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Phytotaxa 559 (2): 176–184
https://www.mapress.com/pt/
Copyright © 2022 Magnolia Press Article PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
176 Accepted by Ruvishika Shehali Jayawardena: 10 Aug. 2022; published: 23 Aug. 2022
https://doi.org/10.11646/phytotaxa.559.2.6
Neodigitodesmium, a novel genus of family Dictyosporiaceae from Sichuan
Province, China
WENHUI TIAN1,2, YANPENG CHEN1,3 & SAJEEWA S. N. MAHARACHCHIKUMBURA1,4*
1 School of Life Science and Technology, Center for Informational Biology, University of Electronic Science and Technology of China,
Chengdu 611731, P.R. China
2
�
wenhuitian@qq.com; https://orcid.org/0000-0001-8892-853X
3
�
yanpengch@qq.com; https://orcid.org/0000-0002-2554-5272
4
�
sajeewa83@yahoo.com; https://orcid.org/0000-0001-9127-0783
*Corresponding author:
�
sajeewa83@yahoo.com
Abstract
A fungus within the family Dictyosporiaceae was isolated from submerged-decaying wood in Sichuan Province, China.
Combined ITS, LSU, SSU and tef1 sequence data showed that the species represented a new genus and is described here as
Neodigitodesmium cheirosporum gen. et sp. nov. Neodigitodesmium distinct from other genera in the family Dictyosporiaceae
based on a hyaline and holoblastic basal connecting conidiogenous cell, conical shaped and smaller with fewer rows (1–4)
of conidia. In addition, phylogenetic analysis and morphological comparison showed that the recently introduced genus
Paradictyocheirospora is congeneric with Digitodesmium. Based on these results, Paradictyocheirospora is reduced to
synonymy under the older generic name Digitodesmium.
Keywords: Asexual morph, Digitodesmium, Dothideomycetes, Paradictyocheirospora, Taxonomy
Introduction
The family Dictyosporiaceae was established within the order Pleosporales (Dothideomycetes) by Boonmee et al.
(2016). The family Dictyosporiaceae mainly accommodates aquatic lignicolous species (Boonmee et al. 2016; Li
et al. 2017; Yang et al. 2018). The type genus Dictyosporium was introduced by Corda (1836), with D. elegans
Corda as the type species. The sexual morph genera in Dictyosporiaceae are characterized by immersed to erumpent,
superficial, brown to black ascomata, globose to subglobose with a rough surface, bitunicate asci with short ocular
chamber, and 1-septate, hyaline, sheathed ascospores (Boonmee et al. 2016; Atienza et al. 2021). The asexual morphs
are hyphomycetous with brown, dictyosporous and cheirosporous conidia, produced by sporodochial conidiomata on
micronematous conidiophores (Abdel-Aziz 2016; Boonmee et al. 2016; Wang et al. 2016).
The Digitodesmium was introduced by Kirk with type D. elegans P.M. Kirk (Kirk 1981), and the genus is
characterised by punctiform, sporodochial conidiomata, conidial secession schizolytic, acrogenous, euseptate,
cheiroid, digitate conidia, with an apical gelatinous cap (Kirk 1981; Nóbrega et al. 2021). Eight Digitodesmium species
epithets are listed in Index Fungorum (http://www.indexfungorum.org/; 18 June 2022). However, only three species,
D. chiangmaiense, D. bambusicola and D. polybrachiatum, have sequence data.
We are studying the diversity of fungi in southwestern China, Sichuan Province, along the Yangtze River, which has
enormous fungal diversity (Liang et al. 2003; Zhou et al. 2014; Yan et al. 2021; Zhou et al. 2021). This study aimed to
introduce a new genus Neodigitodesmium based on morphological and phylogenetic analyses. Paradictyocheirospora
was synonymized under Digitodesmium and provided an updated tree of Dictyosporiaceae based on analysis of the
ITS, LSU, SSU and tef1 sequence data.
A SPECIES OF NEODIGITODESMIUM Phytotaxa 559 (2) © 2022 Magnolia Press • 177
Materials and methods
Isolation and morphology
The sample was collected from Baiyungou in Sichuan Province, China, on September 27, 2021. The specimen was taken
to the laboratory in a paper envelope. The specimen was photographed using the stereoscope 0020Nikon C-FLED2
(Nikon, Tokyo, Japan). Colonies on the substrate were picked with a needle under the Nikon SMZ168-B (Nikon, Tokyo,
Japan) stereomicroscope and examined under the Nikon Eclipse E200 (Nikon, Tokyo, Japan) compound microscope.
The fungal microscopic structures were photographed using the DS-Fi3 camera (Nikon, Tokyo, Japan) installed at the
Nikon Eclipse Ni-U microscope (Nikon, Tokyo, Japan). Measurements were taken by using the software program NIS-
Elements D (Nikon, Tokyo, Japan). Photos were processed by using Adobe Photoshop version 22.0.
Single conidium isolation was made following the method described in Senanayake et al. (2020). Germinated
conidia were individually transferred to potato dextrose agar (PDA) media plates and incubated in the dark at 25
°C. Herbarium specimens were deposited at the Herbarium of Cryptogams, Kunming Institute of Botany Academia
Sciences (HKAS), Kunming, China, and the Herbarium of University of Electronic Science and Technology (HUEST),
Chengdu, China. The living cultures were deposited in the China General Microbiological Culture Collection Center
(CGMCC) in Beijing, China, and the University of Electronic Science and Technology Culture Collection (UESTCC)
in Chengdu, China.
DNA extraction, PCR amplification and sequencing
Genomic DNA was extracted from fungal mycelia using TreliefTM Plant Genomic DNA Kit (TSINGKE Biotech,
Shanghai, P.R. China) according to the manufacturer’s protocol. Four partial loci, the nuclear ribosomal internal
transcribed spacer (ITS), the nuclear ribosomal small subunit rRNA (SSU), the nuclear ribosomal large subunit rRNA
(LSU) and the translation elongation factor 1-alpha (tef1). The polymerase chain reaction (PCR) was carried out in a
25 μL reaction volume containing 12.5 μL PCR Master Mix (Sangon Biotech, Shanghai, P.R. China), 9.5 μL ddH2O, 1
μL of DNA and 1 μL of each primer. The primers used were ITS9mun/ITS4_KYO1 (Egger 1995; Toju et al. 2012) for
ITS, LR0R/LR5 (Vilgalys & Hester 1990) for LSU, PNS1/ NS41 (Hibbett 1996) for SSU and EF1-728F/EF1-2218R
(Carbone & Kohn 1999; Rehner & Buckley 2005) for tef1. The amplification condition for all four loci were consisted
of initial denaturation at 94 °C for 3 min; followed by 35 cycles of 30 s at 94 °C, 30 s at 56 °C, and 1 min at 72 °C,
and a final extension period of 10 min at 72 °C. The PCR products were analyzed by electrophoresis in 1 % agarose
gels. Sanger sequencing was conducted by Beijing Tsingke Biological Engineering Technology and Services Co. Ltd.
(Beijing, P.R. China).
Phylogenetic analyses
The analyzed dataset was consisted of combined LSU, ITS, SSU and tef1 sequence data. Sequence data of relevant
strains were downloaded from the NCBI nucleotide database using the function read. GenBank integrated in R package
ape (Paradis et al. 2004). Multiple sequence alignments were conducted using MAFFT (Katoh & Standley 2013)
version 7.310 with options “--adjustdirectionaccurately --auto”, and the alignment files were further trimmed using
trimAl (Capella et al. 2009) version 1.4 with the option “-gapthreshold 0.5”, which only allows 50% of taxa with a
gap in each site. The best-fit nucleotide substitution models for each locus were selected using PartitionFinder version
2.1.1 (Lanfear et al. 2016) under the Corrected Akaike Information Criterion (AICC). All sequence alignments were
combined using an in-house python script.
Maximum Likelihood (ML) and Bayesian analysis were conducted on individual and combined datasets. ML
phylogenetic trees were obtained using the IQ-TREE version 2.0.3 (Nguyen et al. 2015), and the topology was evaluated
using 1,000 ultrafast bootstrap replicates. The Bayesian analysis was conducted using parallel MrBayes version 3.2.7a
(Huelsenbeck & Ronquist 2000). Two different runs with 50 million generations and four chains were executed, and
the initial 25% of sample trees were treated as burn-in. Tracer v1.7.1 (Meireles et al. 2019) was used to confirm that the
MCMC runs reached convergence with all ESS values above 200. Then, the ML tree was annotated by TreeAnnotator
2.6.4 implemented in beast (Bouckaert et al. 2019) based on MrBayes. MCMC trees with no discard of burn-in and
no posterior probability limit. The tree was visualized using ggtree (Yu 2020) and edited in Adobe Illustrator version
16.0.0.
TIAN ET AL.
178 • Phytotaxa 559 (2) © 2022 Magnolia Press
TABLE 1. Species details and their GenBank accession numbers used in phylogenetic analyses. Type isolates are in bold,
and newly generated sequences are in red.
Species Voucher/Culture GenBank accession numbers
LSU ITS SSU tef1
Aquadictyospora clematidis MFLUCC 17-2080 MT214545 MT310592 MT226664 MT394727
A. lignicola MFLUCC 17-1318 MF948629 MF948621 - MF953164
Aquaticheirospora lignicola HKUCC 10304 AY736378 AY864770 AY736377 -
Cheirosporium triseriale HMAS 180703 EU413954 EU413953 - -
Dendryphiella stromaticola LAMIC 90/16 MK156678 MK829079 - -
D. trisepta COAD 2388 MK277357 MK278898 - -
D. variabilis CBS 584.96 LT963454 LT963453 - -
D. eucalyptorum CBS 137987 KJ869196 KJ869139 - -
D. fasciculata MFLUCC 17-1074 MF399214 MF399213 - -
D. paravinosa CBS 141286 KX228309 KX228257 - -
Dictyocheirospora aquatica KUMCC 15-0305 KY320513 KY320508 - -
D. bannica KH 332 AB807513 LC014543 - AB808489
D. garethjonesii MFLUCC 16-0909 KY320514 KY320509 - -
D. pseudomusae yone 234 AB807520 LC014550 AB797230 AB808496
D. rotunda MFLUCC 14-0293 KU179100 KU179099 KU179101 -
D. vinaya MFLUCC 14-0294 KU179103 KU179102 KU179104 -
Dictyosporium appendiculatum KUMCC 17-0311 MH376715 MH388343 - -
D. digitatum KUMCC 17-0269 MH376716 MH388344 MH388311 MH388378
D. guttulatum KUMCC 17-0288 MH376717 MH388345 MH388312 MH388379
D. hongkongensis KUMCC 17-0268 MH376718 MH388346 MH388313 MH388380
D. pandanicola MFLU 16-1886 MH376720 MH388347 - MH388382
Digitodesmium tectonae
(Paradictyocheirospora tectonae)NFCCI 4878 MW854647 MW854646 - MW854832
Digitodesmium sp. TBRC 10037 MK405232 MK405234 -MK405230
Digitodesmium sp. TBRC 10038 MK405233 MK405235 -MK405231
D. polybrachiatum COAD 3175 MW879317 MW879319 MW879326 -
D. polybrachiatum COAD 3174 MW879316 MW879318 MW879325 -
D. bambusicola CBS 110279 DQ018103 DQ018091 - -
D. chiangmaiense KUN-HKAS 102163 MK571766 - MK571775 -
Gregarithecium curvisporum KT 922 AB807547 AB809644 AB797257 AB808523
Immotthia bambusae KUNHKAS 112012 MW489450 MW489455 MW489461 MW504646
Jalapriya inflata NTOU 3855 JQ267363 JQ267362 JQ267361 -
J. pulchra MFLUCC 15-0348 KU179109 KU179108 KU179110 -
J. pulchra MFLUCC 17-1683 MF948636 MF948628 -MF953171
J. apicalivaginatum HKAS 115801 MZ621168 MZ621167 - -
J. aquaticum HKAS 115807 MZ621169 MZ621152 MZ621170 MZ851995
J. toruloides CBS 209.65 DQ018104 DQ018093 DQ018081 -
Neodendryphiella mali CBS 139.95 LT906657 LT906655 - -
N. mali FMR 17003 LT993735 LT993734 - -
N. michoacanensis FMR 16098 LT906658 LT906660 - -
N. tarraconensis FMR 16234 LT906656 LT906659 - -
Neodigitodesmium cheirosporum UESTCC 22.0020 ON595713 ON595714 ON595712 ON595700
Pseudocoleophoma calamagrostidis KT 3284 LC014609 LC014592 LC014604 LC014614
P. polygonicola KT 731 AB807546 AB809634 AB797256 AB808522
P. typhicola MFLUCC 16-0123 KX576656 KX576655 - -
P. bauhiniae MFLUCC 17-2586 MK347953 MK347736 MK347844 MK360076
P. clematidis MFLUCC 17-2177 MT214548 MT310595 MT226667 MT394730
......continued on the next page
A SPECIES OF NEODIGITODESMIUM Phytotaxa 559 (2) © 2022 Magnolia Press • 179
TABLE 1. (Continued)
Species Voucher/Culture GenBank accession numbers
LSU ITS SSU tef1
Pseudoconiothyrium broussonetiae CPC 33570 NG_066331 NR_163377 - -
Pseudodictyosporium. elegans CBS 688.93 DQ018106 DQ018099 - -
P. indicum CBS 471.95 -DQ018097 - -
P. thailandica MFLUCC 16-0029 KX259522 KX259520 - KX259526
P. wauense NBRC 30078 DQ018105 DQ018098 - -
P. wauense DLUCC 0801 MF948630 MF948622 -MF953165
Vikalpa australiensis HKUCC 8797 - DQ018092 - -
Verrucoccum coppinsii E 00814291 MT918769 MT918785 MT918777 -
V. hymeniicola SPO2343 MT918765 MT918780 MT918773 -
Periconia igniaria CBS 845.96 AB807567 LC014586 AB797277 AB808543
Results
Phylogenetic analyses
A combined dataset included four loci (LSU:1–835, ITS:836–1367, SSU:1368–2380, tef1:2381–3307) from 55 strains
of Dictyosporiaceae and Periconia igniaria (CBS 845.96) as the outgroup taxon. Newly generated sequences were
deposited in the GenBank, and the accession numbers were listed in TABLE 1. A total of 3304 characters, including
gaps were subjected to the phylogenetic analysis. Of these characters, 2407 were constant, 294 were variable and
parsimony-uninformative, and 603 were parsimony-informative. The best-fit evolution models GTR+I+G f for LSU,
ITS, SSU and tef1 were determined. The best-scoring ML consensus tree (lnL = -18399.635) with ultrafast bootstrap
values from ML analyses and posterior probabilities from MrBayes analysis at the node is shown in FIGURE 1.
Phylogenetic analysis based on combined LSU, ITS, SSU and tef1 loci (FIG. 1) confirmed that our isolate UESTCC
22.0020 could not assign to any known genera in Dictyosporiaceae. Therefore, introduced here it as Neodigitodesmium
cheirosporum gen. et sp. nov.
Taxonomy
Neodigitodesmium W. H. Tian & Maharachch. gen. nov.
MycoBank: MB 844432
Etymology: Named after its morphological similarity to Digitodesmium
Saprobic on submerged decayed wood in aquatic habitats. Sexual morph: Undetermined. Asexual morph:
Hyphomycetous. Conidiomata sporodochial on natural substrate, superficial, compact, dark brown to black, with base
attached on surface of substrate. Conidiophores semi-macronmatous, micronematous, reduced to conidiogenous cell.
Conidiogenous cells holoblastic, pale hyaline to brown at the base of the spore. Conidia solitary, dictyosporous, conical
shaped, cheiroid, not complanate, with a hyaline basal connecting cell, brown, subhyaline at the tip of peripheral
rows.
Type species: Neodigitodesmium cheirosporum W. H. Tian & Maharachch. sp. nov.
Neodigitodesmium cheirosporum W. H. Tian & Maharachch. sp. nov. (FIG. 2)
MycoBank: MB 844433
Etymology: The name refers to the cheiroid conidia
Saprobic on submerged decayed wood in aquatic habitats. Sexual morph: Undetermined. Asexual morph:
Hyphomycetous. Conidiomata sporodochial on natural substrate, superficial, compact, dark brown to black, with base
attached on the surface of substrate. Conidiophores micronematous, reduced to conidiogenous cell. Conidiogenous
cells 7–12 × 5–11 μm (x = 9 × 7 μm, n = 15), holoblastic, at the base of the spore, hyaline. Conidia 35–70 × 9–24 μm
TIAN ET AL.
180 • Phytotaxa 559 (2) © 2022 Magnolia Press
(x = 57 ×19 μm, n = 30), solitary, dictyosporous, conical shaped, cheiroid, not complanate, consisting of 1–4 rows with
each row composed of 6–10 cells, with a hyaline basal connecting cell, not easy to separate, brown, subhyaline at the
tip of peripheral rows.
Culture characteristics: Conidia germinating on PDA within 12 h at 25 °C in the dark. Colonies on PDA circular,
woolly at the margin, raised at the center, reverse yellow to brown center with a white margin.
Material examined: CHINA, Sichuan Province, Chengdu, Baiyungou, on submerged decayed wood, 103°24′19″
E, 30°47′52″ N, 27 September 2021, W. H. Tian BY112_3 (HKAS 124014, holotype), ex-type culture CGMCC 3.23623
= UESTCC 22.0020.
FIGURE 1. Phylogram of the best ML tree based on a combined dataset (LSU, ITS, SSU and tef1) of Dictyosporiaceae.
The ML ultrafast bootstrap values/Bayesian PP greater than 75%/0.95 are indicated at the respective nodes. Type
isolates are in bold, and the new taxa and synonymized taxa are indicated in red. The tree is rooted with Periconia
igniaria (CBS 845.96).
A SPECIES OF NEODIGITODESMIUM Phytotaxa 559 (2) © 2022 Magnolia Press • 181
FIGURE 2. Neodigitodesmium cheirosporum (HUEST 22.0020, holotype). a–c Colonies on the substrate d
Squash mount of sporodochium e, f, k Conidiogenous cells g–j Conidia l–n Crushed conidium o Germinating
conidium p, q Colonies on PDA after 30 days. Scale bars: d = 20 μm, e–o = 10 μm, Scale bars of e applies to e
and f, Scale bars of g applies to g–o.
TIAN ET AL.
182 • Phytotaxa 559 (2) © 2022 Magnolia Press
Notes: Combined sequences of LSU, ITS, SSU and tef1 confirmed the isolate of Neodigitodesmium formed a
distinct clade in Dictyosporiaceae. However, it could not assign to any known genera (FIG. 1). In the combined gene
tree, Neodigitodesmium is phylogenetically related to the Vikalpa australiensis (HKUCC 8797), Aquadictyospora
lignicola (MFLUCC 17-1318) and Aquadictyospora clematidis (MFLUCC 17-2080) (FIG. 1), but they are not
clustered on one branch and have obvious morphological differences. Neodigitodesmium is morphologically similar to
Digitodesmium, both having sporodochial conidiomata and acrogenous, cheiroid, digitate conidia (Kirk 1981; Nóbrega
et al. 2021). However, Neodigitodesmium differs from Digitodesmium and other Dictyosporiaceae members in conical-
shaped conidia, composed of 1–4 compactly arranged rows of dark brown cells, with a hyaline basal connecting
conidiogenous cells (FIG. 2). Therefore, Neodigitodesmium is described as a new genus based on phylogeny and
morphological evidence.
Digitodesmium tectonae (Rajeshkumar, R. K. Verma, Boonmee, K. D. Hyde, Chandrasiri & Wijayaw.) W. H. Tian &
Maharachch, comb. nov.
MycoBank: MB 844526
Basionym: Paradictyocheirospora tectonae Rajeshkumar, R. K. Verma, Boonmee, K. D. Hyde, Chandrasiri & Wijayaw, Phytotaxa 509
(3): 259–271 (2021).
Notes: The genus Paradictyocheirospora was introduced by Rajeshkumar et al. (2021), with P. tectonae as the type
species. The phylogenetic analysis based on a combined dataset of LSU, ITS, SSU and tef1 sequences show that P.
tectonae is clustered within Digitodesmium (FIG. 1). In addition, Paradictyocheirospora possesses key characteristics
of Digitodesmium such as punctiform, sporodochial conidiomata, acrogenous, cheiroid, digitate conidia, with an
apical gelatinous cap (Kirk 1981; Rajeshkumar et al. 2021; Nóbrega et al. 2021). Thus, Paradictyocheirospora
and Digitodesmium are congeneric. Digitodesmium is the older name and should take priority, and this selection is
recommended here.
Acknowledgments
This study was supported by the University of Electronic Science and Technology of China.
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