Four New Wood-Inhabiting Fungal Species of Peniophoraceae (Russulales, Basidiomycota) from the Yunnan-Guizhou Plateau, China

Abstract

Four new fungi of the family Peniophoraceae, viz., Peniophora roseoalba, P. yunnanensis, Vararia daweishanensis, and V. fragilis are herein proposed, based on a combination of morphological features and molecular evidence. Peniophora roseoalba is characterized by resupinate, membranaceous basidiomata with a rose pink to pale pinkish grey hymenophore, a monomitic hyphal system with clamped generative hyphae, the presence of cystidia, and ellipsoid basidiospores. However, P. yunnanensis differs in being tuberculate, with a pale cream to cream hymenial surface, small lamprocystidia (18–29 × 4.5–7 µm), and subcylindrical basidiospores. Vararia daweishanensis is characterized by resupinate, membranous basidiomata with a pale yellowish hymenial surface, a dimitic hyphal system with clamped generative hyphae, strongly dextrinoid dichohyphae, and allantoid basidiospores; V. fragilis is characterized by resupinate, brittle basidiomata, with a buff to ochraceous hymenial surface and small ellipsoid basidiospores measuring 3.5–5.5 × 2.5–3.5 µm. Sequences of the ITS and nLSU rRNA markers of the studied samples were generated, and phylogenetic analyses were performed with the maximum likelihood, maximum parsimony, and Bayesian inference methods. The nLSU analysis revealed that the four new species can be clustered into the family Peniophoraceae (Russulales), in the genera Peniophora and Vararia. Further studies based on the ITS dataset showed that four fungi of the family Peniophoraceae were new to science.

Full text

Citation: Zou, L.; Zhang, X.; Deng, Y.;
Zhao, C. Four New Wood-Inhabiting
Fungal Species of Peniophoraceae
(Russulales, Basidiomycota) from the
Yunnan-Guizhou Plateau, China. J.
Fungi 2022,8, 1227. https://doi.org/
10.3390/jof8111227
Academic Editor: Seung-Yoon Oh
Received: 13 October 2022
Accepted: 18 November 2022
Published: 21 November 2022
Publisher’s Note: MDPI stays neutral
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4.0/).
Fungi
Journal of
Article
Four New Wood-Inhabiting Fungal Species of Peniophoraceae
(Russulales, Basidiomycota) from the Yunnan-Guizhou
Plateau, China
Lei Zou 1,2, Xiaolu Zhang 2, Yinglian Deng 2and Changlin Zhao 1,2,3,*
1Key Laboratory for Forest Resources Conservation and Utilization in the Southwest Mountains of China,
Ministry of Education, Southwest Forestry University, Kunming 650224, China
2College of Biodiversity Conservation, Southwest Forestry University, Kunming 650224, China
3Yunnan Key Laboratory for Fungal Diversity and Green Development, Kunming Institute of Botany,
Chinese Academy of Sciences, Kunming 650201, China
*Correspondence: fungi@swfu.edu.cn or fungichanglinz@163.com
Abstract:
Four new fungi of the family Peniophoraceae, viz., Peniophora roseoalba,P. yunnanensis,
Vararia daweishanensis, and V. fragilis are herein proposed, based on a combination of morphological
features and molecular evidence. Peniophora roseoalba is characterized by resupinate, membranaceous
basidiomata with a rose pink to pale pinkish grey hymenophore, a monomitic hyphal system with
clamped generative hyphae, the presence of cystidia, and ellipsoid basidiospores. However, P. yunna-
nensis differs in being tuberculate, with a pale cream to cream hymenial surface, small lamprocystidia
(18–29
×
4.5–7
µ
m), and subcylindrical basidiospores. Vararia daweishanensis is characterized by re-
supinate, membranous basidiomata with a pale yellowish hymenial surface, a dimitic hyphal system
with clamped generative hyphae, strongly dextrinoid dichohyphae, and allantoid basidiospores;
V. fragilis is characterized by resupinate, brittle basidiomata, with a buff to ochraceous hymenial
surface and small ellipsoid basidiospores measuring 3.5–5.5
×
2.5–3.5
µ
m. Sequences of the ITS
and nLSU rRNA markers of the studied samples were generated, and phylogenetic analyses were
performed with the maximum likelihood, maximum parsimony, and Bayesian inference methods.
The nLSU analysis revealed that the four new species can be clustered into the family Peniophoraceae
(Russulales), in the genera Peniophora and Vararia. Further studies based on the ITS dataset showed
that four fungi of the family Peniophoraceae were new to science.
Keywords: Asia; macrofungi; molecular systematics; taxonomy; Yunnan Province
1. Introduction
The family Peniophoraceae (Russulales) is a large and rather heterogeneous family
with seven genera accepted; two genera, Peniophora Cooke and Vararia P. Karst., have the
highest number of taxa in this family, in which they play fundamental ecological roles to
drive carbon cycling in forest soils, acting as decomposers [1,2].
Peniophora, typified by P. quercina (Pers.) Cooke, is characterized by resupinate, mem-
branaceous to ceraceous basidiomata, with smooth to tuberculate hymenophores having
a grey, violaceous, orange, red, or brown hymenial surface, a monomitic hyphal system
with clamped generative hyphae; dendrohyphidia, lamprocystidia, and gloeocystidia are
present or absent; the basidiospores are ellipsoid, cylindrical to allantoid, smooth, thin-
walled, acyanophilous, and without reaction with Melzer [
3
]. Based on the MycoBank
database (http://www.MycoBank.org, accessed on 13 October 2022) and the Index Fungo-
rum (http://www.indexfungorum.org, accessed on 13 October 2022), the genus Peniophora
has 637 specific and registered names, but the actual number of species has reached
191 [4–18]
.Vararia is typified by V. investiens (Schwein.) P. Karst. This genus is character-
ized by resupinate basidiomata, a dimitic hyphal system with clamped or simple-septate
J. Fungi 2022,8, 1227. https://doi.org/10.3390/jof8111227 https://www.mdpi.com/journal/jof

J. Fungi 2022,8, 1227 2 of 23
generative hyphae and often, dextrinoid dichohyphae, the presence of gloeocystidia, and
variously shaped smooth basidiospores with or without an amyloid reaction [
19
–
21
]. The
MycoBank database (http://www.MycoBank.org, accessed on 13 October 2022) and Index
Fungorum (http://www.indexfungorum.org, accessed on 13 October 2022) have registered
96 specific and infraspecific names in Vararia, but the actual number of the species has
reached 72, and they occur mainly in the tropical and subtropical areas of the world [
22
–
31
].
However, Vararia is still poorly studied in China [
32
], from whence eight species, namely, V.
amphithallica Boidin, Lanq. & Gilles, V. bispora S.L. Liu & S.H. He, V. breviphysa Boidin &
Lanq., V. cinnamomea Boidin, Lanq. & Gilles, V. investiens (Schwein.) P. Karst., V. montana
S.L. Liu & S.H. He, V. racemosa (Burt.) D.P. Rogers & H.S. Jacks., and V. sphaericospora Gilb.,
have been reported in this country [32–34].
These pioneering research studies into the Peniophoraceae family were just the prelude
to the molecular systematics period. The phylogenetic diversity displayed by corticioid fun-
gal species, based on 5.8S and 28S nuclear rDNA, revealed that the taxa of Peniophoraceae
are nested in the russuloid clade, which holds a considerable share of the phylogenetic
framework, and include the genera of Peniophora and Vararia [
35
]. The phylogenetic research
about the major clades of mushroom-forming fungi (Homobasidiomycetes) indicated that
the largest resupinate forms divided into the polyporoid, russuloid, and hymenochaetoid
clades, in which Peniophora grouped with Asterostroma Massee and Scytinostroma Donk [
36
].
Molecular phylogenetic analyses of nrITS and nrLSU sequences revealed affinities among
families with the Peniophorales in the Russulales, in which the presence of distinctive
hyphal elements, which are homologous to the defining features of Peniophorales, was
consistent with the phylogenetic evidence, and the Varariaceae were grouped closely with
the Peniophoraceae [37].
During the investigations into wood-inhabiting fungi in Yunnan Province, China, four
new taxa of Peniophoraceae were found that could not be assigned to any described species.
Herein, we present the morphological and molecular phylogenetic evidence that supports
the recognition of these four new species within the Peniophora and Vararia, based on the
internal transcribed spacer (ITS) regions and the large subunit nuclear ribosomal RNA gene
(nLSU) sequences.
2. Materials and Methods
2.1. Morphology
Fresh fruiting bodies of the fungi were collected from Chuxiong, Honghe, Puer, and
Wenshan of Yunnan Province, in China. The specimens were dried in an electric food dehy-
drator at 40
◦
C, then sealed and stored in an envelope bag and deposited in the herbarium
of the Southwest Forestry University (SWFC), Kunming, Yunnan Province, China. The
macromorphological descriptions are based on field notes and photos captured in the field
and laboratory. The color terminology follows the example set by Petersen [
38
–
40
]. Micro-
morphological data were obtained from the dried specimens when observed under a light
microscope, following the method used by Dai [
41
]. The following abbreviations were used:
KOH = 5% potassium hydroxide water solution, CB = Cotton Blue,
CB−= acyanophilous
,
CB+ = cyanophilous, IKI = Melzer’s reagent,
IKI−= both inamyloid and indextrinoid
,
L = mean spore length (arithmetic average for all spores)
, W = mean spore width (arith-
metic average for all spores), Q = variation in the L/Wratios between the specimens studied,
and n= a/b (number of spores (a) measured from a given number (b) of specimens).
2.2. Molecular Phylogeny
The CTAB rapid plant genome extraction kit-DN14 (Aidlab Biotechnologies Co., Ltd.,
Beijing, China) was used to obtain the genomic DNA from the dried specimens using the
manufacturer’s instructions, following a previous study [
42
]. The nuclear ribosomal ITS
region was amplified with the primers ITS5 and ITS4 [
43
]. The nuclear nLSU region was
amplified with the primer pair, LR0R and LR7 (http://lutzonilab.org/nuclear-ribosomal-
dna/; accessed on 13 October 2022). The PCR procedure for ITS was as follows: initial

J. Fungi 2022,8, 1227 3 of 23
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, with a final extension of 72
◦
C for 10 min. The PCR procedure 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, with a final extension of 72
◦
C for 10 min. The
PCR products were purified and sequenced at Kunming Tsingke Biological Technology
Company, Limited (Yunnan Province, China). All of the newly generated sequences were
deposited in the NCBI GenBank (https://www.ncbi.nlm.nih.gov/genbank/; accessed on
13 October 2022) (Table 1).
The sequencer, 4.6 (GeneCodes, Ann Arbor, MI, USA), was used to assemble and edit
the generated sequence reads. The sequences were aligned in MAFFT 7 (https://mafft.
cbrc.jp/alignment/server/; accessed on 13 October 2022), using the “G-INS-i” strategy for
the ITS and nLSU dataset, manually adjusted in BioEdit [
44
]. The sequences of Sistotrema
brinkmannii (Bres.) J. Erikss. and S. coronilla (Höhn.) Donk ex. D.P. Rogers, obtained
from GenBank, were selected as an outgroup for the phylogenetic analysis of the nLSU
phylogenetic tree (Figure 1) [
45
]; the sequences of Dichostereum durum (Bourdot & Galzin)
Pilát and D. effuscatum (Cooke & Ellis) Boidin & Lanq. were selected as an outgroup for
phylogenetic analysis of ITS phylogenetic tree (Figure 2) [
45
]; the sequences of P. incarnata
(Pers.) P. Karst. and P. nuda (Fr.) Bres. were selected as an outgroup in the ITS analysis
(Figure 3), following the method of a previous study [38].
J. Fungi 2022, 8, x FOR PEER REVIEW 8 of 25
Figure 1. A maximum parsimony strict consensus tree, illustrating the phylogeny of four new
species and related genera in the order Russulales, based on nLSU sequences. The branches are
labeled with maximum likelihood bootstrap values of > 70%, parsimony bootstrap values of > 50%,
and Bayesian posterior probabilities of > 0.95, respectively. The new species are marked with
asterisks.
Figure 1.
A maximum parsimony strict consensus tree, illustrating the phylogeny of four new species
and related genera in the order Russulales, based on nLSU sequences. The branches are labeled with
maximum likelihood bootstrap values of >70%, parsimony bootstrap values of >50%, and Bayesian
posterior probabilities of >0.95, respectively. The new species are marked with asterisks.

J. Fungi 2022,8, 1227 4 of 23
J. Fungi 2022, 8, x FOR PEER REVIEW 9 of 25
Figure 2. A maximum parsimony strict consensus tree, illustrating the phylogeny of two new
species and the related species in the genus Peniophora, based on the ITS sequences. The branches
are labeled with maximum likelihood bootstrap values higher than 70%, parsimony bootstrap
proportions that are higher than 50%, and Bayesian posterior probabilities of more than 0.95,
respectively.
Figure 2.
A maximum parsimony strict consensus tree, illustrating the phylogeny of two new species
and the related species in the genus Peniophora, based on the ITS sequences. The branches are labeled
with maximum likelihood bootstrap values higher than 70%, parsimony bootstrap proportions that
are higher than 50%, and Bayesian posterior probabilities of more than 0.95, respectively.

J. Fungi 2022,8, 1227 5 of 23
J. Fungi 2022, 8, x FOR PEER REVIEW 10 of 25
Figure 3. A maximum parsimony strict consensus tree illustrating the phylogeny of two new species
and related species in the genus Vararia, based on ITS sequences. The branches are labeled with a
maximum likelihood bootstrap value of > 70%, a parsimony bootstrap value of > 50%, and Bayesian
posterior probabilities of > 0.95, respectively.
3. Results
3.1. Molecular Phylogeny
The nLSU dataset (Figure 1) included sequences from 55 fungal specimens,
representing 55 species. The dataset had an aligned length of 1415 characters, of which
923 characters are constant, 152 are variable and parsimony-uninformative, and 340 are
parsimony-informative. The maximum parsimony analysis yielded one equally
parsimonious tree (TL = 860, CI = 0.3233, HI = 0.6767, RI = 0.6123, RC = 0.1979). The best
model for the ITS+nLSU dataset, which was 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 analysis and ML analysis resulted in a similar topology to the MP
analysis, with an average standard deviation of split frequencies = 0.009575 (BI); the
effective sample size (ESS) across the two runs is the double of the average ESS (avg ESS)
= 200.5. The phylogeny (Figure 1), based on the combined nLSU sequences, includes six
families within the order of Russulales, which indicated that nine genera, comprising
Asterostroma Massee, Baltazaria Leal-Dutra, Dentinger & G.W. Griff., Gloiothele Bres.,
Lachnocladium Lév., Michenera Berk. & M.A. Curtis, Peniophora, Scytinostroma Donk,
Vararia, and Vesiculomyces E. Hagstr. could be incorporated into the Peniophoraceae
family. Our current four new species can be clustered into the genera of Peniophora and
Vararia, respectively.
Figure 3.
A maximum parsimony strict consensus tree illustrating the phylogeny of two new species
and related species in the genus Vararia, based on ITS sequences. The branches are labeled with a
maximum likelihood bootstrap value of >70%, a parsimony bootstrap value of >50%, and Bayesian
posterior probabilities of >0.95, respectively.
Maximum parsimony (MP), maximum likelihood (ML), and Bayesian inference (BI)
analyses were applied to the three combined datasets, following the technique used in a
previous study [
42
], and the tree construction procedure was performed in PAUP*, version
4.0b10 [
46
]. All the characters were equally weighted, and gaps were treated as missing
data. Trees were inferred using the heuristic search option, with TBR branch swapping and
1000 random sequence additions. The max trees were set to 5000, branches of zero length
were collapsed, and all parsimonious trees were saved. Clade robustness was assessed
using a bootstrap (BT) analysis with 1000 replicates [
47
]. The descriptive tree statistics were
tree length (TL), consistency index (CI), retention index (RI), rescaled consistency index
(RC), and homoplasy index (HI); these were calculated for each maximum parsimonious
tree generated. The multiple sequence alignment was also analyzed using the maximum
likelihood (ML) in RAxML-HPC2, through the Cipres Science Gateway (www.phylo.org;
accessed on 13 October 2022) [
48
]. Branch support (BS) for the ML analysis was determined
by 1000 bootstrap replicates.

J. Fungi 2022,8, 1227 6 of 23
Table 1.
List of species, specimens, and GenBank accession numbers of the sequences used in
this study.
Species Name Specimen No. GenBank Accession No.
References Country
ITS nLSU
Amylostereum
areolatum NH 8041 AF506405 AF506405 [45] Sweden
A. chailletii NH 8031 AF506406 AF506406 [45] Sweden
A. laevigatum NH 12863 AF506407 AF506407 [45] Sweden
Asterostroma
bambusicola He 4132 KY263871 this publication Thailand
A. cervicolor He 2314 KY263869 this publication China
A. laxum EL 33-99 AF506410 AF506410 [45] Sweden
A. vararioides He 4140 KY263870 this publication Thailand
Auriscalpium
vulgare EL 33-95 AF506375 AF506375 [45] Sweden
Baltazaria galactina CBS: 752.86 MH873721 [49] France
B. neogalactina CBS: 755.86 MH873724 MH873724 [49] French
B. occidentalis AFTOL-ID DQ234539 [50] Canada
B. podocarpi Dai 9261 KJ583221 [51] China
Dentipratulum
bialoviesense GG 1645 AF506389 AF506389 [45] Sweden
Dichostereum
durum FG 1985 AF506429 AF506429 [45] Sweden
D. effuscatum GG 930915 AF506390 AF506390 [45] Sweden
Gloeocystidiellum
bisporum KHL 11135 AY048877 AY048877 [45] Sweden
G. clavuligerum FCUG 2159 AF310088 AF310088 [52] Spain
G. purpureum Wu 9310-45 AF441338 AF441338 [45] China
Gloeocystidiopsis
flammea CBS: 324.66 AF506437 AF506437 [45] C. African Rep.
Gloeodontia
columbiensis NH 11118 AF506444 AF506444 [45] Spain
G. discolor KHL 10099 AF506445 AF506445 [45] USA
G. eriobotryae Dai 12080 JQ349103 [53] China
G. pyramidata LR 15502 AF506446 AF506446 [45] Colombia
G. subasperispora KHL 8695 AF506404 AF506404 [45] Norway
G. yunnanensis SWFC 00010504 MN908254 [54] China
Gloeopeniophorella
convolvens KHL 10103 AF506435 AF506435 [45] USA
Gloiothele lactescens EL 8-98 AF506453 AF506453 [45] Sweden
G. lamellosa KHL 11031 AF506454 AF506454 [45] Venezuela
Heterobasidion
annosum 06129/6 KJ583225 [51] Russia
H. parviporum 04121/3 KJ583226 [51] Finland
Lachnocladium
schweinfurthianum KM 49740 MH260051 [38] Cameroon
Lactarius leonis SJ 91016 AF506411 AF506411 [45] Sweden
Lentinellus
cochleatus KGN 960928 AF506417 AF506417 [45] Sweden
L. ursinus EL 73-97 AF506419 AF506419 [45] USA
L. vulpinus KGN 980825 (GB) AF347097 AF347097 [45] Sweden
Megalocystidium
luridum KHL 8635 AF506422 AF506422 [45] Norway
Michenera artocreas GHL-2016-Oct MH204692 [55] USA
M. incrustata He 2630 MH142907 [55] China
Peniophora albobadia
CBS: 329.66 MH858809 MH858809 [49] France
P. bicornis He 3609 MK588763 MK588763 [39] China
P. bicornis He 4767 MK588764 MK588764 [39] China
P. borbonica He 4597 MK588766 MK588766 [39] China
P. borbonica He 4606 MK588765 MK588765 [39] China

J. Fungi 2022,8, 1227 7 of 23
Table 1. Cont.
Species Name Specimen No. GenBank Accession No.
References Country
ITS nLSU
P. cinerea B 1020 MN475151 MN475151 [39] USA
P. crassitunicata CBS: 663.91 MH862292 MH862292 [49] France
P. duplex B 1022 MN475153 MN475153 [39] USA
P. erikssonii CBS: 287.58 MH857788 MH857788 [39] France
P.erikssonii Cui 11871 MK588771 MK588811 [39] China
P. exima B 1011 MN475155 MN475155 [39] USA
P. exima T-523 MK588772 MK588772 [39] USA
P. fasticata CBS: 942.96 MH862624 MH862624 [39] Ethiopia
P. fissilis CBS: 681.91 MH862298 MH862298 [39] France
P. fissilis CBS: 684.91 MH862299 MH862299 [39] Netherlands
P. gabonensis CBS: 673.91 MH862293 MH862293 [39] Gabon
P. gilbertsonii CBS: 357.95 MH862528 MH862528 [39] USA
P. gilbertsonii CBS: 360.95 MH862530 MH862530 [39] USA
P. halimi CBS: 862.84 MH861843 MH861843 [39] France
P. incarnata NH 10271 AF506425 AF506425 [45] Denmark
P. incarnata CBS: 430.72 MH860518 MH872230 [39] Netherlands
P. junipericola He 2462 MK588773 MK588773 this publication China
P. laeta CBS: 256.56 MH857617 MH857617 [39] France
P. laeta CBS: 255.56 MH857616 MH857616 [39] France
P. laxitexta LGMF 1159 JX559580 [39] Brazil
P. laxitexta BAFC 3309 FJ882040 [39] Argentina
P. laxitexta BAFC: 4687 MN518328 [39] Argentina
P. lilacea CBS: 337.66 MH858813 MH858813 [39] Armenia
P. lycii CBS: 264.56 MH857624 MH857624 [39] France
P. lycii CBS: 261.56 MH857621 MH857621 [39] France
P. malaiensis He 4870 MK588775 MK588775 [39] China
P. manshurica He 2956 MK588776 MK588776 [39] China
P. manshurica He 3729 MK588777 MK588777 [39] China
P. meridionalis CBS: 289.58 MH857789 MH857789 [49] France
P. molesta CBS: 678.91 MH862296 MH862296 [39] Cote d’Ivoire
P. molesta CBS: 676.91 MH862294 MH862294 [39] Gabon
P. molesta CBS: 677.91 MH862295 MH862295 [39] Gabon
P. monticola CBS: 649.91 MH862289 MH862289 [39] France
P. nuda AFTOL-ID 660 DQ411533 [39] USA
P. nuda LZ15-07 MT859929 MT859929 this publication China
P. ovalispora CBS: 653.91 MH873971 [39] Netherlands
P. ovalispora CBS: 653.91 MH862290 MH862290 [39] Netherlands
P. parvocystidiata CBS: 716.91 MH862305 MH862305 [39] France
P. parvocystidiata CBS: 717.91 MH862306 MH862306 [39] France
P. piceae B 1010 MN475158 MN475158 this publication USA
P. pilatiana CBS: 269.56 MH857627 MH857627 [39] France
P. pilatiana CBS: 265.56 MH857625 MH857625 [39] France
P. pilatiana CBS: 266.56 MH857626 MH857626 [39] France
P. pini CBS: 273.56 MH857631 MH857631 [39] France
P. pini CBS: 270.56 MH857628 MH857628 [39] France
P. pithya CBS: 275.56 MH857633 MH857633 [49] France
P. polygonia He 3668 MH669233 [56] China
P. polygonia CBS: 404.50 MH856684 MH856684 [39] France
P. proxima CBS: 406.50 MH856686 MH856686 [39] France
P. proxima CBS: 405.50 MH856685 MH856685 [39] France
P. pseudonuda FCUG 2384 GU322866 this publication Sweden
P. pseudonuda FCUG 2390 GU322865 this publication Sweden
P. pseudopini B 1024 MN475163 MN475163 this publication USA
P. pseudoversicolor CBS: 125881 MH864303 MH864303 [39] France
P. quercina CBS: 407.50 MH856687 MH868204 [39] France
P. quercina CBS: 408.50 MH856688 MH856688 [39] France
P. quercina CBS: 409.50 MH856689 MH856689 [39] France

J. Fungi 2022,8, 1227 8 of 23
Table 1. Cont.
Species Name Specimen No. GenBank Accession No.
References Country
ITS nLSU
P. reidii CBS: 397.83 MH861616 MH861616 [39] France
P. rosealba CLZhao 3513 ON786559 OP380690 present study China
P. rosealba CLZhao 9401 * ON786560 present study China
P. rufa B 1014 MN475165 MN475165 this publication USA
P. rufa CBS: 351.59 MH857891 MH869432 [39] Canada
P. rufomarginata CBS: 281.56 MH857639 MH857639 [39] France
P. rufomarginata CBS: 282.56 MH857640 MH857640 [39] France
P. septentrionalis CBS: 294.58 MH857791 MH857791 [39] Canada
P. simulans CBS: 875.84 MH861850 MH861850 [39] France
P. simulans CBS: 874.84 MH861849 MH861849 [39] France
P. subsalmonea CBS: 697.91 MH862303 MH862303 [39] Netherlands
P. subsalmonea CBS: 696.91 MH862302 MH862302 [39] Netherlands
P. taiwanensis Wu 9209-14 MK588794 MK588794 [39] China
P. tamaricicola CBS: 438.62 MH858203 MH858203 [39] Morocco
P. tamaricicola CBS: 439.62 MH858204 MH858204 [39] Morocco
P. tamaricicola CBS: 441.62 MH858205 MH858205 [39] Morocco
P. versicolor CBS: 358.61 MH858082 MH858082 [39] Morocco
P. violaceolivida CBS: 348.52 MH857077 MH857077 [39] France
P. yunnanensis CLZhao 3978 OP380617 OP380689 present study China
P. yunnanensis CLZhao 7347 * OP380616 present study China
P. yunnanensis CLZhao 8135 OP380615 present study China
Russula violacea SJ 93009 AF506465 AF506465 [45] Sweden
Scytinostroma
portentosum EL 11-99 AF506470 AF506470 [45] Sweden
Sistotrema
brinkmannii NH 11412 AF506473 AF506473 [45] Turkey
S. coronilla NH 7598 AF506475 AF506475 [45] Canada
Stereum hirsutum NH 7960 AF506479 AF506479 [45] Romania
Vararia abortiphysa CBS: 632.81 MH861387 MH861387 [49] Gabon
V. ambigua CBS: 634.81 MH861388 MH873137 [49] France
V. amphithallica CBS: 687.81 MH861431 MH861431 [49] France
V. aurantiaca CBS: 642.81 MH861394 MH861394 [49] Gabon
V. aurantiaca CBS: 641.81 MH861393 MH861393 [49] France
V. breviphysa CBS: 644.81 MH861396 MH861396 [49] Gabon
V. calami CBS: 646.81 MH861398 MH861398 [49] France
V. calami CBS: 648.81 MH861399 MH861399 [49] France
V. callichroa CBS: 744.91 MH874000 MH874000 [49] France
V. cinnamomea CBS: 642.84 MH873488 MH873488 [49] Madagascar
V. cinnamomea CBS: 641.84 MH861794 MH861794 [49] Madagascar
V. cremea CBS: 651.81 MH873147 MH873147 [49] France
V. daweishanensis CLZhao 17911 OP380613 OP615103 present study China
V. daweishanensis CLZhao 17936 * OP380614 OP380688 present study China
V. dussii CBS: 655.81 MH861405 MH861405 [49] France
V. dussii CBS: 652.81 MH873148 MH873148 [49] France
V. ellipsospora HHB-19503 MW740328 MW740328 this publication New Zealand
V. fragilis CLZhao 2628 OP380611 present study China
V. fragilis CLZhao 16475 *OP380612 OP380687 present study China
V. fusispora PDD: 119539 OL709443 OL709443 this publication New Zealand
V. gallica CBS: 234.91 MH862250 [49] Canada
V. gallica CBS: 656.81 MH861406 MH873152 [49] France
V. gillesii CBS: 660.81 MH873153 MH873153 [49] Cote d’Ivoire
V. gomezii CBS: 661.81 MH873154 MH873154 [49] French
V. gracilispora CBS: 664.81 MH861412 MH861412 [49] Gabon
V. gracilispora CBS: 663.81 MH861411 [49] Gabon
V. insolita CBS: 668.81 MH861413 MH861413 [49] France
V. intricata CBS: 673.81 MH861418 MH861418 [49] France

J. Fungi 2022,8, 1227 9 of 23
Table 1. Cont.
Species Name Specimen No. GenBank Accession No.
References Country
ITS nLSU
V. investiens FP-151122 MH971976 MH971977 [56] USA
V. malaysiana CBS: 644.84 MH873490 MH873490 [49] Singapore
V. minispora CBS: 682.81 MH861426 MH861426 [49] France
V. ochroleuca CBS: 465.61 MH858109 MH858109 [49] France
V. ochroleuca JS 24400 AF506485 AF506485 [45] Norway
V. parmastoi CBS: 879.84 MH861852 MH861852 [49] Uzbekistan
V. perplexa CBS: 695.81 MH861438 MH861438 [49] France
V. pectinata CBS: 685.81 MH861429 [49] Cote d’Ivoire
V. pirispora CBS: 720.86 MH862016 MH862016 [49] France
V. rhombospora CBS: 743.81 MH861470 MH861470 [49] France
V. rosulenta CBS: 743.86 MH862028 [49] France
V. rugosispora CBS: 697.81 MH861440 MH861440 [49] Gabon
V. sigmatospora CBS: 748.91 MH874001 MH874001 [49] Netherlands
V. sphaericospora CBS: 700.81 MH873185 MH873185 [49] Gabon
V. sphaericospora CBS: 703.81 MH861446 MH861446 [49] Gabon
V. trinidadensis CBS: 651.84 MH861803 MH861803 [49] Madagascar
V. trinidadensis CBS: 650.84 MH873495 MH873495 [49] Madagascar
V. tropica CBS: 704.81 MH861447 MH873189 [49] France
V. vassilievae UC2022892 KP814203 KP814203 this publication USA
V. verrucosa CBS 706.81 MH861449 MH861449 [49] France
Vesiculomyces
citrinus EL 53-97 AF506486 AF506486 [45] Sweden
* indicates the holotype.
MrModeltest 2.3 [
57
] was used to determine the best-fit evolution model for each
dataset, using Bayesian inference (BI), which was performed using MrBayes 3.2.7a, with a
GTR+I+G model of the DNA substitution and a gamma distribution rate variation across
the sites [
58
]. Four Markov chains were run for 2 runs, beginning from random starting
trees for 0.9 million generations for nLSU (Figure 1), for 1.5 million generations for ITS
(Figure 2) with trees, and 1 million generations for ITS (Figure 3) with trees, with the
parameters sampled every 1000 generations. The first one-quarter of all generations were
discarded as the burn-in. The majority rule consensus tree of all the remaining trees was
calculated. Branches were considered significantly supported if they received a maximum
likelihood bootstrap value (BS) > 70%, a maximum parsimony bootstrap value (BT) > 70%,
or Bayesian posterior probabilities (BPP) > 0.95.
3. Results
3.1. Molecular Phylogeny
The nLSU dataset (Figure 1) included sequences from 55 fungal specimens, rep-
resenting 55 species. The dataset had an aligned length of 1415 characters, of which
923 characters are constant, 152 are variable and parsimony-uninformative, and 340 are
parsimony-informative. The maximum parsimony analysis yielded one equally parsimo-
nious tree (
TL = 860
, CI = 0.3233, HI = 0.6767, RI = 0.6123, RC = 0.1979). The best model
for the ITS+nLSU dataset, which was 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 analysis and ML analysis resulted in a similar topology to the MP analysis, with
an average standard deviation of split frequencies = 0.009575 (BI); the effective sample size
(ESS) across the two runs is the double of the average ESS (avg ESS) = 200.5. The phylogeny
(Figure 1), based on the combined nLSU sequences, includes six families within the order
of Russulales, which indicated that nine genera, comprising Asterostroma Massee, Baltazaria
Leal-Dutra, Dentinger & G.W. Griff., Gloiothele Bres., Lachnocladium Lév., Michenera Berk. &
M.A. Curtis, Peniophora,Scytinostroma Donk, Vararia, and Vesiculomyces E. Hagstr. could be

J. Fungi 2022,8, 1227 10 of 23
incorporated into the Peniophoraceae family. Our current four new species can be clustered
into the genera of Peniophora and Vararia, respectively.
The ITS-alone dataset of the genus Peniophora (Figure 2) included the sequences from
83 fungal specimens, representing 52 species. The dataset had an aligned length of 607 char-
acters, of which 353 characters were constant, while 64 were variable and parsimony-
uninformative, and 190 were parsimony-informative. The maximum parsimony analysis
yielded 12 equally parsimonious trees (TL = 1681, CI = 0.3111, HI = 0.6889,
RI = 0.4496
,
RC = 0.1399
). The best model for the ITS dataset that was estimated and applied in the
Bayesian analysis was GTR+I+G (lset nst = 6, rates = invgamma; prset statefreqpr = dirichlet
(1,1,1,1). Bayesian analysis and ML analysis resulted in a similar topology to MP analysis,
with an average standard deviation of split frequencies = 0.009599 (BI). The phyloge-
netic tree indicated that P. roseoalba can be grouped with two close taxa,
P.versicolor
and
P. ovalispora
, whereas P. yunnanensis can be grouped with a clade comprising P. lycii and
P.violaceolivida.
The ITS-only dataset of the genus Vararia (Figure 3) included sequences from 63 fungal
specimens, representing 39 species. The dataset had an aligned length of 1128 characters,
of which 511 characters were constant, 133 were variable and parsimony uninformative,
and 484 were parsimony informative. Maximum parsimony analysis yielded 6 equally
parsimonious trees (TL = 4589, CI = 0.2805, HI = 0.7195, RI = 0.4174, and RC = 0.1171).
The best model for the ITS dataset estimated and applied in the Bayesian analysis was
GTR+I+G. The Bayesian and ML analyses resulted in a similar topology to that of the
MP analysis with split frequencies = 0.0096082 (BI). The phylogram inferred from the ITS
sequences (Figure 3) revealed that Vararia daweishanensis could be grouped with four close
taxa: V.gomezii,V.rhombospora,V.sigmatospora, and V.trinidadensis, whereas the other
species of V.fragilis could be grouped with a clade comprising V. ambigua and V. ellipsospora,
with a low level of support.
3.2. Taxonomy
Peniophora roseoalba L. Zou & C.L. Zhao, sp. nov. (Figures 4and 5).
MycoBank no.: 845758.
Holotype—
China, Yunnan Province, Puer, Jingdong county, the Forest of Pineap-
ple, 24
◦
37
0
N, 100
◦
45
0
E, altitude 2083 m asl., on the fallen branch of an angiosperm,
4 January 2019, CLZhao 9401 (SWFC).
Etymology—Roseoalba
(Lat.): referring to the rose to pale pinkish grey color of the
hymenial surface of the specimens.
Fruiting body—
Basidiomata are annual, resupinate, membranaceous, without odor
and taste when fresh, up to 90 mm long, 20 mm wide, 70–100
µ
m thick. The hymenial
surface is smooth, occasionally cracked, and rose to pale pinkish grey. The sterile margin is
indistinct and is rose to pinkish grey.
Hyphal system—
Monomitic, generative hyphae with clamp connections, colorless,
thin- to thick-walled, moderately branched, 1.5–4.5
µ
m in diameter, CB
−
, IKI
−
; tissues
unchanged in KOH.
Hymenium—
The cystidia are of two types: (1) Gloeocystidia is subcylindrical to
conical, smooth, colorless, thin-walled, 31.5–40.5
×
6.5–7.5
µ
m; (2) Lamprocystidia is
abundant in the hymenium, and is conical, thick-walled, encrusted apical part, colorless,
33–42.5
×
7–10.5
µ
m. The Basidia are subclavate to subcylindrical, slightly constricted in
the middle, with four sterigmata and a basal clamp connection, sized 24–39.5
×
3.5–5.5
µ
m.
Basidiospores—
Basidiospores are ellipsoid, colorless, thin-walled, smooth, IKI
−
,
CB−, 4–6.5 ×3–5 µm, L = 5.19 µm, W = 3.8 µm, Q = 1.26–1.48 (n= 60/2).
Additional specimen examined—
China, Yunnan Province, Puer, Jingdong County,
Wuliangshan National Nature Reserve, 23
◦
57
0
N, 100
◦
22
0
E, altitude 3376 m asl., found on
the fallen branch of an angiosperm, 2 October 2017, CLZhao 3513 (SWFC).

J. Fungi 2022,8, 1227 11 of 23
J. Fungi 2022, 8, x FOR PEER REVIEW 12 of 25
Figure 4. Basidiomata of Peniophora roseoalba (holotype): the front of the basidiome (A); the
characteristic hymenophore (B). Bars: (A) = 1 cm and (B) = 1 mm.
Figure 4.
Basidiomata of Peniophora roseoalba (holotype): the front of the basidiome (
A
); the character-
istic hymenophore (B). Bars: (A)=1cmand(B) = 1 mm.

J. Fungi 2022,8, 1227 12 of 23
J. Fungi 2022, 8, x FOR PEER REVIEW 13 of 25
Figure 5. Microscopic structures of the Peniophora roseoalba (holotype): basidiospores (A); basidia
(B); basidioles (C); lamprocystidia (D); subcylindrical to conical gloeocystidia (E); a section of the
hymenium (F). Bars: (A–F) = 10 µm.
Peniophora yunnanensis L. Zou & C.L. Zhao sp. nov. (Figures 6 and 7).
MycoBank no.: 845760.
Holotype—China, Yunnan Province, Chuxiong, Zixishan Forestry Park, 25°01′ N,
101°24′ E., altitude 2356 m asl., on an angiosperm stump, 2 July 2018, code: CLZhao 7347
(SWFC).
Etymology—Yunnanensis (Lat.): referring to the geographic provenance (Yunnan
Province) of the specimens.
Fruiting body—Basidiomata are annual, resupinate, and coriaceous, without odor
and taste when fresh, up to 100 mm long, 25 mm wide, and 70–100 µm thick. The hymenial
surface is tuberculate and is pale cream to cream. The sterile margin is indistinct and
slightly cream-colored.
Hyphal system—Monomitic, generative hyphae with clamp connections, colorless,
thin- to thick-walled, moderately branched, 2.5–3.5 µm in diameter, IKI−, CB−; tissues are
unchanged in the KOH; the subiculum generative hyphae are dense, with a subparallel
arrangement; the subhymenium is composed of strongly agglutinated vertical hyphae.
Figure 5.
Microscopic structures of the Peniophora roseoalba (holotype): basidiospores (
A
); basidia
(
B
); basidioles (
C
); lamprocystidia (
D
); subcylindrical to conical gloeocystidia (
E
); a section of the
hymenium (F). Bars: (A–F) = 10 µm.
Peniophora yunnanensis L. Zou & C.L. Zhao sp. nov. (Figures 6and 7).
MycoBank no.: 845760.
Holotype—
China, Yunnan Province, Chuxiong, Zixishan Forestry Park, 25
◦
01
0
N,
101
◦
24
0
E., altitude 2356 m asl., on an angiosperm stump, 2 July 2018, code: CLZhao
7347 (SWFC).
Etymology—Yunnanensis
(Lat.): referring to the geographic provenance (Yunnan
Province) of the specimens.
Fruiting body—
Basidiomata are annual, resupinate, and coriaceous, without odor
and taste when fresh, up to 100 mm long, 25 mm wide, and 70–100
µ
m thick. The hymenial
surface is tuberculate and is pale cream to cream. The sterile margin is indistinct and
slightly cream-colored.
Hyphal system—
Monomitic, generative hyphae with clamp connections, colorless,
thin- to thick-walled, moderately branched, 2.5–3.5
µ
m in diameter, IKI
−
, CB
−
; tissues are
unchanged in the KOH; the subiculum generative hyphae are dense, with a subparallel
arrangement; the subhymenium is composed of strongly agglutinated vertical hyphae.
Hymenium—
The cystidia are of two types: (1) Gloeocystidia, which are different in
shape, conical, clavate to fusiform, and subglobose, usually containing refractive materials;
they are colorless, smooth, thin-walled, and 12.5–58
×
5.5–15.5
µ
m; (2) Lamprocystidia
are abundant in the hymenium, the conical, thick-walled, encrusted apical part, colorless,

J. Fungi 2022,8, 1227 13 of 23
and 18–29
×
4.5–7
µ
m. The basidia subclavate changes to subcylindrical, being slightly
constricted in the middle to somewhat constricted, with four sterigmata and a basal clamp
connection, 22.5–39.5 ×4.5–8 µm.
Basidiospores—
The basidiospores are subcylindrical, colorless, thin-walled, and
smooth, with oil drops occasionally found inside, IKI
−
, CB
−
, (5–) 5.5–10 (–11)
×
3–5.5
µ
m,
L = 7.72 µm, W = 4.44 µm, Q = 1.61–1.88 (n= 90/3).
Additional specimens examined (paratypes)—
China, Yunnan Province, Puer, Jing-
dong County, Taizhong Town, Ailaoshan, 24
◦
23
0
N, 120
◦
53
0
E, altitude 3166 m asl.; found
on the fallen branch of an angiosperm, 4 October 2017, CLZhao 3978 (SWFC). Zhenyuan
County, Ailaoshan, Jinshan Original Forestry, 24
◦
00
0
N, 101
◦
10
0
E; altitude 2300 m asl., and
found on the fallen branch of an angiosperm, 21 August 2018, CLZhao 8135 (SWFC).
J. Fungi 2022, 8, x FOR PEER REVIEW 15 of 25
Figure 6. Basidiomata of Peniophora yunnanensis (holotype): the front of the basidiomata (A); the
characteristic hymenophore (B). Bars: (A) = 1 cm and (B) = 1 mm.
Figure 6.
Basidiomata of Peniophora yunnanensis (holotype): the front of the basidiomata (
A
); the
characteristic hymenophore (B). Bars: (A)=1cmand(B) = 1 mm.

J. Fungi 2022,8, 1227 14 of 23
J. Fungi 2022, 8, x FOR PEER REVIEW 16 of 25
Figure 7. Microscopic structures of Peniophora yunnanensis (holotype): basidiospores (A); basidia and
basidioles (B); lamprocystidia (C); the conical, clavate to fusiform, subglobose gloeocystidia (D); a
section of the hymenium (E). Bars: (A–E) = 10 µm.
Vararia daweishanensis L. Zou & C.L. Zhao, sp. nov. (Figures 8 and 9).
MycoBank no.: 845761.
Holotype—China, Yunnan Province, Honghe, Pinbian County, Daweishan National
Forestry Park, 22°53′ N, 103°35′ E, altitude 1670 m asl., found on a fallen angiosperm
branch, 1 August 2019, CLZhao 17936 (SWFC).
Etymology—daweishanensis (Lat.): referring to the provenance (Daweishan) of the
specimens.
Fruiting body—Basidiomata are annual, resupinate, membranous, soft, and adnate,
up to 80 mm long, 16 mm wide, and 70–150 µm thick. The hymenial surface is smooth and
pale yellowish. The sterile margin is distinct, narrow, whitish, and attached.
Hyphal system—Dimitic, generative hyphae with clamp connections, colorless, thin-
to thick-walled, occasionally branched, interwoven, 2–4 µm in diameter, IKI−, CB+, tissues
Figure 7.
Microscopic structures of Peniophora yunnanensis (holotype): basidiospores (
A
); basidia and
basidioles (
B
); lamprocystidia (
C
); the conical, clavate to fusiform, subglobose gloeocystidia (
D
); a
section of the hymenium (E). Bars: (A–E) = 10 µm.
Vararia daweishanensis L. Zou & C.L. Zhao, sp. nov. (Figures 8and 9).
MycoBank no.: 845761.
Holotype—
China, Yunnan Province, Honghe, Pinbian County, Daweishan National
Forestry Park, 22
◦
53
0
N, 103
◦
35
0
E, altitude 1670 m asl., found on a fallen angiosperm
branch, 1 August 2019, CLZhao 17936 (SWFC).
Etymology—daweishanensis
(Lat.): referring to the provenance (Daweishan) of the
specimens.
Fruiting body—
Basidiomata are annual, resupinate, membranous, soft, and adnate,
up to 80 mm long, 16 mm wide, and 70–150
µ
m thick. The hymenial surface is smooth and
pale yellowish. The sterile margin is distinct, narrow, whitish, and attached.
Hyphal system—
Dimitic, generative hyphae with clamp connections, colorless, thin-
to thick-walled, occasionally branched, interwoven, 2–4
µ
m in diameter, IKI
−
, CB+, tissues
are unchanged in KOH; dichohyphae in subhymenium abundant, yellowish, capillary,
distinctly thick-walled; dichotomously to irregularly branched, with the main branches up
to 4
µ
m in diameter and with acute tips, moderately dextrinoid when in Melzer’s reagent;

J. Fungi 2022,8, 1227 15 of 23
more frequently branched with more narrow and shorter branches in the hymenium, with
slightly curved tips and a stronger dextrinoid reaction.
Hymenium—
The gloeocystidia are empty or are filled with a refractive oil-like matter;
they are also subcylindrical. The hymenium is elliptical to ovoid, smooth, colorless, thin-
walled, and 9–23
×
7–10.5
µ
m. The basidia are subcylindrical, with four sterigmata and a
basal clamp connection, 26–46 ×5–8 µm.
Basidiospores—
The basidiospores are allantoid, colorless, thin-walled, and smooth,
with oil droplets inside, IKI
−
, CB
−
, (8.5–) 9–13 (–14)
×
3.5–5
µ
m, L = 10.57
µ
m,
W = 4.23 µm
,
Q = 2.44–2.55 (n= 60/2).
Additional specimens examined (paratypes)—
China, Yunnan Province, Honghe,
Pinbian County, Daweishan National Forestry Park, 22
◦
53
0
N, 103
◦
35
0
E, altitude 1670 m
asl., found on a fallen angiosperm branch, 1 August 2019, CLZhao 17911 (SWFC).
J. Fungi 2022, 8, x FOR PEER REVIEW 17 of 25
are unchanged in KOH; dichohyphae in subhymenium abundant, yellowish, capillary,
distinctly thick-walled; dichotomously to irregularly branched, with the main branches
up to 4 µm in diameter and with acute tips, moderately dextrinoid when in Melzer’s
reagent; more frequently branched with more narrow and shorter branches in the
hymenium, with slightly curved tips and a stronger dextrinoid reaction.
Hymenium—The gloeocystidia are empty or are filled with a refractive oil-like
matter; they are also subcylindrical. The hymenium is elliptical to ovoid, smooth,
colorless, thin-walled, and 9–23 × 7–10.5 µm. The basidia are subcylindrical, with four
sterigmata and a basal clamp connection, 26–46 × 5–8 µm.
Basidiospores—The basidiospores are allantoid, colorless, thin-walled, and smooth,
with oil droplets inside, IKI−, CB−, (8.5–) 9–13 (–14) × 3.5–5 µm, L = 10.57 µm, W = 4.23
µm, Q = 2.44–2.55 (n = 60/2).
Additional specimens examined (paratypes)—China, Yunnan Province, Honghe,
Pinbian County, Daweishan National Forestry Park, 22°53′ N, 103°35′ E, altitude 1670 m
asl., found on a fallen angiosperm branch, 1 August 2019, CLZhao 17911 (SWFC).
Figure 8.
Basidiomata of the Vararia daweishanensis (holotype): the front of the basidiomata (
A
); the
characteristic hymenophore (B). Bars: (A)=1cmand(B) = 1 mm.

J. Fungi 2022,8, 1227 16 of 23
J. Fungi 2022, 8, x FOR PEER REVIEW 18 of 25
Figure 8. Basidiomata of the Vararia daweishanensis (holotype): the front of the basidiomata (A); the
characteristic hymenophore (B). Bars: (A) = 1 cm and (B) = 1 mm.
Figure 9. Microscopic structures of Vararia daweishanensis (holotype): basidiospores (A); basidia and
basidioles (B); subcylindrical, elliptical to ovoid gloeocystidia (C); dichohyphae (D); a section of the
hymenium (E). Bars: (A–E) = 10 µm.
Vararia fragilis L. Zou & C.L. Zhao, sp. nov. (Figures 10 and 11).
MycoBank no.: 845763.
Holotype—China, Yunnan Province, Wenshan, Wenshan National Nature Reserve.
GPS coordinates: found at 23°22′ N, 104°43′ E, altitude 1500 m asl., found on the fallen
branch of an angiosperm, 26 July 2019, CLZhao 16475 (SWFC).
Etymology—fragilis (Lat.): referring to the fragile basidiomata.
Fruiting body—Basidiomata are annual, resupinate, adnate, thin, membranous, and
fragile, without odor and taste when fresh, up to 85 mm long, 40 mm wide, and 30–100
µm thick. The hymenial surface is smooth, buff when fresh, buff to ochraceous on drying
and cracking. The sterile margin is indistinct, attached, and is cream to buff.
Hyphal system—Dimitic, generative hyphae, bearing simple septa, colorless, thin-
to thick-walled, occasionally branched, interwoven, 1.5–3.5 µm in diameter, IKI−, CB+,
and with tissues unchanged in KOH; the dichohyphae in the subhymenium are abundant,
Figure 9.
Microscopic structures of Vararia daweishanensis (holotype): basidiospores (
A
); basidia and
basidioles (
B
); subcylindrical, elliptical to ovoid gloeocystidia (
C
); dichohyphae (
D
); a section of the
hymenium (E). Bars: (A–E) = 10 µm.
Vararia fragilis L. Zou & C.L. Zhao, sp. nov. (Figures 10 and 11).
MycoBank no.: 845763.
Holotype—
China, Yunnan Province, Wenshan, Wenshan National Nature Reserve.
GPS coordinates: found at 23
◦
22
0
N, 104
◦
43
0
E, altitude 1500 m asl., found on the fallen
branch of an angiosperm, 26 July 2019, CLZhao 16475 (SWFC).
Etymology—fragilis (Lat.): referring to the fragile basidiomata.
Fruiting body—
Basidiomata are annual, resupinate, adnate, thin, membranous, and
fragile, without odor and taste when fresh, up to 85 mm long, 40 mm wide, and 30–100
µ
m
thick. The hymenial surface is smooth, buff when fresh, buff to ochraceous on drying and
cracking. The sterile margin is indistinct, attached, and is cream to buff.
Hyphal system—
Dimitic, generative hyphae, bearing simple septa, colorless, thin- to
thick-walled, occasionally branched, interwoven, 1.5–3.5
µ
m in diameter, IKI
−
, CB+, and
with tissues unchanged in KOH; the dichohyphae in the subhymenium are abundant, pre-
dominantly yellowish, capillary, distinctly thick-walled, and dichotomously to irregularly
branched, with the main branches up to 2
µ
m in diameter and with acute tips; moderately

J. Fungi 2022,8, 1227 17 of 23
dextrinoid in Melzer’s reagent; more frequently branched, with more narrow and shorter
branches in the subiculum, with slightly curved tips and a stronger dextrinoid reaction.
Hymenium—
The gloeocystidia are of two types: (1) elliptical to ovoid,
5.8–16 ×3.5–7 µm
;
(2) subulate, usually with a constriction at the tip, smooth, colorless, thin-walled,
16.5–27
×
4–7
µ
m. Basidia subcylindrical, with four sterigmata and a basal simple-septa
connection, 13–23.5 ×3–4.5 µm.
Basidiospores—
The basidiospores are broad from ellipsoid to ellipsoid, colorless,
thin-walled, smooth, IKI
−
, CB
−
, 3.5–5.5 (–6)
×
2.5–3.5
µ
m, L = 4.78
µ
m, W = 3.12
µ
m,
Q = 1.48–1.56 (n= 60/2).
Additional specimen examined (paratype)—
China, Yunnan Province, Yuxi, Xiping
County, Mopanshan National Forestry Park, 24
◦
07
0
N, 101
◦
98
0
E, altitude 2614 m asl., on
the fallen branch of an angiosperm, 20 August 2017, CLZhao 2628 (SWFC).
J. Fungi 2022, 8, x FOR PEER REVIEW 19 of 25
predominantly yellowish, capillary, distinctly thick-walled, and dichotomously to
irregularly branched, with the main branches up to 2 µm in diameter and with acute tips;
moderately dextrinoid in Melzer’s reagent; more frequently branched, with more narrow
and shorter branches in the subiculum, with slightly curved tips and a stronger dextrinoid
reaction.
Hymenium—The gloeocystidia are of two types: (1) elliptical to ovoid, 5.8–16 × 3.5–
7 µm; (2) subulate, usually with a constriction at the tip, smooth, colorless, thin-walled,
16.5–27 × 4–7 µm. Basidia subcylindrical, with four sterigmata and a basal simple-septa
connection, 13–23.5 × 3–4.5 µm.
Basidiospores—The basidiospores are broad from ellipsoid to ellipsoid, colorless,
thin-walled, smooth, IKI−, CB−, 3.5–5.5 (–6) × 2.5–3.5 µm, L = 4.78 µm, W = 3.12 µm, Q =
1.48–1.56 (n = 60/2).
Additional specimen examined (paratype)—China, Yunnan Province, Yuxi, Xiping
County, Mopanshan National Forestry Park, 24°07′ N, 101°98′ E, altitude 2614 m asl., on
the fallen branch of an angiosperm, 20 August 2017, CLZhao 2628 (SWFC).
Figure 10.
Basidiomata of Vararia fragilis (holotype): the front of the basidiomata (
A
); the characteristic
hymenophore (B). Bars: (A)=1cmand(B) = 1 mm.

J. Fungi 2022,8, 1227 18 of 23
J. Fungi 2022, 8, x FOR PEER REVIEW 20 of 25
Figure 10. Basidiomata of Vararia fragilis (holotype): the front of the basidiomata (A); the
characteristic hymenophore (B). Bars: (A) = 1 cm and (B) = 1 mm.
Figure 11. Microscopic structures of Vararia fragilis (holotype): basidiospores (A); basidia and
basidioles (B); fusiform gloeocystidia (C); elliptical to ovoid gloeocystidia (D); a section of the
hymenium (E); dichohyphae (F). Bars: (A) = 5 µm, (B–F) = 10 µm.
4. Discussion
Four genera, Gloiothele, Peniophora, Scytinostroman, and Vararia have been grouped
together and clustered within the family Peniophoraceae, as inferred from a dataset with
178 terminal taxa [37]. In the present study, based on the nLSU data (Figure 1), four new
species were classified in the family Peniophoraceae and were then classified within the
genera of Peniophora and Vararia.
Based on the ITS phylogenetic analysis (Figure 2), two new taxa have been grouped
within the genus Peniophora, named P. roseoalba and P. yunnanensis, in which P. roseoalba
is grouped with two close taxa, P. versicolor and P. ovalispora; P. yunnanensis was grouped
with a clade comprising P. lycii and P. violaceolivida. However, morphologically,
Peniophora versicolor differs from P. roseoalba by its dark brown to reddish brown or
ochraceous hymenophore, smaller lamprocystidia (10–20 × 8–10 µm), and larger
basidiospores (9–11 × 4.5–6 µm) [59]. P. ovalispora is separated from P. roseoalba by having
a cream-colored to salmon or brownish hymenophore, with a pruinose margin [13,60].
Peniophora lycii is separated from P. yunnanensis by its even, greyish lilac to bluish
violaceous hymenial surface, the presence of the dendrohyphidia and the wider
lamprocystidia (22–42 × 14–25 µm) [60]; P. violaceolivida differs in terms of its pale pink,
with a violaceous hymenial surface and a fimbriate margin [60].
In the current study, based on the further ITS phylogenetic tree (Figure 3), two new
taxa have been grouped within the genus Vararia. These are V. daweishanensis and V.
fragilis, in which V. daweishanensis was grouped with four close taxa, namely, V. gomezii,
V. rhombospora, V. sigmatospora and V. trinidadensis, while V. fragilis was grouped with a
Figure 11.
Microscopic structures of Vararia fragilis (holotype): basidiospores (
A
); basidia and basidi-
oles (
B
); fusiform gloeocystidia (
C
); elliptical to ovoid gloeocystidia (
D
); a section of the hymenium
(E); dichohyphae (F). Bars: (A)=5µm, (B–F) = 10 µm.
4. Discussion
Four genera, Gloiothele,Peniophora,Scytinostroman, and Vararia have been grouped
together and clustered within the family Peniophoraceae, as inferred from a dataset with
178 terminal taxa [
37
]. In the present study, based on the nLSU data (Figure 1), four new
species were classified in the family Peniophoraceae and were then classified within the
genera of Peniophora and Vararia.
Based on the ITS phylogenetic analysis (Figure 2), two new taxa have been grouped
within the genus Peniophora, named P.roseoalba and P. yunnanensis, in which P. roseoalba is
grouped with two close taxa, P. versicolor and P. ovalispora;P. yunnanensis was grouped with a
clade comprising P. lycii and P. violaceolivida. However, morphologically, Peniophora versicolor
differs from P. roseoalba by its dark brown to reddish brown or ochraceous hymenophore,
smaller lamprocystidia (10–20
×
8–10
µ
m), and larger basidiospores (
9–11 ×4.5–6 µm
) [
59
].
P. ovalispora is separated from P. roseoalba by having a cream-colored to salmon or brownish
hymenophore, with a pruinose margin [
13
,
60
]. Peniophora lycii is separated from P. yun-
nanensis by its even, greyish lilac to bluish violaceous hymenial surface, the presence of
the dendrohyphidia and the wider lamprocystidia (22–42
×
14–25
µ
m) [
60
]; P. violaceo-
livida differs in terms of its pale pink, with a violaceous hymenial surface and a fimbriate
margin [60].
In the current study, based on the further ITS phylogenetic tree (Figure 3), two new
taxa have been grouped within the genus Vararia. These are V. daweishanensis and V.
fragilis, in which V. daweishanensis was grouped with four close taxa, namely, V. gomezii,V.
rhombospora,V. sigmatospora and V. trinidadensis, while V. fragilis was grouped with a clade
comprising V. ambigua and V. ellipsospora. However, morphologically speaking, V. gomezii
differs from V. daweishanensis in having a pinkish buff to cream hymenial surface and
simple-septate generative hyphae, as well as navicular basidiospores [
20
]. V. rhombospora
is separated from V. daweishanensis by having a fragile basidiomata with a cream to beige
gray hymenial surface, with rhomboid and larger basidiospores (15–17
×
5–6
µ
m) [
61
];

J. Fungi 2022,8, 1227 19 of 23
V. sigmatospora is distinguishable from V. daweishanensis by its simple-septate generative
hyphae and fusiform, narrower basidiospores (13–15.2
×
2.5–3
µ
m) [
62
]; V. trinidadensis
differs in its gray to grayish-white hymenial surface, simple-septate generative hyphae, and
fusiform, narrower basidiospores (13–17
×
2.5–3.2
µ
m) [
63
]. V. ambigua is distinguishable
from V. fragilis by its powdery hymenial surface, as well as by basidia that are swollen at
the base and larger basidia (27–40
×
3.5–4
µ
m) [
21
]; V. ellipsospora differs from V. fragilis in
its fimbriate margin, clamped generative hyphae, wider gloeocystidia (28–48
×
8–11
µ
m)
and larger basidiospores (8–12 ×5.5–6.5 µm) [22].
Morphologically, Peniophora cinerea (Pers.) Cooke, P. laeta (Fr.) Donk, P. laurentii S.
Lundell, P. polygonia (Pers.) Bourdot & Galzin, P. rhodocarpa Rehill & B.K. Bakshi are
similar to P. roseoalba by having encrusted lamprocystidia. However, P. cinerea differs
from P. roseoalba by its smaller lamprocystidia (15–20
×
6–10
µ
m), and subcylindrical to
allantoid basidiospores [
60
]; P. laeta is separated from P. roseoalba by having a hydnoid
to raduloid hymenophore, larger gloeocystidia (60–120
×
8–10
µ
m) and cylindrical to
suballantoid, larger basidiospores (9–15
×
3.5–4.5
µ
m) [
64
]; P. laurentii is distinguished
from P. roseoalba by tuberculate to plicate or merulioid hymenophore, white margin, simple-
septa generative hyphae, as well as longer gloeocystidia (70–150
×
8–12
µ
m) and larger
basidia (50–60
×
6–8
µ
m) [
60
]; P. polygonia is separated from P. roseoalba by having bladder
like, bigger gloeocystidia (60–100
×
15–25
µ
m), presence of dendrohyphidia, and cylindrical
to allantoid, larger basidiospores (10–14
×
2.5–4
µ
m) [
60
]; P. rhodocarpa differs P. roseoalba
by having tuberculate, rimose hymenial surface, larger gloeocystidia (50–90
×
12–18
µ
m)
with larger lamprocystidia (60–100
×
12–18
µ
m), and allantoid, narrower basidiospores
(5–8.5 ×1.7–2.2 µm) [60].
Peniophora yunnanensis is similar to P. aurantiaca (Bres.) Höhn. & Litsch., P. bonariensis
C.E. Gómez, P. junipericola J. Erikss., P. meridionalis Boidin, P. quercina (Pers.) Cooke, based
on having clamped generative hyphae and gloeocystidia. However, Peniophora aurantiaca
is distinguished from P. yunnanensis by its orange-red, reddish to reddish grey hymenial
surface, larger gloeocystidia (70–150
×
10–20
µ
m), larger basidia (60–80
×
10–15
µ
m), and
ellipsoid, larger basidiospores (14–20
×
8–12
µ
m) [
60
]; P. bonariensis can be delimited from
P. yunnanensis by its pinkish grey to greyish violaceous hymenial surface, thick-walled
gloeocystidia and larger lamprocystidia (30–50
×
12–25
µ
m) [
60
]; P. junipericola differs
by having pinkish or greyish red to violaceous hymenial surface, larger lamprocystidia
(
40–80 ×6–18 µm
), and allantoid basidiospores [
60
]; P. meridionalis differs from P. yunnanen-
sis by its ochraceous grey, yellowish brown hymenial surface, presence of dendrohyphidia,
and larger lamprocystidia (35–55
×
8–20
µ
m) [
60
]; P. quercina is separated from P. yunna-
nensis by having the pinkish to pinkish grey or bluish grey to violaceous hymenial surface,
and larger lamprocystidia (30–80 ×10–20 µm) [60].
Peniophora yunnanensis resembles P. gilbertsonii Boidin, P. lilacea Bourdot & Galzin, P.
limitata (Chaillet ex Fr.) Cooke, P. piceae (Pers.) J. Erikss. and P. rufomarginata (Pers.) Bourdot
& Galzin in having a tuberculate hymenial surface. However, Peniophora gilbertsonii is
different from P. yunnanensis in having an ochraceous pink to reddish or brown to grey
hymenial surface and the presence of dendrohyphidia [
60
]; P. lilacea can be delimited from P.
yunnanensis along its pinkish grey to ochraceous violaceous hymenial surface, along with its
thick-walled gloeocystidia in trauma. We recorded the presence of the dendrohyphidia and
ellipsoid, wider basidiospores (9–16
×
6.5–10
µ
m) [
60
]; P. limitata differs from P. yunnanensis
by having pinkish gray or violaceous gray to a dark blue-gray hymenial surface, and wider
lamprocystidia (25–60
×
8–12
µ
m) [
19
]; P. piceae is distinguished from P. yunnanensis by
its reddish grey to grey to a dark violaceous grey hymenial surface, larger lamprocystidia
(40–80
×
6–18
µ
m), and allantoid, narrower basidiospores (6.5–9.5
×
2–2.8
µ
m) [
60
]; P.
rufomarginata is separated from P. yunnanensis by having a pinkish to pinkish gray or
bluish gray hue to the violaceous hymenial surface, along with larger lamprocystidia
(30–80 ×10–20 µm) [60].
Vararia amphithallica Boidin, Lanq. & Gilles, V. bispora S.L. Liu & S.H. He, V. montana
S.L. Liu & S.H. He, V. ochroleuca (Bourdot & Galzin) Donk and V. rugosispora Boidin, Lanq.

J. Fungi 2022,8, 1227 20 of 23
& Gilles resembles V. daweishanensis by having a smooth hymenial surface and clavate to
cylindrical basidia. However, Vararia amphithallica is distinguished from V. daweishanensis
by its fimbriate margin, 2-sterigmata basidia, and ellipsoid to cylindrical basidiospores
(
9–12 ×4–7 µm
) [
31
]; V. bispora differs in V. daweishanensis by having the thick-walled
gloeocystidia, with 2-sterigmata basidia, and larger, fusiform to cylindrical basidiospores
(
16–24 ×6–8 µm
) [
31
]; V. montana is separated from V. daweishanensis by having the brittle
basidiomata, longer gloeocystidia (50–100
×
4–9
µ
m), and broadly ellipsoid, larger ba-
sidiospores (16–24
×
8–14
µ
m) [
31
]; V. ochroleuca differs from V. daweishanensis by having
cream-colored to pallid ochraceous hymenial surface, slightly thick-walled gloeocystidia,
simple-septa generative hyphae, and broadly ellipsoid, to drop-shaped, smaller basid-
iospores (2.6–3.8
×
2–3.2
µ
m) [
65
]. V. rugosispora can be delimited from V. daweishanensis by
its simple-septate generative hyphae and longer basidiospores (12–16 ×7–8 µm) [21].
Vararia breviphysa Boidin & Lanq., V. cinnamomea Boidin, Lanq. & Gilles, V. cremea
Boidin, Lanq. & Gilles, V. gallica (Bourdot & Galzin) Boidin, V. hauerslevii Boidin, and
V. sinapicolor Boidin & Gilles are similar to V. fragilis, based on characteristics such as
the thick-walled dichohyphae, and four sterigmata basidia. However, V. breviphysa dif-
fers from V. fragilis by having the larger gloeocystidia (50–65
×
6–8.5
µ
m), fusiform and
larger basidiospores (15–22
×
4–6
µ
m) [
20
]. V. cinnamomea is distinguished from V. fragilis
by its cinnamon hymenial surface, larger basidia (45–65
×
8–10
µ
m), and larger basid-
iospores (9–13
×
5–7.2
µ
m) [
25
]. V. cremea can be delimited from V. fragilis by the longer
gloeocystidia (40–90
×
7–15
µ
m), and larger basidiospores (15–20
×
2.7–3.5
µ
m) [
21
]. V.
gallica differs from V. fragilis in having a whitish hymenial surface and larger basidiospores
(
9–12 ×3.5–5 µm
) [
19
]. V. hauerslevii is separated from V. fragilis by its larger gloeocys-
tidia (50–60
×
7–9
µ
m) and subfusoid, larger basidiospores (10–15
×
3.5–4.5
µ
m) [
66
]. V.
sphaericospora differs from V. fragilis in having clamped generative hyphae, bigger basidia
(33–45 ×6–7 µm), and larger basidiospores (12.5–14 ×5.2–7 µm) [20,21,23].
The taxa of Peniophora and Vararia are typical examples of wood-rotting fungi, which
is an extensively studied family [
19
,
67
–
70
]. So far, several studies on new wood-decaying
fungi belonging to the Peniophora and Vararia from China have been reported [34,71–75].
Author Contributions:
Conceptualization, C.Z.; methodology, C.Z. and L.Z.; software, C.Z. and L.Z.;
validation, C.Z. and L.Z.; formal analysis, C.Z., X.Z., Y.D. and L.Z.; investigation, C.Z. and L.Z.;
resources, C.Z.; writing—original draft preparation, C.Z., X.Z. and L.Z.; writing—review and editing,
C.Z. and L.Z.; visualization, C.Z.; supervision, C.Z.; project administration, C.Z.; funding acquisition,
C.Z. All 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 Changlin Zhao, Yunnan Fundamental Research Project (Grant No.
202001AS070043) to Changlin Zhao, the High-level Talents Program of Yunnan Province (YNQR-
QNRC-2018-111) to Changlin Zhao.
Institutional Review Board Statement: Not applicable.
Informed Consent Statement: Not applicable.
Data Availability Statement:
Publicly available datasets were analyzed in this study. This data can be
found here: [https://www.ncbi.nlm.nih.gov/;https://www.mycobank.org/page/Simple%20names%
20search;http://purl.org/phylo/treebase, submission ID 28664; all accessed on 13 October 2022].
Conflicts of Interest: The authors declare no conflict of interest.
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