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Fungi
Journal of
Article
Morphological and Phylogenetic Evidence for Recognition of
Two New Species of Phanerochaete from East Asia
Dong-Qiong Wang 1,2 and Chang-Lin Zhao 1,2,3,4,*
Citation: Wang, D.-Q.; Zhao, C.-L.
Morphological and Phylogenetic
Evidence for Recognition of Two New
Species of Phanerochaete from East
Asia. J. Fungi 2021,7, 1063. https://
doi.org/10.3390/jof7121063
Academic Editors: Vladimír Antonín
and Hana Sevcikova
Received: 11 October 2021
Accepted: 9 December 2021
Published: 11 December 2021
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1Key Laboratory for Forest Resources Conservation and Utilization in the Southwest Mountains of China,
Ministry of Education, Southwest Forestry University, Kunming 650224, China;
fungiwangdongqiong@163.com
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
4School of Life Sciences, Tsinghua University, Beijing 100084, China
*Correspondence: fungi@swfu.edu.cn or fungichanglinz@163.com
Abstract:
Two new corticioid fungal species, Phanerochaete pruinosa and P. rhizomorpha spp. nov. are
proposed based on a combination of morphological features and molecular evidence.
Phanerochaete
pruinosa is characterized by the resupinate basidiomata with the pruinose hymenial surface, a monomitic
hyphal system with simple-septate generative hyphae and subcylindrical basidiospores measuring
as 3.5–6.7
×
1.5–2.7
µ
m. Phanerochaete rhizomorpha is characterized by having a smooth hymenophore
covered by orange hymenial surface, the presence of rhizomorphs, subulate cystidia, and narrower
ellipsoid to ellipsoid basidiospores. Sequences of ITS+nLSU nrRNA gene regions of the studied
specimens were generated and phylogenetic analyses were performed with maximum likelihood,
maximum parsimony, and Bayesian inference methods. These phylogenetic analyses showed that
two new species clustered into genus Phanerochaete, in which P. pruinosa was sister to P. yunnanensis
with high supports (100% BS, 100% BT, 1.00 BPP); morphologically differing by a pale orange to
greyish orange and densely cracked hymenial surface. Another species P. rhizomorpha was closely
grouped with P. citrinosanguinea with lower supports; morphologically having yellow to reddish
yellow hymenial surface, and smaller cystidia measuring as 31–48 ×2.3–4.8 µm.
Keywords: corticioid fungi; Phanerochaetaceae; molecular systematics; taxonomy; Yunnan Province
1. Introduction
Corticioid fungi is a large group of Basidiomycota with simpler basidiomata with the
diverse morphological features when compared with polypores, but the phylogenetic di-
versity of this group is less intensively studied [
1
,
2
]. In the subtropical–tropical areas, many
corticioid taxa have not been discovered and described worldwide. The genus Phanerochaete
P. Karst. is a member of the corticioid fungi, which is typified by P. alnea (Fr.) P. Karst. [
3
],
and the genus is characterized by the resupinate, membranaceous basidiomata with or
without rhizomorphs, a monomitic hyphal system with primarily simple-septate gener-
ative hyphae, clavate basidia with four sterigmata, and smooth, thin-walled, inamyloid
basidiospores [
1
,
4
,
5
]. Index Fungorum (http://www.indexfungorum.org; accessed on 9
December 2021) registers 187 specific and infraspecific names in Phanerochaete. The diversity
and taxonomy of Phanerochaete s.l. in China have been studied for the last 30 years [6–19].
Molecular studies involving Phanerochaete based on the ribosomal DNA (rDNA) se-
quences, revealed the phylogenetic distribution of resupinate forms across the major clades
of mushroom-forming fungi, in which P. chrysosporium Burds. nested into phlebioid clade
in Polyporales [
20
]. Revisiting the taxonomy of Phanerochaete (Polyporales, Basidiomy-
cota) using a four gene dataset and extensive ITS sampling indicated that Phanerochaete
sensu lato was polyphyletic and distributed across nine lineages in the phlebioid clade,
J. Fungi 2021,7, 1063. https://doi.org/10.3390/jof7121063 https://www.mdpi.com/journal/jof
J. Fungi 2021,7, 1063 2 of 14
in which six lineages were associated to described genera [
21
]. Miettinen et al. [
22
]. ex-
plored the DNA-phylogeny-based and morphology-based to reconcile the polypores and
genus concepts in the family Phanerochaetaceae, which the macromorphology of fruiting
bodies and hymenophore construction did not reflect monophyletic groups, and Ceriporia
inflata B.S. Jia and B.K. Cui was combined into Phanerochaete. Amplifying nrLSU, nrITS,
and rpb1 genes across the Polyporales revealed that eleven genera clustered into family
Phanerochaetaceae, and two families Hapalopilaceae and Bjerkanderaceae were placed as syn-
onyms of Phanerochaetaceae [
23
]. Recently, the research supported by morphological studies
and the phylogenetic analyses, showed that many new taxa of Phanerochaete s.s. were found
and displayed the taxonomic status for the new taxa within genus Phanerochaete [14,19].
In 2018–2019, we collected the material supposedly belonging to the two undescribed
species of corticioid fungi from Yunnan Province, China. We present the morphological
and molecular phylogenetic evidence that support the recognition of two new species
within the Phanerochaete s.s., based on the internal transcribed spacer (ITS) and regions
nLSU sequences.
2. Materials and Methods
2.1. Morphology
The fruiting bodies were observed growing on the ground of broad-leaved treemixed
forest. The fruiting bodies were dried in an electric food dehydrator at 40
◦
C, then sealed
and stored in an envelope bag. They were then transported to Kunming where microscopic
morphology and phylogeny to be studied at the mycology laboratory of Southwest Forestry
University, Kunming, Yunnan Province, China. The for-study specimens were deposited
at the herbarium of Southwest Forestry University (SWFC), Kunming, Yunnan Province,
China. Macromorphological descriptions were based on field notes and photos captured
in the field and lab. Color terminology follow Petersen [
24
]. Micromorphological data
were obtained from the dried specimens, and observed under a light microscope following
Dai [
25
]. The following abbreviations were used: KOH = 5% potassium hydroxide water
solution, CB = Cotton Blue, CB– = acyanophilous, IKI = Melzer’s reagent, IKI– = both
inamyloid and indextrinoid, L = mean spore length (arithmetic average for all spores),
W = mean spore width (arithmetic average for all spores), Q = variation in the L/W ratios
between the specimens studied, n= a/b (number of spores (a) measured from given
number of specimens (b)).
2.2. Molecular Phylogeny
CTAB rapid plant genome extraction kit-DN14 (Aidlab Biotechnologies Co., Ltd.,
Beijing, China) was used to obtain genomic DNA from dried specimens, according to the
manufacturer’s instructions followed previous study [
26
]. ITS region was amplified with
primer pair ITS5 and ITS4 [
27
]. nLSU region was amplified with primer pair LR0R and
LR7 (http://lutzonilab.org/nuclear-ribosomal-dna/; accessed on 28 September 2021). The
PCR procedure 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 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 directly sequenced
at Kunming Tsingke Biological Technology Limited Company, Kunming, Yunnan Province,
China. All newly generated sequences were deposited in NCBI GenBank (Table 1).
Table 1. List of species, specimens, and GenBank accession numbers of sequences used in this study.
Species Name Specimen No. GenBank Accession No.
References
ITS nLSU
Bjerkandera adusta FP-101236 KP134982 [21]
B. adusta HHB-12826 KP134983 KP135198 [21]
B. fumosa Dai 12674B MW507112 MW520213 [28]
J. Fungi 2021,7, 1063 3 of 14
Table 1. Cont.
Species Name Specimen No. GenBank Accession No.
References
ITS nLSU
B. fumosa Dai 21087 MW507110 [28]
Byssomerulius corium FP-102382 KP135007 KP135230 [21]
B. corium FP-107055 KP135008 [21]
Ceraceomyces serpens HHB-15692-Sp KP135031 KP135200 [21]
C. serpens L-11105 KP135032 [21]
Ceriporia purpurea KKN-223-Sp KP135044 KP135203 [21]
C. purpurea HHB-3964 KP135042 [21]
C. reticulata RLG-11354 KP135041 KP135204 [21]
C. reticulata L-7837 KP135040 [21]
Efibula gracilis FD-455 KP135027 MZ637116 [21]
E. gracilis FP-102052 KP135028 [21]
E. tropica Wu 0809-8 MZ636968 MZ637130 unpublished
E. tropica WEI 18-149 MZ636967 MZ637129 unpublished
Gloeoporus dichrous FP-151129 KP135058 KP135213 [21]
G. pannocinctus L-15726-Sp KP135060 KP135214 [21]
Hyphodermella poroides Dai 12045 KX008367 KX011852 [29]
H. poroides Dai 10848 KX008368 KX011853 [29]
H. rosae FP-150552 KP134978 KP135223 [21]
H. rosae MA-Fungi FN600389 JN939588 [30]
Irpex lacteus FD-9 KP135026 KP135224 [21]
I. lacteus FD-93 KP135025 [21]
Meruliopsis
albostramineus HHB-10729 KP135051 KP135229 [21]
M. albostramineus L-9778 KP135052 [21]
M. taxicola CBS 45548 MH856432 MH867978 [31]
M. taxicola Kuljok 00/75 (GB) EU118648 [32]
Phaeophlebiopsis
caribbeana HHB-6990 KP135415 KP135243 [21]
P. caribbeana FD-442 (TYPE) KP135416 [21]
P. ignerii FD-425 KP135418 [21]
P. peiophoroides FP-150577 KP135417 KP135273 [21]
Phanerochaete. albida FD-31 KP135308 KP135210 [19]
P. alnea OM 8110 KP135171 [21]
P. alnea KHL 12054 EU118653 EU118653 [32]
P. argillacea Wu 9712-18 GQ470656 [13]
P. arizonica RLG-10248-sp KP135170 KP135239 [21]
P. australis He 6013 MT235656 MT248136 [19]
P. australis HHB-7105-sp KP135081 KP135240 [21]
P. australosanguinea 20098 Tell MH233928 [33]
P. australosanguinea 20102 Tell MH233929 [33]
P. bambucicola He 3606 MT235657 MT248137 [19]
P. bambucicola Wu 0707-2 MF399404 MF399395 [15]
P. brunnea He 4192 MT235658 MT248138 [19]
P. brunnea He 1873 KX212220 KX212224 [17]
P. burdsallii He 2066 MT235690 MT248177 [19]
P. burdsallii CFMR: RF9JR KU668973 unpublished
P. burtii HHB-4618-sp KP135117 KP135241 [21]
P. burtii FD-171 KP135116 [21]
P. calotricha Vanhanen-382 KP135107 [21]
P. canobrunnea He 5726 MT235659 MT248139 [19]
P. canobrunnea CHWC 1506-66 LC412095 LC412104 [14]
P. canolutea Wu 9211-105 GQ470641 [13]
P. carnosa He 5172 MT235660 MT248140 [19]
P. carnosa HHB-9195 KP135129 KP135242 [21]
P. chrysosporium HHB-6251 KP135094 KP135246 [21]
P. chrysosporium He 5778 MT235661 MT248141 [19]
J. Fungi 2021,7, 1063 4 of 14
Table 1. Cont.
Species Name Specimen No. GenBank Accession No.
References
ITS nLSU
P. cinerea He 6003 MT248172 [19]
P. citrinosanguinea He 4298 MT235691 MT248178 [19]
P. citrinosanguinea FP-105385-sp KP135100 KP135234 [21]
P. concrescens He 4657 MT235662 MT248142 [19]
P. concrescens Spirin 7322 KP994380 KP994382 [34]
P. cumulodentata He 2995 MT235664 MT248144 [19]
P. cumulodentata LE 298935 KP994359 KP994386 [34]
P. cystidiata He 4224 MT235665 MT248145 [19]
P. cystidiata Wu 1708-326 LC412097 LC412100 [14]
P. deflectens FCUG 2777 GQ470644 [13]
P. ericina He 4285 MT235666 MT248146 [19]
P. ericina HHB-2288 KP135167 KP135247 [21]
P. exilis HHB-6988 KP135001 KP135236 [21]
P. fusca Wu 1409-163 LC412099 LC412106 [14]
P. hymenochaetoides He 5988 MT248173 [19]
P. incarnata He 20120728-1 MT235669 MT248149 [19]
P. incarnata WEI 16-075 MF399406 MF399397 [15]
P. inflata Dai 10376 JX623929 JX644062 [35]
P. inflata Cui 7712 JX623930 JX644063 [35]
P. krikophora HHB-5796 KP135164 KP135268 [21]
P. laevis He 20120917-8 MT235670 MT248150 [19]
P. laevis HHB-15519 KP135149 KP135249 [21]
P. leptocystidiata He 5853 MT235685 MT248168 [19]
P. leptocystidiata Dai 10468 MT235684 MT248167 [19]
P. livescens He 5010 MT235671 MT248151 [19]
P. livescens FD-106 KP135070 KP135253 [21]
P. magnoliae He 3321 MT235672 MT248152 [19]
P. magnoliae HHB-9829-sp KP135089 KP135237 [21]
P. metuloidea He 2565 MT248163 [19]
P. metuloidea He 2766 MT235682 MT248164 [19]
P. minor He 3988 MT235686 MT248170 [19]
P. minor He 3977 MT248169 [19]
P. parmastoi He 4570 MT235673 MT248153 [19]
P. parmastoi Wu 880313-6 GQ470654 [13]
P. porostereoides He 1902 KX212217 KX212221 [17]
P. pruinosa CLZhao 7712 MZ435346 MZ435350 Present study
P. pruinosa CLZhao 7713 MZ435347 MZ435351 Present study
P. pseudomagnoliae PP-25 KP135091 KP135250 [21]
P. pseudosanguinea FD-244 KP135098 KP135251 [21]
P. queletii HHB-11463 KP134994 KP135235 [21]
P. queletii FP-102166 KP134995 [21]
P. rhizomorpha CLZhao 10470 MZ435348 MZ435352 Present study
P. rhizomorpha CLZhao 10477 MZ435349 MZ435353 Present study
P. rhodella FD-18 KP135187 KP135258 [21]
P. robusta Wu 1109-69 MF399409 MF399400 [15]
P. robusta Ghobad 2288 KP127068 KP127069 [16]
P. sanguinea HHB-7524 KP135101 KP135244 [21]
P. sanguineocarnosa FD-359 KP135122 KP135245 [21]
P. sinensis He 4660 MT235688 MT248175 [19]
P. sinensis GC 1809-56 MT235689 MT248176 [19]
P. sordida He 5400 MT235676 MT248157 [19]
P. sordida FD-241 KP135136 KP135252 [21]
P. stereoides He 5824 MT235677 MT248158 [19]
P. subceracea FP-105974-R KP135162 KP135255 [21]
P. subrosea He 2421 MT235687 MT248174 [19]
P. taiwaniana He 5269 MT235680 MT248161 [19]
J. Fungi 2021,7, 1063 5 of 14
Table 1. Cont.
Species Name Specimen No. GenBank Accession No.
References
ITS nLSU
P. taiwaniana Wu 0112-13 MF399412 MF399403 [15]
P. thailandica 2015_07 MF467737 [36]
P. velutina He 3079 MT235681 MT248162 [19]
P. velutina Kotiranta 25567 KP994354 KP994387 [34]
P. xerophila HHB-8509-Sp KP134996 KP135259 [21]
P. xerophila KKN-172 KP134997 [21]
P. yunnanensis He 2719 MT235683 MT248166 [19]
P. yunnanensis He 2697 MT248165 [19]
Phlebiopsis flavidoalba FD-263 KP135402 KP135271 [21]
P. flavidoalba FD-374 KP135403 [21]
P. gigantea FP-70857-sp KP135390 KP135272 [21]
P. gigantea FP-101815 KP135389 [21]
Pirex concentricus OSC-41587 KP134984 KP135275 [21]
P. concentricus Kropp160Bup6-R KP134985 [21]
Rhizochaete filamentosa HHB-3169 KP135410 KP135278 [21]
R. filamentosa FP-105240 KP135411 [21]
R.radicata FD-123 KP135407 KP135279 [21]
Terana caerulea FP-104073 KP134980 KP135276 [21]
T. caerulea T-616 KP134981 [21]
Trametopsis aborigena Robledo 1238 KY655337 [37]
T. aborigena Robledo 1236 KY655336 [37]
T. cervina AJ-185 JN165020 JN164839 [21]
T. cervina AJ-189 JN165021 [21]
New sequences are shown in bold.
Sequences were aligned in MAFFT 7 (https://mafft.cbrc.jp/alignment/server/; ac-
cessed on 28 September 2021) using G-INS-i strategy for ITS+nLSU combined dataset, and
manually adjusted in BioEdit [
38
]. Aligned dataset was deposited in TreeBase (submission
ID 28442). Phlebiopsis gigantea Fr. and Rhizochaete radicata (Henn.) Gresl., Nakasone and
Rajchenb were selected as an outgroup for phylogenetic analyses of combined dataset
following a previous study [
19
]. The taxon sampling strategy for the selection of sequences
for phylogenetic trees was to choose (1) in a larger scale, focusing on the related genera
in the families Phanerochaetaceae and Irpicaceae in Figure 1; (2) the related taxa based on
BLAST search in GenBank within Phanerochaete s.l.; and (3) all species of Phanerochaete s.s.
Maximum parsimony analysis was applied to the combined (ITS+nLSU) dataset. Its
approaches followed Zhao and Wu [
26
], and the tree construction procedure was performed
in PAUP* version 4.0b10 [
39
]. 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 1000 random sequence additions. Max-trees were set to 5000, branches of
zero length were collapsed and all parsimonious trees were saved. Clade robustness was
assessed using bootstrap (BT) analysis with 1000 replicates [
40
]. 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. Datamatrix was also analyzed using Maximum Likelihood (ML) approach
with RAxML-HPC2 through the CIPRES Science Gateway (www.phylo.org; accessed
on 28 September 2021) [
41
]. Branch support (BS) for ML analysis was determined by
1000 bootstrap replicates.
MrModeltest 2.3 [
42
] was used to determine the best-fit evolution model for the
dataset for Bayesian inference (BI). BI was calculated with MrBayes 3.1.7a [
43
]. Four
Markov chains were run for 2 runs from random starting trees for 10 million generations
for ITS+nLSU (Figure 2). The first one-fourth of all generations was discarded as burn-in.
The majority rule consensus tree of all remaining trees was calculated. Branches were
considered as significantly supported if they received maximum likelihood bootstrap
J. Fungi 2021,7, 1063 6 of 14
value (BS) >70%, maximum parsimony bootstrap value (BT) >70%, or Bayesian posterior
probabilities (BPP) >0.95.
J. Fungi 2021, 7, x FOR PEER REVIEW 6 of 16
branch swapping and 1000 random sequence additions. Max-trees were set to 5000,
branches of zero length were collapsed and all parsimonious trees were saved. Clade ro-
bustness was assessed using bootstrap (BT) analysis with 1000 replicates [40]. Descriptive
tree statistics: tree length (TL), consistency index (CI), retention index (RI), rescaled con-
sistency index (RC), and homoplasy index (HI) were calculated for each Maximum Parsi-
monious Tree generated. Datamatrix was also analyzed using Maximum Likelihood (ML)
approach with RAxML-HPC2 through the CIPRES Science Gateway (www.phylo.org; ac-
cessed on 28 September 2021) [41]. Branch support (BS) for ML analysis was determined
by 1000 bootstrap replicates.
J. Fungi 2021, 7, x FOR PEER REVIEW 7 of 16
Figure 1. Maximum Parsimony strict consensus tree illustrating the phylogeny of two new species
and related genera in Phanerochaetaceae and Irpicaceae based on ITS+nLSU sequences. Branches
are labeled with maximum likelihood bootstrap values > 70%, parsimony bootstrap values > 70%
and Bayesian posterior probabilities > 0.95, respectively.
MrModeltest 2.3 [42] was used to determine the best-fit evolution model for the da-
taset for Bayesian inference (BI). BI was calculated with MrBayes 3.1.7a [43]. Four Markov
chains were run for 2 runs from random starting trees for 10 million generations for
ITS+nLSU (Figure 2). The first one-fourth of all generations was discarded as burn-in. The
majority rule consensus tree of all remaining trees was calculated. Branches were consid-
ered as significantly supported if they received maximum likelihood bootstrap value (BS)
> 70%, maximum parsimony bootstrap value (BT) > 70%, or Bayesian posterior probabili-
ties (BPP) > 0.95.
Figure 1.
Maximum Parsimony strict consensus tree illustrating the phylogeny of two new species
and related genera in Phanerochaetaceae and Irpicaceae based on ITS+nLSU sequences. Branches are
labeled with maximum likelihood bootstrap values >70%, parsimony bootstrap values >70% and
Bayesian posterior probabilities >0.95, respectively.
J. Fungi 2021,7, 1063 7 of 14
J. Fungi 2021, 7, x FOR PEER REVIEW 8 of 16
Figure 2. Maximum Parsimony strict consensus tree illustrating the phylogeny of two new species
and related species in Phanerochaete based on ITS+nLSU sequences. Branches are labeled with max-
imum likelihood bootstrap values > 70%, parsimony bootstrap values > 70% and Bayesian posterior
probabilities > 0.95, respectively. The yellow backgrounds indicate new species.
3. Results
3.1. Molecular Phylogeny
The ITS+nLSU dataset (Figure 1) included sequences from 86 fungal specimens rep-
resenting 50 species. The dataset had an aligned length of 2368 characters, of which 1170
Figure 2.
Maximum Parsimony strict consensus tree illustrating the phylogeny of two new species
and related species in Phanerochaete based on ITS+nLSU sequences. Branches are labeled with maxi-
mum likelihood bootstrap values >70%, parsimony bootstrap values >70% and Bayesian posterior
probabilities >0.95, respectively. The yellow backgrounds indicate new species.
3. Results
3.1. Molecular Phylogeny
The ITS+nLSU dataset (Figure 1) included sequences from 86 fungal specimens rep-
resenting 50 species. The dataset had an aligned length of 2368 characters, of which
1170 characters are constant, 598 are variable and parsimony-uninformative, and 600 are
parsimony-informative. Maximum parsimony analysis yielded one equally parsimonious
tree (TL = 3476, CI = 0.3631, HI = 0.6369, RI = 0.7539, RC = 0.3512). Best model for the
J. Fungi 2021,7, 1063 8 of 14
ITS+nLSU dataset 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 = 0.038487 (BI), and the effective sample size (ESS) across the two runs
is the double of the average ESS (avg ESS) = 303.
The phylogeny (Figure 1) based on the combined ITS+nLSU sequences indicated
that both species Phanerochaete pruinose and P. rhizomorpha clustered into Phanerochaete s.s
and then P. pruinose grouped with P. subceracea (Burt) Burds.; P. rhizomorpha was sister to
P. citrinosanguinea Floudas and Hibbett.
The ITS+nLSU dataset (Figure 2) included sequences from 83 fungal specimens
representing 53 taxa. The dataset had an aligned length of 2017 characters, of which
1548 characters are constant, 164 are variable and parsimony-uninformative, and 395 are
parsimony-informative. Maximum parsimony analysis yielded 35 equally parsimonious
trees (TL = 1900, CI = 0.4095, HI = 0.5905, RI = 0.6456, RC = 0.2644). Best model for the
ITS+nLSU dataset 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 = 0.004260 (BI), and the effective sample size (ESS) across the two runs
is the double of the average ESS (avg ESS) = 309.
The phylogram inferred from ITS+nLSU sequences (Figure 2) revealed that two new
species were clustered into genus Phanerochaete s.s.; P. pruinosa sp. nova. was sister to
P. yunnanensis Y.L. Xu and S.H. He with high supports (100% BS, 100% BT, 1.00 BPP),
and then grouped with P. robusta Parmasto without supported data. Another species
P. rhizomorpha sp. nova. was closely grouped with P. citrinosanguinea with lower supports,
and then grouped with P. pseudosanguinea Floudas and Hibbett (–BS, 98% BP and 1.00 BPP)
and P. sanguinea (Fr.) Pouzar (96% BS, 91% BP and 1.00 BPP).
3.2. Taxonomy
Phanerochaete pruinosa C.L. Zhao and D.Q. Wang, sp. nov. Figures 3and 4.
MycoBank no.: MB 841271.
Diagnosis: It differs from P. yunnanensis by its pruinose hymenophore with the white
to slightly cream hymenial surface and lightly darkening in KOH.
Holotype—China, Yunnan Province, Chuxiong, Zixishan National Forestry Park, on
the bark of fallen angiosperms, 101.4◦E, 25.1◦N, 1 July 2018, CLZhao 7113 (SWFC).
Etymology
—
pruinosa
(Lat.): from Latin, referring to the white powder on hymenial
surface of basidiomata.
Fruiting body
—Basidiomata annual, resupinate, adnate, undetachable from substrate,
membranaceous to coriaceous, without odor and taste when fresh, up to 15 cm long,
3 cm wide, 50–100
µ
m thick. Hymenial surface smooth to have small verrucous process,
pruinose, white when fresh, white to slightly cream on drying; lightly darkening in KOH.
Margin sterile, narrow, white, attached.
Hyphal system
—Hyphal system monomitic, generative hyphae simple-septa, color-
less, thick-walled, unbranched, interwoven, 3–4.5
µ
m in diameter, subhymenial hyphae
densely covered by larger crystals, basal hyphae regular; IKI–, CB–; tissues unchanged in
KOH.
Hymenium
—Hymenial cystidia and cystidoles absent; basidia clavate to subcylindri-
cal, with four sterigmata and a simple-septum, 13–24 µm×3.5–4.5 µm.
Spores
—Basidiospores cylindrical, colorless, thin-walled, smooth, IKI–, CB–, (3.3–)
3.5–6.7(–7) µm×1.5–2.7(–2.9) µm, L = 4.42 µm, W = 1.94 µm, Q = 2.21–2.35 (n = 60/2).
Additional specimen examined
—China, Yunnan Province, Zixishan National Forestry
Park, on fallen branch of angiosperm, 101.4
◦
E, 25.1
◦
N, 1 July 2018, C.L. Zhao 7112 (SWFC).
Habitat and ecology
—Climate of the sample collection site is monsoon humid, and
the forest type is evergreen broad-leaved forest, and the samples were collected on an
angiosperm branch.
J. Fungi 2021,7, 1063 9 of 14
J. Fungi 2021, 7, x FOR PEER REVIEW 10 of 16
Figure 3. Basidiomata of Phanerochaete pruinosa (holotype) Bars: (A) = 2 cm and (B) = 1 mm.
Etymology—pruinosa (Lat.): from Latin, referring to the white powder on hymenial
surface of basidiomata.
Fruiting body—Basidiomata annual, resupinate, adnate, undetachable from sub-
strate, membranaceous to coriaceous, without odor and taste when fresh, up to 15 cm
long, 3 cm wide, 50–100 µm thick. Hymenial surface smooth to have small verrucous pro-
cess, pruinose, white when fresh, white to slightly cream on drying; lightly darkening in
KOH. Margin sterile, narrow, white, attached.
Hyphal system—Hyphal system monomitic, generative hyphae simple-septa, color-
less, thick-walled, unbranched, interwoven, 3–4.5 µm in diameter, subhymenial hyphae
Figure 3. Basidiomata of Phanerochaete pruinosa (holotype) Bars: (A)=2cmand(B) = 1 mm.
J. Fungi 2021, 7, x FOR PEER REVIEW 11 of 16
densely covered by larger crystals, basal hyphae regular; IKI–, CB–; tissues unchanged in
KOH.
Hymenium—Hymenial cystidia and cystidoles absent; basidia clavate to subcylin-
drical, with four sterigmata and a simple-septum, 13–24 µm × 3.5–4.5 µm.
Spores—Basidiospores cylindrical, colorless, thin-walled, smooth, IKI–, CB–, (3.3–)
3.5–6.7(–7) µm × 1.5–2.7(–2.9) µm, L = 4.42 µm, W = 1.94 µm, Q = 2.21–2.35 (n = 60/2).
Figure 4. Microscopic structures of Phanerochaete pruinosa (holotype): basidiospores (A), basidia and
basidioles (B), A section of hymenium (C). Bars: (A) = 5 µm, (B,C) = 10 µm.
Additional specimen examined—China, Yunnan Province, Zixishan National For-
estry Park, on fallen branch of angiosperm, 101.4° E, 25.1° N, 1 July 2018, C.L. Zhao 7112
(SWFC).
Habitat and ecology—Climate of the sample collection site is monsoon humid, and
the forest type is evergreen broad-leaved forest, and the samples were collected on an
angiosperm branch.
Phanerochaete rhizomorpha C.L. Zhao and D.Q. Wang sp. nov. Figures 5 and 6.
MycoBank no.: MB 841272.
Diagnosis: It differs from P. citrinosanguinea by its orange hymenial surface and larger
cystidia 48.5–71.5 µm × 3–6.5 µm)
Figure 4.
Microscopic structures of Phanerochaete pruinosa (holotype): basidiospores (
A
), basidia and
basidioles (B), A section of hymenium (C). Bars: (A)=5µm, (B,C) = 10 µm.
J. Fungi 2021,7, 1063 10 of 14
Phanerochaete rhizomorpha C.L. Zhao and D.Q. Wang sp. nov. Figures 5and 6.
J. Fungi 2021, 7, x FOR PEER REVIEW 12 of 16
Figure 5. Basidiomata of Phanerochaete rhizomorpha (holotype) Bars: (A) = 2 cm and (B) = 1 mm.
Holotype—China, Yunnan Province, Dali, Nanjian Country, Lingbaoshan National
Forestry Park, on the fallen branch of angiosperm, 24.7° N, 100.6° E, 10 January 2019, C.L.
Zhao 10,477 (SWFC).
Etymology—rhizomorpha (Lat.): from Latin, referring to the rhizomorphic basidio-
mata of the specimens.
Fruiting body—Basidiomata annual, resupinate, adnate, easily detachable from sub-
strate, membranaceous, up to 5 cm long, 3 cm wide, 200–300 µm thick. Hymenial surface
smooth, slightly orange when fresh, orange upon drying; lightly darkening in KOH. Mar-
gin sterile, buff to slightly orange, up to 1 mm wide, rhizomorphic.
Figure 5. Basidiomata of Phanerochaete rhizomorpha (holotype) Bars: (A)=2cmand(B) = 1 mm.
J. Fungi 2021, 7, x FOR PEER REVIEW 13 of 16
Hyphal system—Hyphal system monomitic, generative hyphae simple-septa, color-
less, thick-walled, frequently branched, interwoven, 3–6.5 µm in diameter, basal hyphae
regular, numerous crystals present among the abhymenium hyphae, IKI–, CB–; tissues
unchanged in KOH.
Hymenium—Hymenium cystidia subulate or tapering, colorless, thick-walled, with
2–4 septa, 48.5–71.5 µm × 3–6.5 µm; basidia subcylindrical, with 4 sterigmata, 18.5–35.5
µm × 3.5–5.5 µm.
Spores—Basidiospores narrower ellipsoid to ellipsoid, colorless, thin-walled,
smooth, with oil 1–2 drops inside, IKI–, CB–, 4.5–5.8(–6) µm × 2.7–3.6(–3.8) µm, L = 5.07
µm, W = 3.19 µm, Q = 1.58–1.60 (n = 62/2).
Figure 6. Microscopic structures of Phanerochaete rhizomorpha (holotype): basidiospores (A), basidia
and basidioles (B), cystidia (C). A section of hymenium (D). Bars: (A) = 5 µm, (B–D) = 10 µm.
Additional specimen examined—China, Yunnan Province, Nanjian Country, Ling-
baoshan National Forestry Park, on fallen branch of angiosperm, 24.7° N, 100.6° E, 10 Jan-
uary 2019, C.L. Zhao 10,470 (SWFC).
Habitat and ecology—Climate of the sample collection site is a transition between
tropical and subtropical climate, and the forest type is the tropical monsoon evergreen
broad-leaved forest, and the samples were collected on an angiosperm trunk.
Figure 6.
Microscopic structures of Phanerochaete rhizomorpha (holotype): basidiospores (
A
), basidia
and basidioles (B), cystidia (C). A section of hymenium (D). Bars: (A)=5µm, (B–D) = 10 µm.
J. Fungi 2021,7, 1063 11 of 14
MycoBank no.: MB 841272.
Diagnosis: It differs from P. citrinosanguinea by its orange hymenial surface and larger
cystidia 48.5–71.5 µm×3–6.5 µm)
Holotype
—China, Yunnan Province, Dali, Nanjian Country, Lingbaoshan National
Forestry Park, on the fallen branch of angiosperm, 24.7
◦
N, 100.6
◦
E, 10 January 2019, C.L.
Zhao 10,477 (SWFC).
Etymology
—
rhizomorpha
(Lat.): from Latin, referring to the rhizomorphic basid-
iomata of the specimens.
Fruiting body
—Basidiomata annual, resupinate, adnate, easily detachable from sub-
strate, membranaceous, up to 5 cm long, 3 cm wide, 200–300
µ
m thick. Hymenial surface
smooth, slightly orange when fresh, orange upon drying; lightly darkening in KOH. Margin
sterile, buff to slightly orange, up to 1 mm wide, rhizomorphic.
Hyphal system
—Hyphal system monomitic, generative hyphae simple-septa, color-
less, thick-walled, frequently branched, interwoven, 3–6.5
µ
m in diameter, basal hyphae
regular, numerous crystals present among the abhymenium hyphae, IKI–, CB–; tissues
unchanged in KOH.
Hymenium
—Hymenium cystidia subulate or tapering, colorless, thick-walled, with
2–4 septa, 48.5–71.5
µ
m
×
3–6.5
µ
m; basidia subcylindrical, with 4 sterigmata, 18.5–35.5
µ
m
×3.5–5.5 µm.
Spores
—Basidiospores narrower ellipsoid to ellipsoid, colorless, thin-walled, smooth,
with oil 1–2 drops inside, IKI–, CB–, 4.5–5.8(–6)
µ
m
×
2.7–3.6(–3.8)
µ
m, L = 5.07
µ
m,
W = 3.19 µm, Q = 1.58–1.60 (n = 62/2).
Additional specimen examined
—China, Yunnan Province, Nanjian Country, Ling-
baoshan National Forestry Park, on fallen branch of angiosperm, 24.7
◦
N, 100.6
◦
E, 10 Jan-
uary 2019, C.L. Zhao 10,470 (SWFC).
Habitat and ecology
—Climate of the sample collection site is a transition between
tropical and subtropical climate, and the forest type is the tropical monsoon evergreen
broad-leaved forest, and the samples were collected on an angiosperm trunk.
4. Discussion
In the present study, two new species, Phanerochaete pruinosa C.L. Zhao and D.Q. Wang
and P.rhizomorpha C.L. Zhao and D.Q. Wang spp. nov., are described based on phylogenetic
analyses and morphological characters. The nucleotide differences of phylogenetically
similar species to Phanerochaete pruinosa and P.rhizomorpha.
Phylogenetically, Xu et al. [
19
] revealed the taxonomy and phylogeny of Phanerochaete
sensu stricto (Polyporales, Basidiomycota) with emphasis on Chinese collections, which
showed that twenty-eight species of Phanerochaete s.s. from China are confirmed by mor-
phology and DNA sequence data. In the present study (Figure 2), two new taxa clustered
into Phanerochaete s.s., in which P. pruinosa was sister to P. yunnanensis, and then grouped
with P. robusta. Another species P. rhizomorpha was closely grouped with P. citrinosanguinea
with lower supports, and then grouped with P. pseudosanguinea and P. sanguinea. However,
morphologically, P. yunnanensis is separated from P. pruinosa by having a pale orange to
greyish orange and densely cracked hymenial surface [
19
]; P. robusta differs in its yellow
basidiomata and two kinds of cystidia without encrustation, larger basidiospores (5.5–7
µ
m
×
2.4–2.9
µ
m) and a boreal distribution [
34
]. Phanerochaete citrinosanguinea differs from
P. rhizomorpha by having yellow to reddish yellow hymenial surface, and smaller cystidia
(31–48
µ
m
×
2.3–4.8
µ
m) [
21
]; P. pseudosanguinea differs P. rhizomorpha in its light red or dark
red hymenial surface, and narrower basidiospores (4–5.5
µ
m
×
2–2.5
µ
m) [
21
]; P. sanguinea
is separated from P. rhizomorpha by having the thin-walled cystidia and the larger basidia
(25–45 µm×4–6 µm) [1]; in addition, there is some coloration of wood as in P. sanguinea.
Morphologically, Phanerochaete pruinosa is similar to P. concrescens Spirin and Volobuev
and P. sordida (P. Karst.) J. Erikss. and Ryvarden, based on presence of white or cream
hymenial surface. However, P. concrescens differs from P. pruinosa by having the large
J. Fungi 2021,7, 1063 12 of 14
basidia (27–39
µ
m
×
4–5
µ
m) [
34
]; P. sordida is separated from P. pruinosa by presence of
cystidia and wider basidiospores (5–7 µm×2.5–3.5 µm) [1].
Phanerochaete rhizomorpha reminds four taxa of Phanerochaete based on the character
by having the rhizomorph, P. burdsallii Y.L. Xu, Nakasone and S.H. He, P. leptocystidiata
Y.L. Xu and S.H. He, P. sinensis Y.L. Xu, C.C. Chen and S.H. He and P. subrosea Y.L. Xu and
S.H. He. However, P.burdsallii differs from P. rhizomorpha by having the cystidia encrusted
with small crystals [
19
]; P. leptocystidiata differs in having a tuberculate hymenial surface
and thin-walled cystidia encrusted at apex (24–30
µ
m
×
4–6
µ
m) [
19
]; P. sinensis differs
in its thin-walled cystidia and the shorter basidia (17–22
µ
m
×
4–5
µ
m) [
19
]; P. subrosea is
separated from P. rhizomorpha by having the thin-walled and smaller cystidia (33–55
µ
m
×
3–5 µm) [19].
Phanerochaete rhizomorpha is similar to P. aurantiobadia Ghob.-Nejh., S.L. Liu, Langer
and Y.C. Dai, P. cumulodentata (Nikol.) Parmasto and P. hymenochaetoides Y.L. Xu and S.H.
He based on the character by the orange hymenial surface. However, P. aurantiobadia
differs from P. rhizomorpha by having the larger basidiospores (5–8.3
µ
m
×
2–3
µ
m) [
16
];
P. cumulodentata differs from P. rhizomorpha by a tuberculate hymenophore and shorter
basidia (16.7–28.3
µ
m
×
3.7–5.2
µ
m) [
34
]; P. hymenochaetoides differs from P. rhizomorpha
by having both smaller cystidia (30–45
µ
m
×
3–4
µ
m) and basidiospores (4–5.2
µ
m
×
2–2.8 µm) [19].
In the ecology and biogeography, the taxa of Phanerochaete are a typical example of
wood-rotting fungi, which are mainly distributed in Asia, Europe, and America, and the
substrata are mostly hardwood [
1
,
25
], and this genus is an extensively studied group
of Basidiomycota; nonetheless, the wood-rotting fungi diversity is still not well known
in the subtropics and tropics [
44
–
48
]. The two new species, Phanerochaete pruinosa and
P.rhizomorpha spp. nov., were found in subtropics, which enriches the diversity of wood-
rotting fungi.
Author Contributions:
Conceptualization, C.-L.Z. and D.-Q.W.; methodology, C.-L.Z. and D.-Q.W.;
software, D.-Q.W.; validation, C.-L.Z. and D.-Q.W.; formal analysis, D.-Q.W.; investigation, D.-Q.W.;
resources, C.-L.Z.; data curation, D.-Q.W.; writing—original draft preparation, C.-L.Z. and D.-Q.W.;
writing—review and editing, C.-L.Z. and D.-Q.W.; visualization, D.-Q.W.; supervision, C.-L.Z.;
project administration, C.-L.Z.; funding acquisition, C.-L.Z. All authors have read and agreed to the
published version of the manuscript.
Funding:
The research was supported by the the National Natural Science Foundation of China
(Project No. 32170004), Yunnan Fundamental Research Project (Grant No. 202001AS070043) and the
High-level Talents Program of Yunnan Province (YNQR-QNRC-2018-111).
Institutional Review Board Statement: Not applicable for studies involving humans or animals.
Informed Consent Statement: Not applicable for studies involving humans.
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/;https://www.treebase.
org/treebase-web/home.html;jsessionid=6440D6056D96C04A8D29290992C18900, submission ID
28442; accessed on 16 November 2021].
Conflicts of Interest: The authors declare no conflict of interest.
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