Front. Cell. Infect. Microbiol. Frontiers in Cellular and Infection Microbiology Front. Cell. Infect. Microbiol. 2235-2988 Frontiers Media S.A. 10.3389/fcimb.2023.1105918 Cellular and Infection Microbiology Original Research Two new corticioid species of Phanerochaetaceae (Polyporales, Basidiomycota) from Southwest China Zhang Qiu-Yue 1 † Liu Zhan-Bo 1 † Liu Hong-Gao 2 Si Jing 1 * 1 Institute of Microbiology, School of Ecology and Nature Conservation, Beijing Forestry University, Beijing, China 2 Faculty of Agronomy and Life Sciences, Zhaotong University, Zhaotong, Yunnan, China

Edited by: Jia-Jia Chen, Jiangsu Vocational College of Agriculture and Forestry, China

Reviewed by: James A. Fraser, The University of Queensland, Australia; Haixia Ma, Chinese Academy of Tropical Agricultural Sciences, China

*Correspondence: Jing Si, jingsi1788@126.com

†These authors have contributed equally to this work and share first authorship

This article was submitted to Fungal Pathogenesis, a section of the journal Frontiers in Cellular and Infection Microbiology

 03 02 2023 2023 13 1105918 23 11 2022 18 01 2023 Copyright © 2023 Zhang, Liu, Liu and Si 2023 Zhang, Liu, Liu and Si

This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.

Two new corticioid fungi in the family Phanerochaetaceae, Phanerochaete shenghuaii and Rhizochaete variegata, are described and illustrated from Southwest China based on morphological characteristics and molecular data. Phanerochaete shenghuaii is characterized by annual, effused, inseparable basidiocarps from substrate, ivory white to cream hymenial surface when juvenile, buff to yellowish brown with age, buff in KOH, a monomitic hyphal system, smooth cystidia, and ellipsoid basidiospores measuring 4.8–6 × 2.5–3.8 µm. Rhizochaete variegata is characterized by annual, effused, easily separable basidiocarps from substrate, buff-yellow to clay-pink fresh hymenial surface becoming cream to buff upon drying, violet in KOH, a monomitic hyphal system, encrusted cystidia, and ellipsoid basidiospores measuring 3–4 × 2.2–3 µm. The phylogenetic analyses based on ITS + nLSU rDNA sequences confirm the placement of the two new species, respectively, in the Phanerochaete clade and the Rhizochaete clade of Phanerochaetaceae. Phylogenetically related and morphologically similar species to these two new species are discussed.

 new taxa phlebioid clade phylogeny taxonomy wood-decaying fungi 32270016, 32070016 National Natural Science Foundation of China10.13039/501100001809 Introduction

A phlebioid clade is a large group of Polyporales, comprising three families (Phanerochaetaceae Jülich, Irpicaceae Spirin & Zmitr., and Meruliaceae Rea), which accommodates massive corticioid fungi (Wu et al., 2010; Dai, 2011; Justo et al., 2017; He et al., 2019). Most members of the phlebioid clade are saprotrophs on dead wood, causing white rot, which plays an essential role in the maintenance of forest ecosystems (Justo et al., 2017; Ryvarden and Melo, 2017). However, compared with the antrodia and core polyporoid fungi in Polyporales, the phlebioid clade, especially corticioid fungi, has not been intensively studied, with some corticioid genera being known as paraphyletic or polyphyletic, and their members are scattered in different lineages, not fully consistent with the morphological features (Ortiz-Santana et al., 2013; Justo et al., 2017; Cui et al., 2019).

Phanerochaete P. Karst., established based on P. velutina (DC.) P. Karst., is the largest corticioid genus with more than 100 described species in Phanerochaetaceae (Burdsall, 1985; Kirk et al., 2008; Wu et al., 2010; Ghobad-Nejhad et al., 2015). The genus has a worldwide distribution and is characterized by white-rot, resupinate, and membranaceous basidiocarps; smooth or tuberculate hymenial surface; a monomitic hyphal system; generative hyphae mostly simple septate; the presence of smooth or encrusted cystidia; and thin-walled, non-amyloid, and acyanophilous basidiospores (Wu, 2000; Wu et al., 2010; Floudas and Hibbett, 2015; Ghobad-Nejhad et al., 2015). The diversity and taxonomy of Phanerochaete s.l. in China have been studied for 30 years (Wu, 1990; Wu, 1995; Wu, 1998; Wu, 2000; Wu, 2004; Wu, 2007; Xiong and Dai, 2009; Wu et al., 2010; Ghobad-Nejhad et al., 2015; Liu and He, 2016; Chen et al., 2018; Wu et al., 2018a; Wu et al., 2018b). Early studies focused on fungi of Taiwan Province and were mostly based solely on morphology. Recent studies have confirmed that the genus is highly polyphyletic and its species are distributed throughout the phlebioid clade, comprising a number of Phanerochaete species assembled in a highly supported clade, referred to as the core Phanerochaete clade, containing the type P. velutina (Wu et al., 2010; Floudas and Hibbett, 2015; Justo et al., 2017; Chen et al., 2021).

Rhizochaete is a small genus introduced by Greslebin et al. (2004), based on R. brunnea Gresl. et al., as a segregate of Phanerochaete, differing mainly by the reaction of basidiocarps and rhizomorphs (hyphal cords) with KOH: basidiocarps of Rhizochaete become red or violet in KOH, while they keep unchanged in Phanerochaete. Rhizochaete is characterized by resupinate, loosely adnate basidiocarps, with smooth to tuberculate hymenophore, usually turning red to violet in KOH, a monomitic hyphal system with simple septa or clamp connections, cylindrical to ellipsoid basidiospores, usually non-amyloid and acyanophilous (Nakasone et al., 2017; Gu and Zhao, 2021). Since Rhizochaete was erected, the number of newly named species is increasing continuously. Based on studying the parenthesome structure of some corticioid fungi, Bianchinotti et al. (2005) reported that three Rhizochaete species had perforate septal dolipore caps or parenthesomes. Nakasone et al. (2017) described a new species of Rhizochaete from Belize and transferred three additional species to the genus based on morphological and molecular data. Gu and Zhao (2021) reported two new species based on a combination of morphological features and molecular evidence. So far, approximately 17 species have been accepted in Rhizochaete worldwide (Greslebin et al., 2004; Chikowski et al., 2016; Nakasone et al., 2017; Gu and Zhao, 2021). Recently, a family-level classification of Polyporales or phlebioid fungi has shown that the genus Rhizochaete nested within Phanerochaetaceae, grouped with Hapalopilus P. Karst., Phaeophlebiopsis Floudas & Hibbett, and Phlebiopsis Jülich (Greslebin et al., 2004; Wu et al., 2010; Ghobad-Nejhad et al., 2015; Chen et al., 2021; Zhao et al., 2021).

During investigations on the diversity of wood-rotting fungi from China, four unknown corticioid specimens were collected from Southwest China, and their morphology corresponded to the concepts of Phanerochaete and Rhizochaete. To confirm their affinity, phylogenetic analyses based on the internal transcribed spacer (ITS) and nLSU rDNA sequences were carried out. Both morphological characteristics and molecular evidence demonstrated that these four corticioid specimens represent two new species of Phanerochaetaceae. So, we describe them in the present paper.

 Materials and methods Morphological studies

The studied specimens are deposited in the herbarium of the Institute of Microbiology, Beijing Forestry University (BJFC). Macro-morphological descriptions are based on field notes and measurements of herbarium specimens. Micro-morphological data and drawings are obtained from the dried specimens and observed under a light microscope following Chen et al. (2021) and Wu et al. (2022b). Color terms followed Petersen (1996). Sections were studied at a magnification up to ×1,000 using a Nikon Eclipse 80i microscope with phase contrast illumination (Nikon, Tokyo, Japan). Drawings were made with the aid of a drawing tube. Microscopic features, measurements, and drawings were made from slide preparations stained with Cotton Blue and Melzer’s reagent. Basidiospores were measured from sections cut from the hymenophore. To present the variation of basidiospores size, 5% of measurements were excluded from each end of the range and are given in parentheses. The following abbreviations are used: IKI = Melzer’s reagent; IKI− = neither amyloid nor dextrinoid; KOH = 5% potassium hydroxide; CB = Cotton Blue; CB− = acyanophilous; L = arithmetic average of all basidiospores length; W = arithmetic average of all basidiospores width; Q = variation in the L/W ratios between the specimens studied, (n = x/y) = the number of basidiospores (x) measured from a given number of specimens (y).

 DNA extraction and sequencing

A cetyltrimethylammonium bromide (CTAB) rapid plant genome extraction kit (Aidlab Biotechnologies, Co., Ltd., Beijing, China) was used to extract DNA (Wu et al., 2020). The following primer pairs were used to amplify the DNA: ITS5 (5′‐GGA AGT AAA AGT CGT AAC AAG G‐3′) and ITS4 (5′‐TCC TCC GCT TAT TGATAT GC‐3′) for the ITS regions (White et al., 1990); LR0R (5′‐ACC CGC TGA ACT TAA GC‐3′) and LR7 (5′‐TAC TAC CAC CAA GAT CT‐3′) for nuclear large subunit rDNA (nLSU) (Vilgalys and Hester, 1990). The PCR products were purified with a Gel Extraction and PCR Purification Combo Kit (Spin-column) at Beijing Genomics Institute (BGI), China. The purified products were then sequenced on an ABI-3730-XL DNA Analyzer (Applied Biosystems, Foster City, CA, USA) using the same primers as in the original PCR amplifications. All newly generated sequences were submitted to GenBank and are listed in Table 1 .

Taxa information and GenBank accession numbers of sequences used in this study.

Species Specimen no. Locality ITS nLSU Literature
Bjerkandera adusta HHB-12826-Sp Alaska, United States KP134983 KP135198 Justo et al. (2017)
B. centroamericana L-13104-sp Costa Rica KY948791 KY948855 Wu et al. (2010)
Hapalopilus eupatorii Dammrich 10744 Germany KX752620 KX752620 Miettinen et al. (2016)
H. nidulans JV0206/2 Sweden KX752623 KX752623 Miettinen et al. (2016)
H. percoctus Miettinen 2008 Botswana KX752597 KX752597 Miettinen et al. (2016)
Phaeophlebiopsis caribbeana HHB-6990 United States KP135415 KP135243 Floudas and Hibbett (2015)
P. himalayensis He 3854 Hainan, China MT386378 MT447410 Zhao et al. (2021)
P. peniophoroides FP-150577 United States KP135417 KP135273 Floudas and Hibbett (2015)
P. ravenelii CBS 411.5 France MH856691 MH868208 Vu et al. (2019)
P. ravenelii FCUG 2216 France – GQ470674 Wu et al. (2010)
Phanerochaete aculeata GC 1703-117 Taiwan, China MZ422785 MZ637177 Chen et al. (2021)
P. aculeata Wu 880701-2 Taiwan, China MZ422787 GQ470636 Chen et al. (2021)
P. albida GC 1407-14 Taiwan, China MZ422788 MZ637179 Chen et al. (2021)
P. albida WEI 18-365 Taiwan, China MZ422789 MZ637180 Chen et al. (2021)
P. allantospora KKN-111-Sp Arizona, United States KP135038 KP135238 Chen et al. (2021)
P. allantospora RLG-10478* Arizona, United States KP135039 – Chen et al. (2021)
P. alnea Larsson 12054 (GB) Norway KX538924 – Floudas and Hibbett (2015)
P. alnea FP-151125 Michigan, United States KP135177 MZ637181 Spirin et al. (2017)
P. alnea ssp. lubrica Spirin 8229 Washington, United States KU893876 – Floudas and Hibbett (2015)
P. alnea ssp. lubrica HHB-13753 Alaska, United States KP135178 – Spirin et al. (2017)
P. alpina Wu 1308-61* Yunnan, China MZ422790 MZ637182 Chen et al. (2021)
P. alpina Wu 1308-77 Yunnan, China MZ422791 MZ637183 Chen et al. (2021)
P. arizonica RLG-10248-Sp United States KP135170 KP135239 Floudas and Hibbett (2015)
P. australis GC 1704-27 Taiwan, China MZ422793 MZ637185 Floudas and Hibbett (2015)
P. australis HHB-7105-Sp United States KP135081 KP135240 Floudas and Hibbett (2015)
P. australosanguinea MA-Fungi 91308 Chile MH233925 MH233928 Phookamsak et al. (2019)
P. australosanguinea MA-Fungi 91309* Chile MH233926 MH233929 Phookamsak et al. (2019)
P. bambusicola Wu 0707-2 Taiwan, China MF399404 MF399395 Wu et al. (2018b)
P. brunnea He 1873 Zhejiang, China KX212220 KX212224 Liu and He (2016)
P. burdsallii FP-101018-sp Minnesota, United States AY219348 – Liu and He (2016)
P. burdsallii He 2066* Wisconsin, United States MT235690 MT248177 de Koker et al. (2003)
P. burtii FD-171 Massachusetts, United States KP135116 – Floudas and Hibbett (2015)
P. burtii HHB-4618-Sp United States KP135117 KP135241 Floudas and Hibbett (2015)
P. calotricha Vanhanen-382 Finland KP135107 – Floudas and Hibbett (2015)
P. canobrunnea CHWC 1506-66 Taiwan, China LC412095 LC412104 Wu et al. (2018a)
P. canolutea Wu 9712-18 Taiwan, China MZ422796 – Chen et al. (2021)
P. canolutea Wu 9211-105* Taiwan, China MZ422795 GQ470641 Chen et al. (2021)
P. carnosa HHB-9195 United States KP135129 KP135242 Floudas and Hibbett (2015)
P. chrysosporium HHB-6251-Sp United States KP135094 KP135246 Floudas and Hibbett (2015)
P. chrysosporium PC139 Taiwan, China MZ422797 MZ637186 Floudas and Hibbett (2015)
P. cinerea He 5998* Hainan, China – MT248171 Xu et al. (2020)
P. cinerea He 6003 Hainan, China – MT248172 Xu et al. (2020)
P. citrinosanguinea FP-105385 Massachusetts, United States KP135100 KP135234 Floudas and Hibbett (2015)
P. citrinosanguinea FD-287* Massachusetts, United States KP135095 – Floudas and Hibbett (2015)
P. citrinosanguinea FP-105385-Sp United States KP135100 KP135234 Floudas and Hibbett (2015)
P. concrescens Spirin 7322 Russia KP994380 KP994382 Volobuev et al. (2015)
P. concrescens CHWC 1507-39 Taiwan, China MZ422798 – Chen et al. (2021)
P. crystallina Chen 3576* Taiwan, China MZ422801 – Chen et al. (2021)
P. crystallina GC 1409-7 Taiwan, China MZ422803 MZ637189 Chen et al. (2021)
P. cumulodentata Wu 1708-91 Liaoning, China MZ422804 MZ637190 Volobuev et al. (2015)
P. cumulodentata LE 298935 Russia KP994359 KP994386 Volobuev et al. (2015)
P. cystidiata GC 1708-358* Liaoning, China LC412096 – Wu et al. (2018a)
P. cystidiata Wu 1708-326 Taiwan, China LC412097 LC412100 Wu et al. (2018a)
P. deflectens FCUG 2777 Turkey – GQ470644 Wu et al. (2010)
P. ericina HHB-2288 United States KP135167 KP135247 Floudas and Hibbett (2015)
P. ericina HHB-2714 North Carolina, United States KP135169 – Floudas and Hibbett (2015)
P. fusca Wu 1409-163 Hubei, China LC412099 LC412106 Wu et al. (2018a)
P. fusca Wu 1409-161* Hubei, China LC412098 LC412105 Wu et al. (2018a)
P. fuscomarginata RLG-10834-Sp New Mexico, United States MZ422806 MZ637192 Chen et al. (2021)
P. ginnsii Wu 9210-22* Hubei, China MZ422807 MZ637193 Chen et al. (2021)
P. granulate GC 1703-5 Hubei, China MZ422809 MZ637195 Chen et al. (2021)
P. granulate Wu 9210-57* Hubei, China MZ422810 MZ637196 Chen et al. (2021)
P. guangdongensis Wu 1809-348* Guangdong, China MZ422813 MZ637199 Chen et al. (2021)
P. guangdongensis Wu 1809-359 Guangdong, China MZ422814 MZ637200 Chen et al. (2021)
P. hymenochaetoides He 5988* Hainan, China – MT248173 Xu et al. (2020)
P. incarnata WEI 16-075 Taiwan, China MF399406 MF399397 Wu et al. (2018b)
P. incarnata WEI 16-078* Taiwan, China MF399407 MF399398 Wu et al. (2018b)
P. inflata Dai 10376 Jiangxi, China JX623929 JX644062 Jia et al. (2014)
P. krikophora GC 1602-73 Taiwan, China MZ422816 MZ637202 Chen et al. (2021)
P. krikophora HHB-6736-Sp Florida, United States MZ422817 MZ637203 Chen et al. (2021)
P. laevis KHL11839 Sweden EU118652 EU118652 Larsson (2007)
P. laevis Wu 0309-40 Jilin, China MZ422818 – Chen et al. (2021)
P. laevis HHB-15519 United States KP135149 KP135249 Floudas and Hibbett (2015)
P. leptocystidiata Dai 10468 Jiangxi, China MT235684 MT248167 Xu et al. (2020)
P. leptocystidiata He 5853* Guangdong, China MT235685 MT248168 Xu et al. (2020)
P. livescens GC 1612-11 Taiwan, China MZ422819 MZ637204 Floudas and Hibbett (2015)
P. livescens FD-106 United States KP135070 KP135253 Floudas and Hibbett (2015)
P. magnoliae HHB-9829-Sp United States KP135089 KP135237 Floudas and Hibbett (2015)
P. metuloidea He 2565* Yunnan, China – MT248163 Xu et al. (2020)
P. metuloidea He 2766 Yunnan, China MT235682 MT248164 Xu et al. (2020)
P. minor He 3977 Hainan, China – MT248169 Xu et al. (2020)
P. minor He 3988* Hainan, China MT235686 MT248170 Xu et al. (2020)
P. parmastoi WEI 16-481 Taiwan, China MZ422822 MZ637207 Chen et al. (2021)
P. parmastoi Wu 880313-6* Taiwan, China MZ422823 GQ470654 Chen et al. (2021)
P. porostereoides He 1902 Shanxi, China KX212217 KX212221 Liu and He (2016)
P. pruinose CLZhao 7112 Yunnan, China MZ435346 MZ435350 Wang and Zhao (2021)
P. pruinose CLZhao 7113* Yunnan, China MZ435347 MZ435351 Wang and Zhao (2021)
P. pseudomagnoliae PP-25 South Africa KP135091 KP135250 Floudas and Hibbett (2015)
P. pseudosanguinea FD-244 United States KP135098 KP135251 Floudas and Hibbett (2015)
P. queletii HHB-11463 Wisconsin, United States KP134994 KP135235 Floudas and Hibbett (2015)
P. queletii FP-102166 Illinois, United States KP134995 – Floudas and Hibbett (2015)
P. rhizomorpha GC 1708-335* Taiwan, China MZ422824 MZ637208 Chen et al. (2021)
P. rhizomorpha GC 1708-354 Taiwan, China MZ422825 MZ637209 Chen et al. (2021)
P. rhodella FD-18 United States KP135187 KP135258 Floudas and Hibbett (2015)
P. robusta Wu 1109-69 Jilin, China MF399409 MF399400 Wu et al. (2018b)
P. sanguinea HHB-7524 United States KP135101 KP135244 Floudas and Hibbett (2015)
P. sanguinea Niemela 7993 Finland KP135105 – Floudas and Hibbett (2015)
P. sanguineocarnosa FD-359 United States KP135122 KP135245 Floudas and Hibbett (2015)
P. shenghuaii Dai 24610* Yunnan, China OP874925 OP874920 Present study
P. shenghuaii Dai 24609 Yunnan, China OP874924 OP874919 Present study
P. sinensis GC 1809-56 Taiwan, China MT235689 MT248176 Xu et al. (2020)
P. sinensis He 4660* Liaoning, China MT235688 MT248175 Xu et al. (2020)
P. sordida FD-241 United States KP135136 KP135252 Floudas and Hibbett (2015)
Phanerochaete s.l. sp. TJV-93-262-T Louisiana, United States KP135021 – Floudas and Hibbett (2015)
Phanerochaete s.l. sp. RLG-13408-Sp Louisiana, United States KP135020 – Floudas and Hibbett (2015)
Phanerochaete sp. FCUG 2777 Turkey MZ422830 – Wu et al. (2010)
P. spadicea Wu 0504-11 Yunnan, China MZ422836 – Chen et al. (2021)
P. spadicea Wu 0504-15* Yunnan, China MZ422837 – Chen et al. (2021)
P. stereoides He 2309 Hunan, China KX212219 KX212223 Liu and He (2016)
P. subceracea FP-105974-R United States KP135162 KP135255 Floudas and Hibbett (2015)
P. subrosea He 2421* Ningxia, China MT235687 MT248174 Xu et al. (2020)
P. taiwaniana Wu 880824-17* Taiwan, China MZ422842 GQ470666 Chen et al. (2021)
P. taiwaniana Wu 0112-13 Taiwan, China MF399412 MF399403 Wu et al. (2018b)
P. thailandica 2015-07* Thailand MF467737 – Chen et al. (2021)
P. thailandica Wu 1710-3 Vietnam MZ422843 MZ637223 Chen et al. (2021)
P. velutina Kotiranta 25567 Russia KP994354 KP994387 Volobuev et al. (2015)
P. xerophila HHB-8509-Sp Arizona, United States KP134996 KP135259 Floudas and Hibbett (2015)
P. xerophila KKN-172 Arizona, United States KP134997 – Floudas and Hibbett (2015)
P. yunnanensis He 2697 Yunnan, China – MT248165 Xu et al. (2020)
P. yunnanensis He 2719* Yunnan, China MT235683 MT248166 Xu et al. (2020)
Phlebiopsis brunneocystidiata Chen 666 Taiwan, China MT561707 GQ470640 Wu et al. (2010)
P. crassa He 5205 Vietnam MT452523 MT447448 Zhao et al. (2021)
P. cylindrospora He 5984* Hainan, China MT386404 MT447445 Zhao et al. (2021)
P. friesii He 5820 Sri Lanka MT452530 MT447415 Zhao et al. (2021)
P. magnicystidiata He 5648* Hunan, China MT386377 MT447409 Zhao et al. (2021)
P. membranacea He 3849* Hainan, China MT386401 MT447441 Zhao et al. (2021)
P. sinensis He 4673* Sichuan, China MT386397 MT447435 Zhao et al. (2021)
P. yunnanensis CLZhao 3990 Yunnan, China MH744141 MH744143 Zhao et al. (2019)
Rhizochaete americana FP-102188 Illinois, United States KP135409 KP135277 Floudas and Hibbett (2015)
R. americana HHB2004 Georgia, United States AY219391 AY219391 Greslebin et al. (2004)
R. belizensis FP150712 Belize KP135408 KP135280 Floudas and Hibbett (2015)
R. borneensis WEI16-426 Taiwan, China MZ637070 MZ637270 Chen et al. (2021)
R. brunnea MR11455 Argentina AY219389 AY219389 Greslebin et al. (2004)
R. filamentosa FP105240 Indiana, United States KP135411 AY219393 Nakasone et al. (2017)
R. filamentosa HHB 3169 Maryland, United States KP135410 KP135278 Floudas and Hibbett (2015)
R. fissurata CLZhao2200 Yunnan, China MZ713640 MZ713844 Gu and Zhao (2021)
R. fissurata CLZhao7965 Yunnan, China MZ713641 MZ713845 Gu and Zhao (2021)
R. fissurata CLZhao10407* Yunnan, China MZ713642 MZ713846 Gu and Zhao (2021)
R. fissurata CLZhao10418 Yunnan, China MZ713643 MZ713847 Gu and Zhao (2021)
R. flava PR 1141 Puerto Rico KY273030 KY273033 Nakasone et al. (2017)
R. flava PR3148 Puerto Rico KY273029 – Nakasone et al. (2017)
R. fouquieriae KKN-121 Arizona, United States AY219390 GU187608 Nakasone et al. (2017)
R. fouquieriae KKN-121sp United States KY948786 KY948858 Justo et al. (2017)
R. grandinosa CLZhao3117* Yunnan, China MZ713644 MZ713848 Gu and Zhao (2021)
R. lutea Wu 880417-5 Taiwan, China MZ637072 GQ470651 Chen et al. (2021)
R. radicata FD123 Massachusetts, United States KP135407 KP135279 Floudas and Hibbett (2015)
R. radicata FD338 Massachusetts, United States KP135406 – Floudas and Hibbett (2015)
R. radicata HHB1909 Highlands, United States AY219392 AY219392 Greslebin et al. (2004)
R. rubescens Wu0910-45 Beijing, China LC387335 MF110294 Chen et al. (2018)
R. sulphurina DLL2014-176 Idaho, United States KY273032 – Nakasone et al. (2017)
R. sulphurina HHB5604 Montana, United States KY273031 GU187610 Nakasone et al. (2017)
R. sulphurina KHL16087 Brazil KT003523 – Chikowski et al. (2016)
R. sulphurina URM87190 Brazil KT003522 KT003519 Chikowski et al. (2016)
R. variegata Dai 24600* Guizhou, China OP874926 OP874921 Present study
R. variegata Dai 24601 Guizhou, China OP874927 OP874922 Present study

New species are in bold with type specimens marked with an asterisk (*).

 Phylogenetic analyses

New sequences, deposited in GenBank (http://www.ncbi.nlm.nih.gov/genbank/) ( Table 1 ), were aligned with additional sequences retrieved from GenBank ( Table 1 ) using BioEdit 7.0.5.3 (Hall, 1999) and ClustalX 1.83 (Thompson et al., 1997), followed by manual adjustment. Sequence alignment was deposited at TreeBase (http://purl.org/phylo/treebase/; submission ID 29897). Sequences of Bjerkandera adusta (Willd.) P. Karst. and B. centroamericana Kout et al. were used as outgroups (Chen et al., 2021). Maximum likelihood (ML) and Bayesian inference (BI) methods were used for the phylogenetic analysis. The GTR + I + G model was estimated as the best-fit evolutionary model by PhyloSuite 1.2.2 (Zhang et al., 2020) using the Akaike information criterion. The ML analysis was carried out with RAxML 8.2.12 (Stamatakis, 2006; Silvestro and Michalak, 2012), and the BI tree reconstruction was carried out with MrBayes 3.2.5 (Ronquist et al., 2012). Four Markov chains were run for two runs from random starting trees for 10 million generations, and trees were sampled every 1,000 generations. The burn-in was set to discard 25% of the trees. A majority rule consensus tree of all the remaining trees was calculated. Branches that received bootstrap support for ML and Bayesian posterior probabilities (BPP) greater than or equal to 75% (ML) and 0.95 (BPP) were considered as significantly supported.

 Results Phylogeny

The ITS + nLSU dataset included 155 fungal collections representing 101 taxa of the family Phanerochaetaceae. PhyloSuite suggested GTR + I + G to be the best-fit models of nucleotide evolution for BI. Bayesian analysis resulted in a concordant topology with an average standard deviation of split frequencies = 0.006701. The ML and BI analyses resulted in nearly identical topologies, and thus, only the ML tree is presented with the ML and BPP when they were greater than or equal to 50% and 0.90, respectively.

The phylogram inferred from ITS + nLSU sequences within the family Phanerochaetaceae highlighted two undescribed species nested in Phanerochaete and Rhizochaete, respectively. Phanerochaete shenghuaii formed an independent lineage with a robust support (ML = 99, BPP = 1.0) and stably nested within the core Phanerochaete clade. Rhizochaete variegata clustered in Rhizochaete clade with high support (ML = 99, BPP = 1.0) and grouped with Rhizochaete radicata (Henn.) Gresl. et al. and R. grandinosa C.L. Zhao & Z.R. Gu.

 Taxonomy

Phanerochaete shenghuaii Q.Y. Zhang, Y.C. Dai & Jing Si, sp. nov., Figures 1 , 2

Basidiocarps of Phanerochaete shenghuaii (holotype, Dai 24610). (A) In situ. (B) Detailed view of the margin. (C) Reaction with KOH. Scale bars: (A) = 1 cm, (B) = 2 mm, (C) = 0.5 cm.

Microscopic structures of Phanerochaete shenghuaii (drawn from the holotype, Dai 24610). (A) Basidiospores. (B) Basidia and basidioles. (C) Cystidia. (D) A vertical section of the subiculum. (E) A vertical section of the hymenium.

MycoBank: 847200

Type — China, Yunnan Province, Zhaotong, Daguan County, Huanglianhe Scenic Spot, on fallen liana branch, 16 July 2022, Dai 24610 (holotype, BJFC038931).

Etymology — Shenghuaii (Lat.): In honor of Professor Sheng-Hua Wu, the Chinese mycologist.

Basidiocarps — Annual, effused, adnate, inseparable from substrate, membranaceous to subceraceous, up to 2.5 cm long, 1.5 cm wide, and 0.2 mm thick in section. Hymenial surface ivory white to cream when juvenile, buff to yellowish brown with age, buff in KOH, smooth, uncracked; margin concolorous with hymenial surface, thinning out, usually rhizomorphic.

Hyphal structure — Hyphal system monomitic; generative hyphae mostly simple septate, occasionally with clamp connections in subiculum, IKI−, CB−; tissue unchanged in KOH.

Subiculum — Subicular hyphae hyaline, slightly thick-walled, frequently simple septate, occasionally with clamp connections, frequently branched, usually strongly encrusted with crystal granules, interwoven, 3–5 μm in diameter.

Hymenophore — Subhymenial hyphae hyaline, thin-walled, smooth, simple septate, frequently branched, interwoven, 2.5–5 μm in diameter; cystidia smooth, immersed or projecting from hymenium, narrowly fusiform or clavate with pointed tips, hyaline, thin-walled, smooth, with a simple septum at the base, 18–35 × 3–5 µm; basidia clavate, with a basal simple septum and four sterigmata, 22–30 × 4–5 µm; basidioles similar to basidia in shape, but slightly smaller.

Basidiospores — Ellipsoid with a distinct apiculus, hyaline, thin-walled, smooth, occasionally with one or two guttules, IKI−, CB−, (4.5–)4.8–6(–6.4) × 2.5–3.8(–4) µm, L = 5.26 µm, W = 3.01 µm, Q = 1.71–1.79 (n = 60/2).

Additional specimen (paratype) examined — China, Yunnan Province, Zhaotong, Daguan County, Huanglianhe Scenic Spot, on fallen angiosperm branch, 16 July 2022, Dai 24609 (BJFC038930).

Rhizochaete variegata Q.Y. Zhang, Y.C. Dai & Jing Si, sp. nov., Figures 3 , 4

Basidiocarps of Rhizochaete variegata (holotype, Dai 24600). (A) In situ. (B) Detailed view of the margin. (C) Reaction with KOH. Scale bars: (A) = 1 cm, (B) = 1 mm, (C) = 0.5 cm.

Microscopic structures of Rhizochaete variegata (drawn from the holotype, Dai 24600). (A) Basidiospores. (B) Basidia and basidioles. (C) Cystidia. (D) A vertical section of the subiculum. (E) A vertical section of the hymenium.

MycoBank: 847201

Type — China, Guizhou Province, Zunyi, Suiyang County, Kuankuoshui Nature Reserve, on fallen angiosperm trunk, 07 July 2022, Dai 24600 (holotype, BJFC038928).

Etymology — Variegata (Lat.): referring to the species having variable cystidia.

Basidiocarps — Annual, effused, loosely adnate, easily separable from substrate, membranaceous, soft, fragile, up to 9 cm long, 3.5 cm wide, and 1 mm thick in section. Hymenial surface buff-yellow to clay-pink when fresh, cream to buff upon drying, violet in KOH, smooth or locally tuberculate, occasionally cracked; margin darker or concolorous with hymenial surface, thinning out, usually rhizomorphic.

Hyphal structure — Hyphal system monomitic; generative hyphae simple septate, IKI−, CB−; tissue unchanged in KOH.

Subiculum — Subicular hyphae hyaline, slightly thick-walled, simple septate, rarely branched, bearing abundant crystal granules, strongly interwoven, 3.5–6 μm in diameter.

Hymenophore — Subhymenial hyphae hyaline, slightly thick-walled, smooth, simple septate, more or less regularly arranged, 3–5 μm in diameter. Hymenium contains a dense palisade of cystidia and basidia, IKI−, CB−; cystidia numerous, immersed or projecting from hymenium, clavate, subfusiform or subulate with an obtuse apex, hyaline, slightly thick-walled, some with thin-walled apex, with a simple septum at the base, some apically or centrally encrusted, 28–52 × 5–8 µm; basidia narrowly clavate, with a basal simple septum and four sterigmata, 30–45 × 4–5 µm; basidioles similar to basidia in shape, but slightly smaller.

Basidiospores — Ellipsoid with a distinct apiculus, hyaline, thin-walled, smooth, occasionally with one or two small guttules, IKI−, CB−, 3–4(–4.2) × (2–)2.2–3(–3.2) µm, L = 3.61 µm, W = 2.72 µm, Q = 1.27–1.38 (n = 60/2).

Additional specimen (paratype) examined — China, Guizhou Province, Zunyi, Suiyang County, Kuankuoshui Nature Reserve, on fallen angiosperm trunk, 07 July 2022, Dai 24601 (BJFC038929).

 Discussion

Southwest China has a complex topography and geography, luxuriant vegetation, and virgin forests and has highly variable weather including tropical, subtropical, and alpine climates, thus providing a favorable region for the growth and reproduction of higher fungi (Yuan and Dai, 2008; Dai et al., 2021; Wu et al., 2022a). The extremely high fungal diversity in this area has attracted much attention from mycologists both at home and abroad (Feng and Yang, 2018). It is worth noting that the two new corticioid species P. shenghuaii and R. variegata were collected from Northeast Yunnan and Northwest Guizhou, respectively, and the type locality of the two new species is in a typical subtropical climate.

Phanerochaete shenghuaii is characterized by white to cream basidiocarps with rhizomorphic margin, encrusted subicular hyphae, and smooth cystidia. Morphologically, three species, Phanerochaete rhizomorpha C.C. Chen et al., P. leptocystidiata Y.L. Xu & S.H. He, and P. sinensis Y.L. Xu et al., are similar to P. shenghuaii by sharing similar basidiocarps, rhizomorphic margin, and smooth cystidia. However, P. rhizomorpha is described from Taiwan Province, China, and differs from P. shenghuaii by its subcapitate to cylindrical cystidia with obtuse apices and smaller basidiospores (3.9–5.3 × 2.1–3 μm vs. 4.8–6 × 2.5–3.8 µm, Chen et al., 2021). Phanerochaete leptocystidiata is widely distributed in South China and differs from P. shenghuaii by its basidiocarps easily separable from substrate and longer cystidia (30–70 μm in length vs. 18–35 µm in length, Xu et al., 2020). Phanerochaete sinensis is distinguished from P. shenghuaii in having longer cystidia (35–50 µm in length vs. 18–35 µm in length) and smaller basidiospores (4–5 × 2–2.5 μm vs. 4.8–6 × 2.5–3.8 µm, Xu et al., 2020).

In addition, the diversity of flora of seed plants and the distinctly diverse climates in Yunnan Province both contribute to the suitable substrates and environments for Phanerochaete species. Recently, a large number of Phanerochaete species have been found in Yunnan Province (Xiong and Dai, 2009; Wu et al., 2010; Xu et al., 2020; Chen et al., 2021; Wang and Zhao, 2021). Among them, Phanerochaete yunnanensis Y.L. Xu & S.H. He is similar to P. shenghuaii by growing on dead liana and fallen angiosperm branches but differs by grandinioid basidiocarps and the absence of cystidia. Phanerochaete pruinose C.L. Zhao and D.Q. Wang is similar to P. shenghuaii by sharing white and smooth hymenophore, but differs by lacking cystidia and having thinner basidiospores (1.5–2.7 μm in width vs. 2.5–3.8 µm in width, Wang and Zhao, 2021). It is still noteworthy that P. rhizomorpha C.L. Zhao and D.Q. Wang described from Yunnan Province is an invalid name, attributed to the priority of P. rhizomorpha C.C. Chen et al. (Chen et al., 2021; Wang and Zhao, 2021). In addition, the two taxa represent two independent species according to their distinctive DNA sequences and morphology.

Our phylogenetic analysis demonstrates that Rhizochaete is monophyletic with a low support and clusters as a sister clade to Hapalopilus, Phaeophlebiopsis, and Phlebiopsis. Two specimens of R. variegata form a lineage with a strong support (ML = 99, BPP = 1.0, Figure 5 ). Rhizochaete variegata is closely related to R. grandinosa and R. radicata (ML = 100, BPP = 1, Figure 5 ), and these three species share curry-yellow hymenial surface, violet in KOH, thick-walled and encrusted subicular hyphae, and similar-sized basidiospores. However, R. variegata has abundant variable and slightly thick-walled cystidia with a thin-walled apex, which can be readily distinguished from R. grandinosa and R. radicata (Greslebin et al., 2004; Gu and Zhao, 2021). Furthermore, there are differences of more than eight base pairs between their sequences, which amounts to 2% nucleotides in the ITS regions. Morphologically, Rhizochaete sulphurosa (Bres.) Chikowski et al. may be confused with R. variegata by sharing yellow basidiocarps, hymenial surface violet in KOH, and thin or slightly thick-walled (<1 µm) cystidia. Nevertheless, R. sulphurosa differs from R. variegata by its longer basidiospores (4.5–5.5 µm in length vs. 3–4 µm in length, Chikowski et al., 2016).

Maximum likelihood (ML) tree illustrating the phylogeny of the two new species in Phanerochaetaceae based on ITS + nLSU sequences. Branches are labeled with ML bootstrap >50% and Bayesian posterior probabilities (BPP) >0.90, respectively. New species are highlighted by red text.

Although more taxa of Phanerochaetaceae have been described (Greslebin et al., 2004; Bianchinotti et al., 2005; Chen et al., 2021), the taxonomy of corticioid fungi in Polyporales is woefully understudied. Many closely related genera are difficult to differentiate based on apparently plesiomorphic morphology, such as Phanerochaete, Rhizochaete, Phaeophlebiopsis, and Hapalopilus (Bianchinotti et al., 2005; Chen et al., 2021). Rhizochaete is separated from Phanerochaete mainly by their basidiocarp reaction with KOH (Greslebin et al., 2004; Chen et al., 2021). Indeed, this character is delimited in most species of the two genera. However, there are still some species of Phanerochaete exhibiting red in KOH, such as P. affinis (Burt) Parmasto and P. aurantiobadia Ghob.-Nejh. et al. (Punugu et al., 1980; Ghobad-Nejhad et al., 2015). Therefore, more samples from worldwide and multigene phylogeny analysis are urgently needed for understanding the diversity of corticioid species of Polyporales.

Southwest China is a hotspot for fungal diversity, and numerous taxa of wood-inhabiting fungi have been described from this area based on morphological and molecular phylogenetic analyses (Dai, 2010; Zhao et al., 2015; Zhou et al., 2016; Dai et al., 2021; Guan and Zhao, 2021; Wang et al., 2021; Wu et al., 2021; Wu et al., 2022b). Notably, the species diversity of corticioid fungi in this area is still not well-known, and therefore, the present paper confirms that more unknown species exist in this area.

 Data availability statement

The datasets presented in this study can be found in online repositories. The names of the repository/repositories and accession number(s) can be found below: https://www.ncbi.nlm.nih.gov/genbank/, OP874919, OP874920, OP874921, OP874922, OP874924, OP874925, OP874926, OP874927.

 Author contributions

Q-YZ, Z-BL, and JS designed the research and contributed to data analysis and interpretation. Q-YZ prepared the samples and drafted the manuscript. Z-BL conducted molecular experiments and analyzed the data. H-GL and JS discussed the results and edited the manuscript. All authors contributed to the article and approved the submitted version.

 Funding

The research was supported by the National Natural Science Foundation of China (Nos. 32270016 and 32070016).

 Acknowledgments

The authors would like to express their deep appreciation to Prof. Yu-Cheng Dai (Beijing Forestry University, China) for allowing them to study his specimens.

 Conflict of interest

The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.

 Publisher’s note

All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.

 References Bianchinotti M. V. Rajchenberg M. Greslebin A. G. (2005). Parenthesome structure of some corticioid fungi. Mycol. Res. 109, 923–926. doi: 10.1017/S0953756205003333 Burdsall H. H. (1985). A contribution to the taxonomy of the genus Phanerochaete . Mycol. Mem. 10, 1–165. Chen C. C. Chen C. Y. Wu S. H. (2021). Species diversity, taxonomy and multi-gene phylogeny of phlebioid clade (Phanerochaetaceae, irpicaceae, meruliaceae) of polyporales. Fungal Divers. 111, 337–442. doi: 10.1007/s13225-021-00490-w Chen C. C. Wu S. H. Chen C. Y. (2018). Hydnophanerochaete and Odontoefibula, two new genera of phanerochaetoid fungi (Polyporales, basidiomycota) from East Asia. MycoKeys 39, 75–96. doi: 10.3897/mycokeys.39.28010 Chikowski R. S. Larsson K. H. Gibertoni T. B. (2016). Three new combinations in Rhizochaete (Agaricomycetes, fungi) and a new record to the Brazilian Amazonia. Nova Hedwig. 102, 185–196. doi: 10.1127/nova_hedwigia/2015/0298 Cui B. K. Li H. J. Ji X. Zhou J. L. Song J. Si J. . (2019). Species diversity, taxonomy and phylogeny of polyporaceae (Basidiomycota) in China. Fungal Divers. 97, 137–392. doi: 10.1007/s13225-019-00427-4 Dai Y. C. (2010). Hymenochaetaceae (Basidiomycota) in China. Fungal Divers. 45, 131–343. doi: 10.1007/s13225-010-0066-9 Dai Y. C. (2011). A revised checklist of corticioid and hydnoid fungi in China for 2010. Mycoscience 52, 69–79. doi: 10.1007/S10267-010-0068-1 Dai Y. C. Yang Z. L. Cui B. K. Wu G. Yuan H. S. Zhou L. W. . (2021). Diversity and systematics of the important macrofungi in Chinese forests. Mycosystema 40, 770–805. doi: 10.13346/j.mycosystema.210036 de Koker T. Nakasone K. K. Haarhof J. Burdsall H. H. Janse B. J. H. (2003). Phylogenetic relationships of the genus Phanerochaete inferred from the internal transcribed spacer region. Mycol. Res. 107, 1032–1040. doi: 10.1017/S095375620300827X Feng B. Yang Z. (2018). Studies on diversity of higher fungi in yunnan, southwestern China: A review. Plant Divers. 40, 165–171. doi: 10.1016/j.pld.2018.07.001 Floudas D. Hibbett D. S. (2015). Revisiting the taxonomy of Phanerochaete (Polyporales, basidiomycota) using a four gene dataset and extensive ITS sampling. Fungal Biol. 119, 679–719. doi: 10.1016/j.funbio.2015.04.003 Ghobad-Nejhad M. Liu S. L. Langer E. Dai Y. C. (2015). Molecular and morphological evidence reveal a new non-cystidiate species belonging to the core Phanerochaete (Polyporales). Mycol. Prog. 14, 1–7. doi: 10.1007/s11557-015-1072-9 Greslebin A. Nakasone K. K. Rajchenberg M. (2004). Rhizochaete, a new genus of phanerochaetoid fungi. Mycologia 96, 260–271. doi: 10.1080/15572536.2005.11832976 Guan Q. X. Zhao C. L. (2021). Two new corticioid species, Hyphoderma sinense and H. floccosum (Hyphodermataceae, polyporales), from southern China. Mycosystema 40, 447–461. doi: 10.13346/j.mycosystema.200382 Gu Z. R. Zhao C. L. (2021). The hidden wood-decaying fungal diversity: Rhizochaete from east Asia. Diversity 13, 503. doi: 10.3390/d13100503 Hall T. A. (1999). Bioedit: A user-friendly biological sequence alignment editor and analysis program for windows 95/98/NT. Nucleic Acids Symp. Ser. 41, 95–98. doi: 10.1021/bk-1999-0734.ch008 He M. Q. Zhao R. L. Hyde K. D. Begerow D. Kemler M. Yurkov A. . (2019). Notes, outline and divergence times of basidiomycota. Fungal Divers. 99, 105–367. doi: 10.1007/s13225-019-00435-4 Jia B. S. Zhou L. W. Cui B. K. Rivoire B. Dai Y. C. (2014). Taxonomy and phylogeny of Ceriporia (Polyporales, basidiomycota) with an emphasis of Chinese collections. Mycol. Prog. 13, 81–93. doi: 10.1007/s11557-013-0895-5 Justo A. Miettinen O. Floudas D. Ortiz-Santana B. Sjokvist E. Lindner D. . (2017). A revised family-level classification of the polyporales (Basidiomycota) . Fungal Biol. 121, 798–824. doi: 10.1016/j.funbio.2017.05.010 Kirk P. M. Cannon P. F. Minter D. W. Stalpers J. A. (2008). Dictionary of the fungi. 10th edn (Wallingford: CABI). Larsson K. H. (2007). Re-thinking the classification of corticioid fungi. Mycol. Res. 111, 1040–1063. doi: 10.1016/j.mycres.2007.08.001 Liu S. L. He S. H. (2016). Phanerochaete porostereoides, a new species in the core clade with brown generative hyphae from China. Mycosphere 7, 648–655. doi: 10.5943/mycosphere/7/5/10 Miettinen O. Spirin V. Vlasák J. Rivoire B. Stenroos S. Hibbett D. (2016). Polypores and genus concepts in phanerochaetaceae (Polyporales, basidiomycota) . MycoKeys 17, 1–46. doi: 10.3897/mycokeys.17.10153 Nakasone K. K. Draeger K. R. Ortiz-Santana B. (2017). A contribution to the taxonomy of Rhizochaete (Polyporales, basidiomycota). Cryptogam. Mycol. 38, 81–99. doi: 10.7872/crym/v38.iss1.2017.81 Ortiz-Santana B. Lindner D. L. Miettinen O. Justo A. Hibbett D. S. (2013). A phylogenetic overview of the antrodia clade (Basidiomycota, polyporales) . Mycologia 105, 1391–1411. doi: 10.3852/13-051 Petersen J. H. (1996). “The Danish mycological society’s colour-chart,” in Greven: Foreningen til svampekundskabens fremme, Denmark, 1–6. Phookamsak R. Hyde K. D. Jeewon R. Bhat D. J. Jones E. B. G. Maharachchikumbura S. S. N. . (2019). Fungal diversity notes 929–1035: taxonomic and phylogenetic contributions on genera and species of fungi. Fungal Divers. 95, 1–273. doi: 10.1007/s13225-019-00421-w Punugu A. Dunn M. T. Welden A. L. (1980). The peniophoroid fungi of the West indies. Mycotaxon 10, 428–454. Ronquist F. Teslenko M. van der Mark P. Avres D. L. Darling A. Höhna S. . (2012). MrBayes 3.2: Efficient Bayesian phylogenetic inference and model choice, across a large model space. Syst. Biol. 61, 539–542. doi: 10.1093/sysbio/sys029 Ryvarden L. Melo I. (2017). Poroid fungi of Europe. Synop. Fungorum 37, 1–431. Silvestro D. Michalak I. (2012). RaxmlGUI: A graphical front-end for RAxML. Org. Divers. Evol. 12, 335–337. doi: 10.1007/s13127-011-0056-0 Spirin V. Volobuev S. Okun M. Miettinen O. Larsson K. H. (2017). What is the type species of Phanerochaete (Polyporales, basidiomycota)? Mycol. Prog. 16, 171–183. doi: 10.1007/s11557-016-1267-8 Stamatakis A. (2006). RAxML-VI-HPC: Maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models. Bioinf. (Oxford England) 22, 2688–2690. doi: 10.1093/bioinformatics/btl446 Thompson J. D. Gibson T. J. Plewniak F. Jeanmougin F. Higgins D. G. (1997). The Clustal_X windows interface: fexible strategies for multiple sequence alignment aided by quality analysis tools. Nucleic Acids Res. 25, 4876–4882. doi: 10.1093/nar/25.24.4876 Vilgalys R. Hester M. (1990). Rapid genetic identification and mapping of enzymatically amplified ribosomal DNA from several Cryptococcus species. J. Bacteriol. 172, 4238–4246. doi: 10.1128/jb.172.8.4238-4246.1990 Volobuev S. Okun M. Ordynets A. Spirin V. (2015). The Phanerochaete sordida group (Polyporales, basidiomycota) in temperate Eurasia, with a note on Phanerochaete pallida . Mycol. Prog. 14, 1–13. doi: 10.1007/s11557-015-1097-0 Vu D. Groenewald M. de Vries M. Gehrmann T. Stielow B. Eberhardt U. . (2019). Large-Scale generation and analysis of filamentous fungal DNA barcodes boosts coverage for kingdom fungi and reveals thresholds for fungal species and higher taxon delimitation. Stud. Mycol. 92, 1–20. doi: 10.1016/j.simyco.2018.05.001 Wang K. Chen S. L. Dai Y. C. Jia Z. F. Li T. H. Liu T. Z. . (2021). Overview of china's nomenclature novelties of fungi in the new century, (2000–2020). Mycosystema 40, 822–833. doi: 10.13346/j.mycosystema.210064 Wang D. Q. Zhao C. L. (2021). Morphological and phylogenetic evidence for recognition of two new species of Phanerochaete from East Asia. J. Fungi 7, 1063. doi: 10.3390/jof7121063 White T. J. Bruns T. D. Lee S. Taylor J. (1990). “Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics,” in PCR protocols: A guide to methods and applications. Eds. Innis M. A. Gelfand D. H. Sninsky J. J. White T. J. (New York Academic Press: US), 315–322. doi: 10.1016/B978-0-12-372180-8.50042-1 Wu S. H. (1990). The corticiaceae (Basidiomycetes) subfamilies phlebioideae, phanerochaetoideae and hyphodermoideae in Taiwan. Acta Bot. Fenn. 142, 1–123. Wu S. H. (1995). A study of the genus Phanerochaete (Aphyllophorales) with brown subicular hyphae. Mycotaxon 54, 163–172. Wu S. H. (1998). Nine new species of Phanerochaete from Taiwan. Mycol. Res. 102, 1126–1132. doi: 10.1017/S0953756298006091 Wu S. H. (2000). Six new species of Phanerochaete from Taiwan. Bot. Bull. Acad. Sin. (Taipei). 41, 165–174. Wu S. H. (2004). Two new species of Phanerochaete from Taiwan. Mycotaxon 90, 423–429. Wu S. H. (2007). Three new species of corticioid fungi from Taiwan. Bot. Stud. 48, 325–330. Wu S. H. Chen C. C. Wei C. L. (2018a). Three new species of Phanerochaete (Polyporales, basidiomycota) . MycoKeys 41, 91–106. doi: 10.3897/mycokeys.41.29070 Wu S. H. Chen Y. P. Wei C. L. Floudas D. Dai Y. C. (2018b). Two new species of Phanerochaete (Basidiomycota) and redescription of P. robusta . Mycol. Prog. 17, 425–435. doi: 10.1007/s11557-017-1368-z Wu F. Li S. J. Dong C. H. Dai Y. C. Papp V. (2020). The genus Pachyma (syn. wolfiporia) reinstated and species clarification of the cultivated medicinal mushroom “Fuling” in China. Front. Microbiol. 11. doi: 10.3389/fmicb.2020.590788 Wu F. Man X. W. Tohtirjap A. Dai Y. C. (2022a). A comparison of polypore funga and species composition in forest ecosystems of China, north America, and Europe. For. Ecosyst. 9, 100051. doi: 10.1016/j.fecs.2022.100051 Wu S. H. Nilsson H. R. Chen C. T. Yu S. Y. Hallenberg N. (2010). The white-rotting genus Phanerochaete is polyphyletic and distributed throughout the phleboid clade of the polyporales (Basidiomycota) . Fungal Divers. 42, 107–118. doi: 10.1007/s13225-010-0031-7 Wu F. Tohtirjap A. Fan L. F. Zhou L. W. Alvarenga R. L. M. Gibertoni T. B. . (2021). Global diversity and updated phylogeny of Auricularia (Auriculariales, basidiomycota) . J. Fungi 7, 933. doi: 10.3390/jof7110933 Wu F. Zhou L. W. Vlasák J. Dai Y. C. (2022b). Global diversity and systematics of hymenochaetaceae with poroid hymenophore. Fungal Divers. 113, 1–192. doi: 10.1007/s13225-021-00496-4 Xiong H. Dai Y. C. (2009). Notes on lignicolous corticioid fungi in China 3. Phanerochaete basidiomycota, polyporales in China. Mycosystem 28, 29–35. doi: 10.13346/j.mycosystema.2009.01.006 Xu Y. L. Cao Y. F. Nakasone K. K. Chen C. C. He S. H. (2020). Taxonomy and phylogeny of Phanerochaete sensu stricto (Polyporales, basidiomycota) with emphasis on Chinese collections and descriptions of nine new species. Mycosphere 11, 1527–1552. doi: 10.5943/mycosphere/11/1/12 Yuan H. S. Dai Y. C. (2008). Polypores from northern and central yunnan province, southwestern China. Sydowia 60, 147–159. Zhang D. Gao F. Jakovlić I. Zou H. Zhang J. Li W. X. . (2020). PhyloSuite: an integrated and scalable desktop platform for streamlined molecular sequence data management and evolutionary phylogenetics studies. Mol. Ecol. Resour. 20, 348–355. doi: 10.1111/1755-0998.13096 Zhao C. L. Cui B. K. Song J. Dai Y. C. (2015). Fragiliporiaceae, a new family of polyporales (Basidiomycota). Fungal Divers. 70, 115–126. doi: 10.1007/s13225-014-0299-0 Zhao Y. N. He S. H. Nakasone K. K. Wasantha Kumara K. L. Chen C. C. Liu S. L. . (2021). Global phylogeny and taxonomy of the wood-decaying fungal genus Phlebiopsis (Polyporales, basidiomycota) . Front. Microbiol. 12. doi: 10.3389/fmicb.2021.622460 Zhao C. L. Liu X. F. Ma X. (2019). Phlebiopsis yunnanensis sp. nov. (Polyporales, basidiomycota) evidenced by morphological characters and phylogenetic analysis. Nova Hedwig. 108, 265–279. doi: 10.1127/nova_hedwigia/2018/0508 Zhou L. W. Vlasák J. Decock C. Assefa A. Stenlid J. Abate D. . (2016). Global diversity and taxonomy of the Inonotus linteus complex (Hymenochaetales, basidiomycota): Sanghuangporus gen. nov., Tropicoporus excentrodendri and T. guanacastensis gen. et spp. nov., and 17 new combinations. Fungal Divers. 77, 335–347. doi: 10.1007/s13225-015-0335-8