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J. Fungi 2022, 8, 720. https://doi.org/10.3390/jof8070720 www.mdpi.com/journal/jof
Article
Morpho-Phylogenetic Evidence Reveals Novel Pleosporalean
Taxa from Sichuan Province, China
Xian-Dong Yu, Sheng-Nan Zhang and Jian-Kui Liu *
School of Life Science and Technology, Center for Informational Biology,
University of Electronic Science and Technology of China, Chengdu 611731, China;
yuxiandong@std.uestc.edu.cn (X.-D.Y.); zhangshengnan@uestc.edu.cn (S.-N.Z.)
* Correspondence: liujiankui@uestc.edu.cn; Tel.: +86-028-6183-1832
Abstract: Pleosporales is the largest and most morphologically diverse order in Dothideomycetes,
including a large proportion of saprobic fungi. During the investigation of microfungi from decay-
ing wood in Sichuan Province, several novel fungal taxa of asexual and sexual morphs were col-
lected, identified, and well-described. Phylogenetic analyses based on SSU, ITS, LSU, RPB2 and
TEF1α gene sequences suggested that these new taxa were related to Pleosporales and distributed
in five families, viz. Amorosiaceae, Bambusicolaceae, Lophiostomataceae, Occultibambusaceae and
Tetraplosphaeriaceae. The morphological comparison and molecular phylogeny evidence justify
the establishment of six new taxa, namely Bambusicola guttulata sp. nov., Flabellascoma sichuanense
sp. nov., Neoangustimassarina sichuanensis gen. et sp. nov., Occultibambusa sichuanensis sp. nov. and
Pseudotetraploa bambusicola sp. nov. Among them, Neoangustimassarina was introduced as the second
sexual morph genus in Amorosiaceae; Bambusicola guttulata, O. sichuanensis and P. bambusicola were
isolated from bamboos, which contributed to the diversity of bambusicolous fungi. The detailed,
illustrated descriptions and notes for each new taxon are provided, as well as a brief note for each
family. The potential richness of fungal diversity in Sichuan Province is also discussed.
Keywords: 6 new taxa; Dothideomycetes; multi-gene; phylogeny; taxonomy
1. Introduction
Pleosporales is the largest order in the class Dothideomycetes [1], and its members
are found worldwide on a variety of host plants as epiphytes, endophytes, saprobes and
parasites [2–4]. In addition, they are commonly found in terrestrial, marine and freshwa-
ter habitats [5–7]. Some of them produce secondary metabolites that can serve as a basis
for developing new antimicrobials, agrochemical pesticides and other useful compounds
[8].
The Pleosporales was invalidly introduced by Luttrell [9], and later revised by Barr
[10], based on the family Pleosporaceae with the type species Pleospora herbarum [11].
Members of Pleosporales usually have perithecioid ascomata, cellular pseudoparaphyses,
bitunicate and fissitunicate asci, and various shaped, aseptate or septate ascospores, with
or without a gelatinous sheath [12–14], and their asexual morphs are coelomycetes and
hyphomycetes [12,13]. For example, asexual morphs of Bambusicola and Pseudotetraploa
are the most common forms of coelomycetes and hyphomycetes in Pleosporales, respec-
tively. Zhang et al. listed 26 families in Pleosporales [12], while Hyde et al. revised Pleo-
sporales and accepted 41 families [13]. Hongsanan et al. redefined the families of Dothide-
omycetes and accepted 91 families in Pleosporales based on morphology and multigene
analysis [15]. Currently, Pleosporales consists of approximately 91 families and 655 gen-
era (including 41 genera incertae sedis) [16].
Fungi have a broad geographical distribution and diversity comparable to plants and
other organisms [17,18]. However, the fungal kingdom, in general, is less well-
Citation: Yu, X.-D.; Zhang, S.-N.;
Liu, J.-K. Morpho-Phylogenetic
Evidence Reveals Novel
Pleosporalean Taxa from Sichuan
Province, China. J. Fungi 2022, 8, 720.
https://doi.org/10.3390/jof8070720
Academic Editor: Philippe Silar
Received: 9 June 2022
Accepted: 7 July 2022
Published: 9 July 2022
Publisher’s Note: MDPI stays neu-
tral with regard to jurisdictional
claims in published maps and institu-
tional affiliations.
Copyright: © 2022 by the authors. Li-
censee MDPI, Basel, Switzerland.
This article is an open access article
distributed under the terms and con-
ditions of the Creative Commons At-
tribution (CC BY) license (http://crea-
tivecommons.org/licenses/by/4.0/).
J. Fungi 2022, 8 2
documented than the plant kingdom in terms of the number of species [19]. As one of the
biodiversity hotspots in China, Sichuan Province (located in southwestern China) has a
variety of complex topography (mountains, hills, plains, basins and plateaus) and climate
conditions, and these are important factors contributing to the biodiversity [20]. However,
little research on fungi has been carried out in this area; meanwhile, the highly variable
climate and lush vegetation are shown to have an important influence on fungal diversity.
Therefore, Sichuan Province is believed to have a large amount of hidden fungal diversity
to be explored and discovered [21,22].
During a survey of micro-fungi from decomposing wood in Sichuan Province, China,
a series of interesting asexual and sexual fungi were collected. In this study, we aim to
describe these new findings and contribute fungal diversity to China. The multi-gene phy-
logeny integrated with morphological comparison was carried out to determine the clas-
sification of these new collections. One new genus and five new species are introduced,
and the establishment of these new taxa is justified by morphology and phylogenetic ev-
idences.
2. Materials and Methods
2.1. Isolation and Morphological Examination
Fungi associated with decaying wood were collected from Sichuan Province, China
in 2021. Specimens were placed in envelopes and taken to the laboratory. Fungal colonies
and fruiting bodies were observed using Motic SMZ 168-B. Fungal structures were exam-
ined and photographed by using a Nikon ECLIPSE Ni-U compound microscope fitted
with a Nikon DS-Ri2 digital camera. The detailed morphological examination approaches
used in this paper were generally based on Senanayake et al. [23]. Single spore isolations
were made following the method in Senanayake et al. [23]. Measurements were made
with the Tarosoft (R) Image Framework program v. 0.9.7, following the procedures out-
lined by Liu et al. [24]. Photo plates representing fungal structures were processed in
Adobe Photoshop CS6 software (Adobe Systems Inc., San Jose, CA, USA). Herbarium
specimens (dry branches with fungal material) were deposited in the herbarium of Cryp-
togams, Kunming Institute of Botany Academia Sinica (HKAS), Kunming, China and the
herbarium of the University of Electronic Science and Technology (HUEST), Chengdu,
China. The isolates obtained in this study were deposited in China General Microbiologi-
cal Culture Collection Center (CGMCC), Beijing, China and the University of Electronic
Science and Technology Culture Collection (UESTCC), Chengdu, China. The names of the
new taxa were registered in MycoBank [25].
2.2. DNA Extraction, PCR Amplification and Sequencing
A Trelief TM Plant Genomic DNA Kit (Beijing TsingKe Biotech Co., Ltd., Beijing,
China) was used to extract total genomic DNA from fresh mycelia, according to the man-
ufacturer’s instructions. DNA amplification was performed by polymerase chain reaction
(PCR). SSU, ITS, LSU, RPB2 and TEF1α gene regions were amplified using the primer
pairs NS1/NS4, ITS5/ITS4, LR0R/LR5, fRPB2-5F/fRPB2-7cR and 983F/2218R, respectively,
[26–29]. The amplification reactions were performed in 25 μL PCR mixtures containing 22
μL PCR MasterMix (Green) (TsingKe Co., Beijing, China), 1 μL DNA template and 1 μL
of each primer (10 μM/L). The PCR thermal cycle program for SSU, ITS, LSU, RPB2 and
TEF1α amplification were listed in Table 1. PCR products were checked on 1% agarose
electrophoresis gels stained with Gel Red. The sequencing reactions were carried out with
primers, mentioned above, by Beijing Tsingke Biotechnology Co., Ltd., Chengdu, China.
J. Fungi 2022, 8 3
Table 1. PCR thermal cycles for SSU, ITS, LSU, RPB2 and TEF1α amplification.
Step SSU ITS, LSU, RPB2 TEF1α
Temperature Time Cycles Temperature Time Cycles Temperature Time Cycles
Initial Denaturation 98 °C 2 min 1 98 °C 2 min 1 98 °C 2 min 1
Denaturation 98 °C 10 s
35
98 °C 10 s
35
98 °C 10 s
35 Annealing 47 °C 10 s 56 °C 10 s 61.7°C 10 s
Extension 72 °C 10 s 72 °C 10 s 72 °C 10 s
Final Extension 72 °C 5 min 1 72 °C 5 min 1 72 °C 5 min 1
Hold 4 °C - - 4 °C - - 4 °C - -
2.3. Phylogenetic Analyses
The chromatograms of the new sequences obtained in this study were viewed in
Finch TV Version 1.4.0 (https://digitalworldbiology.com/FinchTV (accessed on 22 Septem-
ber 2021)). The BLAST searches were performed for finding similar sequences that match
our data. A concatenated dataset of the SSU, ITS, LSU, RPB2 and TEF1α sequences were
used for phylogenetic analyses with the inclusion of reference taxa from GenBank (Table
2). The sequences were aligned by using the online multiple-alignment program MAFFT
v.7 (http://mafft.cbrc.jp/alignment/server/(accessed on 5 January 2022)) [30], and the align-
ment was manually optimized in BioEdit v.7.0.9 [31]. Each gene dataset was concatenated
by Mesquite v. 3.11 (http://www.mesquiteproject.org/(accessed on 15 April 2022)) for
multi-gene phylogenetic analyses. Maximum likelihood (ML) and bayesian inference (BI)
were carried out as detailed in Dissanayake et al. [32]. The programs used in this study
are RAxMLGUI v. 1.0 [33], PAUP v.4.0b10 [34], Mr Modeltest 2.3 [35] and MrBayes v. 3.1.2
[36,37]. The phylogenetic tree was visualized by FigTree v.1.4.0
(http://tree.bio.ed.ac.uk/software/figtree/(accessed on 15 April 2022)).
Table 2. Taxa used in the phylogenetic analyses and their GenBank accession numbers. Newly generated sequences are
indicated with * and the ex-type strains are in bold.
Species Voucher/Strain/Isolate GenBank Accession Number
SSU ITS LSU RPB2 TEF1α
Alfoldia vorosii CBS 145501 MK589346 JN859336 MK589354 N/A MK599320
Alfoldia vorosii REF113 MK589345 JN859333 MK589353 N/A MK599319
Alfoldia vorosii REF117 MK589347 JN859337 MK589355 N/A MK599321
Amorocoelophoma camelliae NTUCC 18-097-1 MT071230 MT112303 MT071279 MT459143 MT743271
Amorocoelophoma camelliae NTUCC 18-097-2 MT071231 MT112304 MT071280 MT459141 MT743272
Amorocoelophoma camelliae NTUCC 18-097-3 MT071232 MT112305 MT071281 MT459142 MT743273
Amorocoelophoma cassiae MFLUCC 17-2283 MK347847 MK347739 MK347956 MK434894 MK360041
Amorocoelophoma neoregeliae CBS 146820 N/A MZ064410 MZ064467 MZ078193 MZ078247
Amorosia littoralis CBS 120399 AM292056 AM292047 AM292055 N/A N/A
Angustimassarina acerina MFLUCC 14-0505 KP899123 KP899132 KP888637 N/A KR075168
Angustimassarina arezzoensis MFLUCC 13-0578 KY501113 KY496743 KY496722 N/A KY514392
Angustimassarina camporesii MFLU 18-0057 MN244173 MN244197 MN244167 N/A N/A
Angustimassarina italica MFLUCC 15-0082 KY501124 KY496756 KY496736 N/A KY514400
Angustimassarina lonicerae MFLUCC 15-0087 N/A KY496759 KY496724 N/A N/A
Angustimassarina populi MFLUCC 13-0034 KP899128 KP899137 KP888642 N/A KR075164
Angustimassarina premilcurensis MFLUCC 15-0074 N/A KY496745 KY496725 KY514404 N/A
Angustimassarina quercicola MFLUCC 14-0506 KP899124 KP899133 KP888638 N/A KR075169
Angustimassarina rosarum MFLUCC 15-0080 N/A MG828869 MG828985 N/A N/A
Angustimassarina sylvatica MFLUCC 18-0550 MK314097 MK307843 MK307844 N/A MK360181
Angustimassarina alni MFLUCC 15-0184 KY548098 KY548099 KY548097 N/A N/A
Angustimassarina coryli MFLUCC 14-0981 N/A MF167431 MF167432 N/A MF167433
Aquatisphaeria thailandica MFLUCC 21-0025 MW890967 MW890969 MW890763 N/A N/A
Bambusicola aquatica MFLUCC 18-1031 MT864293 MT627729 MN913710 MT878462 MT954392
Bambusicola bambusae MFLUCC 11-0614 JX442039 JX442031 JX442035 KP761718 KP761722
Bambusicola didymospora MFLUCC 10-0557 KU872110 KU940116 KU863105 KU940163 KU940188
Bambusicola dimorpha MFLUCC 13-0282 KY038354 KY026582 KY000661 KY056663 N/A
Bambusicola ficuum MFLUCC 17-0872 MT215581 N/A MT215580 N/A MT199326
Bambusicola fusispora MFLUCC 20-0149 MW076529 MW076532 MW076531 MW034589 N/A
Bambusicola guttulata * CGMCC 3.20935 ON332919 ON332909 ON332927 ON383985 ON381177
J. Fungi 2022, 8 4
Bambusicola guttulata * UESTCC 22.0002 ON332920 ON332910 ON332928 ON383986 ON381178
Bambusicola irregulispora MFLUCC 11-0437 JX442040 JX442032 JX442036 KP761719 KP761723
Bambusicola loculata MFLUCC 13-0856 KP761735 KP761732 KP761729 KP761715 KP761724
Bambusicola massarinia MFLUCC 11-0389 JX442041 JX442033 JX442037 KP761716 KP761725
Bambusicola pustulata MFLUCC 15-0190 KU872112 KU940118 KU863107 KU940165 KU940190
Bambusicola sichuanensis SICAUCC 16-0002 MK253528 MK253473 MK253532 MK262830 MK262828
Bambusicola splendida MFLUCC 11-0439 JX442042 JX442034 JX442038 KP761717 KP761726
Bambusicola subthailandica SICAU 16-0005 MK253529 MK253474 MK253533 MK262831 MK262829
Bambusicola thailandica MFLUCC 11-0147 N/A KU940119 KU863108 KU940166 KU940191
Bambusicola triseptatispora MFLUCC 11-0166 N/A KU940120 KU863109 KU940167 N/A
Brunneofusispora clematidis MFLUCC 17-2070 MT226685 MT310615 MT214570 MT394692 MT394629
Brunneofusispora inclinatiostiola CGMCC 3.20403 MZ964884 MZ964866 MZ964875 OK061075 OK061069
Brunneofusispora sinensis KUMCC 17-0030 MH393556 MH393558 MH393557 N/A MH395329
Corylicola italica MFLU 19-0500 MT554923 MT554925 MT554926 MT590776 N/A
Corylicola italica MFLUCC 20-0111 MT633084 MT633085 MT626713 MT635596 MT590777
Crassiclypeus aquaticus CBS 143641 LC312470 LC312499 LC312528 LC312586 LC312557
Crassiclypeus aquaticus CBS 143643 LC312472 LC312501 LC312530 LC312588 LC312559
Ernakulamia krabiensis MFLUCC 18–0237 MK347880 MK347773 MK347990 N/A N/A
Ernakulamia tanakae NFCCI 4615 N/A MN937229 MN937211 N/A N/A
Ernakulamia xishuangbannaensis KUMCC 17-0187 MH260354 MH275080 MH260314 N/A N/A
Exosporium stylobatum CBS 160.30 N/A JQ044428 JQ044447 N/A N/A
Flabellascoma aquaticum KUMCC 15-0258 MN304832 MN304827 MN274564 MN328895 MN328898
Flabellascoma cycadicola CBS 143644 LC312473 LC312502 LC312531 LC312589 LC312560
Flabellascoma fusiforme MFLUCC 18-1584 N/A MN304830 MN274567 N/A MN328902
Flabellascoma minimum CBS 143645 LC312474 LC312503 LC312532 LC312590 LC312561
Flabellascoma minimum CBS 143646 LC312475 LC312504 LC312533 LC312591 LC312562
F
labellascoma sichuanense * CGMCC 3.20936 ON332921 ON332911 ON332929 ON383987 ON381179
Flabellascoma sichuanense * UESTCC 22.0003 ON332922 ON332912 ON332930 ON383988 ON381180
Hysterium rhizophorae NFCCI-4250 MG844280 MG844284 MG844276 MG968956 N/A
Lentistoma bipolare CBS 115375 LC312477 LC312506 LC312535 LC312593 LC312564
Lentistoma bipolare KT 3056 LC312484 LC312513 LC312542 LC312600 LC312571
Leptoparies palmarum CBS 143653 LC312485 LC312514 LC312543 LC312601 LC312572
Leucaenicola aseptata MFLUCC 17-2423 MK347853 MK347746 MK347963 MK434891 MK360059
Leucaenicola camelliae NTUCC 18-093-4 MT071229 MT112302 MT071278 MT743283 MT374091
Leucaenicola phraeana MFLUCC 18-0472 MK347892 MK347785 MK348003 MK434867 MK360060
Lophiomurispora hongheensis KUMCC 20-0216 MW264227 MW264218 MW264197 MW256810 MW256819
Lophiomurispora hongheensis KUMCC 20-0217 MW264225 MW264216 MW264195 MW256808 MW256817
Lophiostoma macrostomum KT 709/HHUF 27293 AB521732 AB433276 AB433274 JN993493 LC001753
Lophiostoma pseudodictyosporium MFLUCC 13-0451 N/A KR025858 KR025862 N/A N/A
Lophiostoma ravennicum MFLUCC 14-0005 KP698415 KP698413 KP698414 N/A N/A
Massarina corticola CBS 154.93 FJ795491 N/A FJ795448 FJ795465 N/A
Neoangustimassarina sichuanensis * CGMCC 3.20937 ON332917 ON332907 ON332925 ON383983 ON381175
Neoangustimassarina sichuanensis * UESTCC 22.0001 ON332918 ON332908 ON332926 ON383984 ON381176
Neooccultibambusa chiangraiensis MFLUCC 12-0559 KU712458 KU712442 KU764699 N/A KU872761
Neooccultibambusa kaiyangensis CGMCC 3.20404 MZ964886 MZ964868 MZ964877 OK061077 OK061071
Neooccultibambusa trachycarpi CGMCC 3.20405 MZ964888 MZ964870 MZ964879 OK061079 OK061073
Neothyrostroma encephalarti CBS 146037 N/A MN562104 MN567612 N/A MN556830
Neothyrostroma encephalarti CPC 35999 N/A MN562105 MN567613 N/A MN556831
Neovaginatispora clematidis MFLUCC 17–2156 MT226676 MT310606 MT214559 N/A MT394738
Neovaginatispora fuckelii KH 161 AB618689 LC001731 AB619008 N/A LC001749
Neovaginatispora fuckelii KT 634 AB618690 LC001732 AB619009 N/A LC001750
Occultibambusa aquatica MFLUCC 11-0006 KX698112 KX698114 KX698110 N/A N/A
Occultibambusa bambusae MFLUCC 13-0855 KU872116 KU940123 KU863112 KU940170 KU940193
Occultibambusa chiangraiensis MFLUCC 16-0380 KX655551 N/A KX655546 KX655566 KX655561
Occultibambusa fusispora MFLUCC 11-0127 N/A KU940125 KU863114 KU940172 KU940195
Occultibambusa hongheensis KUMCC 21-0020 MZ329029 MZ329037 MZ329033 N/A MZ325467
Occultibambusa jonesii GZCC 16-0117 KY628324 N/A KY628322 KY814758 KY814756
Occultibambusa kunmingensis HKAS 102151 MT864342 MT627716 MN913733 MT878453 MT954407
Occultibambusa maolanensis GZCC 16-0116 KY628325 N/A KY628323 KY814759 KY814757
Occultibambusa pustula MFLUCC 11-0502 KU872118 KU940126 KU863115 N/A N/A
Occultibambusa sichuanensis * CGMCC 3.20938 N/A ON332913 ON332931 ON383989 ON381181
Occultibambusa sichuanensis * UESTCC 22.0004 N/A ON332914 ON332932 ON383990 ON381182
Palmiascoma gregariascomum MFLUCC 11-0175 KP753958 KP744452 KP744495 KP998466 N/A
Palmiascoma qujingense KUMCC 19-0201 MT477186 MT477183 MT477185 MT495782 N/A
J. Fungi 2022, 8 5
Parapaucispora pseudoarmatispora KT 2237 LC100018 LC100021 LC100026 N/A LC100030
Paucispora quadrispora KT 843 AB618692 LC001734 AB619011 N/A LC001755
Paucispora versicolor KH 110 LC001721 AB918731 AB918732 N/A LC001760
Podocarpomyces knysnanus CBS 146076 N/A MN562155 MN567662 MN556816 MN556836
Polyplosphaeria fusca KT1616 AB524463 AB524789 AB524604 N/A N/A
Pseudotetraploa curviappendiculata JCM 12852 AB524467 AB524792 AB524608 N/A N/A
Pseudotetraploa javanica JCM 12854 AB524470 AB524795 AB524611 N/A N/A
Pseudotetraploa longissima JCM 12853 AB524471 AB524796 AB524612 N/A N/A
Pseudotetraploa rajmachiensis NFCCI 4618 N/A MN937222 MN937204 N/A N/A
Pseudotetraploa bambusicola * CGMCC 3.20939 ON332923 ON332915 ON332933 ON383991 ON381183
Pseudotetraploa bambusicola * UESTCC 22.0005 ON332924 ON332916 ON332934 ON383992 ON381184
Quadricrura bicornis CBS 125427 AB524472 AB524797 AB524613 N/A N/A
Quadricrura meridionalis CBS 125684 AB524473 AB524798 AB524614 N/A N/A
Seriascoma bambusae KUMCC 21-0021 MZ329031 MZ329039 MZ329035 MZ325470 MZ325468
Seriascoma didymosporum MFLUCC 11-0179 KU872119 KU940127 KU863116 KU940173 KU940196
Seriascoma yunnanense MFLU 19-0690 MN174694 N/A MN174695 MN210324 MN381858
Shrungabeeja aquatica MFLUCC 18-0664 N/A MT627722 MT627663 N/A N/A
Shrungabeeja longiappendiculata BCC 76463 KT376471 KT376474 KT376472 N/A N/A
Shrungabeeja vadirajensis MFLUCC 17-2362 N/A MT627681 MN913685 N/A N/A
Tetraploa aquatica MFLU 19-0995 N/A MT530448 MT530452 N/A N/A
Tetraploa aristata CBS 996.70 AB524486 AB524805 AB524627 N/A N/A
Tetraploa dwibahubeeja NFCCI 4621 N/A MN937226 MN937208 N/A N/A
Tetraploa nagasakiensis JCM 13168 AB524489 AB524806 AB524630 N/A N/A
Triplosphaeria acuta JCM 13171 AB524492 AB524809 AB524633 N/A N/A
Triplosphaeria maxima JCM 13172 AB524496 AB524812 AB524637 N/A N/A
Triplosphaeria yezoensis CBS 125436 AB524497 AB524813 AB524638 N/A N/A
Vaginatispora amygdali CBS 143662 LC312495 LC312524 LC312553 LC312611 LC312582
Vaginatispora appendiculata MFLUCC 16-0314 KU743219 KU743217 KU743218 N/A KU743220
Vaginatispora aquatica MFLUCC 11-0083 KJ591575 KJ591577 KJ591576 N/A N/A
Versicolorisporium triseptatum JCM 14775 AB524501 AB365596 AB330081 N/A N/A
Versicolorisporium triseptatum UESTCC 21.0016 = NMX1222 OL741381 OL741378 OL741318 N/A N/A
3. Results
3.1. Phylogenetic Analyses
Five gene loci SSU, ITS, LSU, RPB2 and TEF1α were used to determine the phyloge-
netic placement of the new collections. The concatenated matrix comprised 124 taxa with
a total of 4633 characters (SSU: 1021 bp; ITS: 684 bp; LSU: 904 bp; RPB2: 1031 bp; TEF1α:
993 bp) including gaps. Maximum likelihood (ML) and Bayesian inference (BI) analyses
were carried out to infer phylogenetic relationships. The best scoring ML tree (Figure 1)
was selected to represent the relationships among taxa, in which a final likelihood value
of –51844.747390 is presented. The matrix had 2418 distinct alignment patterns. Estimated
base frequencies were as follows: A = 0.245989, C = 0.250069, G = 0.270993, T = 0.232949;
substitution rates AC = 1.588007, AG = 3.646881, AT = 1.331328, CG = 1.142338, CT =
7.560715, GT = 1.000000. GTR + I + G is the best-fit model selected by AIC in MrModeltest
based on each gene (SSU, ITS, LSU, RPB2 and TEF1α), which was used for maximum like-
lihood and Bayesian analysis. Six simultaneous Markov chains were run for 1,970,000 gen-
erations and trees were sampled every 1000 generations and 1970 trees were obtained.
The first 394 trees representing the burn-in phase of the analyses were discarded, while
the remaining 1576 trees were used for calculating posterior probabilities in the majority
rule consensus tree (critical value for the topological convergence diagnostic is 0.01).
J. Fungi 2022, 8 6
J. Fungi 2022, 8 7
J. Fungi 2022, 8 8
Figure 1. RAxML tree generated from combined SSU, ITS, LSU, RPB2 and TEF1α sequence data of
targeted five families (Amorosiaceae, Bambusicolaceae, Lophiostomataceae, Occultibambusaceae,
and Tetraplosphaeriaceae) in Pleosporales. Bootstrap values for ML equal to or greater than 75% are
placed above the branches. Branches with Bayesian posterior probabilities (PP) from MCMC analy-
sis equal to or greater than 0.95 are in bold. The tree was rooted with Hysterium rhizophorae (NFCCI
4250). The ex-type strains are indicated in bold and newly generated sequences are indicated in red.
The newly obtained isolates were grouped with pleosporalean families of Amorosi-
aceae, Bambusicolaceae, Lophiostomataceae, Occultibambusaceae and Tetraplosphaeria-
ceae. Two isolates of Neoangustimassarina sichuanensis (CGMCC 3.20937 and UESTCC
22.0001) formed a distinct, well-supported clade (84% ML/1.00 BYPP) in Amorosiaceae.
Two isolates of Bambusicola guttulata (CGMCC 3.20935 and UESTCC 22.0002) were nested
in the genus Bambusicola in Bambusicolaceae. Two isolates of Flabellascoma sichuanense
(CGMCC 3.20936 and UESTCC 22.0003) clustered with Flabellascoma in Lophiosto-
mataceae. Two strains of Occultibambusa sichuanensis (CGMCC 3.20938 and UESTCC
22.0004) formed a distinct branch within Occultibambusaceae, which was closely related
to Occultibambusa hongheensis (KUMCC 21-0020), O. maolanensis (GZCC 16-0116), and Ver-
sicolorisporium triseptatum (JCM 14775 and NMX1222) with statistical support (100%
ML/1.00 BYPP). The other two strains of Pseudotetraploa bambusicola (CGMCC 3.20939 and
UESTCC 22.0005) grouped with Pseudotetraploa species in Tetraplosphaeriaceae.
3.2. Taxonomy
Pleosporales Luttr. ex M.E. Barr, Prodromus to class Loculoascomycetes: 67 (1987)
Amorosiaceae Thambug and K.D. Hyde, Fungal Diversity 74: 252 (2015)
Notes: Amorosiaceae was established by Thambugala et al. [38] and typified by Am-
orosia Mantle and D. Hawksw., which is characterized by micronematous to semi-mac-
ronematous conidiophores, integrated, terminal, or intercalary, monoblastic conidioge-
nous cells, elongate-clavate and 3–4-septate conidia [39]. Six genera were accepted in the
family, viz. Alfoldia D.G. Knapp, Imrefi and Kovács, Amorosia Mantle and D. Hawksw.,
Amorocoelophoma Jayasiri, E.B.G. Jones and K.D. Hyde, Angustimassarina Thambugala,
Kaz. Tanaka and K.D. Hyde, Neothyrostroma Crous and Podocarpomyces Crous [38–42]. An-
gustimassarina is the only genus in the family that represents the sexual morph [38].
Herein, we introduce the second genus with a sexual morph to Amorosiaceae.
Neoangustimassarina X.D. Yu and Jian K. Liu, gen. nov.
Type species: Neoangustimassarina sichuanensis X.D. Yu, S.N. Zhang and Jian K.
Liu
MycoBank: MB 843716
Etymology: The name refers to the similarity to the genus Angustimassarina.
Saprobic on dead wood in terrestrial habitat. Sexual morph: Ascomata solitary, scat-
tered, immersed, visible as pale brown, circular cap with a small central black dot, sub-
globose, uniloculate. Peridium composed of several layers of hyaline to brown cells of tex-
tura angularis. Hamathecium hyphae-like, pseudoparaphyses, embedded in a gelatinous
matrix. Asci 8-spored, bitunicate, fissitunicate, broad clavate to cylindric-clavate, short
pedicellate. Ascospores biseriate, fusiform with obtuse ends, hyaline, 1-septate, guttulate,
smooth-walled, surrounded by a mucilaginous sheath. Asexual morph: Undetermined.
Notes: The monotypic genus was introduced to accommodate Neoangustimassarina si-
chuanensis, which formed a distinct clade within Amorosiaceae (Figure 1). Neoangustimas-
sarina resembles Angustimassarina in forming globose to subglobose ascomata, hyaline,
and septate ascospores surrounded by mucilaginous sheaths [38]. However, Neoangusti-
massarina differs from the latter in having immersed ascomata without a pore opening,
broader asci (broad clavate to cylindric-clavate vs. cylindrical to cylindric-clavate), and
the septa of the ascospores (1-septate vs. 1–3-septate). We, hereby, introduce the new ge-
nus based on the distinctiveness of morphology and multi-gene phylogeny.
J. Fungi 2022, 8 9
Neoangustimassarina sichuanensis X.D. Yu, S.N. Zhang and Jian K. Liu, sp.
nov., Figure 2
MycoBank: MB 843717
Etymology: The epithet refers to Sichuan Province where the fungus was col-
lected.
Holotype: HKAS 123092
Figure 2. Neoangustimassarina sichuanensis (HKAS 123092, holotype) (a–c) Ascomata on host sub-
strate. (d–f) Vertical section of ascoma. (g–k) Asci. (l,m) Structure of peridium. (n) Pseudoparaphy-
ses. (o–s) Ascospores. (t) Germinated ascospore. (u,v) Colonies on PDA, above (u) and below (v).
Scale bars: (d–f) = 100 μm, (g–t) = 20 μm.
J. Fungi 2022, 8 10
Saprobic on dead wood in terrestrial habitat. Sexual morph: Ascomata solitary, scat-
tered, immersed, visible as circular, pale brown to nearly white flat cap, with a small black
dot in the center, in vertical section 135–235 μm high, 190–260 μm diam., subglobose, uni-
loculate, ostiolate. Peridium 10–23 μm wide, composed of several layers of hyaline to
brown cells of textura angularis. Hamathecium 1.9–2.9 μm wide, hyphae-like, pseudo-
paraphyses, embedded in a gelatinous matrix. Asci 78–125 × 20–30 μm ( = 92 × 25 μm, n
= 30), 8-spored, bitunicate, fissitunicate, broad clavate to cylindric-clavate, straight or
slightly curved, short pedicellate to subsessile, rounded at the apex, with an ocular cham-
ber. Ascospores 23–35 × 6.5–10.5 μm ( = 30 × 9 μm, n = 30), overlapping biseriate, fusiform
with obtuse ends, hyaline, 1-septate, constricted at the septum, the upper cell slightly
wider than the lower cell, guttulate when young, smooth-walled, surrounded by a wide
mucilaginous sheath. Asexual morph: Undetermined.
Culture characteristics: Colonies on PDA reaching 50–60 mm after 7 weeks at 25 °C,
circular, dry, the mycelium sparse at the margin, greyish-brown, reverse dark brown.
Material examined: CHINA, Sichuan Province, Chengdu City, Pengzhou County, Hui-
longgou Scenic Area, 31°14′21″ N, 103°47′28″ E, 1135 m Elevation, on dead wood, 28 July
2021, X.D. Yu, HLG3 (HKAS 123092, holotype); ex-holotype living culture CGMCC
3.20937; ibid., HUEST 22.0001, isotype, ex-isotype living culture UESTCC 22.0001.
Bambusicolaceae D.Q. Dai and K.D. Hyde, Fungal Diversity 63: 49 (2013)
Notes: Bambusicolaceae was established by Hyde et al. [13] to accommodate Bam-
busicola, D.Q. Dai and K.D. Hyde [43]. Four genera were accepted in this family, viz. Bam-
busicola [43], Corylicola [44], Leucaenicola [41] and Palmiascoma [45]. Most Bambusicola spe-
cies are parasites or saprobes and have been found on Bamboos (Poaceae) in terrestrial
habitats [43,46–50], except B. aquatica (from a freshwater habitat) and B. ficuum (on Ficus
sp., Moraceae) [51,52]. In this study, a coelomycetous Bambusicola species is introduced.
Bambusicola D.Q. Dai and K.D. Hyde, Cryptogamie Mycologie 33: 367 (2012)
Bambusicola guttulata X.D. Yu, S.N. Zhang and Jian K. Liu, sp. nov., Figure 3.
MycoBank: MB 843718
Etymology: Referring to the conidia with large guttules.
Holotype: HKAS 123091
Saprobic on dead branches of bamboo. Sexual morph: Undetermined. Asexual
morph: Conidiomata 100–170 μm high, 130–250 μm diam., dark brown to black, pycnidial,
usually forming in a linear series on the host surface, solitary, closed when young, becom-
ing erumpent, stromatic, irregular subglobose in section, immersed or semi-immersed,
unilocular, thick-walled. Conidiomatal wall 25–55 μm wide, composed of thick-walled, sub-
hyaline to brown cells of textura angularis. Conidiophores hyaline, cylindrical, branched,
straight or slightly flexuous, septate, and occasionally reduced to conidiogenous cells. Co-
nidiogenous cells 6–16 × 3–5 μm, holoblastic, hyaline, cylindrical to subcylindrical, determi-
nate, terminal, smooth-walled. Conidia 14–21 × 4–6 μm ( = 17 × 5 μm; n = 30), hyaline to
pale brown, unicellular when young, becoming 1-septate at maturity, cylindrical to sub-
cylindrical, sometimes with a narrow and truncate base, straight or slightly curved,
smooth-walled, guttulate.
Culture characteristics: Colonies on PDA reaching 30–40 mm after 7 weeks at 25 °C,
irregular, raised to umbonate, surface rough, dense, edge undulate, greyish-yellow, dry,
reverse dark brown.
Material examined: CHINA, Sichuan Province, Chengdu City, Tianfu New Area, Dalin
Village, 30°16′43″ N, 104°6′44″ E, 500 m Elevation, on dead branches of bamboo, 24 July
2021, X.D. Yu, B2 (HKAS 123091, holotype); ex-holotype living culture CGMCC 3.20935;
ibid., HUEST 22.0002, isotype, ex-isotype living culture UESTCC 22.0002.
Notes: Two isolates of Bambusicola guttulata formed a distinct lineage in Bambusicola
(Figure 1). Morphologically, B. guttulata is most similar to the asexual morph of the generic
type B. massarinia compared to the anamorphic species in the genus Bambusicola [43].
x
x
x
J. Fungi 2022, 8 11
However, B. guttulata has broader conidia than that of B. massarinia (14–21 × 4–6 μm vs.
14–20 × 2–3 μm). The establishment of the new species B. guttulata is justified by morpho-
logical and phylogenetic evidence.
Figure 3. Bambusicola guttulata (HKAS 123091, holotype) (a–c) Conidiomata on surface of dead
bamboo culms. (d–e) Vertical section of conidioma. (f) Wall of conidioma. (g–j) Conidiogenous cells
bearing conidia (the arrows indicated how the conidiogenous cells produce conidia). (k–o) Conidia.
(p) Germinating conidia. (q,r) Colonies on PDA, above and below. Scale bars: (d,e) = 50 μm, (f,g,p)
= 20 μm, (h–o) = 10 μm.
Lophiostomataceae Sacc., Sylloge Fungorum 2: 672 (1883)
J. Fungi 2022, 8 12
Notes: Nitschke [53] introduced “Lophiostomeae” based on the type species of
Lophiostoma macrostomum (Tode) Ces. and De Not. Saccardo formally established the fam-
ily Lophiostomataceae and placed “Lophiostomeae” in the order Pleosporales [54]. Mem-
bers of this family have crest-like ostioles in most cases and easily to be recognized. They
are characterized by immersed to erumpent ascomata, mostly clavate asci, hyaline to dark
brown ascospores with appendages or mucilaginous sheaths [38,55]. With the continuous
increase of new members, the family currently comprises 30 genera [16]. A new species
added to the genus Flabellascoma is identified and described.
Flabellascoma A. Hashim., K. Hiray. and Kaz. Tanaka, Studies in Mycology 90:
167 (2018)
Flabellascoma sichuanense X.D. Yu, S.N. Zhang and Jian K. Liu, sp. nov., Figure
4.
J. Fungi 2022, 8 13
Figure 4. Flabellascoma sichuanense (HKAS 123094, holotype) (a–c) Ascomata on host substrate.
(d) Vertical section of ascoma. (e) Ostiole, showing periphyses. (f) Structure of peridium. (g) Pseu-
doparaphyses. (h–k) Asci. (l–o) Ascospores. (p) Germinated ascospore. (q,r) Colonies on PDA,
above and below. Scale bars: (d,e) = 50 μm, (f,p) = 20 μm, (g–o) = 10 μm.
MycoBank: MB 843719
Etymology: The epithet refers to Sichuan Province where the fungus was col-
lected.
Holotype: HKAS 123094
Saprobic on dead branches of Eriobotrya sp. (Rosaceae). Sexual morph: Ascomata soli-
tary, scattered, rarely clustered, immersed to erumpent, visible as black, crest-like ostiolar
neck on the substrate, in vertical section 150–350 μm high, 190–280 μm diam., subglobose,
uniloculate. Ostiole central, laterally compressed, periphysate. Peridium 25–35 μm wide,
composed of several layers of brown, thick-walled cells of textura angularis. Hamathecium
1.5–3.5 μm wide, hyphae-like, pseudoparaphyses, embedded in a gelatinous matrix. Asci
55–75 × 9.5–13 μm ( = 64 × 11 μm, n = 30), 8-spored, bitunicate, fissitunicate, cylindric-
clavate, straight or slightly curved, shortly pedicellate, rounded at the apex, with an ocular
chamber. Ascospores 15–18 × 5–7 μm ( = 16.5 × 5.5 μm, n = 30), overlapping biseriate,
fusiform, hyaline, 1-septate, constricted at the septum, the upper cell slightly wider than
the lower cell, guttulate, smooth-walled, with a narrow bipolar sheath. Sheath 3.0–7.0 μm
long, 1.5–2.5 μm wide, drawn-out at both ends, with an internal chamber at both ends of
ascospores (Figure 4l). Asexual morph: Undetermined.
Culture characteristics: Colonies on PDA reaching 40–50 mm after 7 weeks at 25 °C,
circular, with dense mycelium on the surface, dark grayish of the inner ring, and brown
of the outer ring; in reverse black of the inner ring, and brown of the outer ring.
Material examined: CHINA, Sichuan Province, Chengdu City, Tianfu New Area, Dalin
Village, 30°16′43″ N, 104°6′44″ E, 500 m Elevation, on dead branches of Eriobotrya sp.
(Rosaceae), 24 July 2021, X.D. Yu, L4 (HKAS 123094, holotype); ex-holotype living culture
CGMCC 3.20936; ibid., HUEST 22.0003, isotype, ex-isotype living culture UESTCC 22.0003.
Notes: The phylogenetic result based on SSU, ITS, LSU, RPB2 and TEF1α sequence
data showed that the new collection Flabellascoma sichuanense nested in Flabellascoma (Fig-
ure 1) and formed a distinct lineage. Morphologically, it fits well with the genus Flabel-
lascoma in having immersed ascomata, bitunicate, fissitunicate, cylindrical, clavate asci
and fusiform, hyaline, 1-septate ascospores with a narrow bipolar sheath [56]. However,
the dimensions of asci and ascospores distinguish F. sichuanense from other species (Table
3).
Table 3. Morphological comparative data of Flabellascoma species.
Taxa Ascomata (μm) Hamathecium (μm) Asci (μm) Ascospores (μm) Sheath (μm) References
F. aquaticum 280–440 × 260–390 1.2–2.0 48.0–72.0 × 8–9 16–18 × 4.3–5.3 4.7–7.0 μm wide [57]
F. cycadicola 490–530 × 600–620 1.0–3.0 67.5–88.0 × 9–12 17–23 × 4.5–7.0 7.0–10 μm long [56]
F. fusiforme 310–420 × 320–380 1.5–3.0 66.0–80.0 × 10–12 15–18 × 4.0–5.0 5.4–8.0 μm wide [57]
F. minimumT 250–320 × 350–500 1.5–3.0 45.0–77.5 × 7.5–12 12–17.5 × 3.5–5 5.5–8.0 μm long [56]
F
. sichuanense 150–350 × 190–280 1.5–3.5 55.0–75.0 × 9.5–13 15–18 × 5.0–7.0 3.0–7.0 μm long,
1.5–2.5 μm wide This study
Occultibambusaceae D.Q. Dai and K.D. Hyde, Fungal Diversity 82: 25 (2017)
Notes: Species of Occultibambusaceae are mostly saprobic and frequently found on
monocotyledons or hardwood trees in terrestrial and aquatic habitats [47,58]. Dai et al. [47]
established this family to accommodate Neooccultibambusa, Occultibambusa, Seriascoma and
Versicolorisporium. Brunneofusispora, typified by Brunneofusispora sinensis, was subse-
quently introduced to this family by Phookamsak et al. [59]. Phylogenetically, the coelo-
mycetous genus Versicolorisporium appeared to be a close relationship with Occultibambusa
x
x
J. Fungi 2022, 8 14
in previous studies [21,52,60,61]. However, they continue to be treated as two distinct gen-
era because the known asexual morph of Occultibambusa is different from Versicolorispo-
rium [60]. In this study, a new Occultibambusa species is introduced.
Occultibambusa D.Q. Dai and K.D. Hyde, Fungal Diversity 82: 25 (2017)
Occultibambusa sichuanensis X.D. Yu, S.N. Zhang and Jian K. Liu, sp. nov., Fig-
ure 5.
Figure 5. Occultibambusa sichuanensis (HKAS 123093, holotype) (a–c) Ascomata on host substrate.
(d,e) Vertical section of ascoma. (f–j) Asci. (k) Structure of peridium. (l) Pseudoparaphyses. (s) As-
cospores. (t) Germinated ascospore. Colonies on PDA, above and below. Scale bars: (d,e) = 50 μm,
(f–k,m–t) = 20 μm, (l) = 10 μm.
MycoBank: MB 843720
J. Fungi 2022, 8 15
Etymology: The epithet refers to Sichuan Province where the fungus was col-
lected.
Holotype: HKAS 123093
Saprobic on dea d branc hes of B amboo. Sexual morph: Ascomata solitary to gregarious,
semi-immersed, visible as black domes on the substrate, in vertical section 130–180 μm
high, 340–440 μm diam., subglobose, coriaceous, uniloculate. Peridium 25–90 μm wide,
composed of several layers of brown, thick-walled cells of textura angularis. Hamathecium
2.8–3.3 μm wide, hyphae-like, cellular pseudoparaphyses, embedded in a gelatinous ma-
trix. Asci 70–100 × 22–27 μm ( = 85 × 24 μm, n = 30), 8-spored, bitunicate, fissitunicate,
obovoid to pyriform, straight or slightly curved, shortly pedicellate, rounded at the apex,
with an ocular chamber. Ascospores 31–41 × 6.5–10 μm ( = 36 × 8 μm, n = 30), 3-seriate,
fusiform, straight to somewhat curved, brown, 1-septate, constricted at the septum, gut-
tulate, smooth-walled, surrounded by a mucilaginous sheath. Asexual morph: Undeter-
mined.
Culture characteristics: Colonies on PDA reaching 40–50 mm after 7 weeks at 25 °C,
circular, with sparse mycelium on the surface, light gray of the inner ring, and brown of
the outer ring; in reverse olive green.
Material examined: CHINA, Sichuan Province, Chengdu City, Pengzhou County, Hui-
longgou Scenic Area, 31°14′21″ N, 103°47′28″ E, 1135 m Elevation, on dead branches of
bamboo in a terrestrial environment, 28 July 2021, X.D. Yu, HLG8 (HKAS 123093, holo-
type); ex-holotype living culture CGMCC 3.20938; ibid., HUEST 22.0004, isotype, ex-iso-
type living culture UESTCC 22.0004.
Notes: The blast search based on LSU sequence data of our new collection showed
that the closest hits were Versicolorisporium triseptatum (HHUF 28815 = JCM14775, identity
99.18%; NMX1222, identity 99.03%), and Occultibambusa bambusae (MFLUCC 11-0394,
identity 98.01%); the closest hits based on ITS sequence were Versicolorisporium triseptatum
(JCM 14775, identity 93.95%; NMX1222, identity 93.72%), and Occultibambusa hongheensis
(KUMCC 21-0020, identity 90.95%); the closest hits based on TEF1α sequence were Oc-
cultibambusa hongheensis (KUMCC 21-0020, identity 97.02%), and O. maolanensis (KUMCC
21-0020, identity 96.91%). Multi-gene phylogeny showed that the new collection grouped
with Occultibambusa and Versicolorisporium. It formed a sister clade with V. triseptatum with
high statistical support (100% ML/1.00 BYPP, Figure 1). However, the morphology of our
collection fits well with Occultibambusa. Further morphological evidence of its association
with Versicolorisporium is somewhat difficult due to the lack of asexual morph in our col-
lection. Therefore, we recognize our new collection as a new species of Occultibambusa,
namely, O. sichuanensis. The morphological comparison of Occultibambusa species was
listed in Table 4.
Table 4. Morphological comparative data of Occultibambusa.
Taxa Ascomata (μm) Asci Ascospores
Refer-
ences
Morphology Size (μm) Morphology Size (μm)
O. aquatica 100–250 × 180–280 Clavate 73–86 × 9–13 Narrowly fusiform, 1-septate, brownish,
with a sheath 19–25 × 3.5–6.5 [62]
O. bambusaeT 150–200 × 400–550 Broadly cylin-
drical
(50–)60–80(−90) × (9.5–
)11.5–14.5(−15)
Slightly broad and fusiform, 1-septate,
dark brown, with a sheath
(22–)23.5–27.5×
4.5–7 [47]
O. chiangraiensis 195–295 × 352–520 Clavate-oblong 47–92 × 12–16
Fusiform, (1–)3-septate, hyaline when
immature, pale brown to red-brown at
maturity, without a sheath
16–24 × 5–7 [62]
O. fusispora 135–185 × 240–275 Clavate to cy-
lindric-clavate
(60–)65–90(−110) × (11–
)12–14(−15)(−16)
Fusiform, mostly 1-septate, rarely 2–3-
septate, light brown, without a sheath
(20–)22–25(−26)
× 5–6(−6.5) [47]
O. hongheensis 180–340 × 400–550 Cylindric-cla-
vate to clavate
(78–)80–130(–137) ×
(18–)19–23(–25)
Fusiform, 1-septate, hyaline when young
and becoming pale brown when mature,
with a sheath
(25–)27–30 ×
(5.5–)8–9(–10) [60]
O. jonesii 196–236 × 200–260 Broadly cylin-
drical to clavate
(65–)75–89(–105) ×
13.5–19
Inequilateral-fusiform, 2-celled, hyaline
when young and becoming brown to
grayish when mature, without a sheath
27–33.5 × 5.5–
6.5 [63]
x
x
J. Fungi 2022, 8 16
O. kunmingensis 110–150 × 220–260 Clavate or cy-
lindric-clavate
110–140(–160) × 13–
16.5
Fusiform, 1-septate, brown, without a
sheath 32–40 × 5–6.5 [52]
O. maolanensis 544–600 diameter
Broadly cylin-
drical to clavate
(66–)77–85(–94) × 17–
20(–24)
Inequality-fusiform, 2-celled, hyaline
when young and become light brown
when mature, without a sheath
25–31 × 8–10 [63]
O. pustula 150–200 × 200–300 Cylindrical 80–105 × 8–12 Slightly broad-fusiform, 1-septate, hya-
line to pale brown, with a sheath 22–25 × 5–5.5 [47]
O. sichuanensis 130–180 × 340–440 Obovoid to
pyriform 70–100 × 22–27 Fusiform, 1-septate, brown, with a
sheath 31–41 × 6.5–10 This
study
Tetraplosphaeriaceae Kaz. Tanaka and K. Hiray, Studies in Mycology 64: 177
(2009)
Notes: Tetraplosphaeriaceae was introduced by Tanaka et al. [64], and typified by
Tetraplosphaeria. The latest taxonomic treatment of the family contains nine genera [1].
Pseudotetraploa is a genus with only known asexual forms, which were commonly associ-
ated with Poaceae (Dendrocalamus stocksii, Pleioblastus chino, Pleioblastus chino, Sasa kurilen-
sis) distributed in Japan or India [64,65]. In this study, a new Pseudotetraploa species asso-
ciated with bamboos from China is introduced.
Pseudotetraploa Kaz. Tanaka and K. Hiray, Studies in Mycology 64: 193 (2009)
Pseudotetraploa bambusicola X.D. Yu, S.N. Zhang and Jian K. Liu, sp. nov., Fig-
ure 6.
J. Fungi 2022, 8 17
Figure 6. Pseudotetraploa bambusicola (HKAS 123095, holotype) (a–c) Colonies on natural sub-
stratum. (d–l) Conidia. (m) Germinating conidium. (n) Colony on PDA from above and below. Scale
bars: (d–m) = 20 μm.
MycoBank: MB 843721
Etymology: Refers to the bamboo host.
Holotype: HKAS 123095
Saprobic on dead branches of Bamboo. Sexual morph: Undetermined. Asexual
morph: Colonies effuse, black. Mycelium superficial. Conidiophores absent. Conidiogenous
cells monoblastic, integrated, usually indistinguishable from superficial hyphae. Conidia
23–41 × 14–24 μm ( = 32 × 19 μm, n = 50), solitary, amygdaliform to ovoid, or obovoid,
dark brown to black, pseudoseptate, consisting of 3–4 columns, with 0–4 setose append-
ages. Appendages 8.85–95 × 2.5–4.5 μm ( = 36 × 3.5 μm, n = 50), 0–4-septate, dark brown,
smooth, unbranched, straight or curved.
Culture characteristics: Colonies on PDA reaching 40–50 mm after 7 weeks at 25 °C,
circular, dry, with dense mycelium, raised, entire at the edge, grayish brown, reverse dark
brown.
Material examined: CHINA, Sichuan Province, Chengdu City, Longquanyi District,
Longquan Mountain Scenic Area, 30°32′47″ N, 104°19′11″ E, 800 m Elevation, on dead
branches of bamboo in a terrestrial environment, 13 August 2021, X.D. Yu, THGL14
(HKAS 123095, holotype); ex-holotype living culture CGMCC 3.20939; ibid., HUEST
22.0005, isotype, ex-isotype living culture UESTCC 22.0005.
Notes: The phylogenetic result (Figure 1) showed that Pseudotetraploa bambusicola
formed a distinct lineage within Pseudotetraploa [64]. Morphologically, Pseudotetraploa
bambusicola resembles P. curviappendiculata, P. javanica, P. longissimi and P. rajma-
chiensis in having monoblastic conidiogenous cells. However, they can be distinguished
by the shape of conidia (obclavate to narrowly obpyriform in P. curviappendiculata and
P. longissimi; ovoid in P. javanica, ovoid to obclavate or obpyriform in P. rajmachiensis;
amygdaliform to ovoid, or obovoid in P. bambusicola) [64].
4. Discussion
The genus Flabellascoma was introduced by Hashimoto et al. [56] to accommodate two
terrestrial species F. cycadicola A. Hashim., K. Hiray and Kaz. Tanaka and F. minimum.
Subsequently, two species F. aquaticum D.F. Bao, Z.L. Luo, K.D. Hyde and H.Y. Su and F.
fusiforme D.F. Bao, Z.L. Luo, K.D. Hyde and H.Y. Su from freshwater habitats were intro-
duced based on multi-gene phylogeny [57]. Members of Flabellascoma have similar mor-
phological features [56,57] and it is difficult to distinguish Flabellascoma species by the size
and shape of asci and ascospores [57]. Bao et al. [57] proposed that the ascomatal features
appear to be remarkable features to distinguish taxa in this genus. Molecular data were
found to be more supportive for the identification of the new species in this study, and
we believed that DNA data provided more objective evidence for the species distinction
of Flabellascoma.
In previous studies, the relationship between Occultibambusa and Versicolorisporium
has not been well resolved due to the asexual morphs of Occultibambusa and Versicolor-
isporium being inconsistent [60]. In our phylogenetic tree, however, the genus Occultibam-
busa is not monophyletic (Figure 1), of which O. fusispora formed an independent lineage
in Occultibambusaceae; this is consistent with recent relevant studies [60,61]. Occultibam-
busa fusispora is the only species in the genus reported with its holomorph [47]; we cannot
solve the problem between Occultibambusa and Versicolorisporium due to the type-of-spe-
cies issue, although O. fusispora has an asexual morph. Therefore, further studies are
needed to provide sexual and asexual links of the type of species of O. bambusae and V.
triseptatum towards the classification of Occultibambusa and Versicolorisporium with more
sampling and taxa population included in the analysis.
x
x
J. Fungi 2022, 8 18
During the investigation of microfungi in Sichuan Province, we randomly sampled
three times in the vicinity of Chengdu city from July to August 2021. Morphological and
phylogenetic results showed that these newly collected interesting taxa were distributed
in five different pleosporalean families. It is worth noting that three new species found on
bamboo are typical bambusicolous fungi. Bamboo is a gramineous plant with economic
and ornamental value, and its culms and leaves are abundant in saprobic fungi [44,66–68].
China has the richest bamboo resources, with a total of 861 species distributes in 43 genera
[69]. Among them, Sichuan has a large area of bamboo forests, with an area of 592,800 ha,
ranking fifth in the country after Fujian, Jiangxi, Zhejiang and Hunan [69]. In recent years,
an increasing number of new species of bambusicolous fungi have been reported and dis-
covered in China [60,67,70,71]. Therefore, the unique natural conditions in Sichuan are of
great potential for the excavation and identification of bamboo fungi.
Author Contributions: Conceptualization, X.-D.Y., S.-N.Z. and J.-K.L.; methodology, X.-D.Y.; for-
mal analysis, X.-D.Y.; resources, X.-D.Y.; data curation, X.-D.Y. and S.-N.Z.; writing—original draft
preparation, X.-D.Y.; writing—review and editing, X.-D.Y., S.-N.Z. and J.-K.L.; supervision, J.-K.L.;
project administration, J.-K.L.; funding acquisition, J.-K.L. All authors have read and agreed to the
published version of the manuscript.
Funding: This study is supported by the Joint Fund of the National Natural Science Foundation of
China and the Karst Science Research Center of Guizhou province (Grant No. U1812401).
Institutional Review Board Statement: Not applicable.
Informed Consent Statement: Not applicable.
Data Availability Statement: The sequences data were submitted to GenBank.
Acknowledgments: Sajeewa Maharachchikumbura is thanked for his helps with sample collection.
Conflicts of Interest: The authors declare no conflict of interest.
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