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Phytotaxa 446 (2): 095–102
https://www.mapress.com/j/pt/
Copyright © 2020 Magnolia Press Article PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
Accepted by Sinang Hongsanan: 19 May 2020; published: 27 May 2020
https://doi.org/10.11646/phytotaxa.446.2.2
95
Pseudodactylaria fusiformis sp. nov. from freshwater habitat in China
YONG-ZHONG LU1,5, JING-YI ZHANG1,2,6, CHUAN-GEN LIN2,7, ZONG-LONG LUO3,8 & JIAN-KUI (JACK)
LIU4,9*
1School of Food and Pharmaceutical Engineering, Guizhou Institute of Technology, Guiyang 550003, P.R. China.
2Center of Excellence in Fungal Research, Mae Fah Luang University, Chiang Rai 57100, Thailand.
3College of Agriculture and Biological Sciences, Dali University, Dali 671003, P.R. China.
4School of Life Science and Technology, University of Electronic Science and Technology of China, Chengdu 611731, P.R. China.
5
�
yzlu86@gmail.com; http://orcid.org/0000-0002-1033-5782
6
�
zjingyi127@gmail.com; https://orcid.org/0000-0003-0606-6169
7
�
chuangenlin@qq.com; https://orcid.org/0000-0003-2750-8720
8
�
514992672@qq.com; https://orcid.org/0000-0001-7307-4885
9
�
liujiankui@uestc.edu.cn; http://orcid.org/0000-0002-9232-228X
*Corresponding author
Abstract
Pseudodactylaria fusiformis sp. nov. was collected during an investigation of freshwater fungi along a north-south latitudinal
gradient in the Asian region. Evidence for the new species is provided by morphological comparison and sequence data
analysis. Pseudodactylaria fusiformis differs from other species in having hyaline conidiophores and fusiform, 0–1-septate
hyaline conidia without a sheath. Phylogenetic analysis based on combined ITS and LSU sequence data was carried out to
determine the phylogenetic placement of the species. Six Pseudodactylaria taxa clustered together and formed a monotypic
clade representing the genus, and five species are well recognized. Pseudodactylaria fusiformis and P. camporesiana share a
sister relationship and they are phylogenetically distinct species. A detailed description and illustration are provided, as well
as the comparisons with similar taxa.
Keywords: 1 new taxon, asexual morph, phylogeny, Sordariomycetes, taxonomy
Introduction
The genus Pseudodactylaria was established by Crous et al. (2017) based on the type species P. xanthorrhoeae Crous,
to accommodate two dactylaria-like species, namely as P. hyalotunicata (K.M. Tsui et al.) Crous and P. xanthorrhoeae.
Pseudodactylariaceae was introduced as a monotypic family by Crous et al. (2017) in Pseudodactylariales. In their study
(Crous et al. 2017), Pseudodactylariales formed a distinct clade and is phylogenetically close to Chaetosphaeriales and
Vermiculariopsiellales within the subclass Sordariomycetidae. Lin et al. (2018) confirmed the taxonomic status of
Pseudodactylariaceae based on phylogenetic analysis of LSU and ITS sequence data, and introduced a new species,
P. brevis C.G. Lin, McKenzie & K.D. Hyde, based on phylogenetic and morphological evidence. Hyde et al. (2020)
described P. camporesiana W. Dong, Doilom & K.D. Hyde from submerged wood in a stream in Thailand.
In this study, we describe the new species, Pseudodactylaria fusiformis, which is the fifth Pseudodactylaria
species, collected from submerged wood from a freshwater stream in China. The phylogenetic analysis based on
combined ITS and LSU sequence data indicates that Pseudodactylaria fusiformis is a distinct species in the genus. A
synopsis of accepted Pseudodactylaria species is provided.
Materials and Methods
Sample collection and specimen examination
Decaying wood specimens were collected from a freshwater stream in Guizhou Province, China. The samples were
processed following the methods reported in Luo et al. (2018). Morphological observations were made using a Motic
LU ET AL.
96 • Phytotaxa 446 (2) © 2020 Magnolia Press
SMZ 168 Series stereomicroscope and photographed using a Nikon E80i microscope-camera system. Measurements
were made with the Tarosoft (R) Image Frame Work (Liu et al. 2010). Figures were processed with Adobe Photoshop
CS6 Extended version 10.0 software (Adobe Systems, USA).
Single spore isolations were obtained using the method described by Liu et al. (2010). Germinating spores were
aseptically transferred to fresh potato dextrose agar (PDA) plates and incubated at 28 °C. Cultures were grown for
2 weeks and morphological characters, such as colour, colony shape, and texture were recorded. Type materials are
deposited in the Herbarium of Mae Fah Luang University (Herb. MFLU), Chiang Rai, Thailand and Herbarium of
Guizhou Academy of Agricultural Sciences (Herb. GZAAS), Guiyang, China. Ex-type living cultures are deposited at
Mae Fah Luang University Culture Collection (MFLUCC). Facesoffungi (Jayasiri et al. 2015) and Index Fungorum
numbers are provided.
DNA extraction, PCR amplification and sequencing
Genomic DNA was extracted from fungal mycelium grown on PDA at 28 °C for 2 weeks. Five gene regions were
amplified with universal primers: the internal transcribed spacer region of ribosomal DNA (ITS: ITS5/ITS4) (White
et al. 1990), the large subunit nuclear ribosomal DNA (LSU: LR0R/LR5) (Vilgalys & Hester 1990), the small subunit
nuclear ribosomal DNA (SSU: NS1/NS4) (White et al. 1990), the RNA polymerase II second largest subunit (RPB2:
fRPB2-5F/fRPB2-7cR) (Liu et al. 1999), and the translation elongation factor 1-alpha gene (TEF1α: EF1-983F/EF1-
2218R) (Rehner & Buckley 2005). The PCR products were purified and sequenced with the same primers. The ITS,
LSU, SSU, RPB2 and TEF1α amplification reactions were carried out using the method described by Lu et al. (2017).
The quality of PCR products was checked on 1% agarose gel electrophoresis stained with ethidium bromide. The PCR
products were sent for sequencing at Sangon Biotech, Shanghai, China.
Phylogenetic analysis
Original sequences were checked using BioEdit version 7.0.5.3 (Hall 1999), along with reference sequences originating
from previous publications (Crous et al. 2017; Lin et al. 2018; Hyde et al. 2020). The remaining homogenous sequences
were obtained by BLAST searches (Altschul et al. 1990) from GenBank. All sequences used in this study are listed
in TABLE 1. Alignments for each locus were done in MAFFT v7.307 online version (Katoh & Standley 2016) and
manually verified in MEGA 6.06 (Tamura et al. 2013). The interleaved NEXUS files for Bayesian inference analyses
were formatted with AliView v1.19-beta1k (Larsson 2014). Bayesian inference (BI), maximum parsimony (MP) and
maximum likelihood (ML) were used for phylogenetic analyses. For Bayesian inference analysis, the best model
of evolution was determined using MrModeltest v2 (Nylander 2004). Bayesian inference analysis was done with
MrBayes v 3.2.6 (Ronquist et al. 2012). Maximum parsimony analysis was performed in PAUP*4.0b10 (Swofford
2002). Maximum likelihood analysis was performed in raxmlGUI v 1.3.1 (Silvestro & Michalak 2012). Phylogenetic
trees were drawn with FigTree v1.4.3 (Rambaut 2012). The sequences are deposited in GenBank (TABLE 1). The
alignment was deposited in TreeBASE (http://www.treebase.org, submission number 25934).
Results
Phylogenetic analysis of combined ITS and LSU sequence data
The aligned sequence matrix comprises ITS (683 bp) and LSU (897 bp) sequence data for 25 taxa (ingroup) and two
outgroup taxa for a total of 1,580 characters after alignment including the gaps, of which 747 characters were constant,
327 variable characters were parsimony-uninformative and 506 characters were parsimony informative.
The dataset consists of eight orders within the subclass Sordariomycetidae. The tree was rooted with Arthrinium
arundinis (AFTOL-ID 951 and CBS 133509). Maximum parsimony analysis resulted in four trees with TL = 2113, CI
= 0.610, RI = 0.573, RC = 0.349, HI = 0.390. For the Bayesian analysis, two parallel runs with six chains were run for
1,000,000 generations and trees were sampled every 100th generation, resulted in 20002 trees from two runs of which
15002 trees were used to calculate the posterior probabilities (each run resulted in 10001 trees of which 7501 trees
were sampled). The MP, ML and BI analyses based on combined LSU and ITS sequence data provided similar tree
topologies, and the result of ML analysis (lnL = -10873.398596) is shown in FIGURE 1.
PSEUDODACTYLARIA FUSIFORMIS SP. NOV. Phytotaxa 446 (2) © 2020 Magnolia Press • 97
The newly obtained isolate of Pseudodactylaria fusiformis clustered together with P. camporesiana and these two
species are phylogenetically distinct (FIGURE 1).
TABLE 1 Taxa used in this study and their GenBank accession numbers for ITS and LSU DNA sequence data.
Taxa Strain/Voucher No. GenBank Accession No.
ITS LSU
Albertiniella polyporicola CBS 457.88 –aAF096185
Apiorhynchostoma curreyi UAMH 11088 NR_120207 JX460989
Arthrinium arundinis CBS 133509 KF144886 KF144930
Arthrinium arundinis AFTOL-ID 951 – DQ471018
Ascovaginospora stellipala P5-13A – U85088
Asteridiella obesa VIC 31239 NR_120256 JX096809
Camarops ustulinoides AFTOL-ID 72 – DQ470941
Cephalotheca foveolata UAMH 11631 KC408422 KC408398
Chaetosphaeria innumera SMH 2748 – AY017375
Coccodiella miconiae ppMP 1342 MF460365 KX430506
Coniochaeta ligniaria C8 – AY198388
Coniochaeta luteoviridis CBS 206.38 NR_154769 FR691987
Coniochaeta ostrea AFTOL-ID 915 – DQ470959
Cornipulvina ellipsoides SMH 1378 – DQ231441
Cryptendoxyla hypophloia CBS 508.70 – NG_058720
Endomeliola dingleyae PDD 98304 GU138865 GU138866
Gelasinospora tetrasperma CBS 178.33 NR_077163 DQ470980
Meliola centellae VIC 31244 NR_137799 JQ734545
Pseudodactylaria brevis MFLUCC 16–0032 MH262308 MH262310
Pseudodactylaria brevis MFLUCC 16–0034 MH262309 MH262311
Pseudodactylaria camporesiana MFLUCC 18–1410 MN796325 MN796326
Pseudodactylaria fusiformis MFLUCC 20–0085 MT184905 MT184906
Pseudodactylaria hyalotunicata HKUCC 2593 – EU107298
Pseudodactylaria xanthorrhoeae CBS 143414 MG386064 MG386117
Sordaria fimicola CBS 508.50 AY681188 AY681160
Sporoschisma hemipsila SMH 2125 – AY346292
Umbrinosphaeria caesariata CBS 102664 – AF261069
Notes: new sequences are in bold.
a No data in GenBank.
Taxonomy
Pseudodactylaria fusiformis Y.Z. Lu, J.Y. Zhang & Jian K. Liu, sp. nov. FIGURE 2
Index Fungorum: IF 557329; Facesoffungi number: FoF 07741
Etymology: ‘fusiformis’ referring to the fusiform conidia of this fungus.
Saprobic on submerged decaying wood in freshwater. Sexual morph: undetermined. Asexual morph:
hyphomycetous. Colonies on the substratum superficial, effuse, gregarious, white. Mycelium composed of partly
immersed, partly superficial, hyaline, septate, branched hyphae. Conidiophores macronematous, mononematous,
solitary, erect, unbranched, subcylindrical, straight to slightly flexuous, hyaline, 1–3-septate, 30–50 × 3.5–4.5 μm (
x
= 38 × 4 μm, n = 20). Conidiogenous cells holoblastic, polyblastic, integrated, terminal, sympodial, subcylindrical,
straight or flexuous, hyaline, 14–28 × 3.5–4.5 μm (
x
= 20 × 4 μm, n = 20), apical part forming a rachis with numerous,
aggregated, cylindrical denticles, 0.9–1.5 × 0.7–1.5 μm. Conidia solitary, acropleurogenous, smooth, prominently
LU ET AL.
98 • Phytotaxa 446 (2) © 2020 Magnolia Press
guttulate, 0–1-septate, fusiform, subtruncate at base, subobtuse at apex, hyaline, 20–25 × 3–4 μm (
x
= 22.5 × 3.5 μm,
n = 30).
FIGURE 1. Phylogenetic tree generated from ML analysis based on combined ITS and LSU sequence data for subclass Sordariomycetidae.
Bootstrap support values for maximum likelihood (ML, first set) and maximum parsimony (MP, second set) greater than 75 % and
Bayesian posterior probability greater than 0.95 (PP) are indicated above the branches. Hyphen (“-”) indicates a value lower than 75% for
ML and MP and a posterior probability lower than 0.95 for PP. The new isolate is in red and ex-type strains are in bold. The tree is rooted
with Arthrinium arundinis (AFTOL-ID 951 and CBS 133509) (Apiosporaceae).
Culture characteristics: Conidia germinating on water agar and producing germ tubes within 12 hours. Colonies
growing on PDA, circular, with flat surface, edge entire, reaching 12 mm in 10 days at 28 °C, greyish white.
Material examined: CHINA, Guizhou province, Zunyi city, Chishui county, Hushi town, Chishui Alsophila Natural
Reserve (28°29’43”N, 106°0’24”E), on submerged decaying bamboo culms in a freshwater stream, 22 September
2019, Yong-Zhong Lu, C1–1 (MFLU 20–0204, holotype; GZAAS 20–0095, isotype); ex-type living culture, MFLUCC
20–0085. Additional sequence: SSU: MT184897, RPB2: MT188555, TEF1α: MT188556.
Notes: Morphologically, Pseudodactylaria fusiformis is similar to P. hyalotunicata and P. xanthorrhoeae in having
unbranched hyaline conidiophores but can be distinguished by its conidia without a sheath (Tsui et al. 1997; Crous
et al. 2017). It is also similar to P. brevis and P. camporesiana in having hyaline, 1-septate fusiform conidia without
a sheath. However, Pseudodactylaria fusiformis can be distinguished from P. brevis by its larger conidia (20–25 μm
PSEUDODACTYLARIA FUSIFORMIS SP. NOV. Phytotaxa 446 (2) © 2020 Magnolia Press • 99
vs. 11.5–17.5 μm) and conidiophores (30–50 μm vs. 11–25 μm) (Lin et al. 2018), and from P. camporesiana by its
hyaline conidiophores while the later is brown at the base (Hyde et al. 2020). Phylogenetically, Pseudodactylaria
fusiformis shares a sister relationship to P. camporesiana with good bootstrap support (100 MLBS/100 MPBS/ 1.00
PP) (FIGURE 1), and the phylogenetic analysis showed that P. fusiformis is distinct from other Pseudodactylaria
species. Following the recommendations of Jeewon & Hyde (2016) for delimitation of new species, we delved into
pairwise dissimilarities of DNA sequences and noted that there are 36 nucleotide differences among 527 nucleotides in
the ribosomal ITS sequences (6.83% nucleotide differences), and 6 base pair differences in LSU (data did not contain
a gap) between Pseudodactylaria fusiformis and P. camporesiana, which confirmed our new taxon is a novel species.
FIGURE 2. Pseudodactylaria fusiformis (MFLU 20–0204, holotype). a Host material. b, c Colony on decaying wood. d–f Conidiophores.
g, h Conidiogenous cells with attached conidia. i Germinating conidium. j–m Conidia. n, o Colony on PDA from above and below. Scale
bars: b = 100 µm, c = 50 µm, d–i = 20 µm, j–m = 10 µm.
LU ET AL.
100 • Phytotaxa 446 (2) © 2020 Magnolia Press
Discussion
Pseudodactylariaceous species can be found from terrestrial and freshwater habitats (Tsui et al. 1997; Crous et al. 2017;
Lin et al. 2018; Hyde et al. 2020). The type genus Pseudodactylaria is characterized by erect, hyaline conidiophores,
integrated conidiogenous cells with a rachis and hyaline, fusiforme conidia, which is similar to Dactylaria sensu lato,
but can be distinguished by its 1-septate conidia covered by a mucoid sheath (Crous et al. 2017). Subsequently, Lin
et al. (2018) reported a new Pseudodactylaria species, P. brevis, whose conidia are 1-septate but lacking a mucoid
sheath. Recently, Hyde et al. (2020) reported the second aquatic Pseudodactylaria species, P. camporesiana, which
was collected from submerged wood in Thailand and its conidia are 1-septate and has no sheath.
In this study, we introduce the third aquatic Pseudodactylaria species, P. fusiformis, which is prominently similar
to P. camporesiana in conidial shape and size, but can be recognized by the colour of conidiophores (Hyde et al.
2020). In addition, Pseudodactylaria fusiformis shares similar conidiophores morphology with P. hyalotunicata and P.
xanthorrhoeae, but can be distinguished by its conidia without a sheath, while the later two have a sheath (Tsui et al.
1997; Crous et al. 2017), and the phylogenetic results (FIG 1) indicates that they are distinct species.
Crous et al. (2017) reported that Pseudodactylaria resembles species of Dactylaria (hyaline conidiophores and
septate, hyaline conidia formed on denticles), and it can be distinguished by having 1-septate conidia encased in
a mucoid sheath, which was absent in species of Dactylaria. However, the conidia of Pseudodactylaria brevis, P.
camporesiana and P. fusiformis are also lacking a sheath (shown in TABLE 2), which revealed that the conidia encased
in a sheath or not, is not the key character for identifying the genera Dactylaria and Pseudodactylaria.
TABLE 2 Synopsis of Pseudodactylaria species
Species Habitats/
Hosts Conidiophores Conidia References
Pseudodactylaria
brevis Terrestrial
Sometimes branched, septate, often
reduced to conidiogenous cells,
hyaline, 11–25 × 2–5 μm
No sheath, 1-septate, 11.5–17.5 ×
2.5–4.0 μm Lin et al. (2018)
Pseudodactylaria
camporesiana Freshwater
Unbranched, brown at the base,
hyaline at upper part, septate, 35–45
× 3.5–5 μm
No sheath, 1-septate, 18–22 ×
3.5–4.5 μm Hyde et al. (2020)
Pseudodactylaria
fusiformis Freshwater Unbranched, hyaline, 1–3-septate,
30–50 × 3.5–4.5 μm
No sheath, 0–1-septate, 20–25 ×
3–4 μm This study
Pseudodactylaria
hyalotunicata Freshwater Unbranched, hyaline, 4–6-septate,
30–60 × 4–5 μm
Surrounded by a gelatinous sheath,
0–1-septate, 20–25 × 2.5–3 μm
Tsui et al. (1997),
as Dactylaria
hyalotunicata
Pseudodactylaria
xanthorrhoeae Terrestrial Unbranched, hyaline, 1–3-septate,
1–3-septate, 20–50 × 4–5 μm
Surrounded by a thin mucilaginous
sheath, 1-septate, 20–33 × 3–4 μm Crous et al. (2017)
Acknowledgments
This work was funded by the National Natural Science Foundation of China (NSFC 31900020) and the Science and
Technology Foundation of Guizhou Province ([2020]1Y058).
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