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Phytotaxa 422 (2): 144–156
https://www.mapress.com/j/pt/
Copyright © 2019 Magnolia Press Article PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
144 Accepted by Sajeewa Maharachchikumbura: 6 Oct. 2019; published: 23 Oct. 2019
https://doi.org/10.11646/phytotaxa.422.2.2
Sporidesmium guizhouense sp. nov. (Sordariomycetes incertae sedis), a new species
from a freshwater habitat in Guizhou Province, China
LING-LING LIU1,2,3, JING YANG3,4, NING-GUO LIU3,5, YA-YA CHEN1,3,6, XIAO-XIA GUI1,3 & ZUO-YI LIU3*
1Guizhou University, Guiyang 5500025, Guizhou, China
2Guizhou Institute of Soil and Fertilizer, Guizhou Academy of Agricultural Sciences, Guiyang 550006, Guizhou, China
3Guizhou Key Laboratory of Agricultural Biotechnology, Guizhou Academy of Agricultural Sciences, Guiyang 550006, Guizhou, China
4Center of Excellence in Fungal Research, Mae Fah Luang University, Chiang Rai 57100, Thailand
5Faculty of Agriculture, Natural Resources and Environment, Naresuan University, Phitsanulok 65000, Thailand
6Guizhou Institute of Crop Germplasm Resources, Guizhou Academy of Agricultural Sciences, Guiyang 550006, Guizhou, China
*Corresponding author: Zuo-Yi Liu, email: gzliuzuoyi@163.com
Abstract
During a survey of freshwater fungi in Guizhou Province, China, a collection from a submerged decaying twig in Baihua
Lake was identified as a new species of Sporidesmium sensu lato based on morphological characters and phylogenetic
analyses of combined LSU, SSU, ITS, TEF1α and RPB2 sequence data. Phylogenetic analyses supported its placement in
Sordariomycetes but the fungus grouped distant from Sporidesmium sensu stricto, and its ordinal or familial position within
the class remained inconclusive. Sporidesmium guizhouense sp. nov. is introduced with an illustrated account and notes on
its taxonomy, phylogeny and systematic position are provided.
Keywords: asexual fungi, phylogeny, Sordariomycetes, taxonomy
Introduction
Freshwater fungi depend on submersion in water for the whole or part of their life cycle playing an important role
in freshwater ecosystems (Wong et al. 1998). This group of fungi has generally been divided into the leaf-inhabiting
freshwater fungi or Ingoldian, the lignicolous freshwater fungi or wood decomposing and the lower fungi (Hyde et
al. 2016b). The leaf-inhabiting fungi were reviewed by Graça et al. (2015), while lignicolous freshwater fungi along
a north–south gradient in the Asian/Australian region are currently being studied (Hyde et al. 2016b). Sporidesmium-
like taxa are common on woody substrates in freshwater habitats (Cai et al. 2002) and have been frequently reported
(Hyde & Goh 1998, Ho et al. 2001, 2002, Cai et al. 2003, Luo et al. 2004, Hyde et al. 2016a, Su et al. 2016, Luo et
al. 2018, Yang et al. 2018).
Sporidesmium Link is a large and complex genus (Subramanian 1992, Wu & Zhuang 2005, Shenoy et al. 2006).
Its asexual morphs are characterized by terminal, monoblastic, determinate or proliferating conidiogenous cells,
macronematous, unbranched conidiophores, and euseptate or distoseptate conidia in variable shapes (cylindrical,
obclavate, subfusiform, lageniform, obpyriform or obturbinate) with a mucilaginous sheath sometimes. (Link
1809, Ellis 1958, 1971, Zhou et al. 2001, Shenoy et al. 2006, Dubey & Pandey 2011, Seifert et al. 2011, Ma et
al. 2012, Dubey & Sengupta 2015, Su et al. 2016, Yang et al. 2018). Zhang et al. (2017) reported a sexual morph
in Sporidesmium sensu stricto based on molecular study. Presently, 484 epithets are referred to the genus in Index
Fungorum (2019). However, many species have been revised and transferred to more than 30 genera (Hughes 1979,
Kirk 1982, Hernández-Gutiérrez & Sutton 1997, Shoemaker & Hambleton 2001, Iturriaga et al. 2008, Delgado et
al. 2018). Sporidesmium was shown to be polyphyletic with species clustered in different families or orders within
Dothideomycetes and Sordariomycetes based on phylogenetic analyses (Shenoy et al. 2006). Within Sordariomycetes,
Su et al. (2016) designated a monoclade as Sporidesmiaceae sensu stricto with species morphologically similar to the
type species of Sporidesmium; Distoseptisporaceae was introduced to accommodate species resembling Ellisembia (Su
et al. 2016); Crous et al. (2017) established Pseudosporidesmiaceae which has percurrently rejuvenating conidiophores
and obclavate, pale brown conidia. Morphological characters do not work well for the generic delimitations of most
SPORIDESMIUM GUIZHOUENSE SP. NOV. Phytotaxa 422 (2) © 2019 Magnolia Press • 145
Sporidesmium-like taxa and therefore phylogenetic-based studies are required for the identification of most species in
Sporidesmium sensu lato (Shenoy et al. 2010, Su et al. 2016).
There have been several studies on the lignicolous freshwater fungi from China (Li et al. 2017, Bao et al. 2018,
Luo et al. 2018) and those in habiting freshwater lakes (Cai et al. 2002, Luo et al. 2004). During a survey of freshwater
fungi in Guizhou Province, China, a novel species of Sporidesmium was obtained from a freshwater lake. Based on
distinct morphological characters and phylogenetic analyses, Sporidesmium guizhouense sp. nov. is introduced with an
illustrated account and molecular support.
Materials & Methods
Collection and examination of specimens
Specimens of submerged decaying wood were collected from Baihua Lake, Guiyang, Guizhou Province, China, in
April 2018. Methods of morphological observation followed Luo et al. (2018). Measurements were performed using
the Tarosoft (R) Image Frame Work program and images used for figures were processed with Adobe Photoshop CS6
software. Single spore isolation was carried out following the process: spore suspension from the specimen was spread
onto water agar (WA), and then the germinating spores were transferred onto potato dextrose agar (PDA) and incubated
at 25°C following the method in Luo et al. (2018). Specimen were deposited in the Herbarium of Guizhou Academy
of Agriculture Sciences (GZAAS), Guiyang, Guizhou Province, China and the Herbarium of Cryptogams, Kunming
Institute of Botany, Academia Sinica (HKAS), Kunming, Yunnan Province, China. Axenic cultures were deposited in
Guizhou Culture Collection (GZCC), Guiyang, Guizhou Province, China and China General Microbiological Culture
Collection Center (CGMCC), Beijing, China. Facesoffungi and MycoBank numbers were registered as outlined in
Jayasiri et al. (2015) and MycoBank (2019).
DNA extraction, PCR amplification and sequencing
A sterile scalpel was used to scrape fresh mycelia from pure cultures growing on PDA for one month at 25°C. Fungal
mycelia were scraped off and transferred to a 1.5 ml microcentrifuge tube using a sterilized lancet for genomic DNA
extraction. Ezup Column Fungi Genomic DNA Purification Kit (Sangon Biotech, Shanghai, China) was used to extract
genomic DNA following the manufacturer’s instructions. Five gene regions, LSU, SSU, ITS, TEF1α and RPB2 were
amplified using the primer pairs LR0R/LR5, NS1/NS4, ITS5/ITS4, EF1-983F/EF1-2218R and RPB2-5F/RPB2-7cR,
respectively (Vilgalys & Hester 1990, White et al. 1990, Liu et al. 1999). The amplification procedure was performed
in a 25 μl reaction volume containing 12.5 μl Taq PCR Master Mix (2 ×, with Blue Dye) (Sangon Biotech, China), 9.5
μl ddH2O, 1 μl of each primer and 1 μl DNA template. The PCR condition for LSU, SSU, ITS and TEF1α consisted of
initial denaturation at 94°C for 3 min, followed by 40 cycles at 94°C for 45 s, at 56°C for 50 s, at 72°C for 1 min and
a final extension period at 72°C for 10 min. The PCR condition for RPB2 were amplified with initial denaturation at
95°C for 5 min followed by 40 cycles at 95 °C for 1 min, at 54°C for 2 min, at 72°C for 90 s, and the final extension
at 72°C for 10 min. The PCR products were examined using 1% agarose electrophoresis gels stained with ethidium
bromide. PCR products were purified and sequenced by Shanghai Sangon Biological Engineering Technology and
Services Co., China.
Phylogenetic analysis
Five gene regions (LSU, SSU, ITS, TEF1α, RPB2) were used for phylogenetic analyses and additional sequence data
were obtained from GenBank (Miller & Huhndorf. 2005, Maharachchikumbura et al. 2015, 2016, Hyde et al. 2016a,
Su et al. 2016, Luo et al. 2018, Yang et al. 2018). The sequences were aligned using the multiple alignment program
MAFFT v.7 (http://mafft.cbrc.jp/alignment/server/) (Katoh & Standley 2013). The alignments were visually checked
and optimized manually via AliView (Larsson 2014) wherever necessary. The phylogeny website tools ALTER (http://
www.sing-group.org/ALTER/) were used to transform the aligned fasta file to Phylip format for Maximum likelihood
(ML) analysis and to Nexus format for Maximum parsimony (MP) and Bayesian analyses (Glez-Peña et al. 2010).
Maximum likelihood (ML) analyses were performed using RAxML v.8 (Stamatakis 2006, Stamatakis et al. 2008)
at the CIPRES Science Gateway v.3.3 (Miller et al. 2010). The tree search covered 1000 non-parametric bootstrap
replicates. The best scoring tree was selected from suboptimal trees under the GTRGAMMA substitution model. The
resulting replicates were plotted on the best scoring tree obtained previously. ML bootstrap values equal or greater than
70% were marked near each node.
LIU ET AL.
146 • Phytotaxa 422 (2) © 2019 Magnolia Press
Maximum parsimony (MP) analyses were performed with the heuristic search in PAUP v. 4.0b10 (Swofford
2003). The gaps in the alignment were treated as missing characters and all characters were unordered. Maxtrees were
unlimited, branches of zero length were collapsed and all multiples, equally parsimonious trees were saved. Clade
stability was assessed using a bootstrap (BT) analysis with 1000 replicates, each with 10 replicates of random stepwise
addition of taxa (Hillis & Bull 1993). MP bootstrap values equal or greater than 70% were marked near each node.
Bayesian analyses were performed in MrBayes 3.2.6 (Ronquist et al. 2012). The program MrModeltest2 v. 2.3
(Nylander 2008) was used to determine the best nucleotide substitution model for each data partition. GTR+I+G
substitution model with gamma rates and dirichlet base frequencies were decided for all LSU, SSU, ITS, TEF1α
and RPB2 sequences. The Markov Chain Monte Carlo (MCMC) sampling approach was used to calculate posterior
probabilities (PP) (Rannala & Yang 1996, Zhaxybayeva & Gogarten 2002). Bayesian analyses of six simultaneous
Markov chains were run for 1271000 generations with trees sampled every 100 generations. The first 20% of trees,
representing the burn-in phase of the analyses, were discarded and the remaining trees were used for calculating PP in
the majority rule consensus tree. (Larget & Simon 1999). PP values equal or greater than 0.95 were marked near each
node.
The final trees were visualized with FigTree v1.4.0 (Rambaut 2008) and the layout was edited using Microsoft
PowerPoint 2010. Sequences generated in this study were deposited in GenBank (TABLE 1).
TABLE 1. Taxa and strains used in this study. Newly generated sequences are indicated in bold.
Species Strains GenBank accession no. References
LSU SSU ITS TEF1α RPB2
Annulatascus
velatisporus
HKUCC 3701 AF132320 – – – – Ranghoo et al. 1999
Annulusmagnus
triseptatus
CBS 128831 GQ996540 JQ429242 – – JQ429258 Réblová et al. 2010,
2012
Cryphonectria
parasitica
CMW 7084 JN940858 JN938760 JN942325 – – Schoch et al. 2012
Cryptadelphia
groenendalensis
SMH 3767 EU528001 – – – – Huhndorf et al. 2008
Cryptadelphia
groenendalensis
SH 12 EU528007 – – – – Huhndorf et al. 2008
Distoseptispora
aquatica
MFLUCC 15-
0374
KU376268 – MF077552 – – Su et al. 2016
Distoseptispora
fluminicola
MFLUCC 15-
0417
KU376270 – MF077553 – – Su et al. 2016
Distoseptispora
guttulata
MFLUCC 16-
0183
MF077554 MF077532 MF077543 MF135651 – Yang et al.2018
Distoseptispora
leonensisi
HKUCC 10822 DQ408566 – – – DQ435089 Shenoy et al. 2006
Distoseptispora
obpyriformis
MFLUCC 17-
1694
MG979764 – – MG988422 MG988415 Luo et al. 2018
Distoseptispora
phangngaensis
MFLUCC 16-
0857
MF077556 MF077534 MF077545 MF135653 – Yang et al. 2018
Distoseptispora
rostrata
MFLUCC 16-
0969
MG979766 – MG979758 MG988424 MG988417 Luo et al. 2018
Distoseptispora
suoluoensis
MFLUCC 17-
0224
MF077557 MF077535 MF077546 MF135654 – Yang et al. 2018
Distoseptispora
tectonae
MFLUCC 12-
0291
KX751713 – KX751711 KX751710 KX751708 Hyde et al. 2016a,
2016b
Fragosphaeria
purpurea
CBS 133.34 AB189154 AF096176 AB278192 – – Suh & Blackwell 1999,
Yaguchi et al. 2006
Gnomonia gnomon CBS 199.53 =
AFTOL-ID 952
AF408361 DQ471019 AY818956 DQ471094 DQ470922 Castlebury et al. 2002,
Sogonov et al. 2005,
Spatafora et al. 2006
Jobellisia fraterna SMH 2863 AY346285 – – – – Huhndorf et al. 2004
...continued on the next page
SPORIDESMIUM GUIZHOUENSE SP. NOV. Phytotaxa 422 (2) © 2019 Magnolia Press • 147
TABLE 1. (Continued)
Species Strains GenBank accession no. References
LSU SSU ITS TEF1α RPB2
Magnaporthe salvinii M 21 JF414887 JF414862 – JF710406 – Zhang et al. 2011
Ophiodiaporthe
cyatheae
BCRC 34961 JX570891 JX570890 JX570889 – JX570893 Fu et al. 2013
Ophiostoma piliferum AFTOL-ID 910 DQ470955 DQ471003 – DQ471074 DQ470905 Spatafora et al. 2006
Pseudoplagiostoma
variabile
CBS 113067 GU973611 – GU973536 – – Cheewangkoon et al.
2010
Pyricularia borealis CBS 461.65 KM009150 KM009210 KM009162 KM009198 – Luo et al. 2015
Sordaria fimicola CBS 508.50 AY681160 – AY681188 – – Cai et al. 2006
Sporidesmium
aquaticivaginatum
MFLUCC 15-
0624
KX710142 MF077541 KX710147 MF135660 MF135647 Hyde et al. 2016a,
Yang et al. 2018
Sporidesmium
aquaticum
MFLUCC 15-
0420
KU376273 – – – – Su et al. 2016
Sporidesmium
guizhouense
CGMCC
3.19605
MK818586 MK818588 MK818587 MK828512 MK828516 this study
Sporidesmium
bambusicola
HKUCC 3578 DQ408562 – – – – Shenoy et al. 2006
Sporidesmium
fluminicola
MFLUCC 15-
0346
KU376271 – – – – Su et al. 2016
Sporidesmium
gyrinomorphum
MFLUCC 16-
0186
MF077560 MF077538 MF077549 MF135656 MF135645 Yang et al. 2018
Sporidesmium
macrurum
HKUCC 2740 DQ408555 – – – DQ435086 Shenoy et al. 2006
Sporidesmium
minigelatinosa
NN 47497 DQ408567 – – – DQ435090 Shenoy et al. 2006
Sporidesmium
olivaceoconidium
MFLUCC 15-
0380
KX710139 MF077542 KX710144 MF135661 MF135648 Hyde et al. 2016a,
Yang et al. 2018
Sporidesmium parvum HKUCC 10836 DQ408558 – – – – Shenoy et al. 2006
Sporidesmium
pyriformatum
MFLUCC 15-
0620
KX710141 – KX710146 MF135662 MF135649 Hyde et al. 2016a,
Yang et al. 2018
Sporidesmium
pyriformatum
MFLUCC 15-
0627
KX710143 – KX710148 MF135663 MF135650 Hyde et al. 2016a,
Yang et al. 2018
Sporidesmium sp. HKUCC 10558 DQ408565 – – – – Shenoy et al. 2006
Sporidesmium
submersum
MFLUCC 15-
0421
KU376272 – – – – Su et al. 2016
Sporidesmium
thailandense
MFLUCC 15-
0617
MF077561 – MF077550 MF135657 – Yang et al. 2018
Sporidesmium
thailandense
MFLUCC 15-
0964
MF374370 – MF374361 MF370957 MF370955 Zhang et al. 2017
Sporidesmium
tropicale
MFLUCC 16-
0185
MF077562 MF077539 MF077551 MF135658 MF135646 Yang et al. 2018
Sporidesmium
tropicale
HKUCC 10838 DQ408560 – – – – Shenoy et al. 2006
Thyridium vestitum AFTOL-ID 172 AY544671 AY544715 – DQ471058 DQ470890 James et al. 2006,
Spatafora et al. 2006
Valsella salicis AR 3514 EU255210 – – EU222018 EU219346 Zhang et al. 2006,
Sogonov et al. 2008
LIU ET AL.
148 • Phytotaxa 422 (2) © 2019 Magnolia Press
Results
Phylogenetic analyses
The analyzed dataset comprised 44 taxa retrieved from GenBank and fresh specimens with Sordaria fimicola (CBS
508.50) as the outgroup taxon (TABLE 1). Partial nucleotide sequences of LSU (814bp), SSU (1024bp), ITS (518bp),
TEF1α (841bp) and RPB2 (971bp) for a total of 4168 characters including gaps were used to determine the phylogenetic
placement of the new taxon. The generated MP, ML and Bayesian trees were similar in topology, and the best scoring
RAxML tree is presented in FIGURE 1.
Sporidesmium guizhouense (CGMCC 3.19605) clustered as a sister taxon to S. aquaticivaginatum (MFLUCC 15–
0624) and S. olivaceoconidium (MFLUCC 15–0380) with strong support (100% MLBS, 1.00 PP and 100% MPBS).
The three species grouped distant from the families Sporidesmiaceae and Distoseptisporaceae. Phylogenetic analysis
also showed that S. guizhouense was distinct from the morphologically similar S. minigelatinosa represented by strain
NN47497.
FIGURE 1. Maximum likelihood majority rule consensus tree based on a dataset of combined LSU, SSU, ITS, TEF1α
and RPB2 sequence data. ML bootstrap support ≥ 70 %, Bayesian PP ≥ 0.95 and MP bootstrap support ≥ 70 % values
are given at the nodes. Branches with 100 % MLBS, 1.0 PP and 100% MPBS are shown in black dots. The tree is
rooted with Sordaria fimicola CBS 508.50. Strain numbers are noted after species names. Sporidesmium taxa are in
orange background. New species are indicated in red and ex-type strains are in bold.
SPORIDESMIUM GUIZHOUENSE SP. NOV. Phytotaxa 422 (2) © 2019 Magnolia Press • 149
Taxonomy
Sporidesmium guizhouense L.L. Liu & Z.Y. Liu, sp. nov. Figure 2 and Figure 3.
MycoBank number: MB 830973; Facesoffungi number: FoF 06112
Etymology: Referring to the type location from Guizhou Province, China.
Holotype: HKAS 104644
FIGURE 2. Sporidesmium guizhouense (HKAS 104644, holotype) a Colonies on natural substrate. b Conidiophores
and conidia. c Conidiophore. d–k Conidia. Scale bars: b = 50 μm, c–k = 25 μm.
LIU ET AL.
150 • Phytotaxa 422 (2) © 2019 Magnolia Press
FIGURE 3. Sporidesmium guizhouense on PDA. a Conidiophore and conidia. b–c Conidiophores and conidiogenous
cells. d, e Conidia. f, g Conidiophores and conidia. h, i Colonies on PDA, h from above, i from below. Scale bars: a,
e–h = 50 μm. b, c = 25 μm.
Saprobic on decaying plant substrates. Sexual morph: Unknown. Asexual morph: Colonies effuse, brown to dark
brown, hairy, velvety, glistening. Mycelium partly superficial, partly immersed, composed of pale brown to hyaline,
septate, branched hyphae, 1–3 μm wide. Conidiophores macronematous, mononematous, solitary, cylindrical, straight
or slightly flexuous, 1–5-septate, brown, smooth, thick-walled, slightly narrower at the apex, 23–59 × 2–4.6 μm ( =
38 × 4 μm, n = 20). Conidiogenous cells monoblastic, integrated, terminal, determinate, subcrylindrical, brown, 13–25
× 3.1–4 μm ( = 18 × 3.7 μm, n = 20), truncate at the apex. Conidia acrogenous, solitary, obclavate or obspathulate,
SPORIDESMIUM GUIZHOUENSE SP. NOV. Phytotaxa 422 (2) © 2019 Magnolia Press • 151
rostrate, tapering towards the apex, 9–16-septate, subhyaline to pale olivaceous green or pale brown, paler to hyaline
towards the apex, 46–86 × 7–11.4 μm ( = 64 × 8.4 μm, n = 20), sometimes with an apical mucilaginous sheath,
truncate at base. Conidial secession schizolytic.
Cultural characteristics: Conidia germinating on WA within 24h and germ tubes growing from the apex. Colonies
on PDA slow growing, reaching 11–13 mm diam. after 2 months at 25°C in dark, circular, wrinkled, reverse brown to
pale brown from the center to the entire margin, sporulation abundant. Mycelium composed of septate, subhyaline to
pale brown, branched, smooth hyphae, 1–4 μm wide. Conidiophores macronematous, solitary, cylindrical, straight or
slightly flexuous, 1–7-septate, brown, smooth, thick-walled, slightly tapering towards the apex, 28–71 × 3–5 μm ( =
46 × 4 μm, n = 20), sometimes reduced to a single conidiogenous cell. Conidiogenous cells monoblastic, integrated,
terminal, doliiform or subcylindrical, brown, 8–13 × 3.7–6 μm ( = 11 × 4 μm, n = 20), truncate at the apex. Conidia
acrogenous, solitary, obclavate or obspathulate, rostrate, tapering towards the apex, 12–33-septate, subhyaline to pale
brown, paler to hyaline towards the apex, 63–199 × 6–11 μm ( = 115 × 8 μm, n = 20).
Material examined: CHINA. Guizhou Province: Guiyang City, Baihua Lake, 26°39.29′N, 106°32.22′E, 1205 m
asl, on submerged decaying twig, 18 April 2018, Lingling Liu, 18B–40 (HKAS 104644 holotype, GZAAS 19–005
isotype), ex-type living cultures CGMCC 3.19605, GZCC 19–005.
Notes: Sporidesmium guizhouense resembles S. aquaticivaginatum J. Yang & K.D. Hyde (Hyde et al. 2016a),
S. bambusae M.B. Ellis (Ellis 1958, 1976, Hernández-Gutiérrez & Sutton 1997), S. minigelatinosa Matush. (Wu &
Zhuang 2005), S. novozymium W.P. Wu (Wu & Zhuang 2005), S. olivaceoconidium J. Yang & K.D. Hyde (Hyde et al.
2016a) and S. palmicola W.P. Wu (Wu & Zhuang 2005) in having morphologically similar obclavate conidia with an
apical mucilaginous sheath. However, conidiophores of S. guizhouense are shorter than those of S. aquaticivaginatum
(60–125 μm long) and S. novozymium (150–200 μm long). Sporidesmium guizhouense differs also from S. bambusae
and S. olivaceoconidium in conidial size. Conidia of S. guizhouense are larger than those of S. olivaceoconidium
(25–50 × 6–10 μm) but smaller than those of S. bambusae (65–100 × 6–10 μm). Synnematous conidiophores were
described in S. palmicola but those were not observed in S. guizhouense.
Discussion
Hyde et al. (2016a) and Yang et al. (2018) accommodated S. aquaticivaginatum (MFLUCC 15–0624) and S.
olivaceoconidium (MFLUCC 15–0380) within Sporidesmium sensu stricto based on a combined ITS-LSU sequence
data comprising 39 taxa and a combined LSU, SSU, TEF1α, RPB2 and ITS dataset including 125 taxa, respectively. In
this study, however, phylogenetic analyses using five gene regions (LSU, SSU, ITS, TEF1α, and RPB2) and including
44 taxa showed different results from previous studies. Sporidesmium guizhouense (CGMCC 3.19605) formed a
strongly-supported monophyletic clade with S. aquaticivaginatum (MFLUCC 15–0624) and S. olivaceoconidium
(MFLUCC 15–0380) but they grouped distant from Sporidesmium sensu stricto. Therefore, the phylogenetic placement
of these three taxa remains incertae sedis within Sordariomycetes. We prefer to avoid proposing a new genus for S.
aquaticivaginatum (MFLUCC 15–0624), S. guizhouense (CGMCC 3.19605), and S. olivaceoconidium (MFLUCC 15–
0380), based on current molecular data, until more collections with DNA sequence data together with morphological
differences become available for study. We therefore accept S. guizhouense (CGMCC 3.19605) within Sporidesmium
sensu lato due to its phylogenetic position and the currently accepted polyphyletic nature of the genus.
Acknowledgments
We would like to thank the Research of Featured Microbial Resources and Diversity Investigation in the Southwest
Karst area (Project No. 2014FY120100) and Science and Technology Fund of Guizhou Province (Project No. Guizhou
[2017]1181). Ling-Ling Liu thanks Professor Jian-Kui Liu for the correction to the manuscript.
References
Bao, D.F., Luo, Z.L., Liu, J.K., Bhat, D.J., Sarunya, N., Li, W.L., Su, H.Y. & Hyde, K.D. (2018) Lignicolous freshwater fungi in China III:
New species and record of Kirschsteiniothelia from northwestern Yunnan Province. Mycosphere 9: 755–768.
LIU ET AL.
152 • Phytotaxa 422 (2) © 2019 Magnolia Press
https://doi.org/10.5943/mycosphere/9/4/4
Cai, L., Tsui, C.K.M., Zhang, K.Q. & Hyde, K.D. (2002) Aquatic fungi from Lake Fuxian, Yunnan, China. Fungal Diversity 9: 57–70.
Cai, L., Zhang, K., McKenzie, E.H.C. & Hyde, K.D. (2003) Freshwater fungi from bamboo and wood submerged in the Liput River in
the Philippines. Fungal Diversity 13: 1–12.
Cai, L., Jeewon, R. & Hyde, K.D. (2006) Phylogenetic investigations of Sordariaceae based on multiple gene sequences and morphology.
Mycological Research 110: 137–150.
https://doi.org/10.1016/j.mycres.2005. 09.014
Castlebury, L.A., Rossman, A.Y., Jaklitsch, W.J. & Vasilyeva, L.N. (2002) A preliminary overview of the Diaporthales based on large
subunit nuclear ribosomal DNA sequences. Mycologia 94: 1017–1031.
https://doi.org/10.1080/15572536.2003.11833157
Cheewangkoon, R., Groenewald, J.Z., Verkley, G.J.M., Hyde, K.D., Wingfield, M.J., Gryzenhout, M., Summerell, B.A., Denman, S.,
Toanun, C. & Crous, P.W. (2010) Re-evaluation of Cryptosporiopsis eucalypti and Cryptosporiopsis-like species occurring on
Eucalyptus leaves. Fungal Diversity 44: 89–105.
https://doi.org/10.1007/s13225-010-0041-5
Crous, P.W., Wingfield, M.J., Burgess, T.I., Carnegie, A.J., Hardy, G.E.St.J., Smith, D, Summerell, B.A., Cano-Lira, J.F., Guarro, J.,
Houbraken, J. Lombard, L. Martín, M.P., Sandoval-Denis, M., Alexandrova, A.V., Barnes, C.W., Baseia, I.G., Bezerra, J.D.P.,
Guarnaccia, V., May, T.W., Hernández-Restrepo, M., Stchigel, A.M., Miller, A.N., Ordoñez, M.E., Abreu, V.P., Accioly, T., Agnello,
C., Agustin Colmán, A., Albuquerque, C.C., Alfredo, D.S., Alvarado, P., Araújo-Magalhães, G.R., Arauzo, S., Atkinson, T., Barili,
A., Barreto, R.W., Bezerra, J.L., Cabral, T.S., Camello Rodríguez, F., Cruz, R.H.S.F., Daniëls, P.P., da Silva, B.D.B., de Almeida,
D.A.C., de Carvalho Júnior, A.A., Decock, C.A., Delgat, L., Denman, S., Dimitrov, R.A., Edwards, J., Fedosova, A.G., Ferreira,
R.J., Firmino, A.L., Flores, J.A., García, D., Gené, J., Giraldo, A., Góis, J.S., Gomes, A.A.M., Gonçalves, C.M., Gouliamova,
D.E., Groenewald, M., Guéorguiev, B.V., Guevara-Suarez, M., Gusmão, L.F.P., Hosaka, K., Hubka, V., Huhndorf, S.M., Jadan,
M., Jurjević, Ž., Kraak, B., Kučera, V., Kumar, T.K.A., Kušan, I., Lacerda, S.R., Lamlertthon, S., Lisboa, W.S., Loizides, M.,
Luangsa-Ard, J.J., Lysková, P., Mac Cormack, W.P., Macedo, D.M., Machado, A.R., Malysheva, E.F., Marinho, P., Matočec, N.,
Meijer, M., Mešić, A., Mongkolsamrit, S., Moreira, K.A., Morozova, O.V., Nair, K.U., Nakamura, N., Noisripoom, W., Olariaga, I.,
Oliveira, R.J.V., Paiva, L.M., Pawar, P., Pereira, O.L., Peterson, S.W., Prieto, M., Rodríguez-Andrade, E., Rojo De Blas, C., Roy, M.,
Santos, E.S., Sharma, R., Silva, G.A., Souza-Motta, C.M., Takeuchi-Kaneko, Y., Tanaka, C., Thakur, A., Smith, M., Th Tkalčec, Z.,
Valenzuela-Lopez, N., van der Kleij, P., Verbeken, A., Viana, M.G., Wang, X.W., Groenewald, J.Z. (2017) Fungal Planet description
sheets: 625–715. Persoonia 39: 270–467.
https://doi.org/10.3767/persoonia.2017.39.11
Delgado, G., Miller, A.N. & Piepenbring, M. (2018) South florida microfungi: Castanedospora, a new genus to accommodate Sporidesmium
pachyanthicola (Capnodiales, Ascomycota). Cryptogamie Mycologie 39: 109–127.
https://doi.org/10.7872/crym/v39.iss1.2018.109
Dubey, R. & Pandey, A.K. (2011) New species of Scytalidium Pesante and Sporidesmium Link ex Fries from India. Nelumbo 53: 1–6.
https://doi.org/10.20324/nelumbo/v53/2011/57494
Dubey, R. & Sengupta, S. (2015) Sporidesmium bilgiriense—a new species of microfungi from Western Ghats of India. Current Research
in Environmental & Applied Mycology 5: 390–393.
https://doi.org/10.5943/cream/5/4/10
Ellis, M.B. (1958) Clasterosporium and some allied dematiaceae-phragmosporae: I. Mycological Papers 70: 1–89.
Ellis, M.B. (1971) Dematiaceous hyphomycetes. Commonwealth Mycological Institute, Kew Pubilshing 604 pp.
Ellis, M.B. (1976) More dematiaceous hyphomycetes. Commonwealth Mycological Institute, Kew Pubilshing 507 pp.
Fu, C.H., Hsieh, H.M., Chen, C.Y., Chang, T.T., Huang, Y.M. & Ju, Y.M. (2013) Ophiodiaporthe cyatheae gen. et sp. nov., a diaporthalean
pathogen causing a devastating wilt disease of Cyathea lepifera in Taiwan. Mycologia 105: 861–872.
https://doi.org/10.3852/12-346
Glez-Peña, D., Reboiro-Jato, M., Maia, P., Rocha, M., Díaz, F. & Fdez-Riverola, F. (2010) AIBench: A rapid application development
framework for translational research in biomedicine. Computer Methods and Programs in Biomedicine 98: 191–203.
https://doi.org/10.1016/j.cmpb.2009.12.003
Graça, M.A.S., Hyde, K.D. & Chauvet, E. (2015) Aquatic hyphomycetes and litter decomposition in tropical - subtropical low order
streams. Fungal Ecology 19: 182–189.
https://doi.org/10.1016/j. funeco.2015.08.001
Hernández-Gutiérrez, A. & Sutton, B.C. (1997) Imimyces and Linkosia, two new genera segregated from Sporidesmium sensu lato, and
redescription of Polydesmus. Mycological Research 101: 201–209.
https://doi.org/10.1017/s0953756296002419
Hillis, D.M. & Bull, J.J. (1993) An empirical test of bootstrapping as a method for assessing confidence in phylogenetic analysis. Systematic
SPORIDESMIUM GUIZHOUENSE SP. NOV. Phytotaxa 422 (2) © 2019 Magnolia Press • 153
Biology 42: 182–192.
https://doi.org/10.1093/sysbio/ 42.2.182
Ho, W.H., Hyde, K.D., Hodgkiss, I.J. & Yanna (2001) Fungal communities on submerged wood from streams in Brunei, Hong Kong, and
Malaysia. Mycological Research 105: 1492–1501.
https://doi.org/10.1017/s095375620100507x
Ho, W.H., Yanna, Hyde, K.D. & Hodgkiss, I.J. (2002) Seasonality and sequential occurrence of fungi on wood submerged in Tai Po Kau
Forest Stream, Hong Kong. Fungal Diversity 10: 21–43.
Hughes, S.J. (1979) Relocation of species of Endophragmia auct. with notes on relevant generic names. New Zealand Journal of Botany
17: 161–163.
https://doi.org/10.1080/0028825X.1979.10426887
Huhndorf, S.M., Miller, A.N. & Fernández, F.A. (2004) Molecular systematics of the Sordariales: the order and the family Lasiosphaeriaceae
redefined. Mycologia 96: 368–387.
https://doi.org/10.2307/3762068
Huhndorf, S.M., Greif, M., Mugambi, G.K. & Miller, A.N. (2008) Two new genera in the Magnaporthaceae, a new addition to
Ceratosphaeria and two new species of Lentomitella. Mycologia 100: 940–955.
https://doi.org/10.3852/08-037
Hyde, K.D. & Goh, T.K. (1998) Fungi on submerged wood in Lake Barrine, north Queensland, Australia. Mycological Research 102:
739–749.
https://doi.org/10. 1017/s0953756297005868
Hyde, K.D., Hongsanan, S., Jeewon, R., Bhat, D.J., McKenzie, E.H.C., Jones, E.B.G., Phookamsak, R., Ariyawansa, H.A., Boonmee, S.,
Zhao, Q., Abdel-Aziz, F.A., Abdel-Wahab, M.A., Banmai, S., Chomnunti, P., Cui, B.K., Daranagama, D.A., Das, K., Dayarathne,
M.C., de Silva, N.I., Dissanayake, A.J., Doilom, M., Ekanayaka, A.H., Gibertoni, T.B., Go’esNeto, A., Huang, S.K., Jayasiri, S.C.,
Jayawardena, R.S., Konta, S., Lee, H.B., Li, W.J., Lin, C.G., Liu, J.K., Lu, Y.Z., Luo, Z.L., Manawasinghe, I.S., Manimohan, P.,
Mapook, A., Niskanen, T., Norphanphoun, C., Papizadeh, M., Perera, R.H., Phukhamsakda, C., Richter, C., de Santiago, A.L.C.M.A.,
Drechsler-Santos, E.R., Senanayake, I.C., Tanaka, K., TMDS, T., Thambugala, K.M., Tian, Q., Tibpromma, S., Thongbai, B., Vizzini,
A., Wanasinghe, D.N., Wijayawardene, N.N., Wu, H.X., Yang, J., Zeng, X.Y., Zhang, H., Zhang, J.F., Bulgakov, T.S., Camporesi, E.,
Bahkali, A.H., Amoozegar, M.A., Araujo-Neta, L.S., Ammirati, J.F., Baghela, A., Bhatt, R.P., Bojantchev, D., Buyck, B., da Silva,
G.A., de Lima, C.L.F., de Oliveira, R.J.V., de Souza, C.A.F., Dai, Y.C., Dima, B., Duong, T.T., Ercole, E., MafaldaFreire, F., Ghosh,
F., Hashimoto, A., Kamolhan, S., Kang, J.C., Karunarathna, S.C., Kirk, P.M., Kytövuori, I., Lantieri, A., Liimatainen, K., Liu, Z.Y.,
Liu, X.Z., Lücking, R., Medardi, G., Mortimer, P.E., TTT, N., Promputtha, I., KNA, R., Reck, M.A., Lumyong, S., Shahzadeh-Fazeli,
S.A., Stadler, M., Soudi, M.R., Su, H.Y., Takahashi, T., Tangthirasunun, N., Uniyal, P., Wang, Y., Wen, T.C., Xu, J.C., Zhang, Z.K.,
Zhao, Y.C., Zhou, J.L. & Zhu, L. (2016a) Fungal diversity notes 367–490: taxonomic and phylogenetic contributions to fungal taxa.
Fungal Diversity 80: 1–270.
https://doi.org/10.1007/s13225-016-0373-x
Hyde, K.D., Fryar, S., Tian, Q., Bahkali, A.H. & Xu, J.C. (2016b) Lignicolous freshwater fungi along a north–south latitudinal gradient
in the Asian/Australian region; can we predict the impact of global warming on biodiversity and function? Fungal Ecology 19:
190–200.
https://doi.org/10.1016/ j.funeco.2015.07.002
Iturriaga, T., Hawksworth, D.L. & Crane, J.L. (2008) ‘Sporidesmium’ lichenicola sp. nov., a new lichenicolous fungus on Leptogium from
Venezuela. Mycologia 100: 392–396.
https://doi.org/10.3852/06-166R
Index Fungorum (2019) Available from: http://www.indexfungorum.org/ (accessed April 2019)
James, T.Y., Kauff, F., Schoch, C., Matheny, P.B., Hofstetter, V., Cox, C.J., Celio, G., Gueidan, C., Fraker, E., Miadlikowska, J., Lumbsch,
T., Rauhut, A., Reeb, V., Arnold, A.E., Amtoft, A., Stajich, J.E., Hosaka, K., Sung, G.H., Johnson, D., O’Rourke, B., Binder, M.,
Curtis, J.M., Slot, J.C., Wang, Z., Wilson, A.W., Schüßler, A., Longcore, J.E., O’Donnell, K., MozleyStandridge, K., Porter, D.,
Letcher, P.M., Powell, M.J., Taylor, J.W., White, M.M., Griffith, G.W., Davies, D.R., Sugiyama, J., Rossman, A.Y., Rogers, J.D.,
Pfister, D.H., Hewitt, D., Hansen, K., Hambleton, S., Shoemaker, R.A., Kohlmeyer, J., Volkmann-Kohlmeyer, B., Spotts, R.A.,
Serdani, M., Crous, P.W., Hughes, K.W., Matsuura, K., Langer, E., Langer, G., Untereiner, W.A., Lücking, R., Büdel, B., Geiser,
D.M., Aptroot, A., Buck, W.R., Cole, M.S., Diederich, P., Printzen, C., Schmitt, I., Schultz, M., Yahr, R., Zavarzin, A., Hibbett, D.S.,
Lutzoni, F., McLaughlin, D.J., Spatafora, J.W. & Vilgalys, R. (2006) Reconstructing the early evolution of the fungi using a six-gene
phylogeny. Nature 443: 818–822.
https://doi.org/10.1038/nature05110
Jayasiri, S.C., Hyde, K.D., Ariyawansa, H.A., Bhat, D.J., Buyck, B., Cai, L., Dai, Y.C., Abd-Elsalam, K.A., Ertz, D., Hidayat, I., Jeewon,
R., Jones, E.B.G., Bahkali, A.H., Karunarathna, S.C., Liu, J.K., Luangsa-ard, J.J., Lumbsch, H.T., Maharachchikumbura, S.S.N.,
LIU ET AL.
154 • Phytotaxa 422 (2) © 2019 Magnolia Press
McKenzie, E.H.C., Moncalvo, J.M., Ghobad-Nejhad, M., Nilsson, H., Pang, K.L., Pereira, O.L., Phillips, A.J.L., Raspé, O., Rollins,
A.W., Romero, A.I., Etayo, J., Selçuk, F., Stephenson, S.L., Suetrong, S., Taylor, J.E., Tsui, C.K.M., Vizzini, A., Abdel-Wahab,
M.A., Wen, T.C., Boonmee, S., Dai, D.Q., Daranagama, D.A., Dissanayake, A.J., Ekanayaka, A.H., Fryar, S.C., Hongsanan, S.,
Jayawardena, R.S., Li, W.J., Perera, R.H., Phookamsak, R., Silva, N.I., Thambugala, K.M., Tian, Q., Wijayawardene, N.N., Zhao,
R.L., Zhao, Q., Kang, J.C. & Promputtha, I. (2015) The faces of fungi database: fungal names linked with morphology, phylogeny
and human impacts. Fungal Diversity 74: 3–18.
https://doi.org/10.1007/s13225-015-0351-8
Katoh, K. & Standley, D.M. (2013) MAFFT multiple sequence alignment software version 7: improvements in performance and usability.
Molecular Biology and Evolution 30: 772–780.
https://doi.org/10.1093/molbev/mst010
Kirk, P.M. (1982) New or interesting microfungi VI. Sporidesmiella gen. nov. (Hyphomycetes). Transactions of the British Mycological
Society 79: 479–489.
https://doi.org/10.1016/s0007-1536(82)80040-5
Larget, B. & Simon, D.L. (1999) Markov Chain Monte Carlo algorithms for the Bayesian analysis of phylogenetic trees. Molecular
Biology and Evolution 16: 750–759.
https://doi.org/10.1093/oxfordjournals.molbev.a026160
Larsson, A. (2014) AliView: a fast and lightweight alignment viewer and editor for large datasets. Bioinformatics 30: 3276–3278.
https://doi.org/10.1093/bioinformatics/btu531
Li, W.L., Luo, Z.L., Liu, J.K., Bhat, D.J., Bao, D.F., Su, H.Y. & Hyde, K.D. (2017) Lignicolous freshwater fungi from China I:
Aquadictyospora lignicola gen. et sp. nov. and new record of Pseudodictyosporium wauense from northwestern Yunnan Province.
Mycosphere 8: 1587–1597.
https://doi.org/10.5943/mycosphere/8/10/1
Link, J.H.F. (1809) Observationes in ordines plantarum naturales. Magazin der Gesellschaft Naturforschender Freunde Berlin 3: 3–42.
Liu, Y.J., Whelen, S. & Hall, B.D. (1999) Phylogenetic relationships among ascomycetes: evidence from an RNA Polymerase II subunit.
Molecular Biology and Evolution 16: 1799–1808.
https://doi.org/10.1093/oxfordjournals. molbev.a026092
Luo, J., Yin, J., Cai, L., Zhang, K.Q. & Hyde, K.D. (2004) Freshwater fungi in Lake Dianchi, a heavily polluted lake in Yunnan, China.
Fungal Diversity 16: 93–112.
Luo, J., Walsh, E. & Zhang, N. (2015) Toward monophyletic generic concepts in Magnaporthales: species with Harpophora asexual states.
Mycologia 107: 641–646.
https://doi.org/10.3852/14-302
Luo, Z.L., Hyde, K.D., Liu, J.K., Bhat, D.J., Bao, D.F., Li, W.L. & Su, H.Y. (2018) Lignicolous freshwater fungi from China II: Novel
Distoseptispora (Distoseptisporaceae) species from northwestern Yunnan Province and a suggested unified method for studying
lignicolous freshwater fungi. Mycosphere 9: 444–461.
https://doi.org/10.5943/mycosphere/9/3/2
Ma, J., Ma, L.G., Zhang, Y.D., Xia, J.W. & Zhang, X.G. (2012) New species and record of Sporidesmium from southern China. Mycotaxon
119: 17–25.
https://doi.org/10.5248/119.103
Maharachchikumbura, S.S.N., Hyde, K.D., Jones, E.B.G., McKenzie, E.H.C., Huang, S.K., Abdel-Wahab, M.A., Daranagama, D.A.,
Dayarathne, M.C., D’souza, M.J., Goonasekara, I.D., Hongsanan, S., Jayawardena, R.S., Kirk, P.M., Konta, S., Liu, J.K., Liu,
Z.Y., Norphanphoun, C., Pang, K.L., Perera, R.H., Senanayake, I.C., Shang, Q.J., Shenoy, B.D., Xiao, Y., Bahkali, A.H., Kang, J.,
Somrothipol, S., Suetrong, S., Wen, T. & Xu, J.C. (2015) Towards a natural classification and backbone tree for Sordariomycetes.
Fungal Diversity 72: 199–301.
https://doi.org/10.1007/s13225-015-0331-z
Maharachchikumbura, S.S.N., Hyde, K.D., Jones, E.B.G., McKenzie, E.H.C., Bhat, D.J., Dayarathne, M., Huang, S.K., Norphanphoun,
C., Senanayake, I.C., Perera, R.H., Shang, Q., Xiao, Y., D’souza, M.J., Hongsanan, S., Jayawardena, R.S., Daranagama, D.A., Konta,
S., Goonasekara, I.D., Zhuang, W.Y., Jeewon, R., Phillips, A.J.L., AbdelWahab, M.A., Al-Sadi, A.M., Bahkali, A.H., Boonmee,
S., Boonyuen, N., Cheewangkoon, R., Dissanayake, A.J., Kang, J., Liu, J.K., Liu, X., Liu, Z.Y., Pang, K.L., Phookamsak, R.,
Promputtha, I., Suetrong, S., Wen, T. & Wijayawardene, N.N. (2016) Families of Sordariomycetes. Fungal Diversity 79: 1–317.
https://doi.org/10.1007/s13225-016-0369-6
Miller, A.N. & Huhndorf, S.M. (2005) Multi-gene phylogenies indicate ascomal wall morphology is a better predictor of phylogenetic
relationships than ascospore morphology in the Sordariales (Ascomycota, Fungi). Molecular Phylogenetics and Evolution 35: 60–
75.
https://doi.org/10.1016/j.ympev.2005.01.007
SPORIDESMIUM GUIZHOUENSE SP. NOV. Phytotaxa 422 (2) © 2019 Magnolia Press • 155
Miller, M.A., Pfeiffer, W. & Schwartz, T. (2010) Creating the CIPRES Science Gateway for inference of large phylogenetic trees. In:
Proceedings of the Gateway Computing Environments Workshop (GCE). New Orleans, pp. 1–8.
https://doi.org/10.1109/GCE.2010.5676129
MycoBank (2019) Available from: http://www.MycoBank.org/ (accessed April 2019)
Nylander, J. (2008) MrModeltest2 v. 2.3 (Program for selecting DNA substitution models using PAUP*). Evolutionary Biology Centre,
Uppsala.
Ranghoo, V.M., Hyde, K.D., Liew, E.C.Y. & Spatafora, J.W. (1999) Family placement of Ascotaiwania and Ascolacicola based on DNA
sequences from the large subunit rRNA gene. Fungal Diversity 2: 159–168.
Rambaut, A. & Drummond, A. (2008) FigTree: Tree figure drawing tool, version 1.2.2. Institute of Evolutionary Biology, University of
Edinburgh.
Rannala, B. & Yang, Z. (1996) Probability distribution of molecular evolutionary trees: a new method of phylogenetic inference. Journal
of Molecular Evolution 43: 304–311.
https://doi.org/10.1007/BF02338839
Réblová, M., Fournier, J. & Hyde, K.D. (2010) Achroceratosphaeria, a new ascomycete genus in the Sordariomycetes, and re-evaluation
of Ceratosphaeria incolorata. Fungal Diversity 43: 75–84.
https://doi.org/10.1007/s13225-010-0032-6
Réblová, M., Seifert, K.A., Fournier, J. & Štěpánek, V. (2012) Phylogenetic classification of Pleurothecium and Pleurotheciella gen.
nov. and its dactylaria-like anamorph (Sordariomycetes) based on nuclear ribosomal and protein-coding genes. Mycologia 104:
1299–1314.
https://doi.org/10.3852/12-035
Ronquist, F., Teslenko, M., Mark, P., Ayres, D.L., Darling, A., Höhna, S., Larget, B., Liu, L., Suchard, M.A. & Huelsenbeck, J.P. (2012)
MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology 61: 539–
542.
https://doi.org/10.1093/sysbio/sys029
Schoch, C.L., Seifert, K.A., Huhndorf, S., Robert, V., Spougea, J.L., Levesqueb, C.A., Chen, W. & Fungal Barcoding Consortiuma
(2012) Nuclear ribosomal internal transcribed spacer (ITS) region as a universal DNA barcode marker for Fungi. Proceedings of the
National Academy of Sciences of the United States America 109: 6241–6246.
https://doi.org/10.1073/pnas.1117018109
Seifert, K, Morgan-Jones, G, Gams, W. & Kendrick, B. (2011) The genera of hyphomycetes. CBS–KNAW Fungal Biodiversity Centre,
Utrecht, Netherlands.
Shenoy, B.D., Jeewon, R., Wu, W.P., Bhat, D.J. & Hyde, K.D. (2006) Ribosomal and RPB2 DNA sequence analyses suggest that
Sporidesmium and morphologically similar genera are polyphyletic. Mycological Research 110: 916–928.
https://doi.org/10.1016/j.mycres.2006.06.004
Shenoy, B.D., Jeewon, R., Wang, H., Amandeep, K., Ho, W.H., Bhat, D.J., Crous, P.W. & Hyde, K.D. (2010) Sequence data reveals
phylogenetic affinities of fungal anamorphs Bahusutrabeeja, Diplococcium, Natarajania, Paliphora, Polyschema, Rattania and
Spadicoides. Fungal Diversity 44: 161–169.
https://doi.org/10.1007/s13225-010-0059-8
Shoemaker, R.A. & Hambleton, S. (2001) Helminthosporium asterinum, Polydesmus elegans, Imimyces, and allies. Canadian Journal of
Botany 79: 592–599.
https://doi.org/10.1139/b01-038
Sogonov, M.V., Castlebury, L.A., Rossman, A.Y., Farr, D.F. & White, J.F. (2005) The type species of genus Gnomonia, G. gnomon, and
the closely related G. setacea. Sydowia 57: 102–119.
Sogonov, M.V., Castlebury, L.A., Rossman, A.Y., Mejía, L.C. & White, J.F. (2008) Leaf-inhabiting genera of the Gnomoniaceae,
Diaporthales. Studies in Mycology 62: 1–77.
https://doi.org/10.3114/sim.2008.62.01
Spatafora, J.W., Sung, G.H., Johnson, D., Hesse, C., O’Rourke, B., Serdani, M., Spotts, R., Lutzoni, F., Hofstetter, V., Miadlikowska, J.,
Reeb, V., Gueidan, C., Fraker, E., Lumbsch, T., Lücking, R., Schmitt, I., Hosaka, K., Aptroot, A., Roux, C., Miller, A.N., Geiser,
D.M., Hafellner, J., Hestmark, G., Arnold, A.E., Büdel, B., Rauhut, A., Hewitt, D., Untereiner, W.A., Cole, M.S., Scheidegger, C.,
Schultz, M., Sipman, H. & Schoch, C.L. (2006) A five-gene phylogeny of Pezizomycotina. Mycologia 98: 1018–1028.
https://doi.org/10.3852/mycologia.98.6.1018
Stamatakis, A. (2006) RAxML-VI-HPC: maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models.
Bioinformatics 22: 2688–2690.
https://doi.org/10.1093/bioinformatics/btl446
Stamatakis, A., Hoover, P. & Rougemont, J. (2008) A rapid bootstrap algorithm for the RAxML web servers. Systematic Biology 57:
LIU ET AL.
156 • Phytotaxa 422 (2) © 2019 Magnolia Press
758–771.
https://doi.org/10.1080/10635150802429642
Su, H.Y., Hyde, K.D., Maharachchikumbura, S.S.N., Ariyawansa, H.A., Luo, Z.L., Promputtha, I., Tian, Q., Lin, C.G., Shang, Q.J., Zhao,
Y.C., Chai, H.M., Liu, X.Y., Bahkali, A.H., Bhat, D.J., McKenzie, E.H.C. & Zhou, D.Q. (2016) The families Distoseptisporaceae
fam. nov., Kirschsteiniotheliaceae, Sporormiaceae and Torulaceae, with new species from freshwater in Yunnan Province, China.
Fungal Diversity 80: 375–409.
https://doi.org/10.1007/s13225-016-0362-0
Subramanian, C.V. (1992) A reassessment of Sporidesmium (hyphomycetes) and some related taxa. Proceedings of the Indian Academy
of Sciences (Plant Sciences) 58: 179–190.
Swofford, D.L. (2003) PAUP*: Phylogenetic analysis using parsimony (*and other methods). Version 4. Sinauer, Sunderland.
Suh, S.O. & Blackwell, M. (1999) Molecular phylogeny of the cleistothecial fungi placed in Cephalothecaceae and Pseudeurotiaceae.
Mycologia 91: 836–848.
https://doi.org/10.2307/3761537
Vilgalys, R. & Hester, M. (1990) Rapid genetic identification and mapping enzymatically amplified ribosomal DNA from several
Cryptococcus species. Journal of Bacteriology 172: 4238–4246.
https://doi.org/10.1128/jb.172.8.4238-4246.1990
White, T.J., Bruns, T., Lee, S. & Taylor, J. (1990) Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics.
In: Innis, M., Gelfand, D., Shinsky, J. & White, T. (Ed.) PCR protocols: a guide to methods and applications. Academic Press, New
York, pp. 315−322.
https://doi.org/10.1016/B978-0-12-372180-8.50042-1
Wong, M.K.M., Goh, T.K., Hodgkiss, I.J., Hyde, K.D., Ranghoo, V.M., Tsui, C.K.M., Ho, W.H., Wong, S.W. & Yuen, T.K. (1998) Role of
fungi in freshwater ecosystems. Biodiversity and Conservation 7: 1187–1206.
https://doi.org/10.1023/A:1008883716975
Wu, W.P. & Zhuang, W.Y. (2005) Sporidesmium, Endophragmiella and related genera from China. Fungal Diversity Research Series 15:
1–351.
Yaguchi, T., Sano, A., Yarita, K., Suh, M.K., Nishimura, K. & Udagawa, S. (2006) A new Cephalotheca isolate from Korean patient.
Mycotaxon 96: 309–322.
Yang, J., Maharachchikumbura, S.S.N., Liu, J.K., Hyde, K.D., Jones, E.B.G., Al-Sadi, A.M. & Liu, Z.Y. (2018) Pseudostanjehughesia
aquitropica gen. et sp. nov. and Sporidesmium sensu lato species from freshwater habitats. Mycological Progress 17: 591–616.
https://doi.org/10.1007/s11557-017-1339-4
Zhang, H., Dong, W., Hyde, K.D., Maharachchikumbura, S.S.N., Hongsanan, S., Bhat, D.J., Al-Sadi, A.M. & Zhang, D. (2017) Towards
a natural classification of Annulatascaceae-like taxa: introducing Atractosporales ord. nov. and six new families. Fungal Diversity
85: 75–110.
https://doi.org/10.1007/s13225-017-0387-z
Zhang, N., Castlebury, L.A., Miller, A.N., Huhndorf, S.M., Schoch, C.L., Seifert, K.A., Rossman, A.Y., Rogers, J.D., Kohlmeyer, J.,
Volkmann-Kohlmeyer, B. & Sung, G.H. (2006) An overview of the systematics of the Sordariomycetes based on a four-gene
phylogeny. Mycologia 98: 1076–1087.
https://doi.org/10.3852/mycologia.98.6.1076
Zhang, N., Zhao, S. & Shen, Q. (2011) A six-gene phylogeny reveals the evolution of mode of infection in the rice blast fungus and allied
species. Mycologia 103: 1267–1276.
https://doi.org/10.3852/11–022
Zhaxybayeva, O. & Gogarten, J.P. (2002) Bootstrap, Bayesian probability and maximum likelihood mapping: exploring new tools for
comparative genome analyses. BMC Genomics 3: 4.
https://doi.org/10.1186/1471-2164-3-4
Zhou, D.Q., Hyde, K.D. & Wu, X.L. (2001) New records of Ellisembia, Penzigomyces, Sporidesmium and Repetophragma species on
bamboo from China. Acta Botanica Yunnanica 23: 45–51.