Morphology and multigene phylogeny reveal a new order and a new species of wood-inhabiting basidiomycete fungi (Agaricomycetes)

Abstract

Dead wood-associated fungi play an important role in wood degradation and the recycling of organic matter in the forest ecological system. Xenasmataceae is a cosmopolitan group of wood-rotting fungi that grows on tropical, subtropical, temperate, and boreal vegetation. In this study, a new fungal order, Xenasmatales, is introduced based on both morphology and multigene phylogeny to accommodate Xenasmataceae. According to the internal transcribed spacer and nuclear large subunit (ITS+nLSU) and nLSU-only analyses of 13 orders, Xenasmatales formed a single lineage and then grouped with orders Atheliales, Boletales, and Hymenochaetales. The ITS dataset revealed that the new taxon Xenasmatella nigroidea clustered into Xenasmatella and was closely grouped with Xenasmatella vaga. In the present study, Xenasmatella nigroidea collected from Southern China is proposed as a new taxon, based on a combination of morphology and phylogeny. Additionally, a key to the Xenasmatella worldwide is provided.

Full text

TYPE Original Research
PUBLISHED 31 August 2022
DOI 10.3389/fmicb.2022.970731
OPEN ACCESS
EDITED BY
Samantha Chandranath Karunarathna,
Qujing Normal University, China
REVIEWED BY
Yusuon Gaorov,
Academy of Science of the Republic of
Uzbekistan, Uzbekistan
Kalani Hapuarachchi,
Guizhou University, China
*CORRESPONDENCE
Chang-Lin Zhao
fungi@swfu.edu.cn
SPECIALTY SECTION
This article was submitted to
Microbe and Virus Interactions with
Plants,
a section of the journal
Frontiers in Microbiology
RECEIVED 16 June 2022
ACCEPTED 08 August 2022
PUBLISHED 31 August 2022
CITATION
Luo K-Y and Zhao C-L (2022)
Morphology and multigene phylogeny
reveal a new order and a new species
of wood-inhabiting basidiomycete
fungi (Agaricomycetes).
Front. Microbiol. 13:970731.
doi: 10.3389/fmicb.2022.970731
COPYRIGHT
©2022 Luo and Zhao. This is an
open-access article distributed under
the terms of the Creative Commons
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distribution or reproduction in other
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or reproduction is permitted which
does not comply with these terms.
Morphology and multigene
phylogeny reveal a new order
and a new species of
wood-inhabiting basidiomycete
fungi (Agaricomycetes)
Kai-Yue Luo1,2,3 and Chang-Lin Zhao1,2,3,4*
1Key Laboratory for Forest Resources Conservation and Utilization in the Southwest Mountains of
China, Ministry of Education, Southwest Forestry University, Kunming, China, 2Yunnan Key
Laboratory of Plateau Wetland Conservation, Restoration and Ecological Services, Southwest
Forestry University, Kunming, China, 3College of Biodiversity Conservation, Southwest Forestry
University, Kunming, China, 4Yunnan Key Laboratory for Fungal Diversity and Green Development,
Kunming Institute of Botany, Chinese Academy of Science, Kunming, China
Dead wood-associated fungi play an important role in wood degradation
and the recycling of organic matter in the forest ecological system.
Xenasmataceae is a cosmopolitan group of wood-rotting fungi that grows
on tropical, subtropical, temperate, and boreal vegetation. In this study, a
new fungal order, Xenasmatales, is introduced based on both morphology
and multigene phylogeny to accommodate Xenasmataceae. According to
the internal transcribed spacer and nuclear large subunit (ITS+nLSU) and
nLSU-only analyses of 13 orders, Xenasmatales formed a single lineage and
then grouped with orders Atheliales, Boletales, and Hymenochaetales. The
ITS dataset revealed that the new taxon Xenasmatella nigroidea clustered into
Xenasmatella and was closely grouped with Xenasmatella vaga. In the present
study, Xenasmatella nigroidea collected from Southern China is proposed
as a new taxon, based on a combination of morphology and phylogeny.
Additionally, a key to the Xenasmatella worldwide is provided.
KEYWORDS
biodiversity, fungal systematics, ITS, LSU, new taxa, wood-decaying fungi,
Xenasmatales, Xenasmatella nigroidea
Introduction
Among eukaryotic microorganisms, wood-decaying fungi interact positively with
dead wood, playing a fundamental ecological role as decomposers of plants in the
fungal tree of life (James et al., 2020). Wood-associated fungi are cosmopolitan and rich
in diversity since they grow on tropical, subtropical, temperate, and boreal vegetation
(Gilbertson and Ryvarden, 1987;Núñez and Ryvarden, 2001;Bernicchia and Gorjón,
2010;Dai, 2012;Ryvarden and Melo, 2014;Dai et al., 2015,2021;Wu et al., 2020).
Xenasmataceae Oberw., a typical wood-associated fungal group mainly distributed
in the tropics was discovered by Oberwinkler (1966), and typified by Xenasma Donk.
Three genera, namely, Xenasma,Xenasmatella Oberw., and Xenosperma Oberw., have
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been accommodated in this family, however, higher-level
classification of the order has not been designated. The
tenth edition of the Dictionary of the Fungi showed that
Xenasmataceae belongs to Polyporales Gäum., and consists
of three genera (Kirk et al., 2008). MycoBank indicates that
Xenasmataceae has a higher classification within Polyporales,
although the Index Fungorum shows that Xenasmataceae
belongs to the order Russulales.
High phylogenetic diversity among corticioid
homobasidiomycetes suggests a close relationship among
Radulomyces M.P. Christ., Xenasmatella, and Coronicium J.
Erikss. and Ryvarden. Xenasma pseudotsugae (Burt) J. Erikss.
nested into the euagarics clade, in which it grouped with
Coronicium and Radulomyces. The three taxa of Radulomyces
grouped together with Phlebiella pseudotsugae (Burt) K.H.
Larss. and Hjortstam and Coronicium alboglaucum (Bourdot
and Galzin) Jülich, and were composed of a rather confusing
group with no obvious morphological features or ecological
specialization to tie these three genera together (Larsson et al.,
2004). The classification of corticioid fungi with 50 putative
families from published preliminary analyses and phylogenies
of sequence data showed that three species of Xenasmatella
formed a single lineage with strong support within the unplaced
Phlebiella family, in which this clade was unclaimed to any
orders (Larsson, 2007). A higher-level phylogenetic classification
of the Kingdom Fungi revealed that the Phlebiella clade and
Jaapia clade do not show affinities within any orders (Hibbett
et al., 2007). An outline of all genera of Basidiomycota with
combined SSU, ITS, LSU, tef1, rpb1, and rpb2 datasets showed
that Xenasmatella was assigned to Xenasmataceae within the
order Russulales (He et al., 2019). Therefore, there is debate on
the classification at the order level for the Xenasmataceae.
Recently, Xenasmatella has been studied deeply on the
basis of morphology and phylogeny. Phlebiella P. Karst. was
deemed to have not been legitimately published previously,
and the name Xenasmatella was accepted (Duhem, 2010;
Larsson et al., 2020;Maekawa, 2021). Molecular systematics
involving Xenasmatella was carried out recently. On the basis of
morphological and molecular identification, Zong et al. (2021)
studied the sequences of 27 fungal specimens representing 24
species between the Xenasmatella clade and related orders; and
the Xenasmatella clade formed a single lineage and three new
species, namely, X. rhizomorpha C.L. Zhao, X. tenuis C.L. Zhao,
and X. xinpingensis C.L. Zhao. Both the MycoBank database
(http://www.MycoBank.org) and Index Fungorum (http://www.
indexfungorum.org, accessed on June 20, 2022) have recorded
41 specific and infraspecific names in Xenasmatella. To date, the
number of Xenasmatella species accepted worldwide has reached
25 (Oberwinkler, 1966;Stalpers, 1996;Hjortstam and Ryvarden,
2005;Bernicchia and Gorjón, 2010;Duhem, 2010;Larsson et al.,
2020;Maekawa, 2021), of which, nine species have been found in
China (Dai et al., 2004;Dai, 2011;Huang et al., 2019;Zong and
Zhao, 2021;Zong et al., 2021).
In the present study, we verified the taxonomy and
phylogeny of Xenasmataceae. In addition, we analyzed the
species diversity of Xenasmataceae and constructed a phylogeny
to the order level of this family on the basis of large subunit
nuclear ribosomal RNA gene (nLSU) sequences, the internal
transcribed spacer (ITS) regions, and ITS+nLSU analyses. Based
on both morphology and phylogeny, we propose a new fungal
order, Xenasmatales and a new species, Xenasmatella nigroidea.
A key to the 25 accepted species of Xenasmatella worldwide is
also provided.
The accepted species list
Xenasma Donk (1957).
1. Xenasma Aculeatum C.E. Gómez (1972).
2. Xenasma Amylosporum Parmasto (1968).
3. Xenasma Longicystidiatum Boidin and Gilles (2000).
4. Xenasma Parvisporum Pouzar (1982).
5. Xenasma Praeteritum (H.S. Jacks.) Donk (1957).
6. Xenasma Pruinosum (Pat.) Donk (1957).
7. Xenasma Pulverulentum (H.S. Jacks.) Donk (1957).
8. Xenasma Rimicola (P. Karst.) Donk (1957).
9. Xenasma Subclematidis S.S. Rattan (1977).
10. Xenasma Tulasnelloideum (Höhn. and Litsch.) Donk
(1957).
11. Xenasma Vassilievae Parmasto (1965).
Xenasmatella Oberwinkler (1966).
1. Xenasmatella Ailaoshanensis C.L. Zhao ex C.L. Zhao and
T.K. Zong (2021).
2. Xenasmatella Alnicola (Bourdot and Galzin) K.H. Larss. and
Ryvarden (2020).
3. Xenasmatella Ardosiaca (Bourdot and Galzin)
Stalpers (1996).
4. Xenasmatella Athelioidea (N. Maek.) N. Maek. (2021).
5. Xenasmatella Bicornis (Boidin and Gilles) Piatek (2005).
6. Xenasmatella Borealis (K.H. Larss. and Hjortstam)
Duhem (2010).
7. Xenasmatella Caricis-Pendulae (P. Roberts) Duhem (2010).
8. Xenasmatella Christiansenii (Parmasto) Stalpers (1996).
9. Xenasmatella Cinnamomea (Burds. and Nakasone)
Stalpers (1996).
10. Xenasmatella Fibrillosa (Hallenb.) Stalpers (1996).
11. Xenasmatella Globigera (Hjortstam and Ryvarden)
Duhem (2010).
12. Xenasmatella Gossypina (C.L. Zhao) G. Gruhn and
Trichies (2021).
13. Xenasmatella Inopinata (H.S. Jacks.) Hjortstam and
Ryvarden (1979).
14. Xenasmatella Insperata (H.S. Jacks.) Jülich (1979).
15. Xenasmatella Nasti Boidin and Gilles ex Stalpers (1996).
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TABLE 1 The list of species, specimens, and GenBank accession numbers of sequences used in this study.
Species Name Specimen No. GenBank Accession No. References
ITS nLSU
Albatrellus confluens PV 10193 – AF506393 Larsson et al., 2004
Aleurobotrys botryosus CBS 336.66 MH858812 MH870451 Vu et al., 2019
Amaurodon viridis TAA 149664 AY463374 AY586625 Larsson et al., 2004
Amphinema byssoides EL 1198 – AY586626 Larsson et al., 2004
Amylostereum areolatum NH 8041 – AF506405 Larsson and Larsson, 2003
Aphanobasidium pseudotsugae NH 10396 – AY586696 Larsson et al., 2004
Auriscalpium vulgare EL 3395 – AF506375 Larsson and Larsson, 2003
Athelia epiphylla EL 1298 AY463382 AY586633 Larsson et al., 2004
Athelopsis subinconspicua KHL 8490 AY463383 AY586634 Larsson et al., 2004
Bondarzewia dickinsii Li 150909/19 KX263721 KX263723 Unpublished
Candelabrochaete septocystidia AS 95 – EU118609 Larsson, 2007
Chaetodermella luna NH 8482 EU118615 – Larsson, 2007
C. luna CBS 305.65 – MH870216 Vu et al., 2019
Chondrostereum purpureum EL 5997 – AY586644 Larsson et al., 2004
Clavulicium delectabile KHL 11147 – AY586688 Larsson et al., 2004
Clavulina cristata EL 9597 AY463398 AY586648 Larsson et al., 2004
Columnocystis abietina KHL 12474 EU118619 – Larsson, 2007
Coronicium alboglaucum NH 4208 – AY586650 Larsson et al., 2004
Cystostereum murrayi KHL 12496 EU118623 – Larsson, 2007
Dacrymyces stillatus CBS 195.48 MH856306 MH867857 Vu et al., 2019
Dacryopinax spathularia Miettinen 20559 MW191976 MW159092 Unpublished
Erythricium laetum NH 14530 AY463407 AY586655 Larsson et al., 2004
Exidia recisa SL Lindberg 180317 – MT664783 Unpublished
Exidiopsis calcea KHL 11075 – AY586654 Larsson et al., 2004
Gloeocystidiellum porosum FCUG 1933 – AF310094 Larsson and Hallenberg, 2001
Haplotrichum conspersum KHL 11063 AY463409 AY586657 Larsson et al., 2004
Hydnocristella himantia KUC 20131001-35 – KJ668382 Unpublished
Hydnomerulius pinastri 412 – AF352044 Jarosch and Besl, 2001
Hydnum repandum 420526MF0827 – MG712372 Unpublished
Hygrophoropsis aurantiaca EL 4299 – AY586659 Larsson et al., 2004
Hymenochaete cinnamomea EL 699 AY463416 AY586664 Larsson et al., 2004
Hyphodermella corrugate KHL 3663 – EU118630 Larsson, 2007
Hyphodontia aspera KHL 8530 AY463427 AY586675 Larsson et al., 2004
Inonotus radiatus TW 704 – AF311018 Wagner and Fischer, 2001
Junghuhnia nitida CBS 45950 – MH868226 Vu et al., 2019
Kavinia alboviridis EL 1698 – AY463434 Larsson et al., 2004
Kavinia himantia LL 98 AY463435 AY586682 Larsson et al., 2004
Lactarius volemus KHL 8267 – AF506414 Larsson and Larsson, 2003
Laetisaria fuciformis CBS 18249 – MH868023 Vu et al., 2019
Lentaria dendroidea SJ 98012 EU118640 EU118641 Larsson, 2007
Lignosus hainanensis Dai 10670 NR154112 GU580886 Cui et al., 2011
Merulicium fusisporum Hjm s.n. EU118647 – Larsson, 2007
Mycoaciella bispora EL 1399 – AY586692 Larsson et al., 2004
Peniophora pini Hjm 18143 – EU118651 Larsson, 2007
(Continued)
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TABLE 1 (Continued)
Species Name Specimen No. GenBank Accession No. References
ITS nLSU
Phanerochaete sordida KHL 12054 – EU118653 Larsson, 2007
Phellinus chrysoloma TN 4008 – AF311026 Wagner and Fischer, 2001
Phlebia nitidula Nystroem 020830 – EU118655 Larsson, 2007
Podoscypha multizonata CBS 66384 – MH873501 Vu et al., 2019
Polyporus tubiformis WD 1839 AB587634 AB368101 Sotome et al., 2011
Porpomyces mucidus KHL 11062 AF347091 – Unpublished
P. mucidus Dai 10726 – KT157839 Wu et al., 2015
Pseudomerulius aureus BN 99 – AY586701 Larsson et al., 2004
Punctularia strigosozonata LR 40885 AY463456 AY586702 Larsson et al., 2004
Rickenella fibula AD 86033 – AY586710 Larsson et al., 2004
Russula violacea SJ 93009 AF506465 AF506465 Larsson and Larsson, 2003
Scopuloides hydnoides WEI 17569 – MZ637283 Chen et al., 2021
Sistotrema alboluteum TAA 167982 AY463467 AY586713 Larsson et al., 2004
Sistotremastrum niveocremeum MAFungi 12915 – JX310442 Telleria et al., 2013
Sistotremastrum suecicum KHL 11849 – EU118667 Larsson, 2007
Sphaerobasidium minutum KHL 11714 – DQ873653 Larsson et al., 2006
Stereum hirsutum NH 7960 AF506479 – Larsson and Larsson, 2003
Tomentellopsis echinospora KHL 8459 AY463472 AY586718 Larsson et al., 2004
Trametes suaveolens CBS 279.28 MH855012 MH866480 Vu et al., 2019
Trechispora farinacea KHL 8793 AF347089 – Larsson et al., 2004
T. farinacea MAFungi 79474 – JX392856 Telleria et al., 2013
Tubulicrinis subulatus KHL 11079 AY463478 AY586722 Larsson et al., 2004
Veluticeps abietina HHB 13663 – KJ141191 Unpublished
Veluticeps berkeleyi HHB 8594 – HM536081 Garcia-Sandoval et al., 2010
Vuilleminia comedens EL 199 AY463482 AY586725 Larsson et al., 2004
Wrightoporia lenta KN 150311 – AF506489 Larsson and Larsson, 2003
Xerocomus chrysenteron EL 3999 AF347103 – Larsson et al., 2004
Xenasma praeteritum ACD 0185 OM009268 Unpublished
Xenasma pruinosum OTU 1299 MT594801 Unpublished
Xenasma rimicola NLB 1571 MT571671 Unpublished
X. rimicola NLB 1449 MT537020 Unpublished
Xenasmatella ailaoshanensis CLZhao 3895 MN487105 – Huang et al., 2019
X. ailaoshanensis CLZhao 4839 MN487106 – Huang et al., 2019
Xenasmatella ardosiaca CBS 126045 MH864060 MH875515 Vu et al., 2019
Xenasmatella borealis UC 2022974 KP814210 – Rosenthal et al., 2017
X. borealis UC 2023132 KP814274 – Rosenthal et al., 2017
Xenasmatella christiansenii TASM YGG 26 MT526341 – Gafforov et al., 2020
X. christiansenii TASM YGG 36 MT526342 – Gafforov et al., 2020
Xenasmatella gossypina CLZhao 4149 MW545958 – Zong and Zhao, 2021
X. gossypina CLZhao 8233 MW545957 – Zong and Zhao, 2021
Xenasmatella nigroidea CLZhao 18300 OK045679 OK045677 Present study
X. nigroidea CLZhao 18333 * OK045680 OK045678 Present study
Xenasmatella rhizomorpha CLZhao 9156 MT832954 – Zong et al., 2021
X. rhizomorpha CLZhao 9847 MT832953 – Zong et al., 2021
Xenasmatella tenuis CLZhao 4528 MT832960 – Zong et al., 2021
(Continued)
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TABLE 1 (Continued)
Species Name Specimen No. GenBank Accession No. References
ITS nLSU
X. tenuis CLZhao 11258 MT832959 – Zong et al., 2021
Xenasmatella vaga KHL 11065 EU118660 EU118661 Larsson, 2007
X. vaga BHI-F 160a MF161185 – Haelewaters et al., 2018
Xenasmatella wuliangshanensis CLZhao 4080 MW545962 – Zong and Zhao, 2021
X. wuliangshanensis CLZhao 4308 MW545963 – Zong and Zhao, 2021
Xenasmatella xinpingensis CLZhao 2216 MT832961 – Zong et al., 2021
X. xinpingensis CLZhao 2467 MT832962 – Zong et al., 2021
*Indicates type materials.
FIGURE 1
A maximum parsimony strict consensus tree illustrating the phylogeny of the new order Xenasmatales and related order in the class
Agaricomycetes based on ITS+nLSU sequences. The orders represented by each color are indicated in the upper left of the phylogenetic tree.
Branches are labeled with a maximum likelihood bootstrap value ≥70%, and a parsimony bootstrap value ≥50, respectively.
16. Xenasmatella Odontioidea Ryvarden and Liberta (1978).
17. Xenasmatella Palmicola (Hjortstam and Ryvarden)
Duhem (2010).
18. Xenasmatella Rhizomorpha C.L. Zhao (2021).
19. Xenasmatella Romellii Hjortstam (1983).
20. Xenasmatella Sanguinescens Svrˇ
cek (1973).
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21. Xenasmatella Subflavidogrisea (Litsch.) Oberw. ex
Jülich (1979).
22. Xenasmatella Tenuis C.L. Zhao (2021).
23. Xenasmatella Vaga (Fr.) Stalpers (1996).
24. Xenasmatella Wuliangshanensis (C.L. Zhao) G. Gruhn and
Trichies (2021).
25. Xenasmatella Xinpingensis C.L. Zhao (2021).
Xenosperma Oberw. (1966).
1. Xenosperma Hexagonosporum Boidin and Gilles (1989).
2. Xenosperma Ludibundum (D.P. Rogers and Liberta) Oberw.
ex Jülich (1979).
3. Xenosperma Murrillii Gilb. and M. Blackw. (1987).
4. Xenosperma Pravum Boidin and Gilles (1989).
Materials and methods
Sample collection and herbarium
specimen preparation
Fresh fruit bodies of fungi growing on the stumps of
angiosperms were collected from Honghe, Yunnan Province,
P.R. China. The samples were photographed in situ, and
macroscopic details were recorded. Field photographs were
taken by a Jianeng 80D camera. All photographs were focus
stacked and merged using Helicon Focus software. Once
the macroscopic details were recorded, the specimens were
transported to a field station where they were dried on an
electronic food dryer at 45◦C. Once dried, the specimens were
FIGURE 2
A maximum parsimony strict consensus tree illustrating the phylogeny of the new order Xenasmatales and related order in the class
Agaricomycetes based on nLSU sequences. The orders represented by each color are indicated in the upper left of the phylogenetic tree.
Branches are labeled with a maximum likelihood bootstrap value ≥70%, a parsimony bootstrap value ≥50%, and Bayesian posterior
probabilities ≥0.95, respectively.
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labeled and sealed in envelopes and plastic bags. The dried
specimens were deposited in the herbarium of the Southwest
Forestry University (SWFC), Kunming, Yunnan Province,
P.R. China.
Morphology
The macromorphological descriptions were based on field
notes and photos captured in the field and laboratory. The
color, texture, taste, and odor of fruit bodies were mostly based
on the authors’ field trip investigations. Rayner (1970) and
Petersen (1996) were used for the color terms. All materials
were examined under a Nikon 80i microscope. Drawings were
made with the aid of a drawing tube. The measurements
and drawings were made from slide preparations stained with
cotton blue (0.1 mg aniline blue dissolved in 60 g pure lactic
acid), melzer’s reagent (1.5 g potassium iodide, 0.5 g crystalline
iodine, 22 g chloral hydrate, and aq. dest. 20 ml), and 5%
potassium hydroxide. Spores were measured from the sections
of the tubes; and when presenting spore size data, 5% of the
measurements excluded from each end of the range are shown
in parentheses (Wu et al., 2022). The following abbreviations
were used: KOH =5% potassium hydroxide water solution,
CB =cotton clue, CB– =acyanophilous, IKI =Melzer’s
reagent, IKI– =both inamyloid and indextrinoid, L =means
spore length (arithmetic average for all spores), W =means
spore width (arithmetic average for all spores), Q =variation
in the L/W ratios between the specimens studied, and n =
a/b (number of spores (a) measured from given number (b)
of specimens).
Molecular phylogeny
The CTAB rapid plant genome extraction kit-DN14 (Aidlab
Biotechnologies Co., Ltd., Beijing, P.R. China) was used to
obtain genomic DNA from the dried specimens following
the manufacturer’s instructions (Zhao and Wu, 2017). The
nuclear ribosomal ITS region was amplified with the primers
ITS5 and ITS4 (White et al., 1990). The nuclear nLSU region
was amplified with the primer pairs LR0R and LR7 (http://
lutzonilab.org/nuclear-ribosomal-dna/, accessed on September
12, 2021). The PCR procedure used for ITS was as follows:
initial denaturation at 95◦C for 3 min, followed by 35 cycles
at 94◦C for 40 s, 58◦C for 45 s, and 72◦C for 1 min, and
a final extension of 72◦C for 10 min. The PCR procedure
used for nLSU was as follows: initial denaturation at 94◦C
for 1 min, followed by 35 cycles at 94◦C for 30 s, 48◦C
for 1 min, and 72◦C for 1.5 min, and a final extension
of 72◦C for 10 min. The PCR products were purified and
sequenced at Kunming Tsingke Biological Technology Limited
Company (Yunnan Province, P.R. China). All the newly
generated sequences were deposited in the National Center
FIGURE 3
A maximum parsimony strict consensus tree illustrating the phylogeny of a new species and related species in Xenasmatella and Xenasma based
on ITS sequences. Branches are labeled with a maximum likelihood bootstrap value ≥70%, a parsimony bootstrap value ≥50%, and Bayesian
posterior probabilities ≥0.95, respectively. The new species are in bold.
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Luo and Zhao 10.3389/fmicb.2022.970731
for Biotechnology Information (NCBI) GenBank (https://www.
ncbi.nlm.nih.gov/genbank/, accessed on September 12, 2021)
(Table 1).
The sequences and alignment were adjusted manually using
AliView version 1.27 (Larsson, 2014). The datasets were aligned
with Mesquite version 3.51. The ITS+nLSU dataset and the
nLSU-only sequence dataset were used to position a new order,
Xenasmatales, and the ITS-only dataset was used to position
a new species among the Xenasmatella-related taxa. Sequences
of Dacrymyces stillatus and Dacryopinax spathularia retrieved
from GenBank were used as the outgroup for the ITS+nLSU
sequences (Figure 1) (He et al., 2019); sequences of Exidia recisa
and Exidiopsis calcea retrieved from GenBank were used as the
outgroup for the nLSU sequences (Figure 2) (Larsson, 2007); and
the sequence of Trametes suaveolens was used as the outgroup
for the ITS-only sequences (Figure 3) (Zong and Zhao, 2021).
The three combined datasets were analyzed using maximum
parsimony (MP), maximum likelihood (ML), and Bayesian
inference (BI), according to Zhao and Wu (2017), and the tree
was constructed using PAUP∗version 4.0b10 (Swofford, 2002).
All characters were equally weighted and gaps were treated as
missing data. Trees were inferred using the heuristic search
option with TBR branch swapping and 1,000 random sequence
FIGURE 4
Basidiomata of Xenasmatella nigroidea (holotype). Bars: (A)
1 cm; (B) 1 mm.
additions. Max-trees were set to 5,000, branches of zero length
were collapsed, and all parsimonious trees were saved. Clade
robustness was assessed using the bootstrap (BT) analysis with
1,000 replicates (Felsenstein, 1985). Descriptive tree statistics—
tree length (TL), consistency index (CI), retention index (RI),
rescaled consistency index (RC), and homoplasy index (HI)—
were calculated for each maximum parsimonious tree generated.
In addition, multiple sequence alignment was analyzed using
ML in RAxML-HPC2 through the Cipres Science Gateway
(Miller et al., 2012). Branch support (BS) for ML analysis was
determined by 1,000 bootstrap replicates.
MrModeltest 2.3 (Nylander, 2004) was used to determine
the best-fit evolution model for each dataset of BI, which was
performed using MrBayes 3.2.7a with a GTR+I+G model of
DNA substitution and a gamma distribution rate variation
across sites (Ronquist et al., 2012). A total of 4 Markov chains
were run for 2 runs from random starting trees for 1 million
generations for the ITS+nLSU dataset (Figure 1), 1.4 million
generations for the nLSU-only sequences (Figure 2), and 0.5
million generations for the ITS-only sequences (Figure 3), with
trees and parameters sampled every 1,000 generations. The
first one-fourth of all generations was discarded as a burn-in.
The majority rule consensus tree of all remaining trees was
calculated. Branches were considered significantly supported if
FIGURE 5
Microscopic structures of Xenasmatella nigroidea (drawn from
the holotype). (A) Basidiospores. (B) Basidia and basidioles. (C) A
section of hymenium. Bars: (A) 5µm; (B,C) 10 µm.
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Luo and Zhao 10.3389/fmicb.2022.970731
they received a maximum likelihood bootstrap value (BS) ≥70%,
a maximum parsimony bootstrap value (BT) ≥70%, or Bayesian
posterior probabilities (BPP) ≥0.95.
Results
Phylogenetic analyses
The ITS+nLSU dataset (Figure 1) included sequences from
45 fungal specimens representing 45 species. The dataset had an
aligned length of 3,095 characters, of which 1,910 characters are
constant, 353 are variable and parsimony uninformative, and
832 are parsimony informative. Maximum parsimony analysis
yielded 45 equally parsimonious trees (TL =3,984, CI =0.4666,
HI =0.5334, RI =0.3909, and RC =0.1824). The best model was
GTR+I+G [lset nst =6, rates =invgamma; prset statefreqpr =
dirichlet (1,1,1,1)]. Bayesian and ML analyses showed a topology
similar to that of MP analysis with split frequencies equal to
0.009126 (BI), and the effective sample size (ESS) across the two
runs is double that of the average ESS (avg ESS) =250.5.
The ITS+nLSU rDNA gene regions (Figure 1) were
based on 13 orders, namely, Agaricales Underw., Atheliales
Jülich, Boletales E.J. Gilbert, Cantharellales Gäum.,
Corticiales K.H. Larss., Gloeophyllales Thorn, Gomphales
Jülich, Hymenochaetales Oberw., Polyporales, Russulales,
Thelephorales Corner ex Oberw., Trechisporales, and
Xenasmatales, while Xenasmatella was separated from the
other orders.
The nLSU-alone dataset (Figure 2) included sequences from
58 fungal specimens representing 58 species. The dataset had an
aligned length of 1,343 characters, of which 726 characters are
constant, 176 are variable and parsimony-uninformative, and
441 are parsimony-informative. Maximum parsimony analysis
yielded 3 equally parsimonious trees (TL =2,864, CI =0.3209,
HI =0.6791, RI =0.4476, and RC =0.1436). The best model
for the ITS dataset estimated and applied in the Bayesian
analysis was GTR+I+G [lset nst =6, rates =invgamma; prset
statefreqpr =dirichlet (1,1,1,1)]. The Bayesian and ML analyses
TABLE 2 Morphological characteristics of the relevant orders used in this study.
Order Name Morphological characteristics References
Agaricales Hymenophore type gilled, poroid, ridged, veined, spinose, papillate, and smooth; spore deposit color white,
pink, brown, purple-brown and black
Fries, 1821–1832,1828,
1857–1863,1874
Atheliales Generally corticioid and athelioid, producing effused, crust like fruiting bodies that are loosely attached to the
substrate and with non-differentiated margins
Eriksson et al., 1978,
1981,1984
Boletales Includes conspicuous stipitate-pileate forms that mainly have tubular and sometimes lamellate hymenophores
or intermediates that show transitions between the two types of hymenophores. Also includes gasteromycetes
(puffball-like forms), resupinate or crust-like fungi that produce smooth, merulioid (wrinkled to warted), or
hydnoid (toothed) hymenophores, and a single polypore-like species, Bondarcevomyces taxi
Gilbert, 1931;Besl and
Bresinsky, 1997;Jarosch,
2001;Larsson et al., 2004
Corticiales Basidiomata resupinata, effuso-reflexa vel discoidea; hymenophora laevia; systema hypharum monomiticum;
dendrohyphidia raro absentia; basidia saepe e probasidiis oriuntur. Cystidia presentia vel absentia. Sporae
hyalinae, tenuitunicatae, albae vel aggregatae roseae.
Hibbett et al., 2007
Gloeophyllales Basidiomata annua vel perennia, resupinata, effuso-reflexa, dimidiata vel pileata; hymenophora laevia,
merulioidea, odontioidea vel poroidea. Systema hypharum monomiticum, dimiticum vel trimiticum. Hyphae
generativae fibulatae vel efibulatae. Leptocystidia ex trama in hymenium projecta, hyalina vel brunnea,
tenuitunicata vel crassitunicata. Basidiosporae laeves, hyalinae, tenuitunicatae, ellipsoideae vel cylindricae vel
allantoideae, inamyloideae. Lignum decompositum brunneum vel album.
Hibbett et al., 2007
Gomphales Basidiomata can be coralloid, unipileate or merismatoid (having a pileus divided into many smaller pilei); the
pileus, if present, can be fan- to funnel-shaped
Gonzalez-Avila et al.,
2017
Hymenochaetales Hymenial structure (corticioid, hydnoid or poroid) and basidiocarps (resupinate, pileate or stipitate); the main
characters are the xanthochroic reaction, the lack of clamps, the frequent occurrence of setae
Tobias and Michael,
2002
Thelephorales Basidiospores tuberosae spinosaeque plus minusve coloratae Oberwinkler, 1975
Trechisporales Basidiomata resupinata, stipitata vel clavarioidea. Hymenophora laevia, grandinioidea, hydnoidea vel poroidea.
Systema hypharum monomiticum vel dimiticum. Hyphae fibulatae, septa hypharum interdum inflata
(ampullata). Cystidia praesentia vel absentia. Basidia 4-6 sterigmata formantia. Sporae laeves vel ornatae.
Species lignicolae vel terricolae.
Hibbett et al., 2007
Xenasmatales Basidiomata resupinate. Hyphal system monomitic, generative hyphae with clamp connections. Basidia pleural.
Basidiospores colorless.
Present study
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Luo and Zhao 10.3389/fmicb.2022.970731
resulted in a topology similar to that of MP analysis with split
frequencies equal to 0.009830 (BI), and the effective sample size
(ESS) across the two runs is double that of the average ESS (avg
ESS) =402.
The nLSU regions (Figure 2) were based on 13 orders,
namely, Agaricales, Atheliales, Boletales, Cantharellales,
Corticiales, Gloeophyllales, Gomphales, Hymenochaetales,
Polyporales, Russulales, Thelephorales, Trechisporales, and
Xenasmatales, while Xenasmatella was separated from the
other orders.
The ITS-alone dataset (Figure 3) included sequences from
26 fungal specimens representing 15 species belonging to
Xenasma and Xenasmatella. The dataset had an aligned length
of 598 characters, of which 267 characters are constant, 74 are
variable and parsimony-uninformative, and 257 are parsimony-
informative. Maximum parsimony analysis yielded 1 equally
parsimonious tree (TL =629, CI =0.7329, HI =0.2671, RI =
0.8301, and RC =0.6084). The best model for the ITS dataset
estimated and applied in the Bayesian analysis was GTR+I+G
[lset nst =6, rates =invgamma; prset statefreqpr =dirichlet
(1,1,1,1)]. The Bayesian and ML analyses resulted in a topology
similar to MP analysis with split frequencies equal to 0.007632
(BI), and the effective sample size (ESS) across the two runs is
double that of the average ESS (avg ESS) =300.5.
In the ITS sequence analysis (Figure 3), a previously
undescribed species was grouped into Xenasmatella with a sister
group to X. vaga (Fr.) Stalpers.
Taxonomy
Xenasmatales K.Y. Luo and C.L. Zhao, ord. nov.
MycoBank no.: MB 842882
Type family: Xenasmataceae Oberw.
Basidiomata resupinate. Hyphal systems are monomitic,
generative hyphae with clamp connections. Basidia pleural.
Basidiospores are colorless.
Xenasmataceae Oberw., Sydowia 19(1–6): 25 (1966).
MycoBank no.: MB 81527
Type genus: Xenasma Donk
Basidiomata resupinate, ceraceous to geletinous.
Hyphal systems are monomitic, generative hyphae
with clamp connections. Basidia pleural usually with 4
sterigmata and a basal clamp connection. Basidiospores
are colorless.
TABLE 3 Morphological characteristic comparison of Xenasmatella nigroidea and other species.
Species name Basidiomata Hymenial surface Basidia Basidiospores References
Xenasmatella nigroidea Thin, very hard to
separate from
substrate
Smooth, byssaceous to reticulate
under the lens
12–18 ×4.5–6 µm Ellipsoid, 3.5–4.5 ×2.5–3.5 µm;
asperulate with blunt spines up
to 0.2 µm long
Present study
X. christiansenii Fragile Smooth, pruinose to farinaceous or
more or less reticulate
6–7 ×4–4.5 µm Ellipsoid, 6–7 ×4–4.5 µm;
asperulate with blunt spines up
to 1 µm long
Bernicchia and
Gorjón, 2010
X. fibrillosa Thin, fragile Porulose to reticulate or formed by
radially arranged, white to pale
yellowish white
12–15 ×4–5 µm Ellipsoid, 4.5–5.5 ×3–3.5 µmBernicchia and
Gorjón, 2010
X. gaspesica Small spots and
becoming a closed
coating, firmly
attached
Resh smooth and somewhat
gelatinous, light gray, dry waxy,
white gray
7–11 ×4–4.5 µm Ellipsoid, 8–10 ×2–2.5 µmGrosse-Brauckmann
and Kummer, 2004
X. gossypina Cotton to flocculent Cream to buff 14–23.5 ×4–7 µm Subglobose to globose, 3.3–4.4
×2.8–4 µm
Zong and Zhao, 2021
X. odontioidea Colliculosa Ceraceo-membranacea 17.5–20 ×4.5–5 µm Ovale-ellipsoid, 2.5–3.5 µmRyvarden and Liberta,
1978
X. rhizomorpha Presence of the
rhizomorph
Clay-buff to cinnamon 10.5–17.5 ×3.5–6.5 µm Ellipsoid, 3.1–4.9 ×2.3–3.3 µmZong et al., 2021
X. subflavidogrisea Thin White to grayish 10–12 ×4–5 µm Ellipsoid, 3.5–4.5 ×2–2.5 µmBernicchia and
Gorjón, 2010
X. vaga Detachable Grandinioid 15–20 ×5–6 µm Ellipsoid, 5–5.5 ×4–4.5 µmBernicchia and
Gorjón, 2010
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Xenasma Donk, Fungus, Wageningen 27: 25 (1957).
MycoBank no.: MB 18755
Type species: Xenasma rimicola (P. Karst.) Donk.
Basidiomata resupinate, adnate, are ceraceous to
gelatinous when fresh, membranaceous when dry, and
have a hymenophore smooth. Hyphal system are monomitic,
generative hyphae with clamp connections. Cystidia and
cystidioles are present. Basidia are cylindrical to subclavate,
pleural, usually with 4 sterigmata and a basal clamp connection.
Basidiospores are globose to cylindrical, colorless, thin-walled,
warted to striate, non-amyloid, and weakly dextrinoid.
Xenosperma Oberw., Sydowia 19(1–6): 45 (1966).
MycoBank no.: MB 18759
Type species: Xenosperma ludibundum (D.P. Rogers and
Liberta) Oberw.
Basidiomata resupinate, closely adnate to the substratum,
are gelatinous when fresh and pruinose when dry. Hyphal
FIGURE 6
The geographic distribution of Xenasmataceae species (holotype) worldwide.
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Luo and Zhao 10.3389/fmicb.2022.970731
systems are monomitic, generative hyphae with clamp
connections. Cystidia are absent. Basidia pleural, usually with
2–4 sterigmata and a basal clamp connection. Basidiospores
are angular, colorless, thin-walled, tetrahedral, with some
protuberances, IKI–, and CB–.
Xenasmatella Oberw., Sydowia 19(1–6): 28 (1966).
MycoBank no.: MB 18756
Type species: Xenasmatella subflavidogrisea (Litsch.)
Oberw. ex Jülich.
TABLE 4 The geographic distribution and host-substratum of Xenasmataceae species (holotype).
Species name Geographic distribution Host-substratum References
Xenasma aculeatum Argentina On fructifications of Hypoxylon Gómez, 1972
X. amylosporum Primorye On rotten trunk of Picea jezoensis Parmasto, 1968
X. longicystidiatum Réunion On Rubus alcaefolius Boidin and Gilles, 2000
X. parvisporum Czech Republic On fallen branch of Quercus petraea Pouzar, 1982
X. praeteritum Ontario On wood Donk, 1957
X. pruinosum Tunisia On oak tree, bared and rotten Donk, 1957
X. pulverulentum Austria On rotten wood Donk, 1957
X. rimicola Finland On cracks in bark Donk, 1957
X. subclematidis Jammu-Kashmir On log Rattan, 1977
X. tulasnelloideum America On very rotten wood Höhnel and Litschauer, 1908
X. vassilievae Khabarovsk On fallen trunk of Taxus cuspidata Parmasto, 1965
Xenasmatella ailaoshanensis Yunnan On trunk of Angiospermae Huang et al., 2019
X. alnicola Allier Sur bois humides, aune, saule blane Bourdot and Galzin, 1928
X. ardosiaca France On decayed wood Bourdot and Galzin, 1928
X. athelioidea Japan On rotten trunk of Quercus Maekawa, 2021
X. bicornis Gabon Among shrubs on shore Boidin and Gilles, 2004
X. borealis Norway On rotten Pinus sylvestris Hjortstam and Larsson, 1987
X. caricis-pendulae Great Britain On dead attached leaf of Carex pendula Roberts, 2007
X. christiansenii Kamchatka On fallen branch of Larix kurilensis var. glabra Parmasto, 1965
X. cinnamomea Florida On Magnolia Burdsall and Nakasone, 1981
X. fibrillosa Iran On decayed wood Hallenberg, 1978
X. globigera Venezuela On hardwood Hjortstam and Ryvarden, 2005
X. gossypina Yunnan On trunk of Angiospermae Zong and Zhao, 2021
X. inopinata Ontario On Tsuga canadensis Jackson, 1950
X. insperata Ontario On bark Jackson, 1950
X. nasti Reunion Under Nastus borbonicus Stalpers, 1996
X. odontioidea Canary On decayed wood Ryvarden and Liberta, 1978
X. palmicola Venezuela On palm Hjortstam and Ryvarden, 2007
X. rhizomorpha Yunnan On trunk of Angiospermae Zong et al., 2021
X. romellii Sweden On deciduous wood Hjortstam, 1983
X. sanguinescens Czech Republic On decayed wood Svrcek, 1973
X. subflavidogrisea Sweden On rotten wood of Pinus sylvestris Jülich, 1979
X. tenuis Yunnan On trunk of Angiospermae Zong et al., 2021
X. vaga Italy On Robinia pseudoacacia Stalpers, 1996
X. wuliangshanensis Yunnan On trunk of Angiospermae Zong and Zhao, 2021
X. xinpingensis Yunnan On trunk of Angiospermae Zong et al., 2021
Xenosperma hexagonosporum France On wood of Platanus acerifolia Boidin and Gilles, 1989
X. ludibundum Massachusetts On bark of Quercus and decayed wood of Chamaecyparis thyoides Jülich, 1979
X. murrillii Florida On branch of Juniperus virginiana Gilbertson and Blackwell, 1987
X. pravum Réunion On dead branch Boidin and Gilles, 1989
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Basidiomata resupinate with a gelatinous. Hyphal system
with clamped generative hyphae. Cystidia are absent. Basidia
pleural, usually with 4 sterigmata and a basal clamp connection.
Basidiospores are hyaline, thin-walled, warted, IKI–, and CB–.
Xenasmatella nigroidea K.Y. Luo and C.L. Zhao, sp. nov.
MycoBank no.: MB 842470, Figures 4,5.
Holotype—China. Yunnan Province, Honghe, Pingbian
County, Daweishan National Nature Reserve, GPS coordinates
23◦42′N, 103◦32′E, altitude 1,500 m asl., on angiosperm stump,
leg. C.L. Zhao, August 3, 2019, CLZhao 18333 (SWFC).
Etymology—nigroidea (Lat.): refers to the black
hymenial surface.
Basidiomata: Basidiomata are annuals, resupinate, thin, very
hard to separate from substrate, odorless or tasteless when fresh,
grayish when fresh, gray to black and brittle when dry, up to
7.5 cm long, 3.5cm wide, 70–150 µm thick. Hymenial is surface
smooth, and byssaceous to reticulate under the lens. Sterile
margin indistinct, black, up to 1 mm wide.
Hyphal system: monomitic, generative hyphae with clamp
connections, thick-walled, unbranched, 2.5–4 µm in diameter,
IKI–, CB–, and tissues unchanged in KOH.
Hymenium: cystidia and cystidioles are absent; basidia are
pleural, clavate, with 4 sterigmata and a basal clamp connection,
12.0–18.0 ×4.5–6 µm; basidioles are shaped similar to basidia
but slightly smaller.
Basidiospores: ellipsoid, colorless, thin-walled, warted
throughout, asperulate with blunt spines up to 0.2 µm long,
with one oil drop inside, IKI–, CB–, 3.5–4.5 ×2.5–3.5 µm, L =
4.07 µm, W =2.87 µm, Q =1.38–1.45 (n=60/2).
Type of rot: White rot.
Additional specimen examined: CHINA, Yunnan Province,
Honghe, Pingbian County, Daweishan National Nature Reserve,
GPS coordinates 23◦40′N, 103◦31′E, altitude 1,500 m asl., on
the angiosperm stump, leg. C.L. Zhao, August 3, 2019, CLZhao
18300 (SWFC).
Discussion
There have been debates among mycologists regarding the
order level taxonomic status of the Xenasmataceae. Corticioid
homobasidiomycetes have a high phylogenetic diversity.
Thus, an accurate place for the taxa of Xenasmataceae has
not been decided. However, it was only assigned to euagarics
clade (Larsson et al., 2004). Later, the Phlebiella family was
proposed by Larsson (2007) on the basis of corticioid fungi;
however, this group was not placed under any order. Recently,
Xenasmataceae was placed under Russulales by He et al.
(2019). Zong et al. (2021) studied the specimens and sequences
from China and treated this group as Xenasmatella as the
phylogenetic datasets showed that this clade does not belong
TABLE 5 Key to 25 accepted species of Xenasmatella worldwide.
1. Gloeocystidia present X. inopinata
1. Cystidia absent 2
2. Basidia with 2, 3 sterigmata X. bicornis
2. Basidia with 4 sterigmata 3
3. Basidia sterigmata >5µm in length X. nasti
3. Basidia sterigmata <5µm in length 4
4. Basidiospores >5µm in length 5
4. Basidiospores <5µm in length 12
5. Basidiospores >4µm in width 6
5. Basidiospores <4µm in width 9
6. Basidiospores globose X. ardosiaca
6. Basidiospores ellipsoid 7
7. Basidia <6µm in width X. vaga
7. Basidia >6µm in width 8
8. Growth on dead angiosperm X. caricis-pendulae
8. Growth on the trunk of gymnosperm X. christiansenii
9. Basidiospores <2µm in width X. athelioidea
9. Basidiospores >2µm in width 10
10. Hymenial margin with fimbriae X. romellii
10. Hymenial margin without fimbriae 11
11. Hymenial surface arachnoid or byssoid X. borealis
11. Hymenial surface smooth X. insperata
12. Basidiospores subglobose to globose 13
12. Basidiospores ellipsoid to subcylindrical 17
13. Basidiospores thick-walled X. globigera
13. Basidiospores thin-walled 14
14. Hymenial surface clay-pink to saffron X. wuliangshanensis
14. Hymenial surface white to grayish or cream to buff 15
15. Generative hyphae thick-walled, unbranched X. xinpingensis
15. Generative hyphae thin-walled, branched 16
16. Hymenial surface gossypine to byssaceous X. gossypina
16. Hymenial surface pruinose to farinaceous X. ailaoshanensis
17. Generative hyphae thick-walled 18
17. Generative hyphae thin-walled 19
18. Hymenial surface gray to black X. nigroidea
18. Hymenial surface clay-buff to cinnamon X. rhizomorpha
19. Growth on palm X. palmicola
19. Growth on other plant 20
20. Growth on the bark of magnolia X. cinnamomea
20. Growth on other wood 21
21. Basidiospores slightly thick-walled X. alnicola
21. Basidiospores thin-walled 22
22. Basidia barrel-shaped X. tenuis
22. Basidia cylindrical 23
23. Basidiomata ochreous X. odontioidea
23. Basidiomata white to gray 24
24. Basidiospores >3µm in width X. fibrillosa
24. Basidiospores <3µm in width X. subflavidogrisea
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Luo and Zhao 10.3389/fmicb.2022.970731
to any order. In the present study (Figure 1), the ITS+nLSU
analyses of 13 orders, namely, Agaricales, Atheliales, Boletales,
Cantharellales, Corticiales, Gloeophyllales, Gomphales,
Hymenochaetales, Polyporales, Russulales, Thelephorales,
Trechisporales, and Xenasmatales showed that the taxa of
Xenasmataceae form a single lineage with the sequences of
Hymenochaetales and Atheliales; and this is similar to the
results of Larsson (2007). In the present study (Figure 2), the
nLSU analysis showed that the taxa of Xenasmataceae form
a single lineage with the sequences of Hymenochaetales and
Boletales; and this is similar to the results of Larsson (2007). In
the present study (Table 2), we have enumerated morphological
differences among the related orders. Therefore, a new fungal
order, Xenasmatales, is proposed on the basis of morphological
and molecular identification.
Phlebiella was not deemed to be a legitimately published
genus (Duhem, 2010), and transferring to Xenasmatella was
proposed. Later, Larsson et al. (2020) studied corticioid fungi
(Basidiomycota and Agaricomycetes) and agreed with Duhem
(2010), who suggested accepting the genus Xenasmatella.
Recently, several mycologists have suggested the replacement
of the invalid genus Phlebiella with Xenasmatella on the basis
of morphology and molecular analyses (Maekawa, 2021;Zong
et al., 2021).
On the basis of ITS dataset, a previous study showed
that nine species of Xenasmatella have been reported, of
which 6 new species were found in China, namely, X.
ailaoshanensis C.L. Zhao ex C.L. Zhao and T.K. Zong, X.
gossypina,X. rhizomorpha,X. tenuis,X. wuliangshanensis, and
X. xinpingensis. According to our sequence data, Xenasmatella
nigroidea was nested into Xenasmatella with strong statistical
support (Figure 3), and formed a sister group with X.
vaga. However, X. nigroidea is morphologically distinguished
from X. vaga by larger basidiospores (5–5.5 ×4–4.5 µm).
In addition, it turns dark red or purplish with KOH
(Bernicchia and Gorjón, 2010).
Morphological comparisons of Xenasmatella nigroidea and
other species are included in Table 3.Xenasmatella nigroidea
is similar to X. christiansenii (Parmasto) Stalpers, X. fibrillosa
(Hallenb.) Stalpers, X. gossypina, and X. rhizomorpha C.L. Zhao
by having gossypine, byssaceous to reticulate hymenial surface,
however, X. christiansenii is distinguished from X. nigroidea
by its larger basidiospores (6–7 ×4–4.5 µm) and asperulate
with blunt spines (up to 1 µm long; Bernicchia and Gorjón,
2010). Xenasmatella fibrillosa differs from X. nigroidea due
to the presence of a white to pale yellowish white hymenial
surface and longer basidiospores (4.5–5.5 µm; Bernicchia and
Gorjón, 2010). Xenasmatella gossypina can be distinguished
from X. nigroidea because it has cotton to flocculent basidiomata
with a cream to buff hymenial surface and subglobose to
globose basidiospores (Zong and Zhao, 2021). Xenasmatella
rhizomorpha is separated from X. nigroidea by the clay-
buff to cinnamon hymenial surface and the presence of the
rhizomorphs (Zong et al., 2021).
Xenasmatella nigroidea is similar to X. gaspesica
(Liberta) Hjortstam, X. odontioidea Ryvarden & Liberta,
X. subflavidogrisea (Litsch.) Oberw. ex Jülich, and X.
vaga (Fr.) Stalpers due to the presence of the ellipsoid or
narrowly ellipsoid basidiospores. However, X. gaspesica
differs from X. nigroidea because it has smaller basidia (7–11
×4–4.5 µm) and larger basidiospores (8–10 ×2–2.5 µm;
Grosse-Brauckmann and Kummer, 2004). Xenasmatella
odontioidea can be distinguished from X. nigroidea by
its colliculosa hymenial surface and shorter basidiospores
(2.5–3.5 µm; Ryvarden and Liberta, 1978). Xenasmatella
subflavidogrisea is separated from X. nigroidea due to the
presence of a white to grayish hymenial surface, turning
dark reddish brown in KOH and narrower basidiospores
(2–2.5 µm; Bernicchia and Gorjón, 2010). Xenasmatella
vaga differs from X. nigroidea due to its grandinioid
hymenial surface and larger basidiospores (5–5.5 ×4–4.5 µm;
Bernicchia and Gorjón, 2010).
Based on the geographical distribution in America, Asia, and
Europe, and ecological habits, white-rot causing Xenasmataceae
have been reported in angiosperms and gymnosperms (Figure 6
and Table 4) (Stalpers, 1996;Dai et al., 2004;Hjortstam
and Ryvarden, 2005;Bernicchia and Gorjón, 2010;Duhem,
2010;Dai, 2011;Huang et al., 2019;Larsson et al., 2020;
Maekawa, 2021;Zong and Zhao, 2021;Zong et al., 2021).
Key to 25 accepted species of Xenasmatella worldwide in
Table 5. Many wood-decaying fungi have been recently reported
worldwide (Zhu et al., 2019;Angelini et al., 2020;Gafforov
et al., 2020;Zhao and Zhao, 2021). According to the
results of our study on Xenasmatella, all these fungi can
be classified into a new taxon (Figure 3). In addition, this
study contributes to the knowledge of the fungal diversity
in Asia.
Data availability statement
The datasets presented in this study can be found in
online repositories. The names of the repository/repositories
and accession number(s) can be found in the
article/supplementary material.
Author contributions
C-LZ: conceptualization, resources, supervision,
project administration, and funding acquisition. C-LZ
and K-YL: methodology, software, validation, formal
analysis, investigation, writing—original draft preparation,
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Luo and Zhao 10.3389/fmicb.2022.970731
writing—review and editing, and visualization. Both
authors have read and agreed to the published version of
the manuscript.
Funding
The research was supported by the National Natural Science
Foundation of China (Project No. 32170004, U2102220) to
C-LZ, the Yunnan Fundamental Research Project (Grant No.
202001AS070043) to C-LZ, the High-level Talents Program
of Yunnan Province (YNQR-QNRC-2018-111) to C-LZ, and
the Yunnan Key Laboratory of Plateau Wetland Conservation,
Restoration, and Ecological Services (202105AG070002)
to K-YL.
Conflict of interest
The authors declare that the research was conducted in the
absence of any commercial or financial relationships that could
be construed as a potential conflict of interest.
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References
Angelini, C., Vizzini, A., Justo, A., Bizzi, A., and Kaya, E. (2020). First report
of a neotropical agaric (lepiota spiculata, agaricales, basidiomycota) containing
lethal α-amanitin at toxicologically relevant levels. Front. Microbiol. 11, 1833.
doi: 10.3389/fmicb.2020.01833
Bernicchia, A., and Gorjón, S. P. (2010). Fungi Europaei 12: Corticiaceae s.l.
Alassio: Edizioni Candusso.
Besl, H., and Bresinsky, A. (1997). Chemosystematics of suillaceae
and gomphidiaceae (suborder Suillineae). Plant Syst. Evol. 206, 223–242.
doi: 10.1007/BF00987949
Boidin, J., and Gilles, G. (1989). Les Corticiés pleurobasidiés (Basidiomycotina)
en France. Cryptogamic Bot. 1, 70–79.
Boidin, J., and Gilles, G. (2000). Basidiomycètes Aphyllophorales de l’ile de La
Reunion. XXI - Suite. Mycotaxon 75, 357–387.
Boidin, J., and Gilles, G. (2004). Homobasidiomycètes Aphyllophorales non
porés à basides dominantes à 2 (3) stérigmates. Bull. Trimest. Soc. Mycol. Fr.
119, 1–17.
Bourdot, H., and Galzin, A. (1928). Hyménomycètes de France : Hétérobasidiés.
Homobasidiés gymnocarpes / par MM. l’abbé H. Bourdot et A. Galzin. Paris:
Bibliothèque nationale de France, département Sciences et techniques, 786.
Burdsall, H. H., and Nakasone, K. K. (1981). New or little known lignicolous
aphyllophorales (Basidiomycotina) from southeastern United States. Mycologia 73,
454–476. doi: 10.1080/00275514.1981.12021368
Chen, C. C., Chen, C. Y., and Wu, S. H. (2021). Species diversity,
taxonomy and multi-gene phylogeny of phlebioid clade (Phanerochaetaceae,
Irpicaceae, Meruliaceae) of polyporales. Fungal Divers. 111, 1–106.
doi: 10.1007/s13225-021-00490-w
Cui, B. K., Du, P., and Dai, Y. C. (2011). Three new species of Inonotus
(Basidiomycota, Hymenochaetaceae) from China. Mycol. Prog. 10, 107–114.
doi: 10.1007/s11557-010-0681-6
Dai, Y. C. (2011). A revised checklist of corticioid and hydnoid fungi in China
for 2010. Mycoscience 52, 69–79. doi: 10.1007/S10267-010-0068-1
Dai, Y. C. (2012). Polypore diversity in China with an annotated checklist of
Chinese polypores. Mycoscience 53, 49–80. doi: 10.1007/s10267-011-0134-3
Dai, Y. C., Cui, B. K., Si, J., He, S. H., Hyde, K. D., Yuan, H. S., et al. (2015).
Dynamics of the worldwide number of fungi with emphasis on fungal diversity in
China. Mycol. Prog. 14, 62. doi: 10.1007/s11557-015-1084-5
Dai, Y. C., Wei, Y. L., and Zhang, X. Q. (2004). An annotated checklist of
non-poroid Aphyllophorales in China. Ann. Bot. Fennici 41, 233–247.
Dai, Y. C., Yang, Z. L., Cui, B. K., Wu, G., Yuan, H. S., Zhou, L. W., et al.
(2021). Diversity and systematics of the important macrofungi in Chinese forests.
Mycosystema 40, 770–805.
Donk, M. A. (1957). Notes on resupinate Hymenomycetes IV. Fungus 27, 1–29.
Duhem, B. (2010). Deux corticiés nouveaux méditerranéens à spores allantoïdes.
Cryptogam. Mycol. 31, 143–152.
Eriksson, J., Hjortstam, K., and Ryvarden, L. (1978). Corticiaceae of North Europe
Volume 5: Mycoaciella-Phanerochaete. Oslo: Fungiflora.
Eriksson, J., Hjortstam, K., and Ryvarden, L. (1981). Corticiaceae of North Europe
Volume 6: Phlebia-Sarcodontia. Oslo: Fungiflora.
Eriksson, J., Hjortstam, K., and Ryvarden, L. (1984). Corticiaceae of North Europe
Volume 7: Schizopora-Suillosporium. Oslo: Fungiflora.
Felsenstein, J. (1985). Confidence intervals on phylogenetics: an approach
using bootstrap. Evolution 39, 783–791. doi: 10.1111/j.1558-5646.1985.tb00
420.x
Fries, E. (1821–1832). Systema Mycologicum, Sistens Fungorum Ordines,
Generaet Species Hucusque Cognitas. Gryphiswaldiae: Ernestus Mauritius.
Fries, E. (1828). Elenchus Fungorum. Vols. I and II. Germany: Greifswald.
Fries, E. (1857–1863). Monographia Hymenomycetum Sueciae. Vols. I and II.
Leffler, C. A. Uppsala: Nabu Press.
Fries, E. (1874). Hymenomycetes Europaei. Berling; Uppsala: Typis descripsip Ed.
p. 755.
Gafforov, Y., Ordynets, A., Langer, E., Yarasheva, M., de Mello Gugliotta, A.,
Schigel, D., et al. (2020). Species diversity with comprehensive annotations of
wood-inhabiting poroid and corticioid fungi in Uzbekistan. Front. Microbiol. 11,
598321. doi: 10.3389/fmicb.2020.598321
Garcia-Sandoval, R., Wang, Z., Binder, M., and Hibbett, D. S. (2010).
Molecular phylogenetics of the Gloeophyllales and relative ages of clades of
Agaricomycotina producing a brown rot. Mycologia 103, 510–524. doi: 10.3852/
10-209
Gilbert, J. E. (1931). Les Bolets, in les livres du Mycologue. Paris: Le Village du
Livre. p. 254.
Gilbertson, R. L., and Blackwell, M. (1987). Notes on wood-rotting fungi on
Junipers in the Gulf Coast region. II. Mycotaxon 28, 369–402.
Gilbertson, R. L., and Ryvarden, L. (1987). North American Polypores 1-2.
Fungiflora; Oslo: Lubrecht and Cramer Ltd. p. 1–433.
Gómez, C. E. (1972). Xenasma y géneros afines de los alrededores de Buenos
Aires (Aphyllophorales). Bol Soc Argent Bot 14, 269–281.
Gonzalez-Avila, A., Contreras-Medina, R., Espinosa, D., and Luna-Vega, I.
(2017). Track analysis of the order Gomphales (Fungi: Basidiomycota) in Mexico.
Phytotaxa 316, 22–38. doi: 10.11646/phytotaxa.316.1.2
Grosse-Brauckmann, H., and Kummer, V. (2004). Fünf bemerkenswerte
Funde corticioider Pilze aus Deutschland. Feddes Repert. 115, 90–101.
doi: 10.1002/fedr.200311029
Haelewaters, D., Dirks, A. C., Kappler, L. A., Mitchell, J. K., Quijada, L.,
Vandegrift, R., et al. (2018). A preliminary checklist of fungi at the Boston Harbor
Islands. Northeast. Nat. 25, 45–76. doi: 10.1656/045.025.s904
Hallenberg, N. (1978). Wood-Fungi (Corticiaceae, Coniophoraceae,
Lachnocladiaceae, Thelephoraceae) in N. Iran. I. Iran. J. Plant Pathol. 14, 38–87.
Frontiers in Microbiology 15 frontiersin.org

Luo and Zhao 10.3389/fmicb.2022.970731
He, M. Q., Zhao, R. L., Hyde, K. D., Begerow, D., Kemler, M., Yurkov, A., et al.
(2019). Notes, outline and divergence times of Basidiomycota. Fungal Divers. 99,
105–367. doi: 10.1007/s13225-019-00435-4
Hibbett, D. S., Binder, M., Bischoff, J. F., Blackwell, M., Cannon, P. F., Eriksson,
O. E., et al. (2007). A higher-level phylogenetic classification of the Fungi. Mycol.
Res. 111, 509–547. doi: 10.1016/j.mycres.2007.03.004
Hjortstam, K. (1983). Notes on Corticiaceae (Basidiomycetes). XII. Mycotaxon
17, 577–584.
Hjortstam, K., and Larsson, K. H. (1987). Additions to Phlebiella (Corticiaceae,
Basidiomycetes), with notes on Xenasma and Sistotrema. Mycotaxon 29, 315–319.
Hjortstam, K., and Ryvarden, L. (2005). New taxa and new combinations in
tropical corticioid fungi, (Basidiomycotina, Aphyllophorales). Synop. Fungorum
20, 33–41.
Hjortstam, K., and Ryvarden, L. (2007). Studies in corticioid fungi from
Venezuela III (Basidiomycotina, Aphyllophorales). Synop. Fungorum 23, 56–107.
Höhnel, F., and Litschauer, V. (1908). Beiträge zur Kenntnis der Corticieen: III.
Sitz. K. Akad. Wiss. Math. Naturw. Klasse Abt. I. 117, 1081–1124.
Huang, R. X., Chen, J. Z., Wu, J. R., and Zhao, C. L. (2019). Phlebiella
ailaoshanensis sp. nov. (Polyporales, Basidiomycota) described from China.
Phytotaxa 419, 105–109. doi: 10.11646/phytotaxa.419.1.8
Jackson, H. S. (1950). Studies of Canadian Thelephoraceae. VII. Some
new species of Corticium, section Athele. Can. J. Res. 28, 716–725.
doi: 10.1139/cjr50c-045
James, T. Y., Stajich, J. E., Hittinger, C. T., and Rokas, A. (2020). Toward
a fully resolved fungal tree of life. Annu. Rev. Microbiol. 74, 291–313.
doi: 10.1146/annurev-micro-022020-051835
Jarosch, M. (2001). Zur molekularen systematik der boletales: coniophorineae,
paxillineae und suillineae. Bibl. Mycol. 191, 1–158.
Jarosch, M., and Besl, H. (2001). Leucogyrophana, a polyphyletic genus of the
order boletales (Basidiomycetes). Plant Biol. 3, 443–448. doi: 10.1055/s-2001-16455
Jülich, W. (1979). Studies in resupinate Basidiomycetes - V. On some new taxa.
Persoonia 10, 325–336.
Kirk, P. M., Cannon, P. F., David, J. C., Minter, D. W., and Stalpers, J. A. (2008).
Ainsworth and bisby’s dictionary of the fungi. 10th ed. Wallingford, Oxon, UK:
CAB International Press, 783.
Larsson, A. (2014). AliView: a fast and lightweight alignment
viewer and editor for large data sets. Bioinformatics 30, 3276–3278.
doi: 10.1093/bioinformatics/btu531
Larsson, E., and Hallenberg, N. (2001). Species delimitation in the
Gloeocystidiellum Porosum-Clavuligerum complex inferred from compatibility
studies and nuclear RDNA sequence data. Mycologia 93, 907–914.
doi: 10.1080/00275514.2001.12063225
Larsson, E., and Larsson, K. H. (2003). Phylogenetic relationships of
Russuloid Basidiomycetes with emphasis on Aphyllophoralean taxa. Mycologia 95,
1037–1065. doi: 10.1080/15572536.2004.11833020
Larsson, K. H. (2007). Re-thinking the classification of corticioid fungi. Mycol.
Res. 111, 1040–1063. doi: 10.1016/j.mycres.2007.08.001
Larsson, K. H., Larsson, E., and Koljalg, U. (2004). High phylogenetic
diversity among corticioid homobasidiomycetes. Mycol. Res. 108, 983–1002.
doi: 10.1017/S0953756204000851
Larsson, K. H., Larsson, E., Ryvarden, L., and Spirin, V. (2020). Some
new combinations of corticioid fungi (Basidiomycota. Agaricomycetes). Synop.
Fungorum 40, 113–117.
Larsson, K. H., Parmasto, E., Fischer, M., Langer, E., Nakasone,
K. K., and Redhead, S. A. (2006). Hymenochaetales: a molecular
phylogeny for the hymenochaetoid clade. Mycologia 98, 926–936.
doi: 10.1080/15572536.2006.11832622
Maekawa, N. (2021). Taxonomy of corticioid fungi in Japan: present status
and future prospects. Mycoscience 62, 345–355. doi: 10.47371/mycosci.2021.
10.002
Miller, M. A., Pfeiffer, W., and Schwartz, T. (2012). The CIPRES science gateway:
enabling high-impact science for phylogenetics researchers with limited resources.
Assoc. Comput. Mach. 39, 1–8. doi: 10.1145/2335755.2335836
Núñez, M., and Ryvarden, L. (2001). East Asian polypores 2. Synop. Fungorum
14, 165–522.
Nylander, J. A. A. (2004). MrModeltest v2. Program Distributed by the Author.
Uppsala: Evolutionary Biology Centre; Uppsala Univeristy.
Oberwinkler, F. (1966). Primitive Basidiomyceten. Revision einiger
Formenkreise von Basidienpilzen mit plastischer Basidie. Sydowia 19, 1–72.
Oberwinkler, F. (1975). Eine agaricoide Gattung der Thelephorales. Sydowia
28, 359–362.
Parmasto, E. (1965). Corticiaceae U.R.S.S. I. Descriptiones taxorum
novarum. Combinationes novae. Eesti NSV Tead. Akad. TOIM. 14, 220–233.
doi: 10.3176/biol.1965.2.06
Parmasto, E. (1968). Conspectus Systematis Corticiacearum. Spain: Euorpa Press.
p. 1–261.
Petersen, J. H. (1996). Farvekort. The Danish Mycological Society’s Colour-Chart.
Greve: Foreningen til Svampekundskabens Fremme.
Piatek, M. (2005). A note on the genus Xenosmatella (Fungi, Basidiomycetes).
Polish Bot. J. 50, 11–13.
Pouzar, Z. (1982). Taxonomic studies in resupinate fungi I. Ceská Mykol.
36, 141–145.
Rattan, S. S. (1977). The resupinate Aphyllophorales of the North Western
Himalayas. Bibl. Mycol. 60, 1–427.
Rayner, R. W. (1970). A Mycological Colour Chart. Commonwealth Mycological
Institute, Kew and British Mycological Society, Kew, United Kingdom. p. 1–34.
Roberts, P. J. (2007). Phlebiella caricis-pendulae: A new corticoid fungus from
Wales. Synop. Fungorum 22, 25–26.
Ronquist, F., Teslenko, M., van der Mark, P., Ayres, D. L., Darling, A.,
Hohna, S., et al. (2012). Mrbayes 3.2: efficient bayesian phylogenetic inference
and model choice across a large model space. Syst. Biol. 61, 539–542.
doi: 10.1093/sysbio/sys029
Rosenthal, L. M., Larsson, K. H., Branco, S., Chung, J. A., Glassman, S.
I., Liao, H. L., et al. (2017). Survey of corticioid fungi in North American
pinaceous forests reveals hyperdi-versity, underpopulated sequence databases,
and species that are potentially ectomycorrhizal. Mycologia 109, 115–127.
doi: 10.1080/00275514.2017.1281677
Ryvarden, L., and Liberta, A. E. (1978). Contribution to the Aphyllophoralles of
the Canary Islands 4. Two new species of Trechispora and Xenmastella.Can. J. Bot.
56, 2617–2619. doi: 10.1139/b78-314
Ryvarden, L., and Melo, I. (2014). Poroid fungi of Europe. Synop. Fungorum
31, 1–455.
Sotome, K., Hattori, T., and Ota, Y. (2011). Taxonomic study on a threatened
polypore, Polyporus pseudobetulinus, and a morphologically similar species, P.
subvarius.Mycoscience 52, 319–326. doi: 10.1007/S10267-011-0111-X
Stalpers, J. A. (1996). The aphyllophoraceous fungi II. Keys to the species of the
Hericiales. Stud. Mycol. 40, 1–185.
Svrcek, M. (1973). Species novae Corticiacearum e Bohemia. Ceská Mykol.
27, 201–206.
Swofford, D. L. (2002). PAUP∗: Phylogenetic Analysis Using Parsimony (∗and
other methods). Version 4.0b10. Sunderland, Massachusetts: Sinauer Associates.
Telleria, M. T., Melo, I., Duenas, M., Larsson, K. H., and Paz Martin, M. P. (2013).
Molecular analyses confirm Brevicellicium in Trechisporales. IMA Fungus 4, 21–28.
doi: 10.5598/imafungus.2013.04.01.03
Tobias, W., and Michael, F. (2002). Classification and phylogenetic relationships
of Hymenochaete and allied genera of the Hymenochaetales, inferred from rDNA
sequence data and nuclear behaviour of vegetative mycelium. Mycol. Prog. 1,
93–104. doi: 10.1007/s11557-006-0008-9
Vu, D., Groenewald, M., de Vries, M., Gehrmann, T., Stielow, B., Eberhardt,
U., et al. (2019). Large-scale generation and analysis of filamentous fungal
DNA barcodes boosts coverage for kingdom fungi and reveals thresholds
for fungal species and higher taxon delimitation. Stud. Mycol. 92, 135–154.
doi: 10.1016/j.simyco.2018.05.001
Wagner, T., and Fischer, M. (2001). Natural groups and a revised system for
the European poroid Hymenochaetales (Basidiomycota) supported by nLSU rDNA
sequence data. Mycol. Res. 105, 773–782. doi: 10.1017/S0953756201004257
White, T. J., Bruns, T., Lee, S., and Taylor, J. (1990). “Amplification and direct
sequencing of fungal ribosomal RNA genes for phylogenetics,” in PCR protocols: A
Guide to Methods And Applications, eds Innis, M. A., Gelfand, D. H., Sninsky, J. J.,
and White, T. J. (San Diego, CA: Academic Press), 315–322.
Wu, F., Yuan, H. S., Zhou, L. W., Yuan, Y., Cui, B. K., and Dai, Y. C. (2020).
Polypore diversity in South China. Mycosystema 39, 653–682.
Wu, F., Yuan, Y., and Zhao, C. L. (2015). Porpomyces submucidus
(Hydnodontaceae, Basidiomycota), a new species from tropical China
based on morphological and molecular evidence. Phytotaxa 230, 61–68.
doi: 10.11646/phytotaxa.230.1.5
Wu, F., Zhou, L. W., Vlasák, J., and Dai, Y. C. (2022). Global diversity and
systematics of Hymenochaetaceae with poroid hymenophore. Fungal Divers. 113,
1–192. doi: 10.1007/s13225-021-00496-4
Frontiers in Microbiology 16 frontiersin.org

Luo and Zhao 10.3389/fmicb.2022.970731
Zhao, C. L., and Wu, Z. Q. (2017). Ceriporiopsis kunmingensis sp. nov.
(Polyporales, Basidiomycota) evidenced by morphological characters and
phylogenetic analysis. Mycol. Prog. 16, 93–100. doi: 10.1007/s11557-016-1259-8
Zhao, W., and Zhao, C. L. (2021). The phylogenetic relationship revealed three
new wood-inhabiting fungal species from genus Trechispora.Front. Microbiol. 12,
650195. doi: 10.3389/fmicb.2021.650195
Zhu, L., Song, J., Zhou, J. L., Si, J., and Cui, B. K. (2019). Species diversity,
phylogeny, divergence time and biogeography of the genus Sanghuangporus
(Basidiomycota). Front. Microbiol. 10, 812. doi: 10.3389/fmicb.2019.
00812
Zong, T. K., Wu, J. R., and Zhao, C. L. (2021). Three new Xenasmatella
(Polyporales, Basidiomycota) species from China. Phytotaxa 489, 111–120.
doi: 10.11646/phytotaxa.489.2.1
Zong, T. K., and Zhao, C. L. (2021). Morphological and molecular identification
of two new species of Phlebiella (Polyporales, Basidiomycota) from southern
China. Nova Hedwig. 112, 501–514. doi: 10.1127/nova_hedwigia/2021/0628
Frontiers in Microbiology 17 frontiersin.org