig. 15.
Etymology: Refers to the host genus Clematis.
Holotype: MFLU 16–2288.
Saprobic on dead stems of Clematis vitalba. Sexual
morph: Undetermined. Asexual morph: Conidiomata
220–375 × 180–290 μm ( x̄ = 280 × 230 μm, n = 5), pycnidial,
aggregated, uniloculate, superficial or semi-immersed, subglobose to depressed globose, cupulate, when dried coriaceous, thick-walled, brown to light brown, with ostioles.
Ostioles central, papillate, opened-like pore. Conidiomatal
wall 10–15(–25 μm at apex) wide, composed of 7–9 layers
of light brown to brown cells of textura angularis, heavily
pigmented at the outer layers, lined with a hyaline layer bearing conidiogenous cells. Conidiophores reduced to conidiogenous cells. Conidiogenous cells 2–6(–12) × 2.3–3.3 μm
( x̄ = 5 × 3 μm, n = 20), enteroblastic, phialidic, determinate,
discrete, ampulliform, smooth-walled, hyaline, arising from
the inner layers of conidiomata. Conidia 4–8 × 2–5 μm
( x̄ = 6 × 3.5 μm, n = 50), oblong-elliptical, oval, slightly
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Fungal Diversity (2020) 102:1–203
curved towards the ends, rounded ends, hyaline, with 1(–2)
guttules in each cell, aseptate, verrucose.
Culture characters: Colonies on MEA reaching 20 mm
diam. after 2 weeks at 16 °C. Cultures from above, grey with
white aerial mycelium, dense, circular, umbonate, papillate,
fluffy, reverse brownish white, cream at the edge.
Material examined: Italy, Forlì-Cesena Province, Monte
Fumaiolo, dead aerial branch of Clematis vitalba, 6 August
2016, E. Camporesi, IT3054 (MFLU 16–2288, holotype);
ex-type living culture, MFLUCC 16–1365.
Host: Clematis vitalba—(This study).
Distribution: Italy—(This study).
GenBank accession numbers: LSU: MT214553; ITS:
MT310600; act: MT394622.
Notes: In the phylogenetic analysis, Xenodidymella clematidis clustered with X. applanata with strong support (1.00
in BYPP). Xenodidymella clematidis has a long ostiole,
with oblong-elliptical or oval conidia, while X. applanata
has a short ostiole and ellipsoidal conidia (Gruyter et al.
2002; Chen et al. 2015). Xenodidymella applanata is commonly a pathogen of raspberry (Rubus sp.). Xenodidymella
clematidis was a saprobe on Clematis vitalba (Fig. 15). In
a BLASTn search of GenBank, the ITS sequence had 98%
similarity (11 nucleotides differences out of 488 nucleotides). The new strain is introduced as a new species of
Xenodidymella herein.
Didymosphaeriaceae Munk
The latest treatment of Didymosphaeriaceae was by Ariyawansa et al. (2014a). The family is typified with Didymosphaeria. Several genera have been introduced to the family
based on morphological and phylogenetic evidence (Tibpromma et al. 2018; Wanasinghe et al. 2018; Wijayawardene
et al. 2018). Mapook et al. (2020) introduced Chromolaenicola Mapook & K.D. Hyde into Didymosphaeriaceae.
Phylogeny and morphological comparisons revealed a
novel species of Chromolaenicola and a new host record
of Didymosphaeria rubi-ulmifolii from Clematis (Fig. 16).
Chromolaenicola Mapook & K.D. Hyde
Mapook et al. (2020) introduced Chromolaenicola (typified with C. nanensis) for a monophyletic clade of fungi
described on Chromolaena odorata. Chromolaenicola is
characterized by immersed to semi-immersed and coriaceous ascomata, cylindrical asci, and uniseriate, ellipsoid,
muriform ascospores (Mapook et al. 2020). The asexual
morph is pycnidial, with enteroblastic, phialidic conidiogenous cells, and oblong or oval to ellipsoid, globose to subglobose conidia (Jayasiri et al. 2019; Mapook et al. 2020).
We introduce a new species Chromolaenicola clematidis
based on morphological comparison (Fig. 17) and phylogenetic analyses (Fig. 16).
Fungal Diversity (2020) 102:1–203
31
Fig. 15 Xenodidymella clematidis (MFLU 16–2288, holotype). a
Appearance of conidiomata on Clematis vitalba. b Vertical section
through conidioma. c Ostiolar canal. d Section of conidioma wall. e,
f Conidiogenous cells and conidia. g Conidia. h Culture characters on
MEA. Scale bars: b = 200 µm, c, d = 50 µm, e–g = 10 µm
Chromolaenicola clematidis Phukhams. & K.D. Hyde, sp.
nov.
Index Fungorum number: IF557133; Facesoffungi number: FoF 07253, Fig. 17.
Etymology: Named after the host genus, Clematis.
Holotype: MFLU 17–1483.
Saprobic on dead stem of Clematis subumbellata. Sexual morph: Undetermined. Asexual morph: Conidiomata
76–145 × 107–128 μm ( x̄ = 121 × 117 μm, n = 10), pycnidial,
solitary, uniloculate, immersed, globose, coriaceous, thin,
brown to light brown, ostiolate. Conidiomatal wall 5–10 μm
wide, uniform, wider at apex, composed of 3–5 layers of
pale brown to bronze cells of textura angularis, lined with
a thin, hyaline layer bearing conidiogenous cells. Conidiophores reduced to conidiogenous cells. Conidiogenous cells
2.6–4.5 × 4–7 μm ( x̄ = 3.5 × 5 μm, n = 20), enteroblastic,
phialidic, determinate, discrete, truncate, hyaline, smooth,
arising from the inner layer of pycnidial wall. Conidia
7–10 × 4.5–7 μm ( x̄ = 8.5 × 6 μm, n = 50), broad oblong or
oval, rounded ends, hyaline when immature, reddish brown
at maturity, 1-septate, with guttule in each cell, roughwalled, verrucose.
Cultural characters: Colonies on MEA reaching 30 mm
diam. after 4 weeks at 25 °C. Cultures from above, dark
brown, radiating outwardly, dense, umbonate, undulate edge,
umbonate, papillate, fluffy, covered with white aerial mycelium; reverse dark brown, undulate.
Material examined: Thailand, Chiang Rai Province, Mae
Sai District, dead stems of Clematis subumbellata, 20 March
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Paraphaeosphaeria michotii MFLUCC 13-0349
Paraphaeosphaeria
Paraphaeosphaeria angularis CBS 167.70
Paraphaeosphaeria minitans CBS 111750
93/1.00
ML/BYPP
100/1.00 Karstenula rhodostoma CBS 690.94
Karstenula
99/1.00
Karstenula rhodostoma CBS 691.94
Paraphaeosphaeria rosicola MFLUCC 15-0042
Paraphaeosphaeria
-75/0.99
Paraphaeosphaeria rosae MFLUCC 17-2547
79/0.95 Austropleospora archidendri CBS 168.77
-Austropleospora
Austropleospora keteleeriae MFLUCC 18-1551
Austropleospora osteospermi LM 2009a
97/0.99 Chromolaenicola nanensis MFLUCC 17-1477
93/1.00 Chromolaenicola nanensis MFLUCC 17-1473
-63/0.99
Chromolaenicola lampangensis MFLUCC 17-1462
Chromolaenicola
90/1.00 Chromolaenicola thailandensis MFLUCC 17-1510
100/0.95
Chromolaenicola thailandensis MFLUCC 17-1475
Chromolaenicola
clematidis
MFLUCC
17-2075
66/-53/0.95
Chromolaenicola chiangraiensis MFLUCC 17-1493
0
100/1.00
Chromolaenicola siamensis MFLUCC 17-2527
69/1.00
Cylindroaseptospora leucaenae MFLUCC 17–2424 Cylindroaseptospora
Verrucoconiothyrium nitidae CBS 119209
Verrucoconiothyrium
r sp. CBS 122320
73/-- Didymosphaeria
Didymosphaeria sp. CBS 122319
Didymosphaeria rubi-ulmifolii MFLUCC 16-1000
Didymosphaeria rubi-ulmifolii CBS 100299
Didymosphaeria
--/0.91
Didymosphaeria rubi-ulmifolii MFLUCC 14-0023
81/-Didymosphaeria rubi-ulmifolii CBS 587.84
76/-Didymosphaeria sp. CBS 115.92
-Didymosphaeria variabile CBS 120014
Pseudocamarosporium pteleae MFLUCC 17-0724
85/1.00
97/1.00
Pseudocamarosporium corni MFLUCC 13-0541
Pseudocamarosporium
Pseudocamarosporium propinquum MFLUCC 13-0544
53/0.99
Paracamarosporium hawaiiense CBS 120025
Paracamarosporium
Paracamarosporium fagi CPC 24890
87/1.00 Paracamarosporium fagi CPC 24892
100/1.00
Paraconiothyrium ungicola CBS 113269
Paraconiothyrium
67/-Paraconiothyrium magnoliae MFLUCC 10-0278
100/1.00 Deniquelata barringtoniae MFLUCC 11-0257
Deniquelata
Deniquelata barringtoniae MFLUCC 11-0422
63/1.00
100/1.00
Tremateia arundicola MFLU 16-1275
Tremateia
Tremateia guiyangensis GZAAS01
-Neokalmusia brevispora KT 2313
Neokalmusia
83/-Neokalmusia scabrispora KT 2202
99/1.00
Pseudopithomyces chartarum UTHSC 04-678
Pseudopithomyces
Pseudopithomyces sp. MUCL 15905
100/1.00
Montagnula aloes CPC 19671
Montagnula
Montagnula opulenta CBS 168.34
(Out group)
Didymosphaeriaceae
97/1.00
98/1.00
0.02
Fig. 16 The best scoring RAxML tree with a final likelihood value
of − 12878.030199 based on combined LSU, ITS, SSU, rpb2 and
tub sequence data of Didymosphaeriaceae. The tree is rooted with
sequences of Montagnula species. Forty-six strains were included in
the combined DNA sequence analyses which comprised 3681 characters (860 characters for LSU, 516 characters for ITS, 913 characters
for SSU, 935 characters for rpb2, 457 characters for tub, including
gap regions). The topology and clade stability of the combined gene
analyses was compared to the single gene analyses. The tree from the
maximum likelihood analysis had similar topology to the Bayesian
analyses. The matrix had 764 distinct alignment patterns, with 9.02%
undetermined characters or gaps propotions. Estimated base frequen-
cies were as follows: A = 0.239562, C = 0.247759, G = 0.271470,
T = 0.241209; substitution rates AC = 1.719408, AG = 2.576378,
AT = 1.444379, CG = 0.958169, CT = 9.083623, GT = 1.000000;
gamma distribution shape parameter α = 0.621443. In our analysis,
GTR + I + G model was used for each partition in Bayesian posterior
analysis. The species determined in this study are indicated in blue.
Bootstrap values (BS) greater than 50% BS (ML, left) and Bayesian
posterior probabilities (BYPP, right) greater than 0.90 are given at the
nodes. Hyphens (-) represent support values less than 50% BS/0.90
BYPP. Thick branches represent significant support values from all
analyses at the genus level (BS ≥ 70%/BYPP ≥ 0.95)
2017, C. Phukhamsakda, CMTH19 (MFLU 17–1483, holotype); ex-type living culture, MFLUCC 17–2075.
Host: Clematis subumbellata—(This study).
Distribution: Thailand—(This study).
GenBank accession numbers: LSU: MT214554; SSU:
MT226671; ITS: MT310601; rpb2: MT394683.
Notes: In our phylogenetic analysis (Fig. 16), Chromolaenicola clematidis (MFLUCC 17–2075) clustered with C.
chiangraiensis Mapook & K.D. Hyde with moderate statistical support (66% ML/0.82 BYPP). A comparison of the ITS
sequences showed three nucleotide differences in 516 nucleotides, while the rpb2 showed five nucleotide differences in
935 nucleotides. Chromolaenicola clematidis has smaller
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Fungal Diversity (2020) 102:1–203
33
Fig. 17 Chromolaenicola clematidis (MFLU 17–1483, holotype). a,
b Appearance of conidiomata on Clematis subumbellata. c Vertical
section through conidioma. d Section of conidioma wall. e–f Con-
idiogenous cells and conidia. g–j Conidia. k–l Culture characters on
MEA. Scale bars: b = 200 µm, c, d = 50 µm, e–j = 5 µm
conidia than C. chiangraiensis (mean 8.5 × 6 vs 11 × 7.5 µm)
with guttules in each cell (Fig. 17, Table 2). The new strain
is introduced as a new species of Chromolaenicola based on
the guidelines proposed by Jeewon and Hyde (2016).
species but a holomorphic connection has not been proven
(Sivanesan 1984; Kirk et al. 2008; Ariyawansa et al. 2014b).
Didymosphaeria is typified by D. futilis, however, fresh collections are needed to confirm its phylogenetic placement
(Ariyawansa et al. 2014a; Wijayawardene et al. 2018). More
than 500 epithets are listed under Didymosphaeria (Index
Fungorum 2020), but only seven species were accepted by
Aptroot (1995). Only three species of Didymosphaeria have
phylogenetic evidence (D. rubi-ulmifolii, D. variabile, and
Didymosphaeria sp. (as Paraconiothyrium brasiliense CBS
115.92, CBS 587.84, CBS 122319 and CBS 122320)) (Verkley et al. 2004). In this study, phylogenetic analyses based
Didymosphaeria Fuckel
Didymosphaeria is characterized by trabeculate pseudoparaphyses (sensu Liew et al. 2000), which anastomose above the cylindrical asci, and uniseriate, 1-septate
ascospores (Aptroot 1995; Ariyawansa et al. 2014b). The
asexual morph of Didymosphaeria has been suggested to
be Ascochyta, fusicladiella-like, Periconia, and phoma-like
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Table 2 A comparison of Chromolaenicola species discussed in this study
Species
Conidiomata (μm)
Conidiogenous cells (μm) Conidia (μm) Host
Chromolaenicola clematidis
76–145 × 107–128 2.6 − 4.5 × 4 − 7
Ch. chiangraiensis (MFLUCC 17–1493) –
3.5–6.5 × 1–2
Ch. siamensis
110–165 × 140–190 6.5–7.4 × 3.2–4.7
(MFLUCC 17–2527)
on a combined dataset of the LSU, ITS, SSU, rpb2, and tub
sequences revealed a new host record for D. rubi-ulmifolii
on Clematis heracleifolia from China (Fig. 18).
Didymosphaeria rubi-ulmifolii Ariyaw., Camporesi & K.D.
Hyde, Phytotaxa 176:111 (2014), new host record
Index Fungorum number: IF808165; Facesoffungi number: FoF 07254, Fig. 18.
Saprobic on dead stem of Clematis heracleifolia. Sexual morph: Ariyawansa et al. (2014a). Asexual morph:
Conidiomata 78–160 × 75–244 μm ( x̄ = 110 × 120 μm,
n = 5), pycnidial, solitary, unilocular or multilocular, scattered, immersed or erumpent, under host epidermis, globose to compressed, brown to dark brown, without ostioles.
Pycnidial wall 12–20(−30 μm at apex) wide, composed
of 4–5 brown cell layers of textura angularis, inner layer
subhyaline, lining bearing conidiogenous cells. Conidiophores reduced to conidiogenous cells. Conidiogenous
cells 1.8–5 × 2–4 μm ( x̄ = 3 × 3 μm, n = 20), enteroblastic,
phialidic, determinate, smooth-walled, hyaline. Conidia
6–11 × 2.5–4.5 μm ( x̄ = 9 × 4 μm, n = 50), ellipsoid, 1 septum, constricted at septum, rounded ends, initially hyaline,
pale brown at maturity, with 1–2 guttules, smooth-walled.
Cultural characters: Colonies on MEA reaching 30 mm
diam. after 4 weeks at 16 °C. Cultures from above, olive
brown at the centre, radiating outwardly, medium dense,
circular, entire edge, umbonate, papillate, fairly fluffy, covered with white aerial mycelium; reverse dark brown at the
centre, faintly zonate slightly present, white mycelium at
the edge.
Material examined: China, Dali, on dead terrestrial stem
of Clematis heracleifolia DC., 8 May 2016, C. Phukhamsakda, CMCN03 (MFLU 17–1460); living culture,
MFLUCC 16–1000.
Hosts: Coffea arabica, Rubus ulmifolius, Clematis heracleifolia—(Verkley et al. 2004; Ariyawansa et al. 2014a;
this study).
Distribution: Brazil, China, Italy—(Verkley et al. 2004;
Ariyawansa et al. 2014a; this study).
GenBank accession numbers: LSU: MT214555; SSU:
MT226672; ITS: MT310602; tef1: MT394734.
Notes: Ariyawansa et al. (2014a) introduced Didymosphaeria rubi-ulmifolii from a collection of Rubus ulmifolius in Italy. The ex-type strain (MFLUCC 14–0023) formed
13
7–10 × 4–7
9–14 × 6–9
7–9 × 5–6
Clematis subumbellata
Chromolaena odorata (Asteraceae)
Leucaena sp. (Fabaceae)
a distinct clade with the type strain of Paraconiothyrium
brasiliense (CBS 100299). Based on the multilocus phylogenetic analyses, our strain (MFLUCC 16–1000) formed
a close relationship with the other D. rubi-ulmifolii strains
with moderate support (Fig. 16), with no significant pairwise
differences. Morphological comparison of D. rubi-ulmifolii
(MFLUCC 16–1000) with the asexual morph report in D.
rubi-ulmifolii (strain CBS 100299) show that they are different in the conidial characters (Fig. 18). Strain MFLUCC
16–1000 is saprobic and has 2-celled conidia while, CBS
100299 has single celled conidia in culture (Verkley et al.
2004). This study is the first record of D. rubi-ulmifolii on
Clematis species.
Hermatomycetaceae Locq. ex A. Hashim. & Kaz. Tanaka
Hermatomycetaceae is typified by Hermatomyces and
currently only known from asexual morph characters
(Tibpromma et al. 2016, 2018; Koukol et al. 2018; Hyde
et al. 2019a). We introduce a novel Hermatomyces species
based on its distinct morphology with phylogenetic support
(Figs. 19, 20).
Hermatomyces Speg.
Hermatomyces tucumanensis is the type species. The
genus has sporodochial conidiomata, and muriform, lenticular, hyaline or dematiaceous conidia of one or two types
(Spegazzini 1911; Chang 1995; Tibpromma et al. 2016;
Hashimoto et al. 2017; Hyde et al. 2019a). Examination of a
Clematis collection revealed a novel species Hermatomyces
clematidis based on its distinct morphological and phylogenetic relationship from other Hermatomyces. This is the first
record of a Hermatomyces species on Clematis (Fig. 21).
Hermatomyces clematidis Phukhams., D.J. Bhat & K.D.
Hyde, sp. nov.
Index Fungorum number: IF557134; Facesoffungi number: FoF 07244, Fig. 21.
Etymology: Name refers to the host plant, Clematis.
Holotype: MFLU 17–1493.
Saprobic on dead stem of Clematis sikkimensis. Sexual
morph: Undetermined. Asexual morph: Hyphomycetous. Colonies on natural substrate forming sporodochial
conidiomata, subiculate, superficial, scattered, circular or
oval, blackish brown, velvety, glistening, orbicular, with
Fungal Diversity (2020) 102:1–203
35
Fig. 18 Didymosphaeria rubi-ulmifolii (MFLU 17–1535). a, b
Appearance of conidiomata on Clematis heracleifolia. c Vertical
section through conidioma. d Section of conidioma wall. e–h Con-
idiogenous cells and conidia. i–l Conidia. Scale bars: a = 500 µm,
b = 200 µm, c = 100 µm, d = 20 µm, e–k = 5 µm, l = 10 µm
abundant sporulation, conidia readily liberated when agitated, 410–565 μm wide ( x̄ = 490 μm, n = 20). Mycelium
2–4 μm wide, mostly superficial, composed of a loose
or compact network of repent, branched, septate, roughwalled, thick-walled, reddish brown to brown hyphae;
subicular hyphae short, densely packed. Conidiophores
22–38 × 2–5 μm, micronematous or semi-macronematous,
mononematous, cylindrical, erect, verruculose, aseptate,
branched, often corresponding to conidiogenous cells, reddish brown to brown. Conidiogenous cells 7–13 × 4–7 μm,
holoblastic, monoblastic, solitary, integrated, terminal,
determinate, cylindrical or slightly subulate, subsphaerical
or ampulliform, reddish brown to brown, sometimes hyaline. Conidia dimorphic solitary, smooth-walled; lenticular
conidia: 30–45 × 24–31 μm ( x̄ = 40 × 28 μm, n = 30), muriform, smooth, disc-shaped, circular to oval in front view,
central cells brown to reddish brown, peripheral cells hyaline to subhyaline, forming a wide and distinct ring, slightly
constricted at the septa, inside view composed of one column of 5–6 cells, end cells subhyaline to pale brown, often
carrying remnant of conidiogenous cell at base; cylindrical
conidia: 29–35 × 12–14 μm ( x̄ = 32 × 13 μm, n = 20), straight
or flexuous, septate, constricted at the septa, consisting of
one or two columns, usually separate at apex, each column
with 4–5 transverse septa, obclavate, apical cell acute, basal
cells rounded, smooth, hyaline.
Culture characters: Colonies on MEA reaching 50 mm
diam. after 4 weeks at 25 °C. Cultures from above, white,
dense, circular, margin erose, umbonate, papillate with fairly
fluffy, wrinkled, folded, pale orange, covered with cream
aerial mycelium; reverse dark brown at the centre, faintly
zonate at the edge, greyish orange radiating outwardly.
Material examined: Thailand, Chiang Rai Province,
on dead stem of Clematis sikkimensis, 2 May 2017, C.
Phukhamsakda & M.V. de Bult, CMTHDT01 (MFLU
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Hermatomyces sphaericus MFLUCC 14-1140 (=H. tectonae HT)
Hermatomyces sphaericus MFLUCC 14-1141 (=H. tectonae)
Hermatomyces bioconisporus KUMCC 17-0183 HT
Hermatomyces sphaericus MFLUCC 16-0266 (=H. saikhuensis HT)
Hermatomyces sphaericus MFLUCC 16-0267 (=H. saikhuensis)
Hermatomyces sphaericus PRC 4104
Hermatomyces sphaericus MFLUCC 14-1142 (=H. tectonae)
Hermatomyces sphaericus PMA 116080
Hermatomyces sphaericus PMA 116081
Hermatomyces sphaericus MFLUCC 16-2819 (=H. chromolaenae HT)
Hermatomyces sphaericus PRC 4100
Hermatomyces sphaericus PRC 4106
Hermatomyces sphaericus PMA 116085
Hermatomyces sphaericus PRC 4116
Hermatomyces sphaericus PRM 946201
Hermatomyces sphaericus PMA 116082
-Hermatomyces sphaericus PRC 4105
Hermatomyces sphaericus KZP462
Hermatomyces sphaericus PRC 4117
96/99
1.00
Hermatomyces pandanicola MFLUCC 16-0251
96/97 Hermatomyces indicus MFLUCC 14-1144
50/68
0.99
0.95
Hermatomyces indicus MFLUCC 14-1143
Hermatomyces indicus MFLUCC 14-1145
82/90
94/97
1.00
Hermatomyces bauhiniae MFLUCC 16-0395
0.99
Hermatomyces nabanheensis KUMCC 16-0149
99/100
1.00
Hermatomyces bifurcatus CCF 5899
84/92
Hermatomyces bifurcatus CCF 5900
0.99
99/100
Hermatomyces megasporus CCF 5898
100/100 1.00
Hermatomyces megasporus CCF 5897
1.00
Hermatomyces reticulatus MFLUCC 15-0843
98/100 Hermatomyces reticulatus CCF 5893
--/53
0.99
-97/100
100/100 Hermatomyces tucumanensis CCF 5912
1.00
1.00
Hermatomyces tucumanensis CCF 5915
100/100
Hermatomyces
tucumanensis CCF 5913
1.00
92/-100/100
-1.00 Hermatomyces trangensis BCC 80741
100/99
1.00
Hermatomyces trangensis BCC 80742
Hermatomyces clematidis MFLUCC 17-2085
92/100 Hermatomyces krabiensis MFLUCC 16-0249
100/100
1.00
1.00
Hermatomyces krabiensis MFLUCC 16-2817
Hermatomyces iriomotensis KT 2016
100/100
1.00
Hermatomyces sphaericoides CCF 5908
57/74
Hermatomyces sphaericoides CCF 5895
0.92
Hermatomyces constrictus CCF 5904
97/100
Hermatomyces verrucosus CCF 5903
1.00
100/100 Hermatomyces verrucosus CCF 5892
1.00
Anteaglonium globosum ANM 925.2
Anteagloniaceae
100/100
Anteaglonium parvulum MFLUCC 14-0821
Hermatomycetaceae
MP/ML
PP
(Outgroup)
1.00
30.0
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37
◂Fig. 19 Phylogram generated from maximum parsimony analysis
based on combined LSU, ITS, tef1 and rpb2 sequence data. Related
sequences are taken from Nuankaew et al. (2019) and retrieved from
GenBank. Forty-nine strains were included in the analysis of the
combined DNA loci and comprise 3309 characters (826 characters for
LSU, 514 characters for ITS, 948 characters for tef1, 1021 characters
for rpb2, including gaps). The tree is rooted with Anteaglonium globosum (ANM 925.2) and A. parvulum (MFLUCC 14-0821) in Anteagloniaceae. Maximum parsimony analysis of 471 parsimony informative characters resulted in a most parsimonious tree (CI = 0.674,
RI = 0.879, RC = 0.593, HI = 0.326). The best scoring RAxML tree
had a final likelihood value of −10433.015131. The matrix had 802
distinct alignment patterns with 28.12% undetermined characters and
gaps. Estimated base frequencies were: A = 0.244372, C = 0.264291,
G = 0.261114, T = 0.230223; substitution rates AC = 0.858179,
AG = 3.949480, AT = 1.122065, CG = 0.761732, CT = 11.028490,
GT = 1.000000; gamma distribution shape parameter α = 0.151089.
In our analysis, GTR + I + G model was used for each partition in
Bayesian posterior analysis. Bootstrap values (BS) greater than 50%
BS (ML, left) and Bayesian posterior probabilities (BYPP, right)
greater than 0.90 are given at the nodes. Hyphens (-) represent support values less than 50% BS/0.90 BYPP. Thick branches represent
significant support values from all analyses (BS ≥ 70%/BYPP ≥ 0.95).
The ex-type strains are in bold and black. The newly generated
sequence is in bold and blue
17–1493, holotype); ex-type living culture, MFLUCC
17–2085.
Host: Clematis sikkimensis—(This study).
Distribution: Thailand—(This study).
Fig. 20 The splits graph from
the pairwise homoplasy index
(PHI) test generated from the
concatenated gene set of LSU,
ITS, tef1 and rpb2 sequence
data of closely related species
using both LogDet transformation and splits decomposition.
PHI test results (Φw) < 0.05
indicates significant recombination within the dataset
GenBank accession numbers: LSU: MT214556; SSU:
MT226673; ITS: MT310603; tef1: MT394735; rpb2:
MT394684.
Notes: Hermatomyces clematidis is introduced as a
new species based on its distinct morphology and phylogenetic results of a combined LSU, ITS, tef1, and rpb2
dataset (Fig. 19). Hermatomyces clematidis matches the
generic concept in having sporodochial conidiomata, with
both lenticular and cylindrical conidia (Doilom et al. 2017;
Hashimoto et al. 2017, Fig. 21). Morphological comparison
with known Hermatomyces species shows it is similar to H.
tucumanensis, however, the conidiophore of H. clematidis
are straighter with larger conidia ((22–)27–35 × 18–25 vs
30–45 × 24–31 µm) (Koukol et al. 2018). In the phylogenetic
analysis, H. clematidis formed a close relationship with H.
trangensis with strong support (100% MP/99% ML/1.00
BYPP). Hermatomyces trangensis is associated with sugar
palm in southern Thailand (Nuankaew et al. 2019). Hermatomyces trangensis differs from H. clematidis because it lacks
cylindrical conidia. In a BLASTn search of GenBank, the
closest match of the LSU sequence of MFLUCC 17–2085
is H. subiculosa (MFLUCC 15–0843) with 97% similarity,
while the closest match of the ITS sequence was H. tectonae
(MFLUCC 14–1140) (NR_154079) with 97% similarity.
Based on current evidence, phylogenetic results from the
concatenated gene loci does not delineate Hermatomyces
H. sphaericus PRC 4104
H. sphaericus PMA 116082
=H. saikhuensis MFLUCC 16-0267
H. sphaericus PRC 4117
H. sphaericus PRC 4105
H. sphaericus PRC 4116
H. sphaericus PRC 4106
H. sphaericus KZP462
=H. tectonae MFLUCC 14-1141
H. sphaericus PMA 16080
=H. tectonae MFLUCC 14-1140 HT
H. sphaericus PRC 4100
=H. saikhuensis MFLUCC 16-0266 HT
H. sphaericus PMA 116081
=H. tectonae MFLUCC 14-1142
H. sphaericus PRM 946201
H. sphaericus PMA 116085
H. sphaericus sensu stricto
1.0
Φw = 0.009
Hermatomyces pandanicola
MFLUCC 16-0251
Hermatomyces biconisporus
KUMCC 17-0183
13
38
species, especially the H. sphaericus clade (Fig. 19). Therefore, we applied the GCPSR concept with the H. sphaericus clade. Seventeen isolates of H. sphaericus formed an
isolated clade with two species (H. biconisporus and H.
pandanicola) forming distant relationship lineage with low
statistic support. However, the pairwise homoplasy index
showed Φw = 0.009 when the degree of genealogical correlation model was applied between neighbouring strains of
the clade (Fig. 20). These results are not congruent with the
phylogenetic lineages shown in Fig. 19, which rather indicate that H. biconisporus, H. pandanicola and H. sphaericus
should currently be treated as the same species.
Leptosphaeriaceae Barr
Leptosphaeriaceae is typified by Leptosphaeria. Ariyawansa et al. (2015b) illustrated the members of Leptosphaeriaceae and included ten genera. Quaedvlieg et al. (2013)
introduced an asexual morph genus, Acicuseptoria Quaedvl., Verkley & Crous to the family, however, a multilocus
phylogenetic analysis (Fig. 22) reveals that Acicuseptoria
clusters with Paraleptosphaeria. Crous and Groenewald
(2017a) introduced Querciphoma isolated from stems and
leaves of woody plants in a terrestrial environment. Members of this family usually have a single, papillate, immersed
or erumpent, perithecial ascomata, with thick, scleroplectenchymatous or plectenchymatous cells and cylindrical asci
with hyaline to brown, transversely septate ascospores (Hyde
et al. 2013; Ariyawansa et al. 2015b). The asexual morphs in
Leptosphaeriaceae can be coelomycetous or hyphomycetous
(Gruyter et al. 2013; Hyde et al. 2013; Crous and Groenewald 2017). We introduce a novel Alloleptosphaeria species and describe a novel genus Sclerenchymomyces based
on distinct morphology and phylogenetic support (Fig. 22).
Alloleptosphaeria Ariyaw., Wanas. & K.D. Hyde
Alloleptosphaeria was introduced as a monotypic genus
to accommodate A. italica from Clematis vitalba in Italy
(Ariyawansa et al. 2015b). It has immersed ascomata and a
thin-walled peridium of reddish brown to dark brown pseudoparenchymatous cells (Ariyawansa et al. 2015b). In our
phylogenetic analysis (Fig. 22), Alloleptosphaeria formed a
distinct clade basal to Leptosphaeria sensu stricto. We introduce the second species of Alloleptosphaeria from Clematis
subumbellata collected in Thailand (Fig. 23).
Alloleptosphaeria clematidis Phukhams. & K.D. Hyde, sp.
nov.
Index Fungorum number: IF557109; Facesoffungi number: FoF 07286, Fig. 23.
Etymology: Epithet reflect the host genus Clematis.
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Fungal Diversity (2020) 102:1–203
Holotype: MFLU 17–1479.
Saprobic on dead stems of Clematis subumbellata.
Sexual morph: Ascomata 210–260 × 125–190 μm ( x̄
= 237 × 160 μm, n = 5), on surface of the host, covered by
a pseudoclypeus, visible as black spots, immersed, solitary, scattered, uniloculate, obpyriform, coriaceous, black
to dark brown, rough-walled, with apical ostioles. Ostioles
70–85 × 50–70 μm ( x̄ = 80 × 60 μm, n = 5), central, pale
brown to dark brown, papillate, opening by a pore, ostioles
with periphyses. Peridium 9–17(–25 μm at apex) wide, thin,
multilayered, pseudoparenchymatous cell type, comprising
4–5 layers of brown to dark brown cells of textura angularis,
inner layers comprising thin, hyaline cells. Hamathecium
composed of numerous, 5–3.5 μm wide ( x̄ = 2.5 μm, n = 50),
dense, filiform branched, anastomosing above asci, reaching
the ostioles, transversely septate, cellular pseudoparaphyses.
Asci 75–125 × 8–15 μm ( x̄ = 105 × 10 μm, n = 40), 8-spored,
bitunicate, fissitunicate, cylindrical, apically rounded, short,
bulbous pedicel, ocular chamber clearly visible when immature. Ascospores 15–25 × 6–9 μm ( x̄ = 18 × 7 μm, n = 50),
uniseriate, partially overlapping, broad fusiform, tapering
towards the ends, round at both ends, with (1–)4–5 transverse septa, 1-longitudinal septum in each ascospore, initially hyaline, yellowish at maturity, slightly constricted
at the septa, deeply constricted at the median septum, cell
above median septum slightly wider than that below, verruculose, with guttule in each cell, with thin mucilaginous
sheath. Asexual morph: Undetermined.
Culture characters: Colonies on MEA reaching 20 mm
diam. after 4 weeks at 25 °C. Culture from above dark brown
radiating outwardly, wrinkled folded in the middle, dense,
circulate, flattened, umbonate, edge irregular, fairly fluffy;
reverse black in the middle and dark brown at the edge,
orange pigment slightly diffusing into the agar.
Material examined: Thailand, Chiang Rai Province, on
dead stems of Clematis subumbellata, 20 March 2017, C.
Phukhamsakda, CMTH15 (MFLU 17–1479, holotype); extype living culture, MFLUCC 17–2071.
Host: Clematis subumbellata—(This study).
Distribution: Thailand—(This study).
GenBank accession numbers: LSU: MT214557; SSU:
MT226674; ITS: MT310604; tef1: MT394736; rpb2:
MT394685.
Notes: Alloleptosphaeria clematidis is the first report
of Alloleptosphaeria from Thailand and the second Alloleptosphaeria species known on Clematis. Alloleptosphaeria clematidis formed a robust clade with A. italica with
strong support (97% ML/1.00 BYPP, Fig. 22). Alloleptosphaeria clematidis is compatible with the generic concept
of Alloleptosphaeria in having immersed ascomata with a
Fungal Diversity (2020) 102:1–203
Fig. 21 Hermatomyces clematidis (MFLU 17–1493, holotype). a, b
Sporodochia on natural substrate. c Vertical section of sporodochia.
d, e Subicular hyphae. f–h Cylindrical conidia. i–l Mature lenticular
39
conidia. m Germinated conidium. n, o Culture characters on MEA.
Scale bars: b = 500 μm, c = 100 μm, d–m = 20 μm
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Plenodomus biglobosa CBS 127249 T
Plenodomus biglobosa CBS 119951
-Plenodomus pimpinellae CBS 101637 T
Plenodomus guttulatus MFLUCC 15-1876 T
-Plenodomus lingam CBS 275.63
100/1.00
Plenodomus lingam CBS 260.94 T
Plenodomus
hendersoniae CBS 139.78 T
Plenodomus
--/0.96
Plenodomus hendersoniae CBS 113702
62/0.99
79/1.00 Plenodomus agnitus CBS 121.89 T
99/1.00
Plenodomus fallaciosus CBS 414.62
86/1.00
Plenodomus lupini CBS 248.92 T
Plenodomus visci CBS 122783
--/1.00
73/1.00
Plenodomus influorescens CBS 143.84 T
96/1.00
Plenodomus lindquistii CBS 381.67 T
Alternariaster bidentis CBS 134185
100/1.00
Alternariaster helianthi CBS 327.69 T
83/1.00
Alternariaster
96/1.00
100/1.00 Alternariaster centaureae-diffusae MFLUCC 14-0992 T
Alternariaster centaureae-diffusae MFLUCC 15-0009
Sphaerellopsis macroconidiale CBS 233.51 T
100/1.00
Sphaerellopsis
Sphaerellopsis macroconidiale CBS 658.78
73/1.00
Sphaerellopsis filum CBS 234.51 T
Sphaerellopsis paraphysata CPC 21841
96/1.00
100/1.00 Subplenodomus valerianae CBS 499.91 T
Subplenodomus valerianae CBS 630.68
--Subplenodomus apiicola CBS 285.72 T
-- 100/1.00 Subplenodomus apiicola CBS 504.91
Subplenodomus
Subplenodomus
drobnjacensis
CBS
269.92
T
90/1.00
Subplenodomus drobnjacensis CBS 270.92
Subplenodomus violicola CBS 306.68 T
-100/1.00 Heterospora chenopodii CBS 448.68
Heterospora chenopodii CBS 115.96 T
99/1.00
Heterospora
Heterospora dimorphospora CBS 345.78
100/1.00 Heterospora dimorphospora CBS 165.78
-- --/0.97 Paraleptosphaeria nitschkei CBS 306.51 T
--/0.99
Paraleptosphaeria macrospora CBS 114198
--/0.99 Paraleptosphaeria rubi MFLUCC 14-0211 T
Paraleptosphaeria praetermissa CBS 114591
Paraleptosphaeria
83/0.99
Paraleptosphaeria dryadis CBS 643.86
---/0.90
Paraleptosphaeria macrospora CBS 114198
-Paraleptosphaeria orobanches CBS 101638
Leptosphaeria sydowii CBS 125976
96/0.90 Leptosphaeria sydowii CBS 385.80 T
71/-- Leptosphaeria macrocapsa CBS 640.93
79/0.99
Leptosphaeria doliolum CBS 155.94 T Leptosphaeria
Leptosphaeria doliolum CBS 541.66
--/0.97
Leptosphaeria cichorium MFLUCC 14-1063 T
Leptosphaeria slovacica CBS 389.80
100/1.00
--/0.91
61/1.00
Leptosphaeria pedicularis CBS 390.80
Leptosphaeria conoidea CBS 616.75
50/0.99
76/-Leptosphaeria sclerotioides CBS 144.84
Alloleptosphaeria italica MFLUCC 14-0934 T Alloleptosphaeria
56/0.99 99/1.00
Alloleptosphaeria clematidis MFLUCC 17-2071 T
--/1.00
Schleroplectenchymyces clematidicola MFLUCC 17-2180 T
100/1.00
62/1.00
Schleroplectenchymyces jonesii MFLUCC 16-1442 T Schleroplectenchymyces
Neoleptosphaeria rubefaciens CBS 223.77 T
Neoleptosphaeria
Leptosphaeriaceae
62/-98/0.99
Neoleptosphaeria rubefaciens CBS 387.80
Pseudoleptosphaeria etheridgei CBS 125980 Pseudoleptosphaeria
Pyrenochaeta protearum CBS 137997
100/1.00
Parapyrenochaetaceae
Pyrenochaeta protearum CBS 131315 T
Didymella exigua CBS 183.55
100/1.00
(Outgroup)
0.02
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◂Fig. 22 The best scoring RAxML tree with a final likelihood value
of − 13918.604336 based on combined LSU, SSU, ITS and tef1
sequence data for Leptosphaeriaceae. The tree is rooted with a member of the Didymellaceae. Seventy strains were included in the combined gene sequence analyses, which comprise 3257 characters (831
characters for LSU, 951 characters for SSU, 566 characters for ITS,
and 909 characters for tef1, including gap regions). The topology
and clade stability of the combined gene analyses was compared to
the single gene analyses. The matrix had 649 distinct alignment patterns, with 45.29% of undetermined characters and gaps. Estimated
base frequencies were as follows; A = 0.244507, C = 0.225295,
G = 0.271500, T = 0.258698; substitution rates AC = 1.503087,
AG = 2.814632, AT = 2.230882, CG = 0.467105, CT = 7.217503,
GT = 1.000000; gamma distribution shape parameter α = 0.541381.
In our analysis, GTR + I + G model was used for every partition in
Bayesian analysis. The species determined in this study are indicated
in blue. Bootstrap values (BS) greater than 50% BS (ML, left) and
Bayesian posterior probabilities (PP, right) greater than 0.90 are given
at the nodes. Hyphens (-) represent support values less than 50%
BS/0.90 BYPP. The supported values from all analyses are BS ≥ 70%/
BYPP ≥ 0.95
thin-walled peridium of brown to dark brown or reddish
brown, pseudoparenchymatous cells (Ariyawansa et al.
2015b). Based on morphology, A. clematidis is distinguished
by its cylindrical asci and the presence of 1 longitudinal
septum in its yellowish ascospore (Fig. 23). In a BLASTn
search of GenBank, the closest match of the ITS region of
MFLUCC 17–2071 is 92.5% similarity to Subplenodomus
iridicola strain CBS 143395 (NR_159068). Pairwise comparison of the ITS region of MFLUCC 17–2071 with A.
italica (MFLUCC 14–0934) shows 97 nucleotide differences
from 566 base pairs (17.13%).
Sclerenchymomyces Phukhams. & K.D. Hyde, gen. nov.
Index Fungorum number: IF557110; Facesoffungi number: FoF 07287, Fig. 24
Etymology: Genus name reflects the characteristic of scleroplectenchyma tissue type.
Saprobic on dead or dead branch of herbaceous or woody
plants in terrestrial habitats. Sexual morph: Ascomata
covered by plant epidermis, located on the surface of host
substrate, black, shiny, superficial to semi-immersed, solitary, globose, black, ostiolate. Ostioles central, short, filled
with hyaline cells. Peridium composed of blackish to dark
brown cells of textura angularis, thick, multilayered, scleroplectenchymatous cells, cells towards the inside lighter.
Hamathecium composed of numerous, branched, septate,
pseudoparaphyses. Asci 8-spored, bitunicate, fissitunicate,
cylindrical, short-pedicellate. Ascospores uniseriate, partial
overlapping, muriform, broad ellipsoidal, narrowed towards
the ends, initially hyaline, becoming brown at maturity, constricted at median septum, guttulate, surrounded by a mucilaginous sheath (Wanasinghe et al. 2016). Asexual morph:
Conidiomata pycnidial, solitary, sometimes aggregated, uniloculate or multiloculate, erumpent or superficial on host
41
substrate, with black shiny ostioles, globose to subglobose,
coriaceous, dark brown to brown. Ostioles central, papillate, oblong. Conidiomatal wall thick-walled, multilayered,
scleroplectenchymatous or pseudoparenchymatous cells, flat
at base, composed of textura angularis, lined with a thick
hyaline layer bearing conidiogenous cells. Conidiophores
reduced to conidiogenous cells. Conidiogenous cells enteroblastic, phialidic, determinate, discrete, sub-cylindrical to
truncate, smooth-walled, hyaline, arising from the inner
layers of conidiomata. Conidia ellipsoid or cylindrical to
oblong, rounded at both ends, slightly curved, hyaline when
immature, yellowish at maturity, aseptate or septate, guttulate, smooth-walled.
Type species: Sclerenchymomyces clematidis Phukhams.
& K.D. Hyde
Notes: Sclerenchymomyces is introduced for a lineage
comprising Sclerenchymomyces clematidis and S. jonesii
(≡ Neoleptosphaeria jonesii) which received strong support (99% ML/1.00 BYPP, Fig. 22). Based on the multigene phylogenetic analyses, Sclerenchymomyces formed a
separate lineage to Alloleptosphaeria, Leptosphaeria sensu
stricto and Neoleptosphaeria (62% ML/1.00 BYPP). The
members of Leptosphaeriaceae are remarkable by their characteristic scleroplectenchymatous or pseudoparenchymatous tissue types (Ariyawansa et al. 2015b). Two isolates
of Sclerenchymomyces share similar characters in having
black, shiny, superficial to semi-immersed ascomata with a
multilayer of scleroplectenchymatous tissue type (Wanasinghe et al. 2016a, Fig. 24). Sclerenchymomyces shares similar
pycnidial characters with Leptosphaeria sensu stricto and
Neoleptosphaeria in the scleroplectenchymatous or plectenchymatous cell type in the peridium (Ariyawansa et al.
2015b). However, Leptosphaeria species only have transverse septa while Sclerenchymomyces has longitudinal septa
in the ascospores (Fig. 24). Sclerenchymomyces has oblong,
brown phragmoconidia in nature and phoma-like characters
in culture (Gruyter et al. 2013; Wanasinghe et al. 2016). The
combined dataset of the LSU, SSU, ITS and tef1 sequences
for Leptosphaeriaceae revealed a lineage of Sclerenchymomyces as a new genus from Clematis (Fig. 22).
Sclerenchymomyces clematidis Phukhams. & K.D. Hyde,
sp. nov.
Index Fungorum number: IF557111; Facesoffungi number: FoF 07288, Fig. 24.
Etymology: Name refers to the host plant, Clematis.
Holotype: MFLU 16–2492.
Saprobic on dead stems of Clematis vitalba. Sexual
morph: Undetermined. Asexual morph: Conidiomata
187–225 × 85–217 μm ( x̄ = 210 × 130 μm, n = 10), pycnidial, solitary, sometimes aggregated, uniloculate or
multiloculate, erumpent or superficial on host substrate,
with black shiny ostioles visible, globose to subglobose,
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◂Fig. 23 Alloleptosphaeria clematidis (MFLU 17–1479, holotype).
a Appearance of ascomata on Clematis subumbellata. b Close up
of ascoma on host substrate. c Vertical section through ascoma.
d Ostiolar canal. e Section of peridium. f Pseudoparaphyses. g–i
Asci. j–p Ascospores (p Ascospore in 10% Indian ink). q Germinated ascospore. r, s Culture characteristics on MEA. Scale bars:
b = 200 µm, c = 100 µm, d, f–i = 50 µm, e = 20 µm, j–p = 10 µm
coriaceous, dark brown to brown, ostiolate. Ostioles central, papillate, oblong. Conidiomatal wall 14–30(–35) μm
wide, multilayered, scleroplectenchymatous cells, flat at
base, outer layer composed of 5–9 layers of light brown to
brown cells of textura angularis, lined with a thick hyaline
layer bearing conidiogenous cells. Conidiophores reduced
to conidiogenous cells. Conidiogenous cells 3–8 × 1.5–4 μm
( x̄ = 5 × 3 μm, n = 30), enteroblastic, phialidic, determinate,
discrete, sub-cylindrical to truncate, smooth-walled, hyaline, arising from the inner layers of conidiomata. Conidia
11–18 × 2.5–5 μm ( x̄ = 14 × 4 μm, n = 50), broad cylindrical
to oblong, rounded at both ends, hyaline when immature,
yellowish at maturity, slightly curved, 3-septate, with 1(–2)
guttules in each cell, smooth-walled.
Culture characters: Colonies on MEA reaching 40 mm
diam. after 4 weeks at 25 °C. Cultures from above, creambrown, spare mycelia, circular, umbonate, papillate with
fluffy, covered with white aerial mycelium; reverse dark
brown at the centre, cream radiating outwardly.
Material examined: Italy, Forlì-Cesena Province, near
Meldola, on dead aerial branch of Clematis vitalba, 15
November 2013, E. Camporesi, 1518C (MFLU 16–2492,
holotype); ex-type living culture, MFLUCC 17–2180.
Host: Clematis vitalba—(This study).
Distribution: Italy—(This study).
GenBank accession numbers: LSU: MT214558; SSU:
MT226675; ITS: MT310605; tef1: MT394737; rpb2:
MT394686.
Notes: Sclerenchymomyces clematidis is distinct from S.
jonesii in conidial characters (Fig. 24). Sclerenchymomyces clematidis has broad cylindrical to oblong, yellowish
conidia with 3 septa and 1(–2) guttules, while S. jonesii has
hyaline, aseptate conidia (Wanasinghe et al. 2016a). In a
BLASTn search of GenBank, the LSU sequence of S. clematidis (strain MFLUCC 17–2180) was 98.7% similar to S.
jonesii (≡ Neoleptosphaeria jonesii), while the ITS sequence
showed 97.28% similarity to NR_152375. Pairwise comparison of the ITS sequence reveals nine bases pair differences
(1.59%) between S. clematidis and S. jonesii (MFLUCC
16–1442). The tef1 region shows seven bases pair difference between S. clematidis and S. jonesii.
Sclerenchymomyces jonesii (Wanasinghe, Camporesi &
K.D. Hyde) Phukhams. & K.D. Hyde, comb. nov.
43
Basionym: Neoleptosphaeria jonesii Wanasinghe, Camporesi & K.D. Hyde, in Wanasinghe, Camporesi & Hu, Mycosphere 7(9): 1373 (2016)
Index Fungorum number: IF552569; Facesoffungi number: FoF 02716
Notes: Wanasinghe et al. (2016a) introduced Neoleptosphaeria jonesii for a fungus whose morphology and phylogeny are related to Neoleptosphaeria rubefaciens (Gruyter
et al. 2013; Wanasinghe et al. 2016a). The analyses of combined LSU, SSU, ITS and tef1 sequence data for Leptosphaeriaceae showed that the ex-type strain of Neoleptosphaeria
jonesii (MFLUCC 16–1442) clustered with Sclerenchymomyces clematidis (MFLUCC 17–2180), a fungal strain
reported from the same host. Pairwise comparison of the
ITS sequence data reveals 85 bases pair difference from 566
(15.9%, including gap region) between N. jonesii (MFLUCC
16–1442) and N. rubefaciens (CBS 223.77 and CBS 387.80).
According to phylogeny (Fig. 22) coupled with morphology,
we transfer N. jonesii to Sclerenchymomyces.
Host: Clematis vitalba—(Wanasinghe et al. 2016a).
Distribution: Italy—(Wanasinghe et al. 2016a).
Longiostiolaceae Phukhams., Doilom & K.D. Hyde, fam.
nov.
Index Fungorum number: IF557086; Facesoffungi number: FoF 07215, Fig. 25.
Saprobic on dead bark. Sexual morph: Ascomata
immersed to semi-immersed, uniloculate, solitary, globose
to subglobose, coriaceous, base flattened, ostiolate. Ostioles
long, obtuse or dolabriform, central. Peridium thick, comprising several layers of scleroplectenchymatous or pseudoparenchymatous cell types, dark brown to black cells
arranged in a textura angularis or textura globosa. Hamathecium composed of numerous, filiform, septate, branched,
cellular pseudoparaphyses. Asci 4–8-spored, bitunicate,
cylindrical to clavate, pedicellate, with ocular chamber.
Ascospores biseriate, partial overlapping, broad fusiform,
hyaline, brownish at the mature state, multi-septate, with
or without mucilaginous sheath. Asexual morph: coelomycetous-like or hyphomycetous-like structures produced
in culture. Conidiomata scattered, globose to subglobose,
ostiolate. Conidiomatal wall thick, comprising of multilayered textura angularis, pale brown to brown cells. Conidiophores reduced to conidiogenous cells. Conidiogenous cells
annellidic, doliiform to ampulliform. Conidia cylindrical,
hyaline, bud scars disjunctors at base, multi-septate, smooth,
without sheath (Matsumura et al. 2018). Colonies on MEA
produce conidia structures on aerial mycelium, subglobose
to ellipsoidal, initially hyaline, becoming black, aseptate,
smooth-walled.
Type genus: Longiostiolum Doilom, Ariyaw. & K.D.
Hyde, in Li et al., Fungal Diversity 78: 55 (2016)
13
44
Fig. 24 Sclerenchymomyces clematidis (MFLU 16–2492, holotype).
a Appearance of conidiomata on Clematis vitalba. b Close up of conidioma on host substrate. c Vertical section through conidiomata. d
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Fungal Diversity (2020) 102:1–203
Ostiole from above. e Section of conidioma wall. f–i Conidiogenous
cells and conidia. j–l Conidia. Scale bars: b = 200 µm, c = 100 µm,
d = 20 µm, e = 50 µm, f–l = 5 µm
Fungal Diversity (2020) 102:1–203
Notes: Longiostiolaceae is introduced to accommodate
two ascomycetous genera, Crassiperidium and Longiostiolum. Longiostiolum was introduced by Li et al. (2016) for
a fungus associated with Tectona grandis from northern
Thailand. Matsumura et al. (2018) reported Crassiperidium
from twigs of Fagus crenata in Japan. Crassiperidium and
Longiostiolum were associated with wood and share characters in having immersed to semi-immersed, uniloculate
ascomata, thick-walled scleroplectenchymatous or pseudoparenchymatous cell, with hyaline ascospores turning
brownish at maturity (Li et al. 2016; Matsumura et al. 2018).
The asexual morphs of these two genera are different, with
Crassiperidium forming pycnidial conidiomata with hyaline
conidia, and Longiostiolum forming a hyphomycetous-like
structure with globose, black conidia in culture (Fig. 25).
Matsumura et al. (2018) found that Crassiperidium was
related to Cyclothyriellaceae (Pleosporales) based on phylogeny of an SSU, LSU and rpb2 sequence dataset. Longiostiolum was assigned as an incertae sedis taxon in Pleosporales based on combined LSU, SSU, rpb2 and tef1 sequence
data (Li et al. 2016). Phylogenetic analysis based on the LSU
regions of Crassiperidium and Longiostiolum formed a clade
together with moderate support (57% ML, data not shown)
and sister to Cyclothyriellaceae. The concatenated dataset
of LSU, SSU, ITS, tef1 and rpb2 showed Crassiperidium
species (C. octosporum and C. quadrisporum) and Longiostiolum tectonae (MFLUCC 12–0562) forming a close
relationship with strong support (98% ML/1.00 BYPP) and
related to Cyclothyriellaceae (Fig. 2). Therefore, we introduce a new family to accommodate this distinct lineage.
Type genus: Longiostiolum Doilom, Ariyaw. & K.D.
Hyde, in Li et al., Fungal Diversity 78: 55 (2016).
Type species: Longiostiolum tectonae Doilom, Bhat &
K.D. Hyde, in Li et al., Fungal Diversity 78: 55 (2016).
Genera included: Crassiperidium Matsum. & Kaz. Tanaka (Matsumura et al. 2018); Longiostiolum Doilom, Ariyaw. & K.D. Hyde (Li et al. 2016).
Hosts: Fagus crenata, Tectona grandis—(Li et al. 2016;
Matsumura et al. 2018).
Distribution: Japan, Thailand—(Li et al. 2016; Matsumura et al. 2018).
Longiostiolum tectonae Doilom, Bhat & K.D. Hyde, in Li
et al., Fungal Diversity 78: 55 (2016)
Index Fungorum number: IF 551900, Facesoffungi number: FoF 01882, Fig. 25.
Notes: Based on the phylogenetic analysis of the combined LSU, SSU, rpb2 and tef1 sequence data, the genus
Longiostiolum tectonae (MFLUCC 12–0562) formed a distinct lineage in Pleosporales with no statistical support (Li
et al. 2016). In a BLASTn search of GenBank, the closest match of the LSU sequence of MFLUCC 12–0562 is
45
Crassiperidium octosporum (strain MAFF 242971) with
97% similarity, while the ITS sequence showed 86.5%
similarity.
Host: Tectona grandis—(Li et al. 2016).
Distribution: Thailand—(Li et al. 2016).
Lophiostomataceae Sacc. [as ‘Lophiostomaceae’]
Lophiostomataceae was accepted by Mugambi and Huhndorf (2009). Members of this family are saprobes in terrestrial and aquatic habitats (Saccardo 1883; Tanaka and
Harada 2003; Thambugala et al. 2015; Hashimoto et al.
2018). The family is characterized by carbonaceous ascomata with a slit-like ostiolar neck or opening (Zhang et al.
2009). A revision of Lophiostomataceae by Thambugala
et al. (2015) based on multi-locus phylogeny along with the
re-examination of holotype specimens revealed the boundaries of the family. Hashimoto et al. (2018) revised the classification based mainly on ascospore appendages and introduced several new genera. Mapook et al. (2020) introduced
Pseudocapulatispora Mapook & K.D. Hyde and currently,
there are 25 genera in the family (Wijayawardene et al.
2020). A phylogenetic analysis based on a combined LSU,
ITS, SSU, tef1 and rpb2 sequence dataset (Figs. 26, 27) of
lophiostomataceous taxa on Clematis from Europe and Asia
revealed novel species of Neovaginatispora, Pseudocapulatispora, Pseudolophiostoma and Sigarispora, which are
introduced with morphological support.
Neovaginatispora Hashim., K. Hiray. & Kaz. Tanaka
Hashimoto et al. (2018) reanalysed Lophiostomataceae
and segregated Neovaginatispora fuckelii (Sacc.) Hashim.,
K. Hiray. & Kaz. Tanaka from the type species of Vaginatispora based on phylogenetic analyses. Neovaginatispora is
distinct in its thin, sub-carbonaceous peridium of uniform
thickness (Thambugala et al. 2015). We introduce a second species of Neovaginatispora from Clematis viticella
(Fig. 28).
Neovaginatispora clematidis Phukhams., D. Ertz & C. Gerstmans & K.D. Hyde, sp. nov.
Index Fungorum number: IF557117; Facesoffungi number: FoF 07289, Fig. 28.
Etymology: Name refers to the host plant, Clematis.
Holotype: MFLU 17–1514.
Saprobic on dead stems of Clematis viticella.
Sexual morph: Ascomata 145–250 × 108–160 μm
( x̄ = 172 × 134 μm, n = 10), solitary, gregarious, semiimmersed to erumpent, globose to compressed, coriaceous,
dark brown to black, ostiolate. Ostioles 50–65 × 76–83 μm
( x̄ = 58 × 80 μm, n = 5), with a crest-like apex, central,
filled with hyaline periphyses. Peridium 13–28 μm wide
( x̄ = 19 μm, n= 20), uniform, wider and heavily pigmented at the apex, composed of 5(–6) layers of somewhat
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46
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Fungal Diversity (2020) 102:1–203
◂Fig. 25 Longiostiolum tectonae (MFLU 15–3532, holotype). a, b
Appearance of ascomata on host substrate. c Vertical section through
ascoma. d Ostiolar canal. e Section of peridium. f Cellular pseudoparaphyses. g–i Asci. j–l Ascospores. m Culture characters on
MEA. n–q Conidiogenous cell. r Conidia. Scale bars: b = 200 µm, c,
d = 100 µm, e–l = 50 µm, n = 10 µm, o–r = 5 µm
flattened, thin-walled cells of textura angularis, cells
towards the inside lighter, inner layer composed of thin
hyaline gelatinous layer. Hamathecium composed of numerous, dense, 2–3 µm wide, filamentous, branched, septate,
pseudoparaphyses, embedded in a gelatinous matrix. Asci
53–105 × 9–12 µm ( x̄ = 78 × 11 µm, n = 40), 8-spored, bitunicate, fissitunicate, cylindrical-clavate to clavate, with short,
bulbous pedicel, apically rounded, with an ocular chamber.
Ascospores 16–19 × 5–7 µm ( x̄ = 16 × 6 µm, n = 40), biseriate or partially overlapping, hyaline, broad fusiform with
acute ends, tapering towards the ends, 1-euseptate, strongly
constricted at the septum, upper cell broader than lower cell,
smooth-walled, with two guttules in each cell, with 2 µm
wide globose appendages at both ends. Asexual morph:
Undetermined.
Culture characters: Colonies on MEA reaching 30 mm
diam. after 4 weeks at 25 °C. Cultures from above, dark
brown, dense, circular, umbonate, rough surface, dull, undulate, radially furrowed, covered with grey aerial mycelium,
oil droplets formed in the middle of cultures; reverse black
radiating outwardly.
Material examined: Belgium, Flemish Brabant, Meise
Botanic Garden, Bouchout Domain, dead stems of Clematis viticella, 13 June 2017, D. Ertz & C. Gerstmans,
BRCV2 (MFLU 17–1514, holotype); ex-type living culture,
MFLUCC 17–2156.
Host: Clematis viticella—(This study).
Distribution: Belgium—(This study).
GenBank accession numbers: LSU: MT214559; SSU:
MT226676; ITS: MT310606; tef1: MT394738.
Notes: Neovaginatispora clematidis is compatible with
the concept of Neovaginatispora in having thin, uniform
peridium layers with short and globose appendages at both
ends (Thambugala et al. 2015). Neovaginatispora clematidis
was found on Clematis viticella in Belgium whereas Neovaginatispora fuckelii has mainly been isolated from herbaceous
plant in Japan except the strain MFLUCC 17–2652 that was
isolated from Mangifera indica in Taiwan. Neovaginatispora
clematidis is somewhat similar to the type specimen of N.
fuckelii (CBS 101952), but being distinguishable in its broad
fusiform ascospores with a single eusepta (Fig. 28). Neovaginatispora fuckelii and N. clematidis are reported from
different continents and closely related by morphology but
distinct over the five gene loci phylogeny.
In the multigene phylogenetic analyses, the strain formed
a strongly supported clade with N. fuckelii (98% ML/1.00
47
BYPP, Fig. 26). This clade contains four isolates of N.
fuckelii and a single isolate of N. clematidis (Fig. 26). The
ITS sequence shows six nucleotide differences while the
tef1 sequence has 10 nucleotide differences. In a BLASTn
search of the GenBank, the ITS sequence has 98% similarity to Vaginatispora aquatica (MFLUCC 11–0083), while
the tef1 sequence has 93.49% similarity to V. scabrispora
(LC312583). The GCPSR concept was applied to clade (a)
for testing significant recombination between these isolates
(Laurence et al. 2014). A pairwise homoplasy index of
Φw = 1.0 showed that there is no significant recombination
of the gene flow in N. clematidis and N. fuckelii isolates
(Fig. 27a). The new species of Neovaginatispora is therefore
introduced based on morphological support and phylogenetic analysis.
Pseudocapulatispora Mapook & K.D. Hyde
Pseudocapulatispora was introduced by Mapook et al.
(2020) with Pseudocapulatispora longiappendiculata as the
type species. The genus is characterized by slit-like ostioles,
a peridium of textura prismatica and ascospores with relatively long appendages at the end of capped sheaths. We
introduce a second species of Pseudocapulatispora from
Clematis subumbellata based on morphology (Fig. 29) and
phylogenetic analysis (Fig. 26).
Pseudocapulatispora clematidis Phukhams. & K.D. Hyde,
sp. nov.
Index Fungorum number: IF557118; Facesoffungi number: FoF 07290, Fig. 29.
Etymology: The epithet reflects the host, Clematis.
Holotype: MFLU 17–1469.
Saprobic on dead stem of Clematis subumbellata.
Sexual morph: Ascomata 197–386 × 160–252 µm
( x̄ = 300 × 209 μm, n = 10), solitary, scattered, immersed,
with only black shiny ostioles visible, subglobose to compressed, base flattened, coriaceous to carbonaceous at the
apex, brown to black, with a developed pseudoclypeus,
ostiolate. Ostioles 84–148 × 56–86 μm ( x̄ = 107 × 66 μm,
n = 5), with a crest-like apex, central, long, elongated and
laterally compressed, surrounded by a small blackened
pseudoclypeus, with irregular wall, filled with hyaline periphyses. Peridium 10–23(–38 µm at apex) wide ( x̄ = 23 µm,
n= 20), uniform, wider at the apex, heavily pigmented at the
apex, composed of 4(–5) layers of textura prismatica, cells
towards the inside lighter, inner layer composed of thin, hyaline gelatinous layer, at the apex fusing and indistinguishable
from the host tissues. Hamathecium composed of numerous, dense, 1.5–3 µm wide, filamentous, branched, septate,
pseudoparaphyses, situated between and above the asci,
embedded in a gelatinous matrix. Asci 68–117 × 12–22 µm
( x̄ = 99 × 18 µm, n = 40), 8-spored, bitunicate, fissitunicate,
oblong to cylindrical-clavate, with short, furcate pedicel,
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Fungal Diversity (2020) 102:1–203
Sigarispora arundinis MAFF 239449
Sigarispora arundinis KT 651
Sigarispora arundinis JCM 13550
Sigarispora rosicola MFLU 15-1888
Sigarispora scrophulariae MFLUCC 17-0689
99/1.00 Sigarispora ononidis MFLUCC 14-0613b
83/0.99
Sigarispora ononidis MFLUCC 14-0613
Sigarispora coronillae MFLUCC 14-0941
98/1.00 Sigarispora junci MFLUCC 14-0938
75/0.90
Sigarispora ravennica MFLUCC14-0005
Sigarispora sp. IT 1486
79/1.00
100/1.00 Sigarispora caudata KT 530
69/-- Sigarispora caudata MAFF 239453
Sigarispora clematidis MFLUCC 16-1368
55/-Sigarispora clematidicola MFLUCC 16-0446
--/0.97
Sigarispora caulium MAFF 239450
100/1.00
Sigarispora caulium JCM 17669
100/1.00
Sigarispora caulium MFLUCC 15-0036
87/0.97
Sigarispora montanae MFLUCC16-0999
--/0.96 Sigarispora thymi MFLU 15-2131
Sigarispora medicaginicola MFLUCC 17-0681
82/0.99
Sigarispora muriformis MFLUCC 13-0744
Sigarispora caryophyllacearum MFLUCC 17-0749
91/1.00
99/1.00 Lophiopoacea winteri KT 764
Lophiopoacea winteri KT 740
89/1.00
Lophiopoacea paramacrostoma MFLUCC 11-0463
Neotrematosphaeria biappendiculata KT 975
100/1.00
Neotrematosphaeria biappendiculata KT 1124
--/0.99
Neopaucispora rosacearum MFLUCC 17-0807
--/0.96
Coelodictyosporium pseudodictyosporium MFLUCC 13-0451
--/0.98
Coelodictyosporium
muriforme MFLUCC 13-0351
--/0.98
--/0.99
Lophiostoma semiliberum KT 828
50/-Lophiostoma macrostomum KT 635
Lophiostoma crenatum CBS 629.86
Lophiostoma caulium CBS 623.86
Lophiohelichrysum helichrysi MFLUCC 15-0701
80/1.00 Platystomum crataegi MFLUCC 14-0925
Platystomum compressum MFLUCC 13-0343
66/0.99
Platystomum salicicola MFLUCC 15-0632
77/0.99
Platystomum actinidiae KT 521
92/0.99 Guttulispora crataegi MFLUCC 14-0993
Guttulispora crataegi MFLUCC 13-0442
Capulatispora sagittiformis KT 1934
Lophiostoma macrostomoides CBS 123097
--/0.99
Lophiostoma heterosporum CBS 644.86
100/1.00 Biappendiculispora japonica KT 573
68/-Biappendiculispora japonica KT 686-1
Pseudoplatystomum scabridisporum BCC 22835
100/1.00 Pseudoplatystomum scabridisporum BCC 22835
85/-- Pseudolophiostoma obtusisporum KT 3119
51/-- Pseudolophiostoma obtusisporum KT 3098
100/1.00 Pseudolophiostoma obtusisporum KH 336
54/0.99
100/1.00
Pseudolophiostoma obtusisporum KH 228
Pseudolophiostoma obtusisporum KT 2838
64/0.99
100/1.00
Pseudolophiostoma clematidis MFLUCC 17–2081
92/0.99
99/1.00
77/1.00
52/--
(c)
Lophiostomataceae
ML/BYPP
Sigarispora
Lophiopoacea
Neotrematosphaeria
Neopaucispora
Coelodictyosporium
Lophiostoma
Lophiohelichrysum
Platystomum
Guttulispora
Capulatispora
Biappendiculispora
Pseudoplatystomum
(b)
100/1.00
100/1.00
Pseudolophiostoma cornisporum KH 322
Pseudolophiostoma chiangraiense MFLUCC 17–2076
Pseudolophiostoma
100/1.00
Pseudolophiostoma tropicum KH 352
Pseudolophiostoma tropicum KT 3134
97/1.00
87/0.99 Pseudolophiostoma vitigenum JCM 17676
54/0.90 Pseudolophiostoma vitigenum HH 26930
100/1.00
98/1.00
Pseudolophiostoma vitigenum HH 26931
Pseudolophiostoma vitigenum MAFF 239459
Alpestrisphaeria terricola SC-12
Alpestrisphaeria
78/0.99
Pseudocapulatispora longiappendiculata MFLUCC 17-1452
55/-100/1.00
Pseudocapulatispora longiappendiculata MFLUCC 17-1457
Pseudocapulatispora
Pseudocapulatispora clematidis MFLUCC 17-2063
Lophiostoma quadrinucleatum GKM 1233
96/0.99 Flabellascoma minimum KT 2040
100/1.00 Flabellascoma minimum KT 2013
Flabellascoma
Flabellascoma minimum MFLUCC 17-1474
100/1.00
Flabellascoma minimum MFLUCC 18-0233
85/-Flabellascoma cycadicola KT 2034
74/1.00
Crassiclypeus aquaticus KH 104
73/0.95 Crassiclypeus aquaticus KH 185
Crassiclypeus
62/1.00
100/1.00
Crassiclypeus aquaticus KT 970
52/0.93
Crassiclypeus aquaticus KH 56
--/0.99
Leptoparies
Leptoparies palmarum KT 1653
Parapaucispora pseudoarmatispora KT 2237
58/1.00
Parapaucispora
69/0.99
Parapaucispora alpigenum GKM 1091b
100/1.00 Paucispora quadrispora KH 448
100/1.00
Paucispora
Paucispora quadrispora KH 448
Paucispora versicolor KH 110
Dimorphiopsis brachystegiae CPC 22679
Dimorphiopsis
Lentistoma bipolare KH 222
Lentistoma bipolare KH 311
100/1.00
Lentistoma bipolare KT 2415
Lentistoma
62/0.99
100/1.00
Lentistoma bipolare KT 3056
Lentistoma bipolare CBS 115370
100/1.00
100/1.00
Vaginatispora scabrispora KT 2443
100/1.00
Vaginatispora amygdali KT 2248
Vaginatispora
Vaginatispora aquatica MFLUCC 11-0083
100/1.00
Vaginatispora appendiculata MFLUCC 16-0314
99/1.00
Neovaginatispora fuckelii JCM 17672
96/1.00
Neovaginatispora fuckelii KH161
83/0.96
Neovaginatispora fuckelii KT634
98/1.00
Neovaginatispora
Neovaginatispora fuckelii CBS 101952
100/1.00
84/0.99
Neovaginatispora clematidis MFLUCC 17–2156
Neovaginatispora sp. MFLUCC 11-0577
Pseudopaucispora brunneospora KH 227
Pseudopaucispora
Teichospora trabicola C134
Teichospora rubriostiolata TR7
Teichosporaceae
68/0.99
100/1.00
(a)
100/1.00
Outgroups
0.04
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Fungal Diversity (2020) 102:1–203
◂Fig. 26 The best scoring RAxML tree with a final likelihood value of
− 33003.460353 based on combined LSU, ITS, SSU, tef1 and rpb2
sequence data. The tree is rooted with Teichospora rubriostiolata
(TR7) and T. trabicola (C134) in Teichosporaceae. One hundred and
two strains were included in the combined sequence analyses which
comprise 5255 characters (1300 characters for LSU, 921 characters
for ITS, 1004 characters for SSU, 992 characters for tef1 and 1038
characters for rpb2, including gaps). The topology and clade stability of the combined gene analyses was compared to the single gene
analyses. The tree from the maximum likelihood analysis had similar topology to the Bayesian analyses. The matrix had 1938 distinct
alignment patterns with 36.36% undetermined characters or gaps proportions. Estimated base frequencies were as follows: A = 0.248970,
C = 0.248285, G = 0.267331, T = 0.235414; substitution rates
AC = 1.347247, AG = 3.679551, AT = 1.191656, CG = 1.272401,
CT = 7.723696, GT = 1.000000; gamma distribution shape parameter α = 0.579807. In our analysis, GTR + I + G model was used for
each partition in Bayesian posterior analysis. The species determined
in this study are indicated in blue. Bootstrap values (BS) greater
than 50% BS (ML, left) and Bayesian posterior probabilities (BYPP,
right) greater than 0.90 are given at the nodes. Hyphens (-) represent
support values less than 50% BS/0.90 BYPP. Thick branches represent significant support values from all analyses at the genus level
(BS ≥ 70%/BYPP ≥ 0.95)
apically rounded, with an ocular chamber. Ascospores
22–28 × 7–12 µm ( x̄ = 25 × 9 µm, n = 50), biseriate or partially overlapping, ellipsoid, tapering towards the ends,
acute ends, hyaline, 1-euseptate, strongly constricted at
the septum, with guttule in each cell, slightly swollen near
median septum, sheath drawn out from both ends to form
polar appendages, 14–25 × 3–5 µm ( x̄ = 19 × 4 µm, n = 50),
end caps visible at the ends of the appendages, with a lateral
pad-like structure, up to 4 µm wide at side. Asexual morph:
Undetermined.
Culture characters: Colonies on MEA reaching 30 mm
diam. after 4 weeks at 25 °C. Cultures from above, pale
green to yellow brown in the middle, dense, circular,
umbonate, surface rough, dull, fimbriate, radially furrowed,
covered with yellow aerial mycelia, oil droplets formed in
the middle of the culture; reverse cream radiating outwardly,
yellow pigment diffusing in the agar.
Material examined: Thailand, Phayao Province, Phu Sang
District, dead stems of Clematis subumbellata, 20 March
2017, C. Phukhamsakda, CMTH05 (MFLU 17–1469, holotype); ex-type living culture, MFLUCC 17–2063.
Hosts: Clematis subumbellata—(This study).
Distribution: Thailand—(This study).
GenBank accession numbers: LSU; MT214560; SSU:
MT226677; ITS: MT310607; tef1: MT394739; rpb2:
MT394687.
Notes: In the phylogeny (Fig. 26), this species clustered
with the type species, Pseudocapulatispora longiappendiculata (Mapook et al. 2020). Morphological comparison
of P. clematidis and P. longiappendiculata revealed that
P. clematidis (Fig. 29) has larger ascomata (300 × 209 vs
240 × 135 µm), with shorter polar appendages (19 × 4 vs
49
25 × 4.5 µm). The tef1 sequence of Pseudocapulatispora
clematidis (MFLU 17–1469) had 95% similarity to P. longiappendiculata (5% nucleotide differences in the tef1 region).
Therefore, the new strain is introduced as a new species
based on morphology and phylogenetic evidence.
Pseudolophiostoma Thambug., Kaz. Tanaka & K.D. Hyde
Pseudolophiostoma vitigenum is the type species. The
genus was introduced for a lophiostomataceous taxon that
formed a distinct clade from the type species of Lophiostoma (Hirayama and Tanaka 2011; Thambugala et al. 2015).
Five species are listed in Index Fungorum (Index Fungorum 2020). We introduce two new species namely; P. chiangraiensis and P. clematidis, to accommodate species from
Thailand which occurred on Clematis species (Figs. 30, 31).
Pseudolophiostoma chiangraiense Phukhams. & K.D.
Hyde, sp. nov.
Index Fungorum number: IF557119; Facesoffungi number: FoF 07291, Fig. 30.
Etymology: The epithet reflects the location where the
fungus was collected.
Holotype: MFLU 17–1484.
Saprobic on dead stem of Clematis fulvicoma. Sexual
morph: Ascomata 276–293 × 94–294 μm ( x̄ = 286 × 239 μm,
n = 10), solitary, scattered, sometimes gregarious, immersed,
with only black shiny ostioles visible, subglobose or compressed, flattened base, coriaceous, carbonaceous at the
apex, dark brown to black, with a well-developed clypeus,
indistinguishable from substrate, ostiolate. Ostioles
74–127 × 88–129 μm ( x̄ = 94 × 103 μm, n = 5), with opening
by a pore, central, elongated and laterally compressed, irregular wall, carbonaceous, black, filled with hyaline periphyses. Peridium 14–32 μm wide ( x̄ = 23 μm, n= 20), uniform,
wider at the apex, heavily pigmented at the apex, composed
of 3(–4) layers of textura angularis and textura prismatica,
thin-walled cells, cells towards inside lighter than outside,
somewhat flattened, inner layer composed of thin, hyaline
gelatinous layer, at the apex fusing and indistinguishable
from the host tissues. Hamathecium composed of numerous,
dense, 2–3 µm wide, filamentous, branched, septate, cellular
pseudoparaphyses, embedded in a gelatinous matrix. Asci
78–119 × 11–16 µm ( x̄ = 96 × 13 µm, n = 40), 8-spored, bitunicate, fissitunicate, broad cylindrical to cylindrical-clavate,
with furcate pedicel, apically rounded, with an ocular chamber. Ascospores 20–32 × 4–7 µm ( x̄ = 26 × 6 µm, n = 50),
uniseriate, overlapping, hyaline, becoming pale brown at
senescence, broad-fusiform, tapering towards the ends,
acute ends, 1-euseptate, strongly constricted at the septum,
with 1(–2) guttules in each cell, slightly swollen near median
septum, with 6–10 µm wide sheath drawn-out to form polar
appendages, sheath drawn out, with a pad-like structure, up
to 4 μm wide at side. Asexual morph: Undetermined.
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50
Fig. 27 The splits graph from
the pairwise homoplasy index
(PHI) test generated from
the concatenated gene set of
LSU, ITS, SSU, tef1 and rpb2
sequence data of closely related
species of Neovaginatispora
(a), Pseudolophiostoma (b)
and Sigarispora (c) using both
LogDet transformation and
splits decomposition. PHI test
results (Φw) < 0.05 indicates
significant recombination
within the dataset. The strains
determined in this study are in
bold and blue
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Culture characters: Colonies on MEA reaching 30 mm
diam. after 4 weeks at 25 °C. Cultures from above, olivebrown to dark brown in the middle, dense, circular, convex with papillate surface, fluffy, rough surface, radially
furrowed, covered with grey aerial mycelium, oil droplets
formed in the middle of culture; reverse dark brown, radiating outwardly.
Material examined: Thailand, Chiang Rai Province, Mae
Sai District, dead stems of Clematis fulvicoma, 20 March
2017, C. Phukhamsakda & M. van de Bult, CMTH21
(MFLU 17–1484, holotype); ex-type living culture,
MFLUCC 17–2076.
Host: Clematis fulvicoma—(This study).
Distribution: Thailand—(This study).
GenBank accession numbers: LSU: MT214561; SSU:
MT226678; ITS: MT310608; tef1: MT394740; rpb2:
MT394688.
Notes: In the phylogenetic analysis, Pseudolophiostoma chiangraiensis (MFLUCC 17–2076) formed a close
relationship with P. cornisporum KH 322 (100% ML/1.00
BYPP, Fig. 30). We compared P. chiangraiensis (MFLUCC
17–2076) with P. cornisporum (KH 322) and both have
similar characters in being immersed, with only black shiny
ostioles visible on the host, and broad-fusiform, 1-euseptate
ascospores, with acute ends (Fig. 30). Ascomata of P. chiangraiensis are smaller than P. cornisporum (286 × 239 vs
650–700 × 580–650 μm). In a BLASTn search of GenBank,
the ITS sequence had 99% similarity to P. cornisporum KH
322 (NR_158930), while the rpb2 gene was 98% similar to
LC312602.
Pseudolophiostoma clematidis Phukhams. & K.D. Hyde,
sp. nov.
Index Fungorum number: IF557120; Facesoffungi number: FoF 07292, Fig. 31.
Etymology: The epithet reflects the host, Clematis.
Holotype: MFLU 17–1489.
Saprobic on dead stem of Clematis fulvicoma.
Sexual morph: Ascomata 352–370 × 139–243 μm
( x̄ = 358 × 220 μm, n = 10), solitary, scattered, immersed,
with only black shiny ostioles visible, with globose to compressed, flattened base, coriaceous to carbonaceous at the
apex, dark brown to black, with a developed pseudoclypeus,
ostiolate. Ostiole 140–159 × 60–80 μm ( x̄ = 150 × 70 μm,
n = 5), with a crest-like apex and with opening by a pore,
central, elongated and laterally compressed, surrounded by
a small blackened pseudoclypeus, irregular wall, filled with
hyaline periphyses. Peridium 9–28 μm wide ( x̄ = 14 μm,
n= 20), uniform, wider at the apex, heavily pigmented at the
apex, composed of 4(–5) layers of textura prismatica, cells
towards the inside lighter, somewhat flattened, inner layer
composed of thin, hyaline gelatinous layer, at the apex fusing
and indistinguishable from the host tissues. Hamathecium
51
composed of numerous, dense, 2–3 µm wide, filamentous,
branched, septate, pseudoparaphyses, situated between
and above the asci, embedded in a gelatinous matrix. Asci
67–106 × 10–14 µm ( x̄ = 88 × 12 µm, n = 40), 8-spored, bitunicate, fissitunicate, oblong to cylindrical-clavate, with short,
furcate pedicel, apically rounded, with an ocular chamber.
Ascospores 23–31 × 5–9 µm ( x̄ = 26 × 7 µm, n = 50), biseriate
or partially overlapping, hyaline, broad-fusiform, tapering
towards the ends, round at the end, 1-euseptate, strongly
constricted at the septum, with 2(–3) guttules in each cell,
slightly swollen near median septum, with 6–10 µm sheath
drawn out to form polar appendages, with a lateral pad-like
structure, up to 3 μm wide. Asexual morph: Undetermined.
Culture characters: Colonies on MEA reaching 50 mm
diam. after 4 weeks at 25 °C. Cultures from above, olive
gray to dark brown in the middle, dense, circular, umbonate,
surface rough, dull, fimbriate, radially furrowed, covered
with grey aerial mycelium, oil droplets formed in the middle
of culture; reverse: dark brown radiating outwardly.
Material examined: Thailand, Chiang Rai Province, Mae
Sai District, dead stems of Clematis fulvicoma, 20 March
2017, C. Phukhamsakda & M. van de Bult, CMTH27
(MFLU 17–1489, holotype); ex-type living culture,
MFLUCC 17–2081.
Host: Clematis fulvicoma—(This study).
Distribution: Thailand—(This study).
GenBank accession numbers: LSU: MT214562; SSU:
MT226679; ITS: MN393004; tef1: MT394741; rpb2:
MT394689.
Notes: Pseudolophiostoma clematidis (MFLUCC
17–2081) formed a sister clade to P. obtusisporum strains
(100% ML/1.00 BYPP, Fig. 26). Pseudolophiostoma obtusisporum is commonly reported on herbaceous plants or
palms in Japan (Hashimoto et al. 2018). The morphology of
P. clematidis resembles P. obtusisporum except for the larger
ascomata with thinner peridium (Fig. 31). A comparison
of sequence data revealed that the two species differ in all
studied loci (2 bp differences in ITS, 15 bp in tef1 and 13 bp
in rbp2). In a BLASTn search of GenBank, the ITS sequence
had 99% similarity to the type specimen of Pseudolophiostoma obtusisporum (HHUF 30583), while rpb2 sequence
had 98% similarity to LC312605, derived from the same
voucher specimen.
This strain was further evaluated for secondary metabolites and biological activities. Pseudolophiostoma clematidis
(MFLUCC 17–2081) showed moderate growth of microbes
and cytotoxicity in vitro (Macabeo et al. 2020).
Sigarispora Thambug. & K.D. Hyde
Sigarispora was introduced by Thambugala et al. (2015)
to accommodate Lophiostoma ravennicum and some lophiostomataceous taxa that formed a separate clade from the
type species of Lophiostoma. Sigarispora is characterized
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◂Fig. 28 Neovaginatispora clematidis (MFLU 17–1514, holotype). a
Appearance of ascomata on host surface. b Close up of ascoma on
host substrate. c Vertical section through ascoma. d Ostiolar canal. e
Section of peridium. f Pseudoparaphyses. g–i Asci. j–m Ascospores.
n Culture characteristic on MEA. Scale bars: b = 200 µm, c = 100 µm,
d = 50 µm, e–i = 20 µm, j–n = 10 µm
by its immersed to semi-immersed ascomata, with a small
crest-like ostioles, and brown cigar-shaped, multi-septate
ascospores (Thambugala et al. 2015; Wanasinghe et al.
2018). Fourteen species are listed in Index Fungorum (Jayasiri et al. 2015; Index Fungorum 2020). We introduce three
new species of Sigarispora from Clematis species, S. clematidicola, S. clematidis and S. montana (Figs. 32, 33, 34).
Sigarispora clematidicola Phukhams., Camporesi & K.D.
Hyde, sp. nov.
Index Fungorum number: IF557121; Facesoffungi number: FoF 07293, Fig. 32.
Etymology: The epithet reflects the host Clematis.
Holotype: MFLU 20–0419.
Saprobic on dead stems of Clematis vitalba. Sexual morph: Ascomata 255–288 × 217–254 μm
( x̄ = 276 × 238 μm, n = 10), solitary, scattered immersed,
with only black shiny ostioles present, dark brown to black,
globose to compressed, coriaceous, rough-walled, sometimes with dark brown hyphae projecting from the peridium,
pseudoclypeus, ostiolate. Ostioles 60–135 × 67–156 μm ( x̄
= 97 × 102 μm, n = 5), with a crest-like apex, central, elongated and laterally compressed, irregular wall, filled with
hyaline periphyses. Peridium 13–48 μm wide ( x̄ = 29 μm,
n= 20), wider at the apex, thinner at the base, with 6–7 layers of lightly pigmented light brown to brown, thick-walled
cells of textura angularis, lighter pigmented cells towards
inside, somewhat flattened, inner layer composed of hyaline gelatinous layer, fusing and indistinguishable from
the host tissues. Hamathecium numerous, dense, 2–3 µm
wide, filamentous, branched, septate, pseudoparaphyses,
embedded in a gelatinous matrix. Asci 101–125 × 12–19 µm
( x̄ = 115 × 16 µm, n = 40), 8-spored, bitunicate, fissitunicate, broad cylindrical to clavate, with furcate pedicel,
rounded at the apex, with an ocular chamber. Ascospores
22–29 × 7–9 µm ( x̄ = 26 × 8 µm, n = 50), biseriate or partially
overlapping, initially hyaline, becoming yellowish brown at
maturity, broad fusiform, tapering towards the end, mostly
curved (3–)5-transversely euseptate, constricted at the septa,
cells above central septum swollen, guttulate, indentations
present, without or with 2–4 µm sheath drawn out to form
polar appendages. Asexual morph: Undetermined.
Culture characters: Colonies on MEA, slow-growing,
reaching 20 mm diam. after 4 weeks at 25 °C. Culture centrally black dense, circular, flat, umbonate, surface rough;
reverse: mycelium strongly radiating into agar, black.
53
Material examined: Italy, Forlì-Cesena Province, Viale
Salinatore—Forlì, dead aerial branch of Clematis vitalba, 23
February 2015, E. Camporesi, IT2389-A (MFLU 20–0419,
holotype); ex-type living culture, MFLUCC 16–0446.
Host: Clematis vitalba—(This study).
Distribution: Italy—(This study).
GenBank accession numbers: LSU: MT214563; SSU:
MT226680; ITS: MT310609; tef1: MT394742.
Notes: Based on phylogenetic evidence, the isolates of
Sigarispora caudata (KT 530, MAFF 239453), S. clematidicola (MFLUCC 16–0446) and S. clematidis (MFLUCC
16–1368) formed a moderately supported clade (Fig. 26).
Sigarispora clematidicola (Fig. 32) and S. clematidis
(Fig. 33) share morphological similarity such as brown
ascospores with polar appendages, while S. caudata lacks
polar appendages (Table 3). Sigarispora clematidicola, differs from S. clematidis in having smaller ascomata, asci
and ascospores (Table 3). A pairwise comparison of tef1
sequences of S. clematidis and S. caudata showed 98% similarity with 11 nucleotide differences. The tef1 sequences
of S. clematidicola and S. clematidis showed 30 nucleotide
differences. A pairwise homoplasy index showed Φw = 1.0
when genealogical correlation model was applied between
neighboring strains of clade (c) (Fig. 27c). The result is
congruent with the phylogenetic lineages shown in Fig. 26.
Thus, S. caudata (KT 530, MAFF 239453), S. clematidicola
(MFLUCC 16–0446) and S. clematidis (MFLUCC 16–1368)
are significantly different from each other based on molecular as well as morphological data.
Sigarispora clematidis Phukhams., & K.D. Hyde, sp. nov.
Index Fungorum number: IF557122; Facesoffungi number: FoF 07294, Fig. 33
Etymology: The epithet reflects the host Clematis.
Holotype: MFLU 20–0417
Saprobic on dead stem of Clematis vitalba. Sexual morph: Ascomata 416–493 ×337–386 μm
( x̄ = 444 × 369 μm, n = 10), solitary, scattered, immersed,
with only black shiny ostioles visible, globose to compressed, coriaceous, dark brown to black, rough-walled, ostiolate. Ostioles 151–187 × 107–170 μm ( x̄ = 167 × 129 μm,
n = 5), with a crest-like apex and with opening by a pore,
central, elongated and laterally compressed, irregular wall,
filled with hyaline periphyses, pseudoparenchymatous
cells. Peridium 6–46(–76 μm at apex) wide ( x̄ = 32 μm,
n = 20), wider at the apex, thinner at the base, with lightly
pigmented, thick-walled cells of textura angularis, cell
towards the inside lighter, somewhat flattened, inner layer
composed of thick, hyaline gelatinous layer. Hamathecium
composed of numerous, dense, 1.5–2.5 µm wide, filamentous, branched, septate, pseudoparaphyses, situated between
and above the asci embedded in a gelatinous matrix. Asci
96–145 × 10–18 µm ( x̄ = 118 × 14 µm, n = 40), 8-spored,
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◂Fig. 29 Pseudocapulatispora clematidis (MFLU 17–1469, holo-
type). a Appearance of ascoma on host surface. b Close up of
ascoma on host substrate. c Vertical section through ascoma. d Ostiolar canal. e Section of peridium. f Pseudoparaphyses. g–i Asci. j–m
Ascospores. n Ascospore in cotton blue showing the end chambers. o,
p Culture characteristic on MEA. Scale bars: b = 500 µm, c = 200 µm,
d–i = 50 µm, j–n = 20 µm
bitunicate, fissitunicate, clavate, with furcate pedicel,
rounded at the apex, with an ocular chamber. Ascospores
22–30 × 6–9 µm ( x̄ = 24 × 7 µm, n = 60), biseriate or partially
overlapping, broad fusiform, tapering towards the ends, initially hyaline, becoming yellowish brown at maturity, acute
ends, mostly curved, 5–6 transversely eusepta, slightly constricted at the septa, cells above central septum swollen,
guttulate, indentations present, with 5–10 µm long sheath
drawn out to form polar appendages. Asexual morph:
Undetermined.
Culture characters: Colonies on MEA reaching 50 mm
diam. after 4 weeks at 25 °C. Cultures from above, centrally
black, dense, circular, flat, umbonate, surface rough, dull,
fimbriate, radially furrowed, slightly covered with white
aerial mycelium; reverse: mycelium strongly radiating into
the agar, black with radiating brown outwardly.
Material examined: UK, Hampshire, Swanick Lake,
dead stems of Clematis vitalba, 9 July 2016, E.B.G. Jones,
GJ307 (MFLU 20–0417, holotype); ex-type living culture,
MFLUCC 16–1368.
Host: Clematis vitalba—(This study).
Distribution: UK—(This study).
GenBank accession numbers: LSU: MT214564; SSU:
MT226681; ITS: MT310610; tef1: MT394743.
Notes: See note under Sigarispora clematidicola.
Sigarispora montanae Phukhams., Sue, K.D. Hyde, sp. nov.
Index Fungorum number: IF557124; Facesoffungi number: FoF 07295, Fig. 34.
Etymology: The epithet reflects the host species, Clematis
montana.
Holotype: MFLU 20–0418.
Saprobic on dead stems of Clematis montana.
Sexual morph: Ascomata 180–230 × 140–200 μm
( x̄ = 200 × 160 μm, n = 5), solitary, scattered, semiimmersed, with black shiny ostioles, globose, coriaceous,
partial carbonaceous at the apex, dark brown to black,
rough-walled, forming a clypeus like character, ostiolate.
Ostioles (40–)70–100 × 30–60 μm ( x̄ = 80 × 50 μm, n = 5),
with a crest-like apex, central, elongated and laterally compressed, irregular wall, filled with hyaline periphyses. Peridium 13–30 μm wide ( x̄ = 20 μm, n= 30), wider at the apex,
thinner at the base, with 6–7 layers of lightly pigmented light
brown to dark brown, thick-walled cells of textura angularis,
cells towards the inside lighter, at the outside darker, inner
layer composed of thick hyaline gelatinous layer, fusing
55
and indistinguishable from the host tissues. Hamathecium
composed of numerous, dense, 1.2–1.5 µm ( x̄ = 1.3 μm,
n= 30), filamentous, branched, septate, anastomosing,
pseudoparaphyses, embedded in a gelatinous matrix. Asci
105–130 × 10–14 µm ( x̄ = 120 × 15 µm, n = 20), 8-spored,
bitunicate, fissitunicate, broad cylindrical to clavate, with
a long, with furcate pedicel, rounded at the apex, with an
ocular chamber. Ascospores 20–26 × 5–7 µm ( x̄ = 22 × 5 µm,
n = 50), biseriate or partially overlapping, broad fusiform,
tapering towards the ends, initially hyaline, becoming yellowish brown at maturity, mostly curved, 3(–5) transversely
eusepta, constricted at the septa, cells above central septum
swollen, indentations present, with 3–5 µm sheath drawn out
to form polar appendages. Asexual morph: Undetermined.
Culture characters: Colonies on MEA, reaching 25 mm
diam. after 4 weeks at 16 °C. Cultures from above, centrally grey, bearing cream outwardly, dense, circular, flat,
umbonate, surface rough; reverse cream with dark brown.
Material examined: China, Yunnan Province, Dali District, on dead stems of Clematis montana, 20 May 2016,
C. Phukhamsakda, CMCR1 (MFLU 20–0418, holotype);
ex-type living culture, MFLUCC 16–0999.
Host: Clematis montana—(This study).
Distribution: China—(This study).
GenBank accession numbers: LSU: MT214565; SSU:
MT226682; ITS: MT310611; tef1: MT394744.
Notes: In the phylogenetic analysis, Sigarispora montanae (Fig. 34) clustered basal to Sigarispora caulium with
strong support (100% ML/1.00 BYPP, Fig. 26). Sigarispora
montanae has relatively small ascomata and longer pedicels
when compared with S. caulium (Thambugala et al. 2015,
Table 3). In a BLASTn search of GenBank, the closest
matches of tef1 sequence of MFLUCC 16–0999 is 96.5%
similar to S. thymi strain MFLU 15–2131 (MG829241).
The tef1 sequences of S. montanae had 98% similarity to
Sigarispora caulium (MFLUCC 15–0036) with 15 nucleotide differences in the tef1 region (Thambugala et al. 2015;
Wanasinghe et al. 2018).
Melanommataceae Winter (= Pseudodidymellaceae)
Melanomma Nitschke ex Fuckel is the generic type (Winter 1885). Members of this family can occur on twigs or bark
of various woody plants in terrestrial, marine or freshwater
habitats. Melanommataceae is characterized by carbonaceous or coriaceous, gregarious, immersed to erumpent,
globose to subglobose, papillate or epapillate, thick-walled,
pseudoparenchymatous cells, trabeculate pseudoparaphyses, ascospores uniseriate or biseriate, fusoid to ellipsoidal,
or muriform, hyaline or brown, 1 to multi-septate, with or
without a mucilaginous sheath. Their asexual morphs can
be hyphomycetes or coelomycetes (Sivanesan 1984; Huhndorf 1993; Liew et al. 2000; Tian et al. 2015; Wanasinghe
et al. 2018). Wijayawardene et al. (2018) accepted 24 genera,
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◂Fig. 30 Pseudolophiostoma chiangraiense (MFLU 17–1484, holo-
type). a Appearance of ascoma on host surface. b Close up of ascoma
on host substrate. c Vertical section of ascoma. d Ostiolar canal. e
Section of peridium. f Pseudoparaphyses. g–i Asci. j–m Ascospores.
n Senescent spores. o Culture characteristics on MEA. Scale bars: b,
c = 200 µm, d–i = 50 µm, j–n = 10 µm
thereafter Wanasinghe et al. (2018) introduced five more
genera to Melanommataceae. An analysis of fungal collections on Clematis vitalba revealed a novel genus, Neobyssosphaeria based on a multi-gene phylogeny of LSU, SSU
and ITS sequence data for Melanommataceae (Fig. 35).
Neobyssosphaeria Wanas., E.B.G. Jones & K.D. Hyde, gen.
nov.
Index Fungorum number: IF557189; Facesoffungi number: FoF 07281, Fig. 36.
Etymology: Name refers to the similarity of its morphology to Byssosphaeria.
Saprobic on decaying wood or herbaceous plants in terrestrial habitats. Sexual morph: Ascomata immersed, ostioles orange, solitary or gregarious, globose to depressedglobose, coriaceous, ostiolate. Ostioles central, papillate,
opening by a pore, filled with periphyses with orange pigment around the pore. Peridium thick, multilayered, outer
layer composed of reddish brown cells of textura angularis,
inner layer composed of thin and hyaline cells of textura
angularis. Hamathecium composed of numerous, dense,
filiform, branched, anastomosing, transversely septate, trabeculate pseudoparaphyses. Asci 8-spored, bitunicate, fissitunicate, cylindrical-clavate, pedicelate with an occular
chamber. Ascospores biseriate, broadly fusiform, hyaline,
constricted at the septa, guttulate in each cell, with or without a mucilaginous sheath. Asexual morph: Undetermined.
Type species: Neobyssosphaeria clematidis Wanas.,
Phukhams., E.B.G. Jones & K.D. Hyde
Notes: Neobyssosphaeria is established as a monotypic
genus with N. clematidis as the type species. Based on multigene analyses, isolate MFLUCC 17–0794 formed a basal lineage to Byssosphaeria, but this placement is not supported
by the statistical analyses. Byssosphaeria is characterized
by superficial ascomata with bright yellow, orange or red
colouration around the ostioles, hairy hypha protruding from
the outside of peridium, long pedicellate asci, and hyaline
or pale brown ascospores (Barr 1990; Tian et al. 2015).
Neobyssosphaeria is similar to Byssosphaeria in its orange
apex (Zhang et al. 2012; Hyde et al. 2013). However, Neobyssosphaeria is distinguished by its immersed ascomata
with central papilla filled with periphyses, cellular pseudoparaphyses and broad fusiform and hyaline ascospores
(Fig. 36). Our taxon failed to produce an asexual morph
in culture and therefore it is not possible to compare the
57
asexual characteristics with species of Byssosphaeria. In
the BLASTn search of GenBank, the closest match of the
LSU region of MFLUCC 17–0794 is Uzbekistanica yakutkhanika (strain MFLUCC 17–0842) with 94.48% similarity
(accession number MG829090). Uzbekistanica is however
phylogenetically not closely related to our new collection
(Fig. 35). Therefore, we believe it is taxonomically prudent
to name our collection in a new genus until further studies
are carried out with further taxonomic sampling and DNA
based sequence analyses.
Neobyssosphaeria clematidis Wanas., Phukhams., E.B.G.
Jones & K.D. Hyde, sp. nov.
Index Fungorum number: IF557190; Facesoffungi number: FoF 07282, Fig. 36.
Etymology: Named after the host genus, Clematis.
Holotype: MFLU 17–0614.
Saprobic on dead stems of Clematis vitalba. Sexual morph: Ascomata 525–550 × 500–520 μm
( x̄ = 535 × 510 μm, n = 5), immersed, ostiole part orange, solitary or gregarious, globose to depressd-globose, coriaceous,
indistinguishable from host tissue, brown to pale brown,
rough-walled, ostiolate. Ostioles 235–270 × 190–230 μm
( x̄ = 250 × 200 μm, n = 5), central, papillate, opening by a
pore, filled with periphyses, with orange pigment around
pore. Peridium 30–50(–70 μm at apex) wide, thick, multilayered, outer layer composed of heavily pigmented, reddish brown cells of textura angularis, inner layer composed
of thin and hyaline cells of textura angularis. Hamathecium composed of numerous, 1.6–3 μm ( x̄ = 2.3 μm,
n = 50), dense, filiform, branched, anastomosing, septate,
cellular pseudoparaphyses. Asci 160–210 × 20–30 μm
( x̄ = 185 × 25 μm, n = 20), 8-spored, bitunicate, fissitunicate, cylindrical, long pedicellate with a furcated base,
clavate, apically rounded, with ocular chamber. Ascospores
55–75 × 8–14 μm ( x̄ = 60 × 11 μm, n = 30), biseriate, partially overlapping, broad fusiform, sometimes inequilateral, with acute ends, hyaline, 7-euseptate, constricted at
the septa, cell above median septum enlarged, with guttules
in each cell, rough-walled, without mucilaginous sheath.
Asexual morph: Undetermined.
Culture characters: Colonies on MEA reaching 20 mm
diam. after 4 weeks at 16 °C. Above cream with orange
in the middle, with white edge, medium dense, flattened,
umbonate, floccose; reverse: cream, thin, flat, circular.
Material examined: UK, Hampshire, Botley wood, on
dead stems of Clematis vitalba, 25 May 2016, E.B.G. Jones,
GJ 298 (MFLU 17–0614, holotype); ex-type living culture,
MFLUCC 17–0794.
Host: Clematis vitalba—(This study).
Distribution: UK—(This study).
GenBank accession numbers: LSU: MT214566; SSU:
MT408594.
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◂Fig. 31 Pseudolophiostoma clematidis (MFLU 17–1489, holotype).
a Appearance of ascoma on host surface. b Close-up of ascoma on
host substrate. c Vertical section of ascoma. d Ostiolar canal. e Section of peridium. f Pseudoparaphyses. g–h Asci. i–l Ascospores. m
Germinated ascospore n, o Culture characteristics on MEA. Scale
bars: a = 500 µm, b, c = 200 µm, d, e = 100 µm, f–h = 50 µm, i–
l = 10 µm
Notes: The new fungus morphologically resembles many
genera in Pleosporales (e.g. Angustimassarina, Aquastroma,
Aquilomyces, Carinispora, Falciformispora, Keissleriella,
Lophiopoacea, Lophiostoma, Parabambusicola, Quintaria) in its cylindrical-clavate asci and hyaline, fusiform
ascospores with large guttule in each cell (Zhang et al. 2012;
Tanaka et al. 2015; Thambugala et al. 2015). However,
these taxa are phylogenetically not closely related to Neobyssosphaeria clematidis (see Notes Neobyssosphaeria for
further details). Neobyssosphaeria clematidis differs from
Byssosphaeria species by its immersed ascomata, lacking
hairy hypha protruding from the outside of the peridium
(Tian et al. 2015).
Neomassarinaceae Mapook & K.D. Hyde
The type genus of Neomassarinaceae is Neomassarina.
The family is phylogenetically related to Sporormiaceae but
the characters of the sexual morphs are different. Neomassarinaceae can be characterized by its crest-like ostiolar
necks, with a carbonaceous texture, cylindrical to cylindricclavate asci and broad fusiform ascospores surrounded by a
mucilaginous sheath (Thambugala et al. 2015; Tibpromma
et al. 2017; Mapook et al. 2020). In this study, a dataset
of LSU, SSU, ITS, tef1 and rpb2 were used for phylogenetic analyses (Fig. 2). A collection associated with Clematis formed a clade related to Neomassarinaceae. Thus, we
introduce a new genus Pseudohelminthosporium and a first
report of the asexual morph in Neomassarinaceae (Fig. 37).
Pseudohelminthosporium Phukhams. & K.D. Hyde, gen.
nov.
Index Fungorum number: IF557191; Facesoffungi number: FoF 07283, Fig. 37.
E t y m o l o g y : Re fe r r i n g t o i t s s i m i l a r i t y t o
Helminthosporium.
Saprobic on decaying wood or herbaceous plant material in terrestrial habitats. Sexual morph: Undetermined.
Asexual morph: Colonies on host substrates, effuse, black,
hairy, scattered, dark brown. Mycelium immersed from the
substrate forming dark brown stroma-like aggregations.
Conidiophores macronematous, simple, solitary, branched
at the apex, stripes straight or flexuous, cylindrical, dark
brown to reddish brown, multi-septate, with well-defined
small pores at the apex, smooth or verruculose. Conidiogenous cells monotretic or polytretic, integrated, terminal on
conidiophores, doliiform to oblong, pale brown. Conidia
59
phragmosporous, acrogenous, broad fusiform or obclavate,
dark brown to reddish brown, distoseptate when young,
becoming euseptate at maturity, verrucose, dark brown bud
scars disjunctions present at the basal position, hyaline, elongate cells at the upper end of conidia, with guttules in each
cell.
Type species: Pseudohelminthosporium clematidis
Phukhams. & K.D. Hyde
Notes: Pseudohelminthosporium is established as a
monotypic genus. Based on the multi-gene analysis (Fig. 2),
the isolate MFLUCC 17–2086 formed a basal lineage with
Neomassarina species with moderate support (0.93 BYPP).
Pseudohelminthosporium is morphologically similar to Helminthosporium (Massarinaceae) in having brown to dark
brown phragmosporous conidia (Voglmayr and Jaklitsch
2017). Helminthosporium species have obclavate to rostrate,
pale golden brown to brown conidia, with distoseptate and
angular lumina. Pseudohelminthosporium is distinguishable
by its solitary stipes with monotretic or polytretic conidiogenous cells, phragmosporous, broad fusiform or obclavate
conidia, distoseptate when young, becoming euseptate at
maturity, with hyaline, elongate cells at the upper end of the
conidia and with large guttule in each cell (Fig. 37).
Pseudohelminthosporium clematidis Phukhams. & K.D.
Hyde, sp. nov.
Index Fungorum number: FoF 07284; Facesoffungi number: FoF 07284, Fig. 37.
Etymology: The epithet “clematidis” refers to the host
substrate.
Holotype: MFLU 17–1494.
Saprobic on decaying wood or herbaceous plant material in terrestrial habitats. Sexual morph: Undetermined.
Asexual morph: Colonies on Clematis sikkimensis effuse,
black, hairy, scattered, dark brown. Mycelium immersed,
on the substrate surface forming stroma-like aggregations
of dark brown sheet. Conidiophores 125–435 × 9–20 μm
( x̄ = 230 × 12 μm, n = 20), macronematous, simple, solitary,
branched at the apex, stipes straight or flexuous, cylindrical, erect, septate, smooth, dark brown to reddish brown,
8–14 septa, brown, well-defined small pores with dark scar
at the apex, smooth or verruculose. Conidiogenous cells
17–60 × 9–12 μm ( x̄ = 40 × 12 μm, n = 10), monotretic or
polytretic, integrated, terminal, becoming intercalary, doliiform to oblong, pale brown. Conidia 63–142 × 16–26 μm
( x̄ = 87 × 20 μm, n = 20), phragmosporous, acrogenous,
broad fusiform or obclavate, distoseptate at the early state,
3–5-euseptate at maturity, slightly constricted at septa, dark
brown to reddish brown, verrucose, gradually tapering to
13 μm at the distal end, with a dark brown to black scar
present at the base, subhyaline, elongate cells at the upper
end of conidia, with (1–)2 guttules in each cell.
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◂Fig. 32 Sigarispora clematidicola (MFLU 20–0419, holotype). a
Appearance of ascoma on host surface. b Close up of ascoma. c Vertical section of ascoma. d Ostiolar canal. e Peridium. f Pseudoparaphyses. g–i Asci. j–o Ascospores (o Ascospore in 10% Indian ink;
note the boundary of polar appendages). Scale bars: c, d = 100 µm, e,
f = 50 µm, g–i = 20 µm, j–o = 10 µm
Culture characters: Colonies on MEA reaching 40 mm
diam. after 4 weeks at 25 °C. Culture from above, brownish
grey, dark green, white in the center, forming cream fluffy
mycelium at the edge, dense, umbonate, raised with concave
edge, rough, dull, lobate, radially furrowed, with brown pigment slightly diffusing into the agar; reverse dark brown.
Material examined: Thailand, Chiang Rai Province, Mae
Sai District, dead stems of Clematis sikkimensis, 24 June
2017, C. Phukhamsakda & M. van de Bult, CMTHDT02
(MFLU 17–1494, holotype); ex-type living culture,
MFLUCC 17–2086.
Host: Clematis sikkimensis—(This study).
Distribution: Thailand—(This study).
GenBank accession numbers: LSU: MT214567; SSU:
MT226683; ITS: MT310612; tef1: MT394627; rpb2:
MT394690.
Notes: In a BLASTn search of GenBank, the closest
match for the LSU sequence of MFLUCC 17–2086 was
Preussia terricola strain CBS 317.65 (GQ203725) with
96.77% similarity. The closest match for the ITS region
was Forliomyces uniseptata strain MFLUCC 15–0765
(NR_154006). Based on the multi-locus phylogenetic support (Fig. 2), Pseudohelminthosporium formed a separate
lineage related to Neomassarina species but lacked backbone support. Therefore, Pseudohelminthosporium is treated
as a distinct genus in Neomassarinaceae.
Nigrogranaceae Jaklitsch & H. Voglmayr
Nigrogranaceae was erected to accommodate a wellsupported clade of Nigrograna in Pleosporales, with
Nigrograna as the generic type (Jaklitsch and Voglmayer
2016). Nigrogranaceae was isolated from the bark of wood
and is characterized by its immersed to erumpent ascomata,
surrounded by a subiculum with only papillate ostioles seen
on the host substrate. The remarkable characteristics of
Nigrogranaceae include broad-fusiform to narrowly ellipsoid
ascospores with 1–3-euseptate, and pale to chocolate brown
ascospores. The asexual morph has pseudoparenchymatous
pycnidia and rod-like to ellipsoid, 1-celled, hyaline or subhyaline conidia (Gruyter et al. 2013). Only Nigrograna is
accepted in Nigrogranaceae (Wijayawardene et al. 2017,
2018, Fig. 38).
Nigrograna Gruyter, Verkley & P.W. Crous
Gruyter et al. (2013) introduced Nigrograna for isolates
reported as infectious human pathogen (Serrano et al. 1998;
61
Ahmed et al. 2018). The genus is typified by Nigrograna
mackinnonii (Gruyter et al. 2013), and includes 13 species
(Index Fungorum 2020) based on phylogenetic analyses
of a concatenated dataset of LSU, SSU, ITS and tef1 data
(Fig. 38) coupled with morphology. Nigrograna chromolaenae and N. oblique, on Clematis species, are new host
records (Figs. 39, 40).
Nigrograna chromolaenae Mapook & K.D. Hyde, Fungal
Divers (2020), new host record
Facesoffungi number: FoF 07297, Fig. 39.
Saprobic on dead stems of Clematis fulvicoma. Sexual
morph: Ascomata 135–251 × 87–238 μm ( x̄ = 178 × 141 μm,
n = 10), on the surface of the host, solitary, gregarious,
immersed, the surface of host slightly swollen, subglobose to
depressed, coriaceous, dark brown to brown, smooth-walled,
papillate. Ostioles 119 × 95 μm, central, dark brown, filled
with periphyses. Peridium 9–15 μm wide, multilayered,
composed of 5–7 layers of dark brown cells of textura angularis, heavily pigmented at the outer layer, the inner layer
composed of hyaline and thin layer. Hamathecium composed
of numerous, 0.8–1.5 μm wide ( x̄ = 0.8 μm, n = 50), dense,
filiform, branched, septate, pseudoparaphyses, anastomosing
above asci. Asci 45–60 × 7–9 μm ( x̄ = 46 × 7 μm, n = 30),
8-spored, bitunicate, fissitunicate, cylindrical-clavate to
clavate, with furcate pedicel, apically rounded, ocular chamber visible when immature. Ascospores 11–13 × 3–5 μm
( x̄ = 12 × 4 μm, n = 50), biseriate, ellipsoid to broad fusiform,
sometimes inequilateral, with rounded ends, cognac brown,
(1–)3-septate, constricted at septum, cell above median septum enlarged, with guttule in each cell, smooth-walled, without mucilaginous sheath. Asexual morph: Undetermined.
Culture characters: Colonies on MEA reaching 30 mm
diam. after 4 weeks at 25 °C. Cultures from above, dark
brown, covered with white mycelia in the center, dense,
irregular, umbonate, lobate, velvety, with flat parchmentlike stromatic sheets; reverse dark brown, lobate.
Material examined: Thailand, Chiang Rai Province, on
dead branches of Clematis fulvicoma, 20 April 2017, C.
Phukhamsakda, CMTH25 (MFLU 17–1487); living culture,
MFLUCC 17–2079.
Hosts: Chromolaena odorata, Clematis fulvicoma—
(Mapook et al. 2020; this study).
Distribution: Thailand—(Mapook et al. 2020; this study).
GenBank accession numbers: LSU: MT214568; SSU:
MT226684; ITS: MT310613; tef1: MT394628; rpb2:
MT394691.
Notes: Based on phylogenetic analyses (Fig. 38),
MFLUCC 17–2079 clustered with Nigrograna chromolaenae (MFLUCC 17–1437). Mapook et al. (2020) introduced
N. chromolaenae from a stem of Chromolaena odorata collected in Thailand. Characters of the ex-type strain such as
ascomata, asci and ascospores were reported in Mapook
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◂Fig. 33 Sigarispora clematidis (MFLU 20–0417, holotype). a
Appearance of ascoma on host surface. b Close up of ascoma. c Section of ascoma. d Ostiolar canal. e Section of peridium. f Pseudoparaphyses. g–i Asci. j–m Ascospores. n Culture characteristics on MEA.
Scale bars: b = 500 µm, c = 200 µm, d–i = 50 µm, j–m = 10 µm
et al. (2020) and were similar to our collection (Fig. 39). A
comparison of the ITS and tef1 sequence data revealed no
significant difference between our new collection and the
ex-type strain. Therefore, we introduce Nigrograna chromolaenae on Clematis as a new host record.
Nigrograna obliqua Jaklitsch & H. Voglmayr, Studies in
Mycology 85: 59 (2016), new host record
Index Fungorum number: IF817783; Facesoffungi number: FoF 07298, Fig. 40.
Saprobic on dead stems of Clematis vitalba. Sexual morph: Ascomata 193–450 × 188–294 μm
( x̄ = 266 × 258 μm, n = 5), on the surface of the host, solitary, sometimes gregarious, immersed, only black shiny
ostioles visible, subglobose to depressed, coriaceous,
brown to reddish brown, smooth-walled, papillate. Ostioles
80–165 × 74–96 μm, central, dark brown, filled with periphyses. Peridium 11–27 μm wide, with brown hyphae projecting from the outer layer, composed of 8–10 layers of dark
brown cells of textura angularis, heavily pigmented at outer
layer, the inner layer hyaline and thin. Hamathecium composed of numerous, 1–1.3 μm ( x̄ = 1.2 μm, n = 50), dense,
filiform, branched, transverse septate, pseudoparaphyses.
Asci 60–101 × 10–16 μm ( x̄ = 86 × 12 μm, n = 20), 8-spored,
bitunicate, fissitunicate, clavate to broad cylindrical, with
furcate pedicel, apically rounded, ocular chamber visible
when immature. Ascospores 16–20 × 5–8 μm ( x̄ = 18 × 6 μm,
n = 50), biseriate, overlapping, ellipsoid to broad fusiform,
sometimes inequilateral, with rounded ends, initially hyaline, pale brownish to brown at the maturity, (1–)3-eusepta,
constricted at septa, cell above median septum enlarged, the
second cell slightly wider than others, with guttule in each
cell, smooth-walled, without mucilaginous sheath. Asexual
morph: Undetermined.
Culture characters: Colonies on MEA reaching 30 mm
diam. after 4 weeks at 16 °C. Cultures from above, dark
brown, covered with greyish orange mycelia on the surface, dense, irregular, umbonate, lobate, velvety, flat parchment-like stromatic sheets developed, reverse dark brown,
fimbriae.
Material examined: Italy, Forlì-Cesena Province, Corniolo—Santa Sofia, dead aerial branch of Clematis vitalba, 17
February 2014, E. Camporesi, IT 1726 (MFLU 16–0190);
living culture, MFLUCC 14–0945.
Hosts: Clematis vitalba, Ribes uva-crispa, Salix caprea,
Sambucus nigra, S. racemosa—(Jaklitsch et al. 2016; this
study).
63
Distribution: Austria, France, Italy, UK—(Jaklitsch et al.
2016; this study).
GenBank accession numbers: LSU: MT214569; ITS:
MT310614.
Notes: Our new collection is morphologically similar and
phylogenetically related to the type species of Nigrograna
obliqua (strain MF2). MFLUCC 14–0945 is not very different from that of the type species (Fig. 40). Jaklitsch and
Voglmayr (2016) introduced four strains of N. obliqua
from various shrubs and trees. In the phylogenetic analyses MFLUCC 14–0945 formed a close relationship with the
ex-type strain reported from Salix caprea (84% ML/0.99
BYPP). Interestingly, three strains of N. obliqua (BW4, KE
and MRP) formed a separate clade from the type strain.
However, the morphological data of N. obliqua strains
BW4, KE and MRP were not available for comparison. We
report N. obliqua on Clematis vitalba as a new host record
(Fig. 40).
Occultibambusaceae Dai & K.D. Hyde
Occultibambusaceae is typified by Occultibambusa D.Q.
Dai & K.D. Hyde. The species in this family are generally
associated with monocotyledon such as Occultibambusa,
Seriascoma, and Versicolorisporium. Doilom et al. (2017)
reported Neooccultibambusa, another species from Tectona
grandis and Brunneofusispora from dead wood in China.
Occultibambusaceae is characterized by immersed, solitary
to gregarious ascomata with ostioles, broadly cylindrical to
clavate asci with broad-fusiform, hyaline to dark brown usually septate ascospores. Pycnidia are reported in the asexual
morph of this family (Hatakeyama et al. 2008; Dai et al.
2017).
Brunneofusispora Huang & K.D. Hyde
Brunneofusispora sinensis is the type species. The genus
was recorded from dead wood near a river and is characterized by immersed, uniloculate ascomata, cylindrical to
clavate and short pedicellate asci, and hyaline to brown,
broadly fusiform ascospores (Phookamsak et al. 2019).
Brunneofusispora clematidis is reported as a second species of Brunneofusispora on Clematis and is illustrated. An
updated phylogenetic analysis of a concatenated dataset
of LSU, tef1, ITS and SSU sequence data is presented in
Fig. 41.
Brunneofusispora clematidis Phukhams. & K.D. Hyde, sp.
nov.
Index Fungorum number: IF557194; Facesoffungi number: FoF 07299, Fig. 42.
Etymology: Name refers to the host Clematis.
Holotype: MFLU 17–1478.
Saprobic on dead stems of Clematis subumbellata.
Sexual morph: Ascomata 160–210 × 165–186 μm ( x̄
13
64
Fig. 34 Sigarispora montanae (MFLU 20–0418, holotype). a
Appearance of ascomata on host surface. b Vertical section of
ascoma. c Ostiolar canal. d Section of peridium. e Pseudoparaphy-
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ses. f, g Asci. h–k Ascospores. l Germinated ascospore. m, n Culture characteristics on MEA. Scale bars: b = 100 µm, c–g = 50 µm,
h–k = 10 µm
Fungal Diversity (2020) 102:1–203
65
Table 3 Morphological comparison of known Sigarispora species
Species
Ascomata (μm)
Asci (μm)
320–416 × 250– 94–
310
112.5 × 12.5–
14.5
S. caryophylla- 420–500 × 400– 80–120 × 10–12
cearum
500
S. caudata
145–210 × 210– 86–
305
112.5 × 10.5–
13
S. caulium
180–340 × 200– 75–100 × 12–14
280
S. clematidicola 255–288 × 217– 101–
254
125 × 12–19
S. clematidis
416–493 ×337– 96–145 × 10–18
386
S. coronillae
350–400 × 390– 120–
450
140 × 14–17
Sigarispora
arundinis
Ascospores
Septa
Appendages
(μm)
22–32.5 × 6.3–
7.9
5
Both ends
Phragmites
australis
Thambugala et al.
(2015)
30–40 × 7–8.5
5–9
Not present
23.5–
34.5 × 5.5–7
(4)–5–(6)
Not present
Caryophyllaceae
Dactylis glomerata
Wanasinghe et al.
(2018)
Thambugala et al.
(2015)
18–25 × 5–8
5
Both ends
22–29 × 7–9
(3–)5
2–4
Thambugala et al.
(2015)
This study
22–30 × 6–9
5–6
5–10
20–26 × 8–10
4–5 transversely
eusepta, 2–4
vertical septa
5–7
Not present
Herbaceous
Rosa canina
Clematis
vitalba
Clematis
vitalba
Coronilla
emerus
Juncus sp.
Wanasinghe et al.
(2018)
Medicago
falcata
Clematis montana
Helichrysum
italicum
Wanasinghe et al.
(2018)
This study
200–300 × 200– 120–
250
140 × 14–18
S. medicaginicola
S. montanae
400–500 × 300– 80–120 × 10–15 24–28 × 6.5–7
350
180–230 × 140– 105–
20–26 × 5–7
200
130 × 10–14
425–660 × 335– 84–130 × 12–18 18–28 × 7–12
560
S. ononidis
S. ravennica
References
Size (μm)
S. junci
S. muriformis
Host
26–33 × 6.5–8
240 × 311
96–169 × 17–19 27–34 × 11–12
211–282 × 121– 55–70 × 9–11
18–21 × 4–6
187
400–500 × 300– 80–120 × 10–15 17–23 × 5–6
350
S. scrophulariae 280–350 × 250– 70–90 × 10–12 18–22 × 5–6
300
S. thymi
600–700 × 450– 80–120 × 12–16 23–33 × 6–7
550
S. rosicola
= 185 × 176 μm, n = 5), on the surface of the host, solitary,
gregarious, uniloculate, semi-immersed, shiny, globose to
depressed, coriaceous, dark brown to black, rough-walled,
papillate, ostiolate. Ostioles 96–105 × 86–110 μm, central, dark brown, filled with short periphyses. Peridium
8–17(–28) μm wide, multilayered, composed of 4–5 layers
of dark brown cells of textura prismatica, heavily pigmented
5–6
5–8 μm of
mucilaginous
sheath
8–10
3(–5)
3–5
5–8 transversely
eusepta, 2–3
longitudinal
septa
3–5
6
Not present
This study
Thambugala et al.
(2015)
Tibpromma et al.
(2017)
Ononis spinosa
Poaceae
Li et al. (2016)
Thambugala et al.
(2015)
5
Not present
Not present,
sometimes
with terminal
appendages
2–5
Rosa sp.
4–5
2–4
4–6
Not present
Scrophularia
donetzica
Thymus marshallianus
Wanasinghe et al.
(2018)
Wanasinghe et al.
(2018)
Wanasinghe et al.
(2018)
at outer layer, the inner layer hyaline and thin. Hamathecium
composed of medium dense, 1.7–4.5 μm wide ( x̄ = 2.7 μm,
n = 50), filiform, branched, septate, cellular pseudoparaphyses. Asci 45–117 × 12–24 μm ( x̄ = 73 × 17 μm, n = 20),
8-spored, bitunicate, fissitunicate, clavate, with furcate
pedicelate, apically rounded, ocular chamber visible when
immature. Ascospores 17–35 × 5–10 μm ( x̄ = 27 × 9 μm,
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◂Fig. 35 The best scoring RAxML tree with a final likelihood value of
− 9510.121854 based on combined LSU, SSU and ITS sequence data
for Melanommataceae. The tree is rooted with members of the Cyclothyriellaceae. Seventy-one strains were included in the combined
sequence analyses which comprised 2564 characters (921 characters
for LSU, 1061 characters for SSU, 582 characters for ITS, including
gap regions). The matrix had 602 distinct alignment patterns, with
31.43% of undetermined characters and gaps. Estimated base frequencies were as follows; A = 0.254800, C = 0.214111, G = 0.278600,
T = 0.252489; substitution rates AC = 2.132797, AG = 2.875535,
AT = 1.394124, CG = 0.797579, CT = 9.867341, GT = 1.000000;
gamma distribution shape parameter α = 0.481484. In our analysis,
GTR + I + G model was used for each partition in Bayesian posterior
analysis. The species determined in this study, is indicated in blue.
Bootstrap values (BS) greater than 50% BS (ML, left) and Bayesian posterior probabilities (BYPP, right) greater than 0.90 are given
at the nodes. Hyphens (-) represent support values less than 50%
BS/0.90 BYPP. The supported values from all analyses are BS ≥ 70%/
BYPP ≥ 0.95
n = 50), biseriate, broad fusiform to ellipsoid, sometimes
inequilateral, ends acute, initially hyaline, pale brown at
maturity, 1-euseptate, constricted at septum, with guttule in
each cell, smooth-walled with mucilaginous sheath. Asexual
morph: Undetermined.
Culture characters: Colonies on MEA reaching 30 mm
diam. after 4 weeks at 25 °C. Cultures from above, black,
with greyish orange aerial mycelia on the surface, dense,
irregular, umbonate, lobate, velvety, flat, parchment-like;
reverse black, fimbriae.
Material examined: Thailand, Chiang Rai Province, on
dead branches of Clematis subumbellata, 20 April 2017,
C. Phukhamsakda, CMTH 14 (MFLU 17–1478, holotype);
ex-type living culture, MFLUCC 17–2070.
Host: Clematis subumbellata—(This study).
Distribution: Thailand—(This study).
GenBank accession numbers: LSU: MT214570; SSU:
MT226685; ITS: MT310615; tef1: MT394629; rpb2:
MT394692.
Notes: In the phylogenetic analysis, Brunneofusispora
clematidis clustered with the type species B. sinensis with
strong support (100% ML/1.00 BYPP, Fig. 41). Brunneofusispora clematidis can be distinguished from B. sinensis
by its thinner peridium layer (8–17(–28) vs 20–45 μm) and
longer ascospores (Phookamsak et al. 2019). Comparison of
607 nucleotides across the ITS region reveals 74 bp (12.2%)
differences between B. clematidis and B. sinensis. Therefore,
B. clematidis is introduced as a new species (Fig. 42).
Paradictyoarthriniaceae Doilom, Liu & K.D. Hyde
Paradictyoarthriniaceae was established to accommodate
a hyphomycetes genus Paradictyoarthrinium (P. diffractum,
type species) in aquatic and terrestrial habitats (Matsushima
1996; Liu et al. 2015, 2018). Multilocus phylogenetic analyses revealed a rock-inhibiting hyphomycete, Coniosporium
olivaceum Link (≡ Sirodesmium olivaceum CBS 395.59) that
67
is related to Paradictyoarthriniaceae (Ruibal et al. 2009).
Wanasinghe et al. (2018) introduced the first sexual morph,
Xenomassariosphaeria to Paradictyoarthriniaceae. We
introduce an additional species of Xenomassariosphaeria
on Clematis vitalba in Italy (Figs. 2, 43).
Xenomassariosphaeria Jayasiri, Wanas. & K.D. Hyde
Xenomassariosphaeria was introduced for massariosphaeria-like species that formed a close relationship within
Paradictyoarthriniaceae (Tanaka and Harada 2004; Wanasinghe et al. 2018). The genus is characterized by semiimmersed to erumpent, short papillate ascomata, unequal
peridium of thick pseudoparenchymatous cells, cylindrical
to cylindric-clavate, subsessile to short pedicellate asci, with
broad fusiform, hyaline to brown, asymmetric, multiseptate
ascospores (Wanasinghe et al. 2018). Based on a multigene
analysis of LSU, SSU, ITS, tef1 and rpb2 sequence data
(Fig. 2), Xenomassariosphaeria clematidis formed a clade
with the type species X. rosae with strong support (100%
ML/1.00 BYPP, Fig. 2).
Xenomassariosphaeria clematidis Wanas., Phukhams.,
Camporesi & K.D. Hyde, sp. nov.
Index Fungorum number: IF557112; Facesoffungi number: FoF 07350, Fig. 43.
Etymology: Epithet reflects the host Clematis.
Holotype: MFLU 16–0119.
Saprobic on dead stems of Clematis vitalba. Sexual morph: Ascomata 273–283 × 184–208 μm
( x̄ = 275 × 190 μm, n = 5), solitary, scattered, immersed,
with only black spot visible on the host surface, globose,
coriaceous, dark brown to black, rough-walled, ostiolate.
Ostioles centrally located, short papillate, with periphysoids.
Peridium 12–37 μm wide, composed of 5(–7 at apex) layers of dark brown outer layers of textura angularis, inner
layer composed of hyaline gelatinous cells. Hamathecium
composed of numerous, 2.3–3.5 μm wide ( x̄ = 2.7 μm,
n = 50), dense, filiform, septate, branched, cellular pseudoparaphyses. Asci 91–120 × 20–28 μm ( x̄ = 107 × 25 μm,
n = 20), 8-spored, bitunicate, fissitunicate, clavate to
cylindric-clavate, with furcate pedicel, with ocular chamber visible when immature. Ascospores 27–33 × 7–11 μm
( x̄ = 30 × 9 μm, n = 50), biseriate, naviculate, narrow towards
the lower ends, initially hyaline, becoming yellowish to
brown at maturity, 7–8 transversely eusepta, slightly constricted at the septa, third cells from apex usually enlarged,
smooth-walled, guttulate and indentations present, without
mucilaginous sheath. Asexual morph: Undetermined.
Culture characters: Colonies on MEA reaching 20 mm
diam. after 4 weeks at 18 °C. Cultures from above, brown
radiating yellowish towards the edge, dense, circular, flat,
dull, fimbriate, radially furrowed, and slightly covered with
13
68
Fig. 36 Neobyssosphaeria clematidis (MFLU 17–0614, holotype).
a, b Appearance of ascomata on Clematis vitalba. c Vertical section
through ascoma. d Close up of ostiolar canal. e Cellular pseudopara-
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physes. f, g Asci. h–l Ascospores. Scale bars: a = 1 cm, b = 200 µm,
c = 100 µm, d, f–g = 50 µm, e = 5 µm, h–l = 20 µm
Fungal Diversity (2020) 102:1–203
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Fig. 37 Pseudohelminthosporium clematidis (MFLU 17–1494, holotype). a, b Conidiophores on natural substrate (Clematis sikkimensis). c, d Mononematous conidiophores. e, f Conidiogenous cells and
conidia. g–j Conidia (Black basal conidia). k Culture characteristics
on MEA. Scale bars: c, d = 500 μm, e = 100 μm, f–j = 50 μm
white aerial mycelium; reverse black with radiating brown
mycelium.
Material examined: Italy, Forlì-Cesena Province, Poggio
alla Lastra—Bagno di Romagna, dead aerial stems of Clematis vitalba, 19 January 2013, E. Camporesi, IT1019 (MFLU
16–0119, holotype); ex-type living culture, MFLUCC
14–0923.
Hosts: Clematis vitalba—(This study).
Distribution: Italy—(This study).
GenBank accession numbers: LSU: MT214571; ITS:
MT310616; tef1: MT394630.
Notes: In the phylogenetic analyses (Fig. 2), strain
MFLUCC 14–0923 formed a close relationship with the type
species Xenomassariosphaeria rosae (MFLUCC 15–0179)
with strong support (100% ML/1.00 BYPP). Xenomassariosphaeria clematidis shares common features with Xenomassariosphaeria in having immersed ascomata with thick
papilla, cylindrical or clavate asci, and brown, multiseptate
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Nigrograna mackinnonii CBS110022
Nigrograna mackinnonii dH24524
Nigrograna mackinnonii dH24494
Nigrograna mackinnonii CBS674.75
100/1.00
Nigrograna mackinnonii E9303e
Nigrograna mackinnonii E5202H
74/0.99
Nigrograna chromolaenae MFLUCC 17-1437
99/1.00
Nigrograna chromolaenae MFLUCC 17–2079
Nigrograna
marina CY1228
95/1.00
Nigrograna fuscidula MF1
Nigrograna fuscidula MF1a
Nigrograna fuscidula MF3
--/1.00
100/1.00 Nigrograna fuscidula MF7
Nigrograna fuscidula MF8
Nigrograna fuscidula MF9
-Nigrograna carollii CCF4484
Nigrograna antibiotica CCF4378
100/1.00
-Nigrograna antibiotica CCF4998
Nigrograna yasuniana YU.101026
100/0.99 Nigrograna obliqua BW4
91/1.00 Nigrograna obliqua MRP
-84/1.00
Nigrograna obliqua KE
Nigrograna obliqua MFLUCC 14-0945
84/0.99
99/1.00
Nigrograna obliqua MF2
100/0.97 Nigrograna mycophila MF5
-Nigrograna mycophila TDK
100/1.00
Nigrograna mycophila MF6
100/1.00
Nigrograna cangshanensis MFLUCC 15-0253
Nigrograna thymi MFLUCC 14-1096
Nigrograna norvegica CBS141485
100/1.00
Nigrograna norvegica TR8
Nigrograna peruviensis CCF4485
Paradictyoarthrinium diffractum MFLUCC 13-0466 Paradictyoarthriniaceae
(Outgroup)
Paradictyoarthrinium tectonicola MFLUCC 13-0465
Nigrogranaceae
92/0.91
100/1.00
100/1.00
0.01
Fig. 38 The Bayesian 50% majority-rule consensus phylogram based
on combined LSU, ITS, SSU and tef1 sequence data for Nigrogranaceae. The topology and clade stability of the combined gene analyses was compared to the single gene analyses. The tree is rooted
with members of Paradictyoarthriniaceae. Thirty-four strains were
included in the combined gene sequence analyses which comprised
3202 characters (851 characters for LSU, 474 characters for ITS,
1027 characters for SSU, 850 characters for tef1, including gap
regions). The tree from the maximum likelihood analysis had similar topology to the Bayesian analyses. The best scoring RAxML tree
had a final likelihood value of − 8012.999545. The matrix had 487
distinct alignment patterns with 20.65% of undetermined characters
13
and gaps. Estimated base frequencies were as follows; A = 0.244997,
C = 0.242630, G = 0.268950, T = 0.243423; substitution rates
AC = 1.469986, AG = 2.972149, AT = 1.574187, CG = 0.656303,
CT = 11.538665, GT = 1.000000; gamma distribution shape parameter α = 0.62491. In our analysis, GTR + I + G model was used for
each partition in Bayesian posterior analysis. The species determined
in this study are indicated in blue. Bootstrap values (BS) greater than
70% BS (ML, left) and Bayesian posterior probabilities (BYPP, right)
greater than 0.90 are given at the nodes. Hyphens (-) represent support values less than 70% BS/0.90 BYPP. Thick branches represent
significant support values from all analyses (BS ≥ 70%/BYPP ≥ 0.95)
Fungal Diversity (2020) 102:1–203
ascospores (Tanaka and Harada 2004; Wanasinghe et al.
2018). Xenomassariosphaeria clematidis is distinguished
by its unique obovoid ascospores that are narrow towards
the lower end, brown and multi-septate (Fig. 43).
In a BLASTn search of GenBank, the LSU sequence of
X. clematidis (strain MFLUCC 14–0923) is 97.5% similar
to X. roumeguerei (strain CBS 612.86, MH873692). The
ITS region of X. clematidis (strain MFLUCC 14–0923) had
91.5% similarity with X. roumeguerei (strain CBS 612.86,
MH862004). Thus, we introduce a novel species, X. clematidis based on morphological and phylogenetic evidence.
Periconiaceae Nann.
Tanaka et al. (2015) verified Periconiaceae belonged in
Pleosporales based on modern fungal systematics. Although
sequence data is not available for the type species of Periconia (P. lichenoides), the morphological characters of extant
Periconia species correspond to P. lichenoides (Mason and
Ellis 1953; Tanaka et al. 2015). Periconiaceae is characterized by scattered, immersed to erumpent ascomata, pseudoparaphyses, oblong to cylindrical asci, and broadly fusiform,
1-septate, hyaline ascospores. Asexual characters include
synnemata or noosia-like, macronematous, mononematous conidiomata, monoblastic to polyblastic, discrete and
branched conidiogenous cells, and globose to ellipsoidal,
aseptate, catenate, brown conidia (Tanaka et al. 2015).
Periconia Tode
Over 190 species are listed under Periconia (Index Fungorum 2020). Based on phylogenetic analyses of a concatenated dataset of LSU, ITS, and tef1 sequence data coupled
with morphological characters, P. verrucosa is described as
a new species from Clematis viticella (Figs. 44, 45).
Periconia verrucosa Phukhams, Ertz, Gerstmans & K.D.
Hyde, sp. nov.
Index Fungorum number: IF557143; Facesoffungi number: FoF 07296, Fig. 45.
Etymology: The epithet ‘‘verrucosa’’ refers to the surface
of conidia being verrucose.
Holotype: MFLU 17–1516.
Saprobic on dead stems of Clematis viticella. Sexual
morph: Undetermined. Asexual morph: Colonies effuse
on the natural substrate, scattered, hairy, dark brown.
Mycelium partly superficial, semi-immersed, branched,
composed of pale brown, septate hyphae. Conidiophores
170–296 × 10–12 μm ( x̄ = 225 × 11 μm, n = 20), macronematous, mononematous, solitary, gregarious, scattered,
erect, stipes straight or slightly flexuous, with 3–4 short
branches at the apex, cylindrical, smooth, dark brown,
2–4-septate, smooth or verruculose. Conidiogenous cells
11–26 × 6–14 μm ( x̄ = 16 × 7 μm, n = 20), monoblastic or
polyblastic, acropetally proliferating, integrated, terminal,
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oblong, retrogressive, pale brown. Conidia 7–15 μm ( x̄
= 12 μm, n = 50), in branched chains, acrogenous, globose,
aseptate, thick-walled, hyaline when immature, dark brown
to reddish brown at maturity, verrucose, bud scars or disjunctors present at the site of attachment, easily separating.
Culture characters: Colonies on MEA reaching 30 mm
diam. after 2 weeks at 25 °C. Cultures from above, cream
or white, mycelia medium dense, circular, umbonate, papillate, fluffy, slightly radiating outwardly; reverse: cream at
the centre, radiating outwardly.
Material examined: Belgium, Flemish Brabant, Meise
Botanic Garden, Bouchout Domain, on dead stems of
Clematis viticella, 13 June 2017, D. Ertz & C. Gerstmans,
BRCV4 (MFLU 17–1516, holotype); ex-type living culture,
MFLUCC 17–2158.
Hosts: Artemisia sp., Sasa kurilensis, Clematis viticella—
(Tanaka et al. 2015; this study).
Distribution: Belgium, Japan—(Tanaka et al. 2015; this
study).
GenBank accession numbers: LSU: MT214572; SSU:
MT226686; ITS: MT310617; tef1: MT394631.
Notes: Phylogenetic analysis included LSU, ITS and tef1
sequence data with related sequences retrieved from GenBank (Fig. 44). Periconia verrucosa (MFLUCC 17–2158)
formed a strongly supported clade (96% ML/0.99 BYPP)
with three unnamed Periconia strains (KT 1820A, KT 1825
and S-900, Tanaka et al. 2015). These strains do not have
morphological characters for comparison. A comparison of
the ITS nucleotide bases shows that P. verrucosa (MFLUCC
17–2158) has one nucleotide difference with strain S-900
and one nucleotide difference with KT 1820A and KT 1825
in tef1 region. This is regarded as not significant (Jeewon
and Hyde 2016). We introduce P. verrucosa as a new species
to accommodate this clade of Periconia (Fig. 45).
Phaeoseptaceae Boonmee, Thambug. & K.D. Hyde
Phaeoseptaceae was established for lignicolous fungal
lineages on wood. Five genera are included in the family:
Decaisnella, Lignosphaeria, Phaeoseptum (generic type),
Pleopunctum and putative strains of Thyridaria macrostomoides (Abdel-Wahab and Jones 2003; Zhang et al. 2012;
Ariyawansa et al. 2015a; Thambugala et al. 2015; Hyde
et al. 2018a; Liu et al. 2019; Phukhamsakda et al. 2019).
Phaeoseptaceae members have ascomata immersed in
host tissues, short papillate, anastomosed pseudoparaphyses, cylindrical-clavate, long pedicellate asci and broadly
fusoid, single or multi-transverse septa, and hyaline to brown
ascospores (Hyde et al. 2018a). Phaeoseptaceae had pycnidial or sporodochial characters as asexual morph (Liu et al.
2019). We describe a novel species of Pleopunctum recorded
from Clematis in Thailand, based on a multi-gene analysis
(Fig. 46) coupled with comparable morphology (Fig. 47).
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◂Fig. 39 Nigrograna chromolaenae (MFLU 17–1487). a, b Appear-
ance of ascomata on Clematis fulvicoma. c Vertical section through
ascoma. d Section of peridium. e Pseudoparaphyses. f–h Asci. i–n
Ascospores. o Germinated ascospore. p, q Culture characteristics on
MEA. Scale bars: b = 200 µm, c = 100 µm, d–h = 20 µm, i–o = 10 µm
Pleopunctum Liu, K.D. Hyde & J.K. Liu
Liu et al. (2019) introduced the first asexual morph of
Pleopunctum (typified by P. ellipsoideum) in Phaeoseptaceae from decaying wood collected in China. The genus is
characterized by sporodochial conidiomata, macronematous,
mononematous conidiophores, monoblastic conidiogenous
cells and muriform conidia that have a globose basal cell.
The multilocus phylogeny (LSU, SSU, ITS, tef1 and rpb2)
revealed a new species of Pleopunctum from Clematis based
on the morphology and well supported values from multigene phylogeny (Fig. 46).
Pleopunctum clematidis Phukhams., Bhat & K.D. Hyde,
sp. nov.
Index Fungorum number: IF557139; Facesoffungi number: FoF 07301, Fig. 47.
Etymology: The epithet “clematidis” refers to the host
plant, Clematis.
Holotype: MFLU 17–1499.
Saprobic on dead branch of Clematis sikkimensis Sexual morph: Undetermined. Asexual morph: Colonies
on natural substrate forming sporodochial conidiomata,
168–278 μm wide, superficial, scattered, gregarious, oval,
brown, velvety, glistening, orbicular, conidia readily liberated when agitated. Mycelium immersed, septate, smoothwalled, thin-walled, yellowish brown to brown hyphae,
3.5–4.5 μm wide, subicular hyphae short, medium packed.
Conidiophores 6.5–15.5 × 2–5 μm, micronematous, mononematous, cylindrical or truncate, erect, smooth or finely
verruculose, aseptate, unbranched, often reduced to conidiogenous cells, initially hyaline, brown at maturity. Conidiogenous cells 3–8 × 4–9 μm, holoblastic, monoblastic,
integrated, terminal, determinate, cylindrical or slightly
truncate, subspherical or ampulliform, hyaline. Conidia
dimorphic, solitary, smooth-walled; lenticular conidia
16–33 × 15–23 μm ( x̄ = 25 × 20 μm, n = 50), muriform,
smooth, broadly ellipsoidal to oval in front view, yellowish
brown to brown, slightly constricted at the median septa,
inner view composed of one column of 5–7 cells, end cells
subhyaline to pale brown, often carrying remnant of conidiogenous cell at base; cylindrical conidia 15–35 × 6–11 μm
( x̄ = 20 × 8 μm, n = 30), straight or flexuous, septate, constricted at the septa, consisting of one column, 2–3-septate, doliiform, broad clavate, narrow towards apex, apex
rounded, basal cells globose or subglobose, smooth, hyaline.
Culture characters: Colonies on MEA at room temperature (25 °C) reaching 7 cm after 2 weeks. Cultures from
73
above, circular with lobate margin, olive mycelium, white
at the margin, white aerial mycelium, smooth at the surface
and raised; reverse beige, not sporulating.
Material examined: Thailand, Chiang Rai Province, Doi
Tung, on dried stem of Clematis sikkimensis, 2 May 2017,
C. Phukhamsakda & M.V. de Bult, CMTHDT08 (MFLU
17–1499, holotype); ex-type living culture, MFLUCC
17–2091.
Host: Clematis sikkimensis—(This study).
Distribution: Thailand—(This study).
GenBank accession numbers: LSU: MT214573; ITS:
MT310618; tef1: MT394632; rpb2: MT394693.
Notes: Pleopunctum clematidis (MFLUCC 17–2091) is
similar to extant species of Pleopunctum in having sporodochial conidiomata, holoblastic, monoblastic conidiogenous cells and muriform lenticular conidia (Liu et al. 2019).
The strain is distinguishable from other Pleopunctum species by its yellowish brown and smaller lenticular conidia.
Additionally, P. clematidis has dimorphic conidia on the
natural substrate. The dimorphic conidia type have been
documented in Hermatomycetaceae, however, P. clematidis was without subicular hyphae (Tibpromma et al. 2018;
Hyde et al. 2019a, Fig. 47). The multi-gene phylogeny of
LSU, SSU, ITS, tef1 and rpb2 sequence data revealed that
P. clematidis formed a sister lineage to P. ellipsoideum
(MFLUCC 19–0390) and P. pseudoellipsoideum (MFLUCC
19–0391) with strong support (100% MP/100% ML/1.00
BYPP, Fig. 46). In a BLASTn search of GenBank, the closest match to MFLUCC 17–2091 is Lignosphaeria fusispora
(strain MFLUCC 11–0377, KP888646) with 97.78% similarity in the LSU locus, while the closest match with the ITS
sequence is 85.42% similar to KP899140.
Phaeosphaeriaceae Barr
In this family, the taxa are mostly endophytes, pathogens
or saprobes in various habitats (Quaedvlieg et al. 2013;
Phookamsak et al. 2014). We follow the treatment of Hyde
et al. (2020a) and report a novel genus and four new species based on molecular data coupled with morphological
evidence.
Chaetosphaeronema Moesz
Chaetosphaeronema hispidulum is the type species (Moesz 1915; Clements and Shear 1931). Chaetosphaeronema is characterized by immersed pycnidia with
minute ostioles, enteroblastic, phialidic, determinate, discrete conidiogenous cells and cylindrical, hyaline, 1-septate conidia (Sutton 1980; Hyde et al. 2016). Ophiobolus
cirsii (MFLUCC13–0218) is closely related to Chaetosphaeronema as suggested by Zhang et al. (2012). Five species are accepted under Chaetosphaeronema with two strains
having sequence data (Phookamsak et al. 2019). Our phylogenetic analysis (Fig. 48) revealed two new species of
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◂Fig. 40 Nigrograna obliqua (MFLU 16–0190). a Appearance of
ascomata on Clematis vitalba. b Vertical section through ascoma. c
Section of peridium. d Pseudoparaphyses. e–h Asci. i–m Ascospores.
n Germinated ascospore. o, p Culture characteristics on MEA. Scale
bars: b = 100 µm, c–h = 20 µm, i–n = 10 µm
Chaetosphaeronema from Clematis species, with the sexual
morph reported for the genus (Figs. 49, 50).
Chaetosphaeronema clematidicola Phukhams., Ertz, Gerstmans & & K.D. Hyde, sp. nov.
Index Fungorum number: IF557196; Facesoffungi number: FoF 07303, Fig. 49.
Etymology: The epithet reflects the host Clematis.
Holotype: MFLU 17–1508.
Saprobic on Clematis patens. Sexual morph: Ascomata
275–480 × 260–405 μm ( x̄ = 394 × 330 μm, n = 5), scattered or sometimes clustered, gregarious, semi-immersed,
erumpent through host tissue, visible as raised, with only
black shiny ostioles visible on the host surface, uniloculate,
subglobose to compressed, dark brown to black, ostiolate,
papillate. Ostioles 132–248 × 97–120 µm ( x̄ = 174 × 110 μm,
n = 10), central, campanulate, composed of dark brown to
black walled cells, rounded at the apex, with periphyses,
ostioles filled with orange pigment at the pore. Peridium
13–35 µm wide ( x̄ = 23 μm, n= 20), thicker at apex, composed of several layers of pale brown to dark brown cells of
a textura angularis, inner layer lined with subhyaline cells
of textura prismatica. Hamathecium of numerous, 2–3.5 µm
wide ( x̄ = 2.6 μm, n= 50), filamentous, cellular pseudoparaphyses, with distinct septa, embedded in mucilaginous
matrix, anastomosing at the apex. Asci 139–208 × 6–7 µm
( x̄ = 176 × 6 μm, n = 20), 8-spored, bitunicate, broadly filiform to cylindrical, short, with furcate pedicel, apically
rounded, ocular chamber visible when young. Ascospores
134–188 × 1.5–7 µm ( x̄ = 162 × 2.5 μm, n = 30), fasciculate,
in parallel or spiral, scolecosporous, sometimes breaking at
the septa, hyaline to yellowish brown, (18–)20(–23)-septate,
not constricted at the septa, smooth-walled, with minute guttules in each cell. Asexual morph: Undetermined
Culture characters: Colonies growing on MEA reaching
40 mm after 4 weeks at 25 °C. Cultures from above, sparse,
irregular, filamentous, flattened, smooth surface, with fimbriae edge, cream at the margin, white with pale yellowish
at the centre; reverse colony cream at the margin and dark
brown at the centre.
Material examined: Belgium, Flemish Brabant, Meise
Botanic Garden, Bouchout Domain, dead stems of Clematis patens C. Morren & Decne., 13 June 2017, D. Ertz & C.
Gerstmans, BRCP4 (MFLU 17–1508, holotype); ex-type
living culture, MFLUCC 17–2151.
Host: Clematis patens—(This study).
Distribution: Belgium—(This study).
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GenBank accession numbers: LSU: MT214574; SSU:
MT226687; ITS: MT310619; tef1: MT394633; rpb2:
MT394694.
Notes: Chaetosphaeronema clematidicola grouped with
Chaetosphaeronema species (Fig. 48) with strong support
(100% ML/1.00 BYPP). Chaetosphaeronema clematidicola
(MFLUCC 17–2151) is similar to Leptospora (L. rubella)
and Pseudoophiobolus based on their ascospore morphology, but MFLUCC 17–2151 differs in having orange colouration at the ostioles (Shoemaker 1976; Phookamsak et al.
2017). A morphological comparison of Chaetosphaeronema
clematidicola with Ophiobolus cirsii (MFLUCC 13–0218)
showed that it is similar in having fasciculate, cylindrical ascospores (Fig. 49). However, Ophiobolus cirsii has
erumpent ascomata without pigment at the ostioles. Petrak
(1944) mentioned that the ophiobolus-like characteristic is
usually associated with Chaetosphaeronema species. Zhang
et al. (2012) showed that Chaetosphaeronema is genetically
related to Phaeosphaeriaceae. Phookamsak et al. (2014) subsequently confirmed the taxonomy placement and mentioned
that ophiobolus-like species could be the sexual morph of
Chaetosphaeronema. The asexual morph of C. clematidicola
could not be obtained for morphological comparison.
In the phylogenetic analysis, C. clematidicola (MFLUCC
17–2151) grouped with C. clematidis (MFLUCC 17–2147),
another species also on Clematis. A comparison of the ITS
region (including of the 5.8S region) showed three nucleotide differences (588/599—98% with no gaps). A comparison of the tef1 region revealed 12 base pair differences
(842/874—96% with no gaps). Thus, we keep these isolates
as distinct species.
Chaetosphaeronema clematidis Phukhams., Ertz, Gerstmans & K.D. Hyde, sp. nov.
Index Fungorum number: IF557195; Facesoffungi number: FoF 07302, Fig. 50
Etymology: The epithet name “clematidis” refers to the
host substrate.
Holotype: MFLU 17–1504
Saprobic on dead branches of Clematis orientalis.
Sexual morph: Undetermined. Asexual morph: Conidiomata 175–348 × 115–234 μm ( x̄ = 234 × 159 μm, n = 5),
pycnidial, solitary, sometimes aggregated, uniloculate,
immersed, with only black shiny ostioles visible, globose to
compressed, brown to dark brown, coriaceous, thick-walled,
ostiolate, with minute papilla. Ostioles 41–139 × 66–85 μm
( x̄ = 105 × 73 μm, n = 5), central, oblong, lined with periphyses. Conidiomatal wall 12–32 μm wide ( x̄ = 20 μm,
n= 20), comprises dark brown to light brown cells of textura
angularis, lined with a hyaline layer bearing conidiogenous
cells. Conidiophores reduced to conidiogenous cells. Conidiogenous cells 6–14 × 1.5–3 μm ( x̄ = 10 × 2.5 μm, n = 70),
enteroblastic, phialidic, determinate, discrete, cylindrical to
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Occultibambusa bambusae MFLUCC 11-0394
98/-100/0.99
96/1.00
Occultibambusa bambusae MFLUCC 13-0855
Occultibambusa chiangraiensis MFLUCC 16-0380
Occultibambusa pustula MFLUCC 11-0502
100/1.00
Occultibambusa
96/1.00
78/1.00
65/0.99
Occultibambusa jonesii GZCC 16-0117
Occultibambusa maolanensis GZCC 16-0116
Occultibambusa fusispora MFLUCC 11-0127
85/1.00
--
Versicolorisporium triseptatum HHUF 28815
Versicolorisporium
Seriascoma didymospora MFLUCC 11-0179
100/1.00
89/1.00
Seriascoma didymospora MFLUCC 11-0194
Seriascoma
Neooccultibambusa chiangraiensis MFLUCC 12-0559
--
Neooccultibambusa jonesii MFLUCC 16-0643
70/--
Occultibambusaceae
Occultibambusa aquatica MFLUCC 11-0006
Neooccultibambusa
Neooccultibambusa pandanicola KUMCC 17-0179
70/--
Neooccultibambusa thailandensis MFLUCC 16-0274
100/1.00 56/-100/1.00
Brunneofusispora sinensis KUMCC 17-0030 Brunneofusispora
Brunneofusispora clematidis MFLUCC 17-2070
100/1.00
Nigrograna obliqua BW4
Nigrograna obliqua MF2
Nigrogranaceae
100/1.00
Nigrograna mackinnonii E5202H
100/1.00
Ohleria modesta MGC
Ohleria modesta OM
Ohleriaceae
(Outgroup)
0.02
Fig. 41 Bayesian 50% majority-rule consensus phylogram based on
combined LSU, tef1, ITS and SSU sequence data for Occultibambusaceae. The topology and clade stability of the combined gene
analyses was compared to the single gene analyses. The tree is rooted
with members of Ohleriaceae. Twenty-two strains were included in
the combined sequence analyses which comprised 3213 characters (851 characters for LSU, 730 characters for tef1, 607 characters
for ITS, 1025 characters for SSU, including gap regions). The tree
from the maximum likelihood analysis had a similar topology to the
Bayesian analyses. The best scoring RAxML tree had a final likelihood value of − 8012.999545. The matrix had 763 distinct alignment patterns with 26.03% undetermined characters and gaps. Esti-
mated base frequencies were as follows; A = 0.244085, C = 0.246189,
G = 0.275394, T = 0.234331; substitution rates AC = 1.902831,
AG = 2.660713, AT = 1.395112, CG = 1.125830, CT = 7.375307,
GT = 1.000000; gamma distribution shape parameter α = 0.601072.
In our analysis, GTR + I + G model was used for each partition in
Bayesian posterior analysis. The species determined in this study is
indicated in blue. Bootstrap values (BS) greater than 50% BS (ML,
left) and Bayesian posterior probabilities (BYPP, right) greater than
0.90 are given at the nodes. Hyphens (-) represent support values less
than 50% BS/0.90 BYPP. Thick branches represent significant support values from all analyses (BS ≥ 70%/BYPP ≥ 0.95)
subcylindrical, hyaline, canal and collarette minute, smoothwalled, arising from the inner layers of conidioma. Conidia
10–15 × 4–7 μm ( x̄ = 12 × 5 μm, n = 100), cylindrical, hyaline, slightly curved, with 1(–2) guttules in each cell, aseptate to 1 septum, smooth-walled.
Culture characters: Colonies on MEA reaching 20 mm
diam. after 3 weeks at 25 °C. Cultures from above, cream
to pale yellow in the middle, with medium sparse mycelia,
circular, umbonate, papillate fairly fluffy, covered with grey
aerial mycelium, radially furrowed, dark brown pigment diffusing in the agar; reverse: dark brown at the centre, cream
radiating outwardly.
Material examined: Belgium, Flemish Brabant, Meise
Botanic Garden, Bouchout Domain, dead stems of
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Clematis orientalis L., 13 June 2017, D. Ertz & C. Gerstmans, BRCO1 (MFLU 17–1504, holotype); ex-type living
culture, MFLUCC 17–2147.
Host: Clematis orientalis—(This study).
Distribution: Belgium—(This study).
GenBank accession numbers: LSU: MT214575; SSU:
MT226688; ITS: MT310620; tef1: MT394634; rpb2:
MT394695.
Notes: Chaetosphaeronema clematidis is similar to other
Chaetosphaeronema species in having immersed pycnidia,
unilocular, with enteroblastic, phialidic, cylindrical, a channel and collarette, minute conidiogenous cells and cylindrical,
hyaline conidia (Sutton 1980; Hyde et al. 2016, Fig. 50). It is
phylogenetically close to C. achilleae Huang & K.D. Hyde and
C. hispidulum (Corda) Moesz, but differs by the lack of setae
on top of the ostioles and by larger conidia. In the phylogenetic
analysis, the strain formed a close relationship with C. clematidicola (see under C. clematidicola notes for more details).
Dematiopleospora Wanas., Camporesi, E.B.G. Jones & K.D.
Hyde
Dematiopleospora is typified by D. mariae Wanas., Camporesi, E.B.G. Jones & K.D. Hyde. The genus is characterized
by brown setae filling the ostiolar canal, superficial ascomata
and muriform ascospores with pale end cells. There are seven
species in the genus (Wanasinghe et al. 2018, Fig. 51).
Dermatiopleospora mariae Wanas., Camporesi, E.B.G.
Jones & K.D. Hyde, in Wanasinghe, et al., Cryptog. Mycol.
35(2): 110 (2014), new host record
Index Fungorum number: IF550536; Facesoffungi number: FoF 07304, Fig. 52.
Saprobic on dead stems of Clematis vitalba. Sexual morph: Ascomata 131–210 × 156–300 μm
( x̄ = 195 × 240 μm, n = 5), superficial, solitary, scattered, subglobose, flattened at base, dark brown to black, coriaceous,
cupulate when dry, ostiolate. Ostioles 35–42 × 50–66 μm,
papillate, brown, smooth, comprising short, light brown
setae. Peridium 11–24 μm wide ( x̄ = 19 μm, n = 20), thick,
with 7–9 layers, outer layer heavily pigmented, comprising
reddish to dark brown cells of textura angularis, inner layer
composed of hyaline thin-walled cells of textura angularis.
Hamathecium composed of numerous, 2–4 μm wide, filamentous, branched, septate, cellular pseudoparaphyses. Asci
100–125 × 13–19 µm ( x̄ = 111 × 16 µm, n = 40), 8-spored,
bitunicate, fissitunicate, cylindrical to cylindric-clavate,
pedicellate, thick-walled at the apex, with minute ocular
chamber. Ascospores 20–26 × 7–10 µm ( x̄ = 23 × 9 µm,
n = 40), partially overlapping, 1–2-seriate, muriform, ellipsoidal to broad fusiform, slightly curved, ends acute, upper
part wider than the lower part, 5–6-transversely septate, with
1–3 vertical septa, deeply constricted at the central septum,
initially hyaline, becoming yellowish brown at maturity,
77
ends remaining lighter and without a mucilaginous sheath.
Asexual morph: Undetermined.
Culture characters: Colonies on MEA reaching 20 mm
diam. after 4 weeks at 18 °C. Culture from above, medium
dense, circular, margin smooth, white, flat, surface rough;
reverse cream radiating outwardly, dark brown in the middle.
Material examined: Italy, Forlì-Cesena Province, Fiumana di Predappio dead and hanging branches of Clematis
vitalba, 31 January 2013, E. Camporesi, IT 1037 (MFLU
16–0121).
Hosts: Clematis vitalba, Ononis spinosa—(Wanasinghe
et al. 2014; this study).
Distribution: Italy—(Wanasinghe et al. 2014; this study).
GenBank accession numbers: LSU: MT214576; SSU:
MT226689; ITS: MT310621; tef1: MT394635.
Notes: Dermatiopleospora mariae MFLU 16–0121
(Fig. 52) grouped with the type strain of D. mariae
(MFLUCC 13–0612) with moderate statistical support of
60% ML (Fig. 51). Dermatiopleospora mariae (MFLUCC
13–0612) was originally described from Ononis spinosa in
Italy. Morphological characters of our collection are similar to the type strain (Wanasinghe et al. 2014). The ITS
sequence of our collection shows five base pair differences,
however, the tef1 data is identical to D. mariae (MFLUCC
13–0612).
Leptospora Rabenh.
Leptospora is typified by L. rubella and clustered in Phaeosphaeriaceae (Hyde et al. 2016). Morphological characters
of the holotype of Leptospora mentioned that the fungus
stains host tissue red to purple and is red at the apical part
of ostiolar canal (Rabenhorst 1857; Shoemaker 1976; Crous
et al. 2006). Phylogenetic analyses of a combined LSU, SSU,
ITS and tef1 dataset revealed one new species and a new host
record of Leptospora from Clematis species (Figs. 53, 54).
Leptospora clematidis Phukhams., Ertz, Gerstmans & K.D.
Hyde, sp. nov.
Index Fungorum number: IF557197; Facesoffungi number: FoF 02441, Fig. 53.
Etymology: The specific name “clematidis” refers to the
host.
Holotype: MFLU 17–1505.
Saprobic on Clematis patens. Sexual morph: Ascomata 95–245 × 127–247 μm ( x̄ = 159 × 202 μm, n = 5),
dark brown to black, scattered, sometimes semi-immersed,
erumpent through host epidermis, only black shiny dots are
visible on the host surface, uniloculate, subglobose, compressed, dark brown to black, coriaceous, ostiolate. Ostioles
28–40 × 38–87 µm, central, pseudoclypeus, with periphyses
filling the ostiolar canal, pale brown to brown, with light
orange pigment at the pore. Peridium 10–32 µm wide, uniform, composed of 4(–5) layers of cells arranged in textura
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Fungal Diversity (2020) 102:1–203
◂Fig. 42 Brunneofusispora clematidis (MFLU 17–1478, holotype).
a, b Appearance of ascomata on Clematis subumbellata. c Vertical
section through ascoma. d Ostiolar canal. e Section of peridium. f
Pseudoparaphyses. g–i Asci. j–m Ascospores. n Culture characteristics on MEA. Scale bars: b = 200 µm, c = 100 µm, d, e = 50 µm, f,
g–i = 20 µm, j–m = 10 µm
angularis, brown to dark brown, inner layer lined with subhyaline cells of textura angularis. Hamathecium composed
of numerous, 2–4 µm wide, filamentous, cellular pseudoparaphyses, with distinct septa, embedded in mucilaginous
matrix, anastomosing at the apex. Asci 67–96 × 7–11 µm
( x̄ = 78 × 10 μm, n = 30), 8-spored, bitunicate, clavate,
with short, furcate pedicel, apically rounded, ocular chamber present when young. Ascospores 17–33 × 3–6 µm
( x̄ = 26 × 5 μm, n = 30), narrowly turbinate, rounded at apex,
acute at the bottom, hyaline to yellowish brown, 3-septate,
cell above median septa slightly enlarged, slightly constricted at the septa, smooth-walled, with minute guttule in
each cell, polar appendages visible when immature. Asexual
morph: Undetermined.
Culture characters: Colonies on MEA reaching 30 mm
diam. after 4 weeks at 25 °C, from above: sparse, circulate, flattened, surface smooth, with fimbriate edge, cream
at the margin, white with pale yellowish at the centre; colony
below brown at the margin and centre.
Material examined: Belgium, Flemish Brabant, Meise
Botanic Garden, Bouchout Domain, dead stems of Clematis patens, 13 June 2017, D. Ertz & C. Gerstmans, BRCP1
(MFLU 17–1505, holotype); ex-type living culture,
MFLUCC 17–2148, ibid. (MFLU 17–1509, paratypes); exparatype living culture, MFLUCC 17–2152.
Host: Clematis patens—(This study).
Distribution: Belgium—(This study).
GenBank accession numbers: MFLUCC 17–2148:
LSU: MT214577; SSU: MT226690; ITS: MT310622;
tef1: MT394636; rpb2: MT394696. MFLUCC 17–2152:
LSU: MT214578; SSU: MT226691; ITS: MT310623; tef1:
MT394637; rpb2: MT394697.
Notes: Leptospora clematidis shares common characters with Leptospora in having uniloculate ascomata,
ostioles with light orange pigment, a peridium of thinwalled cells arranged in textura angularis, and light yellow ascospores (Hyde et al. 2016). Leptospora clematidis
has similar morphology to L. galii (KUMCC 15–0521),
the strain recorded from Galium sp. in Italy. However, our
new species differs from L. galii in having larger ascomata
that are subglobose with clavate, furcate asci (Hyde et al.
2016, Fig. 53).
Phylogeny (Fig. 48) reveals that L. clematidis forms a
close relationship with L. galii (KUMCC 15–0521) with
strong support (100% ML/1.00 BYPP, Fig. 48). A comparison of the ITS region (including 5.8S region) showed 10
79
nucleotide differences (585/594—98%, with a single gap).
A comparison of the tef1 region revealed 15 base pair differences (822/837—98%, with no gaps). Thus, we describe
L. clematidis as a distinct species.
Leptospora thailandica Phukhams. & K.D. Hyde, in Hyde
et al. Fungal Diversity 80: 100 (2016), new host record
Index Fungorum number: IF552239; Facesoffungi number: FoF 02381, Fig. 54.
Saprobic on dead branches of Clematis subumbellata. Sexual morph: Ascomata 188–229 × 159–179 μm,
immersed to erumpent through host tissue, only black shiny
dots are visible on the host surface, solitary, scattered,
globose to compressed, smooth, brown to dark brown.
Ostioles 120–132 × 91–95 μm ( x̄ = 125 × 92 μm, n = 5),
papillate, oblong, dark brown to light brown, heavily pigmented outer layer, smooth, filled with periphyses, reddish to orange pigment around pore. Peridium 12–18 μm
wide, up to 30 μm at the apex, thin-walled, brown to dark
brown, pseudoparenchymatous cells, composed of 5–7 layers of textura angularis, inner layers composed of hyaline
gelatinous cells. Hamathecium composed of numerous,
2.8–5.5 μm wide (n = 30), broad, branched, filamentous,
septate, cellular pseudoparaphyses. Asci 63–107 × 8–13 μm
( x̄ = 77 × 10 μm, n = 30), 8-spored, bitunicate, cylindrical
to cylindrical-clavate, with short furcate pedicel, apically
rounded, ocular chamber visible when immature. Ascospores
40–77 × 2–5 μm ( x̄ = 58 × 4 μm, n = 40), fasciculate, scolecosporous, tapering towards the ends, hyaline when immature, pale brown at maturity, with minute guttule in each
cell, with (7–)8–17-septate, slightly constricted at the septa,
with 3–5 µm sheath drawn out to form bipolar appendages,
with a pad-like structure at the apices. Asexual morph:
Undetermined.
Culture characters: Colonies on MEA reaching 30 mm
diam. after 4 weeks at 25 °C. Culture from above brown,
sparse, circular, faintly zonate, convex with moderate aerial
mycelium, downy, with slightly irregular at margins; reverse
brown at the edge, light brown at the centre, dense, margin
rough, not pigmented.
Material examined: Thailand, Phayao Province, on dead
branches of Clematis subumbellata, 20 March 2017, C.
Phukhamsakda, CMTH10 (MFLU 17–1474); living culture,
MFLUCC 17–2066.
Hosts: Chromolaena odoratain, Clematis patens, Duranta
sp.—(Hyde et al. 2016; Mapook et al. 2020; this study).
Distribution: Thailand—(Hyde et al. 2016; Mapook et al.
2020; this study).
GenBank accession numbers: LSU: MT214579; SSU:
MT226692; ITS: MT310624; tef1: MT394638.
Notes: Our new collection of Leptospora thailandica
(MFLUCC 17–2066) is morphologically similar to the
type species (MFLUCC 16–0385) which was reported from
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◂Fig. 43 Xenomassariosphaeria clematidis (MFLU 16–0119, holo-
type). a Appearance of ascomata on host surface. b Vertical section
of ascoma. c Pseudoparaphyses. d–f Asci. g–l Ascospores. m Germinated ascospore. n, o Cultures characteristics on MEA. Scale bars:
b = 100 µm, c–f = 20 µm, g–m = 10 µm
Duranta sp. (Hyde et al. 2016). The collection MFLUCC
17–2066 stains the host substrate pinkish red (Fig. 54). Phylogenetic analysis of combined sequence data indicated that
L. thailandica (MFLUCC 17–2066) clusters together with
the type strain and the strain reported from Chromolaena
odorata with strong support (100% ML/1.00 BYPP, Fig. 48).
The ITS sequence of our collection shows two base pair differences from the type strain (from 577 characters, including
gap regions), while the tef1 region is 100% identical.
Longispora Phukhams. & K.D. Hyde, gen. nov.
Index Fungorum number: IF557198; Facesoffungi number: FoF 07305, Fig. 55.
Etymology: The generic epithet referring to the long
ascospores.
Saprobic on herbaceous plant in terrestrial habitats.
Sexual morph: Ascomata solitary, immersed to erumpent,
subglobose to compressed, cupulate when dry, brown to dark
brown, with brown hyphae projecting from the peridium,
ostiolate. Ostioles papillate, oblong, brown to light brown,
heavily pigmented at outer layer, smooth, filled with periphyses, with a reddish to orange pigment around the pore.
Peridium thick-walled, wider at the apex, comprising brownwalled cells of textura angularis. Hamathecium composed of
numerous, branched, filamentous, transversely septate, cellular pseudoparaphyses. Asci 8-spored, bitunicate, cylindricalclavate, with short pedicel, with a visible ocular chamber.
Ascospores fasciculate, scolecosporous, ends rounded, hyaline when immature, pale brown at maturity, multi-septate,
deeply constricted at the swollen cell, slightly constricted
at the other septa, not separating into part spores. Asexual
morph: Undetermined.
Type species: Longispora clematidis Phukhams. & K.D.
Hyde
Notes: Longispora is established as a monotypic genus
with L. clematidis as the type species. The genus is typical of Phaeosphaeriaceae in having compressed globose,
coriaceous, brown to dark brown ascomata, with a reddish to orange pigments around the ostiolar pore, cellular
pseudoparaphyses and fasciculate, scolecosporous, pale
brown and multi-septate ascospores (Rabenhorst 1857;
Crous et al. 2006; Phookamsak et al. 2014). Longispora
has morphological characters similar to Leptospora and the
sexual morph character of Chaetosphaeronema and Neosetophoma (N. camporesii) in having pigmentation in the
ostiolar canal (Hyde et al. 2016, 2020; this study). Moreover,
the fasciculate, scolecosporous, ascospores are common in
81
Phaeosphaeriaceae such as in Ophiobolus, Ophiosphaerella
or Pseudoophiobolus (Phookamsak et al. 2017). Longispora
is distinguishable from other genera having scolecospores in
Phaeosphaeriaceae in its cupulate ascomata with colouration
around the ostiolar pore, and asci that are cylindrical-clavate,
short with a bulbose pedicel, and ascospores that are deeply
constricted at the swollen cell.
Based on the multi-gene phylogenetic analysis (Fig. 48),
MFLU 20–0420 formed a basal lineage to Leptospora
and Populocrescentia with strong support (96% ML/1.00
BYPP, Fig. 48). A BLAST result of the nucleotide sequences
showed 98.84% similarity to Phaeosphaeria oryzae (CBS
110110) in the LSU region, while the ITS region showed
89.24% similarity to Populocrescentia forlicesenensis
(MFLUCC 14–0651).
Longispora clematidis Phukhams. & K.D. Hyde, sp. nov.
Index Fungorum number: IF557199; Facesoffungi number: FoF 07306, Fig. 55.
Etymology: The epithet reflects the host Clematis.
Holotype: MFLU 20–0420.
Saprobic on dead stems of Clematis vitalba. Sexual morph: Ascomata 324–340 × 362–368 μm ( x̄
= 332 × 365 μm, n = 5), solitary, scattered, immersed to
erumpent through host tissue, only black shiny dots are
visible on the host surface, subglobose to compressed,
cupulate when dry, brown to dark brown, with brown
hyphae projecting from the peridium, ostiolate. Ostioles
79–105 × 75–85 μm ( x̄ = 94 × 80 μm, n = 5), papillate,
oblong, brown to light brown, heavily pigmented at outer
layer, smooth, filled with periphyses, with a reddish to
orange colouration around the pore. Peridium 15–30(–35)
μm wide, up to 50 μm at apex, wider at the apex, comprising 6(–8)-layers of brown-walled cells of textura angularis,
outer layer heavily pigmented, with inner region comprising
hyaline cells of textura angularis. Hamathecium composed
of numerous, 2–4 μm wide (n = 50), branched, filamentous,
septate, cellular pseudoparaphyses. Asci 97–157 × 9–15 μm
( x̄ = 128 × 12 μm, n = 35), 8-spored, bitunicate, cylindricalclavate, with short bulbose pedicel, apically rounded, with
visible ocular chamber. Ascospores 93–124 × 2–5 μm ( x̄
= 110 × 4 μm, n = 40), fasciculate, scolecosporous, ends
rounded, hyaline when immature, pale brown at maturity,
with minute guttule in each cell, (17–)19–23-septate, swollen near the septa between the 11th or 13th or 14th cell,
deeply constricted at the swollen cell, slightly contricted at
the other septa, not separating into part spores, indentations
present. Asexual morph: Undetermined.
Culture characters: Colonies on MEA reaching 20 mm
diam. after 4 weeks at 16 °C. Culture slow growing, black,
dense, rhizoid, raised with concave edge, rough, irregular at
the margins; reverse: black, dense, not pigmented.
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Fungal Diversity (2020) 102:1–203
Periconia “verrucosa” KT 1820A
Periconia “verrucosa” S-900
96/0.99
Periconia verrucosa MFLUCC 17–2158
65/1.00
Periconia “verrucosa” KT 1825
97/--
Periconia cookei MFLUCC 17-1679
92/1.00
Periconia cookei MFLUCC 17-1399
Periconia delonicis MFLUCC 17-2584
75/1.00
80/--
Periconia elaeidis MFLUCC 17-0087
--/0.97
Periconia digitata CBS 510.77
99/1.00
Periconia pseudodigitata KT 1195A
99/1.00 Periconia cortaderiae MFLUCC 15-0453
73/-Periconia cortaderiae MFLUCC 15-0457
58/1.00
100/1.00
Periconia cortaderiae MFLUCC 15-0451
Periconia cortaderiae MFLUCC 18-0667
98/--
74/0.91
Periconia cortaderiae MFLUCC 18-0668
Periconia homothallica KT 916
82/1.00
Noosia banksiae CBS 129526
Bambusistroma didymosporum MFLUCC 13-0862
100/1.00
99/0.96
94/1.00
92/1.00
Periconia igniaria CBS 379.86
79/1.00
57/1.00
100/1.00
100/1.00
Periconia prolifica DBOF74
Periconia prolifica DBOF23
Periconia igniaria CBS 845.96
Periconia macrospinosa REF144
66/0.96
Periconia macrospinosa CBS 135663
Periconiaceae
Periconia cyperacearum CPC 32138
Periconia minutissima MFLUCC 15-0245
Periconia variicolor SACCR64
69/0.98
100/1.00
Flavomyces fulophazii CBS 135761
Flavomyces fulophazii CBS 135664
Periconia byssoides MAFF 243872
95/1.00
99/1.00
87/0.99
100/1.00
Periconia byssoides MAFF 243873
Periconia pseudobyssoides MAFF 243868
98/1.00 Periconia pseudobyssoides DLUCC 0850
Periconia thailandica MFLUCC 17-0065
87/1.00
Periconia submersa MFLUCC 16-1098
97/1.00
87/--
Periconia aquatica MFLUCC 16-0912
Massarina eburnea CBS 473.64
Massarina cisti CBS 266.62
58/0.98
Paraphaeosphaeria michotii MFLUCC 13-0349
100/1.00
Didymosphaeria rubi-ulmifolii CBS 100299
100/1.00
100/1.00
Massarinaceae
Didymosphaeriaceae
Lentithecium clioninum KT1220
Lentithecium clioninum KT1149A
Lentithecium fluviatile CBS 122367
Lentitheciaceae
(Outgroup)
0.02
Material examined: Italy, Forlì-Cesena Province, Viale
Salinatore—Forlì City, on dead aerial branch of Clematis
vitalba, 23 February 2015, E. Camporesi, IT2389B (MFLU
20–0420, holotype).
13
Hosts: Clematis vitalba—(This study).
Distribution: Italy—(This study).
GenBank accession numbers: LSU: MT214580; SSU:
MT226693; ITS: MT310625; tef1: MT394639.
Fungal Diversity (2020) 102:1–203
◂Fig. 44 The best scoring RAxML tree with a likelihood value of
− 10209.184183 based on combined LSU, ITS and tef1 sequence
data for Periconiaceae. The tree is rooted with species of Lentitheciaceae. Forty-seven strains were included in the combined gene
sequence analyses which comprised 2974 characters (1320 characters
for LSU, 620 characters for ITS, 1034 characters for tef1, including
gap regions). The topology and clade stability of the combined gene
analyses was compared to the single gene analyses. The tree from
the maximum likelihood analysis had similar topology to Bayesian
50% majority-rule consensus phylogram. The matrix had 865 distinct alignment patterns with 33.43% undetermined characters and
gaps. Estimated base frequencies were as follows; A = 0.229606,
C = 0.269654, G = 0.273255, T = 0.227485; substitution rates
AC = 1.424251, AG = 2.110026, AT = 1.548277, CG = 1.179391,
CT = 7.719698, GT = 1.000000; gamma distribution shape parameter
α = 0.673748. In our analysis, a GTR + I + G model was used for each
partition in the Bayesian posterior analysis. The species determined
in this study is indicated in blue. Bootstrap values (BS) greater than
50% BS (ML, left) and Bayesian posterior probabilities (BYPP, right)
greater than 0.90 are given at the nodes. Hyphens (-) represent support values less than 50% ML/0.90 BYPP. Thick branches represent
significant support values from all analyses (ML ≥ 70%/BYPP ≥ 0.95)
Notes: Longispora clematidis is characterized by having
sessile, cupulate ascomata, reddish orange ostioles, cellular pseudoparaphyses, cylindric-clavate asci with a bulbose
pedicel and filiform ascospores, which are multi-septate and
swollen between the 11th–14th cells (Fig. 55). We introduce L. clematidis based on morphological and phylogenetic
analyses.
Pseudoophiobolus Phookamsak, Wanas., S.K. Huang,
Camporesi & K.D. Hyde
Pseudoophiobolus is typified by P. mathieui (Westend.)
Phookamsak, Wanas., S.K Huang, Camporesi & K.D. Hyde.
The genus was introduced to accommodate an ophioboluslike taxon that is phylogenetically distant from Ophiobolus
sensu stricto (Phookamsak et al. 2017). Pseudoophiobolus is
distinguishable from other ophiobolus-like species in having
semi-immersed to erumpent, papillate ascomata with pseudoparenchymatous cells, arranged in a textura angularis to
textura prismatica, cellular pseudoparaphyses, and fasciculate, scolecosporous multi-septate ascospores with a swollen
cell, that do not split into part spores. A new host record of
P. rosae on Clematis is presented (Fig. 56).
Pseudoophiobolus rosae Phookamsak, Wanas., Phukhams.,
Camporesi & K.D. Hyde in Phookamsak et al. Fungal Diversity 87: 330 (2017), new host record
Index Fungorum number: IF553928; Facesoffungi number: FoF 03805, Fig. 56.
Saprobic on dried stems of Clematis vitalba. Sexual
morph: Ascomata 290–345 × 240–305 μm ( x̄ = 317 × 274 μm,
n = 5), uniloculate, scattered, solitary, semi-immersed to
superficial, ampulliform, globose, cupulate when dried,
dark brown to black, with dark brown, septate mycelium at
the base, ostiolate. Ostioles 117–135 × 100–117 μm, oblong,
83
apex rounded, short papillate, composed of several layers of
dark pseudoparenchymatous cells, with opening by a pore,
filled with hyaline periphyses. Peridium 12–30(–36) μm
wide, slightly thickened, composed of 6(–9) layers of dark
brown cells arranged in textura angularis, pseudoparenchymatous cells, inner layers comprising 2 layers of hyaline cells,
arranged in textura angularis. Hamathecium composed of
dense, 2–4 μm wide ( x̄ = 2.5, n = 50), wide, broad, branched,
filamentous, septate, cellular pseudoparaphyses, anastomosing at the apex, embedded in a hyaline gelatinous matrix.
Asci 64–153 × 9–14 μm ( x̄ = 116 × 12 μm, n = 30), 8-spored,
bitunicate, fissitunicate, cylindric-clavate, with short bulbose pedicel, apically rounded, with well-developed ocular
chamber. Ascospores (40–)75–110 × 3–5 μm ( x̄ = 88 × 4 μm,
n = 40), fasciculate, scolecosporous, curved, pale yellowish
to yellowish, with rounded ends, tapered towards the lower
cells, swollen at the 8th or 10th cell, 17(–22)-septate, not
constricted at the septa, smooth-walled. Asexual morph:
Undetermined.
Culture characters: Colonies on MEA reaching 40 mm
diam. after 4 weeks at 16 °C. Culture medium dense, circular, umbonate, surface smooth, edge erose, thinly hairy,
green at the edge, yellowish to cream at the centre; reverse:
cream at the margin, brown at the centre, not producing pigmentation in agar.
Material examined: Italy, Arezzo Province, Quota—
Poppi City, dead aerial branch of Clematis vitalba, 5 June
2016, E. Camporesi, IT 2983A (MFLU 15–1014); living
culture, MFLUCC 16–1364.
Hosts: Clematis vitalba, Rosa canina—(Phookamsak
et al. 2017; This study).
Distribution: Italy—(Phookamsak et al. 2017; This
study).
GenBank accession numbers: LSU: MT214581; SSU:
MT226694; ITS: MT310626; tef1: MT394640.
Notes: Our collection, MFLUCC 16–1364 formed a
clade with the type strain of Pseudoophiobolus rosae
(MFLUCC 17–1786) with strong support (100% ML/0.99
BYPP, Fig. 48). Our strain is morphologically similar to the
type strain of P. rosae which was reported on Rosa canina
(Phookamsak et al. 2017, Fig. 56). A nucleotide comparison
of the ITS and tef1 regions of MFLUCC 16–1364 and the
type strain are 100% identical. Thus, we report a new host
record of Pseudoophiobolus rosae on Clematis.
Wojnowiciella Crous, Hern.-Restr. & M.J. Wingf.
Wojnowiciella eucalypti is the type species. The genus is
characterized by a papillate, ostiolate conidiomata, ampulliform, phialidic conidiogenous cells, hyaline, aseptate microconidia and brown, multi-septate macroconidia (Wijayawardene et al. 2013; Crous et al. 2015a). The genus has
morphological resemblance to Wojnowicia which was introduced by Saccardo (1892). However, Crous et al. (2015a)
13
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Fungal Diversity (2020) 102:1–203
Fig. 45 Periconia verrucosa (MFLU 17–1516, holotype). a, b Sporodochia on natural substrate. c Close up of sporodochia. d, e Conidiophores.
f–i Conidiogenous cells. j–m Conidia. Scale bars: b = 500 μm, c–e = 250 μm, f, g = 50 μm, h, i = 10 μm, j–m = 5 μm
13
Fungal Diversity (2020) 102:1–203
85
Thyridaria macrostomoides GKM 1033
94/---
Thyridaria macrostomoides GKM 224N
95/68
1.00
Thyridaria macrostomoides GKM 1159
100/100
1.00
Lignosphaeria thailandica MFLUCC 11-0376
100/100
1.00
99/100
1.00
100/100
1.00
Lignosphaeria fusispora MFLUCC 11-0377
Pleopunctum ellipsoideum MFLUCC 19-0390
Pleopunctum pseudoellipsoideum MFLUCC 19-0391
100/100
1.00
100/100
1.00
Pleopunctum clematidis MFLUCC 17-2091
Phaeoseptum aquaticum CBS 123113
100/100
1.00
50/81
0.93
Phaeoseptum terricola MFLUCC 10-0102
Phaeoseptaceae
MP/ML
PP
Phaeoseptum mali MFLUCC 17-2108
100/92
1.00
Decaisnella formosa BCC 25616
Decaisnella formosa BCC 25617
100/100
1.00
Lophiostoma arundinis CBS 621.86
Lophiostoma crenatum CBS 629.86
Lophiostomataceae
(Outgroup)
50.0
Fig. 46 Phylogram generated from maximum parsimony analysis of
Phaeoseptaceae based on combined LSU, SSU, ITS, tef1 and rpb2
sequence data. Related sequences are taken from Liu et al. (2019)
and retrieved from GenBank. Fifteen strains were included in the
analysis of the combined loci and comprised 4077 characters (810
characters for LSU, 1017 characters for SSU, 480 characters for ITS,
895 characters for tef1, 875 characters for rpb2, including gaps). The
tree is rooted with Lophiostoma arundinis (CBS 621.86) and L. crenatum (CBS 629.86) in Lophiostomataceae. Maximum parsimony
analysis of 717 parsimony informative characters resulted in a most
parsimonious tree (CI = 0.797, RI = 0.757, RC = 0.603, HI = 0.203).
The best scoring RAxML tree received a final likelihood value
of − 12889.704989. The matrix had 769 distinct alignment pat-
terns, with 37.49% undetermined characters and gaps. Estimated
base frequencies were: A = 0.241361, C = 0.260527, G = 0.275500,
T = 0.222613; substitution rates AC = 1.309734, AG = 3.237095,
AT = 1.407451, CG = 1.436805, CT = 9.313126, GT = 1.000000;
gamma distribution shape parameter α = 0.742589. In our analysis,
GTR + I + G model was used for each partition in Bayesian posterior
analysis. Bootstrap values (BS) from maximum parsimony (MP, left),
maximum likelihood (ML, right) higher than 50% BS and Bayesian
posterior probabilities (BYPP, below) greater than 0.90 are given
at the nodes. Hyphens (-) represent support values less than 50%
BS/0.90 BYPP. Thick branches represent significant support values
from all analyses (BS ≥ 70%/BYPP ≥ 0.95). The ex-type strains are in
bold and black. The newly generated sequence is in bold and blue
discussed that the type species of Wojnowicia, W. hirta is
compatible with the generic concept of Septoriella. Thus,
Wojnowicia was synonymised as a member of Septoriella.
Our collection associated with Clematis viticella revealed a
novel species W. clematidis from Belgium (Fig. 57).
( x̄ = 236 × 240 μm, n = 5), solitary, gregarious, scattered,
immersed to erumpent through host epidermis, black shiny
dots are visible on the host surface, uniloculate, subglobose,
compressed, dark brown to black, coriaceous, apapillate,
ostiolate. Ostioles central, pseudoclypeate, dark brown.
Peridium 15–45 µm wide, composed of 7(–9) layers of
cells arranged in textura angularis, brown to dark brown,
inner layer lined with sub-hyaline cells of textura angularis. Hamathecium composed of numerous, 2–4 µm wide,
filamentous, cellular pseudoparaphyses, with distinct septa,
embedded in a mucilaginous matrix, anastomosing at the
apex. Asci 97–136 × 12–17 µm ( x̄ = 116 × 15 μm, n = 30),
8-spored, bitunicate, broadly cylindrical to cylindricalclavate, with short, furcate pedicel, apically rounded, with
Wojnowiciella clematidis Phukhams., Ertz, Gerstmans &
K.D. Hyde, sp. nov.
Index Fungorum number: IF557200; Facesoffungi number: FoF 07307, Fig. 58.
Etymology: The epithet refers to the host plant, Clematis.
Holotype: MFLU 17–1517.
Saprobic on dried stems of Clematis viticella.
Sexual morph: Ascomata 228–240 × 223–250 μm,
13
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Fungal Diversity (2020) 102:1–203
Fig. 47 Pleopunctum clematidis (MFLU 17–1499, holotype). a, b
Sporodochia on natural substrate. c Vertical section of sporodochia.
d Subicular hyphae. e–f Cylindrical conidia and lenticular conidia
on host substrate. g–i Cylindrical conidia. j–n Mature lenticular
conidia. o Culture characteristics on MEA. Scale bars = b = 500 μm,
c = 100 μm, d–n = 20 μm
well-developed ocular chamber. Ascospores 24–38 × 4–9 µm
( x̄ = 29 × 7 μm, n = 30), biseriate, partially overlapping, obovoid to sub-fusiform, rounded at apex, acute at the ends,
hyaline to yellowish brown, 4–6-septate, with an oblique or
longitudinal septum in the central 2–3 cells, above median
septum slightly enlarged, constricted at the cell above
median septum, smooth-walled, with minute guttule in
each cell, without a mucilaginous sheath. Asexual morph:
Undetermined.
Culture characters: Colonies on MEA reaching 30 mm
diam. after 4 weeks at 25 °C. Culture from above sparse,
circular, flattened, surface smooth, with fimbriae at the edge,
13
Fungal Diversity (2020) 102:1–203
87
Sclerostagonospora lathyri MFLUCC 14-0958 T
Sclerostagonospora opuntiae CBS 118224
Sclerostagonospora rosae MFLU 18-0115 T
-- Melnikia anthoxanthii MFLUCC 14-1011 T
-Phoma aloes CPC 21549
Camarosporioides phragmitis MFLUCC 13-0365 T
87/1.00
Stagonospora neglecta CBS 343.86
-94/1.00 Parastagonospora minima MFLUCC 13-0376 T
98/1.00
Parastagonospora dactylidis MFLUCC 13-0375 T
94/1.00
Parastagonospora italica MFLUCC 13-0377 T
-Parastagonospora uniseptata MFLUCC 13-0387 T
-Phaeoseptoriella zeae CBS 144614 T
Diederichomyces ficuzzae CBS 128019 T
53/0.99
87/0.99
Didymocyrtis xanthomendozae CBS 129666 T
Neostagonospora caricis CBS 135092 T
67/0.99
Neostagonospora phragmitis MFLUCC 16-0493
65/0.99
Neostagonospora arrhenatheri MFLUCC15-0464 T
Amarenomyces dactylidis MFLUCC 14-0207 T
96/1.00
68/0.99
Amarenomyces dactylidis KUMUCC 18-0154
56/0.99
Dactylidina dactylidis MFLUCC 13-0618 T
--/0.98
Dactylidina shoemakeri MFLUCC 14-0963
---/0.99
Septoriella allojunci MFLUCC 15-0701 T
--/0.99 Septoriella phragmitis CPC 24118 T
Poaceicola forlicesenica MFLUCC 15-0470 T
-99/1.00 Poaceicola arundinis MFLUCC 15-0702 T
--/0.90
Hydeopsis verrucispora MFLUCC 19-0163 T
Vagicola vagans CBS 604.86 T
--/0.99 94/1.00
Amarenographium ammophilae MFLUCC 16-0293 T
-Allophaeosphaeria muriformia MFLUCC 13-0349 T
--/0.93
Neostagonospora elegiae CBS 135101
--/0.99
51/0.99 Neostagonospora artemisiae MFLUCC 17-0693 T
Phaeopoacea phragmiticola CBS 459.84
Phaeopoacea festucae MFLUCC 17-0056 T
91/0.99
Juncaceicola luzulae MFLUCC 13-0780 T
100/1.00
100/1.00
Juncaceicola typharum CBS 296.54
Juncaceicola italica MFLUCC 13-0750 T
--/0.99
--/0.96
Embarria clematidis MFLUCC 14-0976 T
100/1.00
Embarria clematidis MFLUCC 14-0652
--/0.98
Italica luzulae MFLUCC 14-0932 T
100/1.00
Italica achilleae MFLUCC 14-0955 T
Phaeosphaeria
thysanolaenicola MFLUCC 10-0563 T
90/0.99
Phaeosphaeria oryzae CBS 110110 T
100/1.00
Phaeosphaeria chiangraina MFLUCC 13-0231 T
53/0.98
Phaeosphaeria musae MFLUCC 11-0151 T
-99/1.00 Neosetophoma samarorum CBS 138.96 T
99/1.00
Neosetophoma garethjonesii MFLUCC 14-0528 T
Neosetophoma rosaena MFLUCC 17-0768 T
73/0.99 Wojnowiciella spartii MFLUCC 13-0402 T
--/0.96
Wojnowiciella rosicola MFLUCC 15-0128 T
Wojnowiciella lonicerae MFLUCC 13-0737 T
80/0.95
-Wojnowiciella clematidis MFLUCC 17-2159 T
61/0.98
Wojnowiciella
dactylidis MFLUCC 13-0735 T
98/1.00
Wojnowiciella eucalypti CPC 25024
78/-Wojnowiciella viburni MFLUCC 12-0733
Galiicola baoshanensis KUN-HKAS 102234 T
100/1.00
87/1.00
Galiicola pseudophaeosphaeria MFLU 14-0524 T
Murichromolaenicola
chromolaenae MFLUCC 17-1489 T
100/1.00
-Murichromolaenicola chiangraiensis MFLUCC 17-1488 T
Tintelnotia destructans CBS 127737 T
100/1.00
Tintelnotia opuntiae CBS 376.91 T
-Yunnanensis
phragmitis MFLUCC 17-0315
100/1.00
-Yunnanensis phragmitis MFLUCC 17-1361 T
98/1.00
100/1.00 Yunnanensis chromolaenae MFLUCC 17-1487 T
Yunnanensis chromolaenae MFLUCC 17-1486
97/1.00
100/1.00 Pseudostaurosphaeria chromolaenicola MFLUCC 17-1497
Pseudostaurosphaeria chromolaenicola MFLUCC 17-1491 T
100/1.00
Pseudostaurosphaeria chromolaenae MFLUCC 17-1490 T
Parastagonosporella fallopiae CCTU 1151.1
100/1.00
88/1.00
Parastagonosporella fallopiae CBS 135981 T
Paraphoma
radicina CBS 111.79 T
100/1.00
85/0.99
Paraphoma chrysanthemicola CBS 522.66
Xenoseptoria neosaccardoi CBS 120.43 T
100/1.00
95/1.00
Xenoseptoria neosaccardoi CBS 128665
97/1.00
Setomelanomma holmii CBS 110217
Pseudophaeosphaeria rubi MFLUCC 14-0259 T
O hi b l
di CBS 650 86
72/0.96
65/0.90
Fig. 48 The best scoring RAxML tree with a final likelihood value of
− 34209.688899 based on a combined LSU, SSU, ITS and tef1 dataset for Phaeosphaeriaceae. One hundred and fifty-seven strains were
included in the combined genes sequence analyses which comprised
3290 characters (822 characters for LSU, 603 characters for ITS, 991
characters for SSU and 874 characters for tef1, including gap regions).
The tree is rooted with Staurosphaeria. The topology and clade stability of the combined gene analyses was compared to the single gene
analyses. The tree from the maximum likelihood analysis had similar
topology to the Bayesian 50% majority-rule consensus phylogram. The
matrix had 1355 distinct alignment patterns, with 23.28% of undeter-
Phaeosphaeriaceae
--
Wojnowiciella
mined characters and gaps. Estimated base frequencies were as follows;
A = 0.241166, C = 0.264679, G = 0.235933, T = 0.258221; substitution
rates AC = 1.039932, AG = 3.142919, AT = 2.044232, CG = 0.762621,
CT = 4.795230, GT = 1.000000; gamma distribution shape parameter
α = 0.455752. In our analysis, GTR + I + G model was used for each
partition in Bayesian posterior analysis. The species determined in this
study are indicated in blue. Bootstrap values (BS) greater than 50% BS
(ML, left) and Bayesian posterior probabilities (BYPP, right) greater
than 0.90 are given at the nodes. Hyphens (-) represent support values
less than 50% BS/0.90 BYPP. The significant support values from all
analyses are BS ≥ 70%/BYPP ≥ 0.95
13
88
Fungal Diversity (2020) 102:1–203
p
p
Ophiobolus rudis CBS 650.86
Ophiobolus rossicus MFLU 17-1639 T
Ophiobolus italica MFLUCC 14-0526 T
Ophiobolus senecionis MFLUCC 13-0575 T
89/1.00
87/1.00
Ophiobolus disseminans MFLUCC 17-1787 T
97/1.00
Ophiobolus malleolus MFLUCC 15-1077 T
Ophiobolus artemisiae MFLU 15-1966
100/1.00
65/0.99
Ophiobolus artemisiae MFLUCC 14-1156 T
Chaetosphaeronema clematidicola MFLUCC 17-2151 T
97/0.93
Chaetosphaeronema clematidis MFLUCC 17–2147 T
100/1.00
-Chaetosphaeronema hispidulum CBS 216.75 T
Chaetosphaeronema
96/1.00 Chaetosphaeronema achilleae MFLUCC 16-0476 T
54/1.00
Muriphaeosphaeria galatellae MFLUCC 15-0769
100/1.00
Muriphaeosphaeria galatellae MFLUCC 14-0614 T
Ophiobolopsis italica MFLUCC 17-1791 T
100/1.00 Nodulosphaeria senecionis MFLU 15-1297 T
--/0.91
-Nodulosphaeria sambuci MFLUCC 15-0068 T
Nodulosphaeria italica MFLU 16-1359 T
--/1.00 97/1.00
Nodulosphaeria
hirta MFLUCC 13-0867 T
-Nodulosphaeria scabiosae MFLUCC 14-1111 T
100/1.00
--/0.91
87/1.00 Nodulosphaeria aconiti MFLUCC 13-0728 T
Nodulosphaeria spectabilis MFLUCC 14-1112 T
100/0.99 Pseudoophiobolus rosae MFLU 15-1014 T
Pseudoophiobolus rosae MFLUCC 16-1364
89/-100/1.00
Pseudoophiobolus achilleae MFLU 17-0925 T
--/0.97
Pseudoophiobolus
Pseudoophiobolus galii MFLUCC 17-2257 T
86/1.00100/1.00
Pseudoophiobolus mathieui MFLUCC 17-1785 T
Pseudoophiobolus italicus MFLUCC 17-2255 T
Paraophiobolus plantaginis MFLUCC 17-0245 T
100/1.00
52/0.97
Paraophiobolus arundinis MFLUCC 17-1789 T
67/1.00
Dematiopleospora mariae MFLUCC 13-0612 T
89/1.00
Dermatiopleospora mariae MFLU 16-0121
100/1.00
Dematiopleospora
Dematiopleospora fusiformis MFLU 15-2133 T
Dematiopleospora cirsii MFLUCC 15-0615 T
100/1.00
--/0.99
100/1.00 Hydeomyces desertipleosporoides SQUCC 15259 T
100/1.00
Hydeomyces desertipleosporoides SQUCC 15260
Hydeomyces pinicola GZCC 19-0003 T
53/0.98 Dlhawksworthia lonicera MFLUCC 14-0955 T
76/1.00 100/1.00 Dlhawksworthia alliariae MFLUCC 13-0070 T
Dlhawksworthia clematidicola MFLUCC 14-0910 T
70/0.99
Pseudoophiosphaerella huishuiensis MFLUCC 17-1471
100/1.00
77/1.00
Pseudoophiosphaerella huishuiensis MLUCC 19-0164 T
100/1.00 Neoophiobolus chromolaenae MFLUCC 17-1467
-Neoophiobolus chromolaenae MFLUCC 17-1449 T
89/1.00
Loratospora aestuarii JK 5535B
Paraleptospora chromolaenicola MFLUCC 17-1450 T
100/1.00
Paraleptospora chromolaenae MFLUCC 17-1481 T
99/1.00 Leptospora thailandica MFLUCC 17-1468
100/1.00 Leptospora thailandica MFLUCC 17-2066
97/0.99
Leptospora thailandica MFLUCC 16-0385 T
77/1.00
Leptospora phraeana MFLUCC 17-1478
90/1.00
Leptospora chromolaenae MFLUCC 17-1460 T
99/1.00
Leptospora
Leptospora hydei GZCC 19-0004
-Leptospora rubella CPC 11006 T
78/-Leptospora clematidis MFLUCC 17-2148 T
53/-100/1.00
Leptospora clematidis MFLUCC 17-2152
-Leptospora galii KUMCC 15-0521
100/1.00 Populocrescentia rosacea MFLU 17-0128 T
96/1.00
Populocrescentia ammophilae MFLUCC 17-0665 T
99/1.00
Populocrescentia forlicesenensis MFLUCC 14-0651 T
100/1.00
-Longispora clematidis MFLU 20–0420 T
Longispora
Phaeosphaeriopsis triseptata MFLUCC 13-0271 T
100/1.00
Phaeosphaeriopsis glaucopunctata MFLUCC 13-0265 T
95/1.00
-Phaeosphaeriopsis dracaenicola MFLUCC 11-0157 T
Jeremyomyces labinae CBS 144617 T
100/1.00
Acericola italica MFLUCC 13-0609 T
100/1.00 Ophiosphaerella agrostidis IGM35
Ophiosphaerella agrostidis MFLUCC 11-0152
100/1.00
Ophiosphaerella agrostidis MFLUCC 12-0007
100/1.00
55/-Ophiosphaerella aquatica MFLUCC 14-0033 T
Equiseticola fusispora MFLUCC 14-0522
100/1.00 Neostagonosporella sichuanensis MFLUCC 18-1228 T
100/1.00
Neostagonosporella sichuanensis MFLUCC 18-1223
Kwanghwaensis miscanthi FU31017
54/1.00
Setophoma terrestris CBS 335.29 T
Setophoma sacchari MFLUCC 11-0154
--/1.00
Neosulcatispora agaves CPC 26407 T
70/0.99
--/0.94
84/0.99
Vrystaatia aloeicola CBS 135107 T
-Bhatiellae rosae MFLUCC 17-0664 T
Piniphoma wesendahlina CBS 145032 T
Staurosphaeria rhamnicola MFLUCC 17-0813 T
100/1.00
Staurosphaeria rhamnicola MFLUCC 17-0814
Phaeosphaeriaceae
100/1.00
80/1.00
80/1.00
( Outgroup )
0.03
Fig. 48 (continued)
cream at the margin, white at the centre, colony from below
brown.
Material examined: Belgium, Flemish Brabant, Meise
Botanic Garden, Bouchout Domain, dead stems of Clematis viticella, 13 June 2017, D. Ertz & C. Gerstmans,
13
BRCV5 (MFLU 17–1517, holotype); ex-type living culture,
MFLUCC 17–2159.
Host: Clematis viticella—(This study).
Distribution: Belgium—(This study).
Fungal Diversity (2020) 102:1–203
89
Fig. 49 Chaetosphaeronema clematidicola (MFLU 17–1508, holotype). a Appearance of ascomata on Clematis patens. b Close up of
ascoma on host substrate. c Vertical section through ascoma. d Osti-
olar canal. e Section of peridium. f Pseudoparaphyses. g–h Asci. i–j
Ascospores (j Ascospore in 10% India ink). Scale bars: b = 500 µm,
c = 200 µm, d–h = 100 µm, i–j = 50 µm
GenBank accession numbers: LSU: MT214582; SSU:
MT226695; ITS: MT310627; tef1: MT394641; rpb2:
MT394698.
Notes: Wojnowiciella clematidis (strain MFLUCC
17–2159) is introduced as a novel species based on its
distinctive morphology and phylogeny (Figs. 57, 58). The
species is similar to other Wojnowiciella species in having
immersed to erumpent, dark brown to black, coriaceous,
ostiolate, apapillate ascomata, and hyaline to yellowish brown ascospores (Crous et al. 2015a). Wojnowiciella
13
90
Fungal Diversity (2020) 102:1–203
Fig. 50 Chaetosphaeronema clematidis (MFLU 17–1504, holotype). a Appearance of conidiomata on Clematis orientalis. b Close
up of conidioma on host substrate. c Vertical section through conidioma. d Ostiolar canal. e Section of conidiomatal wall. f–i Conidiog-
enous cells and conidia (f, g conidiogenous cells in cotton blue). j–l
Conidia. Scale bars: b = 500 µm, c = 200 µm, d = 50 µm, e = 20 µm,
f–l = 10 µm
clematidis has similar characters to W. italica (MFLUCC
13–0447), but differs in having cylindrical to cylindricalclavate asci and obovoid to sub-fusiform ascospore (Hyde
et al. 2016). Phylogeny (Fig. 57) revealed that W. clematidis (MFLUCC 17–2159) formed a close relationship
with W. dactylidis (CBS 145077) with good support (77%
MP/84% ML/1.00 BYPP). Wojnowiciella dactylidis strain
CBS 145077 formed a separate clade from the type species (W. dactylidis MFLUCC 13–0735), however the strain
CBS 145077, which was isolated from Dypsis sp. only forms
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60/-79/-89/1.00
Dermatiopleospora mariae MFLU 16-0121
Dematiopleospora fusiformis MFLU 15-2133 T
Dematiopleospora donetzica MFLU 15-1975
99/1.00
Dematiopleospora donetzica MFLU 15-2199 T
--/1.00
50/--
and concatenated phylogenetic analysis of LSU, ITS, SSU
and gadph sequence data (Fig. 59).
Dermatiopleospora mariae MFLUCC 15-0612 T
Dematiopleospora cirsii MFLUCC 13-0615 T
Dematiopleospora
ML/PP
91
Dematiopleospora rosicola MFLU 16-0232 T
50/-Dematiopleospora salsolae TASM 6123 T
Dlhawksworthia alliariae MFLUCC 13-0070 T
0.005
Fig. 51 The Bayesian 50% majority-rule consensus phylogram based
on combined LSU, SSU, ITS and tef1 sequence data for Dermatiopleospora species. Nine strains were included in the combined analyses which comprised 3407 characters (884 characters for LSU, 1033
characters for SSU, 617 characters for ITS and 873 characters for tef1,
including gap regions). The tree is rooted with Dlhawksworthia alliariae (MFLUCC 13-0070). Single gene analyses were also performed
to compare the topology and clade stability with combined gene analyses. Tree topology generated under the maximum likelihood analysis was similar to Bayesian analyses. The best scoring RAxML tree
was obtained with a final likelihood value of − 5814.019143. The
matrix had 162 distinct alignment patterns, with 24.34% of undetermined characters and gaps. Estimated base frequencies were as follows; A = 0.252414, C = 0.229288, G = 0.264353, T = 0.253945;
substitution rates AC = 0.248381, AG = 2.072002, AT = 1.746797,
CG = 1.084349, CT = 5.244757, GT = 1.000000; gamma distribution
shape parameter α = 0.904425. In our analysis, GTR + I + G model
was used for each partition in the Bayesian posterior analysis. The
species determined in this study is indicated in blue. Bootstrap values (BS) from maximum likelihood (ML, left) of more than 50% BS
and Bayesian posterior probabilities (BYPP, right) greater than 0.90
are given at the nodes. Hyphens (-) represent support values less than
50% BS/0.90 BYPP. The significantly supported values from all analyses are BS ≥ 70%/BYPP ≥ 0.95
microconidia in culture (Crous et al. 2019). A comparison
of nucleotides between W. clematidis (MFLUCC 17–2159)
and W. dactylidis (CBS 145077) showed that the ITS region
(including the 5.8S region) has a single nucleotide difference (550/561—98%, including gaps). The comparison of
the tef1 region revealed three base pair differences (869/440,
no gaps). Based on current evidence and the lack of asexual
morph character from this study, these species are considered as distinct.
Pleosporaceae Nitschke
The family was revisited by Ariyawansa et al. (2015c),
with additional taxa and resegregation in Wanasinghe et al.
(2017). We introduce a new species and new host records of
Stemphylium from Clematis vitalba, based on a morphology
Stemphylium Wallr.
Stemphylium is a dematiaceous hyphomycete genus and
the asexual morph has been recorded in Pleospora allies
(Ariyawansa et al. 2015c; Woudenberg et al. 2017). Based
on morphological studies, multi-gene phylogeny analyses,
and ecological evidence, the use of Stemphylium over Pleospora has been recommended (Köhl et al. 2009; McNeill
et al. 2012; Rossman et al. 2015; Woudenberg et al. 2017;
Wijayawardene et al. 2018). Stemphylium is typified with
S. botryosum. Phylogenetic placement of S. botryosum is
verified by Woudenberg et al. (2017). Based on the phylogenetic analysis of the combined LSU, ITS and gapdh dataset
(Fig. 59), we report a new host record for S. vesicarium and
describe a new species S. clematidis from Clematis vitalba
(Figs. 60, 61).
Stemphylium clematidis Wanas., Camporesi & K.D. Hyde,
sp. nov.
Index Fungorum number: IF557305; Facesoffungi number: FoF 07327, Fig. 60.
Etymology: Refers to the name of the host.
Holotype: MFLU 16–0176.
Saprobic on dead stem of Clematis vitalba. Sexual morph: Ascomata 190–230 × 290–320 µm
( x̄ = 220 × 303 µm, n = 20), solitary, scattered, uniloculate,
under epidermal layer of host to superficial, compressed
globose to subglobose, coriaceous, black, ostiolate. Ostioles central, papillate, with variable walls, opening by a
pore, filled with hyaline periphyses. Peridium 30–43 μm
wide ( x̄ = 36 μm, n= 20), uniform, composed of 7 layers
of textura angularis, black, heavy pigment at outer layer,
cells towards the inside lighter, inner layer thin, hyaline
gelatinous. Hamathecium composed of numerous, dense,
3–4 µm wide, filamentous, branched, septate, cellular pseudoparaphyses, situated between and above the asci embedded in a gelatinous matrix. Asci 127–172 × 29–38 µm
( x̄ = 150 × 35 µm, n = 30), 8-spored, bitunicate, fissitunicate, clavate to broad cylindrical-clavate, with short, with
furcate pedicel, apically rounded, with an ocular chamber.
Ascospores 24–37 × 12–18 µm ( x̄ = 32 × 13 µm, n = 50),
biseriate or partially overlapping, broad-fusiform, tapering
towards the rounded ends, dictyosporous, 1(–2)-longitudinal euseptate, 7–9-transverse euseptate, constricted at the
septa, strongly constricted at the median septum, cells above
median septum wider than below, yellowish, with 8–10 µm
wide mucilaginous drawn out sheath. Asexual morph:
Undetermined.
Culture characters: Colonies on MEA reaching 30 mm
diam. after 4 weeks at 16 °C. Culture from above cream,
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◂Fig. 52 Dermatiopleospora mariae (MFLU 16–0121). a Appearance
of ascomata on Clematis vitalba. b Close up of ascoma on host substrate. c Vertical section through ascoma. d Ostiolar canal. e Section
of peridium. f Pseudoparaphyses. g–j Asci. k–p Ascospores. Scale
bars: a = 500 µm, b = 100 µm, c–j = 20 µm, k–p = 10 µm
orange aerial mycelium, dense, fluffy at the edge, umbonate,
rough, lobate; reverse cream radiating white outwardly.
Material examined: Italy, Forli-Cesena Province, Marsignano-Predappio, dead stems of Clematis vitalba, 31 March
2015, E. Camporesi, IT 1537 (MFLU 16–0176, holotype);
ex-type living culture, MFLUCC 14–0937.
Host: Clematis vitalba—(This study).
Distribution: Italy—(This study).
GenBank accession numbers: LSU: MT214583; SSU:
MT226696; ITS: MT310628; gapdh: MT394626.
Notes: Stemphylium clematidis has a pleospora-like sexual morph. The isolate formed a sister clade with S. majusculum, however the species is distinguishable by somewhat
larger ascospores (Ramsey 1934; Simmons 1969, Fig. 60).
The asexual morph was not obtained from culture. A pairwise comparison of the ITS sequence showed six nucleotide
differences out of 555 nucleotides in the ITS regions (including gaps), while gapdh showed five nucleotide differences
out of 595 nucleotides. The new strain is introduced as a
new species of Stemphylium based on guidelines proposed
by Jeewon and Hyde (2016).
Stemphylium rosae (Wanas. et al.) Phukhams. & K.D.
Hyde, comb. nov.
Index Fungorum number: IF557603; Facesoffungi number: FoF 0404
Basionym: Pleospora rosae Wanas., Camporesi, E.B.G.
Jones & K.D. Hyde, in Wanasinghe et al., Fungal Diversity
[153] (2018).
Notes: Based on analyses of combined LSU, ITS and
gadph sequence data for Stemphylium (Fig. 60), two isolates of Pleospora rosae and P. rosae-caninae clustered with
Stemphylium sensu stricto. These two strains are compatible
with the Stemphylium concept. Thus, we synonymize Pleospora rosae under Stemphylium rosae and Pleospora rosaecaninae under Stemphylium rosae-caninae. The nomenclature change is based on one fungus = one name protocol.
Host: Rosa canina—(Wanasinghe et al. 2018).
Distribution: Italy—(Wanasinghe et al. 2018).
Stemphylium rosae-caninae (Wanas. et al.) Phukhams. &
K.D. Hyde, comb. nov.
Index Fungorum number: IF557604; Facesoffungi number: FoF 04047
Basionym: Pleospora rosae-caninae Wanas., Camporesi,
E.B.G. Jones & K.D. Hyde, in Wanasinghe et al., Fungal
Diversity [157] (2018).
93
Notes: See notes under Stemphylium rosae-caninae.
Host: Rosa canina—(Wanasinghe et al. 2018).
Distribution: Italy—(Wanasinghe et al. 2018).
Stemphylium vesicarium (Wallr.) E.G. Simmons, Mycologia 61(1): 9 (1969), new host record
Index Fungorum number: IF339660; Facesoffungi number: FoF 04472, Figs. 61, 62.
Saprobic on dead stem of Clematis vitalba. Sexual morph: Ascomata 170–257 × 148–250 μm
( x̄ = 220 × 205 µm, n = 10), solitary, scattered, uniloculate,
under epidermal layer of host, to superficial, globose to
compressed, coriaceous, black, ostiolate. Ostioles central,
papillate, with variable walls, opening by a pore, filled with
hyaline periphyses. Peridium 25–49 μm wide ( x̄ = 40 μm,
n= 20), thick, uniform, composed of 7–12 layers of textura
angularis, black, heavy pigmented at the outer layer, cells
toward the inside cells of lighter, inner layer composed of
thin hyaline gelatinous layer. Hamathecium composed of
numerous, dense, 1.6–2.25 µm wide ( x̄ = 1.8 μm, n= 20),
filamentous, branched, septate, cellular pseudoparaphyses,
situated between and above the asci embedded in a gelatinous matrix. Asci 103–138 × 16–26 µm ( x̄ = 118 × 23 µm,
n = 30), 8-spored, bitunicate, fissitunicate, oblong to cylindrical-clavate with short, furcate pedicel, apically rounded,
with an ocular chamber. Ascospores 20–30 × 6–12 µm
( x̄ = 26 × 10 µm, n = 50), biseriate or partially overlapping,
muriform, tapering towards the ends, ends rounded, dictyosporous, 1(–2)-longitudinal euseptate, 4–10-transversely
euseptate, constricted at the septa, strongly constricted at
the median septum, yellowish, with 5–12 µm wide drawn
out mucilaginous sheath. Asexual morph: Colonies effuse
on the surface of culture, scattered, hairy, and fluffy. Mycelium white, with 2–5 µm wide, effuse, hyaline, septate.
Conidiophores 10–57 × 3–5 μm ( x̄ = 35 × 4 μm, n = 20),
macronematous, mononematous, simple, branched, stipes
straight or flexuous, cylindrical, erect, septate, smooth, dark
brown to brown, 3(–6)-septate, with 1–2 primary branches,
irregularly branched at the upper parts, brown, smooth.
Conidiogenous cells 5–12 × 4–10 μm ( x̄ = 9 × 7 μm, n = 20),
monotretic, integrated or terminal, on conidiophores, doliiform to oblong, pale brown. Conidia 14–31 × 13–32 μm
( x̄ = 21 × 21 μm, n = 30), in branched chains, acrogenous,
folded, muriform, dictyosporous, 1(–2)-transversely euseptate, 1–3-horizontal euseptate, constricted at septa, dark
brown to brown, bud scars or disjunctors present at the site
of attachment, easily separated.
Culture characters: Colonies on MEA reaching 30 mm
diam. after 4 weeks at 16 °C. Cultures from above, dark
green, fluffy, with dark green aerial mycelium, dense, fluffy
at the edge, umbonate, rough, lobate, faintly zonate; reverse:
dark green, radiating.
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◂Fig. 53 Leptospora clematidis (MFLU 17–1505, holotype). a
Appearance of ascomata on Clematis patens. b Vertical section
through ascoma. c Ostiolar canal. d Section of peridium. e Pseudoparaphyses. f–h Asci. i–l Ascospores. Scale bars: b = 100 µm, c, f–
h = 20 µm, d, e = 50 µm, i–l = 10 µm
Material examined: Italy, Forli-Cesena Province, Viale
Salinatore—Forlì, dead stems of Clematis vitalba, 23 February 2015, E. Camporesi, IT 2983M (MFLU 16–1109); living
culture, MFLUCC 16–0998.
Hosts: Abies sp., Allium cepa, A. sativum, Asparagus
officinalis, Brassica nigra, Brassica pekinensis, Cirsium sp.,
Citrus sp., Clematis vitalba, Cremanthodium discoideum,
Cremanthodium discoideum, Dahlia pinnata, Dianthus caryophyllus, Lathyrus odoratus, Leucadendron sp., Linum usitatissimum, Lunaria annua, Lunaria rediviva, Malus domestica, Malus sieversii, Mangifera indica, Medicago sativa,
Phaseolus vulgaris, Pisum sativum, Populus tomentosa,
Pyrus sinkiangensis, Sedum spectabile, Solanum lycopersicum, Tamaric sp., Trigonella foenum-graecum—(Simmons
1969; Simonyan 1981; Câmara et al. 2002; Köhl et al. 2009;
Arzanlou et al. 2012; Ariyawansa et al. 2015c; Woudenberg
et al. 2017; Farr and Rossman 2020; this study).
Distribution: Australia, Canada, China, Denmark, Germany, India, Italy, Netherlands, New Zealand, Portugal,
South Africa, Tunisia, UK (England), USA (California)—
(Simmons 1969; Simonyan 1981; Câmara et al. 2002; Köhl
et al. 2009; Arzanlou et al. 2012; Ariyawansa et al. 2015c;
Woudenberg et al. 2017; Farr and Rossman 2020; this
study).
GenBank accession numbers: LSU: MT214584; SSU:
MT226697; ITS: MT310629; tef1: MT394642.
Notes: Stemphylium vesicarium (= Pleospora herbarum)
is a plant pathogen distributed on a range of wild and cultivated hosts (Köhl et al. 2009; Arzanlou et al. 2012; Woudenberg et al. 2017). Phylogenetic analysis of combined LSU,
ITS and gapdh regions show that our isolate clusters with
S. vesicarium (Fig. 59). The morphological comparison of
our isolate (MFLUCC 16–0998) with S. vesicarium (CPC
29939) which was reported in Woudenberg et al. (2017)
showed that our collection has 4–10-transverse eusepta
ascospores (Figs. 61, 62). Our collection is similar to the
type strian of S. vesicarium (STR) which was described by
Simmons (1969). A pairwise comparison of the ITS regions
show that our isolate is 100% identical to the type strain of
S. vesicarium (CBS 192.86). This is the first report of S.
vesicarium on Clematis.
Pseudoberkleasmiaceae Phukhams. & K.D. Hyde
Pseudoberkleasmiaceae was introduced to accommodate
a berkleasmium-like species that has phylogenetic stability in Pleosporales (Hyde et al. 2019a). The family comprises Pseudoberkleasmium acacia, Pseudoberkleasmium
95
chiangmaiense and P. pandanicola (the generic type). We
describe the first record of P. chiangmaiense on Clematis
from Thailand (Figs. 2, 63).
Pseudoberkleasmium Tibpromma & K.D. Hyde
The monotypic genus Pseudoberkleasmium was introduced with P. pandanicola Tibpromma & K.D. Hyde as
the type species. Pseudoberkleasmium is characterized by
hyaline, subglobose conidiogenous cells and acrogenous,
broadly ellipsoidal to obovoid, muriform, brown or olivaceous green, and guttulate conidia. There is no sexual morph
report in this family. A collection on Clematis sikkimensis
reveals an additional strain of Pseudoberkleasmium chiangmaiense according to phylogenetic and morphological
evidence (Figs. 2, 63).
Pseudoberkleasmium chiangmaiense Y.Z. Lu & K.D.
Hyde, in Hyde et al., Fungal Diver (2019), new host record
Index Fungorum number: IF555595; Facesoffungi number: FoF 05310, Fig. 63.
Saprobic on dead stems of Clematis sikkimensis. Sexual
morph: Undetermined. Asexual morph: Hyphomycetous,
colonies on natural substrate forming sporodochial conidiomata, 109–460 μm wide, superficial, scattered, gregarious, oval, brown, velvety, glistening, orbicular, conidia
readily liberated when agitated. Mycelium immersed in the
substrate, composed of septate, branched, smooth, hyaline to pale brown, 2.5 μm wide hyphae. Conidiophores
10–25 × 2–5 μm, micronematous, mononematous, cylindrical or truncate, erect, hyaline, smooth-walled. Conidiogenous cells 9–15 × 6–15 μm ( x̄ = 12 × 11 μm, n = 20),
holoblastic, monoblastic, integrated, terminal, determinate,
subglobose, cylindrical or slightly truncate, guttulate, hyaline. Conidia 24–32 × 13–18 μm ( x̄ = 27 × 16 μm, n = 50),
acrogenous, solitary, broadly ellipsoidal to obovoid, flattened, dictyosporous, muriform, apex rounded, basal cells
globose or subglobose, in side view composed of one column of 4–6 cells, guttulate, smooth-walled, brown, usually
with conidiogenous cell attached, bud scars or disjunctors
present at the site of attachment.
Culture characters: Colonies on MEA reaching 30 mm
diam. after 4 weeks at 25 °C. Cultures from above black,
dense, circular, umbonate, papillate, fluffy, slightly radiating, wrinkled folded; reverse black, lifting media up in the
centre, wrinkled.
Material examined: Thailand, Chiang Rai Province, Doi
Tung, on dried stem of Clematis sikkimensis, 2 May 2017,
C. Phukhamsakda & M.V. de Bult, CMTHDT05 (MFLU
17–1496); living culture, MFLUCC 17–2088.
Hosts: Clematis fulvicoma, Undetermined decaying
wood—(Hyde et al. 2019a; this study).
Distribution: Thailand—(Hyde et al. 2019a; this study).
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96
Fig. 54 Leptospora thailandica (MFLU 17–1474). a Appearance
of ascomata on Clematis subumbellata. b Vertical section through
ascoma. c Ostiolar canal. d Section of peridium. e Pseudoparaphyses.
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f–h Asci. i, j Ascospores. k, l Culture characteristics on MEA. Scale
bars: b = 100 µm, c–h = 50 µm, i, j = 20 µm
Fungal Diversity (2020) 102:1–203
Fig. 55 Longispora clematidis (MFLU 20–0420, holotype). a, b
Appearance of ascomata on Clematis vitalba. c Vertical section
through ascoma. d Ostiolar canal. e Section of peridium. f Pseudo-
97
paraphyses. g–i Asci. j–m Ascospores (m Ascospore in cotton blue).
Scale bars: c = 100 µm, d–f, j–m = 20 µm, g–i = 50 µm
13
98
Fig. 56 Pseudoophiobolus rosae (MFLU 15–1014). a Appearance of
ascomata on Clematis vitalba. b Vertical section through ascoma. c
Ostiolar canal. d Section of peridium. e Pseudoparaphyses. f–g Asci.
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h–j Ascospores (j in cotton blue). k Germinated ascospore. l, m Culture characteristics on MEA. Scale bars: a = 500 µm, b = 200 µm,
c–k = 50 µm
Fungal Diversity (2020) 102:1–203
99
98/100
1.00
MP/ML
BYPP
Wojnowiciella kunmingensis KUMCC 18-0159 T
Wojnowiciella kunmingensis KUMCC 18-0160
-55/70
1.00
Wojnowiciella dactylidis MFLU 16-1366
77/84
1.00
Wojnowiciella dactylidis CBS 145077
--
Wojnowiciella clematidis MFLUCC 17-2159 T
Wojnowiciella cissampeli CPC 27455 T
--/66
-Wojnowiciella rosicola MFLUCC 15-0128
--/90
---/89
Wojnowiciella italica MFLU 14-0732 T
0.90
--/52
-Wojnowiciella spartii MFLUCC 13-0402 T
70/70
1.00
84/87
1.00
100/100
1.00
T
Wojnowiciella
Wojnowiciella dactylidis MFLUCC 13-0735 T
58/67
0.98
Wojnowiciella lonicerae MFLUCC 13-0737 T
Wojnowiciella eucalypti CBS 139904 T
Wojnowiciella leptocarpi CBS 115684 T
Wojnowiciella viburni MFLUCC 12-0733 T
100/100
1.00
Galiicola baoshanensis KUN-HKAS 102234 T
Galiicola pseudophaeosphaeria MFLUCC 14-0524 T
0.005
Fig. 57 Phylogram obtained from maximum likelihood based on
combined LSU, SSU, ITS, tef1 and rpb2 sequence data. The tree
is rooted with Galiicola baoshanensis (KUN-HKAS 102234) and
Galiicola pseudophaeosphaeria (MFLUCC 14-0524). Related
sequences are retrieved from GenBank with 16 strains included in
the analysis of the combined loci and comprises 4177 characters
(820 characters for LSU, 1012 characters for SSU, 561 characters
for ITS, 905 characters for tef1, 879 characters for rpb2, including
gaps). The best scoring RAxML tree had a final likelihood value of
− 7456.746869. The matrix had 277 distinct alignment patterns with
31.92% undetermined characters and gaps. Estimated base frequencies were: A = 0.244763, C = 0.241642, G = 0.263061, T = 0.250533;
substitution rates AC = 1.095229, AG = 1.632728, AT = 1.392298,
CG = 0.793970, CT = 6.972828, GT = 1.000000; gamma distribution
shape parameter α = 6.674062. Maximum parsimony analysis of 111
parsimony informative characters resulted in a most parsimonious
tree (CI = 0.916, RI = 0.858, RC = 0.786, HI = 0.084). In our analysis,
GTR + I + G model was used for each partition in Bayesian posterior
analysis. Bootstrap values (BS) from maximum parsimony (MP, left),
maximum likelihood (ML, right) higher than 50% BS and Bayesian
posterior probabilities (BYPP, below) greater than 0.90 are given
at the nodes. Hyphens (-) represent support values less than 50%
ML/0.90 BYPP. Thick branches represent significant support values
from all analyses (ML ≥ 70%/BYPP ≥ 0.95). The ex-type strains are in
bold and black. The newly generated sequence is in bold and blue
GenBank accession numbers: LSU: MT214585; SSU:
MT226698; ITS: MT310630; tef1: MT394643; rpb2:
MT394699.
Notes: Pseudoberkleasmium chiangmaiense (MFLUCC
17–2088) is similar to the type except for culture characters
(Hyde et al. 2019a, Fig. 63), however, our collection was
grown on MEA while the type was cultured on PDA. Phylogenetically, P. chiangmaiense (strain MFLUCC 17–2088)
formed a clade with the type strain (MFLUCC 17–1809)
with strong support (100% ML/1.00 BYPP, Fig. 2). The type
strain was isolated from decaying wood collected in Chiang
Mai, therefore, we present a new host record of P. chiangmaiense on Clematis sikkimensis (Fig. 63).
Pseudomassarinaceae Phukhams. & K.D. Hyde, fam. nov.
Index Fungorum number: IF557104; Facesoffungi number: FoF 07212, Fig. 64.
Saprobic on dried herbaceous plants. Sexual morph:
Ascomata immersed, only ostioles visible, uniloculate,
obpyriform to subglobose, light brown to brown, coriaceous.
Ostioles central, brown to dark brown, carbonaceous, papillate. Peridium thin, multilayered, comprising thin-walled,
light brown to brown cells of textura angularis, inner layers
comprising hyaline cells. Hamathecium of dense, filiform,
branched, pseudoparaphyses. Asci 8-spored, bitunicate, fissitunicate, thick-walled, oblong, apically rounded, with furcate
pedicel. Ascospores biseriate, overlapping, hyaline, broad
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Fungal Diversity (2020) 102:1–203
◂Fig. 58 Wojnowiciella clematidis (MFLU 17–1517, holotype). a, b
Appearance of ascomata on host surface. c Vertical section through
ascoma. d Section of peridium. e Pseudoparaphyses. f–h Asci. i–n
Ascospores. Scale bars: b, c = 200 µm, d–h = 50 µm, i–n = 20 µm
fusiform, tapering towards the ends, acute at both ends, with
transverse septa, with mucilaginous sheath. Asexual morph:
Undetermined.
Type genus: Pseudomassarina Phukhams. & K.D. Hyde
Notes: The new family Pseudomassarinaceae is introduced to accommodate a monotypic genus, Pseudomassarina (Fig. 64). Phylogenetic analysis which included related
families in Pleosporales, showed that Pseudomassarinaceae
formed a distinct lineage related to Amorosiaceae, Halotthiaceae, Lophiostomataceae, Neomassarinaceae, Phaeoseptaceae and Sporormiaceae (Fig. 2). The family is morphologically similar to Lophiostomataceae, Neomassariaceae and
Neomassarinaceae in having hyaline, ellipsoid to fusiform,
septate ascospores (Thambugala et al. 2015; Ariyawansa
et al. 2018; Mapook et al. 2020). Pseudomassarinaceae is
distinguished by its immersed, coriaceous ascomata with
crest-like, carbonaceous ostioles, oblong and short pedicellate asci and wide mucilaginous sheath surrounding the
ascospores.
Pseudomassarina Phukhams. & K.D. Hyde, gen. nov.
Index Fungorum number: IF557097; Facesoffungi number: FoF 07213, Fig. 64.
Etymology: The genus epithet reflects its morphological
similarity to Massarina species.
Saprobic on decaying plants in terrestrial habitats. Sexual
morph: Ascomata on surface of the host, visible as black
spots, immersed, solitary, uniloculate, obpyriform, brown,
rough-walled, coriaceous, with apical ostioles. Ostioles central, brown to dark brown, partially carbonaceous, papillate,
filled with periphyses. Peridium multilayered, comprising
thin-walled, light brown to brown cells of textura angularis, inner layers comprising hyaline cells. Hamathecium of
dense, filiform, branched, transverse septa, pseudoparaphyses. Asci 8-spored, bitunicate, fissitunicate, thick-walled,
oblong, apically rounded, short pedicellate, with an ocular
chamber. Ascospores biseriate, overlapping, hyaline, broad
fusiform, tapering towards the ends, with transverse septum, guttulate, smooth-walled, with mucilaginous sheath.
Asexual morph: Undetermined.
Type species: Pseudomassarina clematidis Phukhams,
Camporesi & K.D. Hyde
Notes: Pseudomassarina, is typified by P. clematidis
and formed a distinct lineage in Pleosporales (Fig. 2). The
morphological comparison showed that our collection has
unique characters among related taxa in Pleosporales (Zhang
et al. 2012; Hyde et al. 2013; Thambugala et al. 2015; Ariyawansa et al. 2018; Mapook et al. 2020). In a BLASTn
101
search of GenBank, the closest match of the LSU sequence
of MFLU 16–0493 is Preussia terricola (strain CBS 317.65)
with 97.43% similarity. The closest match with the ITS
sequence is Preussia polymorpha (strain CBS 117679) with
84.49% similarity.
Pseudomassarina clematidis Phukhams, Camporesi & K.D.
Hyde sp. nov.
Index Fungorum number: IF557098; Facesoffungi number: FoF 07214, Fig. 64.
Etymology: The epithet reflects Clematis.
Holotype: MFLU 16–0493.
Saprobic on dead stems of Clematis vitalba. Sexual
morph: Ascomata 150–220 × 80–130 μm ( x̄ = 190 × 110 μm,
n = 5), on surface of the host, visible as black spots,
immersed, only ostioles visible, solitary, scattered, uniloculate, obpyriform to subglobose, light brown to brown, roughwalled, coriaceous, with apical ostioles. Ostioles central,
65–70 × 50–55 μm, brown to dark brown, papillate, opening
by a pore, filled with periphyses. Peridium 10–20 μm wide,
multilayered, comprising of 4–5 layers of thin-walled light
brown to brown cells of textura angularis, inner layers comprising hyaline cells. Hamathecium of dense, 1.3–1.5 μm
wide ( x̄ = 1.3 μm, n = 50), filiform, branched, septate, pseudoparaphyses. Asci 55–80 × 9–15 μm ( x̄ = 60 × 10 μm,
n = 30), 8-spored, bitunicate, fissitunicate, thick-walled,
oblong, apically rounded, with short, furcate pedicel, ocular chamber clearly visible. Ascospores 16–25 × 3–7 μm
( x̄ = 20 × 5 μm, n = 50), biseriate, overlapping, hyaline, broad
fusiform, tapering towards the ends, acute at both ends, with
1 transverse septum, with large guttules in each cell, deeply
constricted at the septum, cell above septum longer and
wider than below cell, smooth-walled, with 4–6 μm wide
mucilaginous sheath. Asexual morph: Undetermined.
Material examined: Italy, Forlì-Cesena Province, Fiumicello-Premilcuore, on dead aerial branch of Clematis vitalba,
20 March 2016, E. Camporesi, IT 2335 (MFLU 16–0493,
holotype).
Host: Clematis vitalba—(This study).
Distribution: Italy—(This study).
GenBank accession numbers: LSU: MT214586; SSU:
MT226699, MT310631; ITS: MT415397; tef1: MT394644;
rpb2: MT394700.
Notes: See note under Pseudomassarina.
Pseudolophiotremataceae K.D. Hyde & S. Hongsanan
Pseudolophiotremataceae was introduced by Hongsanan
et al. (2018) with Pseudolophiotrema elymicola as the
generic type. The multi-locus analysis (Fig. 2) showed that
Clematidis italica (MFLUCC 15–0084) clustered with P.
elymicola. Thus, we place Clematidis italica (Fig. 65) in
Pseudolophiotremataceae.
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Fungal Diversity (2020) 102:1–203
Stemphylium rosae-caninae MFLU 16-0245
Stemphylium rosae MFLU 16-0254
Stemphylium vesicarium MFLUCC 16-0998
Stemphylium vesicarium IT 1785B
Stemphylium vesicarium MFLUCC 14-0261
Stemphylium vesicarium MFLUCC 13–0344
--/0.90
Stemphylium vesicarium MFLUCC 14-0261
Stemphylium vesicarium CBS 109843
Stemphylium vesicarium CBS 109844
53/0.91
Stemphylium vesicarium CBS 192.86
Stemphylium vesicarium CBS 191.86
Stemphylium armeriae CBS 338.73
53/-Stemphylium vesicarium CBS 122640
93/0.92 Stemphylium majusculum CBS 133424
Stemphylium majusculum CBS 717.68
Stemphylium clematidis MFLUCC 14–0937
Stemphylium gracilariae CBS 115180
Stemphylium lucomagnoense CBS 116601
Stemphylium astragali CBS 116583
72/0.95
Stemphylium eturmiunum CBS 109845
Stemphylium botryosum CBS 714.68
82/0.99 Stemphylium halophilum CBS 410.73
82/0.99
Stemphylium halophilum CBS 337.73
Stemphylium lycii CBS 125241
56/-Stemphylium herbarum MFLUCC 14-0920
55/-Stemphylium amaranthi CBS 124746
-94/0.99
Stemphylium canadense CBS 116602
Stemphylium paludiscirpi CBS 109842
73/0.98
Stemphylium triglochinicola CBS 718.68
Stemphylium sarciniforme CBS 335.33
100/1.00
92/0.99
53/-Stemphylium sarciniforme CBS 136810
55/-Stemphylium loti CBS 407.54
Stemphylium
beticola CBS 141024
93/0.99
100/1.00
Stemphylium simmonsii CBS 716.68
52/0.98
Stemphylium chrysanthemicola CBS 117255
89/0.99
Stemphylium trifolii CBS 116580
Stemphylium drummondii CBS 346.83
--/0.90
Stemphylium novae-zelandiae CBS 138295
Stemphylium ixeridis CBS 124748
Stemphylium lancipes CBS 133314
100/1.00
Stemphylium lycopersici CBS 122803
Stemphylium symphyti CBS 115268
Stemphylium callistephi CBS 527.50
77/-Stemphylium solani CBS 408.54
Curvularia heteropogonis CBS 284.91
100/1.00
Curvularia
Curvularia ravenelii BRIP 13165
Stemphylium
66/--
0.02
Fig. 59 The Bayesian 50% majority-rule consensus phylogram based
on combined LSU, ITS and gadph data for Stemphylium. The topology and clade stability of the combined gene analyses was compared
to the single gene analyses. The tree is rooted with Curvularia species. Forty-six strains were included in the combined gene sequence
analyses which comprised 2052 characters (902 characters for LSU,
555 characters for ITS, 595 characters for gadph, including gap
regions). The best scoring RAxML tree had a likelihood value of
− 6557.926637. The matrix had 441 distinct alignment patterns with
29.84% undetermined characters and gaps. Estimated base frequencies were as follows; A = 0.245289, C = 0.253669, G = 0.255798,
13
T = 0.245244; substitution rates AC = 1.484723, AG = 3.274866,
AT = 0.996508, CG = 1.050822, CT = 5.664441, GT = 1.000000;
gamma distribution shape parameter α = 0.781417. In our analysis,
GTR + I + G model was used for each partition in Bayesian posterior
analysis. The species determined in this study are indicated in blue.
Bootstrap values (BS) greater than 50% BS (ML, left) and Bayesian
posterior probabilities (BYPP, right) greater than 0.90 are given at the
nodes. Hyphens (-) represent support values less than 50% BS/0.90
BYPP. Thick branches represent significant support values from all
analyses (BS ≥ 70%/BYPP ≥ 0.95)
Fungal Diversity (2020) 102:1–203
103
Fig. 60 Stemphylium clematidis (MFLU 16–0176, holotype). a
Appearance of ascomata on host surface. b Close up of ascomata. c
Vertical section though ascoma. d Peridium. e Pseudoparaphyses. f–h
Asci. i–m Ascospores (m Ascospore stained in 10% India ink). Scale
bars: a = 500 µm, b = 200 µm, c = 100 µm, d–h = 20 µm, i–m = 10 µm
Clematidis italica Tibpromma, Camporesi & K.D. Hyde, in
Li et al., Fungal Diversity 78: 60 (2016)
Index Fungorum number: IF 551867, Facesoffungi number: FoF 01813, Fig. 65.
Notes: Li et al. (2016) introduced Clematidis italica
(strain MFLUCC 15–0084) based on analyses of combined
LSU and SSU sequence data. Subsequently, Hashimoto et al.
(2017) introduced Pseudolophiotrema elymicola (MAFF
239600) based on analyses of a SSU, ITS, LSU, tef1 and
rpb2 dataset (Jaklitsch et al. 2018). In our LSU phylogenetic
tree for Pleosporales, Clematidis and Pseudolophiotrema
formed a closely related clade with strong support (94%
ML/0.98 BYPP, data not shown). Consequently, in a combined dataset of LSU, SSU, ITS, tef1 and rpb2, Clematidis
italica (MFLUCC 15–0084) and Pseudolophiotrema elymicola (JCM 13090) formed a well-supported clade (100%
ML/1.00 BYPP) basal to Anteagloniaceae (Fig. 2). A similar
topology is shown in Hashimoto et al. (2017) wherein P.
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Fungal Diversity (2020) 102:1–203
Fungal Diversity (2020) 102:1–203
◂Fig. 61 Stemphylium vesicarium (MFLU 16–1109). a Appear-
ance of ascomata on host surface. b Vertical section of ascoma.
c Ostiolar canal. d Section of peridium. e Pseudoparaphyses. f–h
Asci. i–n Ascospores (n Ascospore in 10% Indian ink). o Germinated ascospore. p, q Culture characteristics on MEA. Scale bars:
a = 500 µm, b = 200 µm, c–h = 50 µm, i–n = 10 µm, o = 20 µm, p,
q = 50 mm
elymicola formed a distinct clade from Lophiotremataceae.
As the epithet “clematidis” is similar to the plant family
Clematidaceae/Clemataceae, it is difficult to introduce a
generic epithet. Therefore, Pseudolophiotrema elymicola
is selected as a generic type for Pseudolophiotremataceae
(Hongsanan et al. 2020).
Roussoellaceae Liu, Phookamsak, Dai & K.D. Hyde
Roussoellaceae was introduced by Liu et al. (2014) and
family members have steadily increased (Jaklitsch and Voglmayr 2016; Tibpromma et al. 2017, 2018; Wanasinghe et al.
2018; Jiang et al. 2019; Karunarathna et al. 2019; Phookamsak et al. 2019; Mapook et al. 2020). We provide an updated
phylogenetic analysis for Roussoellaceae based on a concatenated LSU, ITS, tef1, rpb2, and SSU sequence dataset.
New host records and new species occurring on Clematis
species in Thailand are described based on morphological
characteristics coupled with multigene phylogenetic analyses
(Fig. 66).
Neoroussoella Liu et al.
Neoroussoella was introduced with the single species, N.
bambusae Phookamsak, J.K. Liu & K.D. Hyde. Phylogenetic placement of the species in Roussoellaceae was demonstrated by Karunarathna et al. (2019) and Mapook et al.
(2020). A multigene phylogenetic analysis (Fig. 66) reveals
two novel species N. clematidis and N. fulvicomae, and the
first record of N. heveae from a Clematis species.
Neoroussoella clematidis Phukhams. & K.D. Hyde, sp. nov.
Index Fungorum number: IF557113; Facesoffungi number: FoF 07328, Fig. 67.
Etymology: Name refers to the host genus, Clematis.
Holotype: MFLU 17–1467
Saprobic on dried stems of Clematis subumbellata. Sexual morph: Ascomata 230–250 × 155–160 μm
( x̄ = 235 × 158 μm, n = 5), only ostioles present on the surface of host, solitary, gregarious, erumpent, semi-immersed,
globose to subglobose, black to rust brown, coriaceous,
rough-walled, ostiolate. Ostioles central, 84–90 × 67–70 μm,
dark brown to black, papillate, ostiolar canal lined with periphyses. Peridium 12–29 μm wide, outer layer composed of
5–7 layers of brown to chestnut brown cells of textura angularis, with thin, hyaline inner layer. Hamathecium of dense,
0.8–1.5 μm wide ( x̄ = 1.3 μm, n = 40), filiform, branched,
septate, trabeculate pseudoparaphyses. Asci 62–82 × 4–7 μm
105
( x̄ = 71 × 5 μm, n = 30), (2–)8-spored, bitunicate, fissitunicate, cylindrical, apically rounded, with bulbose pedicel, ocular chamber visible when young. Ascospores
9–11 × 3–5 μm ( x̄ = 10 × 4 μm, n = 50), uniseriate, partially
overlapping, ellipsoid with rounded ends, olive to yellowish brown, 1-septate, constricted at septum, guttulate in
each cell, rough-walled, at maturity, slightly longitudinally
ribbed, without a mucilaginous sheath. Asexual morph:
Undetermined.
Culture characters: Colonies on MEA reaching 20 mm
diam. after 4 weeks at 25 °C. Culture from above, brown,
radiating outwards, dense, irregular in shape, umbonate,
dull, edge erose, downy, covered with fairly white mycelium; reverse brown at the middle dark brown at the edge,
with domes on the media.
Material examined: Thailand, Chiang Rai Province, on
dead branches of Clematis subumbellata, 20 March 2017,
C. Phukhamsakda, CMTH03 (MFLU 17–1467, holotype);
ex-type living culture, MFLUCC 17–2061.
Host: Clematis subumbellata—(This study).
Distribution: Thailand—(This study).
GenBank accession numbers: LSU: MT214587; SSU:
MT226700; ITS: MT310632; tef1: MT394645; rpb2:
MT394701.
Notes: Neoroussoella clematidis shares common characters with Neoroussoella in having uniloculate ascomata
without a clypeus, cylindrical to broad filiform asci with
bulbose pedicel and 1-septate ascospores (Liu et al. 2014).
Our collection is distinguishable by having immersed and
globose ascomata with papillate ostioles (Fig. 67). In the
phylogenetic analysis, N. clematidis (MFLUCC 17–2061)
formed a strongly supported clade (100% ML/1.00 BYPP,
Fig. 66) with N. fulvicoma (MFLUCC 17–2073) which was
an asexual morph (Fig. 68). A comparison of the ITS region
(including 5.8S region) showed 7.7% base pair differences
(with gaps) and 3.4% base pair differences (with gaps) in the
tef1 region, which is evidence for new species rank.
The isolate MFLUCC 17–2061 was evaluated for the
potential of secondary metabolites against Bacillus subtillis,
Escherichia coli, Mucor plumbeus and Schizosaccharomyces
pombe. The strain demonstrated moderate inhibitory activities against Bacillus subtillis and against conidia development in Mucor plumbeus. This isolate is suitable for further
evaluation of secondary metabolites.
Neoroussoella fulvicomae Phukhams. & K.D. Hyde, sp.
nov.
Index Fungorum number: IF557114; Facesoffungi number: FoF 07329, Fig. 68.
Etymology: Name refers to Clematis fulvicoma.
Holotype: MFLU 17–1481.
Saprobic on dried stems of Clematis fulvicoma. Sexual
morph: Undetermined. Asexual morph: Conidiomata
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Fig. 62 Stemphylium vesicarium (MFLUCC 16–0998). a Appearance of asexual morph on the surface of MEA. b, c Close up of conidiophore.
d–k Conidia. Scale bars: a = 200 µm, b = 100 µm, c, d = 50 µm, e–k = 20 µm
140–241 × 137–155 μm ( x̄ = 207 × 146 μm, n = 5), pycnidial,
solitary, sometimes aggregated, uniloculate, immersed, visible as minute, black, shiny ostioles, subglobose, coriaceous,
thick-walled, dark brown to brown, with papilla, ostiolate.
Ostioles 51–60 × 49–55 μm, central, oblong, papillate. Conidiomatal wall 12–18 μm wide ( x̄ = 15 μm, n= 20), thick,
outer layer composed of 5–7 layers of brown to light brown
cells of textura angularis, lined with a hyaline layer bearing conidiogenous cells. Conidiophores reduced to conidiogenous cells. Conidiogenous cells 2.5–5 × 1.5–2 μm
( x̄ = 4 × 1.7 μm, n = 20), enteroblastic, phialidic, annellidic, determinate, discrete, truncate, smooth-walled, hyaline, arising from the inner layers of conidiomata. Conidia
3–6 × 2–3 μm ( x̄ = 4.5 × 2.5 μm, n = 100), oval, slightly
13
curved, hyaline when immature, brown at maturity, with
1(–2) guttules in each cell, aseptate, smooth-walled.
Culture characters: Colonies on MEA reaching 50 mm
diam. after 4 weeks at 25 °C. Cultures from above, black,
dense, circular, margin undulate, umbonate, fluffy, wrinkled,
folded, covered with brown aerial mycelium; reverse black
at the centre radiating outwardly.
Material examined: Thailand, Chiang Rai Province, on
dead branches of Clematis fulvicoma, 20 March 2017, C.
Phukhamsakda, CMTH17 (MFLU 17–1481, holotype); extype living culture, MFLUCC 17–2073.
Host: Clematis fulvicoma—(This study).
Distribution: Thailand—(This study).
Fungal Diversity (2020) 102:1–203
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Fig. 63 Pseudoberkleasmium chiangmaiense (MFLU 17–1496). a
Sporodochia on natural substrate. b Vertical section through sporodochia. c Hyphae. d–g Mature lenticular conidia. h Germinated
conidia. i, j Culture characteristics on MEA. Scale bars = a = 1 cm,
b = 100 μm, c, g–h = 20 μm, d–f = 10 μm
GenBank accession numbers: LSU: MT214588; SSU:
MT226701; ITS: MT310633; tef1: MT394646; rpb2:
MT394702.
Notes: In the phylogenetic analysis of combined sequence
data Neoroussoella fulvicomae clustered in a separate clade
with N. clematidis (100% ML/1.00 BYPP, Fig. 66). Neoroussoella fulvicomae is similar to Neoroussoella in having
immersed, uniloculate, globose to subglobose pycnidia,
annellidic conidiogenesis cells, and aseptate, 2-guttulate
conidia (Liu et al. 2014). Neoroussoella fulvicomae is distinguishable by having papillate ostioles and short conidiogenesis cells and brown conidia with 1–2 guttules in each
cell (Fig. 68). Neoroussoella fulvicomae was evaluated for
potential of secondary metabolites in the same manner as N.
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108
Fig. 64 Pseudomassarina clematidis (MFLU 16–0493, holotype).
a Appearance of ascomata on host surface. b Vertical section of
ascoma. c Ostiolar canal. d Section of peridium. e Pseudoparaphy-
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ses. f–h Asci. i–m Ascospores. n Ascospore in 10% Indian ink. Scale
bars: b = 100 µm, c = 50 µm, d–h = 20 µm, i–n = 10 µm
Fungal Diversity (2020) 102:1–203
109
Fig. 65 Clematidis italica (MFLU 14–0669, holotype). a Appearance of ascomata on host substrate. b Vertical section of ascoma. c Section of
peridium. d Pseudoparaphyses. e–g Asci. h–j Ascospores. Scale bars: a = 200 μm, b = 50 μm, c = 10 μm, d, h–j = 5 μm, e–g = 20 μm
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Fungal Diversity (2020) 102:1–203
Roussoella japanensis MAFT 239636
Roussoella hysterioides CBS 546.94
69/-Roussoella verrucispora CBS 125434
99/1.00
Roussoella scabrispora MFLUCC 11-0624
Roussoella scabrispora RSC
66/0.99
Roussoellopsis tosaensis KT 1659
100/1.00
98/1.00
Roussoellopsis tosaensis MAFF 239638
Roussoellopsis macrospora MFLUCC 12-0005
73/1.00
Roussoellopsis sp. KT 1710
100/1.00 Roussoella intermedia KT 2303
Roussoella intermedia NBRC 106245
Roussoella
99/1.00
Roussoella pustulans KT 1709
77/1.00
Roussoella
nitidula
MFLUCC
11-0634
100/1.00
72/1.00
Roussoella nitidula MFLUCC 11-0182
65/1.00
Roussoella pustulata MFLUCC 13-0854
100/1.00 Roussoella pseudohysterioides MFLUCC 13-0852
100/1.00
Roussoella pseudohysterioides KMUCC 18-0111
Roussoella thailandica MFLUCC 11-0621
Roussoella neopustulans MFLUCC 12-0003
100/1.00
Roussoella neopustulans MFLUCC 11-0609
--/1.00
Roussoella kunmingensis MFLUCC 18-0128
Roussoella angustior MFLUCC 15-0186
73/0.93
86/1.00
Roussoella magnatum MFLUCC 15-0185
76/1.00
Roussoella chiangraina MFLUCC 10-0556
Roussoella siamensis MFLUCC 11-0149
68/-Roussoella yunnanensis MFLUCC 18-0115
51/0.95
Roussoella doimaesalongensis MFLUCC 14-0584
95/0.97 Setoarthopyrenia chromolaenae MFLUCC 17-1444
Setoarthopyrenia
Arthopyrenia sp. UTHSC DI16-362
Arthopyrenia salicis UTHSC DI16-220
94/-71/-Roussoella intermedia CBS 170.96
98/1.00
Arthopyrenia salicis CBS 368.94
Roussoella mexicana CPC 25355
Neoroussoella entadae MFLUCC 17-0920
52/0.97
Neoroussoella entadae MFLUCC 18-0243
52/0.98
--/0.99
Neoroussoella entadae MFLUCC 15-0098
Neoroussoella solani CPC 26331
85/1.00
Neoroussoella leucaenae KT 3264
61/0.98
94/1.00
Neoroussoella leucaenae MFLUCC 17-0927
65/0.99
Neoroussoella leucaenae MFLUCC 18-1544
Neoroussoella
--/1.00
Neoroussoella lenispora GZCC 16-0020
100/1.00
Neoroussoella clematidis MFLUCC 17-2061
61/0.99
Neoroussoella fulvicomae MFLUCC 17-2073
100/0.99
Neoroussoella heveae MFLUCC 17-2069
--/0.99
Neoroussoella heveae MFLUCC 17-0338
100/1.00 Neoroussoella alishanense FU31018
Neoroussoella alishanense FU31016
100/1.00
Neoroussoella bambusae MFLUCC 11-0124
Pseudoroussoella elaeicola MFLUCC 15-0276b
Pseudoroussoella elaeicola MFLUCC 17-1483
97/0.99
Pseudoroussoella elaeicola MFLUCC 15-0276a
100/1.00
Pseudoroussoella elaeicola MFLUCC 17-2059 Pseudoroussoella
96/0.92 Pseudoroussoella chromolaenae MFLUCC 17-1492
83/0.99
Pseudoroussoella chromolaenae MFLUCC 17-2062
Pseudoneoconiothyrium rosae MFLUCC 15-0052
99/1.00
Pseudoneoconiothyrium euonymi CBS 143426 Pseudoneoconiothyrium
99/1.00
99/1.00
Arthopyrenia sp.UTHSC DI16-334
Xenoroussoella
98/0.99
Xenoroussoella triseptata MFLUCC 17-1438
96/1.00
95/1.00
77/1.00
100/1.00
100/1.00
99/0.96
100/1.00
100/1.00
100/1.00
0.06
13
Pararoussoella mukdahanensis KUMCC 18-0121
Pararoussoella mukdahanensis MFLUCC 11-0201
Pararoussoella juglandicola CBS 145037
Pararoussoella
Pararoussoella rosarum MFLU 17-0654
Pararoussoella mangrovei MFLUCC 16-0424
Thyridaria broussonetiae TB
Thyridaria broussonetiae TB1
Ohleria modesta OM
100/1.00
Ohleria modesta MGC
Torula hollandica CBS 220.69
Torula herbarum CPC 24114
100/1.00
Roussoellaceae
ML/BYPP
Thyridariaceae
Ohleriaceae
Torulaceae
(Outgroup)
Fungal Diversity (2020) 102:1–203
◂Fig. 66 Best scoring RAxML tree with a final likelihood value of
− 27212.762040 based on combined LSU, ITS, tef1, rpb2, and SSU
sequence data for Roussoellaceae. The tree is rooted with members of
Torulaceae. Sixty-nine strains were included in the combined analyses which comprised 4382 characters (819 characters for LSU, 567
characters for ITS, 906 characters for tef1, 1052 characters for rpb2,
1038 characters for SSU, including gaps). The topology and clade stability of the combined gene analyses was compared to the single gene
analyses. The tree from the maximum likelihood analysis had similar
topology to the Bayesian 50% majority-rule consensus phylogram.
The matrix had 1501 distinct alignment patterns with 38.60% of
undetermined characters and gaps. Estimated base frequencies were
as follows: A = 0.245133, C = 0.255324, G = 0.268618, T = 0.230925;
substitution rates AC = 1.766964, AG = 5.243182, AT = 2.353818,
CG = 1.339749, CT = 11.449776, GT = 1.000000; gamma distribution
shape parameter α = 0.473912. In our analysis, GTR + I + G model
was used for each partition in Bayesian posterior analysis. The isolates of this study are in blue. Bootstrap values (BS) greater than 50%
BS (ML, left) and Bayesian posterior probabilities (BYPP, right)
greater than 0.90 are given at the nodes. Hyphens (-) represent support values less than 50% BS/0.90 BYPP. Thick branches represent
significant support values from all analyses (BS ≥ 70%/BYPP ≥ 0.95)
at the genus and family levels
clematidis. Interestingly, isolate MFLUCC 17–2073 does not
produce growth inhibitory activity against Bacillus subtillis
and Mucor plumbeus.
Neoroussoella heveae Senwanna, Phookamsak & K.D.
Hyde, in Phookamsak et al., Fungal Divers [66] (2019), new
host record
Index Fungorum number: IF555287; Facesoffungi number: FoF 07330, Fig. 69.
Saprobic on dried stems of Clematis subumbellata. Sexual morph: Undetermined. Asexual morph: Conidiomata
128–263 × 142–304 μm ( x̄ = 177 × 200 μm, n = 5), pycnidial, solitary, gregarious, uniloculate, immersed, visible
only as minute ostioles, globose, coriaceous, thick-walled,
dark brown to brown, ostiolate. Ostioles 40 × 77 μm, central, papillate, oblong, opening by a pore. Conidiomatal
wall 24–31 μm wide ( x̄ = 26 μm, n= 20), thick, 7–8 layers,
outer layer composed of brown cells of textura angularis,
lined with a hyaline layer bearing conidiogenous cells. Conidiophores reduced to conidiogenous cells. Conidiogenous
cells 2–4 × 1.5–2 μm ( x̄ = 3 × 2 μm, n = 50), enteroblastic,
phialidic, determinate, discrete, truncate, smooth-walled,
hyaline, arising from the inner layers of the conidiomata.
Conidia 3.5–4.5 × 2.3–3 μm ( x̄ = 4 × 2.5 μm, n = 100), oval,
hyaline when immature, olive brown at maturity, slightly
curved at the ends, with 1(–2) guttules in each cell, aseptate,
smooth-walled.
Culture characters: Colonies on MEA reaching 50 mm
diam. after 4 weeks at 25 °C. Cultures from above, dark
brown to brown, dense, irregular, margin lobate, umbonate,
wrinkled folded, thinly hairy, covered with brown aerial
mycelium; reverse black at the centre, radiating outwardly.
111
Material examined: Thailand, Chiang Rai Province, on
dead stems of Clematis subumbellata, 20 March 2017, C.
Phukhamsakda, CMTH13 (MFLU 17–1477); living culture,
MFLUCC 17–2069.
Hosts: Clematis subumbellata, Hevea brasiliensis—
(Phookamsak et al. 2019; this study).
Distribution: Thailand—(Phookamsak et al. 2019; this
study).
GenBank accession numbers: LSU: MT214589; SSU:
MT226702; ITS: MT310634; tef1: MT394647; rpb2:
MT394703.
Notes: Neoroussoella heveae was described from a twig
of Hevea brasiliensis from northern Thailand (Phookamsak
et al. 2019). Isolate MFLUCC 17–2069, recorded on Clematis formed a close relationship (100% ML/1.00 BYPP) with
the type strain of N. heveae (MFLUCC 17–0338). Our collection (MFLU 17–1477, Fig. 69) had larger conidia than the
type (128–263 × 142–304 vs 90–130 × 115–180 μm). Comparison of ITS sequence data revealed only 1 base pair difference between our isolate and the type. Unfortunately, the
tef1 region of the type strain is not available for comparison.
Neoroussoella heveae was evaluated for potential secondary metabolite production in the same manner as N. clematidis. Isolate MFLUCC 17–2069 demonstrated weak inhibitory activities against Bacillus subtillis.
Pararoussoella Wanas., E.B.G. Jones & K.D. Hyde
Pararoussoella (type species P. rosarum) was introduced
for species that are distantly related to the type species of
Roussoella (R. nitidula Sacc. & Paol.). The genus is characterized by having immersed and globose ascomata with
minute black dots of ostioles, cellular pseudoparaphyses,
central, papillate, black ostioles, cylindrical to oblong asci,
and uniseriate, brown to dark brown ascospores, with irregular, longitudinal striations (Wanasinghe et al. 2018). Multigene phylogenetic analysis (Fig. 66) showed that Roussoella
mangrovei is related to Pararoussoella, thus a new combination is proposed.
Pararoussoella mangrovei (Phukhams. & K.D. Hyde)
Phukhams. & K.D. Hyde, comb. nov.
≡ Roussoella mangrovei Phukhams. & K.D. Hyde, in
Hyde et al., Mycosphere 9(2): 339 (2018)
Index Fungorum number: IF557372; Facesoffungi number: FoF 03923
Notes: Pararoussoella mangrovei formed a related lineage with other Roussoella species. Karunarathna et al.
(2019) showed that the strain is related to Pararoussoella
(Fig. 66). Therefore, Roussoella mangrovei is transferred to
Pararoussoella.
Pseudoneoconiothyrium Wanas., Phukhams., Camporesi
& K.D. Hyde
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Fig. 67 Neoroussoella clematidis (MFLU 17–1467, holotype). a, b
Appearance of ascomata on Clematis subumbellata. c Vertical section through ascoma. d Ostiolar canal. e Vertical section through
peridium. f Trabeculate pseudoparaphyses. g–i Asci. j-n Ascospores.
o, p Culture characters on MEA. Scale bars: b = 200 µm, c = 100 µm,
d–e = 50 µm, f–i = 20 µm, j–n = 5 µm
Pseudoneoconiothyrium was described as Neoconiothyrium for fungi associated with Rosaceae plants (Wanasinghe et al. 2018). Neoconiothyrium has been assigned for
the fungal strains in Coniothyriaceae (Crous et al. 2017;
Hawksworth et al. 2018). Thus, Pseudoneoconiothyrium was
proposed for Neoconiothyrium species in Roussoellaceae
and is typified by P. rosae (Phukhams., et al.) Phukhams.,
Camporesi & K.D. Hyde (Fig. 66).
13
Pseudoneoconiothyrium euonymi (Crous & Akulov)
Phukhams. & K.D. Hyde, comb. nov.
≡ Roussoella euonymi Crous & Akulov, in Crous et al.,
Fungal Systematics and Evolution 1: 204 (2018)
Fungal Diversity (2020) 102:1–203
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Fig. 68 Neoroussoella fulvicomae (MFLU 17–1481, holotype). a
Appearance of conidiomata on Clematis fulvicoma. b Close up of
conidioma on host substrate. c Vertical section through conidioma. d
Ostiolar canal. e Section of conidioma wall. f–h Conidiogenous cells
and conidia. i–m Conidia. n Culture characteristics on MEA. Scale
bars: b = 200 µm, c = 100 µm, d, e = 20 µm, f–m = 5 µm
Index Fungorum number: IF557605; Facesoffungi number: FoF 07331
Notes: Pseudoneoconiothyrium euonymi originally
clustered with Pararoussoella mukdahanensis (≡ Roussoella mukdahanensis). Based on the phylogenetic analysis
(Fig. 66), Pseudoneoconiothyrium euonymi is transferred as
a second species of Pseudoneoconiothyrium.
Pseudoroussoella chromolaenae Mapook & K.D. Hyde,
Mapook et al., Fungal Divers (2020), new host record
Index Fungorum number: IF557116; Facesoffungi number: FoF 07332, Fig. 70.
Saprobic on dead stems of Clematis subumbellata. Sexual morph: Undetermined. Asexual morph: Conidiomata
86–100 × 107–112 μm ( x̄ = 90 × 110 μm, n = 5), pycnidial,
solitary, sometimes aggregated, uniloculate, immersed,
with black shiny ostioles visible, globose to subglobose,
dark brown to brown, coriaceous, thick-walled, ostiolate.
Ostioles 55–70 × 50–67 μm, central, papillate, ovoid. Conidiomatal wall 6–14(–27) μm wide, outer layer composed
of 8–10 layers of brown to light brown cells of textura
angularis, lined with a hyaline layer bearing conidiogenous
cells. Conidiophores reduced to conidiogenous cells. Conidiogenous cells 3–7 × 1.5–3 μm ( x̄ = 4.5 × 2 μm, n = 30),
enteroblastic, phialidic, determinate, discrete, cylindrical
Pseudoroussoella Mapook & K.D. Hyde
Mapook et al. (2020) introduced Pseudoroussoella as a
separate lineage for P. euonymi and P. chromolaenae. Phylogenetic analyses (Fig. 66) coupled with morphological
characters of collections on Clematis species revealed a first
record of P. chromolaenae and P. euonymi on Clematis species (Figs. 70, 71).
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Fig. 69 Neoroussoella heveae (MFLU 17–1477). a Appearance
of conidiomata on Clematis subumbellata specimen. b Close up of
conidioma on host substrate. c Vertical section of conidioma. d Ostiolar canal. e Section of conidioma wall. f–i Conidiogenous cells
and conidia. j–l Conidia. m Germinated conidium. n, o Cultures
characters on MEA. Scale bars: c = 200 µm, d = 100 µm, e = 20 µm,
f–l = 5 µm
to sub-cylindrical, smooth-walled, hyaline. Conidia
4.5–8 × 3–5 μm ( x̄ = 6 × 4 μm, n = 50), oval, hyaline when
immature, yellowish brown at maturity, slightly curved, with
1(–2) guttules in each cell, aseptate, smooth-walled.
Culture characters: Colonies on MEA reaching 20 mm
diam. after 2 weeks at 25 °C. Cultures from above, greyish brown, dense, circular, umbonate, fluffy, covered with
white aerial mycelium; reverse dark brown at the central,
mycelium cream radiating outwardly.
Material examined: Thailand, Chiang Rai Province, on
dead branches of Clematis subumbellata, 20 March 2017, C.
Phukhamsakda, CMTH04 (MFLU 17–1468); living culture,
MFLUCC 17–2062.
Hosts: Chromolaena odorata, Clematis subumbellata—
(Mapook et al. 2020; this study).
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Fig. 70 Pseudoroussoella chromolaenae (MFLU 17–1468). a
Appearance of conidiomata on Clematis subumbellata. b Close up of
conidioma on host substrate. c Vertical section through conidioma. d
Ostiolar canal. e Section of conidioma wall. f–i Conidiogenous cells
and conidia (f conidiogenous cells in cotton blue). j–m Conidia n
Geminated conidium. o, p Cultures characters on MEA. Scale bars:
b = 500 µm, c = 200 µm, d = 100 µm, e = 20 µm, f–m = 5 µm
Distribution: Thailand—(Mapook et al. 2020; this study).
GenBank accession numbers: LSU: MT214590; SSU:
MT226703; ITS: MT310635; tef1: MT394648; rpb2:
MT394704.
Notes: Phylogenetic analysis (Fig. 66) has shown that the
newly collected strain (MFLUCC 17–2062), is related to
the ex-type strain of P. chromolaenae (MFLUCC 17–1492)
with strong support (100% ML/1.00 BYPP, Fig. 66). The
conidiomata of our collection (Fig. 70) are slightly smaller
than the ex-type strain (90 × 110 vs 165 × 195 µm). Pairwise comparison of the ITS region (including 5.8S region)
showed 100% similarity, while the tef1 region had two base
pair differences; these findings are not significantly distinct
for the introduction of the strain as a new species (Jeewon
and Hyde 2016). Pseudoroussoella chromolaenae strain
MFLUCC 17–2062 was evaluated for secondary metabolite
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Fig. 71 Pseudoroussoella elaeicola (MFLU 17–1465). a–c Appearance of ascomata on Clematis subumbellata. d Vertical section
through ascoma. e Ostiolar canal. f, g Section of peridium. h Pseudoparaphyses. i–l Asci. m–r Ascospores (q verruculose surface). s
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Germinated ascospore. t, u Culture characters on MEA. Scale bars:
a = 500 µm, b–d = 200 µm, e = 100 µm, f, g = 20 µm, h–l = 50 µm,
m–r = 10 µm
Fungal Diversity (2020) 102:1–203
production. The collection showed weak inhibitory activities
against the growth of Bacillus subtillis.
Pseudoroussoella elaeicola (Konta & K.D. Hyde) Mapook
& K.D. Hyde, new host record
Index Fungorum number: IF555291; Facesoffungi number: FoF 07333, Fig. 71.
Saprobic on dead stems of Clematis subumbellata.
Sexual morph: Ascomata 375–554 × 375–462 μm ( x̄
= 453 × 430 μm, n = 10), on the surface of the host, solitary, gregarious, erumpent, semi-immersed, dark brown
hyphae radiating outwards from the peridium wall, globose
to depressed-globose, black to dark brown, coriaceous,
rough-walled, ostiolar. Ostioles 149–157 × 123–140 μm,
central, dark brown to black, papillate, opening by a pore,
ostiolate with periphyses. Peridium 17–54(–83 μm at apex)
wide, outer layer composed of 8–10 layers of dark brown
to light brown cells of textura angularis, the inner layer
comprising thin, hyaline layers. Hamathecium of dense,
0.8–1.5 μm wide ( x̄ = 1.3 μm, n = 50), filiform, branches,
anastomosing above asci, septate, pseudoparaphyses. Asci
60–132 × 7–10 μm ( x̄ = 103 × 8 μm, n = 20), (2–)8-spored,
bitunicate, fissitunicate, cylindrical to cylindrical-clavate,
apically rounded with ocular chamber. Ascospores
10–18 × 5–10 μm (x̅ = 13 × 7 μm, n = 50), uniseriate, sometimes partially overlapping, oval with round ends, brown to
yellowish brown, uni-septate, constricted at septum, with
guttules in each cell, verruculose, with mucilaginous sheath.
Asexual morph: Undetermined.
Culture characters: Colonies on MEA reaching 30 mm
diam. after 4 weeks at 25 °C. Culture above dark green radiating, edge white, dense, circular, flattened, umbonate, edge
entire, fluffy; reverse light brown at the middle, dark brown
at the edge.
Material examined: Thailand, Chiang Rai Province, on
dead branches of Clematis subumbellata, 20 March 2017, C.
Phukhamsakda, CMTH01 (MFLU 17–1465); living culture,
MFLUCC 17–2059.
Hosts: Chromolaena odorata, Clematis subumbellata,
Elaeis guineensis—(Phookamsak et al. 2019; Mapook et al.
2020; this study).
Distribution: Thailand—(Phookamsak et al. 2019;
Mapook et al. 2020; this study).
GenBank accession numbers: LSU: MT214591; SSU:
MT226704; ITS: MH744730; tef1: MH750239; rpb2:
MT394705.
Notes: Pseudoroussoella elaeicola (≡ Roussoella elaeicola) was collected from Elaeis guineensis (Phookamsak
et al. 2019). Mapook et al. (2020) recorded this fungus
on Chromolaena odorata. In the phylogenetic analysis, R.
elaeicola formed a separate clade from the type species of
Roussoella (R. nitidula Sacc. & Paol.), thus Mapook et al.
(2020) synonymized R. elaeicola under Pseudoroussoella
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elaeicola. Our new isolate (MFLUCC 17–2059) grouped
with another collections reported from Chromolaena odorata and Elaeis guineensis (97% ML/0.99 BYPP, Fig. 70).
Our collection is identical to the other collections of this species (Fig. 71). Comparison of the ITS region revealed only
one base pair difference between our isolate and the ex-type
strain (MFLUCC 15–0276). The pairwise comparison of the
available tef1 region with MFLUCC 17–1483 also showed
one base pair difference. Therefore, we introduce our collection as a new host record.
Pseudoroussoella elaeicola (MFLUCC 17–2059) was
evaluated for the potential of secondary metabolite production. The isolate showed inhibitory activity on biofilm formation by Staphylococcus aureus and showed weak cytotoxicity on L929 murine fibroblasts and human KB3-1 cancer
cells (Phukhamsakda et al. 2018).
Sulcatisporaceae Tanaka & K. Hirayama
Sulcatisporaceae was erected for a well-separated clade
which included Magnicamarosporium, Neobambusicola and
Sulcatispora in Pleosporales (Tanaka et al. 2015). Rupcic
et al. (2018) added Pseudobambusicola to Sulcatisporaceae
which was collected from a twig of an unidentified plant in
Thailand based on the morphology and phylogeny analyses.
Sulcatisporaceae is characterized by immersed to erumpent,
subglobose to hemisphaerical ascomata, short ostiolar neck,
trabecular pseudoparaphyses (Liew et al. 2000), clavate,
short pedicellate asci, and broadly fusiform, hyaline, septate ascospores with mucilaginous appendages (Tanaka et al.
2015). Asexual morph is pycnidial conidiomata with various
conidia characters (Tanaka et al. 2015; Phukhamsakda et al.
2017a; Rupcic et al. 2018). We provide an updated tree of
Sulcatisporaceae based on the combined dataset of LSU,
ITS, SSU and tef1 sequence data and propose two new genera from Clematis species collected in Thailand. Additionally, the biological activity of their secondary metabolites is
preliminarily reported (Fig. 72).
Anthosulcatispora Phukhams. & K.D. Hyde, gen. nov.
Index Fungorum number: IF557201; Facesoffungi number: FoF 07340, Fig. 73.
Etymology: Anthos-meaning flower, Anthosulcatispora
referring to the occurrence on flowering plants.
Saprobic on dead stems of herbaceous plants. Sexual
morph: Ascomata semi-immersed, blackish, irregular, scattered, uniloculate, glabrous, ostiolate, apapillate. Ostioles
dark, circular and sunken. Peridium two-layered, outer layer
irregular, comprising dark brown cells of textura angularis and inner layer irregular comprising light brown cells.
Hamathecium composed of numerous, filamentous, branched
or simple, septate, cellular pseudoparaphyses, embedded in
a hyaline gelatinous matrix. Asci 4- or 8-spored, bitunicate,
cylindrical to cylindrical-clavate, short pedicellate, apically
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Magnicamarosporium diospyricola MFLUCC 16-0419
Magnicamarosporium iriomotense KT 2822
98/1.00
Pseudobambusicola thailandica BCC 79462
--/1.00
Parasulcatispora clematidis MFLUCC 17–2082
51/1.00
Sulcatispora acerina KT 2982
100/1.00
--/0.90
98/1.00
Sulcatispora berchemiae KT 1607
Neobambusicola strelitziae CBS 138869
Anthosulcatispora subglobosum MFLUCC 17−2065
100/1.00
--/1.00
Sulcatisporaceae
96/1.00
Anthosulcatispora brunnea MFLU 18−1393
100/1.00
Bambusicola bambusae MFLUCC 11-0614
Bambusicola splendida MFLUCC 11-0439
100/1.00
Bambusicolaceae
91/1.00
Bambusicola massarinia MFLUCC 11-0389
Latorua grootfonteinensis CBS 369.72
100/1.00
Latorua caligans CBS 576.65
100/1.00
Latoruaceae
Polyschema terricola CBS 301.65
100/1.00
Polyschema congolense CBS 542.73
Didymosphaeria rubi-ulmifolii MFLUCC 14-0024
91/1.00
Montagnula aloes CBS 132531
Didymosphaeriaceae
(Outgroup)
0.02
Fig. 72 Bayesian 50% majority-rule consensus phylogram based
on combined LSU, ITS, SSU and tef1 sequence data for Sulcatisporaceae. The topology and clade stability of the combined gene analyses was compared to the single gene analyses. The tree is rooted
with members of the Didymosphaeriaceae. Eighteen strains were
included in the combined analyses which comprised 3390 characters
(843 characters for LSU, 1025 characters for SSU, 585 characters for
ITS, 937 characters for tef1, including gap regions). The best scoring RAxML tree had a final likelihood value of − 11220.180237. The
matrix had 709 distinct alignment patterns, with 31.84% of undetermined characters and gaps. Estimated base frequencies were as fol-
lows; A = 0.238159, C = 0.264197, G = 0.261408, T = 0.236236;
substitution rates AC = 0.958607, AG = 2.023708, AT = 1.033715,
CG = 0.741292, CT = 5.717037, GT = 1.000000; gamma distribution
shape parameter α = 0.485427. In our analysis, GTR + I + G model
was used for each partition in Bayesian posterior analysis. The species determined in this study are indicated in blue. Bootstrap values (BS) greater than 50% BS (ML, left) and Bayesian posterior
probabilities (BYPP, right) greater than 0.90 are given at the nodes.
Hyphens (-) represent support values less than 50% BS/0.90 BYPP.
Thick branches represent significant support values from all analyses
(BS ≥ 70%/BYPP ≥ 0.95)
rounded, with an ocular chamber. Ascospores 1-seriate,
brown to dark brown, oblong to ellipsoidal, with rounded
ends, 1-septate, slightly constricted at the septum, smoothwalled (Phookamsak et al. 2019). Asexual morph: Conidiomata pycnidial, solitary or gregarious, unilocular, scattered, immersed or erumpent, base flattened, subglobose to
compressed, coriaceous, dark brown to reddish brown, with
or without ostioles. Pycnidial wall multilayered with cells
textura angularis, inner layer bearing conidiogenous cells.
Conidiophores reduced to conidiogenous cells. Conidiogenous cells enteroblastic, phialidic, determinate, elongated
cylindrical or truncate, smooth-walled, hyaline. Conidia
oblong, ends rounded, hyaline, aseptate, guttulate, smoothwalled, with mucilaginous sheath.
Type species: Anthosulcatispora subglobosa Phukhams.
& K.D. Hyde
Notes: Anthosulcatispora is introduced for a collection
occurring on stems of herbaceous plants (Phookamsak
et al. 2019; this study). The genus formed a basal lineage
in Sulcatisporaceae with strong support (98% ML/1.00
BYPP, Fig. 72). The sexual morph of Anthosulcatispora was
described as Neobambusicola brunnea (Phookamsak et al.
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2019). Asexual morphs in Sulcatisporaceae comprised Magnicamarosporium, Neobambusicola, Pseudobambusicola
and Sulcatispora (Tanaka et al. 2015; Phukhamsakda et al.
2017a; Rupcic et al. 2018). Anthosulcatispora is distinct
within Sulcatisporaceae in having brown ascospores, while
the sexual morph in Sulcatispora has hyaline, broadly fusiform ascospores with an entire sheath (Tanaka et al. 2015).
The asexual morph of Anthosulcatispora shares similar
characters with Neobambusicola and Pseudobambusicola.
They all have solitary, unilocular pycnidial, phialidic conidiogenesis and hyaline conidia. Anthosulcatispora is distinct from Neobambusicola and Pseudobambusicola in its
subglobose conidiomata, elongated cylindrical or truncate
conidiogenous cells and oblong and aseptate conidia, while
Neobambusicola and Pseudobambusicola have globose
conidiomata with two types of conidia (Crous et al. 2014b;
Rupcic et al. 2018).
Anthosulcatispora brunnea (Chen & C. Norphanphoun)
Phukhams. & K.D. Hyde, comb. nov.
Index Fungorum number: IF557202; Facesoffungi number: FoF 05708
Basionym: Neobambusicola brunnea Chen & Norphanphoun, in Phookamsak et al., Fungal diversity 95:1–273
(2019)
Notes: Neobambusicola brunnea (MFLU 18–1393) was
described based on phylogenetic analysis of a combined
LSU and ITS dataset (Phookamsak et al. 2019). The strain
formed a clade with the type species of Neobambusicola
strelitziae (CBS 138869) with moderate support (87% ML).
In our study, the phylogenetic analysis based on the combined LSU, SSU ITS, and tef1 sequence data showed different topology. The strain formed a separate clade from N.
strelitziae (CBS 138869) and clustered with our new species
Anthosulcatispora subglobosa with strong support (100%
ML/1.00 BYPP, Fig. 72). Therefore, Neobambusicola brunnea is transferred to a new genus.
Hosts: Dead stem of herbage—(Phookamsak et al. 2019).
Distribution: China—(Phookamsak et al. 2019).
Anthosulcatispora subglobosa Phukhams. & K.D. Hyde,
sp. nov.
Index Fungorum number: IF557203; Facesoffungi number: FoF 07341, Fig. 73.
Etymology: Refers to the subglobose conidiomata.
Holotype: MFLU 17–1473.
Saprobic on dried stems of Clematis subumbellata.
Sexual morph: Undetermined. Asexual morph: Conidiomata 160–210 × 265–345 μm ( x̄ = 178 × 291 μm, n = 5),
pycnidial, solitary, unilocular, scattered, shiny, immersed
or erumpent, flattened base, subglobose to compressed,
coriaceous, dark brown to reddish brown, lacking ostioles. Pycnidial wall 18–32 μm wide, thick, multilayered,
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composed of 10–14 brown layers of textura angularis,
inner layer subhyaline, lining bearing conidiogenous cells.
Conidiophores reduced to conidiogenous cells. Conidiogenous cells 5–17(–30) × 2–3.5 μm ( x̄ = 13 × 2.5 μm, n = 30),
enteroblastic, phialidic, determinate, elongated cylindrical
or truncate, smooth-walled, hyaline. Conidia 7–10 × 3–4 μm
( x̄ = 8 × 4 μm, n = 50), oblong, rounded ends, hyaline, aseptate, with 1–3 guttules in each cell, smooth-walled, with
mucilaginous sheath.
Cultural characters: Colonies on MEA reaching 30 mm
diam. after 4 weeks at 25 °C. Cultures from above, cream at
the centre, radiating, dense, circular, edge lobate, umbonate,
papillate, fairly fluffy, covered with white aerial mycelium,
black oil drops produced on the surface of cultures; reverse
dark brown at the centre, faintly zonate, white at the edge.
Material examination: Thailand, Phayao Province, Phu
Sang District, dead stems of Clematis subumbellata, 20
March 2017, C. Phukhamsakda, CMTH09 (MFLU 17–1473,
holotype); ex-type living culture, MFLUCC 17–2065.
Host: Clematis subumbellata—(This study).
Distribution: Thailand—(This study).
GenBank accession numbers: LSU: MT214592; SSU:
MT226705; ITS: MT310636; tef1: MT394649; rpb2:
MT394706.
Notes: In a BLASTn search of GenBank, the closest
match for LSU sequence of Anthosulcatispora subglobosa
MFLUCC 17–2065 was Neobambusicola strelitziae (strain
CBS 138869), (NG_058125) with 97% similarity, while the
closest match for the ITS sequence of MFLUCC 17–2065
was Magnicamarosporium iriomotense (strain HHUF
30125), (NR_153445) with 89% similarity. Pairwise comparison of the ITS region showed that A. subglobosa differs from A. brunnea with 12% differences (76 base pairs
difference of 585 base pairs, with gaps). The collection
has oblong, hyaline, guttulate, and aseptate, with rounded
ends conidia but microconidia were not observed in culture
(Crous et al. 2014b; Rupcic et al. 2018). We also noted the
formation of oil droplets in culture as a notable character for
A. subglobosa (Fig. 73).
Strain MFLUCC 17–2065 was evaluated for potential
secondary metabolite production with Bacillus subtillis,
Escherichia coli, Mucor plumbeus and Schizosaccharomyces pombe astest organisms. The isolate showed moderate
growth inhibitory activities against Bacillus subtillis and
Mucor plumbeus, and is thus a suitable candidate for further evaluation.
Parasulcatispora Phukhams. & K.D. Hyde, gen. nov.
Index Fungorum number: IF557204; Facesoffungi number: FoF 01686, Fig. 74.
Etymology: Refers to the characteristic features similar
to Sulcatispora.
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Fig. 73 Anthosulcatispora subglobosa (MFLU 17–1473, holotype).
a Appearance of conidiomata on Clematis subumbellata. b Vertical
section through conidioma. c Section of conidioma wall. d–g Conid-
iogenous cells and conidia. h–j Conidia. k Culture characteristics on
MEA. Scale bars: b = 100 µm, c = 50 µm, d–j = 10 µm
Saprobic on dried stem of herbaceous plants. Sexual
morph: Ascomata solitary, gregarious, semi-immersed
to erumpent, subglobose to compressed, coriaceous, dark
brown to black, ostiolate. Ostioles central, filled with hyaline periphyses. Peridium thin, uniform of flattened cells
of textura angularis, thin-walled, cells towards the inside
lighter, inner layer composed of thin, hyaline, gelatinous
layer. Hamathecium composed of numerous, filamentous, branched, septate, anastomosing, trabeculate pseudoparaphyses. Asci 8-spored, bitunicate, fissitunicate,
cylindrical-clavate to clavate, asymmetric, with furcate pedicel, apically rounded, with an ocular chamber. Ascospores
biseriate or partially overlapping, broad fusiform, hyaline,
tapering towards the ends, 1-euseptate, constricted at the
septum, smooth-walled, with guttules in each cell, with
mucilaginous sheath. Asexual morph: Undetermined.
Type species: Parasulcatispora clematidis Phukhams. &
K.D. Hyde
Notes: Parasulcatispora is established as a monotypic
genus. In a BLASTn search of GenBank, the closest match
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Fig. 74 Parasulcatispora clematidis (MFLU 17–1490, holotype). a
Appearance of ascomata on host surface. b Close up of ascoma on
host substrate. c Vertical section through ascoma. d Ostiolar canal.
e Section of peridium. f Trabeculate pseudoparaphyses. g–h Asci.
i–l Ascospores. m Culture characteristics on MEA. Scale bars:
b = 200 µm, c = 100 µm, d = 20 µm, e–h = 50 µm, i–l = 10 µm
for LSU and ITS sequences of MFLUCC 17–2082 is Sulcatispora berchemiae (HHUF 29097, NG_059390) with
98% similarity, while the ITS sequence had 88% similarity to NR_153444. Based on the multi-gene (LSU, SSU,
ITS and tef1 regions) analyses, MFLUCC 17–2082 formed
a separate lineage within Sulcatisporaceae (Fig. 72). The
genus matches Sulcatisporaceae species in having immersed,
subglobose to hemisphaerical ascomata, short ostioles, trabeculate pseudoparaphyses (Liew et al. 2000), and fusiform,
hyaline, septate ascospores with mucilaginous appendages
(Tanaka et al. 2015). Parasulcatispora is similar to Sulcatispora but lacks a pseudoclypeus and has small flattened ascomata with narrower asci and ascospores. The asexual morph
failed to develop in culture. We introduce the new genus
based on morphology and phylogenetic evidence.
Parasulcatispora clematidis Phukhams. & K.D. Hyde, sp.
nov.
Index Fungorum number: IF557205; Facesoffungi number: FoF 07342, Fig. 74.
13
Etymology: The specific name “clematidis” refers to the
host substrate.
Holotype: MFLU 17–1490.
Saprobic on dead stems of Clematis fulvicoma.
Sexual morph: Ascomata 160–230 × 230–320 μm
( x̄ = 190 × 305 μm, n = 5), solitary, gregarious, semiimmersed to erumpent, subglobose to compressed, coriaceous, dark brown to black, only ostiole visible, ostiolate.
Ostioles 84–115 × 116–133 μm ( x̄ = 100 × 124 μm, n = 5),
shiny, central, smooth-walled, filled with hyaline periphyses.
Peridium 22–37 μm wide ( x̄ = 19 μm, n= 20), uniform, wider
at the apex, composed of 3–4 layers of somewhat flattened
cells of textura angularis, thin-walled, cells towards the
inside lighter, inner layer composed of thin hyaline gelatinous layer. Hamathecium composed of numerous, dense,
2–3 µm wide, filamentous, branched, septate, anastomosing, trabecular pseudoparaphyses, embedded in a gelatinous matrix. Asci 53–88 × 8–17 µm ( x̄ = 72 × 13 µm, n = 30),
8-spored, bitunicate, fissitunicate, cylindrical-clavate to
clavate, asymmetric, with furcate pedicel, apically rounded,
with an ocular chamber. Ascospores 16–21 × 4–6 µm
( x̄ = 17 × 5 µm, n = 40), biseriate or partially overlapping,
broad fusiform with acute ends, tapering towards the ends,
1-euseptate, constricted at the septum, upper cell broader
than lower cell, smooth-walled, with two guttules in each
cell, hyaline, with 4–7 µm wide mucilaginous appendages.
Asexual morph: Undetermined.
Culture characters: Colonies on MEA reaching 30 mm
diam. after 4 weeks at 25 °C. Culture from above, greyish
brown, covered with grey fluffy mycelia, dense, circular,
umbonate, dull, undulate, radially furrowed, yellow oil droplets formed in the middle of cultures; reverse grey, radiating
outwardly.
Material examined: Thailand, Chiang Rai Province, Mae
Sai District, dried stems of Clematis fulvicoma, 20 March
2017, C. Phukhamsakda, CMTH28 (MFLU 17–1490, holotype); ex-type living culture, MFLUCC 17–2082.
Host: Clematis fulvicoma—(This study).
Distribution: Thailand—(This study).
GenBank accession numbers: LSU: MT214593; ITS:
MT310637; tef1: MT394650.
Notes: Parasulcatispora clematidis can be distinguished
morphologically (Fig. 74) and is supported by the phylogenetic analyses of combined LSU, SSU, ITS and tef1 data
(Tanaka et al. 2015, Fig. 72). The isolate MFLUCC 17–2082
also produces oil droplets in culture, and was further evaluated for the possibility of bioactive secondary metabolite
production. The isolate MFLUCC 17–2082 inhibited the
growth of Bacillus subtillis and partially inhibited conidial
production of Mucor plumbeus, but did not reach significant
values (data not shown).
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Teichosporaceae Barr
Teichosporaceae is characterized by erumpent to superficial ascomata, that are obpyriform, obovoid, or globose,
with blunt ostioles filled with pseudoparaphyses or periphyses, and a multilayered peridium (Barr 2002; Jaklitsch
et al. 2016). The family was treated under Floricolaceae
(Thambugala et al. 2015; Wijayawardene et al. 2016), but
Jaklitsch et al. (2016) synonymized Floricolaceae under Teichosporaceae. Wijayawardene et al. (2018) accepted twelve
genera in Teichosporaceae. Based on molecular analyses and
morphological comparisons, four of our collections from
Clematis clustered in Teichosporaceae (Fig. 75).
Floricola Kohlm. & Volkm.-Kohlm.
Floricola was introduced with F. striata as the type species. This genus is characterized by immersed, ostiolate,
pycnidial conidiomata with a peridium of thick-walled cells
with 5–6 cell layers, phialidic conidiogenesis cells, and
septate, brown conidia (Ariyawansa et al. 2015a; Thambugala et al. 2015). The genus occurs on old inflorescences
(Kohlmeyer and Volkmann-Kohlmeyer 2000; Ariyawansa
et al. 2015a). We introduce F. clematidis on Clematis vitalba
from Italy (Fig. 76).
Floricola clematidis Phukhams., Camporesi & K.D. Hyde,
sp. nov.
Index Fungorum number: IF557180; Facesoffungi number: FoF 07343, Fig. 76.
Etymology: The specific name “clematidis” refers to the
host.
Holotype: MFLU 17–1535.
Saprobic on dead stems of Clematis vitalba. Sexual
morph: Undetermined. Asexual morph: Conidiomata
Fig. 75 Best scoring RAxML tree with a final likelihood value
of − 21485.053484 based on combined LSU, ITS, tef1 and rpb2
sequence data for Teichosporaceae. The topology and clade stability of the combined gene analyses was compared to the single gene
analyses. The tree is rooted with members of the Sporormiaceae.
Sixty-one strains were included in the combined gene sequence analyses which comprised 3719 characters (1084 characters for LSU, 626
characters for ITS, 924 characters for tef1 and 1085 characters for
rpb2, including gap regions). The tree from the maximum likelihood
analysis had similar topology to the Bayesian 50% majority-rule consensus phylogram. The matrix had 1574 distinct alignment patterns
with 42.64% undetermined characters and gaps. Estimated base frequencies were as follows; A = 0.240047, C = 0.261812, G = 0.279530,
T = 0.218610; substitution rates AC = 1.269781, AG = 3.374707,
AT = 1.851379, CG = 1.247283, CT = 9.170687, GT = 1.000000;
gamma distribution shape parameter α = 0.475238. In our analysis,
GTR + I+G model was used for each partition in Bayesian posterior
analysis. The species determined in this study are indicated in blue.
Bootstrap values (BS) greater than 50% BS (ML, left) and Bayesian
posterior probabilities (BYPP, right) greater than 0.90 are given at the
nodes. Hyphens (-) represent support values less than 50% BS/0.90
BYPP. Thick branches represent significant support values from all
analyses (BS ≥ 70%/BYPP ≥ 0.95)
◂
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Fungal Diversity (2020) 102:1–203
123
Teichosporaceae
Magnibotryascoma rubriostiolata C158X
Magnibotryascoma rubriostiolata TR5
91/0.97 Magnibotryascoma rubriostiolata TR7
Magnibotryascoma rubriostiolata MFLUCC 16-1366
74/0.99
Magnibotryascoma rubriostiolata MFLU 16-1247
62/0.99
Magnibotryascoma
Magnibotryascoma uniseriata ANM 909
86/1.00
Magnibotryascoma acaciae CPC 24801
Magnibotryascoma melanommoides MP5
--/0.99
Aurantiascoma minima ANM 60
Aurantiascoma minima ANM 933
71/-Aurantiascoma
Aurantiascoma minima GKM 169N
Aurantiascoma
minima
SMH
2448
75/0.99
Teichospora quercus CBS 143396
Teichospora austroafricana CBS 119330
61/0.99
Teichospora austroafricana CBS 122674
99/1.00
Teichospora proteae CBS 122675
100/1.00
80/1.00
Teichospora pusilla C140
Teichospora
84/0.99
Teichospora trabicola C160
--/0.99
Teichospora grandicipis CPC 1852
98/1.00
Teichospora grandicipis CPC 1853
98/1.00
Teichospora claviformis GKM 1210
85/0.98
Macrodiplodiopsis desmazieri MFLUCC 12-0088
Magnibotryascoma mali MFLUCC 17-0933
--/0.99
Teichospora nephelii CPC 27539
Misturatosphaeria aurantonotata ATCC 42522
100/1.00
61/0.93
Misturatosphaeria aurantonotata SMH 4330
Misturatosphaeria
71/0.98
Misturatosphaeria aurantonotata GKM 1238
100/1.00
Misturatosphaeria aurantonotata GKM 1280
73/-Misturatosphaeria sp. SMH 3747
100/1.00 Pseudoaurantiaascoma kenyense GKM 1195
Pseudoaurantiaascoma
Pseudoaurantiaascoma kenyense GKM L100N
100/1.00
Pseudoaurantiaascoma kenyense GKM 234N
Asymmetrispora mariae C159
Asymmetrispora mariae CBS 124079
69/---/0.98
Asymmetrispora mariae C139
100/0.99
Asymmetrispora
Asymmetrispora mariae C144
Asymmetrispora trabicola C134m
87/0.90
Asymmetrispora tennesseensis ANM 911
--/-Paulkirkia
Paulkirkia arundinis MFLU 13-0315
73/-Teichospora kingiae CPC 29104
Pseudomisturatosphaeria cruciformis SMH 5151 Pseudomisturatosphaeria
100/1.00
100/1.00 Floricola striata JK 5603K
100/1.00 79/1.00
Floricola striata JK 5678I
Floricola
Floricola festucae MFLU 17-0773
97/1.00
Floricola viticola MFLUCC 15-0039
100/1.00
Floricola clematidis MFLUCC 17–2182
Ramusculicola thailandica MFLUCC 10-0126
--/0.90
100/1.00
Ramusculicola thailandica MFLUCC 13-0284
Ramusculicola
100/1.00
Ramusculicola thailandica MFLUCC 16-1366
Ramusculicola clematidis MFLUCC 17-2146
98/0.99 Thyridaria macrostomoides GKM 1033
51/1.00
100/1.00
Thyridaria macrostomoides GKM 1159
100/1.00
Lignosphaeria fusispora MFLUCC 11-0377
Phaeoseptaceae
Phaeoseptum
aquaticum CBS 123113
100/1.00
--/1.00
Phaeoseptum terricola MFLUCC 10-0102
Melanomma pulvis-pyrius CBS 124080
Melanommataceae
Amorosia littoralis NN 6654
100/1.00
Amorosiaceae
Exosporium stylobatum CBS 160.30
70/1.00
Sporormia fimetaria UPS:Dissing Gr.81.194
Sporormiaceae
Preussia funiculata CBS 659.74
84/1.00
(Outgroup)
Westerdykella ornata CBS 379.55
0.03
13
124
Fig. 76 Floricola clematidis (MFLU 17–1535, holotype). a, b
Appearance of conidiomata on Clematis vitalba. c Vertical section
through conidioma. d Section of conidioma wall. e–h Conidiog-
13
Fungal Diversity (2020) 102:1–203
enous cells and conidia. i–k Conidia. l, m Culture characteristic on
MEA. Scale bars: a = 500 µm, b = 200 µm, c = 100 µm, d, k = 20 µm,
e–j = 5 µm, l, m = 25 mm
Fungal Diversity (2020) 102:1–203
119–135 × 133–166 μm ( x̄ = 127 × 146 μm, n = 10), pycnidial, solitary, aggregated, uniloculate, flat, semi-immersed,
covered by a pseudoclypeus, globose, coriaceous, thickwalled, dark brown to reddish brown, without ostioles. Conidiomatal wall 11–24 μm wide, uniform, outer layer composed 4–5 layers of reddish brown cells of textura angularis,
lined with a thin, hyaline layer bearing conidiogenous cells.
Conidiophores reduced to conidiogenous cells. Conidiogenous cells 4–12 × 2–4 μm ( x̄ = 7 × 3 μm, n = 30), enteroblastic, phialidic, determinate, discrete, truncate or doliiform,
hyaline. Conidia 13–21 × 5–7 μm ( x̄ = 17 × 6 μm, n = 50),
broad oblong or ellipsoid, slightly curved towards the ends,
ends rounded, bud scars or disjunctors present at the site of
attachment, hyaline when immature, reddish brown at maturity, with longitudinal striations visible even while attached
to conidiogenous cells, 3-euseptate, with prominent longitudinal striations, smooth-walled.
Culture characters: Colonies on MEA reaching 20 mm
diam. after 2 weeks at 25 °C. Cultures from above, cream to
orangish-white, dense, circular, umbonate, papillate, fluffy,
slightly radiating in the lower part; reverse orange at the
centre, light brown, radiating outwardly.
Material examined: Italy, Forlì-Cesena Province, CabelliSanta Sofia, dead aerial branch of Clematis vitalba, 23 January 2014, E. Camporesi, IT 1668 (MFLU 17–1535, holotype); ex-type living culture, MFLUCC 17–2182.
Host: Clematis vitalba—(This study).
Distribution: Italy—(This study).
GenBank accession numbers: LSU: MT214594; SSU:
MT226706; ITS: MT310638; tef1: MT394651.
Notes: Floricola clematidis constitutes a strongly supported clade (100% ML/1.00 BYPP) with F. viticola
(MFLUCC 15–0039) (Fig. 75). Floricola clematidis is
similar to F. viticola. Both species are similar in having
semi-immersed conidiomata, but F. clematidis has smaller
conidiomata (127 × 146 vs 237 × 177 μm) and larger, broad
oblong or ellipsoid conidia (17 × 6 vs 9 × 4 μm). Floricola
clematidis also has prominent, longitudinal striations on
the conidia (Ariyawansa et al. 2015a, Fig. 76). A BLASTn
search of GenBank showed ITS sequence had 99.26%
similarity to F. viticola (7 nucleotide difference in 626
nucleotides).
Magnibotryascoma Thambug. & K.D. Hyde
Magnabotrioscoma, typified by M. uniseriatum was
introduced to accommodate a separate lineage of Misturatosphaeria uniseriata in Teichosporaceae. Magnibotryascoma
is characterized by erumpent to superficial ascomata with
short ostioles, and fusiform to elliptical, septate, guttulate
ascospores. The asexual morph is pycnidial with aseptate
and brown conidia (Crous et al. 2015b; Jaklitsch et al. 2016).
We redefine the Teichosporaceae, based on multigene phylogenetic analyses, as four species formed a strongly supported
125
clade with Magnibotryascoma sensu stricto (Fig. 75). We
introduce the first report of Magnibotryascoma rubriostiolata on Clematis vitalba and document the characters of the
asexual morph (Fig. 77).
Magnibotryascoma rubriostiolata (Jaklitsch & Voglmayr)
Phukhams., E.B.G. Jones & K.D. Hyde, comb. nov. and
new host record
Index Fungorum number: IF557181; Facesoffungi number: FoF 07344, Fig. 77
Basionym: Teichospora rubriostiolata Jaklitsch & Voglmayr, Mycological Progress 15(31):13 (2016)
Saprobic on dead stems of Clematis vitalba. Sexual
morph: Described in Jaklitsch et al. 2016. Asexual morph:
Conidiomata 110–188 × 93–164 μm ( x̄ = 160 × 131 μm,
n = 5), pycnidial, solitary, aggregated, uniloculate, immersed,
globose to subglobose, coriaceous, dark brown to brown,
with papilla ostiolate. Ostioles 25 × 19 μm, central, papillate. Conidiomatal wall 15–29 μm wide, thick, multilayered,
outer layer composed of 6–7 layers of light brown to brown
cells of textura angularis, lined with a hyaline layer bearing
conidiogenous cells. Conidiophores reduced to conidiogenous cells. Conidiogenous cells 4–11 × 2–3 μm ( x̄ = 7 × 2 μm,
n = 30), enteroblastic, discrete, integrated, oblong, cylindrical, hyaline, arising from the inner layer of pycnidium wall.
Conidia 3–5 × 1.8–3 μm ( x̄ = 3.7 × 2.5 μm, n = 50), subglobose, oval, with guttule in each cell, hyaline when immature,
pale brown at maturity, aseptate, smooth-walled.
Culture characters: Colonies on MEA reaching 30 mm
diam. after 2 weeks at 25 °C. Cultures from above, greenish-brown, dense, circular, umbonate, papillate, fluffy, covered with white aerial mycelium, droplets formed at room
temperature; reverse orange at the centre, brown radiating
outwardly.
Material examined: UK, Carmarthenshire, Laugharne,
on dead stems of Clematis vitalba, 29 July 2016, E. B. G.
Jones, GJ 311 (MFLU 17–1539); living culture, MFLUCC
16–1366.
Hosts: Clematis vitalba, Ribes sanguineum, Robinia
pseudoacacia, Vaccinium myrtillus—(Jaklitsch et al. 2016;
this study).
Distribution: Belgium, Germany, Norway, UK (England)—(Jaklitsch et al. 2016; this study).
GenBank accession numbers: LSU: MT214595; ITS:
MT310639.
Notes: Isolate MFLUCC 16–1366 formed a strongly
supported clade (91% ML/0.97 BYPP) with four isolates
of Magnibotryascoma rubriostiolata (Fig. 75). Magnibotryascoma rubriostiolata was introduced as Teichospora
rubriostiolata by Jaklitsch et al. (2016) on bark and wood
of trees, shrubs, and vineyard poles distributed in Europe.
A collection on twigs of Ribes sanguinea was recorded as
coniothyrium-like, but without a full description, although
13
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Fungal Diversity (2020) 102:1–203
◂Fig. 77 Magnibotryascoma rubriostiolata (MFLU 17–1539). a,
b Appearance of conidiomata on Clematis vitalba. c Vertical section through conidiomata. d Section of conidioma wall. e–h Conidiogenous cells and conidia. i–k Conidia. l Germinating conidium.
m, n Culture characteristics on MEA. Scale bars: a, b = 200 µm,
c = 100 µm, d = 50 µm, e–l = 5 µm
Jaklitsch et al. (2016) illustrated the sexual morph. A comparison of the ITS and tef1 DNA sequences of our isolate
(MFLUCC 16–1366) with four isolates showed 100% similarity. This is the first record of M. rubriostiolata on Clematis with illustration of their asexual morph (Fig. 77). We also
provide sequence data and phylogenetic analyses (Fig. 75).
Ramusculicola Thambug. & K.D. Hyde
Ramusculicola is typified by R. thailandica Thambug. &
K.D. Hyde. The genus is characterized by semi-immersed to
partially erumpent, coriaceous ascomata, short pedicellate
asci, and fusiform to cylindrical, hyaline ascospores with
polar appendages with blunt ends, and a lateral pad-like
structure of sheath that surrounds the ascospores (Thambugala et al. 2015). Based on phylogenetic analyses our collection isolated from Clematis species from Thailand revealed
a new species, Ramusculicola clematidis (Fig. 75).
Ramusculicola clematidis Phukhams. & K.D. Hyde, sp.
nov.
Index Fungorum number: IF557183; Facesoffungi number: FoF 07345, Fig. 78.
Etymology: The specific name “clematidis” refers to the
host.
Holotype: MFLU 17–1503
Saprobic on dead stem of Clematis sikkimensis.
Sexual morph: Ascomata 155–205 × 135–200 μm
( x̄ = 175 × 160 µm, n = 5), solitary or scattered, uniloculate,
semi-immersed to erumpent, ampulliform to compressed
subglobose, coriaceous, black to dark brown, ostiolate. Ostioles 75–95 × 110–130 μm, central, composed of 6–10 layers
of textura prismatica, papillate, with variable walls, with
pore-like opening, filled with hyaline periphyses. Peridium
15–30 μm wide ( x̄ = 20 μm, n= 20), uniform, multilayered,
composed of 7(–9) layers of textura prismatica, cells towards
the inside lighter, somewhat flattened, inner layer composed
of thin hyaline gelatinous layer. Hamathecium composed
of numerous, dense, 1.5–3.5 µm ( x̄ = 2 µm, n = 30), wide,
filamentous, branched, septate, anastomosing, cellular
pseudoparaphyses, situated between and above the asci,
embedded in a gelatinous matrix. Asci 55–100 × 8–13 µm
( x̄ = 80 × 10 µm, n = 30), 8-spored, bitunicate, fissitunicate,
oblong to cylindrical-clavate, short pedicellate, apically
rounded, with an ocular chamber, arising from the basal
ascoma. Ascospores 20–30 × 4–8 µm ( x̄ = 25 × 6 µm, n = 50),
biseriate or partially overlapping, broad-fusiform, tapering
127
towards the ends, rounded at upper end, acute towards the
lower end, (1–)2(–3)-transversely euseptate, strongly constricted at the septa, with (1–)2 guttules in each cell, hyaline,
swollen near median septum, with 3–8 µm of sheath drawn
out at both ends, forming polar appendages with blunt ends,
with a lateral pad-like structure, up to 2 μm wide at the sides.
Asexual morph: Undetermined.
Culture characters: Colonies on MEA reaching 20 mm
diam. after 2 weeks at 25 °C. Culture from above, brownish grey, dark green in the middle, forming cream, fluffy
mycelium at the edge, dense, umbonate, raised with concave
edge, rough, dull, lobate, radially furrowed, slight brown
pigment diffused into the agar, oil droplets formed in the
middle of cultures; reverse dark brown radiating light brown
with white margin.
Material examined: Thailand, Chiang Rai Province, Mae
Sai District, dead stems of Clematis sikkimensis, 24 June
2017, C. Phukhamsakda & M. van de Bult, CMTHDT16
(MFLU 17–1503, holotype); ex-type living culture,
MFLUCC 17–2146.
Host: Clematis sikkimensis—(This study).
Distribution: Thailand—(This study).
GenBank accession numbers: LSU: MT214596; SSU:
MT226707; ITS: MT310640; tef1: MT394652; rpb2:
MT394707.
Notes: Based on phylogenetic analysis, Ramusculicola clematidis (MFLUCC 17–2146) clustered with R.
thailandica with strong support (100% ML/1.00 BYPP,
Fig. 77). Ramusculicola clematidis can be distinguished
by its broad-fusiform ascospores, tapering towards the
ends, rounded at the upper end and acute towards the
lower end, while R. thailandica has ascospores that are
acute at both ends (Thambugala et al. 2015; this study).
Comparison of 551 nucleotides (without gaps) in the ITS
region revealed 23 bp (4.1%) differences, while the tef1
showed 27/920 bp (3%) differences from R. thailandica,
respectively. Therefore, R. clematidis is introduced as a
new species (Fig. 78).
Ramusculicola thailandica Thambug. & K.D. Hyde, in
Fungal Diversity 74: 251 (2015), new host record
Index Fungorum number: IF551265; Facesoffungi number: FoF 01092, Fig. 79.
Saprobic on dead stem of Clematis sikkimensis. Sexual
morph: Ascomata 174–180 × 170–215 μm, solitary or scattered, uniloculate, semi-immersed to erumpent, ampulliform to compressed globose, coriaceous, dark brown
to black, ostiolate. Ostioles 43–62 × 79–120 μm, central,
thick, composed of 20 layers of textura prismatica, papillate, with variable walls, with pore-like opening, filled with
hyaline periphyses. Peridium 15–48 μm wide ( x̄ = 26 μm,
n= 20), uniform, multilayered, composed of 13(–15) layers of textura angularis, thick-walled cells lighter towards
13
128
Fig. 78 Ramusculicola clematidis (MFLU 17–1503, holotype). a
Appearance of ascomata on host surface. b Close up of ascoma on
host substrate. c Vertical section of ascoma. d Ostiolar canal. e Sec-
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Fungal Diversity (2020) 102:1–203
tion of peridium. f Pseudoparaphyses. g–i Asci. j–n Ascospores.
Scale bars: b = 200 µm, c = 100 µm, d = 50 µm, e–i = 20 µm, j–
n = 10 µm
Fungal Diversity (2020) 102:1–203
the inside, somewhat flattened, inner layer composed of
thin hyaline gelatinous layer. Hamathecium composed of
numerous, dense, 1.6–3 µm wide, filamentous, branched,
septate, cellular pseudoparaphyses, situated between and
above the asci, embedded in a gelatinous matrix. Asci
60–91 × 8–13 µm ( x̄ = 74 × 10 µm, n = 30), 8-spored, bitunicate, fissitunicate, oblong to cylindrical-clavate, short
pedicellate, apically rounded, with an ocular chamber.
Ascospores 18–30 × 4–7 µm ( x̄ = 24 × 6 µm, n = 50), biseriate
or partially overlapping, broad-fusiform, tapering towards
the ends, ends acute, 1-transversely euseptate, strongly constricted at the septum, with 2–3(–4) guttules in each cells,
hyaline, swollen near median septum, with 3–6 µm of sheath
drawn out at both ends forming polar appendages, with a lateral pad-like structure, up to 2 μm wide at the sides. Asexual
morph: Undetermined.
Culture characters: Colonies on MEA reaching 40 mm
diam. after 4 weeks at 25 °C. Culture from above, brownish
grey, forming zonate greyish orange, fluffy mycelium at the
edge, dense, umbonate, raised with concave edge, rough,
dull, lobate, radially furrowed, brown pigment slightly diffused in the agar, oil droplets formed in the middle of culture; reverse dark brown, radiating, light orange outwardly.
Material examined: Thailand, Chiang Rai Province, Mae
Sai District, dead stems of Clematis sikkimensis, 24 June
2017, C. Phukhamsakda & M. van de Bult, CMTHDT13
(MFLU 17–1502); living culture, MFLUCC 17–2093.
Hosts: Fallen twig of deciduous tree, Clematis sikkimensis—(Thambugala et al. 2015; this study).
Distribution: Thailand—(Thambugala et al. 2015; this
study).
GenBank accession numbers: LSU: MT214597; SSU:
MT226708; ITS: MT310641; tef1: MT394653; rpb2:
MT394708.
Notes: Ramusculicola thailandica was first recorded by
Thambugala et al. (2015). The comparison of morphological characters and pairwise comparisons in the ITS and tef1
regions confirm the similarity of MFLUCC 17–2093 with
collections reported in Thambugala et al. (2015). This is the
first record of R. thailandica on Clematis species (Fig. 79).
We also provide sequence data and phylogenetic analyses
(Fig. 75).
Thyridariaceae Tian & K.D. Hyde
We follow Hyde et al. (2013) and Tibpromma et al.
(2017) to treat Thyridariaceae as a separate family in Pleosporales. Seven genera are listed in Thyridariaceae (Hyde
et al. 2013; Tibpromma et al. 2017; Devadatha et al. 2018;
Wanasinghe et al. 2018; Mapook et al. 2020). The phylogenetic tree, based on combined LSU, ITS, tef1 and rpb2
sequence data is presented in Fig. 80.
129
Parathyridaria Jaklitsch & H. Voglmayr
Parathyridaria is typified with P. ramulicola. Parathyridaria is characterized by immersed and globose ascomata,
a pseudoparenchymatous peridium, papilla with or without
orange colouration and fusoid, septate, hyaline to greyish
brown ascospores, with a pycnidial asexual morph (Jaklitsch and Voglmayr 2016; Wanasinghe et al. 2018). The
concatenated LSU, ITS, tef1, and rpb2 dataset showed that
fungal collections from Clematis species grouped with the
type strain of Parathyridaria with moderate support (51%
ML/1.00 BYPP, Fig. 80). The new species Parathyridaria
clematidis, P. serratifoliae and P. virginiana are introduced
from Clematis collections from the European continent.
Parathyridaria clematidis Phukhams., Camporesi & K.D.
Hyde, sp. nov.
Index Fungorum number: IF557206; Facesoffungi number: FoF 07347, Fig. 81.
Etymology: The epithet refers to the host plant, Clematis.
Holotype: MFLU 16–0061
Saprobic on dead stems of Clematis. Sexual morph: Ascomata 180–310 × 205–330 μm ( x̄ = 253 × 263 μm, n = 15), on
the surface of the host, solitary, gregarious, immersed, only
orange ostioles are visible, globose to depressed-globose,
coriaceous, black to dark brown, rough-walled, papillate, ostiolate. Ostioles 67–107 × 122–183 μm, central, black to reddish
brown, papillate, opening by a pore, filled with periphyses,
orange around pore. Peridium 17–33(–50 μm at apex) wide,
multilayered, outer layer composed of 7–10 layers of dark
brown to light brown cells of textura angularis, heavily pigmented at outer layer with thin, hyaline inner layer. Hamathecium composed of numerous, dense, 0.5–1.3 μm wide
( x̄ = 1 μm, n = 60), filiform, branched, septate, trabeculate
pseudoparaphyses. Asci 77–112 × 12–15 μm ( x̄ = 93 × 14 μm,
n = 20), (4–)8-spored, bitunicate, fissitunicate, broad fusiform,
clavate, apically rounded with an ocular chamber. Ascospores
16–24 × 5–8 μm ( x̄ = 20 × 6 μm, n = 50), biseriate, partial overlapping, oval to broad fusiform, sometimes inequilateral, with
rounded ends, hyaline, 1(− 2)-septate, constricted at septa,
cell above median septum enlarged, granulate in each cell,
rough-walled, with mucilaginous sheath. Asexual morph:
Coelomycetous, pycnidia produced on mycelium in MEA.
Conidiomata 135–170 × 102–195 μm diam., pycnidial, dark
brown to black, covered by dense vegetative hyphae, superficial, uniloculate, solitary to scattered, globose. Conidiomatal
wall 15–17 μm wide, thin, black to brown, with cells of
textura angularis. Conidiophores reduced to conidiogenous
cells. Conidiogenous cells 4–8(− 18) × 2–6 μm ( x̄ = 7 × 3 μm,
n = 30), enteroblastic, annellidic, sometimes a sympodial.
Conidia 5–7 × 3–5 μm ( x̄ = 6 × 4 μm, n = 100), ellipsoid, pale
brown, aseptate, guttulate, smooth-walled.
Culture characters: Colonies on MEA reaching 30 mm
diam. after 4 weeks at 25 °C. Culture from above, cream
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◂Fig. 79 Ramusculicola thailandica (MFLU 17–1502). a Appearance
of ascomata on host surface. b Close up of ascoma on host substrate.
c Vertical section of ascoma. d Ostiolar canal. e Section of peridium.
f Pseudoparaphyses. g–i Asci. j–m Ascospores. n Culture characters on MEA. Scale bars: b = 200 µm, c = 100 µm, d, g–i = 50 µm,
e–f = 20 µm, j–m = 10 µm
with lilac in the middle, white edge, dense, radially striated
with lobate edge, flattened, umbonate, fluffy; reverse cream.
Asexual morph formed in culture similar to those occurring
on the host substrate.
Material examined: Italy, Forlì-Cesena Province, near
Meldola, on dead aerial stems of Clematis vitalba, 27
December 2015, E. Camporesi, IT 2759 (MFLU 16–0061,
holotype); ex-type living culture, MFLUCC 17–2185; Belgium, Flemish Brabant, Meise Botanic Garden, Bouchout
Domain, on dead stems of C. viticella, 13 June 2017, D.
Ertz & C. Gerstmans, BRCV6 (MFLU 17–1518, paratype);
ex-paratype culture, MFLUCC 17–2160; ibid., BRCV3
(MFLU 17–1515, paratype); ex-paratype culture, MFLUCC
17–2157; on dead stems of C. serratifolia, 13 June 2017, D.
Ertz & C. Gerstmans, BRCS3 (MFLU 17–1511, paratype);
ex-paratype culture, MFLUCC 17–2154.
Hosts: Clematis serratifolia, C. vitalba, C. viticella—
(This study).
Distribution: Belgium, Italy—(This study).
GenBank accession numbers: MFLUCC 17–2185:
LSU: MT214598; SSU: MT226709; ITS: MT310642;
tef1: MT394654; rpb2: MT394709. MFLUCC 17–2160:
LSU: MT214599; SSU: MT226710; ITS: MT310643;
tef1: MT394655; rpb2: MT394710. MFLUCC 17–2157:
LSU: MT214600; SSU: MT226711; ITS: MT310644;
tef1: MT394656; rpb2: MT394711. MFLUCC 17–2154:
LSU: MT214601; SSU: MT226712; ITS: MT310645; tef1:
MT394657; rpb2: MT394712.
Notes: Four isolates of Parathyridaria clematidis were
recovered from Clematis species collected from Europe.
Morphological characters such as immersed ascomata, an
orange ostiolar canal, trabeculate pseudoparaphyses and
fusoid ascospores matches the concept of Parathyridaria
(Jaklitsch and Voglmayr 2016). Parathyridaria clematidis
is similar to P. robiniae in having hyaline ascospores, however, P. robiniae has larger ascomata (450–470 × 255–270
vs 182–310 × 205–329 μm, Table 4) with one septate
ascospores (Tibpromma et al. 2017). Our four isolates of P.
clematidis grouped together with strong statistical support
(96% ML/1.00 BYPP, Fig. 80). A comparison of the tef1
region of P. robiniae (MFLUCC 14–1119) with our new
strains revealed seven base pair differences, therefore, we
identify our isolates as a new species, Parathyridaria clematidis (Fig. 81).
131
Parathyridaria serratifoliae Phukhams., Ertz, Gerstmans &
K.D. Hyde, sp. nov.
Index Fungorum number: IF557208, Facesoffungi number: FoF 07348, Fig. 82.
Etymology: The species epithet refers to a species of the
host species Clematis serratifolia.
Holotype: MFLU 17–1512
Saprobic on dead stems of Clematis serratifolia. Sexual morph: Ascomata 288–342 × 235–301 μm
( x̄ = 322 × 265 μm, n = 10), on the surface of the host,
solitary, gregarious, immersed, orange ostioles visible,
globose to depressed-globose, coriaceous, dark brown to
brown, rough-walled, papillate, ostioles central. Ostioles
72–103 × 106–142 μm ( x̄ = 83 × 124 μm, n = 5), central,
black to reddish brown, papillate, opening by a pore, filled
with periphyses. Peridium 17–30(–50 μm at apex) wide,
outer layer composed of 5–7 layers of reddish brown cells
of textura angularis, heavily pigmented at outer layer, the
inner layer composed of 7 layers of hyaline and thin-walled
cells. Hamathecium of dense, 0.9–1.5 μm wide ( x̄ = 1.3 μm,
n = 50), filiform, branched, septate, trabeculate pseudoparaphyses. Asci 97–145 × 12–19 μm ( x̄ = 110 × 17 μm,
n = 20), 8-spored, bitunicate, fissitunicate, broad fusiform,
cylindrical-clavate, apically rounded with ocular chamber.
Ascospores 20–38 × 5–9 μm ( x̄ = 24 × 7 μm, n = 50), biseriate, partially overlapping, ellipsoid to broad fusiform, sometimes inequilateral, with rounded ends, hyaline, (1–)3-septate, constricted at the septa, cell above median septum
enlarged, granulate in each cell, rough-walled, with mucilaginous sheath. Asexual morph: Undetermined.
Culture characters: Colonies on MEA reaching 30 mm
diam. after 4 weeks at 25 °C. Culture from above, cream,
brown in the middle, edge white, dense, flattened, umbonate,
floccose; reverse cream, thin, with flat parchment-like sheets.
Material examined: Belgium, Flemish Brabant, Meise
Botanic Garden, Bouchout Domain, on dead stems of Clematis serratifolia, 13 June 2017, D. Ertz & C. Gerstmans,
BRCS4 (MFLU 17–1512, holotype); ex-type living culture,
MFLUCC 17–2210.
Host: Clematis serratifolia—(This study).
Distribution: Belgium—(This study).
GenBank accession numbers: LSU: MT214602; SSU:
MT226713; ITS: MT310646; tef1: MT394658; rpb2:
MT394713.
Notes: In our phylogenetic analysis from LSU, ITS, tef1,
and rpb2 sequence data, Parathyridaria serratifoliae formed
a basal clade with P. clematidis and P. robiniae with strong
support (94% ML/1.00 BYPP, Fig. 80). Parathyridaria serratifoliae can be distinguished from P. clematidis and P.
robiniae based on spore dimensions and having 3-septate
ascospores (Table 4). To further support the establishment
of the new taxon as per the guidelines of Jeewon and Hyde
(2016), we checked the nucleotide differences. Within the
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Fungal Diversity (2020) 102:1–203
Parathyridaria clematidis MFLUCC 17-2160
60/1.00
97/1.00
ML/BYPP
Parathyridaria clematidis MFLUCC 17-2157
96/1.00
Parathyridaria clematidis MFLUCC 17-2154
Parathyridaria clematidis MFLUCC 17-2185
94/1.00
100/1.00 42/--
Parathyridaria serratifoliae MFLUCC 17-2210
Parathyridaria robiniae MFLUCC 14-1119
Parathyridaria philadelphi CBS 143432
100/1.00
Parathyridaria
Parathyridaria rosae MFLUCC 17-0800
100/1.00
Parathyridaria ramulicola MRR1
100/1.00
100/1.00
Parathyridaria ramulicola MF4
Parathyridaria percutanea CBS 868.95
Parathyridaria percutanea CBS 128203
100/1.00
--/0.99
Cycasicola leucaenae MFLUCC 17-0914
Cycasicola goaensis MFLUCC 17-0754
91/0.93
100/1.00
Liua muriformis KUMCC 18-0177
Liua
Liua muriformis KUN-HKAS 102241
100/1.00
100/1.00
Thyridariella mangrovei NFCCl 4213
Thyridariella mangrovei NFCCI4214
79/0.99
Thyridariella
Chromolaenomyces appendiculatus MFLUCC 17-1455
99/1.00
100/1.00
100/1.00
100/1.00
Cycasicola
Thyridariaceae
Parathyridaria virginiana MFLUCC 17-2163
100/1.00
51/1.00
Pseudothyridariella mahakashae NFCCl4215
Pseudothyridariella
Thyridariella sp.VVS 2018a
Pseudothyridariella chromolaenae MFLUCC 17-1472
Thyridaria broussonetiae TB
100/1.00
99/--
Thyridaria broussonetiae TB1a
100/1.00
Thyridaria broussonetiae TB1
100/1.00
Thyridaria
Thyridaria broussonetiae TB2
“Thyridaria” acaciae CBS 138873
100/1.00
74/1.00
100/1.00
Ohleria modesta OM
Ohleria modesta MGC
Torula herbarum CBS 111855
Torula herbarum CPC 24114
Ohleriaceae
Torulaceae
(Outgroup)
Torula hollandica CBS 220.69
0.05
tef1 regions there were 10 base pair differences from P. robiniae. Therefore, P. serratifoliae on Clematis serratifolia is
introduced as a new species (Fig. 82).
13
Parathyridaria virginianae Phukhams., Ertz, Gerstmans &
K.D. Hyde, sp. nov.
Index Fungorum number: IF557209; Facesoffungi number: FoF 07349, Fig. 83.
Fungal Diversity (2020) 102:1–203
133
◂Fig. 80 The best scoring RAxML tree with a final likelihood value
GenBank accession numbers: LSU: MT214603; SSU:
MT226714; ITS: MT310647; tef1: MT394659; rpb2:
MT394714.
Notes: According to morphology and phylogenetic
analysis (Fig. 83), Parathyridaria virginianae (MFLUCC
17–2163) is closely related to P. rosae (MFLUCC 17–0800)
which was collected from Rosa sp. in the USA (Wanasinghe
et al. 2018). The asexual morph in Parathyridaria has pycnidia, with phialidic conidiogenous cells and ellipsoid, unicellular, hyaline to pale brown conidia (Crous et al. 2018),
which resembles our collection. Parathyridaria rosae was
reported as a sexual morph whilst our collection showed
only the asexual morph, thus the morphology could not
be compared. Pairwise comparison of ITS (including 5.8S
region) region showed five base pair differences (of 534 base
pairs), however, the tef1 and rpb2 regions are not available
for comparison. Phylogenetic analyses show strong support
for the two species (100% ML/1.00 BYPP, Fig. 80), thus, we
introduce a new species, P. virginianae.
Etymology: The epithet refers to Clematis virginiana.
Holotype: MFLU 17–1521
Saprobic on dead branches of Clematis virginiana. Sexual morph: Undetermined. Asexual morph: Conidiomata
145–207 × 204–259 μm ( x̄ = 175 × 230 μm, n = 5), pycnidial,
solitary, aggregated, uniloculate, immersed, covered by a
pseudoclypeus, globose to subglobose, coriaceous, thickwalled, dark brown to brown, papillate, ostiolate. Ostioles 31 × 50 μm, central. Conidiomatal wall 13–24 μm
wide, outer layer composed of 5–7 layers of brown to light
brown cells of textura angularis, lined with a hyaline layer
bearing conidiogenous cells. Conidiophores single, short,
densely aggregated, straight or flexuous, cylindrical, hyaline,
erect, septate, 1–3-septate, smooth. Conidiogenous cells
4–21 × 2–4 μm ( x̄ = 7 × 3 μm, n = 30), enteroblastic, phialidic, annellidic, hyaline, discrete, arising from the inner layer
of pycnidium wall. Conidia 3–5 × 1.8–3 μm ( x̄ = 4 × 2.5 μm,
n = 50), oval or subglobose, slightly curved, with 1(–2) guttules in each cell, hyaline when immature, pale brown at
maturity, aseptate, smooth-walled.
Culture characters: Colonies on MEA reaching 20 mm
diam. after 2 weeks at 25 °C. Cultures from above, white,
dense, circular, umbonate, covered with white aerial mycelium; reverse yellow at the central, cream.
Material examined: Belgium, Flemish Brabant, Meise
Botanic Garden, Bouchout Domain, on dead stems of Clematis virginiana L., 13 June 2017, D. Ertz & C. Gerstmans,
BRVir3 (MFLU 17–1521, holotype); ex-type living culture,
MFLUCC 17–2163.
Hosts: Clematis virginiana—(This study).
Distribution: Belgium—(This study).
Torulaceae Corda
Torulaceae, typified by Torula was introduced by Sturm
(1829) for an asexual morph with mononematous conidiophores (Seifert et al. 2011). Placement of Torulaceae was
verified in Crous et al. (2015a) with two accepted genera,
Dendryphion and Torula. Subsequently, Sporidesmioides,
Neotorula and Rostriconidium were introduced to the family based on modern taxonomic classification. Torulaceae
currently contains five genera which are asexual morphs
(Crous et al. 2015a; Hyde et al. 2016; Su et al. 2016; Tibpromma et al. 2018). In this study, a combined dataset of
LSU, ITS, tef1, and rpb2 sequence data was used to follow
the recent treatment in Su et al. (2016), with first report
of Dendryphion europaeum and Torula chromolaenae on
Clematis (Fig. 84).
of − 16490.277529 based on combined LSU, ITS, tef1 and rpb2
sequence data for Thyridariaceae. The tree is rooted with members
of the Torulaceae. Thirty-three strains were included in the combined
genes sequence analyses which comprised 3466 characters (890 characters for LSU, 534 characters for ITS, 964 characters for tef1 and
1078 characters for rpb2, including gap regions). The topology and
clade stability of the combined gene analyses was compared to the
single gene analyses. The tree from the maximum likelihood analysis had similar topology to the Bayesian 50% majority-rule consensus phylogram. The matrix had 1267 distinct alignment patterns, with
27.03% of undetermined characters and gaps. Estimated base frequencies were as follows; A = 0.243395, C = 0.269803, G = 0.270965,
T = 0.215837; substitution rates AC = 1.430051, AG = 3.703624,
AT = 1.723186, CG = 1.129553, CT = 9.368763, GT = 1.000000;
gamma distribution shape parameter α = 0.456291. In our analysis,
GTR + I + G model was used for each partition in Bayesian posterior
analysis. The species determined in this study are indicated in blue.
Bootstrap values (BS) greater than 50% BS (ML, left) and Bayesian
posterior probabilities (BYPP, right) greater than 0.90 are given at the
nodes. Hyphens (-) represent support values less than 50% BS/0.90
BYPP. Thick branches represent significant support values from all
analyses (BS ≥ 70%/BYPP ≥ 0.95)
Dendryphion Wallr.
Dendryphion (typified with D. comosum) is commonly
associated with dead stems of herbaceous plants and decaying wood, but also recorded from freshwater (Ellis 1971; Su
et al. 2016). The genus is distinguishable from other members in Torulaceae in having apically branched, polytretic or
enteroblastic conidiophores with dark pores, septate, brown
conidia in chains with bud scars (Seifert et al. 2011; Crous
et al. 2014a; Su et al. 2016). Based on morphological characters and phylogenetic results, we report the first record
of Dendryphion europaeum on Clematis species (Fig. 85).
Dendryphion europaeum Crous & R.K. Schumacher, in
Persoonia 32: 243 (2014), new host records
Index Fungorum number: IF808927; Facesoffungi number: FoF 07346, Fig. 85.
13
134
Fig. 81 Parathyridaria clematidis (MFLU 16–0061, holotype). a,
b Appearance of ascomata on Clematis vitalba. c Vertical section
through ascoma. d Ostiolar canal. e Section of peridium. f Trabeculate pseudoparaphyses. g–i Asci. j-o Ascospores (i Arrow indicated
mucilaginous sheath). p Germinated ascospore. q Culture character-
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Fungal Diversity (2020) 102:1–203
istics on MEA. r Conidiomata forming on agar on rice straw media
after 8 weeks. s–x Conidiogenous cells and developing conidia
(t Conidiogenous cells). y Conidia. Scale bars: a, b, r = 500 µm,
c = 200 µm, d = 100 µm, e–i = 50 µm, j–o, t = 10 µm, s, u–y = 5 µm,
q = 10 cm
Fungal Diversity (2020) 102:1–203
135
Table 4 Synopsis of sexual morph of Parathyridaria species
Species
Parathyridaria
clematidis
MFLUCC
17–2185
P. serratifoliae
MFLUCC
17–2210
P. ramulicola
MRR1
P. robiniae
MFLUCC
14–1119
P. rosae
MFLUCC
17–0800
Ascomata (μm)
Asci (μm)
Ascospores
Size (μm)
Septa
Colour
Host
References
182–310 × 205–
329
77–112 × 12–15
16–24 × 5–8
1(−2)
Hyaline
Clematis sp.
This study
288–342 × 235–
301
97–145 × 12–19
20–38 × 5–9
(1–)3
Hyaline
Clematis serratifolia
This study
Ribes rubrum
and Sambucus
nigra
Jaklitsch and
Voglmayer
(2016)
Robinia pseudacacia
Tibpromma et al.
(2017)
Brown
14.0–16.2(– 3(− 4) trans(200–)260–400(– (67–)75–89(–
96) × 10.0–11.7
19.5) × 4.8– verse, some460) × (200–
times 1-vertical
5.6
)210–335(–
375)
450–470 × 255– 105–115 × 10–15 15–20 × 4–7 1
Hyaline
270
300–400 × 150–
200
80–100 × 12.5–16 18–24 × 6–8
Saprobic on dead stems of Clematis species. Sexual
morph: Undetermined. Asexual morph: Colonies on the
natural substrate effused, scattered, hairy, dark brown.
Mycelium semi-immersed to superficial, composed of pale
brown, septate hyphae. Conidiophores 198–310 × 9–19.6 μm
( x̄ = 240 × 13 μm, n = 20), macronematous, mononematous,
single or in groups of 2–4, branched at the apex, stipes
straight or flexuous, subcylindrical to cylindrical, erect,
septate, smooth, dark brown to reddish brown, 10–19-septate, with 3–5 primary branches, irregularly branched at the
upper parts, brown, smooth or verruculose. Conidiogenous
cells 10–16 × 6–7 μm ( x̄ = 15 × 6 μm, n = 20), monotretic or
polytretic, integrated, terminal on conidiophores, doliiform
to oblong, pale brown. Conidia 14–26(–28) × 2.5–8 μm ( x̄
= 26 × 7 μm, n = 20), phragmosporous, in branched chains,
acrogenous, cylindrical to oblong, 2–5 septa, deeply constricted at septa, dark brown to reddish brown, verrucose,
rounded ends, bud scars or disjunctors present at the site of
attachment, easily separating.
Culture characters: Colonies on MEA reaching 30 mm
diam. after 2 weeks at 25 °C. Cultures from above, cream to
orangish-white, medium, circular, umbonate, fluffy, slightly
radiating outwardly, reverse dark brown at the centre, cream,
radiating outwardly.
Material examined: Belgium, Flemish Brabant, Meise
Botanic Garden, Bouchout Domain, on dead stems of
Clematis virginiana, 13 June 2017, D. Ertz & C. Gerstmans, BRCVir2 (MFLU 17–1520); living culture, MFLUCC
17–2162; UK, Botley wood, Hampshire, on dead stems of
C. vitalba, 16 April 2016, E.B.G. Jones, GJ 265 (MFLU
17–1462); living culture, MFLUCC 16–1003.
1
Pale brown Rosa sp.
Wanasinghe et al.
(2018)
Hosts: Clematis virginiana, C. vitalba, Hedera helix,
Heracleum sphondylium—(Crous et al. 2014a; this study).
Distribution: Belgium Germany, Netherlands, UK (England)—(Crous et al. 2014a; this study).
GenBank accession numbers: MFLUCC 17–2162:
LSU: MT214604; SSU: MT226715; ITS: MT310648;
tef1: MT394660; rpb2: MT394715. MFLUCC 16–1003:
LSU: MT214605; SSU: MT226716; ITS: MT310649; tef1:
MT394661.
Notes: Dendryphion europaeum was described by Crous
et al. (2014a) from Hedera helix (Araliaceae) and Heracleum sphondylium (Apiaceae). A phylogram generated
from multi-locus phylogeny analysis of our collections on
Clematis is shown in Fig. 84. Isolates MFLUCC 16–1003
and MFLUCC 17–2162 formed a strongly supported clade
with the ex-type strain of D. europaeum (92% ML/1.00
BYPP). Based on morphological and geological comparisons, our isolates are identical to those reported by Crous
et al. (2014a). A pairwise comparison of the ITS region of
MFLUCC 16–1003 showed two nucleotide difference from
the other isolates that clustered in the same clade (2/497
base pairs, 0.4%). According to the recommendation for new
taxon establishment proposed by Jeewon and Hyde (2016),
this is regarded as not statistically significant. Therefore, we
name our new isolates as new host records (Fig. 85).
Torula Pers.
Torula is typified by T. herbarum (= T. monilis) (Scott
et al. 2007). Torula species are characterized by asexual
morph characters having monoblastic or polyblastic, intact
or cupulate conidiogenous cells, and acropetal chains of dark
phragmoconidia (Persoon 1795; Scott et al. 2007). More
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Fungal Diversity (2020) 102:1–203
Fig. 82 Parathyridaria serratifoliae (MFLU 17–1512, holotype). a,
b Appearance of ascomata on Clematis serratifolia. c Vertical section
through ascoma. d Ostiolar canal. e Section of peridium. f Trabec-
ulate pseudoparaphyses. g, h Asci. i–l Ascospores. m Ascospore in
10% Indian ink. Scale bars: b, c = 100 µm, d–h = 50 µm, i–l = 10 µm,
m = 20 µm
than 500 epithets are listed in Index Fungorum (2020). We
report T. chromolaenae from Clematis fulvicoma in Thailand
(Fig. 86).
Index Fungorum number: IF819536; Facesoffungi number: FOF 02713, Fig. 86.
Saprobic on dried stems of Clematis fulvicoma. Sexual
morph: Undetermined. Asexual morph: Colonies on the
natural substrate effuse, scattered, hairy, powdery, dark
brown to black. Mycelium 1.7 μm wide, immersed to superficial, initially hyaline, later pale brown, septate, branched
Torula chromolaenae Li, Phookamsak, Mapook & K.D.
Hyde, in Mycological Progress 16 (4): 454 (2017), new
host record
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Fungal Diversity (2020) 102:1–203
137
Fig. 83 Parathyridaria virginianae (MFLU 17–1521, holotype). a
Appearance of conidiomata on Clematis virginiana. b Close up of
conidioma on host substrate. c Vertical section through conidioma. d
Ostiolar canal. e Section of conidioma wall. f–j Conidiogenous cells
and conidia. k–n Conidia. Scale bars: b = 200 µm, c = 100 µm, d,
e = 20 µm, f–n = 5 µm
hyphae. Conidiophores up to 5 μm long, micronematous, reduced to conidiogenous cells, with hyaline to
pale brown ampulliform supporting cell. Conidiogenous
cells 1.15–4 × 1.5–2 μm ( x̄ = 2.5 × 2.3 μm, n = 20) diam.,
mono to polyblastic, solitary, cylindrical, integrated, terminal on conidiophores, hyaline to pale brown. Conidia
9.5–14 × 4–6.5 μm ( x̄ = 11 × 5 μm, n = 50), phragmosporous, in short branched chains, acrogenous, dry, cylindrical,
2–3-septate, deeply constricted at septa, brown to reddish
brown, verrucose, rounded at both ends, bud scars or disjunctors present at the site of attachment, easily separating
into segments.
Culture characters: Colonies on MEA reaching 30 mm
diam. after 4 weeks at 25 °C. Cultures from above, olive
brown at the centre, faintly zonate edge, fluffy, grey, dense,
lobate, raised with concave edge, convex surface, downy,
brown radiating to the media; reverse dark brown at the centre, burnt umber radiating outwardly.
Material examined: Thailand, Nan Province, on dead
stems of Clematis fulvicoma, 20 March 2017, C. Phukhamsakda, CMTH23 (MFLU 17–1486); living culture,
MFLUCC 17–2078.
Hosts: Chromolaena odorata, Clematis fulvicoma, Pandanus tectorius—(Li et al. 2017; Tibpromma et al. 2018;
this study).
Distribution: China, Thailand—(Li et al. 2017; Tibpromma et al. 2018; this study).
GenBank accession numbers: LSU: MT214606; SSU:
MT226717; ITS: MT310650; tef1: MT394662; rpb2:
MT394716.
Notes: Torula chromolaenae was described from Chromolaena odorata collected from Thailand (Li et al. 2017).
Subsequently, Tibpromma et al. (2018) recorded the species
from Pandanus tectorius in China. Based on the phylogenetic analyses (Fig. 84), isolate MFLUCC 17–2078 formed
a strongly supported clade with T. chromolaenae KUMCC
16–0036 and KMUCC 17–0174. Our collection is similar
13
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Fungal Diversity (2020) 102:1–203
Torula ficus CBS 595.96
Torula ficus KUMCC 15-0428
97/1.00
Torula ficus MFLUCC 16-1259
Torula ficus MFLUCC 18-0112
90/0.99
100/1.00 Torula aquatica MFLUCC 16-1115
Torula aquatica DLUCC 0699
100/1.00
Torula gaodangensis MFLUCC 17-0234
69/0.95
Torula herbarum CBS 140066
Torula pluriseptata MFLUCC 14-0437
Torula
Torula
sp. CBS 246.57
99/1.00
Torula hollandica CBS 220.69
100/1.00
Torula chiangmaiensis KUMCC 16-0039
73/0.95
100/1.00 Torula polyseptata KUMCC 18-0131
Torula chromolaenae MFLUCC 17-2078
60/0.91
Torula chromolaenae KUMCC 16-0036
100/1.00 100/1.00 Torula chromolaenae KUMCC 17-0174
Torula breviconidiophora KUMCC 18-0130
100/1.00
Torula mackenziei MFLUCC 13-0839
Torula acaciae CPC 29737
93/1.00
Torula masonii CBS 245.57
Dendryphion europaeum MFLUCC 17-2162
67/Dendryphion europaeum MFLUCC 16-1003
92/1.00 Dendryphion europaeum CPC 22943
91/1.00
88/1.00
Dendryphion europaeum CPC 23231
Dendryphion aquatica MFLUCC 15-0257
--/0.98
Dendryphion
Dendryphion fluminicola DLUCC 0849
97/1.00
99/1.00
Dendryphion fluminicola KUMCC 15-0321
Dendryphion submersa MFLUCC 15-0271
77/0.97 Dendryphion submersum KUMCC15-0455
100/1.00
Dendryphion nanum MFLUCC 16-0975
100/1.00
67/-98/-- Dendryphion nanum HKAS 84010
Dendryphion nanum MFLUCC 17-1319
93/1.00
100/1.00
Dendryphion nanum DLUCC 0745
100/1.00 Neotorula submersa KUMCC 15-0280
Neotorula
100/1.00
Neotorula aquatica MFLUCC 15-0342
Rostriconidium aquaticum MFLUCC 16-1113
96/1.00
100/1.00
Rostriconidium aquaticum KUMCC 15-0297
Rostriconidium
Rostriconidium pandanicola KUMCC 17-0176
Sporidesmioides
thailandica KUMCC160012
100/1.00
Sporidesmioides
Sporidesmioides thailandica MFLUCC 13-0840
Ohleria modesta MGC
100/1.00
Ohleriaceae
Ohleria modesta CBS 141480
(Out group)
Torulaceae
82/0.97
83/0.99
0.04
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Fungal Diversity (2020) 102:1–203
◂Fig. 84 The best scoring RAxML tree with a final likelihood value
of − 12765.550785 based on combined LSU, ITS, tef1 and rpb2
sequence data for Torulaceae. The topology and clade stability of the
combined gene analyses was compared with the single gene analyses. The tree is rooted with members of the Ohleriaceae. Fourty-two
strains were included in the combined genes sequence analyses which
comprised 3000 characters (808 characters for LSU, 497 characters
for ITS, 854 characters for tef1 and 841 characters for rpb2, including gap regions). The tree from the maximum likelihood analysis had
similar topology to the Bayesian 50% majority-rule consensus phylogram. The matrix had 907 distinct alignment patterns, with 29.34% of
undetermined characters and gaps. Estimated base frequencies were
as follows; A = 0.240459, C = 0.270154, G = 0.275197, T = 0.214190;
substitution rates AC = 2.096044, AG = 4.720144, AT = 2.201665,
CG = 1.169889, CT = 12.100709, GT = 1.000000; gamma distribution shape parameter α = 0.542434. In our analysis, GTR + I + G
model was used for each partition in Bayesian posterior analysis.
The species determined in this study are indicated in blue. Bootstrap
values (BS) greater than 50% BS (ML, left) and Bayesian posterior
probabilities (BYPP, right) greater than 0.90 are given at the nodes.
Hyphens (-) represent support values less than 50% BS/0.90 BYPP.
Thick branches represent significant support values from all analyses
(BS ≥ 70%/BYPP ≥ 0.95)
to the type species of T. chromolaenae (KUMCC 16-0036)
but differ in size of the conidia (23 × 8 vs 11 × 5 μm) (Tibpromma et al. 2018). A pairwise comparison of DNA
sequences of ITS and tef1 regions of our isolate (MFLUCC
17–2078) with T. chromolaenae strains (KUMCC 16–0036
and KMUCC 17–0174) do not show significant differences.
Therefore, we name our collection as a new host record.
Isolate MFLUCC 17–2078 was further evaluated for bioactive secondary metabolite production. The strain inhibited
Bacillus subtillis and partially inhibited conidia production
of Mucor plumbeus, but did not reach significant values
(data not shown).
Dothideomycetes, family incertae sedis
Dyfrolomycetales Pang, Hyde & E.B.G. Jones
We follow the treatment by Hyde et al. (2013) and
Wijayawardene et al. (2017).
Pleurotremataceae Watson
Pleurotremataceae was introduced for Pleurotrema polysemum, a species lacking fissitunicate dehiscence asci in
Sordariomycetes (Watson 1929; Barr 1994). The isotype
specimens of P. polysemum were re-examined by Maharachchikumbura et al. (2016). Based on morphological evidence,
P. polysemum has similar characters with species of Saccardoella and Dyfrolomyces in Dyfrolomycetaceae (Dothideomycetes). Subsequently, P. polysemum was excluded
from Sordariomycetes and transferred to Dyfrolomycetaceae
(Dothideomycetes). Dyfrolomycetaceae is currently synonymized under Pleurotremataceae (Maharachchikumbura
et al. 2016). Three genera accepted in Pleurotremataceae
are Dyfrolomyces, Melomastia and Pleurotrema (Wijayawardene et al. 2017).
139
Melomastia Nitschke ex Sacc.
Norphanphoun et al. (2017) verified the phylogenetic
placement of Melomastia. Melomastia is accommodated in
Pleurotremataceae with more than 30 epithets recorded in
Index Fungorum (2020). Based on combined LSU, SSU and
tef1 sequence data (Fig. 87), we introduce two novel species,
M. clematidis and M. fulvicoma, from Clematis species in
Thailand.
Melomastia clematidis Phukhams., & K.D. Hyde, sp. nov.
Index Fungorum number: IF557210; Facesoffungi number: FoF 07334, Fig. 88.
Etymology: Refers to the host genus, Clematis.
Holotype: MFLU 17–1500
Saprobic on dead stems of Clematis sikkimensis. Sexual morph: Ascomata 300–510 × 260–445 μm
( x̄ = 420 × 400 μm, n = 10), only ostioles visible at the
surface of host, solitary, gregarious, semi-immersed to
immersed, globose to compressed globose, carbonaceous,
dark brown to black, rough-walled, ostiolate. Ostioles
80–160 × 90–116 μm, central, oblong, carbonaceous, dark
brown to black, papillate, periphyses filling the ostiolar
canal. Peridium 24–50(–90 at apex) μm wide, outer layer
carbonaceous, composed of 10–15 layers of dark brown cells
of textura prismatica mixed with cells of textura angularis, inner layer comprising thin hyaline layers. Hamathecium composed of numerous, dense, 1.9–3 μm ( x̄ = 2.5 μm,
n = 40), filiform, unbranched, septate, cellular pseudoparaphyses. Asci 115–160 × 4–7 μm ( x̄ = 140 × 6 μm, n = 20),
8-spored, bitunicate, broad filiform, apically rounded with
simple and short pedicel, ocular chamber visible. Ascospores
13–20 × 3.8–5 μm ( x̄ = 15 × 4.7 μm, n = 50), uniseriate, partially overlapping, broad fusiform with acute ends, hyaline,
3-septate,constricted at septa, guttulate, smooth-walled, with
thin mucilaginous sheath. Asexual morph: Undetermined.
Culture characters: Colonies on MEA reaching 30 mm
diam. after 4 weeks at 25 °C. Culture from above, white,
radiating outwardly, with yellow ring in the middle, oil
droplets produced in the cultures, medium dense, circular, umbonate, dull, edge erose, downy, covered with fluffy
cream mycelium; reverse brown in the middle with orange
ring, dark yellow, radiating outwardly.
Material examined: Thailand, Chiang Rai Province, on
dead branches of Clematis sikkimensis, 24 June 2017, C.
Phukhamsakda & M. van de Bult, CMTHDT09 (MFLU
17–1500, holotype); ex-type living culture, MFLUCC
17–2092.
Host: Clematis sikkimensis—(This study).
Distribution: Thailand—(This study).
GenBank accession numbers: LSU: MT214607; SSU:
MT226718; ITS: MT310651; tef1: MT394663.
Notes: In the phylogenetic analysis (Fig. 87), Melomastia
clematidis formed a close relationship with M. fulvicoma
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Fungal Diversity (2020) 102:1–203
◂Fig. 85 Dendryphion europaeum (MFLU 17–1520). a–c Appear-
ance of conidiophores on Clematis. d, e Mononematous conidiophore. f, g Conidiogenous cells and conidia. h–o Conidia. p Germinated conidia. q, r Culture characters on MEA. Scale bars: b,
c = 200 μm, d, e = 100 μm, f, g = 50 μm, h–o = 10 μm
(MFLUCC 17–2083), M. italica (MFLUCC 15–0160)
and M. maolanensis (GZCC 16–0102). Morphologically,
our collection matched with the generic concept of Melomastia in having large ascomata with cylindrical, bitunicate asci and hyaline, symmetrical and septate ascospores
(Pang et al. 2013). Our collection is distinct in having large
ascomata with 3-septate ascospores. In a BLASTn search
of GenBank, the closest match for the LSU region of strain
MFLUCC 17–2092 is D. thailandica (MFLUCC 15–0945)
with 96.79% similarity (NG_059714). The closest match for
the tef1 region is M. maolanensis strain GZCC 16–0102 with
93.74% similarity (KY814762). Therefore, we introduce M.
clematidis as a new species.
Melomastia clematidis strain MFLUCC 17–2083 was
evaluated for secondary metabolite production. However,
the isolate did not show inhibitory activity against the tested
strains.
Melomastia fulvicomae Phukhams., & K.D. Hyde, sp. nov.
Index Fungorum number: IF557211; Facesoffungi number: FoF 07335, Fig. 89.Etymology: The epithet refers to the
species of the host substrate, Clematis fulvicoma.
Holotype: MFLU 17–1491
Saprobic on dead stems of Clematis fulvicoma. Sexual morph: Ascomata 275–375 × 130–250 μm diam.
( x̄ = 310 × 185 μm, n = 5), only ostioles visible on the surface
of host, solitary, gregarious, semi-immersed, globose, subcarbonaceous, black to rust brown, rough-walled, ostiolate.
Ostioles 140–180 × 90–110 μm, central, oblong, carbonaceous, dark brown to black, papillate, periphyses filling ostiole. Peridium 15–30(–40 μm at apex) wide, outer layer composed of 5–7 layers of black cells of textura prismatica, the
inner layer comprised of hyaline cells and thin. Hamathecium composed of numerous, 1.5–2.5 μm ( x̄ = 2 μm, n = 40),
dense, filiform, unbranched, septate, cellular pseudoparaphyses. Asci 70–90 × 4–6 μm ( x̄ = 85 × 5 μm, n = 30), 8-spored,
bitunicate, cylindrical, apically rounded, with simple
and short pedicel, ocular chamber visible when young.
Ascospores 9–15 × 3.5–5.5 μm ( x̄ = 13 × 4 μm, n = 50), uniseriate, partial overlapping, broad fusiform with rounded
ends, ends acute, hyaline, 2–3-septate, constricted at the
septa, with guttules in each cell, smooth-walled with mucilaginous sheath. Asexual morph: Undetermined.
Culture characters: Colonies on MEA reaching 30 mm
diam. after 4 weeks at 25 °C. Culture from above, yellowish
radiating outwardly, with yellow ring in the middle, medium
dense, circular in shape, umbonate, dull, edge erose, downy,
141
covered with fairly cream mycelium; reverse: brown in the
middle with orange ring, dark yellow radiating outwardly.
Material examined: Thailand, Chiang Rai Province, on
dead branches of Clematis fulvicoma, 20 March 2017, C.
Phukhamsakda, CMTH29 (MFLU 17–1491, holotype); extype living culture, MFLUCC 17–2083.
Hosts: Clematis fulvicoma—(This study).
Distribution: Thailand—(This study).
GenBank accession numbers: LSU: MT214608; SSU:
MT226719; ITS: MT310652; tef1: MT394664.
Notes: Melomastia fulvicomae (MFLUCC 17–2146) was
associated with Clematis fulvicoma in Thailand. Melomastia fulvicomae can be distinguished from M. clematidis by
its smaller ascomata (310 × 185 vs 420 × 400 μm) and asci
(85 × 5 vs 140 × 6 μm). The ascospores of M. fulvicomae
are 2-septate, while those of M. clematidis are 3-septate
(Fig. 89). Multi-gene analysis showed that M. fulvicoma
(MFLUCC 17–2083) clustered with other Melomastia species with moderate statistical support (67% ML/1.00 BYPP,
Fig. 87).
The new strain was evaluated for secondary metabolite production with Bacillus subtillis, Escherichia coli,
Mucor plumbeus and Schizosaccharomyces pombe. Isolate
MFLUCC 17–2083 had moderate growth inhibitory activity
against Bacillus subtillis and partial development inhibition
against Mucor plumbeus. This strain is a suitable candidate
for further evaluation.
Class Lecanoromycetes Erikss. & K. Winka
Subclass Ostropomycetidae Reeb, Lutzoni & Cl. Roux
Ostropales Nannf.
Taxa of Ostropales are highly diverse and contain
lichenized, lichenicolous and non-lichenized clades of fungi.
Ascomata are apothecial or perithecial (Nannfeldt 1932;
Lumbsch et al. 2007).
Stictidaceae Fr. [as ‘Stictei’]
Taxa of Stictidaceae are parasitic, lichenized or lichenicolous and characterized by crustose thalli with chlorococcoid photobionts, apothecial ascomata or perithecia, filiform
unbranched paraphyses, J-, cylindrical asci and ellipsoid to
filiform ascospores and, sometimes formed by fragmentation
(Crous et al. 2017).
Fitzroyomyces Crous
Fitzroyomyces is a monotypic genus (Fig. 90). To date,
sexual morphs have not been recorded, however, we found
the sexual morph on Clematis. Asexual morphs are characterised by pycnidial conidiomata (Crous et al. 2017).
Fitzroyomyces cyperacearum Crous, Persoonia 39:389
(2017), new host record
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◂Fig. 86 Torula chromolaenae (MFLU 17–1486). a, b Appearance of
conidiophores on Clematis fulvicoma. c, d Conidiophores and conidia
attached to conidiogenous cells. e–j Conidia. k Culture characters on
MEA. Scale bars: c–j = 5 μm
Index Fungorum number: IF 823923; Facesoffungi number: FoF 05807, Fig. 91.
Saprobic on dead stem of Clematis subumbellata.
Sexual morph: Apothecia 201–260 × 210–310 μm
( x̄ = 230 × 260 μm, n = 10), arising singly or in small groups,
sessile, immersed in substrate, under the clypeus, cupulate.
Hypothecium convex. Disc 270–370 μm wide ( x̄ = 320 μm,
n = 10), whitish to cream. Margins white. Hymenium hyaline,
enclosed in a thick gelatinous matrix. Epithecium absent.
Excipulum 17–70 µm wide, composed of cells of textura
intricata. Paraphyses 1.3–3 µm wide ( x̄ = 2 μm, n = 50) at
the apex, numerous, filiform, aseptate, unbranched. Asci
110–150 × 10–20 µm ( x̄ = 130 × 12 μm, n = 10), 8-spored,
long, broad cylindrical, rounded at the apex, short sessile.
Ascospores 100–145 × 2.5–3.5 µm, fasciculate, spiraled
in ascus, thread-like, filiform, ends rounded, 17–21 transversely euseptate, slightly constricted at the septa, hyaline.
Asexual morph: Undetermined.
Culture characters: Colonies on MEA reaching 20 mm
diam. after 4 weeks at 25 °C. Culture from above, brown,
radiating, yellowish towards the edge, erumpent, spreading,
surface folded, margins lobate, dense, circular, flat, dull, fimbriate, radially furrowed, and slightly covered with white
aerial mycelia; reverse pale orange, with radiating cream
mycelia.
Material examined: Thailand, Chiang Rai Province, on
dead stems of Clematis subumbellata, 20 March 2017, C.
Phukhamsakda, CMTH16 (MFLU 17–1480); living culture,
MFLUCC 17–2072.
Hosts: woody stem, Epilobium (= Chamaenerion) angustifolium, Clematis subumbellata—(Crous et al. 2017; this
study).
Distribution: Australia, Thailand—(Crous et al. 2017;
this study).
GenBank accession numbers: LSU: MT214609; SSU:
MT226720; ITS: MT310653; tef1: MT394665.
Notes: Fitzroyomyces cyperacearum (MFLUCC 17–2072)
was found on the stem of Clematis subumbellata in Thailand. The strain formed a clade with F. cyperacearum from
Australia and the UK (Fig. 90). Isolate MFLUCC 17–2072
clustered with three Fitzroyomyces strains with strong support and the ITS data of our strain was 99% similar to the ITS
data of the type species of F. cyperacearum (CBS 143170).
Fitzroyomyces was introduced based on its pycnidial characters (Crous et al. 2017). However, the asexual morph was
not observed in our collection, therefore it could not be
compared to the asexual morph of the type strain. Taking
into consideration the genetic similarity and phylogenetic
143
results, we name our collection as the sexual morph of F.
cyperacearum. This is the first record of F. cyperacearum
from Thailand and on Clematis (Fig. 91).
Fitzroyomyces cyperacearum (MFLUCC 17–2072) was
evaluated for secondary metabolite production with Bacillus subtillis, Escherichia coli, Aspergillus niger and Schizosaccharomyces pombe. It showed weak inhibitory activities against Bacillus subtillis and completely inhibited the
development of Aspergillus niger with an inhibition zone
diameter of 17 mm. This strain is a suitable candidate for
further evaluation.
Neostictis Ekanayaka, Camporesi & K.D. Hyde, gen. nov.
Index Fungorum number: IF557307; Facesoffungi number: FoF 07338, Fig. 92.
Etymology: Name refers to the similarity to Stictis.
Saprobic on decaying wood material or herbaceous plant
in terrestrial habitats. Sexual morph: Apothecia arising singly or in small groups, sessile, immersed in substrate, under
the clypeus, cupulate. Hypothecium convex. Disc blackish.
Margins brownish. Hymenium hyaline, enclosed in a thick
gelatinous matrix. Excipulum composed of multi-layer of
cells of textura intricata. Paraphyses numerous, filiform,
aseptate. Asci 8-spored, short sessile, broad cylindrical to
oblong, rounded at the apex. Ascospores fasciculate, threadlike, spiraled, filiform, rounded ends, multi-septate, transversely eusepta, hyaline, sometimes breaking into small
fragments. Asexual morph: Undetermined.
Type species: Neostictis nigricans Ekanayaka,
Phukhams., Camporesi & K.D. Hyde
Notes: Our collection MFLU 17–1540, from Italy, formed
a well-supported clade close to Phacidiella podocarpi. Our
new species is morphologically similar to Fitzroyomyces
cyperacearum by having immersed apothecia, long cylindrical asci and filiform ascospores (Crous et al. 2017). However,
our new species differs by having blackish margins, smaller
disc and asci and ascospores (Fig. 92). The ascospores of our
collection failed to germinate and therefore, we were unable
to compare the asexual morph characters of our collection
with Fitzroyomyces cyperi and Phacidiella podocarpi.
Neostictis nigricans Ekanayaka, Phukhams., Camporesi &
K.D. Hyde, sp. nov.
Index Fungorum number: IF557308; Facesoffungi number: FoF 07339, Fig. 92.
Etymology: The epithet refers to the apothecial morphology. The disc and the margins of the apothecium is black.
Holotype: MFLU 17–1540
Saprobic on dead stem of Clematis vitalba. Sexual
morph: Apothecia 0.4–1 mm wide, arising singly or in
small groups, sessile, immersed in substrate, under the
clypeus, cupulate. Hypothecium convex. Disc blackish.
Margins blackish. Hymenium hyaline, enclosed in a thick
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Fungal Diversity (2020) 102:1–203
--
67/1.00
75/1.00
Melomastia maolanensis GZCC 16-0102
Melomastia
Melomastia fulvicoma MFLUCC 17-2083
Melomastia clematidis MFLUCC 17-2092
Dyfrolomyces thailandica MFLUCC 15-0945
97/1.00
75/1.00
Dyfrolomyces rhizophorae JK 5456A
Dyfrolomyces sinensis MFLU 17-0777
94/1.00
100/1.00
Dyfrolomyces
Dyfrolomyces thamplaensis MFLUCC 15-0635
73/1.00
Dyfrolomyces phetchaburiensis MFLUCC 15-0951
Dyfrolomyces chromolaenae MFLUCC 17-1434
100/1.00
98/1.00
Pleurotremataceae (=Dyfrolomycetaceae
(
Melomastia italica MFLUCC 15-0160
71/1.00
Dyfrolomyces tiomanensis NTOU3636
Acrospermum compressum M151
--/1.00
Acrospermum gramineum M152
Acrospermaceae
99/1.00
Acrospermum adeanum M133
Dendryphiopsis atra AFTOL-ID 273
100/1.00
Kirschsteiniothelia aethiops MFLUCC 15-0424
Kirschsteiniotheliaceae
(Outgroup)
0.02
Fig. 87 The Bayesian 50% majority-rule consensus phylogram based
on combined LSU, SSU and tef1 sequence data for Pleurotremataceae
and related taxa. The topology and clade stability of the combined
gene analyses was compared to the single gene analyses. The tree is
rooted with members of the Kirschsteiniotheliaceae. Sixteen strains
were included in the combined genes sequence analyses which comprised 3269 characters (1297 characters for LSU, 1049 characters for
SSU, 923 characters for tef1, including gap regions). The tree from
the maximum likelihood analysis had similar topology to the Bayesian analyses. The best scoring RAxML tree had a final likelihood
value of − 8012.999545. The matrix had 707 distinct alignment patterns, with 31.71% of undetermined characters and gaps. Estimated
13
base frequencies were as follows; A = 0.236800, C = 0.268407,
G = 0.273839, T = 0.220954; substitution rates AC = 0.626404,
AG = 2.767488, AT = 0.626269, CG = 1.143373, CT = 7.006679,
GT = 1.000000; gamma distribution shape parameter α = 0.740235.
In our analysis, GTR + I + G model was used for each partition in
Bayesian posterior analysis. The species determined in this study are
indicated in blue. Bootstrap values (BS) greater than 50% BS (ML,
left) and Bayesian posterior probabilities (BYPP, right) greater than
0.90 are given at the nodes. Hyphens (-) represent support values less
than 50% BS/0.90 BYPP. Thick branches represent significant support values from all analyses at family level (BS ≥ 70%/BYPP ≥ 0.95)
Fungal Diversity (2020) 102:1–203
145
Fig. 88 Melomastia clematidis (MFLU 17–1500, holotype). a, b
Appearance of ascomata on Clematis sikkimensis. c Vertical section
through ascoma. d Ostiolar canal. e Section of peridium. f Pseudo-
paraphyses. g–i Asci. j Ascospores. k Culture characteristics on
MEA. Scale bars: b, c = 200 µm, d–i = 50 µm, j = 10 µm
gelatinous matrix. Excipulum 25–35 µm wide, composed of
multi-layer cells of textura intricata. Paraphyses 1.3–2 µm
wide at the apex, numerous, filiform, aseptate, unbranched.
Asci 80–100 × 8–12 µm, 8-spored, short sessile, broad
cylindrical to oblong, rounded at the apex. Ascospores
35–50 × 2.5–3.5 µm, fasciculate, weakly spiral, thread-like,
filiform, ends rounded, multi-septate, approximately
32-transversely euseptate, hyaline, sometimes breaking into
small fragments. Asexual morph: Undetermined.
Material examined: Italy, Forlì-Cesena Province, Fiumicello di Premilcuore, Bouchout Domain, dead aerial branch
of Clematis vitalba, 18 January 2014, E. Camporesi, IT1655
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Fig. 89 Melomastia fulvicomae (MFLU 17–1491, holotype). a, b
Appearance of ascomata on Clematis fulvicoma. c Vertical section
through ascoma. d Ostiolar canal. e Section of peridium. f Pseudo-
paraphyses. g, h Asci. i-k Ascospores. l Germinated ascospore. m, n
Culture characteristics on MEA. Scale bars: b = 500 µm, c = 200 µm,
d, e = 50 µm, f–h = 20 µm, i–k = 10 µm
(MFLU 17–1540, holotype); ibid., 10 October 2013 (MFLU
16–0483 = MFLU 16–0595).
Host: Clematis vitalba—(This study).
Distribution: Italy—(This study).
GenBank accession numbers: LSU: MT214610; SSU:
MT226721; ITS: MT310654; tef1: MT394666.
Notes: See note under Neostictis.
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147
61
66
Fitzroyomyces cyperacearum GJ339B
Fitzroyomyces cyperacearum MFLUCC 17-2072
Fitzroyomyces cyperacearum GJ 317
97
Fitzroyomyces cyperacearum CBS 143170
54
Phacidiella podocarpi CBS 138904
100 Neostictis nigricans MFLU 17–1540
61
Neostictis nigricans MFLU 16–0483
84 Stictis confusum Wedin 7070
100 Stictis confusum
99
Conotrema populorum GG2353
Conotrema populorum GG2618
Carestiella socia GG2410
Phacidiella ucalypti CBS 120255
Stictis brunnescens GG2359
100 Schizoxylon albescens UM23
96
Schizoxylon albescens UM24
Schizoxylon albescens L560
Ostropa barbara SW071
Cryptodiscus rhopaloides EB100
96
Stictis urceolatum AFTOL-ID 96
92
Conotrema urceolatum LT21500
Robergea cubicularis
Glomerobolus gelineus AFTOL-ID 1349
Stictis radiata GG2449a
Acarosporina microspora AFTOL-ID 78
96
Cryptodiscus pini EB181
51
Cryptodiscus incolor S-F116574
Cryptodiscus cladoniicola RP160
58
Cryptodiscus epicladonia H Ahti 70348a
Absconditella rubra
100
1/4x
Absconditella lignicola EB211
57
69
Cryptodiscus tabularum CO205
97
Ingvariella bispora BCNLich 17183
87
Ingvariella bispora MALich 15288
Xyloschistes platytropa AFTOL-ID 4891
66
Geisleria sychnogonoides 70627
Absconditella sphagnorum M24
99
Micropeltis dendrophthoes MFLUCC 15-0599
63
Micropeltis zingiberacicola
100 Cyanodermella oleoligni CBS 140345
85
Cyanodermella oleoligni DTO 301-G1
Cyanodermella asteris 03HOR06-2-4
Lecanora contractula AFTOL-ID 877
Lecanoraceae
Stictidaceae
100
(Outgroup)
0.2
Fig. 90 The best scoring RAxML tree with a final likelihood value of
− 13518.083608 for ITS and LSU sequence data. The topology and
clade stability of the combined gene analyses was compared to the
single gene analyses. The tree is rooted with Lecanora contractula
(AFTOL-ID 877). The matrix had 958 distinct alignment patterns
with 48.65% undetermined characters and gaps. Estimated base frequencies were as follows; A = 0.252, C = 0.244, G = 0.276, T = 0.228;
substitution rates AC = 1.173341, AG = 1.808670, AT = 1.760640,
CG = 0.888202, CT = 5.267472, GT = 1.000000; gamma distribution
shape parameter α = 0.339348. Ex-type strains are in bold black and
the species determined in this study are indicated in blue. Bootstrap
values (BT) (over 50% BT) from maximum likelihood are given at
the nodes
Class Sordariomycetes Erikss. & K. Winka
We follow Hyde et al. (2020b) which is the latest treatment of this class.
We follow the treatment of the order of Hyde et al.
(2020b).
Subclass: Sordariomycetidae Erikss. & K. Winka
Chaetosphaeriales Huhndorf, Mill. & F.A. Fernández
Chaetosphaeriales, genera incertae sedis
Neoleptosporella Phukhams. & K.D. Hyde, gen. nov.
Index Fungorum number: IF557081; Facesoffungi number: FoF 07246, Fig. 94.
Etymology: The generic epithet refers to its similarity
with Leptosporella.
13
148
Fig. 91 Fitzroyomyces cyperacearum (MFLU 17–1480). a Appearance of apothecia on Clematis subumbellata. b Close up of apothecia on host substrate. c Vertical section through an apothecium. d
Excipulum layers. e Aseptate paraphyses. f, g Cylindrical asci. h-j
13
Fungal Diversity (2020) 102:1–203
Fusiform ascospores (j Ascospore in cotton blue). k Culture characteristics on MEA. Scale bars: b = 500 µm, c = 250 µm, d = 100 µm,
e–j = 50 µm
Fungal Diversity (2020) 102:1–203
Fig. 92 Neostictis nigricans (MFLU 17–1540, holotype). a Appearance of apothecia on Clematis vitalba. b, c Close up of apothecia
on substrate. d Vertical section of an apothecia. e Vertical section of the peridium. f Aseptate paraphyses. g–j Asci. k, l Needle-
149
like ascospores. m Ascospores released from an ascus. n Secondary ascospores. Scale bars: b = 500 µm, c = 200 µm, d = 300 µm,
e = 100 µm, f–j = 30 µm, k–n = 20 µm
13
150
Fungal Diversity (2020) 102:1–203
Dinemasporium nelloi MFLU 14-C0811
Rattania setulifera GUFCC 15501
Thozetella nivea MH863280
Menisporopsis theobromae MFLUCC 15-0055
94
Phialosporostilbe scutiformis MFLUCC 17-0227
Dictyochaeta siamensis MFLUCC 15-0614
Codinaea pini CBS 138866
98
Calvolachnella guaviyunis CBS 134695
Infundibulomyces oblongisporus BCC13400
Tainosphaeria siamensis MFLUCC 15-0607
Neopseudolachnella acutispora HHUF 29727
Pseudodinemasporium fabiforme HHUF 29716
74
Dendrophoma cytisporoides CBS 223.95
Brunneodinemasporium brasiliense CBS 112007
Pseudolachnea fraxini CBS 113701
Pseudolachnella longiciliata HHUF 27528
73
Dictyochaeta curvispora CBS 114070
Menispora tortuosa AFTOL-ID 278
Chaetosphaeriaceae
Multiguttulispora sympodialis MFLU 19-0218
Anacacumisporium appendiculatum HMAS 245593
78
Striatosphaeria codinaeophora SMH 1524
Sporoschisma hemipsilum MFLUCC 15-0615
100
100
Sporoschisma palauense MFLUCC 15-0616
Chloridium aquaticum HKAS 96226
100
Chloridium aseptatum MFLU 11-1051
Adautomilanezia caesalpiniae LAMIC010212
Lecythothecium duriligni CBS 101317
100
Pyrigemmula aurantiaca CBS 126743
Umbrinosphaeria caesariata CBS 102664
99
83
Exserticlava vasiformis AMA 450
Nawawia filiformis MFLUCC 16-0853
Zanclospora iberica CBS 130426
Eucalyptostroma eucalypti CPC 28764
Chaetosphaeria fuegiana ICMP 15153
Cryptophiale udagawae MFLUCC 18-0422
88
Cryptophialoidea fasciculata MFLUCC 17-2119
Conicomyces pseudotransvaalensis HHUF 29956
100
Rimaconus coronatus SMH5212
Rimaconus jamaicensis SMH4782
Erythromada lanciospora SMH1526
Neoleptosporella clematidis MFLUCC 17-2074
98
Synaptospora plumbea SMH3962
Ruzenia spermoides ANM163
100
Chaetosphaeriales
incertae sedis
Helminthosphaeriaceae
Helminthosphaeria hyphodermae SMH4192
Echinosphaeria canescens SMH4666
92
81
Helminthosphaeria clavariarum SMH4609
98
Hilberina caudata SMH1542
Endophragmiella dimorphospora FMR 12150
Neolinocarpon rachidis MFLUCC 15-0332
86
76
79
Neolinocarpon arengae MFLUCC 15-0323
Linocarpaceae
Neolinocarpon phayaoense MFLUCC 17- 0074
Linocarpon cocois MFLUCC 15-0812
93
89
Linocarpon arengae MFLUCC 15-0331
Linocarpon pandanicola HKUCC 4385
99
98
100
Leptosporella gregaria SMH 4290
Leptosporella gregaria SMH 4673
Leptosporella cocois MFLUCC 15-0816
100
100
Leptosporella arengae MFLUCC 15-0330
Leptosporella bambusae MFLUCC 12-0846
Leptosporellaceae
Caudatispora biapiculata SMH1873
Neonawawia malaysiana CBS 125544
Lasiosphaeriella nitida SMH1290
100
Gelasinospora tetrasperma CBS 178.34
Neurospora crassa MUCL 19026
0.2
13
Chaetosphaeriales
incertae sedis
Sordariales
(Outgroup)
Chaetosphaeriales
76
94
Sporoschisma longicatenatum MFLUCC 16-0180
Fungal Diversity (2020) 102:1–203
◂Fig. 93 The best scoring RAxML tree with a final likelihood value
of − 19621.398134 of LSU and ITS sequence data. The topology
and clade stability of the combined gene analyses was compared
to the single gene analyses. Sixty-five strains were included in the
combined genes sequence analyses which comprised 1655 characters (1060 characters for LSU and 595 characters for ITS, including
gap regions). Thistree includes selected taxa of Chaetosphaeriales
and rooted with Gelasinospora tetrasperma (CBS 178.34) and Neurospora crassa (MUCL 19026) in Sordariales. The matrix had 990
distinct alignment patterns, with 30.05% of undetermined characters
and gaps. Estimated base frequencies were as follows; A = 0.230333,
C = 0.267483, G = 0.312066, T = 0.190118; substitution rates
AC = 1.241984, AG = 1.768569, AT = 1.330285, CG = 1.057213,
CT = 5.757832, GT = 1.000000; gamma distribution shape parameter
α = 0.462145. Ex-type strains are in bold black and the species determined in this study is indicated in blue. Bootstrap values (over 70%
BT) from maximum likelihood are given at the nodes
Saprobic on stem of terrestrial herbaceous plants. Sexual
morph: Ascomata solitary, immersed, only black shiny ostioles visible, immersed beneath small clypeus, appear as disc
around the neck, coriaceous, subglobose to depressed globose, as raised blister-like areas, ostiolate. Ostioles central,
carbonaceous, black, filled with periphyses. Peridium outer
cells merging with the host epidermal cells, dark brown
to black cells of textura angularis. Paraphyses comprising numerous, hyaline, branched, septate. Asci 8-spored,
unitunicate, broad cylindrical, long-pedicellate, with a J-,
wedge-shaped, subapical ring. Ascospores fasciculate, fusiform, straight or curved, C-shaped or sigmoid, aseptate, ends
acute, without polar appendages, smooth-walled, with guttules in each cell. Asexual morph: Undetermined.
Type species: Neoleptosporella clematidis Phukhams.,
Konta & K.D. Hyde
Notes: Neoleptosporella is introduced for a taxon on
Clematis subumbellata and is distinguishable from Leptosporella in having immersed ascomata which are partial
carbonaceous at the apex, the lower part is coriaceous, subglobose to depressed globose, not flattened at the base and
without the cover of a pseudoclypeus. The apical part of
the asci is wedge-shaped, and J-, with fusiform, aseptate
ascospore with acute ends (Huhndorf and Miller 2011; Dai
et al. 2017; Konta et al. 2017; Hyde et al. 2020a). The strain
forms a lineage with three strains that have uncertain placement in Chaetosphaeriales, but received low statistical support (less than 75% ML, Fig. 93).
Neoleptosporella clematidis Phukhams., Konta & K.D.
Hyde, sp. nov.
Index Fungorum number: IF557060; Facesoffungi number: FoF 07246, Fig. 94.
Etymology: named after the host genus, Clematis.
Holotype: MFLU 17–1482.
Saprobic on dead branches of Clematis subumbellata. Sexual morph: Ascomata (including neck)
150–200 × 100–120 μm ( x̄ = 170 × 115 μm, n = 10), solitary,
151
immersed, only black shiny ostioles visible, immersed
beneath small clypeus, appearing as a disc around the neck,
coriaceous, subglobose to depressed globose, as raised blister-like areas, ostiolate. Ostioles central, 60 × 55 μm, carbonaceous, black, filled with periphyses. Peridium 5–10(–22)
μm wide, outer cells merging with the host epidermal cells,
composed of 3–4(–8 at apex) layers of dark brown to black
cells of textura angularis. Paraphyses of numerous, 6–16 µm
wide ( x̄ = 4 μm, n = 10), hyaline, branched, septate, longer
than asci. Asci 60–86 × 6–10 μm ( x̄ x̄ = 75 × 8 μm, n = 30),
8-spored, unitunicate, broad cylindrical, long-pedicellate, with a J-, wedge-shaped, subapical ring. Ascospores
32–50 × 2–4 μm ( x̄ = 38 × 3 μm, n = 50), filiform, straight or
curved, C-shaped or sigmoid, aseptate, acute ends, without
polar appendages, smooth-walled, with minute guttules in
each cell. Asexual morph: Undetermined.
Culture characters: Colonies on MEA reaching 30 mm
diam. after 4 weeks at 25 °C, Culture from above, cream,
radiating, wrinkled, folded in the middle, dense, lobate, flattened, umbonate, edge irregular, fluffy; reverse black in the
middle and cream at the edge, radially striate with lobate
edge.
Material examined: Thailand, Chiang Rai Province, on
dead stems of Clematis subumbellata, 20 March 2017, C.
Phukhamsakda, CMTH18 (MFLU 17–1482, holotype); extype living culture, MFLUCC 17–2074.
Host: Clematis subumbellata—(This study).
Distribution: Thailand—(This study).
GenBank accession numbers: LSU: MN628626; tef1:
MN629286; rpb2: MN628628.
Notes: In the phylogenetic analysis, Neoleptosporella
clematidis (strain MFLUCC 17–2074) formed a distinct
clade from other Leptosporella species (Fig. 93). The asexual morph of our taxon was not obtained from culture. In
a BLASTn search of GenBank, the closest match of the
LSU sequence of MFLUCC 17–2074 is Leptosporella
bambusae with 85% similarity to strain MFLUCC 12–0846
(NG_059674). This is the first record of Leptosporellaceae
on Clematis (Fig. 94).
Isolate MFLUCC 17–2074 was evaluated for bioactive
secondary metabolite production. The strain inhibits the
growth of Bacillus subtillis but not at significant values (data
not shown).
Sordariales Chadef. ex Hawksw. & O.E. Erikss.
This order comprises three families Chaetomiaceae, Sordariaceae and Lasiosphaeriaceae sensu lato (Zhang et al.
2006; Maharachchikumbura et al. 2016).
Chaetomiaceae Winter
Chaetomiaceae is a highly diverse family that can be
found in several environments as fungicolous, saprobes, or
parasites in plant or animals (Mukerji and Manoharachary
13
152
Fig. 94 Neoleptosporella clematidis (MFLU 17–1482, holotype). a
Appearance of ascomata on Clematis subumbellata. b Vertical section through ascoma. c Ostiole. d Peridium. e Paraphyses. f–h Asci
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Fungal Diversity (2020) 102:1–203
(h asci in 5% KOH presenting a J-reaction of apical ring, wedgeshaped). i–m Ascospores. n Culture characteristics on MEA. Scale
bars: b = 100 μm, c, d = 50 μm, e–h = 20 μm, i–m = 10 μm
Fungal Diversity (2020) 102:1–203
2010; Ahmed et al. 2016). Wang et al. (2019) made a major
revision for Chaetomiaceae, accepting 37 genera.
Dichotomopilus Wang, Samson & Crous
Dichotomopilus was erected by Wang et al. (2016) for
a group characterised by dichotomously branched terminal
ascomatal hairs. Dichotomopilus is typified by D. indicus,
with 12 species listed in Index Fungorum (2020). Based on
both morphological and phylogenetic analysis of combined
ITS, LSU, tef1 and rpb2 sequence data (Fig. 95), we introduce the first record of Dichotomopilus ramosissimum on
Clematis (Fig. 96).
Dichotomopilus ramosissimum (X. Wei Wang & L. Cai)
Wei Wang & Samson, Studies in Mycology 84: 217 (2016)
new host record
Basionym: Chaetomium ramosissimum X. Wei Wang &
L. Cai, Mycol. Prog. 13: 725. (2014).
Index Fungorum number: IF 801734; Facesoffungi number: FoF 07245, Fig. 96.
Saprobic on dead branches of Clematis vitalba. Sexual
morph: Ascomata 70–225 × 80–220 μm ( x̄ = 150 × 140 μm,
n = 5), erumpent to superficial, with thick aerial hyphae
or exposed ostiolate, covered with black ascomatal hairs,
sphaerical or ovate, without papilla. Peridium reddish brown
to brown, composed of hypha-like cells, textura intricata
mixed with textura epidermoidea in surface view. Terminal hairs erect, 120–290 × 3–6 μm ( x̄ = 170 × 5 μm, n = 20),
rigid, dark brown, copiously dichotomously branched
more than four times at the side, straight angles, starting
from the lower half, punctulate or verrucose. Lateral hairs
50–100 × 3–4 μm ( x̄ = 70 × 4 μm, n = 20), unbranched,
setae-like, tapering towards the end. Asci 30–40 × 8–15 μm
( x̄ = 35 × 10 μm, n = 10), 8-spored, spathulate, long pedicellate, apically rounded. Ascospores 5–8 × 2.5–5 μm ( x̄
= 7 × 4 μm, n = 50), overlapping, ovate to subglobose, ends
acute, hyaline, becoming brown with age, aseptate, verruculose, with slit-like germ pores at maturity. Asexual morph:
Undetermined.
Material examined: UK, Hampshire, Botley wood,
Hampshire, on dead stems of Clematis vitalba, 16 April
2016, E.B.G. Jones, GJ264 (MFLU 16–2137).
Hosts: The rhizosphere of Panax notoginseng, soil sample, Clematis vitalba—(Wang et al. 2014; this study).
Distribution: China, UK—(Wang et al. 2014; this study).
GenBank accession numbers: LSU: MT214611; SSU:
MT226722; ITS: MT310655; tef1: MT394667.
Notes: Dichotomopilus ramosissimum was described as
Chaetomium ramosissimum. Later, with more taxon sampling of Chaetomiaceae, the species formed a separate lineage and was synonymized under Dichotomopilus. Our collection formed a close relationship with the type strain of D.
ramosissimum (CGMCC 3.14183, 100% ML/1.00 BYPP).
153
The morphological characters of our collection are similar to those reported by Wang et al. (2014). In a BLASTn
search of GenBank, the closest match of the ITS sequence
of strain MFLU 16–2137 was D. erectus strain CBS 140.56
with 98.5% similarity (MH857548), while the closest match
of the tef1 sequence was D. ramosissimum strain CGMCC
3.14183 (KC485021) with 100% similarity.
Sordariaceae Winter
Sordariaceae is typified by Sordaria, and species in this
family mainly occur as coprophilous or saprobes (Cai et al.
2006; Zhang et al. 2006; Arif and Saleem 2017). Sordariaceae is characterized by coriaceous ascomata and hyaline or
brown ascospores with sheaths (Maharachchikumbura et al.
2016). Eight genera are accepted in this family (Maharachchikumbura et al. 2016; Wijayawardene et al. 2017).
Sordaria Ces. & De Not
Sordaria is typified with Sordaria fimicola. There are
more than 50 epithets listed in Index Fungorum (2020), but
only 15 strains have sequence data (Cai et al. 2006). Most
species are reported from dung, soil, and seed pods (Furtado 1969; Watanabe 1989; Mungai et al. 2012). Sordaria is
characterized by perithecioid, semi-immersed to superficial,
coriaceous ascomata, paraphyses, cylindrical asci, with a
lobate pedicel, and a prominent apical ring, and uniseriate
ascospores that are ellipsoidal to ovoid, surrounded by a
gelatinous sheath. Based on both morphological and phylogenetic analysis of combined LSU, ITS and tub sequence
data (Fig. 97), we introduce the first record of Sordaria, S.
clematidis on Clematis species (Fig. 98).
Sordaria clematidis Phukhams. & K.D. Hyde, sp. nov.
Index Fungorum number: IF557306; Facesoffungi number: FoF 07337, Fig. 98.
Etymology: Refers to the host genus, Clematis.
Holotype: MFLU 16–2138
Saprobic on dead stems of Clematis vitalba. Sexual morph: Ascomata 220–315 × 200–350 μm
( x̄ = 270 × 300 μm, n = 5), perithecial, single, superficial,
solitary, scattered, ampulliform or pyriform, globose, with
a sphaerical body, coriaceous, smooth or nearly, often glossy
and black, ostiolate. Ostioles central, oblong, dark brown
to black, papillate, periphyses filling ostioles. Peridium
20–30 μm wide, composed of 6–7 layers of textura angularis
of thin-walled, brown cells, inner layer lined with thin hyaline layers. Paraphyses of numerous, 10–17 μm ( x̄ = 14 μm,
n = 40), catenophyses, branched, transversely septate, hyaline. Asci 115–160 × 4–7 μm ( x̄ = 180 × 20 μm, n = 20), unitunicate, oblong, 8-spored, with a thin but rather persistent
wall, with a disc and J-ring at the truncate apex, simple pedicel. Ascospores 21–26 × 11–15 μm ( x̄ = 25 × 13 μm, n = 20),
uniseriate, ovate, aseptate, smooth-walled, hyaline or pale
13
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Fungal Diversity (2020) 102:1–203
Dichotomopilus subfunicola CGMCC 3.12892 T
50/---
Dichotomopilus subfunicola CBS 332.72
Dichotomopilus pseudofunicola CBS 142033
75/1.00
Dichotomopilus variostiolatus CBS 179.84 T
100/1.00 62/--
Dichotomopilus funicola CBS 159.52 T
88/1.00
Dichotomopilus indicus CGMCC 3.14184 e T
Dichotomopilus pratensis CBS 133396 T
Dichotomopilus ramosissimums CGMCC 3.14183 T
100/1.00
Dichotomopilus ramosissimums CGMCC 3.12930
84/0.99
Dichotomopilus ramosissimums MFLU 16-2137
Dichotomopilus
Dichotomopilus pseudoerectus CBS 252.75 T
100/1.00100/1.00
100/1.00
Dichotomopilus erectum CBS 140.56 T
Dichotomopilus erectum CGMCC 3.2900
Dichotomopilus dolichotrichus CBS 162.48 T
85/1.00
59/0.96
90/1.00
Dichotomopilus reflexus CBS 157.49 T
Dichotomopilus fusus CBS 372.66 T
Chaetomium angustispirale CBS 137.58
100/1.00
Chaetomium subglobosum CBS 483.73
--
Chaetomium subfimeti CBS 370.66 T
95/0.99
Chaetomium ascotrichoides CBS 113.83
98/1.00
100/1.00
Chaetomium globosum CBS 371.66 T
Chaetomium
Chaetomium globosum CBS 148.51
83/1.00
100/1.00
Chaetomium novozelandicum CBS 124555 T
Achaetomium umbonatum IMI 381871
77/0.99
51/0.98
Achaetomium macrosporum CBS 152.97 T
Achaetomium globosum CBS 332.67 T
52/0.99
Achaetomium strumarium CBS 333.67 T
99/1.00
100/1.00
Achaetomium
Achaetomium cristalliferum CBS 781.84
Achaetomium lippiae URM7547
79/1.00
Achaetomium luteum CBS 544.83 T
100/1.00
100/1.00
82/1.00
88/1.00
100/1.00
Ovatospora mollicella CBS 583.83 T
Ovatospora pseudomollicella CBS 251.75 T
Ovatospora brasiliensis CBS 126169
Ovatospora
Ovatospora medusarum CBS 148.67 T
Ovatospora senegalensis CBS 728.84 T
Collariella bostrychodes CBS 163.73 T
100/1.00
Collariella carteri CBS 128.85 T
0.03
13
Collariella
(Outgroup)
Chaetomiaceae
98/1.00 99/--
Fungal Diversity (2020) 102:1–203
◂Fig. 95 Phylogram generated from maximum likelihood analy-
sis based on combined ITS, LSU, tef1 and rpb2 sequence data representing related taxa in Chaetomiaceae. Related sequences were
taken from to Wang et al. (2019) and 33 strains were included in
the combined analyses which comprised 2972 characters (581 characters for ITS, 855 characters for LSU, 936 characters for tef1, 600
characters for rpb2, including gap regions). Collariella bostrychodes
(CBS 163.73) and Collariella carteri (CBS 128.85) are used as the
outgroup taxa. The tree from the maximum likelihood analysis had
similar topology to the Bayesian analyses. The best sorting RaxML
tree had a final likelihood value of − 12501.711733 is presented.
The matrix had 710 distinct alignment patterns with 6.40% undetermined characters or gaps. Estimated base frequencies were as follows: A = 0.235341, C = 0.273490, G = 0.285063, T = 0.206106;
substitution rates AC = 1.347902, AG = 3.116716, AT = 1.526141,
CG = 1.846122, CT = 7.547565, GT = 1.000000; gamma distribution
shape parameter α = 1.039485. In our analysis, GTR + I + G model
was used for each partition in Bayesian posterior analysis. The species determined in this study is indicated in blue. Bootstrap values
(BS) greater than 50% BS (ML, left) and Bayesian posterior probabilities (BYPP, right) greater than 0.90 are given at the nodes. Hyphens
(-) represent support values less than 50% BS/0.90 BYPP. Thick
branches represent significant support values from all analyses at the
genus level (BS ≥ 70%/BYPP ≥ 0.95)
brown, guttulate, with 3–7 μm thick mucilaginous sheath
with upper pore. Asexual morph: Undetermined.
Material examined: UK, Hampshire, Botley Wood, on
dead branches of Clematis vitalba, 25 May 2016, E.B.G.
Jones, GJ289 (MFLU 16–2138, holotype).
Hosts: Clematis vitalba—(This study).
Distribution: UK—(This study).
GenBank accession numbers: LSU: MT214612; SSU:
MT226723; ITS: MT310656; tef1: MT394668; rpb2:
MT394717.
Notes: Sordaria clematidis (Fig. 98) has characters typical of Sordaria in having single perithecial, superficial,
coriaceous ascomata with papillate ostioles, unitunicate,
oblong asci with uniseriate, ovate, aseptate ascospores
with a mucilaginous sheath. In the phylogenetic analysis
(Fig. 97), S. clematidis formed a close relationship with the
generic type S. fimicola (CBS 508.50), but received relatively low statistical support. Sordaria species are similar
in their morphological characters, however, S. clematidis is
unique in having pale yellow ascospores with upper pore in
mucilaginous sheath and a germ-slit is absent (Maharachchikumbura et al. 2016). In a BLASTn search of GenBank, the
closest match of the LSU sequence of strain MFLU 16–2138
is S. fimicola (strain CBS 485.64) with 98.00% similarity
(MH870123). The closest match with the ITS sequence was
Sordaria fimicola strain CBS 485.64 with 99.06% similarity (MH858489). Therefore, we introduce S. clematidis as
a new species.
155
Subclass Diaporthomycetidae Senan., Maharachch. & K.D.
Hyde
Diaporthales Nannf.
Diaporthales is one of the most diverse orders within
Diaporthomycetidae and comprises 24 families (Senanayake
et al. 2017; Guterres et al. 2019). The order includes plantassociated fungi, some of which are economically important
pathogens, parasites, others are endophytes or saprobes on
various substrates (Alvarez et al. 2016). Diaporthales taxa
are characterized by having pseudo- or ascostromata, brown
to black perithecial fruiting bodies, asci with J-, refractive
apical ring and hyaline to brown ascospores. The asexual
states are usually coelomycetous with conidiophores arising from the inner side of the peridium layer, enteroblastic,
holoblastic, phialidic or annellidic conidiogenous cells, and
hyaline to brown conidia (Sogonov et al. 2008; Senanayake
et al. 2017).
Diaporthaceae Höhn. ex Wehm.
Diaporthaceae taxa have a cosmopolitan distribution
occurring as pathogens, saprobes or endophytes of terrestrial
plants (Udayanga et al. 2014; Rossman et al. 2017; Thambugala and Hyde 2018). The sexual morphs of this family are
characterized by perithecial, immersed to erumpent, papillate ascomata, 8-spored, asci with J-, refractive apical ring,
and 1 septum, hyaline or dark to blackish brown ascospores
with or without polar appendages. Asexual morphs are characterized by acervular or pycnidial conidiomata, ampulliform, cylindrical, septate conidiophores, enteroblastic,
phialidic, determinate conidiogenous cells and 0–2-septate,
hyaline or brown conidia sometimes with short appendages
(Voglmayr and Jaklitsch 2014; Dissanayake et al. 2017;
Senanayake et al. 2017, 2018). The family comprises 14
genera accepted by Wijayawardene et al. (2018).
Diaporthe Nitschke
Diaporthe (= Phomopsis) is an economically important
plant pathogen that also includes endophytes or saprobes
on a broad range of plant hosts (Udayanga et al. 2014; Hyde
et al. 2016). Over 1000 epithets are listed under Diaporthe
and most were introduced based on morphological characters and host association (Aa et al. 1990). In view of the
genetic diversity and complexity of the known species, identifications of Diaporthe species currently relies on multilocus phylogenetic analyses (Senanayake et al. 2018; Manawasinghe et al. 2019). Phylogenetic analysis resulting from
combined ITS, tef1, cal and tub2 sequence data of selected
Diaporthe species are presented in Figs. 99 and 101. The
ambiguous species that are related to the newly described
species were analysed separately using Genealogical Concordance Phylogenetic Species Recognition (GCPSR) (Taylor et al. 2000) by performing a pairwise homoplasy index
(Φw) test and the results are shown in Figs. 100 and 103.
13
156
Fig. 96 Dichotomopilus ramosissimum (MFLU 16–2137). a Appearance of perithecia on Clematis vitalba. b, c Close up of ascomata on
host surface. d Ascoma. e, f Dichotomously branched terminal hairs
13
Fungal Diversity (2020) 102:1–203
and setae-like lateral hairs. g Structure of peridium in surface view.
h Asci. i–m Ascospores. Scale bars: b = 200 µm, c, d = 100 µm, e,
f = 50 µm, g, h = 20 µm, i–m = 5 µm
Fungal Diversity (2020) 102:1–203
157
Sordaria lappae CBS 154.97
64/--
Sordaria superba CBS 784.96
Sordaria arctica CBS 143.68
Sordaria sibutii CBS 768.96
--/0.97
Sordaria macrospora ATCC MYA-4828
Sordaria nodulifera TW 85-72 T
100/1.00
Sordaria tamaensis NBRC 32552 T
51/--
80/1.00
Sordaria fimicola CBS 508.50
Sordaria clematidis MFLU 16–2138 T
68/--
Sordaria cf. macrospora DB5751
Sordaria
Sordaria humana ATCC 22796
Sordaria alcina CBS 109460
91/0.99
97/1.00
Sordaria tomentoalba CBS 260.78
Sordaria sclerogenia FGSC 2741
96/1.00
Sordaria brevicollis FGSC 1904
99/1.00
Sordaria prolifica CBS 567.72
99/1.00
Sordaria conoidea CBS 563.72
Neurospora santi-florii CBS 571.72
91/0.98
Neurospora sitophila FGSC 8770
0.006
Fig. 97 The Bayesian 50% majority-rule consensus phylogram for
Sordaria members based on combined LSU, ITS and tub sequence
data. The topology and clade stability of the combined gene analyses was compared to the single gene analyses. The tree is rooted
with Neurospora santi-florii (CBS 571.72) and Neurospora sitophila
(FGSC 8770). Nineteen strains were included in the combined analyses which comprised 2436 characters (833 characters for LSU, 601
characters for ITS, 1002 characters for tub, including gap regions).
The tree from the maximum likelihood analysis had similar topology to the Bayesian analyses. The best scoring RAxML tree had a
final likelihood value of − 5155.259712. The matrix had 275 distinct alignment patterns with 36.05% undetermined characters and
gaps. Estimated base frequencies were as follows; A = 0.239918,
C = 0.256198, G = 0.278423, T = 0.225461; substitution rates
AC = 1.231551, AG = 4.410425, AT = 1.886486, CG = 1.711684,
CT = 9.592785, GT = 1.000000; gamma distribution shape parameter α = 1.270759. In our analysis, GTR + I + G model was used for
each partition in Bayesian posterior analysis. The species determined
in this study is indicated in blue. Bootstrap values (BS) greater than
50% BS (ML, left) and Bayesian posterior probabilities (BYPP, right)
greater than 0.90 are given at the nodes. Hyphens (-) represent support values less than 50% BS/0.90 BYPP. Thick branches represent
significant support values from all analyses (BS ≥ 70%/BYPP ≥ 0.95)
at the genus level
13
158
Fig. 98 Sordaria clematidis (MFLU 16–2138, holotype). a, b
Appearance of ascomata on Clematis vitalba. c Vertical section
through ascoma. d Ostioles. e Partial peridium part. f Paraphyses. g
Presenting of apical ring. h–j Asci (j Ascus in 5% KOH with J-reac-
13
Fungal Diversity (2020) 102:1–203
tion of wedge-shaped apical ring. k–n Ascospores (n Ascospore in
10% Indian ink). Scale bars: a = 500 μm, b = 200 μm, c = 100 μm,
d–f, k–n = 20 μm, g = 10 μm, h–j = 50 μm
Fungal Diversity (2020) 102:1–203
Fig. 99 Phylogram generated
from maximum parsimony
analysis based on combined
ITS, tef1, cal and tub2 sequence
data for selected members of
Diaporthe. Related sequences
are taken from Hyde et al.
(2019) and retrieved from GenBank. The species tree included
all members of Diaporthe and
was performed for placement
determination. Forty-two strains
were included in the analysis of the combined loci and
comprised 2359 characters (598
characters for ITS, 393 characters for tef1, 541 characters
for cal, 827 characters for tub2,
including alignment gaps). The
tree was rooted with Diaporthe
crataegi (CBS 114435). Maximum parsimony analysis of 590
parsimony informative characters resulted in a single most
parsimonious tree (CI = 0.585,
RI = 0.681, RC = 0.398,
HI = 0.415). The best scoring
RAxML tree had a final likelihood value of − 14560.846477.
GTRGAMMA bootstrapping
model was applied to the matrix
which had 1152 distinct alignment patterns, with 27.96% of
undetermined characters and
gaps. Estimated base frequencies were: A = 0.218108,
C = 0.320856, G = 0.237387,
T = 0.223649; substitution rates
AC = 1.282937, AG = 3.695471,
AT = 1.225056, CG = 1.103739,
CT = 4.789247, GT = 1.000000;
gamma distribution shape
parameter α = 0.888152. In our
analysis, GTR + I + G model
was used for each partition in
Bayesian posterior analysis.
Bootstrap values (BS) from
maximum parsimony (MP,
left), maximum likelihood (ML,
right) higher than 50% BS and
Bayesian posterior probabilities
(BYPP, below) greater than 0.90
are given at the nodes. Hyphens
(-) represent support values
less than 50% BS/0.90 BYPP.
The ex-type strains are in bold
and black. The newly generated
sequence is in bold and blue
159
-/88
99/100
1.00
Diaporthe celastrina CBS 139.27
Diaporthe maritima DAOM 695742
-/73
0.98
Diaporthe
Diaporthe bicincta CBS 121004
54/70
0.99
MP/ML
PP
Diaporthe alleghaniensis CBS 495.72
81/89
1.00
Diaporthe vaccinii CBS 160.32
Diaporthe biguttusis CGMCC 3.17081
-/55
0.99
Diaporthe eres CBS 138594
100/100
1.00
Diaporthe lonicerae MFLUCC 17-0963
78/89
1.00
-/0.94
66/85
1.00
Diaporthe longicolla CGMCC 3.17089
Diaporthe camptothecicola CFCC 51632
70/89
1.00
Diaporthe momicola MFLUCC 16-0113
70/76
1.00
Diaporthe phragmitis CBS 138897
69/60
1.00
68/79
1.00
Diaporthe celeris CBS 143349
Diaporthe cf. nobilis CBS 338.89
51/88
-
Diaporthe nitschkei AR5211
60/67
0.99
Diaporthe mahothocarpus MFLUCC 15-0749
-/79
-
56/84
0.99
Diaporthe cf. nobilis RG-2013 CBS 124030
Diaporthe betulae CFCC 50469
Diaporthe alnea CBS 146.46
54/81
0.99
100/100
1.00
Diaporthe neilliae CBS 144.27
Diaporthe heterophyllae CPC 26215
62/91
1.00
85/98
1.00
85/98
1.00
Diaporthe virgiliae CMW 40755
Diaporthe penetriteum LC 3353
-/0.99
Diaporthe subclavata CGMCC 3.17257
-/0.98
100/100
1.00
Diaporthe citrichinensis CBS 134242
Diaporthe oraccinii LC3166
Diaporthe citri CBS 135422
94/89
1.00
Diaporthe apiculatum LC3418
100/100
1.00
71/75
1.00
98/99
1.00
Diaporthe gardeniae CBS 288.56
Diaporthe charlesworthii BRIP 54884
77/75
1.00
Diaporthe sennicola CFCC 51634
Diaporthe nomurai CBS 157.29
84/95
1.00
100/100
1.00
Diaporthe clematidina MFLUCC 17–2060
54/81
0.98
72/85
1.00
Diaporthe aquatica IFRDCC 3015
Diaporthe psoraleae-pinnatae CPC 21638
99/100
1.00
Diaporthe incompleta CGMCC 3.18288
Diaporthe rhoina CBS 146.27
89/58
1.00
78/70
1.00
Diaporthe aspalathi CBS 117169
Diaporthe woodii CBS 558.93
86/98
1.00
-/89
-
87/98
1.00
Diaporthe crotalariae CBS 162.33
Diaporthe litoricola MFLUCC 17-1657
Diaporthe caulivora CBS 127268
Diaporthe crataegi CBS 114435
0.03
13
160
Fig. 100 The splits graph from
the pairwise homoplasy index
(PHI) test generated from the
concatenated gene set of ITS,
tef1, cal and tub2 sequence data
of closely related species using
both LogDet transformation
and splits decomposition. In
the PHI test results (Φw) < 0.05
indicates significant recombination within the dataset. The
strain determined in this study
is in bold and blue
Fungal Diversity (2020) 102:1–203
Diaporthe incompleta
CGMCC 3.18288
Diaporthe aquatica IFRDCC 3015
Diaporthe incompleta
LC6754
Diaporthe clematidina
MFLUCC 17-2060
Φw = 0.42
0.01
Diaporthe psoraleae-pinnatae
CPC 21638
Diaporthe rhoina CBS 146.27
Diaporthe clematidina Phukhams., M.V. de Bult & K.D.
Hyde, sp. nov.
Index Fungorum number: IF557300; Facesoffungi number: FoF 07262, Fig. 101.
Etymology: Named after the host genus, Clematis.
Holotype: MFLU 17–1466.
Saprobic on Clematis subumbellata. Sexual morph:
Ascomata 280–367 × 167–296 μm ( x̄ = 325 × 250 μm, n = 5),
perithecial, solitary to aggregated, immersed, papillate only
visible on the host substrate, obpyriform to globose, coriaceous, brown to dark brown, heavily pigmented at apex,
papillate. Ostioles 125–221 × 47–50 μm ( x̄ = 180 × 50 μm,
n = 5), papillate, central or eccentric, broad oblong, filled
with periphyses. Peridium 12–31 μm wide ( x̄ = 21 μm,
n = 20), composed of 2–3 layers of thin-walled cells of
textura globosa mixed with textura angularis, brown to reddish brown, thin at inner layer, hyaline. Paraphyses dense,
2–3 μm ( x̄ = 2.5 μm, n = 30), septate, constricted at septa,
broad filiform, tapering above asci. Asci 32–54 × 7–10 μm
( x̄ = 46.5 × 9 μm, n = 20), 8-spored, unitunicate, oblong
to broad oblong, thin-walled, short or apedicellate, with
a refractive, J-, apical ring. Ascospores 12–15 × 3–5 μm
( x̄ = 14 × 4 μm, n = 40), partially overlapping, oblongellipsoidal, hyaline, uniseptate, with two guttules in each
cell. Asexual morph: Pycnidial on PDA media. Conidiomata 140–350 × 100–220 μm ( x̄ = 225 × 155 μm, n = 5),
globose, eustromatic, multilocular, gregarious, erumpent or
superficial, covered by dense vegetative hyphae, occasionally with ostiolate necks. Pycnidial wall parenchymatous,
consisting of brown, thick-walled cells of textura angularis.
Conidiophores short, hyaline, smooth, densely aggregated,
13
cylindrical, straight. Conidiogenous cells 8–14 × 8–12 μm,
phialidic, cylindrical, terminal, with slight tapering towards
apex, hyaline, formed from the inner layer of pycnidial wall.
Alpha conidia 5–8 × 2–3 μm ( x̄ = 6 × 2.5 μm, n = 20), aseptate, hyaline, smooth, ovate to ellipsoidal, biguttulate. Beta
conidia 17.5–30 × 1–2 μm ( x̄ = 25 × 1.7 μm, n = 5), aseptate,
hyaline, smooth, fusiform, tapering towards both ends, apex
conical, base subtruncate. Gamma conidia not observed.
Culture characters: Colonies growing on PDA reaching 20 mm within 4 weeks at 25 °C. Culture from above,
white, radiating, to the edge, faintly zonate, margin undulate,
dense, flat or umbonate; reverse light brown, white radiating
outwardly.
Material examined: Thailand, Chiang Rai Province, on
dead stems of Clematis subumbellata, 20 March 2017, C.
Phukhamsakda, CMTH2 (MFLU 17–1466, holotype); extype living culture, MFLUCC 17–2060.
Host: Clematis subumbellata—(This study).
Distribution: Thailand—(This study).
GenBank accession numbers: LSU: MT214613; SSU:
MT226724; ITS: MT310657; tef1: MT394669, MT394625;
rpb2: MT394718; cal: MT394624; tub2: MT394623.
Notes: Phylogenetic analysis of Diaporthe clematidina
based on the combined ITS, tef1, cal and tub2 sequence
data, shows that the strain clusters in a well-supported clade
(84% MP, 95% ML support and 1.00 BYPP) sister to D.
aquatica (IFRDCC 3015) as shown in Fig. 99. Diaporthe
clematidina differs from D. aquatica in its shorter neck
(1100–2250 × 80–120 vs 125–221 × 47–50 μm), and broad
oblong ascospores with conical ends. Diaporthe clematidina
was found in a terrestrial habitat, while D. aquatica was
Fungal Diversity (2020) 102:1–203
Fig. 101 Diaporthe clematidina (MFLU 17–1466, holotype). a,
b Appearance of ascomata on Clematis subumbellata. c Close up
of ascoma on the host. d Vertical section through ascoma. e Ostiolar canal. f Peridium. g Paraphyses. h–j Asci. k–n Ascospores. o,
161
p Culture characteristics on PDA. q Pycnidia produced in culture. r
Conidiogenous cell with an alpha conidium. s Conidiogenous cell
with a beta conidium. t Alpha and beta conidia. Scale bars: b–d,
q = 200 µm, e = 100 µm, f, g = 50 µm, h–j = 20 µm, k–n, r–t = 10 µm
13
162
Fig. 102 Phylogram generated
from maximum parsimony
analysis based on combined
ITS, tef1, cal and tub2 sequence
data for selected Diaporthe
species. Related sequences were
selected from Hyde et al. (2019)
and retrieved from GenBank.
The species tree was performed
for placement determination.
Sixty-four strains were included
in the combined analyses, which
comprised 2380 characters (603
characters for ITS, 409 characters for tef1, 542 characters
for cal, 826 characters for tub2,
including alignment gaps). The
tree was rooted with Diaporthe
toxica (CBS 534.93). Maximum
parsimony analysis of 736
parsimony informative characters resulted in a single most
parsimonious tree (CI = 0.555,
RI = 0.815, RC = 0.453,
HI = 0.445). The best scoring
RAxML tree had a final likelihood value of − 18803.562906.
The matrix had 1329 distinct
alignment patterns, with 32.77%
of undetermined characters and
gaps. Estimated base frequencies were: A = 0.224328,
C = 0.306223, G = 0.237767,
T = 0.231682; substitution rates
AC = 1.005463, AG = 2.446938,
AT = 0.976921, CG = 0.744774,
CT = 3.4857627,
GT = 1.000000; gamma
distribution shape parameter
α = 0.765064. In our analysis,
GTR + I + G model was used
for each partition in Bayesian
posterior analysis. The species
determined in this study are
indicated in blue. Bootstrap
values (BS) from maximum
parsimony (MP, left), maximum
likelihood (ML, right) higher
than 50% BS and Bayesian
posterior probabilities (BYPP,
below) greater than 0.90 are
given at the nodes. Hyphens (-)
represent support values less
than 50% BS/0.90 BYPP. The
ex-type strains are in bold and
black
Fungal Diversity (2020) 102:1–203
Diaporthe
MP/ML
PP
90/99
1.00
0.98
--/63
0.99
59/63
0.99
68/87
100/100 0.99
1.00
80/83
1.00
83/97
1.00
72/94
1.00
73/94
1.00
Diaporthe ternstroemia CGMCC 3.15183
Diaporthe fusicola CGMCC 3.17087
Diaporthe sterilis CBS 136969
Diaporthe ovoicicola CGMCC 3.17092
Diaporthe garethjonesii MFLUCC 12-0542
Diaporthe amygdali CBS 126679
Diaporthe obtusifoliae CPC 32336
Diaporthe acaciigena CBS 129521
Diaporthe pustulata CBS 109742
--/65
0.99
68/-1.00
97/87
1.00
100/100
1.00
64/71
0.95
86/97
1.00
64/71
1.00
Diaporthe ravennica MFLUCC 15-0479, Tamarix sp.
Diaporthe ravennica MFLUCC 17-1029, Salvia sp.
Diaporthe ravennica MFLUCC 15-0480, Tamarix sp.
Diaporthe ravennica MFLUCC 16-0997, Clematis vitalba
Diaporthe baccae CBS 136972
Diaporthe foeniculacea CBS 111553
Diaporthe chamaeropis CBS 454.81
Diaporthe cytosporella FAU 461
Diaporthe dorycnii MFLUCC 17-1015
Diaporthe cinerascens CBS 719.96
100/100
Diaporthe macintoshii BRIP 55064
1.00
53/74
Diaporthe vangueriae CPC 22703
---/---/-- Diaporthe maytenicola CPC 21896
0.93
0.98
Diaporthe parapterocarpi CPC 22729
-54/56
Diaporthe cissampeli CPC 27302
-67/61
Diaporthe hickoriae CBS 145.26
0.99
57/61
0.90
Diaporthe saccarata CBS 116311
Diaporthe oncostoma CBS 589.78
-57/61
0.90
Diaporthe canthii CBS 132533
99/98
Diaporthe anacardii CBS 720.97
-1.00
Diaporthe velutina CGMCC 3.18286
-100/100
--/51
Diaporthe inconspicua CBS 133813
1.00
-Diaporthe pterocarpi MFLUCC 10-0571
100/100
Diaporthe isoberliniae CPC 22549
1.00
Diaporthe elaeagni CBS 504.72
--/51
Diaporthe stictica CBS 370.54
-66/90
0.99
Diaporthe elaeagni-glabrae CGMCC 3.18287
--/57
Diaporthe diospyricola CPC 21169
-Diaporthe psoraleae CPC 21634
Diaporthe crataegi CBS 114435
Diaporthe toxica CBS 534.93
60.0
13
Diaporthe rudis CPC 28268, Vitis vinifera
Diaporthe rudis STE.U 5683, Vitis vinifera
Diaporthe rudis CBS 266.85, Rosa rugosa
Diaporthe rudis CPC 28425, Vitis vinifera
Diaporthe rudis CPC 29658, Vitis vinifera
Diaporthe rudis CBS 109292, Brugmansia sp.
Diaporthe rudis AR3654, Rosa canina
Diaporthe rudis AR3422, Laburnum anagyroides
Diaporthe rudis CPC 29320, Vitis vinifera
Diaporthe rudis CBS 114436, Sambucus cf. racemosa
66/100
Diaporthe rudis CBS 113201, Vitis vinifera
0.98
Diaporthe rudis MFLUCC 17-2153, Clematis serratifolia
68/99
Diaporthe rudis MFLUCC 17-2500, Anthoxanthum odoratum
0.99
Diaporthe rudis ICMP 16419, Castanea sativa
91/99
Diaporthe asheicola CBS 136967
1.00
Diaporthe australafricana CBS 111886
66/99
1.00
Diaporthe cynaroidis CBS 122676
85/100
57/68
Diaporthe salicicola BRIP 54825
0.99
0.91
Diaporthe pseudotsugae MFLU 15-3228
91/99
1.00
Diaporthe beckhausii CBS 138.27
72/93
Diaporthe cassines CBS 136440
0.99
Diaporthe nothofagi BRIP 54801
98/99
Diaporthe cf. heveae CBS 852.97
1.00
Diaporthe ocoteae CPC 26217
65/63
Fungal Diversity (2020) 102:1–203
163
Diaporthe chamaeropis CBS 454.81
Diaporthe cytosporella FAU 461
Diaporthe ravennica MFLUCC 15-0479, Tamarix sp.
Diaporthe ravennica MFLUCC 17-1029, Salvia sp.
Diaporthe ravennica MFLUCC 16-0997, Clematis vitalba
Diaporthe ravennica MFLUCC 15-0480, Tamarix sp.
Diaporthe foeniculacea
CBS 111553
Diaporthe baccae CBS 136972
Diaporthe baccae CPC 29659
Diaporthe baccae CBS 143343
Diaporthe baccae CPC 29639
0.001
a Φw = 0.15
Diaporthe rudis CBS 109292, Brugmansia sp.
Diaporthe rudis AR3422, Laburnum anagyroides
Diaporthe rudis AR3654, Rosa canina
0.001
Diaporthe rudis CBS 266.85, Rosa rugosa
Diaporthe rudis CPC 28425, Vitis vinifera
Diaporthe rudis CPC 29658, Vitis vinifera
Diaporthe rudis CPC 29320, Vitis vinifera
Diaporthe rudis CPC 28268, Vitis vinifera
Diaporthe rudis STE.U 5683, Vitis vinifera
Diaporthe rudis CBS 114436, Sambucus cf. racemosa
Diaporthe rudis CBS 113201, Vitis vinifera
Diaporthe rudis ICMP 16419, Castanea sativa
Diaporthe rudis MFLUCC 17-2153, Clematis serratifolia
Diaporthe rudis MFLUCC 17-2500, Anthoxanthum odoratum
b Φw = 0.05
Fig. 103 The splits graph from the pairwise homoplasy index (PHI)
test generated from the concatenated gene set of closely related Diaporthe species a D. ravennica, b D. rudis using both LogDet transfor-
mation and splits decomposition. The PHI test (Φw) < 0.05 indicates
significant recombination within the dataset. The strains determined
in this study are in blue
13
164
13
Fungal Diversity (2020) 102:1–203
Fungal Diversity (2020) 102:1–203
associated with submerged wood in a small ditch (Hu et al.
2012). Diaporthe clematidina is phylogenetically related,
but differentiated from D. aquatica by 49 nucleotides in
ITS. However, the other gene regions for D. aquatica are
not available for nucleotide comparisons. When applying the
GCPSR concept to fulfil the genetic differentially independence of the combined dataset within the closely related taxa
(Fig. 99), we detected no significant recombination between
these isolates (Φw = 0.42, Fig. 100). Thus, D. aquatica and
D. clematidina represents two distinct species.
Diaporthe ravennica Thambug., Camporesi & K.D. Hyde,
Fungal Diversity 82: 296 (2016), new host record
Index Fungorum number: IF552100; Facesoffungi number: FoF02171, Fig. 104.
Saprobic on Clematis vitalba. Sexual morph:
Un d et e r m i n e d . A s ex u a l m o r p h : C o n i d i o m a ta
250–450 × 220–300 μm ( x̄ = 330 × 250 μm, n = 5), scattered,
uniloculate or multi-loculate, immersed to slightly erumpent,
gregarious, globose, occasionally with ostiolate necks.
Ostioles 75–190 × 52–170 μm ( x̄ = 121 × 124 μm, n = 5),
papillate, central or eccentric, broad oblong, lined with
periphyses. Pycnidial wall 12–20 μm ( x̄ = 14.6 μm, n = 20),
parenchymatous, walls consisting of dark brown, thick,
walled cells of textura angularis. Paraphyses not observed.
Conidiophores 5–14 × 1.5–2.5 μm ( x̄ = 8.5 × 2.0 μm, n = 30),
long, hyaline, smooth, densely aggregated, unbranched,
cylindrical, straight. Conidiogenous cells 1.8–2 × 2–3 μm,
phialidic, cylindrical, terminal, slightly tapering towards
apex, hyaline, formed from the inner layer of pycnidium
wall. Alpha conidia 8–11.5 × 2–4 μm ( x̄ = 9.5 × 2.8 μm,
n = 40), aseptate, hyaline, smooth, ovate to ellipsoidal,
biguttulate or multi-guttulate. Beta and gamma conidia not
observed on Clematis vitalba or in culture.
Culture characters: Colonies growing on PDA reaching
40 mm within 4 weeks at 16 °C. Culture from above, white,
radiating to the edge, margin undulate, medium dense, flat or
umbonate; reverse, cream, radiating white outwardly.
Material examined: Italy, Forlì-Cesena Province, San
Lorenzo in Noceto-Forlì, on dead aerial branch Clematis
vitalba, 20 March 2016, E. Camporesi, IT2893C (MFLU
20–0422); living culture, MFLUCC 16–0997.
Hosts: Clematis vitalba, Tamarix sp., Salvia sp.—(Thambugala et al. 2017; Dissanayake et al. 2017; this study).
◂ Fig. 104 Diaporthe ravennica (MFLU 20–0422). a, b Appearance
of conidiomata on Clematis vitalba. c Close up of conidioma on host
substrate. d Vertical section through conidioma. e Ostiolar canal. f
Peridium. g-j Conidiogenous cells and developing states of conidia.
k-o Alpha conidia. p, q Culture characteristics on PDA. Scale bars:
b = 500 µm, c, d = 200 µm, e, f = 50 µm, g–o = 10 µm
165
Distribution: Italy—(Thambugala et al. 2017; Dissanayake et al. 2017; this study).
GenBank accession numbers: LSU: MT214614; SSU:
MT226725; ITS: MT310658; tef1: MT394670.
Notes: Diaporthe ravennica (MFLUCC 16–0997) clusters
with other D. ravennica isolates from Tamarix sp. and Salvia
sp. (Thambugala et al. 2017; Dissanayake et al. 2017). The
description of D. ravennica by Thambugala et al. (2017)
stated that the strain produced beta conidia in natural substrates, but beta conidia were not observed in the strain
associated with Clematis vitalba branches or in culture. A
phylogenetic tree derived from an alignment of combined
ITS, tef1, cal and tub2 sequence data showed that the D.
ravennica clade received 65% support from ML and 0.99
BYPP (Fig. 102). The GCPSR concept was applied to the
concatenated gene set of the closely related taxa (D. baccae, D. chamaeropis, D. foeniculacea, D. ravennica) are
presented (Fig. 103). A phi-test of the four loci detected nonsignificant recombination between these isolates (Φw = 0.15,
Fig. 103a). Thus, the four isolates of D. ravennica including
our collection are clearly demarcated and delimited from the
other taxa (Fig. 104).
Diaporthe rudis (Fr.) Nitschke, in Pyrenomycetes Germanici 2: 282 (1870), new host record
Index Fungorum number: IF552100; Facesoffungi number: FoF02171, Fig. 105.
Saprobic on Clematis serratifolia. Sexual morph: Ascomata 320–420 × 190–275 μm ( x̄ = 376 × 246 μm, n = 5),
perithecial, solitary to aggregated, immersed, only papilla
visible on the host substrate, obpyriform to globose, coriaceous, dark brown to black, lighter pigment at apex, papillate. Ostioles 130–255 × 120–135 μm ( x̄ = 205 × 125 μm,
n = 5), papillate, central or eccentric, broad oblong, lined
with periphyses. Peridium 18–30(–45 μm at apex) wide,
composed of 2–3 layers of thin-walled cells of textura
angularis, brown to reddish brown, thin at inner layer, hyaline. Paraphyses sparse, 5–8 μm ( x̄ = 6 μm, n = 30), septate,
constricted at septa, broad filiform, tapering above asci.
Asci 50–72 × 9–13 μm ( x̄ = 60 × 10 μm, n = 20), 8-spored,
unitunicate, oblong to broad oblong, thin-walled, short or
apedicellate, with a refractive, J-, apical ring. Ascospores
7.5–13 × 2–5 μm ( x̄ = 7 × 3 μm, n = 40), partially overlapping, biseriate, oblong-ellipsoidal, hyaline, uniseptate, with
two guttules in each cell, with approximately 2.5 μm long
polar appendages. Asexual morph: See Udayanga et al.
(2014).
Culture characters: Colonies growing on PDA reaching 20 mm within 4 weeks at 25 °C. Culture from above,
white, radiating outwardly to the edge, circular, dense, flat or
umbonate; reverse cream, white, radiating outwardly.
Material examined: Belgium, Flemish Brabant; Meise
Botanic Garden, Bouchout domain, on dead branch of
13
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Fungal Diversity (2020) 102:1–203
Fig. 105 Diaporthe rudis (MFLU 17–1510). a Appearance of ascomata on Clematis serratifolia. b Close up of ascoma on host substrate. c Vertical section through ascoma. d Ostiolar canal. e Perid-
ium. f Paraphyses. g–h Asci. i–l Ascospores. Scale bars: b = 500 µm,
c = 200 µm, d, e = 100 µm, f = 50 µm, g, h = 20 µm, i–l = 5 µm
Clematis serratifolia, 13 June 2017, D. Ertz & C. Gerstmans, BRCS1 (MFLU 17–1510); living culture, MFLUCC
17–2153.
Hosts: Acer sp., Anthoxanthum odoratum, Asphodelus
albus, Aucuba japonica, Brugmansia, Carlina vulgaris,
Caragana arborescens, Castanea sp., Clematis serratifolia,
Cornus sp., Corylus sp., Dioscorea communis, Dipsacus fullonum, Epilobium sp., Eucalyptus sp., Fagus sp., Fraxinus
sp., Genista sp., Holcus sp., Hydrangea sp., Ileostylis sp.,
Laburnum sp., Lupinus, Malus sp., Protea sp., Pyrus sp.,
Rosa sp., Sambucus cf. racemosa, Salix sp., Vaccinium sp.,
Vitis vinifera—(Dissanayake et al. 2017; Guarnaccia and
Crous 2017; Rossman et al. 2017; Guterres et al. 2019; Farr
and Rossman 2020; this study).
Distribution: Australia, Austria, Belgium, Canada, Chile,
Czech Republic, France, Germany, Italy, Latvia, Japan,
Poland, Portugal, Spain, Sweden, Switzerland, Netherlands,
New Zealand, South Africa, UK, Ukraine, USA—(Dissanayake et al. 2017; Guarnaccia and Crous 2017; Rossman
et al. 2017; Guterres et al. 2019; Farr and Rossman 2020;
this study).
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Fungal Diversity (2020) 102:1–203
Phomatosporales Senan., Maharachch & K.D. Hyde
Senanayake et al. (2016) introduced Phomatosporales for
fungal strains that formed a distinct lineage in Diaporthomycetidae. The order comprises one family and the phylogeny
of Phomatosporaceae is shown in Fig. 106.
Phomatosporaceae Senan. & K.D. Hyde
Phomatosporaceae was formally reinstated by Senanayake et al. (2016) and comprises three genera, Lanspora, Phomatospora and Tenuimurus (Fig. 106). Phomatosporaceae
is typified by Phomatospora, which was placed in Sordariomycetes genera incertae sedis by Lumbsch and Huhndorf
(2007). We establish a novel species of Phomatospora from
Clematis in Thailand based on molecular data.
Phomatospora Sacc.
Phomatospora can be an endophyte or saprobe in both
aquatic and terrestrial habitats (Fallah and Shearer 1998;
Senanayake et al. 2016; Guarnaccia and Crous 2017). Phomatospora is typified by P. berkeleyi Sacc. and is characterized by immersed, globose or subglobose ascomata, pseudoparenchymatous cells forming a textura angularis to textura
prismatica peridium cell type, a paraphyses of hypha-like,
filamentous structures, cylindrical or oblong-fusiform,
and short pedicellate, unitunicate asci, with J-apical ring,
93/1.00
95/1.00
Phomatospora bellaminuta AFTOL-ID 766
Phomatospora viticola MFLU 16-1973 T
Phomatospora striatigera CBS 133932 T
Phomatospora biseriata MFLU 16-1970
--/1.00
97/1.00
50/1.00
58/0.93
--/1.00
Phomatospora biseriata MFLUCC 14-0832 T
Phomatospora uniseriata MFLUCC 17-2068 T
Lanspora coronata JK 5839A T
Phomatosporaceae
GenBank accession numbers: LSU: MT214615; SSU:
MT226726; ITS: MT310659; tef1: MT394671; rpb2:
MT394719.
Notes: Diaporthe rudis has been reported from a broad
range of hosts such as cultivated crops, ornamental plants
and is associated with grapevine trunk diseases worldwide
(Lombard et al. 2014; Guarnaccia and Crous 2017). Phylogenetic analysis represented by 14 isolates of D. rudis from
ten different hosts and 14 countries show species variation
of host and geographic ranges (Dissanayake et al. 2017).
The monophyletic clade has strong support with 66% MP
and 100% ML support and 1.00 BYPP (Fig. 102). Diaporthe
rudis was described as Sphaeria rudis on Laburnum anagyroides collected in France (Udayanga et al. 2014; Guarnaccia
and Crous 2017). Our strain (Fig. 105), which was isolated
from Clematis serratifolia in Belgium, differs from the holotype by having a thinner peridium (Udayanga et al. 2014).
Due to the genetic complexity especially in the ITS region of
D. rudis, the recombination parameters among selected taxa
were calculated as shown in Figs. 103b. The GCPSR result
of the concatenated gene set showed borderline insignificant
recombination between the isolates (Φw = 0.05, Fig. 103b).
Thus, we confirm the status of D. rudis according to pairwise homoplasy index and the phylogenetic results, which
highlights the high degree of genetic variation and distribution in a wide host range (Quaedvlieg et al. 2014; Guarnaccia and Crous 2017).
167
Tenuimurus clematidis MFLU 16-1971
100/1.00
Tenuimurus clematidis MFLUCC 14-0833 T
Ophiostoma gemellus CMW23059 T
0.02
Fig. 106 The Bayesian 50% majority-rule consensus phylogram
based on combined LSU, SSU and ITS sequence data for Phomatosporaceae members. The topology and clade stability of the combined
gene analyses was compared to the single gene analyses. The tree is
rooted with Ophiostoma gemellus (CMW23059). Ten strains were
included in the combined analyses which comprised of 2467 characters (883 characters for LSU, 1049 characters for SSU, 535 characters
for ITS, including gap regions). The tree from the maximum likelihood analysis had similar topology to the Bayesian analyses. The best
scoring RAxML tree had a final likelihood value of − 5571.410827.
The matrix had 292 distinct alignment patterns, with 31.28% of undetermined characters and gaps. Estimated base frequencies were as
follows; A = 0.253538, C = 0.239531, G = 0.293422, T = 0.213509;
substitution rates AC = 3.147737, AG = 3.686222, AT = 2.035694,
CG = 1.607623, CT = 10.739856, GT = 1.000000; gamma distribution shape parameter α = 0.540032. In our analysis, GTR + I + G
model was used for each partition in Bayesian posterior analysis.
The species determined in this study is indicated in blue. Bootstrap
values (BS) greater than 50% BS (ML, left) and Bayesian posterior
probabilities (BYPP, right) greater than 0.90 are given at the nodes.
Hyphens (-) represent support values less than 70% BS/0.90 BYPP.
Thick branches represent significant support values from all analyses
(BS ≥ 70%/BYPP ≥ 0.95) in genus level
ellipsoidal to fusiform, hyaline, aseptate to multi-septate
ascospores, with a rounded bipolar gelatinous caps, and
sporothrix-like asexual morphs reported from culture (Fallah and Shearer 1998; Fournier and Lechat 2010). Based on
the phylogenetic results of a combined LSU, SSU and ITS
dataset, the fungus collected on Clematis in Thailand clustered with other Phomatospora species (Fig. 106).
Phomatospora uniseriata Phukhams., M.V. de Bult & K.D.
Hyde, sp. nov.
Index Fungorum number: IF557302; Facesoffungi number: FoF 06577, Fig. 107.
Etymology: Based on the uniseriate arrangement of the
ascospores.
Holotype: MFLU 17–1476
13
168
Saprobic on Clematis subumbellata. Sexual morph:
Ascomata 185–220 × 115–185 μm ( x̄ = 195 × 175 μm, n = 5),
solitary to aggregated, immersed and becoming erumpent
with age, obpyriform to subglobose, coriaceous, brown to
dark brown, heavily pigmented at apex, ostiolate. Ostioles
50–60 × 50–55 μm ( x̄ = 57 × 53 μm, n = 5), papillate, central or eccentric, broadly conical, filled with periphyses.
Peridium 8–12 μm wide ( x̄ = 10 μm, n = 10), composed of
5–6 layers of thin-walled cells of textura prismatica mixed
with textura angularis, brown to reddish brown, comprising inner, hyaline, thick-walled cells. Paraphyses sparse,
1.6–3.4 μm ( x̄ = 2 μm, n = 30), hypha-like, thin-walled, septate, constricted at septa, tapering above and shorter than
the asci. Asci 90–135 × 3–5 μm ( x̄ = 103 × 4 μm, n = 20),
8-spored, unitunicate, cylindrical, thin-walled, pedicellate,
with a refractive, J-, apical ring. Ascospores 5.5–9 × 2–4 μm
( x̄ = 7 × 3 μm, n = 40), uniseriate, partially overlapping,
oblong-ellipsoidal, hyaline, pale brown when mature, unicellular, bi-guttulate, guttules located at the ends of the cell,
longitudinally striate. Asexual morph: Undetermined.
Culture characters: Colonies on MEA reaching 20 mm
within 4 weeks at 25 °C. Culture from above, cream, radiating to the edge, wrinkled, folded, margin undulate, medium
dense, flat or umbonate, and covered with spare grey aerial
mycelium; reverse white radiating outwardly.
Material examined: Thailand, Chiang Rai Province, on
dead stems of Clematis subumbellata, 20 March 2017, C.
Phukhamsakda, CMTH12 (MFLU 17–1476, holotype); extype living culture, MFLUCC 17–2068.
Host: Clematis subumbellata—(This study).
Distribution: Thailand—(This study).
GenBank accession numbers: LSU: MT214616; SSU:
MT226727; ITS: MT310660; tef1: MT394672; rpb2:
MT394720.
Notes: Phomatospora uniseriata (MFLUCC 17–2068)
formed a close relationship with P. biseriata which was
also recorded on Clematis vitalba in Italy (Senanayake et al.
2016). Phomatospora biseriata is similar to P. uniseriata
in having unitunicate, cylindrical, thin-walled, pedicellate
asci, with J-apical ring, overlapping uniseriate as ellipsoid,
hyaline, unicellular, bi-guttulate and longitudinally striate
ascospores (Senanayake et al. 2016). Phomatospora uniseriata has uniseriate, oblong ascospores that become pale
brown at senescence (Fig. 107). Phomatospora uniseriata
is distinguished from P. biseriata by its smaller asci and
ascospores (Table 5).
In the phylogenetic analysis, P. uniseriata (MFLUCC
17–2068) formed a close relationship with P. biseriata (50%
ML/1.00 BYPP, Fig. 106). A comparison of the ITS region
(including of the 5.8S region) showed 1.5% nucleotide (eight
nucleotide difference from 535 base pairs). This justifies
these two isolates as being two distinct taxa.
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Diaporthomycetidae, family incertae sedis
Distoseptisporaceae Hyde & McKenzie
Distoseptisporaceae contains Distoseptispora Hyde, E.
McKenzie & S. Maharachchikumbura as the generic type
(Fig. 108). We follow the latest treatment and updated
accounts of Distoseptisporaceae by Maharachchikumbura
et al. (2016) and Wijayawardene et al. (2017).
Dist osep tispora Hyde & E. McKenzie & S.
Maharachchikumbura
Distoseptispora is characterized by macronematous, septate, unbranched, short olivaceous to brown conidiophores,
monoblastic, determinate, terminal conidiogenous cells and
acrogenous, brown, euseptate or distoseptate conidia with
a basal cell with cross walls and a basal scar (Yang et al.
2018b). There are 16 species listed in Distoseptispora (Hyde
et al. 2016; Su et al. 2016; Tibpromma et al. 2017; Luo et al.
2018; Yang et al. 2018b). The combined gene phylogenetic
analyses revealed a new species, Distoseptispora clematidis, which constitutes the first record of Distoseptispora
on Clematis from Thailand (Fig. 108).
Distoseptispora clematidis Phukhams., M.V. de Bult & K.D.
Hyde, sp. nov.
Index Fungorum number: IF557301; Facesoffungi number: FoF 07261, Fig. 109.
Etymology: In reference to the host genus, Clematis.
Holotype: MFLU 17–1501
Saprobic on dried branches of Clematis sikkimensis. Colonies on the substratum superficial, effuse, scattered, hairy or
velvety, black. Mycelium immersed, composed of branched,
septate, smooth, dark brown hyphae. Sexual morph: Undetermined. Asexual morph: Conidiophores 22–40 × 4–10 μm
( x̄ = 25 × 7 μm, n = 10), macronematous, mononematous,
septate, unbranched, single or in groups of 2 or 3, erect,
straight or flexuous, 3–5-septate, smooth, dark brown to
brown, cylindrical, robust at the base. Conidiogenous cells
3–10 × 2–8 μm ( x̄ = 6 × 5 μm, n = 20), monoblastic, integrated, determinate, terminal, cylindrical, light brown.
Conidia 120–210 × 12–20 μm ( x̄ = 175 × 17 μm, n = 20),
acrogenous, solitary, dry, oblong, obclavate, cylindrical or
rostrate, elongated, straight or curved, truncate at the base,
rounded at the apex, 28–35-distoseptate, smooth, brown with
green tinge, lighter towards the apex, thick-walled, bud scars
or disjunctors present at the site of attachment.
Culture characters: Colonies on MEA reaching 30 mm
diam. after 4 weeks at 25 °C. Cultures from above, dark
brown, medium dense, circular, umbonate, fluffy; reverse
dark brown at the centre, black, radiating outwardly.
Material examined: Thailand, Chiang Rai Province, Doi
Tung, on dried stem of Clematis sikkimensis, 2 May 2017,
C. Phukhamsakda & M.V. de Bult, CMTHDT12 (MFLU
Fungal Diversity (2020) 102:1–203
Fig. 107 Phomatospora uniseriata (MFLU 17–1476, holotype). a
Appearance of ascomata on Clematis subumbellata. b Close up of
ascoma on host substrate. c Vertical section through ascoma. d Osti-
169
olar canal. e Peridium. f Paraphyses. g, h Asci i–l Ascospores. m Culture characteristics on MEA. Scale bars: b = 100 µm, c–h = 50 µm,
i–l = 5 µm
13
Mare Anglaise beach, Seychelles
(Hyde and Jones 1986)
20–46 × 7–11, unitunicate,
cylindrical or oblong-ventricose, short pedicellate
160–320 × 200–360, solitary,
globose to subglobose,
coriaceous
Lanspora coronata (AFTOLID 736)
13
Driftwood
90–135 × 3–4, unitunicate,
cylindrical, J-apical ring
185–220 × 115–185, solitary,
immersed, obpyriform to
subglobose, coriaceous
P. uniseriata (MFLU 17–1476)
Clematis subumbellata Chiang Rai, Thailand (this
study)
Clematis vitalba
25–29 × 9–11.5, overlapping
biseriate, ellipsoidal, hyaline,
unicellular, longitudinally
striate
5.5–9 × 2–4, overlapping, unicellular, oblong-ellipsoidal,
hyaline, pale brown at senescent states, longitudinally
striate
10–15 × 2.5–4, uni or biseriate,
ellipsoidal, 1-celled, hyaline,
with longitudinal wall striations, crown-like appendage
at both ends
200–230 × 19–23, unitunicate,
cylindrical to fusiform,
J-apical ring
115–170 × 125–200, solitary,
immersed, globose to subglobose, coriaceous
Phomatospora biseriata
(MFLU 16–1970)
Italy (Senanayake et al. 2016)
Host
Ascospores (μm)
Asci (μm)
Ascomata (μm)
Species
Table 5 A synopsis comparison of Phomatosporaceae species discussed in this study
Location/reference
Fungal Diversity (2020) 102:1–203
Fig. 108 Best scoring RAxML tree with a final likelihood value
of − 16025.131561 based on combined LSU, SSU, ITS and rpb2
sequence data for Distoseptisporaceae. The tree is rooted with Crytadelphia members. Thirty-seven strains were included in the combined
analyses which comprised 3444 characters (905 characters for LSU,
1022 characters for SSU, 454 characters for ITS, 1063 characters for
rpb2, including gap regions). The topology and clade stability of the
combined gene analyses was compared to the single gene analyses.
The tree from the maximum likelihood analysis had similar topology to the Bayesian 50% majority-rule consensus phylogram. The
matrix had 1081 distinct alignment patterns, with 40.90% undetermined characters and gaps. Estimated base frequencies were as follows; A = 0.252755, C = 0.239940, G = 0.285179, T = 0.222126;
substitution rates AC = 1.420436, AG = 2.672673, AT = 1.211428,
CG = 1.349930, CT = 6.537630, GT = 1.000000; gamma distribution shape parameter α = 0.699416. In our analysis, GTR + I + G
model was used for each partition in Bayesian posterior analysis.
The species determined in this study is indicated in blue. Bootstrap
values (BS) greater than 50% BS (ML, left) and Bayesian posterior
probabilities (BYPP, right) greater than 0.90 are given at the nodes.
Hyphens (-) represent support values less than 50% BS/0.90 BYPP.
Thick branches represent significant support values from all analyses
(BS ≥ 70%/BYPP ≥ 0.95) at genus and family level
17–1501, holotype); ex-type living culture, MFLUCC
17–2145.
Host: Clematis sikkimensis—(This study).
Distribution: Thailand—(This study).
GenBank accession numbers: LSU: MT214617; SSU:
MT226728; ITS: MT310661; rpb2: MT394721.
Notes: Distoseptispora clematidis (MFLUCC 17–2145)
is introduced based on both morphological (Fig. 109) and
phylogenetic evidence (Fig. 108). The morphology of D.
clematidis is similar to D. xishuangbannaensis, but distinguishable by the characters of the conidia. Distoseptispora
clematidis has smaller conidia (175 × 17 vs 244 × 11.5 μm)
and up to 35 septa, while D. xishuangbannaensis has 40
septa (Tibpromma et al. 2017). In the phylogenetic analysis,
D. clematidis forms a related lineage with D. fluminicola,
D. xishuangbannaensis and D. tectonigena with moderate
support (69% ML/0.99 BYPP). In a BLASTn search of GenBank, the closest match of the LSU sequence of MFLUCC
17–2145 was D. tectonigena (MFLUCC 12–0292) with
99.26% similarity to NG_059144, while the closest match
with the ITS sequence was 97.36% similarity to NR_154018.
This is the first record of Distoseptispora on Clematis.
Subclass: Xylariomycetidae Erikss. & W. Winka
The latest treatment of subclass Xylariomycetidae is by
Senanayake et al. (2015).
Amphisphaeriales Hawksw. & O.E. Erikss.
Several studies treated Amphisphaeriales as a synonym of
Xylariales. Senanayake et al. (2015) revised the taxonomic
relationship within Amphisphaeriales, and currently, 14
families are accepted by Hyde et al. (2020).
◂
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171
Distoseptispora fluminicola MFLUCC 15-0417 T
Distoseptispora fluminicola DLUCC 0999
Distoseptispora fluminicola DLUCC 0391
Distoseptispora clematidis MFLUCC 17-2145 T
Distoseptispora xishuangbannaensis KUMCC 17-0290 T
100/1.00
Distoseptispora tectonigena MFLUCC 12-0292 T
Distoseptispora submersa MFLUCC 16-0946 T
Distoseptispora tectonae MFLUCC 12-0291 T
Distoseptispora phangngaensis MFLUCC 16-0183 T
Distoseptispora cangdhanensis MFLUCC 16-0970 T
Distoseptispora thailandica MFLUCC 16-0270
Distoseptispora multiseptata MFLUCC 15-0609 T
69/0.99
--/0.98
65/0.98
91/0.99
--/0.95
61/-67/0.94
84/0.98
--
76/---/0.98
100/1.00
Distoseptispora multiseptata MFLU 15-1144
Distoseptispora multiseptata MFLU 17-0856
Distoseptispora adscendens HKUCC 10820
Distoseptispora cangshanensis MFLUCC 16-0970 T
100/1.00
Distoseptispora cangshanensis MFLU 18-0474
Distoseptispora rostrata DLUCC 0885
50/--
89/0.90
94/0.98
61/-60/--
79/0.98
Distoseptispora
Distoseptispora rostrata MFLU 18-0479
Distoseptispora rostrata MFLUCC 16-0969 T
Distoseptispora obpyriformis MFLUCC 17-1694 T
Distoseptispora obpyriformis DLUCC 0867
Distoseptisporaceae
98/0.99
98/1.00
Distoseptispora thysanolaenae KUN-HKAS 102247 T
Distoseptispora
guttulata MFLUCC 16-0183 T
58/0.96
Distoseptispora guttulata MFLU 17-0852
54/0.91
Distoseptispora sp. MFLUCC 18-0417
-100/0.99 Distoseptispora sp. MFLUCC 18-1045
Distoseptispora aquatica WHXM 13-1-S-036
100/1.00
Distoseptispora suoluoensis MFLUCC 17-1305
100/1.00
-Distoseptispora suoluoensis MFLUCC 17-0224 T
Aquapteridospora lignicola MFLU 15-1172 T
100/1.00
Distoseptispora leonensis HKUCC 10822 T
Sporidesmium
pyriformatum MFLUCC 15-0620 T
100/1.00
Sporidesmium
thailandense
MFLUCC 15-0964 T
Sporidesmiaceae
-Sporidesmium tropicale MFLUCC 16-0185 T
Cryptadelphia groenendalensis SMH3767 T
(Outgroup)
100/1.00
Cryptadelphia groenendalensis SH 12
65/--
--
0.03
13
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Fig. 109 Distoseptispora clematidis (MFLU 17–1501, holotype). a, b. Colonies on the substratum. c Conidiophore and conidia. d Conidiophore. e, f Conidia. g, h Culture characteristics on MEA. Scale bars: b = 200 μm, c, e–f = 100 μm, d = 20 μm
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Sporocadaceae Corda [as “Sporocadeae”], Icon. Fung.
(Prague) 5: 34. 1842
Sporocadaceae was resolved by Liu et al. (2019) and
comprises 30 genera. Members of Sporocadaceae are saprobic or pathogenic on various plants (Senanayake et al. 2015).
The family is recognised by having immersed ascomata and
uniseriate, brown, ovoid to elliptic, straight or inequilateral,
septate ascospores. The asexual morph characters of some
Sporocadaceae are usually termed “pestaloid”, referring to
acervular or pycnidial conidiomata, ellipsoid to clavate, or
fusiform conidia that are hyaline, pale olivaceous or brown,
septate, with polar appendages (Guba 1961; Maharachchikumbura et al. 2014; Senanayake et al. 2015; Liu et al.
2019).
Pestalotiopsis Stewart
Pestalotiopsis is characterized by acervular conidioma
and fusoid to oval, 4-euseptate, pigmented conidia, usually two to five apical appendages arising as tubular extensions from the apical cell, with a centric basal appendage
(Maharachchikumbura et al. 2014). Pestalotiopsis occurs as
an endophyte or saprobe in various plants and often causes
disease symptoms on leaves and fruit in many plants (Liu
et al. 2006; Maharachchikumbura et al. 2012; Chen et al.
2018). Pestalotiopsis verruculosa was associated with Clematis vitalba branches in Italy. The multigene phylogenetic
analysis of ITS and tub sequence data for Pestalotiopsis is
shown in Fig. 110.
Pestalotiopsis verruculosa Maharachch. & K.D. Hyde, in
Maharachchikumbura et al. Fungal Diversity 56(1): 123
(2012), new host record
Index Fungorum number: 800527; Facesoffungi number:
FoF 07336, Fig. 111.
Saprobic on dead stem of Clematis vitalba. Conidiomata
pycnidial in culture on PDA, globose to oval, solitary or
aggregated in clusters, semi-immersed, black, 120–320 μm
diam., exuding globose, black, glistening, conidial masses.
Conidia 21–33 × 6–15 μm ( x̄ = 27 × 9 μm, n = 40), fusiform
or clavate-fusiform, straight or slightly curved, 4-septate,
concolorous; basal cell conical, hyaline, thin and verruculose, 2–5 μm long ( x̄ = 3.4 μm, n = 15), three median cells
doliiform, olivaceous, concolorous, veruculose, 14–16 μm
long, septa and periclinal walls darker than the rest of the
cell (second cell from base 2–6 μm; third cell 5–9 μm; fourth
cell 3.4–6.9 μm long); apical cell 1.6–5.4 μm long, hyaline, cylindrical to subcylindrical with 3–6 tubular apical
appendages (mostly 3), arising from crown of the apical cell,
173
unbranched, filiform 11–34 μm ( x̄ = 20 μm, n = 60) long; single, unbranched, filiform basal appendage present, 6–10 μm
long ( x̄ = 7.5 μm, n = 60).
Culture characters: Colonies on PDA reaching 30 mm
diam. after 7 days at 25 °C. Culture from above,edge undulate, white to pale yellow, aerial mycelium with black fruiting bodies, concentric; reverse of culture, yellow to pale
orange.
Material examined: Italy, Forlì-Cesena Province, Santa
Sofia–near Corniolo, on dead and terrestrial stem of
Clematis vitalba, 5 February 2013, E. Camporesi (MFLU
14–0624), living culture MFLUCC 14–1091.
Host: Clematis vitalba, Rhododendron sp.—(Maharachchikumbura et al. 2012; this study).
Distribution: China, Italy—(Maharachchikumbura et al.
2012; this study).
GenBank accession numbers: ITS: MT186199; tub2:
MT199995; tef1: MT199996.
Notes: This strain of Pestalotiopsis verruculosa has somewhat smaller conidia (21–33 × 6–15 μm) when compared
with the type strain of P. verruculosa (28–35 × 9–11 μm)
and shorter apical appendages (11–34 vs 25–40 μm) (Maharachchikumbura et al. 2012). In the phylogenetic analyses,
our strain clustered with the type strain of P. verruculosa
(Fig. 111). Pestalotiopsis verruculosa has been recorded
only from Rhododendron sp. in China as an endophyte
(Maharachchikumbura et al. 2012). This study provides the
first report of P. verruculosa from Clematis vitalba.
Xylariales Nannf.
The Xylariales is a large order in Sordariomycetes, with
20 accepted families. We follow the treatments by Maharachchikumbura et al. (2016), Li et al. (2017), and Hyde
et al. (2020).
Diatrypaceae Nitschke
Species of Diatrypaceae are highly diverse and widespread on woody plants in terrestrial, aquatic or marine
(Trouillas and Gubler 2010; Abdel-Wahab et al. 2014; Liu
et al. 2015; Dayarathne et al. 2016; Senwanna et al. 2017;
Shang et al. 2017). Based on the latest treatment and updated
accounts of Diatrypaceae, the family contains 20 genera
(Maharachchikumbura et al. 2015, 2016; Shang et al. 2017;
Phookamsak et al. 2019). Diatrypaceae is characterized by
perithecial ascomata, with poorly or well-developed ascostromata, immersed to erumpent in the host substrates, with
papillate ascomata (Fuckel 1870). Based on phylogenetic
analysis of a combined ITS and tub dataset (Fig. 112), we
13
Fungal Diversity (2020) 102:1–203
--/60/1.00
--/100/1.00
--/90/1.00
Pestalotiopsis trachicarpicola IFRDCC 2240 T
Pestalotiopsis rhodomyrtus HGUP4230 T
58/70/0.98
Pestalotiopsis oryzae CBS 353.69 T
ML/MP/BYPP
Pestalotiopsis kenyana CBS 442.67 T
Pestalotiopsis australasiae CBS 114126 T
50/--/1.00 60/57/1.00 Pestalotiopsis telopeae CBS 114161 T
Pestalotiopsis grevilleae CBS 114127 T
-Pestalotiopsis knightiae CBS 114138 T
Pestalotiopsis sp. LC3637
Pestalotiopsis biciliata CBS 124463 T
Pestalotiopsis brachiata LC2988 T
-- 84/81/1.00 Pestalotiopsis rhizophorae MFLUCC 17-0416 T
Pestalotiopsis rhizophorae MFLUCC 17-0417
79/90/1.00
Pestalotiopsis thailandica MFLUCC 17-1616 T
51/--/1.00
Pestalotiopsis thailandica MFLUCC 17-1617
Pestalotiopsis formosana NTUCC 17-009 T
84/71/1.00 63/99/0.99
Pestalotiopsis formosana NTUCC 17-010
100/100/1.00
Pestalotiopsis parva CBS 265.37
100/100/1.00 Pestalotiopsis parva CBS 278.35 T
Pestalotiopsis digitalis ICMP 5434 T
Pestalotiopsis rosea MFLUCC 12-0258 T
--/72/1.00
Pestalotiopsis dracontomelon MFUCC 10-0149 T
61/70/0.96
Pestalotiopsis colombiensis CBS 118553 T
76/90/1.00
75/72/1.00
Pestalotiopsis jinchanghensis LC6636 T
83/79/1.00
Pestalotiopsis aggestorum LC6301 T
Pestalotiopsis licualacola HGUP4057 T
Pestalotiopsis adusta ICMP 6088 T
83/79/1.00
Pestalotiopsis papuana CBS 331.96 T
--/70/0.99
Pestalotiopsis malayana CBS 102220 T
Pestalotiopsis diploclisiae CBS 115587 T
99/100/1.00
Pestalotiopsis humus CBS 336.97 T
76/61/1.00
Pestalotiopsis shorea MFLUCC 12-0314 T
50/68/-Pestalotiopsis neolitseae NTUCC 17-011 T
Pestalotiopsis neglecta TAP1100 T
100/100/1.00
Pestalotiopsis distincta LC3232 T
100/100/1.00 Pestalotiopsis intermedia MFLUCC 12-0259 T
Pestalotiopsis linearis MFLUCC 12-0271 T
74/70/1.00
Pestalotiopsis unicolor MFLUCC 12-0276 T
58/--/-Pestalotiopsis chamaeropis CBS 186.71 T
85/90/1.00
Pestalotiopsis jiangxiensis LC4399 T
Pestalotiopsis australis CBS 114193 T
74/89/1.00
Pestalotiopsis scoparia CBS 176.25 T
98/100/1.00
Pestalotiopsis brassicae CBS 170.26 T
Pestalotiopsis hollandica CBS 265.33 T
77/--/0.96
Pestalotiopsis italiana MFLUCC 12-0657 T
54/--/1.00
Pestalotiopsis verruculosa MFLUCC 12-0274 T
98/100/1.00
99/90/1.00 95/80/1.00 Pestalotiopsis verruculosa MFLUCC 14-1091
Pestalotiopsis monochaeta CBS 144.97 T
93/91/1.00
Pestalotiopsis lushanensis LC4344 T
97/89/1.00
Pestalotiopsis rhododendri IFRDCC 2399 T
99/100/1.00
Pestalotiopsis clavata MFLUCC 12-0268 T
Pestalotiopsis inflexa MFLUCC 12-0270 T
100/100/1.00
Pestalotiopsis portugalica CBS 393.48 T
Pestalotiopsis anacardiacearum IFRDCC 2397 T
100/100/1.00
Pestalotiopsis hawaiiensis CBS 114491 T
56/58/1.00
Pestalotiopsis arengae CBS 331.92 T
100/90/1.00
--/70/-Pestalotiopsis pallidotheae MAFF 240993 T
50/70/-Pestalotiopsis arceuthobii CBS 434.65 T
Pestalotiopsis ericacearum IFRDCC 2439 T
100/--/1.00
Pestalotiopsis gaultheria IFRD 411.014 T
100/100/1.00
--/70/1.00
99/100/1.00
Pestalotiopsis gibbosa NOF 3175 T
50/50/-Pestalotiopsis spathulata CBS 356.86 T
90/100/1.00
Pestalotiopsis diversiseta MFLUCC 12-0287 T
Pestalotiopsis novae-hollandiae CBS 130973 T
Pestalotiopsis camelliae MFLUCC 12-0277 T
--/80/1.00
Pestalotiopsis yanglingensis LC4553 T
--/80/1.00
Pestalotiopsis furcata MFLUCC 12-0054 T
Pestalotiopsis longiappendiculata LC3013 T
Pestalotiopsis jesteri CBS 109350 T
Pestalotiopsis montellica MFLUCC 12-0279 T
Neopestalotiopsis saprophytica MFLUCC 12-0282 T
40.0
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Pestalotiopsis
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Fungal Diversity (2020) 102:1–203
◂Fig. 110 One of the 1000 most parsimonious trees obtained from a
heuristic search of combined ITS and tub sequence data for Pestalotiopsis. The tree is rooted at Neopestalotiopsis saprophytica (MFLUCC
12-0282). Maximum parsimony and maximum likelihood support ≥ 50%, Bayesian posterior probabilities ≥ 0.90 (ML/MP/BYPP)
are given at the nodes. The species obtained in this study is in blue.
Ex-type (T) taxa from other studies are in bold black
report a new record of N. baoshanensis on Clematis vitalba
from Europe.
Neoeutypella Raza, Shang, Phookamsak & L. Cai
Neoeutypella was introduced as a monotypic genus for
eutypella-like strains that clustered among Diatrypella
species but are distantly related to Eutypella sensu stricto
(Dayarathne et al. 2016). Neoeutypella is characterized
by entostromatic, carbonaceous ascostromata, perithecial,
black, immersed to semi-immersed, papilla filled with
periphyses, thick peridium, 8-spored spindle-shaped, and
long pedicellate. Asexual morph of Neoeutypella produce
hyphae-like, branched, filiform, hyaline or pale brown
conidia in culture conditions (Crous et al. 2016; Phookamsak et al. 2019). Phylograms generated from maximum likelihood and Bayesian probability analyses based on combined
ITS and tub sequence data (Fig. 112) showed that our collection from Clematis vitalba clustered with the type species of
Neoeutypella. Thus, the new host record of N. baoshanensis
is reported (Fig. 113).
Neoeutypella baoshanensis Raza, Shang, Phookamsak &
L. Cai, new host record
Synonym: Eutypella caricae (De Not.) Berl.: [1] (1902)
Index Fungorum number: IF555372; Facesoffungi number: FoF 04928, Fig. 113.
Saprobic on dead stem of Clematis vitalba. Sexual
morph: Stromata poorly developed, flask-shaped, multiloculate, with 8–12 locules forming groups in stromata,
black, erumpent or semi-immersed in the surface layers of the host tissue. Ascomata partially or deeply
immersed in wide-spread stroma, 550–620 × 350–600 μm
( x̄ = 585 × 500 μm, n = 10), globose, reddish brown to black,
single or gregarious and sometimes confluent, ostiolate.
Ostioles 110–140 × 90–125 μm ( x̄ = 130 × 110 μm, n = 10),
oblong-conical, with papillate, immersed in a wide-spread
entostroma which protrudes above the substrate surface,
periphyses. Peridium 13–40(–80 μm at apex) wide, comprising an outer, brown, thick-walled layer of polygonal
melanized cells, interspersed with cells of the substrate and
several inner, hyaline, thick-walled cell layers of textura
angularis. Paraphyses composed of numerous, 1.8–3.5 μm
wide, aseptate, paraphyses, narrowing and tapering towards
the apex. Asci 75–120 × 7–12 μm ( x̄ = 100 × 8 μm, n = 20),
multi-spored, unitunicate, with narrow, thin-walled pedicel,
40–70 μm long, with cylindrical, swollen upper portion,
175
apex flat, with J-, cylindrical, conspicuous ring. Ascospores
10–15 × 2–4 μm ( x̄ = 13 × 3 μm, n = 50), hyaline or pale reddish brown to brown, allantoid, straight or slightly curved,
unicellular, thin-walled with 2 small globules in each cell,
smooth-walled. Asexual morph: Undetermined.
Culture characters: Colonies on MEA reaching 30 mm
diam. after 4 weeks at 25 °C. Culture from above, white,
yellowish towards the edge, erumpent, spreading, surface
folded, margins lobate, medium dense, circular, flat, dull,
fimbriate, radially furrowed and slightly covered with white
aerial mycelium; reverse yellowish with radiating cream
mycelium.
Material examined: UK, Hampshire, Botley Wood, on
dead stems of Clematis vitalba, 16 April 2016, E.B.G.
Jones, GJ 279 (MFLU 17–1461); living culture, MFLUCC
16–1002.
Hosts: Clematis vitalba, Ficus carica, Pinus armandii—
(Acero et al. 2004; Phookamsak et al. 2019; this study)
Distribution: China, France, UK—(Acero et al. 2004;
Phookamsak et al. 2019; this study)
GenBank accession numbers: LSU: MT214618; SSU:
MT226729; ITS: MT310662; tef1: MT394673.
Notes: Our new isolate MFLUCC 16–1002 clustered
with the type species of Neoeutypella baoshanensis strain
LC 12111 (Fig. 112) with moderate support (98% ML/0.95
BYPP). Isolate MFLUCC 16–1002 is similar to the type
species reported by Phookamsak et al. (2019), Fig. 113. In
the phylogenetic analysis, Diatrype macowaniana strain
CBS 214.87 clustered among Neoeutypella baoshanensis
strains. Although Acero et al. (2004) mentioned that N.
baoshanensis (= Eutypella caricae) and Diatrype macowaniana have no sequence divergence; morphological
comparison of Diatrype macowaniana and N. baoshanensis were distinct in having smaller asci (32–36 × 6
vs 77.5 × 8 μm) and hyaline ascospores (10.8 × 2.4 vs
5–7 × 1.5–2) (Saccardo 1882). Thus, we maintain the strain
as Diatrype macowaniana until the paratype specimen is
re-collected. Diatrypella banksiae CPC 29054 and CPC
29118 formed a strongly supported clade with Neoeutypella. However, Diatrypella banksiae is only present as
an asexual morph, thus we maintain the strain under the
current name (Crous et al. 2016).
Hypocreomycetidae Erikss. & K. Winka
Glomerellales Chadef. ex Réblová et al.
Glomerellales includes Australiascaceae, Glomerellaceae,
Malaysiascaceae, Plectosphaerellaceae and Reticulascaceae
(Réblová et al. 2011; Maharachchikumbura et al. 2016; Tibpromma et al. 2018).
Plectosphaerellaceae Gams, Summerb. & R. Zare
Plectosphaerellaceae was introduced by Zare et al.
(2007) to accommodate a monophyletic lineage within
13
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Fungal Diversity (2020) 102:1–203
Fig. 111 Pestalotiopsis verruculosa (MFLU 14-0624) a, b Appearance of conidiomata on Clematis vitalba. c–e Conidia with concolorous
median cells. f Colony on PDA (upper view) g Colony on PDA (reverse view). Scale bars: c–e = 10 μm
Glomerellales. Plectosphaerellaceae taxa are fungicolous, insecticolous, plant pathogens, saprobic, and cause
soil-borne diseases (Cannon et al. 2012; Duc et al. 2009;
Grum-Grzhimaylo et al. 2016; Giraldo et al. 2019). The
family is mainly represented by asexual morphs with only
a few sexual morphs described by Carlucci et al. (2012),
Grum-Grzhimaylo et al. (2013, 2016) and Phookamsak et al.
(2019). Phylogenetic relationships of the intra-members of
Plectosphaerellaceae were analysed based on a LSU, ITS,
tef1 and rpb2 dataset (Fig. 114). Combining morphological
and molecular data, one new genus Xenoplectosphaerella
and one new combination Fuscohypha kunmingensis are
introduced below.
Fuscohypha Giraldo López & P. Crous
Fuscohypha is typified by F. expansa Giraldo & Crous.
The genus is characterized by branched conidiophores with
ellipsoidal to cylindrical conidia, pale brown in mass, with
slimy heads. Fuscohypha expansa was isolated from soil
and tuber of Dioscorea. In our study, Fuscohypha formed a
separate clade basal to Plectosphaerella and Brunneochlamydosporium (Fig. 114).
Fuscohypha kunmingensis (Phookamsak, J.F. Li & K.D.
Hyde) Jayaward., Phukhams., & K.D. Hyde, comb. nov.
13
Index Fungorum number: IF557607; Facesoffungi number: FoF 05716
Basionym: Plectosphaerella kunmingensis Phookamsak,
J.F. Li & K.D. Hyde, in Phookamsak et al., Fungal Diversity
95:1–273(2019)
Notes: Fuscohypha kunmingensis was described as Plectosphaerella kunmingensis based on the morphological
comparison and the strain formed a basal clade to Plectosphaerella species (Phookamsak et al. 2019). Phylogenetic
inference in this study places the ex-type strain of Fuscohypha kunmingensis (KMUCC 18–0181) in a close relationship with the type species of Fuscohypha (F. expansa) with
good support (81% ML/100% MP/0.99 BYPP, Fig. 114).
Fuscohypha kunmingensis was a contaminant in the laboratory. The strain is similar to Fuscohypha as they both lack
hyphal coils and hyaline to yellowish brown conidiophores
(Giraldo et al. 2019).
Host: Air-bone fungus—Phookamsak et al. (2019).
Distribution: China—Phookamsak et al. (2019).
Xenoplectosphaerella Jayaward., Phukhams., & K.D. Hyde,
gen. nov.
Index Fungorum number: IF557303; Facesoffungi number: FOF 01334, Fig. 115.
Fungal Diversity (2020) 102:1–203
177
Diatrypella vulgaris HVFRA02
99/-100/--
Diatrypella vulgaris HVGRF03
Diatrypella frostii UFMGCB 1917
Diatrypella delonicis MFLU 16-1032
93/--
99/0.98
Diatrypella delonicis MFLUCC 15-1014 T
Diatrypella major 1058
93/0.99
96/1.00
62/--
Diatrypella
Diatrypella atlantica HUEFS 194228
Diatrypella atlantica HUEFS 136873
Diatrypella heveae MFLUCC 17-0368 T
78/0.97
Diatrypella tectonae MFLUCC 12-0172a T
97/0.99
Diatrypella tectonae MFLUCC 12-0172b
Diatrypella verruciformis UCROK1467 T
100/1.00
66/0.97
71/0.96
Diatrypella verruciformis UCROK754
Neoeutypella baoshanensis CBS 274.87
Neoeutypella baoshanensis MFLUCC 16-1002
62/0.95
98/0.95
--
Neoeutypella baoshanensis LC 12111 T
Diatrype macowaniana CBS 214.87 T Neoeutypella
Neoeutypella baoshanensis GL08362
100/1.00
100/1.00
96/1.00
Diatrypella banksiae CPC 29054 T
Diatrypella banksiae CPC 29118
Allodiatrype mangrovie MFLUCC 17-0391
97/0.99
100/1.00
Diatrype enteroxantha HUEFS 155116
100/1.00
100/1.00
Allodiatrype
Allodiatrype mangrovie MFLUCC 17-0412 T
Diatrype enteroxantha HUEFS 155114
Pedumispora rhizophorae BCC 44877 T
Diatrype
Pedumispora
Pedumispora rhizophorae BCC 44878
Halodiatrype salinicola MFLUCC 15-1277 T
100/1.00
Halodiatrype avcenniae MFLUCC 15-0948 T
0.03
Fig. 112 Best scoring RAxML tree with a final likelihood value
of − 4428.491036 based on combined ITS and tub sequence data
of related species. The tree is rooted with Halodiatrype avcenniae
(MFLUCC 15–0948) and H. salinicola (MFLUCC 15–1277) in Diatrypaceae. Twenty-eight strains were included in the combined gene
analyses which comprised 1393 characters (623 characters for ITS,
770 characters for tub, including gap regions). The tree from the
maximum likelihood analysis had similar topology to the Bayesian
analysis. The matrix had 396 distinct alignment patterns with 40.26%
undetermined characters or gaps. Estimated base frequencies were as
follows: A = 0.224095, C = 0.268057, G = 0.236876, T = 0.270973;
substitution rates AC = 1.104749, AG = 3.483519, AT = 0.878988,
CG = 1.108511, CT = 4.234084, GT = 1.000000; gamma distribution
shape parameter α = 0.276482. In our analysis, GTR + I + G model
was used for each partition in Bayesian posterior analysis. The species determined in this study is indicated in bold blue. Bootstrap
values (BS) greater than 50% BS (ML, left) and Bayesian posterior
probabilities (BYPP, right) greater than 0.90 are given at the nodes.
Hyphens (-) represent support values less than 50% BS/0.90 BYPP.
Thick branches represent significant support values from all analyses
at genus level (BS ≥ 70%/BYPP ≥ 0.95)
13
178
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Fungal Diversity (2020) 102:1–203
◂Fig. 113 Neoeutypella baoshanensis (MFLU 17–1461). a Appear-
ance of stromata on Clematis vitalba. b Close up of stromata on host
substrate. c Horizontal section through stroma. d Vertical section
through stroma. e Vertical section through ascoma. f Ostiolar canal. g
Peridium. h Aseptate paraphyses. i, j Asci. k–m Ascospores. n Germinated ascospore. o, p Culture characteristics on MEA. Scale bars:
a = 1 mm, b–e = 500 µm, f = 200 µm, g–j = 50 µm, k–m = 10 µm
Etymology: Name refers to the similarity of its morphology to Plectosphaerella.
Saprobic on herbaceous plant in terrestrial habitats. Sexual morph: Ascomata single or gregarious, erumpent or
immersed under the host epidermal layer, visible as black
spot on host substrate, papillate. Papilla filled with periphyses. Peridium composed of thin-walled cells of textura
angularis mixed with textura globosa, multilayered, inner
layers, hyaline. Paraphyses numerous, septate. Asci unitunicate, broad clavate to spathulate, with simple pedicel,
conspicuous apical or subapical, J-, ring. Ascospores biseriate, overlapping, fusiform-elliptical to ellipsoid, 1 septum,
hyaline, constricted at the septum, straight or slightly curved
towards the apex. Asexual morph: Undetermined.
Type species: Xenoplectosphaerella clematidis Jayaward., Phukhams., & K.D. Hyde
Notes: Xenoplectosphaerella is introduced as a monotypic genus in Plectosphaerellaceae (Fig. 115). The genus
was associated with a herbaceous plant in Thailand and
formed obpyriform, coriaceous ascomata with papilla, with
paraphyses, and uniquely spathulate asci (Carlucci et al.
2012; Grum-Grzhimaylo et al. 2016; Giraldo et al. 2019).
In the BLASTn search of GenBank, the closest match of
the LSU region of the new species was Musicillium theobromae (CBS 968.72) with 97.22% similarity (accession
number LR025907). The closest match of the ITS region of
MFLUCC 17–2067 was also M. theobromae (CBS 968.72)
with 90.83% similarity (accession number NR_156205).
However, Musicillium is phylogenetically distant from our
new isolate (Fig. 114). Phylogenetic analyses of sequence
data of taxa within Plectosphaerellaceae resolves Xenoplectosphaerella as basal to Musicillium and Paramusicillium,
which are strains isolated from soil and monocotyledons or
the mushroom genus Lactarius sp. (Giraldo et al. 2019).
We name our collection as Xenoplectosphaerella clematidis
MFLUCC 17–2067, a new genus which formed a distinct
lineage from other genera in the family (Fig. 114).
Xenoplectosphaerella clematidis Jayaward., Phukhams., &
K.D. Hyde, sp. nov.
Index Fungorum number: IF557304; Facesoffungi number: FoF 07326, Fig. 115.
Etymology: Name refers to the host plant, Clematis.
Holotype: MFLU 17–1475
179
Saprobic on dead stems of Clematis subumbellata. Sexual
morph: Ascomata 90–100 × 90–110 μm ( x̄ = 98 × 105 μm,
n = 5), single or gregarious, immersed under the host epidermal layer, visible as black spots on host substrate, obpyriform, coriaceous with minute papilla. Ostioles papillate,
filled with periphyses. Peridium 7–12 μm wide, composed
of 7–10 layers of thin-walled cells of textura angularis
mixed with textura globosa, inner layers, hyaline, thickwalled. Paraphyses numerous, 1.5–3.5 μm ( x̄ = 2.5 μm,
n = 30), septate, narrowed and tapering towards the apex.
Asci 31–50 × 10–15 μm ( x̄ = 38 × 12 μm, n = 20), unitunicate,
broad clavate to broad fusiform, simple pedicel, apex flat,
with J-, conspicuous apical or subapical ring. Ascospores
10–17 × 4–5 μm ( x̄ = 13 × 4 μm, n = 50), biseriate, overlapping, fusiform-elliptical to ellipsoid, hyaline, 1 septum, constricted at the median septum, straight or slightly curved
towards the apex, thin-walled, with two guttules in each cell,
smooth-walled. Asexual morph: Undetermined.
Culture characters: Colonies on MEA reaching 20 mm
diam. after 4 weeks at 25 °C. Culture from above, black,
radiating, wrinkled, folded, margin undulate, dense, flat or
umbonate, and slightly covered with grey aerial mycelium;
reverse black with radiating, brown mycelium.
Material examined: Thailand, Chiang Rai Province, on
dead stems of Clematis subumbellata, 20 March 2017, C.
Phukhamsakda, CMTH11 (MFLU 17–1475, holotype); extype living culture, MFLUCC 17–2067.
Host: Clematis subumbellata—(This study).
Distribution: Thailand—(This study).
GenBank accession numbers: LSU: MT214619; SSU:
MT226730; ITS: MT310663; tef1: MT394674; rpb2:
MT394722.
Notes: The novel species resembles other genera in Plectosphaerellaceae (e.g. Fuscohypha and Plectosphaerella) by
its coriaceous ascomata and fusiform-elliptical to ellipsoid, 1
septum ascospores (Carlucci et al. 2012; Giraldo et al. 2019;
Phookamsak et al. 2019). The species has unique characters such as paraphyses and spathulate, simple pedicel asci.
Moreover, Fuscohypha and Plectosphaerella are phylogenetically not closely related (Fig. 114).
Hypocreales Lindau
We followed recent treatment by Hyde et al. (2020b).
Nectriaceae Tul. & C. Tul. [as ‘Nectriei’]
Nectriaceae is a species-rich family which includes
important plant and human pathogens (Lombard et al. 2015).
Members of the family are distinguishable by characters
such as densely gregarious, uniloculate with yellow, orangered to purple ascomata (Rossman et al. 1999; Lombard et al.
2015). Asexual morph are reported as synnematous, sporodochial or pycnidial with phialidic conidiogenesis (Rossman
13
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Fungal Diversity (2020) 102:1–203
Plectosphaerella pauciseptata CBS 131745
Plectosphaerella humicola CBS 423.66
Plectosphaerella plurivora CBS 131742
97/92/1.00
Plectosphaerellaramiseptata CBS 131861
84/78/0.99
87/92/1.00
Plectosphaerella oligotrophica CBS 440.90
93/100/1.00
Plectosphaerellacitrullae CBS 131741
76/--/0.89
Plectosphaerella cucumerina CBS 137.37
Plectosphaerella oratosquillae NJM 0662
98/94/1.00
Plectosphaerella melonis CBS 489.96
-Plectosphaerella delsorboi CBS 116708
95/90/1.00
54/--/-Plectosphaerella alismatis CBS 113362
-Plectosphaerella sinensis ACCC 39145
Plectosphaerella populi CBS 139623
98/90/1.00
Brunneochlamydosporium terrestre CBS 112777
Brunneochlamydosporium nepalense
100/90/1.00 100/90/1.00
Brunneochlamydosporium cibotii CBS 109240
Brunneochlamydosporium macroclavatum CBS 101249
Fuscohypha kunmingensis KUMCC 18-0181
81/100/0.99
Fuscohypha expansa CBS 418.89
100/90/1.00
Musidium stromaticum CBS 863.73
62/64/0.99
--/52/-Sayamraella subulata BCC 78964
77/75/1.00
Summerbellia oligotrophica CBS 657.94
58/55/0.99
Theobromium fuscum CBS 112271
95/77/1.00
Chordomyces albus CBS 987.87
100/100/1.00
Chordomyces antarcticus CBS 120045
97/90/1.00
Furcasterigmium furcatum CBS 122.42
Paragibellulopsis chrysanthemi MAFF 242621
Gibellulopsis aquatica CBS 117131
-Phialoparvum bifurcatum CBS 299.70B
Musicillium tropicale CBS 120009
72/97/0.99
100/100/1.00
86/77/1.00
Musicillium elettariae CBS 252.80
100/100/1.00
Musicillium theobromae CBS 968.72
53/50/0.99
Paramusicillium asperulatum CBS 120158
Xenoplectosphaerella clematidis MFLUCC 17-2067
s
Plectosphaerellaceae
81/90/0.99
99/100/1.00
ML/MP/BYPP
--/--/1.00
Chlamydosporiella restricta CBS 178.40
Nigrocephalum collariferum CBS 124586
Stachylidium bicolor CBS 121802
91/100/1.00
Stachylidium pallidum BCC 79031
Brunneomyces brunnescens CBS 559.73
Lectera capsici CBS142534
Verticillium albo-atrum CBS 130340
Acrostalagmus annulatus CBS 121.84
Acrostalagmus luteoalbus CBS 577.78B
Sodiomyces alcalophilus CBS 114.92
--/100/1.00
68/67/0.99
63/50/1.00
51/50/0.97
100/100/1.00
99/90/1.00
97/99/1.00
100/--/--
CBS 379.77
CBS 869.66
--
Monilochaetes infuscans
0.1
Fig. 114 The best scoring RAxML tree with a final likelihood value
of − 23259.637010 of ITS, LSU, tef1 and rpb2 sequence data for
Plectosphaerellaceae. The topology and clade stability of the combined gene analyses was compared to the single gene analyses. The
tree is rooted at Monilochaetes infuscans (CBS 379.77 and CBS
869.66). The matrix had 1093 distinct alignment patterns, with
13.22% of undetermined characters and gaps. Estimated base frequencies were as follows; A = 0.224235, C = 0.297727, G = 0.281301,
T = 0.196738; substitution rates AC = 0.820050, AG = 1.988895,
13
AT = 1.053617, CG = 0.783458, CT = 5.160854, GT = 1.000000;
gamma distribution shape parameter α = 0.2928059. The new species
in this study are indicated in blue. Bootstrap values (BS) greater than
50% BS (maximum likelihood (left); maximum parsimony (middle))
and Bayesian posterior probabilities (BYPP, right) greater than 0.90
are given at the nodes. Hyphens (-) represent support values less than
50% BS/0.90 BYPP. Thick branches represent significant support values from all analyses (BS ≥ 70%/BYPP ≥ 0.95) at the genus level
Fungal Diversity (2020) 102:1–203
Fig. 115 Xenoplectosphaerella clematidis (MFLU 17–1475, holotype). a Appearance of ascomata on Clematis subumbellata. b Close
up of ascoma on host substrate. c Vertical section through ascoma. d
181
Peridium. e Paraphyses (in cotton blue). f, g Asci. h–j Ascospores. k,
l Culture characteristics on MEA. Scale bars: b = 100 µm, c = 50 µm,
d–g = 20 µm, h–j = 10 µm
13
182
Fungal Diversity (2020) 102:1–203
Fusarium
Fusarium acaciae mearnsii NRRL 26755
F. graminearum NRRL 31084
F. asiaticum NRRL 13818
92
F. culmorum NRRL 25475
F. pseudograminearum NRRL 28062
F. dactylidis NRRL 29298
F. praegraminearum NRRL 39664
F. langsethiae NRRL 54940
98
F. sibiricum NRRL 53430
F. sporotrichioides NRRL 3299
85
F.goolgardi NRRL 66250
99
F. palustre NRRL 54056
F. armeniacum NRRL 6227
97
F. venenatum CBS 458.93
F. roseum NRRL 22187
100
F. poae NRRL 13714
F. kyushuense NRRL 25349
F. transvaalense CBS 144211
F. brachygibbosum NRRL 13829
70
F. longipes NRRL 13368
F. lacertarum NRRL 20423
100
F. equiseti NRRL 13405
F. scirpi NRRL 13402
F. nelsonii NRRL 13338
F. tjaynera NRRL 66246
F. aywerte NRRL 25410
98
F. celtidicola MFLUCC 16-0751
F. celtidicola MFLUCC 17–2176
100
98
F. salinense CBS 142420
F. citricola CBS 142421
100
78
83
F. tricinctum NRRL 25481
F. acuminatum NRRL 36147
F. avenaceum NRRL 54939
F. torulosum NRRL 22748
100
F. petersiae CBS 143231
F. flocciferum NRRL 25473
F. nurragi NRRL 36452
F. heterosporum NRRL 20693
F. anguioides NRRL 25385
F. babinda NRRL 25539
0.03
Fig. 116 The best scoring RAxML tree with a final likelihood value
of − 52171.145500 of rpb1 and rpb2 sequence data. The topology
and clade stability of the combined gene analyses was compared to
the single gene analyses. The tree is rooted with Fusarium anguioides
(NRRL 25385) and F. babinda NRRL (25539). The matrix had 1722
distinct alignment patterns with 15.24% undetermined characters
and gaps. Estimated base frequencies were as follows; A = 0.260064,
13
C = 0.246883, G = 0.252626, T = 0.240426; substitution rates
AC = 1.307580, AG = 5.230737, AT = 1.086622, CG = 0.958821,
CT = 10.666584, GT = 1.000000; gamma distribution shape parameter α = 0.761212. Ex-type strains are in bold black and the species
determined in this study is indicated in blue. Bootstrap values (BT)
(over 70% BT) from maximum likelihood are given at the nodes
Fungal Diversity (2020) 102:1–203
et al. 1999; Lombard et al. 2015). The concept of the family
was addressed by Lombard et al. (2015), Maharachchikumbura et al. (2016) and Hyde et al. (2020b) and include 69
genera (Wijayawardene et al. 2020).
Fusarium Link
Fusarium is a large genus in Nectriaceae, with more
than 1500 epithets (Index Fungorum 2020). The genus
was introduced with Fusarium roseum as the type species
(Wijayawardene et al. 2018). Phylogenetic analysis based
on rpb1 and rpb2 nucleotide sequences resulted in a wellsupported phylogenetic clade of the newly isolated strain
clustering with F. celtidicola (Hyde et al. 2014; Shang
et al. 2018). Partial sequences of DNA-directed RNA polymerase II subunit RPB1 (rpb1) and DNA-directed RNA
polymerase II subunit RPB2 (rpb2) gene regions were used
for phylogenetic analysis (Fig. 116) and supported by morphological observations for species level characterization
(Fig. 117).
Fusarium celtidicola Shang, Camporesi & K.D. Hyde, Phytotaxa 361 (3): 255 (2018), new host record
Index Fungorum number: IF 553845; Facesoffungi number: FoF 02453, Fig. 117.
Saprobic on dead stem of Clematis vitalba. Sexual morph: Ascomata 210–255 × 180–280 μm
( x̄ = 230 × 225 μm, n = 10), superficial on host substrate,
single or gregarious, sometimes confluent, globose, black,
ostiolate, papillate. Papilla 75 × 95 μm, conical, with periphysoids, with a small pore. Peridium 25–42 μm wide,
comprising an outer purple layer, composed of 5–7 layers of thin-walled cells of textura angularis, inner layers,
hyaline, thick-walled, several cell layers of textura angularis. Catenophyses numerous, 13–20 μm ( x̄ = 17 μm,
n = 30), septate, narrowing and tapering towards the apex.
Asci 72–100 × 7–14 μm ( x̄ = 70 × 10 μm, n = 20), unitunicate, simple pedicel, broad fusiform to oblong, apex flat,
with J-, cylindrical, conspicuous apical ring. Ascospores
15–30 × 4–9 μm ( x̄ = 17 × 7 μm, n = 50), biseriate, partialoverlapping, hyaline, oval, (1–)3-septate, constricted at the
median septum, slightly constricted at the septa, straight or
slightly curved towards the apex, thin-walled, with a minute
guttule in each cell, smooth-walled. Asexual morph: See
Shang et al. (2018).
Culture characters: Colonies on MEA reaching 30 mm
diam. after 4 weeks at 25 °C. Culture from above, purple
or white radiating outwardly to the edge, undulate margin,
medium dense, flat, dull, fimbriate, and slightly covered with
white aerial mycelium; reverse brown with radiating cream
mycelium.
183
Material examined: Italy, Forlì-Cesena, Casone—Dovadola, on dead aerial stems of Clematis vitalba, 9 February
2013, E. Camporesi, IT1061 (MFLU 17–1537), living culture, MFLUCC 17–2176.
Hosts: Celtis australis, Clematis vitalba—(Shang et al.
2018; this study).
Distribution: Italy—(Shang et al. 2018; this study).
GenBank accession numbers: LSU: MT214620; SSU:
MT226731; ITS: MT310664; tef1: MT394675; rpb2:
MT394723.
Notes: Based on the phylogenetic analysis our strain
MFLUCC 17–2176 clustered with the ex-type strain of
Fusarium celtidicola (MFLUCC 16–0526), which was
recorded from Italy on Celtis australis (Shang et al. 2018).
Our collection is morphologically identical to F. celtidicola
(MFLUCC 16–0526) (Fig. 117).
Sarocladiaceae Lombard
Crous et al. (2018) introduced Sarocladiaceae to accommodate Parasarocladium and Sarocladium. Sarocladiaceae
is characterized by its elongated phialides rising solitary on
vegetative hyphae or on conidiophores that are sparsely or
repeatedly branched, with elongated conidia (Giraldo et al.
2015). Phylogenetic analysis of the combined dataset of the
LSU, ITS, and act sequences reveals a novel Sarocladium
species (Fig. 118).
Sarocladium Gams & D. Hawksw.
Sarocladium species have been reported as human pathogens, plant pathogens and saprobes (Gams 1975; Yeh and
Kirschner 2014; Giraldo et al. 2015). The genus is typified
by S. oryzae, a plant pathogen causing sheath-rot disease of
rice (Ayyadurai et al. 2005; Seifert et al. 2011). Currently, 21
epithets are listed in Index Fungorum (2020). Sarocladium
can be distinguished by its elongate phialides rising solitary
on vegetative hyphae or on conidiophores that are sparsely
or repeatedly branched, and cylindrical conidia. The characters of Sarocladium are mostly reported from culture, while
our collection is associated with a dried stem of Clematis
from Belgium (Fig. 119). The combined dataset of the LSU,
ITS, and act sequences based on a multi-locus phylogenetic
analysis revealed Sarocladium clematidis as a novel species
(Figs. 118, 119).
Sarocladium clematidis Phukhams., Ertz, Gerstmans &
K.D. Hyde, sp. nov.
Index Fungorum number: IF556744; Facesoffungi number: FoF 06267, Fig. 119.
Etymology: Named after the host genus, Clematis.
Holotype: MFLU 17–1507
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184
13
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Fungal Diversity (2020) 102:1–203
◂Fig. 117 Fusarium celtidicola (MFLU 17–1461). a Appearance of
ascomata on Clematis vitalba. b Close up of ascomata on host substrate. c Vertical section through an ascoma. d, e Ostiolar canal.
f Peridium. g Catenophyses. h, i Asci. j–n Ascospores. o Culture
characteristics on MEA. Scale bars: a = 500 µm, b = 200 µm, c, f,
g = 100 µm, d, h–i = 50 µm, e = 20 µm, j–n = 10 µm
Saprobic on dried stems of Clematis patens. Sexual
morph: Undetermined. Asexual morph: Colonies effuse
on the natural substrate, scattered, hairy, dark brown.
Mycelium partly immersed, branched, composed of pale
brown, septate hyphae. Conidiophores 85–320 × 12–19 μm
( x̄ = 150 × 15 μm, n = 20), macronematous, synnematous,
composed of 7–12 hyphae in each stipe, tree-like, parallel
and unbranched in the stipe, gregarious, or scattered, erect,
stripes straight or slightly flexuous, 6–17-septate, constricted
at septa, 15–30 μm wide at base, tapering towards apex,
6–14 μm wide at apex, irregularly branched, cylindrical,
smooth, dark brown. Conidiogenous cells 7–15 × 2–5 μm
( x̄ = 10 × 2.5 μm, n = 20), polyphialidic, integrated or terminal, cylindrical, straight to slightly curved, slightly narrowing at apex, acropetally proliferating, hyaline to pale brown,
verrucose. Conidia 4–9 × 2–4 μm ( x̄ = 6 × 2.5 μm, n = 40),
unicellular, fusiform to broadly fusiform, aseptate, thickwalled, hyaline, verrucose, bud scars or disjunctors present
at the site of attachment, 1–2-guttulate.
Culture characters: Colonies on PDA reaching 30 mm
diam. after 4 weeks at 25 °C. Cultures from above, cream or
white, medium dense, circular, umbonate, flat, at first glabrous becoming powdery at centre, wrinkled, folded, slimy
in the middle, slightly radiating outwardly; reverse: cream
at the centre, mycelium radiating outwardly. Asexual morph
formed in culture with morphology similar to that on the
natural substrates.
Material examined: Belgium, Flemish Brabant, Meise
Botanic Garden, Bouchout Domain, on dead stems of
Clematis patens, 13 June 2017, D. Ertz & C. Gerstmans,
BRCP3 (MFLU 17–1507, holotype); ex-type living culture,
MFLUCC 17–2150.
Host: Clematis patens—(This study).
Distribution: Belgium—(This study).
GenBank accession numbers: LSU: MN629285; SSU:
MN629284; ITS: MN629287; tef1: MN628625; rpb2:
MN628627.
Notes: Sarocladium clematidis formed synnemata with
7–12 hypha in each stipe on the natural substrate (Fig. 119).
Sarocladium clematidis (MFLUCC 17–2150) is morphologically similar to Phaeoisaria clematidis, however they
are phylogenetically distant (Yeh and Kirschner 2014).
Morphological characters of Sarocladium have mainly been
reported from culture (Seifert et al. 2011). Sarocladium
clematidis has solitary, elongate phialides, with branched
185
conidiophores and fusiform conidia (Seifert et al. 2011). In
phylogenetic analysis of combined sequence data (Fig. 118),
S. clematidis formed a close relationship with S. dejongiae
(CBS 144929) with strong support (100% ML/1.00 BYPP).
In a BLASTn search of GenBank, the closest match of the
ITS sequence of S. clematidis is S. dejongiae with 98% similarity to the S. dejongiae (511/519 with five gaps). Based
on its distinct characters and the phylogenetic support, we
introduce a novel species herein.
Stachybotryaceae Lombard & P. Crous
Stachybotryaceae was introduced by Crous et al. (2014a)
and comprises 36 genera (Wijayawardene et al. 2018).
Species are saprobes, plant and animal pathogens, or airborne (Crous et al. 2014a; Lombard et al. 2016; Rennberger
2018). Stachybotryaceae members have sporodochial to
synnematous conidiomata, phialidic conidiogenous cells,
and 0–1-septate conidia (Lombard et al. 2016). In this study,
phylogenetic analyses of the combined LSU, ITS, tef1, rpb2
and tub dataset reveals a new record of Memnoniella from
Clematis collected from Thailand (Fig. 120).
Memnoniella Höhn.
Memnoniella was synonymised under Stachybotrys (Galloway 1933). However, the distinctiveness of Memnoniella
was confirmed by morphological characters and phylogenetic evidence (Lombard et al. 2016). Memnoniella is distinguished by mostly smooth, thick-walled and unbranched
conidiophores giving rise to conidia in dry chains, whereas
Stachybotrys has hyaline phialides, brightly coloured, or
dark phialophores, producing 1-celled, dark conidia, accumulating in a slimy cluster (Wang et al. 2015). Some species
of Memnoniella (e.g., M. echinata and M. longistipitata)
produce dimorphic conidia (Li et al. 2003). We report a new
collection of M. oblongispora from a Clematis species from
Thailand which produces dimorphic conidia, a feature not
known in this species (Fig. 121).
Memnoniella oblongispora Lin, McKenzie, Wang & K.D.
Hyde, new host record and dimorphic characters report
Index Fungorum number: IF 552085; Facesoffungi number: FoF 02081, Fig. 121.
Saprobic on dried stems of Clematis subumbellata.
Sexual morph: Undetermined. Asexual morph: Hyphomycetous. 1) Colonies of dry conidial chains, effuse on the
natural substrate, scattered, hairy, dark brown. Mycelium
semi-immersed to superficial, composed of pale brown,
septate hyphae. Conidiophores 30–70(–120) × 3–6 μm
( x̄ = 50 × 5 μm, n = 20), macronematous, mononematous,
erect, simple, stipes straight or flexuous, mostly unbranched,
but irregularly branched at the upper parts,, sub-cylindrical to cylindrical, bearing a crown of phialides at apex,
mammiform at the base cell, 3–4-septate, brown at the
13
186
Sarocladium terricola CBS 243.59 T
51/0.97
Sarocladium bacillisporum CBS 425.67 T
100/1.00
Sarocladium subulatum MUCL 9939 T
60/1.00
Sarocladium glaucum CBS 796.69 T
100/1.00
81/1.00 71/0.99
Sarocladium bifurcatum UTHSC 05-3311
Sarocladium gamsii CBS 707.73 T
Sarocladium implicatum CBS 959.72 T
100/1.00
62/1.00
Sarocladium ochraceum CBS 428.67 T
Sarocladium clematidis MFLUCC 17–2150 T
100/1.00
Sarocladium dejongiae CBS 144929 T
61/--
Sarocladium summerbellii CBS 430.70 T
Sarocladium strictum CBS 346.70 T
99/1.00
Sarocladium bactrocephalum CBS 749.69 T
97/1.00
Sarocladium pseudostrictum UTHSC 02-1892
--
Sarocladiaceae
Fig. 118 Best scoring RAxML
tree with a final likelihood
value of − 9149.858797 of
LSU, ITS, and act sequence
data. The tree is rooted with
Kiflimonium curvulum (CBS
430.66). The matrix had 613
distinct alignment patterns
with 18.90% undetermined
characters and gaps. Estimated base frequencies were
as follows; A = 0.223643,
C = 0.286812, G = 0.271932,
T = 0.217614; substitution rates
AC = 1.580411, AG = 2.743783,
AT = 2.611458, CG = 0.755231,
CT = 8.336114, GT = 1.000000;
gamma distribution shape
parameter α = 0.532334. Extype strains are in bold black
and the species determined
in this study is indicated in
blue. Bootstrap values (BS)
greater than 50% BS (maximum
likelihood (left)) and Bayesian posterior probabilities (PP,
right) greater than 0.90 are
given at the nodes. Hyphens (-)
represent support values less
than 50% BS/0.90 BYPP. Thick
branches represent significant
support values from all analyses
(BS ≥ 70%/BYPP ≥ 0.95) at the
genus and family level
Fungal Diversity (2020) 102:1–203
Sarocladium brachiariae XBL 2015
--
80/1.00
Sarocladium oryzae CBS 180.74 T
Sarocladium spinificis Z0106
--/0.95
Sarocladium zeae CBS 801.69 T
50/--
Sarocladium kiliense CBS 122.29 T
--
Sarocladium hominis UTHSC 04-1034
Kiflimonium curvulum CBS 430.66
0.05
apex, hyaline at the base, smooth or verruculose. Conidiogenous cells 5–10 × 2.5–3.5 μm ( x̄ = 8 × 3 μm, n = 10),
monophialidic, discrete, determinate, terminal, clustered
at the apex of conidiophores, oblong, pale brown. Conidia
3–5 × 2.5–4.5 μm ( x̄ = 4 × 4 μm, n = 20), acrogenous, aseptate, globose, olivaceous, brown to dark brown, verrucose,
thick-walled, dark brown to reddish brown, verrucose,
formed in long chains, easily separated. 2) Colonies of
slimy conidial chains produced in culture on MEA media.
Conidiophores 32–125 × 4–7 μm ( x̄ = 70 × 4.5 μm, n = 10),
macronematous, mononematous, erect, simple, straight or
flexuous, unbranched, smooth, thick-walled, septate, subcylindrical, of elongated phialide base, apex bearing a crown
of phialidic conidiogenous cells, hyaline at the base, olive
grey at the apex. Conidiogenous cells 8–17 × 4.8–5.6 μm
( x̄ = 10 × 5 μm, n = 20), monophialidic, discrete, determinate,
terminal, clustered at the apex of conidiophores, pyriform,
obovate, ellipsoidal, clavate or reniform, smooth, brown.
Conidia 8–14 × 4.5–7 μm ( x̄ = 9 × 5 μm, n = 50), aggregated
in large, slimy, black and glistening heads, acrogenous,
simple, spherical, oblong, immature hyaline, dark brown at
13
maturity, two guttules present when immature, unicellular,
verrucose surface.
Culture characters: Colonies on MEA reaching 30 mm
diam. after 2 weeks at 25 °C. Cultures from above, cream
to yellowish white, medium to sparse, circular, fimbriate,
wrinkled and folded, slightly radiating outwardly; reverse
orange at the centre, cream radiating, dimorphic characters
produced after 3 weeks of incubation, characters of dry
conidial chains similar to those on natural substrates.
Material examined: Thailand, Phayao Province, Phu Sang
District, dead stems of Clematis subumbellata, 20 March
2017, C. Phukhamsakda, CMTH06 (MFLU 17–1470); living
culture, MFLUCC 17–2064.
Hosts: Clematis subumbellata, decaying Quercus leaf—
(Lin et al. 2016; this study)
Distribution: Thailand—(Lin et al. 2016; this study)
GenBank accession numbers: LSU: MT214621; SSU:
MT226732; ITS: MT310665; tef1: MT394676; rpb2:
MT394724.
Notes: Memnoniella oblongispora (MFLUCC 17–2064)
formed a clade with the type strain of M. oblongispora
Fungal Diversity (2020) 102:1–203
(MFLUCC 15–1074) with moderate support (75% ML/0.92
BYPP). Memnoniella species may produce dimorphic asexual characters (Li et al. 2003). Memnoniella oblongispora
was described as producing slimy conidial masses in a black
and glistening head, with oblong, verrucose, olive green to
black conidia (Lin et al. 2016). Our collection on a natural
substrate, produced dry chains of sphaerical conidia. After
three weeks in culture, glistening conidial masses were
observed on the surfaces of white mycelia. In culture, both
dry chains and stachybotrys-type slimy conidia were formed,
which correlated with the morphology reported by Li et al.
(2003).
The analysis (Fig. 120) showed that our collection formed
a clade with the ex-type strain of M. oblongispora and a
strain of M. longistipitata (CBS 136197). However, the
characters of M. longistipitata (CBS 136197) have not been
reported (Lombard et al. 2016). Memnoniella oblongispora
(MFLUCC 17–2063) and the type strain of M. longistipitata (ATCC 22699) are distinguishable by conidiophores
characters. Memnoniella longistipitata has very long conidiophores in culture (260–460 × 3.6–4.7 μm; Li et al. 2003),
while Memnoniella oblongispora (MFLUCC 17–2064) has
shorter conidiophores based on examination of their characters on natural substrates and in culture (32–125 × 4–7 μm;
Lin et al. 2016; this study).
Pairwise comparison of the ITS sequence data in Memnoniella oblongispora (strains MFLUCC 17–2064 and
MFLUCC 15–1074) are identical. Therefore, we report a
new record of M. oblongispora from Clematis in Thailand,
with dimorphic conidial (Fig. 121).
Discussion
In this paper we provide data on collections of fungi associated with typically vigorous, woody, often invasive,
climbing vines in Ranunculaceae, Clematis. The genus is
distributed worldwide as a native plant and as a group that
is commercially cultivated (Grey-Wilson 2000). In the present study, nine Clematis species were sampled from Asia
(China, Thailand), Europe (Belgium, Italy, UK). These are
Clematis fulvicoma, C. orientalis, C. patens, C. serratifolia,
C. sikkimensis, C. subumbellata, C. virginiana, C. vitalba,
C. viticella, additionally with C. ligusticifolia. Seventy-three
taxa were added to the known fungi associated with Clematis. The classification of each species is justified by a consolidated species concept approach (Quaedvlieg et al. 2014).
Applying a consolidated species concept
to delineating taxa
The implementation of a comprehensive approach for confidently introducing new fungal taxa has been strongly
187
recommended in several studies. Quaedvlieg et al. (2014)
introduced the “consolidated species concept” that distinguished fungal taxa based on the combination of the “biological species concept”, “ecological species concept”,
“morphological species concept”, “phylogenetic species
concept” and the application of “genealogical concordance phylogenetic species recognition” (Taylor et al. 2000).
The approach was supplemented by performing a pairwise
homoplasy index, followed by the interpretation of nucleotide difference based on its importance as highlighted by
Jeewon and Hyde (2016). Phylogenetic species concept
is currently regarded as an effective molecular tool to differentiate a common ancestor in the fungal community.
Molecular-clock age estimates have been used as additional
evidence to explore the evolution of lineages and divergence
dates in fungal taxa (Hongsanan et al. 2017; Liu et al. 2017;
Phukhamsakda et al. 2017b; Phillips et al. 2019; Samarakoon et al. 2019; Hyde et al. 2020b). In some complex species with high similarity in the phenotype and the evolution of these sequences cannot be described by a single tree
and therefore the pairwise homoplasy index was applied
to distinguish the genotype group and justify the species
boundaries of phylogenetically closely related taxa (Dettman et al. 2003; Quaedvlieg et al. 2014). This method has
been widely used to determine the possibility of inbreeding
within the population and establish the genetic differentiation in lineages from phylogenetically closely related species
(Laurence et al. 2014). Although the analysis was recommended for sexual organisms, recombination sciences have
also applied this method for historical recombination (Burnett 2003; Turner et al. 2013). Diaporthaceae (Diaporthales),
Hermatomycetaceae, and Lophiostomataceae (Pleosporales)
are used as a case-study in the present work.
Diaporthaceae (Diaporthales): Diaporthe is a complex
genus that presents high genetic variation within one species and can occur on a wide range of hosts. To introduce
a novel species within Diaporthe, consolidated evidence is
needed. We apply genealogical concordance phylogenetic
species recognition to study the nucleotide distances of the
common ancestors within a targeted clade (Fig. 100). The
genealogical concordance phylogenetic species recognition is relatively reliable and can be useful in the identification of species in Diaporthe. Diaporthe rudis showed
the genetic complexity especially in the ITS region and the
recombination parameters among selected taxa are shown
in Fig. 103b. The strains probably contain high intraspecific variation in the pseudogene of the rDNA copy (Stadler
et al. 2020). The GCPSR result of the concatenated gene
set shows insignificant recombination between the isolates
(Φw = 0.05, Fig. 103b). We demonstrate that some D. rudis
isolates have undergone recombination of species based on
the evolutionary dynamics (Quaedvlieg et al. 2014; Guarnaccia and Crous 2017).
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Fungal Diversity (2020) 102:1–203
189
◂Fig. 119 Sarocladium clematidis (MFLU 17–1507, holotype). a
Appearance of synnemata on Clematis patens. b, c Close up of conidiophores. d–g Synnemata. h, i Conidiogenous cells and conidia. j
Conidia. k Germinated conidia. l, m Culture characteristics on PDA.
n–r Asexual morph produced in culture. Scale bars: b = 500 μm, c,
n = 50 μm, d = 200 μm, e–g = 100 μm, h–j, q, r = 5 μm, o = 20, p = 10
Hermatomycetaceae (Pleosporales): This family lacks a
sexual morph and the genealogical concordance phylogenetic species recognition result of Hermatomyces sphaericus
(Fig. 21) contradicts the phylogeny (especially H. biconisporus and H. pandanicola). Phylogenies from maximum
likelihood, maximum parsimony and Bayesian statistics
did not give good support for the separate clade between
H. sphaericus sensu stricto and H. pandanicola. This result
is supported by the genealogical concordance phylogenetic
species recognition value that does not significantly support
Memnoniella echinata CBS 216.32 T
Memnoniella echinata DAOM 235365
99/1.00 Memnoniella echinata CBS 343.50
Memnoniella echinata CBS 627.66
100/1.00 Memnoniella echinata DAOM 173162
Memnoniella echinata CBS 344.39
Memnoniella echinata CBS 406.80
51/-Memnoniella
echinata CBS 304.54
100/1.00
Memnoniella
oblongispora MFLUCC 15-1074 T
75/0.92
69/-- Memnoniella oblongispora MFLUCC 17-2064
78/-Memnoniella longistipitata CBS 136197
100/1.00
Memnoniella longistipitata ATCC 22699 T
100/1.00
Memnoniella chromolaenae MFLUCC 17-1507 T
Memnoniella dichroa CBS 123800
100/1.00
Memnoniella dichroa CBS 526.50
-Memnoniella dichroa ATCC 18913 T
92/1.00
Memnoniella oenanthes CBS 388.73
100/1.00
Memnoniella oenanthes ATCC 22844 T
Memnoniella pseudonilagirica CBS 136405 T
100/1.00
Memnoniella nilagirica MFLUCC 15-0660 T
-Memnoniella humicola CBS 463.74 T
Memnoniella ellipsoidea CBS 136199
100/1.00
Memnoniella ellipsoidea CBS 136200
100/1.00
Memnoniella ellipsoidea CBS 136201 T
Memnoniella ellipsoidea CBS 136202
67/1.00
Memnoniella
putrefolia CBS 101177 T
98/1.00
100/1.00
Memnoniella putrefolia CBS 136171
Memnoniella brunneoconidiophora CBS 109477
100/1.00 Memnoniella brunneoconidiophora CBS 136191 T
Peethambara sundara CBS 646.77
1.00
Memnoniella
55/--
0.03
Fig. 120 Bayesian 50% majority-rule consensus phylogram based
on combined LSU, ITS, tef1, rpb2 and tub sequence data for Memnoniella species. The topology and clade stability of the combined
gene analyses was compared to the single gene analyses. The tree is
rooted with Peethambara sundara strain CBS 646.77. Thirty strains
were included in the combined genes sequence analyses which
comprised 3057 characters (798 for LSU, 618 for ITS, 554 for tef1,
721 for rpb2, 366 for tub, including gap regions). The tree from the
maximum likelihood analysis had similar topology to the Bayesian
analyses. The best scoring RAxML tree had a final likelihood value
of − 11438.749144. The matrix had 794 distinct alignment patterns, with 19.09% of undetermined characters and gaps. Estimated
base frequencies were: A = 0.236362, C = 0.265886, G = 0.272223,
T = 0.225529; substitution rates AC = 1.092531, AG = 3.397943,
AT = 1.269553, CG = 0.955550, CT = 9.625921, GT = 1.000000;
gamma distribution shape parameter α = 0.547123. In our analysis,
GTR + I + G model was used for each partition in Bayesian posterior
analysis. The species determined in this study is indicated in blue.
Bootstrap values (BS) greater than 50% BS (maximum likelihood
(left)) and Bayesian posterior probabilities (PP, right) greater than
0.90 are given at the nodes. Hyphens (-) represent support values less
than 50% BS/0.90 BYPP. Thick branches represent significant support values from all analyses (BS ≥ 70%/BYPP ≥ 0.95) in genus level
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Fungal Diversity (2020) 102:1–203
◂Fig. 121 Memnoniella oblongispora (MFLU 17–1470). a, b Appear-
ance of dry chains of conidiophores on Clematis subumbellata. c
Conidiophores. d–h Conidiogenous cells and conidia. i Conidia. j
Culture characteristics on MEA. k Slimy chains on MEA. l–m Dry
chains and slimy chains of M. oblongispora (MFLUCC 17–2064)
developed in the same culture on MEA media (red arrow indicates
slimy chain; blue arrow indicates dry chains of conidia). n–p Dry
chain of conidiophores, conidiogenous cells and conidia in culture.
q–u Slimy chain with conidiophores, conidiogenous cells and conidia
in culture. Scale bars: b, l–m = 200 μm, c, n, q–s = 20 μm, d–e, o–p,
t–u = 10 μm, f–i = 5 μm
its speciation. Hermatomyces sphaericus occurs in countries that lie close to the equator, such as Cuba, Panama and
Thailand where the climate is mostly hot with tropical conditions. These isolates were reported as saprobes on dicotyledonous and monocotyledonous hosts. The genus lacks a
known sexual morph, thus the establishment of sexual compatibility in mating-type genes such as MAT-1 and MAT-2
genes are recommended for the determination of reproductive differentiation into a sexual morph of Hermatomyces
(Giraud et al. 2008; Mageswari et al. 2016).
Lophiostomataceae (Pleosporales): To support the recognition of the morphological species concept and phylogenetic species concept with numerous taxa being discovered
within Lophiostomataceae, GCPSR was applied to delimit
species within Lophiostomataceae using data of five gene
loci (Fig. 28). There are no conflicting results in this family and we provide consolidated evidence by phylogenetic
relatedness in a multi-locus dataset and the interpretation
of nucleotide differences (Turner et al. 2013; Quaedvlieg
et al. 2014; Jeewon and Hyde 2016). We also compared our
new isolates with documented fungal taxa recorded in the
database and discuss the ecological species concept.
Host–fungal interactions
It is hypothesized that modifications of the divergent plate
boundaries 50 million years ago (Mya) resulting in speciation and adaptation of land plants and living organisms
(Navaud et al. 2018). These caused dramatic change in
continents, temperatures, peregrine, and communities of
living organism and may have impacted the distribution
of plant symbioses (Cawood et al. 2018). Most plants are
inhabited by fungal symbionts as endophytes, mycorrhizae and saprobes. The arbuscular mycorrhizal fungi were
reported to colonize the early diverging vascular plant lineages since 500 Mya (Field et al. 2012). Host–fungal interactions occurred as coevolution events with leading changes
in genetic succession over generations (Naranjo-Ortiz and
Gabaldón 2019). The cospeciation of the fungus with the
original host consistently leads to a decrease in fitness on
the original host and increased distribution of fungi species worldwide. The induction of foreign plants by global
191
weather changes immigration of animal and anthropogenic
movement resulted in the introduction of fungal strains
resident in plants to adapt and expand the range of hosts
(Antonovics et al. 2002). Host jumps commonly occur in
fungal species with a higher rate than the host-fungus cospeciation or coevolution (Roy 2001). This event most commonly occurs in plant pathogens that occasionally change to
another lifestyle (Sillo et al. 2015). Among many examples
of host-fungal interactions, we demonstrate some observed
in this study.
Advances in the understanding of host–pathogen specificity and co-evolutionary interactions with specific plant
genera or families has been demonstrated with Didymellaceae (Aveskamp et al. 2009, 2010; Chen et al. 2015, 2017).
Cospeciation is expected to have congruent phylogenies in
the host and fungal species with similar divergence times
(De Vienne et al. 2013). Species of Didymellaceae associated with Clematis are Calophoma clematidis-rectae
(≡ Phoma clematidis-rectae), C. clematidina (≡ P. clematidina), and P. herbarum (Foister 1961; Tai 1979; Woudenberg
et al. 2009; Aveskamp et al. 2010). Soleimani et al. (2018)
estimated the crown age of genera within Didymellaceae by
using molecular dating. The molecular evolution revealed
that the species of Didymellaceae diverged in the Miocene
period (35.7 Mya). These results are in congruence with
the relative radiation of the crown ages of Clematis species
that had also a diversification event in the Miocene (Xie
et al. 2011). However, the hypotheses of coevolution and
host specificity in genera of Didymellaceae needs further
addressing.
The introduction of a foreign host lends a significant accidental movement of fungi into new habitats and undergo host
jumps (Herrera et al. 2016). The saprobic fungus Nigrograna
chromolaenae was found on Chromolaena odorata, an invasive weed in Thailand, and was also associated with Clematis fulvicoma in this country (Mapook et al. 2020). Thus,
the species is associated with dicotyledons plant in tropical
climates. Dictyocheirospora xishuangbannaensis which was
reported from Pandanaceae (monocots) from Yunnan Province in China was also observed from Clematis sikkimensis
from Thailand. It can be hypothesized that D. xishuangbannaensis generally occur in tropical climates. Stemphylium
vesicarium which causes leaf blight and leaf spots on several
agricultural plants worldwide was associated with Clematis
vitalba in Italy as a saprobe. Based on current trends, many
fungal species in this study also have relatively broad host
range with the consideration of co-factors such as environments and ecological species concept. An increasing number
of fungi are likely to have an ability of switch host and then
speciation might occur with this host plant.
Over 560 fungal species are listed from Clematis
sp. worldwide (Farr and Rossman 2020). To assess the
fungal biodiversity, each individual group, host, and
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192
specific environmental factor could represent an alternative
approaches to study fungal biodiversity (Mueller and Schmit
2007; Hyde et al. 2018b). Blackwell (2011) estimated the
number of fungi to plants by more than 6 to 1. Therefore,
there are many potential fungal habitats that need to be further studied and modern molecular methods could support
the discovery of species (Hawksworth 1991, 2001).
Acknowledgements Chayanard Phukhamsakda would like to thank the
Royal Golden Jubilee PhD Program under Thailand Research Fund
(RGJ) and the German Academic Exchange Service (DAAD) for a
joint TRF-DAAD (PPP 2017–2018) academic exchange Grant to K.D.
Hyde and M. Stadler and the RGJ for a personal grant to C. Phukhamsakda (The scholarship no. PHD/0020/2557 to study towards a PhD).
Dr. Shaun Pennycook is thanked for checking and suggesting Latin
names of the new taxa. Dr. Olivier Raspé, Dr. Rajesh Jeewon, Sirinapa
Konta, Milan Samarakoon, Indunil C. Senanayake, Dr. Sinang Hongsanan, Chuan-Gen Lin, Qiu-Ju Shang and Pranami D. Abeywickrama
are thanked for their valuable suggestions on the phylogenetic analysis
and for sequencing. Mr. Martin van de Bult, Cyrille Gerstmans, Prof.
Hong-Yan Su, Tian Qing, Dr. Zong-Long Luo are gratefully acknowledged for sample collection guidance in China and Thailand. Kevin D.
Hyde thanks the future of specialist fungi in a changing climate: baseline data for generalist and specialist fungi associated with ants, Rhododendron species and Dracaena species (Grant No: DBG6080013),
Impact of climate change on fungal diversity and biogeography in
the Greater Mekong Subregion (Grant No: RDG6130001). Alan J.
L. Phillips acknowledges the support from UID/MULTI/04046/2019
Research Unit grant from FCT, Portugal to BioISI. E. B. Gareth Jones
is supported under the Distinguished Scientist Fellowship Program
(DSFP), King Saud University, Kingdom of Saudi Arabia. D. N.
Wanasinghe would like to thank the CAS President’s International Fellowship Initiative (PIFI) for funding his postdoctoral research (Number 2019PC0008), the National Science Foundation of China and the
Chinese Academy of Sciences for financial support under the following Grants: 41761144055, 41771063 and Y4ZK111B01. Wanasinghe
also thanks the 64th batch of China Postdoctoral Science Foundation
(Grant No.: Y913083271). Saowaluck Tibpromma would like to thanks
the International Postdoctoral Exchange Fellowship Program (Number Y9180822S1), CAS President’s International Fellowship Initiative
(PIFI) (Number 2020PC0009), China Postdoctoral Science Foundation
and the Yunnan Human Resources, and Social Security Department
Foundation for funding her postdoctoral research. Mingkwan Doilom
would like to thank the 5th batch of Postdoctoral Orientation Training Personnel in Yunnan Province (Grant No.: Y934283261) and the
64th batch of China Postdoctoral Science Foundation (Grant No.:
Y913082271).
Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long
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copy of this licence, visit http://creativecommons.org/licenses/by/4.0/.
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Affiliations
Chayanard Phukhamsakda1,2,3 · Eric H. C. McKenzie4 · Alan J. L. Phillips5 · E. B. Gareth Jones9,10 · D. Jayarama Bhat14 ·
Stadler Marc16 · Chitrabhanu S. Bhunjun3 · Dhanushka N. Wanasinghe2,3,8,15 · Benjarong Thongbai16 ·
Erio Camporesi11,12,13 · Damien Ertz17,20 · Ruvishika S. Jayawardena3 · Rekhani Hansika Perera3,18 ·
Anusha H. Ekanayake3 · Saowaluck Tibpromma2,8,19 · Mingkwan Doilom2,8,19 · Jianchu Xu2,19,8 · Kevin D. Hyde1,2,3,6,7,8
* Kevin D. Hyde
kdhyde3@gmail.com
1
2
Institute of Plant Health, Zhongkai University of Agriculture
and Engineering, Haizhu District, Guangzhou 510225,
People’s Republic of China
Key Laboratory for Plant Diversity and Biogeography
of East Asia, Kunming Institute of Botany, Chinese
Academy of Sciences, Kunming 650201, Yunnan,
People’s Republic of China
3
Center of Excellence in Fungal Research, Mae Fah Luang
University, Chiang Rai 57100, Thailand
4
Landcare Research Manaaki Whenua, Private Bag 92170,
Auckland, New Zealand
5
University of Lisbon, Faculty of Sciences, Biosystems
and Integrative Sciences Institute (BioISI), Campo Grande,
1749-016 Lisbon, Portugal
6
Mushroom Research Foundation, 128 M.3 Ban Pa Deng
T. Pa Pae, A. Mae Taeng, Chiang Mai 50150, Thailand
7
Department of Biology, Faculty of Science, Chiang Mai
University, Chiang Mai 50200, Thailand
8
State Key Laboratory for Conservation and Utilization
of East and Central Asia Regional Office, World
Agroforestry Centre (ICRAF), Kunming 650201, Yunnan,
People’s Republic of China
9
33 B St. Edwards Road Southsea Hants,
Hampshire PO5 3DH, UK
10
Department of Botany and Microbiology, College of Science,
King Saud University, P.O Box 2455, Riyadh 11451,
Kingdom of Saudi Arabia
11
A.M.B. Gruppo, Micologico Forlivese “Antonio Cicognani”,
Viale Roma 18, 47121 Forli, Italy
12
A.M.B, Circolo Micologico “Giovanni Carini”, C.P. 101,
25121 Brescia, Italy
13
Societá per gli Studi Naturalistici della Romagna, C.P. 143,
48012 Bagnacavallo, RA, Italy
14
No. 128/1-J, Azad Housing Society, Curca, P.O. Goa Velha,
403108, India
15
Center for Yunnan Plateau Biological Resources Protection
and Utilization, College of Biological Resource and Food
Engineering, Qujing Normal University, Qujing,
Yunnan 655011, People’s Republic of China
16
Department of Microbial Drugs, Helmholtz Centre
for Infection Research and German Centre for Infection
Research (DZIF), partner site Hannover/Braunschweig,
Inhoffenstrasse 7, 38124 Brunswick, Germany
17
Department of Research, Meise Botanic Garden, 1860 Meise,
Belgium
18
Guizhou Key Laboratory of Agricultural Biotechnology,
Guizhou Academy of Agricultural Sciences, Guiyang,
Guizhou Province 550006, People’s Republic of China
19
Centre for Mountain Futures (CMF), Kunming
Institute of Botany, Kunming 650201, Yunnan,
People’s Republic of China
20
Fédération Wallonie-Bruxelles, Service Général de
l’Enseignement Supérieur et de la Recherche Scientifique,
Rue A. Lavallée 1, 1080 Brussels, Belgium
13