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Research Article
Four new species of Phanerochaete (Polyporales, Basidiomycota) from China
expand article infoKai-Yue Luo‡, Xin Zhang‡, Yu-Cheng Dai‡, Yuan Yuan‡
‡ Beijing Forestry University, Beijing, China

Corresponding author: Yuan Yuan ( yuanyuan1018@bjfu.edu.cn )

Academic editor: Jennifer Luangsa-ard
© 2024 Kai-Yue Luo, Xin Zhang, Yu-Cheng Dai, Yuan Yuan.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation: Luo K-Y, Zhang X, Dai Y-C, Yuan Y (2024) Four new species of Phanerochaete (Polyporales, Basidiomycota) from China. MycoKeys 111: 41-64. https://doi.org/10.3897/mycokeys.111.133093
Open Access

Abstract

Four new wood-inhabiting fungi viz. Phanerochaete castanea, P. citrinoalba, P. citrinorhizomorpha, and P. wuyiensis spp. nov. – are proposed based on a combination of morphological features and molecular evidence. Phanerochaete castanea is characterized by soft coriaceous basidiomata detachable from the substrate, becoming reddish brown in KOH, subulate cystidia with an obtuse apex. Phanerochaete citrinoalba is characterized by the coriaceous basidiomata with smooth, cracking hymenial surface, sterile margins with yellowish to whitish rhizomorphs, a monomitic hyphal system, generative hyphae mostly with simple septa and occasionally with clamp connections at basal hyphae. Phanerochaete citrinorhizomorpha is characterized by soft coriaceous basidiomata with a salmon to peach hymenial surface, a sterile margin with yellowish rhizomorphs, simple septate generative hyphae, and clavate to subfusiform or subulate cystidia with an obtuse apex. Phanerochaete wuyiensis is characterized by membranaceous basidiomata with smooth or locally tuberculate hymenial surface and the whitish rhizomorphs, generative hyphae with both simple septa and clamp connections at basal hyphae, cystidia projecting above hymenium. DNA sequences of the ITS and LSU markers of the studied samples were generated, and phylogenetic analyses were performed with Maximum Likelihood and Bayesian Inference methods. The phylogenetic tree inferred from the concatenated ITS+nLSU dataset highlighted the placement of the four new species in the genus Phanerochaete (Phanerochaetaceae, Polyporales). Phylogenetically related and morphologically similar species to these four new species are discussed. Furthermore, an identification key to accepted species of Phanerochaete in China is given.

Key words

Molecular phylogeneny, Polyporales, taxonomy, wood-decaying fungi, white rot

Introduction

The taxa in the family Phanerochaetaceae are mostly corticioid fungi, especially Phanerochaete, as a major member of the family. It is a large genus with diverse morphological features, and it is widely distributed from boreal to tropical forests. It causes a white rot on all kinds of wood and plays an important role in carbon cycling (Burdsall 1985; Larsson et al. 2004; Bernicchia and Gorjón 2010; Ryvarden and Melo 2014; Ghobad-Nejhad et al. 2015; Nagy et al. 2017; Wu 2000; Xu et al. 2020; Yuan et al. 2023).

Phanerochaete P. Karst. is typified by P. alnea (Fr.) P. Karst. and has a worldwide distribution (Spirin et al. 2017). It is characterized by resupinate, membranaceous basidiomata, smooth or tuberculate hymenial surface, a monomitic hyphal system, generative hyphae mostly simple septate, the presence of smooth or encrusted cystidia, thin-walled, non-amyloid, and acyanophilous basidiospores, and causing a white-rot (Wu 2000; Wu et al. 2010; Floudas and Hibbett 2015; Ghobad-Nejhad et al. 2015). Currently, MycoBank (https://www.mycobank.org/page/Simple%20names%20search) and Index Fungorum (https://www.indexfungorum.org/Names/Names.asp?pg=1) have registered 214 records and 202 records in Phanerochaete, respectively. About 100 species are currently accepted in Phanerochaete worldwide, of which 51 have been found in China (including the four new species presented here) as of June 2024 (Xu et al. 2020; Boonmee et al. 2021; Chen et al. 2021; Wang and Zhao 2021; Li et al. 2023; Yu et al. 2023; Zhang et al. 2023; Deng et al. 2024).

Taxonomists used the membranaceous nature of the basidiomata, the monomitic hyphal system, the presence of clamp connections (simple, double or multiple clamps per septum) or simple septa, and the presence of cystidia as characters for the delimitation of the genus. The simplicity of the morphological features characterise Phanerochaete and the existence of species with basidiomata that fulfil only some of these morphological criteria render the limits of the genus uncertain (Floudas and Hibbett 2015).

Recent phylogenetic studies show that Phanerochaete s.l. is polyphyletic, and several new genera have been introduced, some of them placed into different families, or even orders (Greslebin et al. 2004; Wu et al. 2010; Binder et al. 2013; Floudas and Hibbett 2015). Most Phanerochaete species are still retained in a monophyletic lineage (Phanerochaetaceae sensu Larsson 2007) within Polyporales, along with genera such as Hyphodermella, Phlebiopsis, and Rhizochaete (Larsson 2007; Wu et al. 2010; Floudas and Hibbett 2015; Miettinen et al. 2016). Most members of the genus are nested in the phlebioid clade, comprising a number of Phanerochaete species assembled in a highly supported clade, referred to as the core Phanerochaete clade (Wu et al. 2010; Floudas and Hibbett 2015; Justo et al. 2017; Chen et al. 2021).

During investigations on the wood-inhabiting fungi in the Xizang Autonomous Region, Zhejiang, and Yunnan Province of China, samples corresponding to Phanerochaete were collected, and four species were initially identified as potentially new by morphology. To clarify the placement and relationships of the four species, we carried out a phylogenetic and morphological studies on Phanerochaete in China.

Materials and methods

Morphological studies

The studied specimens were collected from wild forests and are deposited in the Fungarium of the Institute of Microbiology, Beijing Forestry University (BJFC). Morphological descriptions are based on field notes and voucher specimens. The microscopic analysis follows Spirin et al. (2017). Freehand sections were made from dried basidiomata and mounted in 2% (w/v) potassium hydroxide (KOH) to observe color changes. Sections were studied at a magnification of up to 1000× using a Nikon Eclipse 80i microscope and phase contrast illumination. Microscopic features and measurements were made from slide preparations stained with Cotton Blue and Melzer’s reagent. To represent the variation in the size of spores, 5% of measurements were excluded from each end of the range and are given in parentheses. In the description: KOH = 5% potassium hydroxide, IKI = Melzer’s reagent, IKI+ = amyloid or dextrinoid, IKI- = neither amyloid nor dextrinoid, CB = Cotton Blue, CB+ = cyanophilous in Cotton Blue, CB- = acyanophilous in Cotton Blue, L = arithmetic average of spore length, W = arithmetic average of spore width, Q = L/W ratios and n = number of basidiospores measured from given number of specimens. Colour terms follow Anonymous (1969) and Petersen (1996).

DNA extraction, amplification, and sequencing

A CTAB rapid plant genome extraction kit-DN14 (Aidlab Biotechnologies Co., Ltd, Beijing) was used to obtain DNA from dried specimens and to perform the polymerase chain reaction (PCR) according to the manufacturer’s instructions with some modifications (Shen et al. 2019; Sun et al. 2020). The internal transcribed spacer (ITS) and large subunit nuclear ribosomal RNA gene (nLSU) were amplified using the primer pairs ITS5/ITS4 and LR0R/LR7 (White et al. 1990; Hopple and Vilgalys 1999) (https://sites.duke.edu/vilgalyslab/rdna_primers_for_fungi/).

The PCR procedure for ITS was as follows: initial denaturation at 95 °C for 3 min, followed by 34 cycles at 94 °C for 40 s, annealing at 54 °C for 45 s and extension at 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 34 cycles of denaturation at 94 °C for 30 s, annealing at 50 °C for 1 min, and extension at 72 °C for 1.5 min, and a final extension at 72 °C for 10 min. The PCR products were purified and sequenced at the Beijing Genomics Institute (BGI), China, with the same primers. DNA sequencing was performed at the Beijing Genomics Institute and the newly generated sequences were deposited in GenBank (Sayers et al. 2024). All sequences analysed in this study are listed in Table 1. Sequences generated from this study were aligned manually with additional sequences downloaded from GenBank using AliView version 1.27 (Larsson 2014). The final ITS and nLSU datasets were subsequently aligned using MAFFT v.7 under the E-INS-i strategy with no cost for opening gaps and equal cost for transformations (command line: mafft –genafpair –maxiterate 1000) (Katoh and Standley 2013) and visualised in AliView. Alignments were spliced and transformed formats in Mesquite v.3.2. (Maddison and Maddison 2017). Multiple sequence alignments were trimmed by trimAI v.1.2 using the -htmlout-gt 0.8 -st option to deal with gaps, when necessary (Capella-Gutierrez et al. 2009).

Table 1.
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Names, specimen numbers, references, and corresponding GenBank accession numbers of the taxa used in the phylogenetic analysis of this study. [New species are shown in bold, * type material; type specimens of other species are shown in bold].

Species name Specimen No. GenBank accession No. Country References
ITS LSU
Phanerochaete aculeata Wu 1809278 MZ422786 MZ637178 China Chen et al. (2021)
P. aculeata GC 1703117 MZ422785 MZ637177 China Chen et al. (2021)
P. albida WEI 18365 MZ422789 MZ637180 China Chen et al. (2021)
P. albida GC 140714 MZ422788 MZ637179 China Chen et al. (2021)
P. alnea K. H. Larsson 12054 KX538924 — Norway Spirin et al. (2017)
P. alpina Wu 130861 MZ422790 MZ637182 China Chen et al. (2021)
P. alpina Wu 130877 MZ422791 MZ637183 China Chen et al. (2021)
P. arizonica RLG 10248 KP135170 KP135239 USA Floudas and Hibbett (2015)
P. australis He 6013 MT235656 MT248136 China Xu et al. (2020)
P. australis HHB 7105 KP135081 KP135240 USA Floudas and Hibbett (2015)
P. australosanguinea MA Fungi 91308 MH233925 MH233928 Chile Phookamsak et al. (2019)
P. australosanguinea MA Fungi 91309 MH233926 MH233929 Chile Phookamsak et al. (2019)
P. bambusicola He 3606 MT235657 MT248137 China Chen et al. (2021)
P. bambusicola Wu 0707-2 MF399404 MF399395 China Xu et al. (2020)
P. brunnea He 4192 MT235658 MT248138 China Phookamsak et al. (2019)
P. burdsallii RF9JR KU668973 — USA Yu et al. (2023)
P. burdsallii He 2066 MT235690 MT248177 USA Xu et al. (2020)
P. burtii HHB 4618 KP135117 KP135241 USA Floudas and Hibbett (2015)
P. burtii FD 171 KP135116 — USA Floudas and Hibbett (2015)
P. calotricha Vanhanen 382 KP135107 — USA Floudas and Hibbett (2015)
P. canobrunnea He 5726 MT235659 MT248139 Sri Lanka Phookamsak et al. (2019)
P. canobrunnea CHWC 150666 LC412095 LC412104 China Phookamsak et al. (2019)
P. canolutea LWZ 202109214a ON897909 ON885366 China Unpublished
P. canolutea TNM F14823 NR175166 NG153829 China Chen et al. (2021)
P. carnosa He 5172 MT235660 MT248140 China Phookamsak et al. (2019)
P. carnosa HHB 9195 KP135129 KP135242 USA Floudas and Hibbett (2015)
P. castanea Dai 24915* PP960566 PP960569 China Present study
P. castanea Dai 24916 PP960567 PP960570 China Present study
P. chrysosporium He 5778 MT235661 MT248141 Sri Lanka Phookamsak et al. (2019)
P. chrysosporium HHB 6251 KP135094 KP135246 USA Floudas and Hibbett (2015)
P. citrinoalba Dai 26584* PP779892 PP779887 China Present study
P. citrinorhizomorpha Dai 20753 PP960568 PP960571 China Present study
P. citrinorhizomorpha Dai 26417* PP779891 PP779886 China Present study
P. citrinosanguinea FP 105385 KP135100 KP135234 USA Floudas and Hibbett (2015)
P. concrescens He 4657 MT235662 MT248142 China Chen et al. (2021)
P. concrescens H Spirin 7322 KP994380 KP994382 Russia Volobuev et al. (2015)
P. conifericola OM 8110 KP135171 — Finland Floudas and Hibbett (2015)
P. crystallina Chen 3823 MZ422802 MZ637188 China Chen et al. (2021)
P. crystallina Chen 3576 MZ422801 — China Chen et al. (2021)
P. cumulodentata He 2995 MT235664 MT248144 China Xu et al. (2020)
P. cumulodentata LERUS 298935 KP994359 KP994386 Russia Volobuev et al. (2015)
P. cystidiata He 4224 MT235665 MT248145 China Phookamsak et al. (2019)
P. cystidiata Wu 1708-326 LC412097 LC412100 China Wu et al. (2018)
P. ericina HHB 2288 KP135167 KP135247 USA Floudas and Hibbett (2015)
P. ericina He 4285 MT235666 MT248146 China Phookamsak et al. (2019)
P. fusca Wu 1409-163 LC412099 LC412106 China Wu et al. (2018)
P. guangdongensis Wu 1809-348 MZ422813 MZ637199 China Chen et al. (2021)
P. guangdongensis Wu 1809-319 MZ422811 MZ637197 China Chen et al. (2021)
P. hainanensis He 3562 MT235692 MT248179 China Boonmee et al. (2021)
P. incarnata He 201207281 MT235669 MT248149 China Phookamsak et al. (2019)
P. incarnata WEI 16075 MF399406 MF399397 China Xu et al. (2020)
P. krikophora HHB 5796 KP135164 KP135268 USA Floudas and Hibbett (2015)
P. laevis He 20120917-8 MT235670 MT248150 China Phookamsak et al. (2019)
P. laevis HHB 15519 KP135149 KP135249 USA Floudas and Hibbett (2015)
P. leptocystidiata He 5853 MT235685 MT248168 China Xu et al. (2020)
P. leptocystidiata Dai 10468 MT235684 MT248167 China Xu et al. (2020)
P. livescens He 5010 MT235671 MT248151 China Phookamsak et al. (2019)
P. magnoliae He 3321 MT235672 MT248152 China Phookamsak et al. (2019)
P. metuloidea He 2766 MT235682 MT248164 China Phookamsak et al. (2019)
P. minor He 3988 MT235686 MT248170 China Phookamsak et al. (2019)
P. parmastoi He 4570 MT235673 MT248153 China Phookamsak et al. (2019)
P. pruinosa CLZhao 7112 MZ435346 MZ435350 China Wang and Zhao (2021)
P. pruinosa CLZhao 7113 MZ435347 MZ435351 China Wang and Zhao (2021)
P. porostereoides He 1902 KX212217 KX212221 China Liu and He (2016)
P. pseudomagnoliae PP 25 KP135091 KP135250 South Africa Floudas and Hibbett (2015)
P. pseudosanguinea FD 244 KP135098 KP135251 USA Floudas and Hibbett (2015)
P. rhizomorpha GC 1708-335 MZ422824 MZ637208 China Chen et al. (2021)
P. rhizomorpha GC 1708-354 MZ422825 MZ637209 China Chen et al. (2021)
P. rhodella FD 18 KP135187 KP135258 USA Floudas and Hibbett (2015)
P. sanguineocarnosa FD-359 KP135122 KP135245 USA Floudas and Hibbett (2015)
P. sinensis He 4660 MT235688 MT248175 China Xu et al. (2020)
P. sinensis GC 180956 MT235689 MT248176 China Xu et al. (2020)
P. singularis He1873 KX212220 KX212224 China Liu and He (2016)
P. spadicea Wu 0504-15 MZ422837 MZ637219 China Chen et al. (2021)
P. spadicea Wu 0504-11 MZ422836 — China Chen et al. (2021)
P. stereoides He 5824 MT235677 MT248158 Sri Lanka Phookamsak et al. (2019)
P. stereoides He 2309 KX212219 KX212223 China Liu and He (2016)
P. subcarnosa Wu 9310-3 MZ422841 GQ470642 China Wu et al. (2010)
P. subcarnosa GC 1809-90 MZ422840 MZ637222 China Chen et al. (2021)
P. subceracea HHB-9434 KP135163 — USA Floudas and Hibbett (2015)
P. subrosea He 2421 MT235687 MT248174 China Phookamsak et al. (2019)
P. subsanguinea CLZhao 10470 MZ435348 MZ435352 China Wang and Zhao (2021)
P. subsanguinea CLZhao 10477 MZ435349 MZ435353 China Wang and Zhao (2021)
P. subtuberculata CLZhao F5130 OP605484 OQ195088 China Yu et al. (2023)
P. subtuberculata CLZhao F6838 OP605485 OQ195087 China Yu et al. (2023)
P. taiwaniana He 5269 MT235680 MT248161 VietNam Phookamsak et al. (2019)
P. taiwaniana Wu 011213 MF399412 MF399403 China Xu et al. (2020)
P. subtropica CLZhao F2763 OP605518 OQ195090 China Yu et al. (2023)
P. subtropica CLZhao F8716 OP605486 OQ195089 China Yu et al. (2023)
P. velutina He 3079 MT235681 MT248162 China Phookamsak et al. (2019)
P. velutina Kotiranta 25567 KP994354 KP994387 Russia Volobuev et al. (2015)
P. wuyiensis Dai 25530* PP779888 PP779883 China Present study
P. wuyiensis Dai 26246 PP779889 PP779884 China Present study
P. wuyiensis Dai 26250 PP779890 PP779885 China Present study
P. yunnanensis He 2719 MT235683 MT248166 China Xu et al. (2020)
Riopa metamorphosa Viacheslav Spirin 2395 KX752601 KX752601 Russia Miettinen et al. (2016)
R. pudens Otto Miettinen 8772 KX752598 — USA Miettinen et al. (2016)

Phylogenetic analyses

The two-marker DNA multiple sequence alignment (ITS+nLSU) was used to determine the phylogenetic position of the new species. The multiple sequence alignments and the retrieved topologies were deposited in Figshare (https://figshare.com/) under accession DOI: 10.6084/m9.figshare.27683265. Sequences of Riopa metamorphosa (Fuckel) Miettinen & Spirin and R. pudens Miettinen, obtained from GenBank, were used as the outgroups (Miettinen et al. 2016). The phylogenetic analyses followed the approach of Han et al. (2015) and Zhu et al. (2019). Maximum Likelihood (ML) and Bayesian Inference (BI) analyses were performed based on the ITS+nLSU datasets.

Sequences were analysed using Maximum Likelihood (ML) with RAxML-HPC2 through the CIPRES Science Gateway (www.phylo.org; Miller et al. 2010). Branch support (BT) for ML analysis was determined by 1,000 bootstrap replicates. Bayesian phylogenetic inference and Bayesian Posterior Probabilities (BPP) were computed with MrBayes 3.2.6 (Ronquist and Huelsenbeck 2003). Four Markov chains were run for 1.6 million generations (two-marker dataset) until the split deviation frequency value was less than 0.01 and trees were sampled every 100 generations. The first 25% of the sampled trees were discarded as burn-in and the remaining ones were used to reconstruct a majority rule consensus and calculate Bayesian Posterior Probabilities (BPP) of the clades. All trees were viewed in FigTree v. 1.4.3 (http://tree.bio.ed.ac.uk/software/figtree/). Branches that received bootstrap support for ML (≥ 75% (ML-BS)) and BPP (≥ 0.95 BPP) were considered as significantly supported. The ML bootstrap (ML) ≥ 50% and BBP (BPP) ≥ 0.90 are presented on topologies from ML analysis, respectively.

Results

Molecular phylogeny

The combined two-marker dataset (ITS+nLSU) included sequences from 97 samples representing 61 taxa. The phylogenetic reconstruction performed with Maximum Likelihood (ML) and Bayesian Inference (BI) analyses for the combined dataset showed similar topology and few differences in statistical support. The best model-fit applied in the Bayesian analysis was GTR+I+G, lset nst = 6, rates = invgamma and prset statefreqpr = dirichlet (1, 1, 1, 1). Bayesian analysis resulted in a nearly congruent topology with an average standard deviation of split frequencies = 0.014712 to ML analysis and thus, only the ML tree is shown (Fig. 1).

Figure 1. 

ML analysis of Phanerochaete based on dataset of ITS+nLSU. ML bootstrap values equal to or higher than 50% and Bayesian posterior probabilities values equal to or higher than 0.90 are shown. New taxa are in bold, * represents type material and in blue colour. Type specimens for all species are in bold.

The phylogenetic tree inferred from the ITS+nLSU sequences indicated that the four new species belonged to Phanerochaete (Fig. 1). In addition, Phanerochaete castanea grouped together with P. metuloidea Y.L. Xu & S.H. He with high support (ML = 100, BPP = 1.00); P. citrinoalba was sister to P. burtii (Romell ex Burt) Parmasto, with a low support (65% BS); P. citrinorhizomorpha grouped together with P. minor Y.L. Xu & S.H. He, with a high support (ML = 100, BPP = 1.00); and P. wuyiensis was sister to P. subtropica J. Yu & C.L. Zhao with high support (ML = 100, BPP = 1.00).

Taxonomy

Phanerochaete castanea K.Y. Luo, Yuan Yuan, Y.C. Dai & Xin Zhang, sp. nov.

MycoBank No: 854762
Figs 2, 3

Holotype

China • Zhejiang Province, Jinhua, Wuyi County, Niutoushan Forest Park; on a rotten bamboo; 17.VI.2023; Y.C. Dai 24915 (BJFC042468).

Figure 2. 

Basidiomata of Phanerochaete castanea (Holotype, Dai 24915). Scale bar: 0.5 cm.

Etymology

Castanea (Lat.) refers to the colour of new species’ basidiomata turning reddish brown in KOH.

Figure 3. 

Microscopic structures of Phanerochaete castanea (Holotype, Dai 24915) a basidiospores b cystidia c basidia and basidioles d vertical section of the subiculum e vertical section of the hymenium and subhymenium.

Description

Basidiomata annual, resupinate, adnate, soft coriaceous, without odor and taste when fresh, detachable from substrate, up to 2 cm long, 1.5 cm wide, 100–200 µm thick. Hymenial surface smooth, whitish to yellowish when fresh, yellowish brown upon drying, becoming reddish brown in KOH. Sterile margins paler than hymenial surface, thinning out, usually with whitish rhizomorphs, up to 0.2 cm.

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

Subicular hyphae hyaline, thick-walled, up to 0.1 µm thick, simple septate, occasionally bearing clamp connections, occasionally branched, parallel to interwoven, 3–5 µm in diameter. Subhymenial hyphae hyaline, thick-walled, clampless, 1.5–3 µm in diameter.

Cystidia mostly subulate with a blunt or acute apex, 45–60 × 6–8 µm, hyaline, thick-walled, up to 0.1 µm thick, with a simple septum at the base, mostly encrusted with crystal granules at apical part, some with smooth apex, projecting up to 15 µm above the hymenial layer; cystidioles absent. Basidia clavate, with four sterigmata and a basal simple septum, 19–27 × 3–5.5 µm; basidioles of similar shape to basidia, but smaller.

Basidiospores ellipsoid, hyaline, thin-walled, smooth, usually with one or two medium guttules, IKI–, CB–, (3.7–)3.9–5.1(–5.2) × (2.2–)2.3–3.1 µm, L = 4.45 μm, W = 2.66 μm, Q = 1.67–1.68 (n=60/2).

Type of rot

White rot.

Additional specimen examined (paratype)

China • Zhejiang Province, Jinhua, Wuyi County, Niutoushan Forest Park; on a rotten angiosperm wood; 17.VI.2023; Y.C. Dai 24916 (BJFC042469).

Phanerochaete citrinoalba K.Y. Luo, Yuan Yuan, Y.C. Dai & Xin Zhang, sp. nov.

MycoBank No: 854772
Figs 4, 5

Holotype

China • Xizang Autonomous Region, Nyingchi, Sejila Mountain; on dead bamboo; 23.X.2023; Dai 26584 (BJFC044134).

Figure 4. 

Basidiomata of Phanerochaete citrinoalba (Holotype, Dai 26584). Scale bar: 1 cm.

Etymology

Citrinoalba (Lat.) refers to the species having yellowish to whitish rhizomorphs.

Figure 5. 

Microscopic structures of Phanerochaete citrinoalba (Holotype, Dai 26584) a basidiospores b basidia and basidioles c vertical section of the subiculum d vertical section of the hymenium and subhymenium.

Description

Basidiomata annual, resupinate, adnate, coriaceous, without odour and taste when fresh, up to 9 cm long, 1.5 cm wide, 70–130 µm thick. Hymenial surfaces smooth, cracking, white to cream when fresh, cream to slightly buff upon drying. Becoming greyish brown in KOH. Sterile margins distinct, concolorous with hymenial surface, usually with yellowish to whitish rhizomorphas, and up to 5 mm.

Hyphal system monomitic, generative hyphae mostly with simple septa, occasionally with clamp connections in subiculum, IKI–, CB–; tissue unchanged in KOH.

Subicular hyphae hyaline, thin- to thick-walled, frequently simple septate, occasionally bearing clamp connections, branched at acute angles, subparallel to interwoven, 3–6 µm in diameter. Subhymenial hyphae hyaline, thin- to thick-walled, clampless, 2–3.5 µm in diameter, encrusted with crystal granules.

Cystidia and cystidioles absent. Subhymenium frequently with crystal granules. Basidia clavate, with four sterigmata and a basal simple septum, 17–25 × 4–7 µm; basidioles of similar shape to basidia, but smaller.

Basidiospores ellipsoid to oblong ellipsoid, hyaline, thin-walled, smooth, some with a medium guttule, IKI–, CB–, (4.7–)4.9–6.3(–6.4) × (2.2–)2.4–3(–3.1) µm, L = 5.49 μm, W = 2.66 μm, Q = 2.06 (n = 30/1).

Type of rot

White rot.

Phanerochaete citrinorhizomorpha K.Y. Luo, Yuan Yuan, Y.C. Dai & Xin Zhang, sp. nov.

MycoBank No: 854773
Figs 6, 7

Holotype

China • Zhejiang Province, Jinhua, Wuyi County, Xinzhai, Daozhi Village; on dead bamboo; 14.X.2023; Y.C. Dai 26417 (BJFC043967).

Figure 6. 

Basidiomata of Phanerochaete citrinorhizomorpha (Holotype, Dai 26417). Scale bar: 2 cm.

Etymology

Citrinorhizomorpha (Lat.) refers to the new species having yellowish rhizomorphs.

Figure 7. 

Microscopic structures of Phanerochaete citrinorhizomorpha (Holotype, Dai 26417) a basidiospores b basidia and basidioles c cystidia d vertical section of the subiculum e vertical section of the hymenium and subhymenium.

Description

Basidiomata annual, resupinate, adnate, soft coriaceous, without odor and taste when fresh, up to 7 cm long, 5 cm wide, 100–200 µm thick. Hymenial surfaces flesh-pink when juvenile, salmon to peach with age. Becoming purple in KOH. Sterile margins paler than hymenial surface, thinning out, usually with yellowish rhizomorphs, and up to 4 cm.

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

Subicular hyphae hyaline, thick-walled, up to 0.1 µm thick, simple septate, frequently branched at acute angles, interwoven, 2–6 µm in diameter. Subhymenial hyphae hyaline, thick-walled, 1.5–3 µm in diameter, encrusted with crystal granules.

Cystidia mostly subulate with an obtuse apex, hyaline, thick-walled, up to 0.1 µm thick, with a simple septum at the base, usually encrusted with crystal granules, projecting above hymenium, projecting up to 17 µm above the hymenial layer, 18–36 × 4–6 µm; cystidioles absent. Basidia clavate, with four sterigmata and a basal simple septum, 12–17 × 4–6 µm; basidioles of similar shape to basidia, but smaller.

Basidiospores ellipsoid, hyaline, thin-walled, smooth, usually with a medium guttule, IKI–, CB–, (3.3–)3.5–4.5(–4.9) × (1.8–)1.9–2.8(–3.2) µm, L = 3.94 μm, W = 2.37 μm, Q = 1.54–1.80 (n = 60/2).

Type of rot

White rot.

Additional specimen examined (paratype)

China • Yunnan Province, Honghe, Jinping County, Fenshuiling Nature Reserve; on a fallen angiosperm branch; 18.VIII.2019; Y.C. Dai 20753 (BJFC032420).

Phanerochaete wuyiensis K.Y. Luo, Yuan Yuan, Y.C. Dai & Xin Zhang, sp. nov.

MycoBank No: 854774
Figs 8, 9

Holotype

China • Zhejiang Province, Jinhua, Wuyi County, Shiehu Scenic Spot; on a fallen branch of Pinus massoniana; 11.VIII.2023; Y.C. Dai 25530 (BJFC043078).

Figure 8. 

A basidioma of Phanerochaete wuyiensis (Holotype, Dai 25530). Scale bar: 2 cm.

Etymology

Wuyiensis (Lat.) refers to “Wuyi County, Zhejiang Province, East China,” where the holotype was found.

Figure 9. 

Microscopic structures of Phanerochaete wuyiensis (Holotype, Dai 25530) a basidiospores b basidia and basidioles c cystidia d vertical section of the subiculum e vertical section of the hymenium and subhymenium.

Description

Basidiomata annual, resupinate, adnate, detachable from substrate, membranaceous, without odour and taste when fresh, up to 8 cm long, 3 cm wide, 200–300 µm thick. Hymenial surfaces smooth or locally tuberculate, uncracked, whitish when fresh and upon drying. Becoming lemon-yellow in KOH. Sterile margins distinct, concolorous with hymenial surface, with whitish rhizomorphs, and up to 1 cm.

Hyphal system monomitic, generative hyphae with simple septa and clamp connections, IKI–, CB–; tissue unchanged in KOH.

Subicular generative hyphae hyaline, thick-walled with simple septa and clamp connections, usually constricted at simple septa, sometimes with three branches at a single septum, interwoven, 3–16 µm in diameter. Subhymenial hyphae hyaline, thick-walled clampless, branched present, 2–3.5 µm in diameter.

Cystidia clavate to fusiform, hyaline, thin-walled, with a simple septum at the base, some apically encrusted with crystal granules, projecting above hymenium, 22–44 × 6–10 µm; cystidioles absent. Basidia clavate, with four sterigmata and a basal simple septum, 13–25 × 4–7 µm; basidioles of similar shape to basidia, but smaller.

Basidiospores ellipsoid, hyaline, thin-walled, smooth, with one or two medium guttules, IKI–, CB–, (3.3–)3.6–4.6(–5.4) × (1.8–)2.1–3.2(–3.4) µm, L = 4.07 μm, W = 2.48 μm, Q = 1.61–1.68 (n = 90/3).

Type of rot

White rot.

Additional specimens examined (paratype)

China • Zhejiang Province, Wuyi County, Shiehu Scenic Spot; on a fallen branch of Pinus massoniana; 12.X.2023; Y.C. Dai 26246 (BJFC043796) • ibid. on rotten wood of Pinus massoniana; 12.X.2023; Y.C. Dai 26250 (BJFC043800).

Discussion

Taxa of the genus Phanerochaete are important components of woody plant ecosystems, and they have the ability to decompose rotten wood in forest or bamboo ecosystems. Many species in the genus have been described from the subtropics and tropics in recent years (Chen et al. 2021; Li et al. 2023; Yu et al. 2023; Zhang et al. 2023; Deng et al. 2024). In the present study, four new species, viz. Phanerochaete castanea, P. citrinoalba, P. citrinorhizomorpha and P. wuyiensis, are described based on a combination of morphological features and molecular evidence.

Phylogenetically, based on ITS+nLSU topology (Fig. 1), four new species were nested in the Phanerochaete clade. Among them, P. castanea grouped together with P. metuloidea, however, P. metuloidea is delimited from P. castanea by having greyish orange, brownish orange to light brown hymenophore and longer basidia (40–70 × 5–8.5 μm vs. 19–27 × 3–5.5 µm, Xu et al. 2020). Phanerochaete citrinoalba was sister to P. burtii, but P. burtii differs from P. citrinoalba by having cylindric cystidia (Parmasto 1967). P. citrinorhizomorpha grouped together with P. minor, however, P. minor is different from P. citrinorhizomorpha by having membranaceous basidiomata and apically encrusted cystidia (Xu et al. 2020). Phanerochaete wuyiensis was sister to P. subtropica, but P. subtropica differs from P. wuyiensis by its coriaceous basidiomata (Yu et al. 2023).

Morphologically, Phanerochaete castanea resembles P. burdsallii Y.L. Xu et al., P. hymenochaetoides Y.L. Xu & S.H. He and P. laevis (Fr.) J. Erikss. & Ryvarden by sharing hymenophore becoming reddish brown or red in KOH. However, P. burdsallii is different from P. castanea by its membranaceous basidiomata and longer basidiospores (5.3–6 × 2.5–3 μm vs. 3.9–5.1 × 2.3–3.1 µm, Xu et al. 2020); P. hymenochaetoides is distinguished from P. castanea by its basidiomata without rhizomorphs and subicular hyphae encrusted with yellow resinous granules (Xu et al. 2020); P. laevis is different from P. castanea by its membranaceous basidiomata, ochraceous to cinnamon hymenial surface, and longer basidia (30–50 × 4–5 μm vs. 19–27 × 3–5.5 µm, Eriksson et al. 1978).

Phanerochaete citrinoalba resembles P. daliensis J. Yu & C.L. Zhao, P. subtuberculata J. Yu & C.L. Zhao and P. tongbiguanensis Y.L. Deng & C.L. Zhao by sharing a coriaceous basidiomata. However, P. daliensis is different from P. citrinoalba by its grandinioid hymenial surface and thick-walled basidiospores (Yu et al. 2023); P. subtuberculata is distinguished from P. citrinoalba by having tuberculate hymenial surface, generative hyphae with simple septa, and clavate cystidia (Yu et al. 2023); and P. tongbiguanensis is different from P. citrinoalba by its generative hyphae with simple septa, subclavate cystidia and bigger basidiospores (6–9 × 3–4.5 μm vs. 4.9–6.3 × 2.4–3 µm, Deng et al. 2024).

Phanerochaete citrinorhizomorpha is similar to P. cinerea Y.L. Xu & S.H. He, P. guangdongensis C.C. Chen et al., and P. spadicea C.C. Chen & Sheng H. Wu by sharing generative hyphae with simple septa. However, P. cinerea is distinguished from P. citrinorhizomorpha by having grey, brownish grey to greyish brown hymenophore, the absence of cystidia, and longer basidiospores (4.8–5.6 × 2–2.5 μm vs. 3.5–4.5 × 1.9–2.8 µm, Xu et al. 2020); P. guangdongensis is different from P. citrinorhizomorpha by its membranaceous to subceraceous basidiomata, buff to yellowish brown hymenial surface, and longer basidiospores (6.9–7.8 × 2.6–3 μm vs. 3.5–4.5 × 1.9–2.8 µm, Chen et al. 2021); P. spadicea is distinguished from P. citrinorhizomorpha by having membranaceous basidiomata, buff to pale brown hymenial surface, and longer basidiospores (4.5–5 × 1.9–2.2 μm vs. 3.5–4.5 × 1.9–2.8 µm, Chen et al. 2021).

Phanerochaete wuyiensis is similar to P. burdsallii Y.L. Xu et al., P. crystallina C.C. Chen et al., and P. subrosea Y.L. Xu & S.H. He by sharing membranaceous basidiomata. However, P. burdsallii is different from P. wuyiensis by its hymenophore becoming reddish brown in KOH and longer basidiospores (5.3–6 × 2.5–3 µm vs. 3.6–4.6 × 2.1–3.2 µm, Xu et al. 2020); P. crystallina is different from P. wuyiensis by having cream to ochraceous-buff hymenial surface and longer basidiospores (5.1–5.7 × 2.2–2.5 µm vs. 3.6–4.6 × 2.1–3.2 µm, Chen et al. 2021); and P. subrosea is readily distinguished from P. wuyiensis by its hymenophore turning purple in KOH and longer basidiospores (5–6 × 2.5–3 µm vs. 3.6–4.6 × 2.1–3.2 µm, Xu et al. 2020).

Xizang Autonomous Region, Yunnan Province in southwest China, and Zhejiang Province in eastern China are very rich for wood-inhabiting fungi. Numerous taxa of such fungi have been described from these areas based on morphological and molecular phylogenetic analyses (Volobuev et al. 2015; Chen et al. 2018; Ordynets et al. 2018; Cui et al. 2019; Dai et al. 2021; Wu et al. 2022a, 2022b; Zhang et al. 2023; Zhou et al. 2023; Zhao et al. 2024). DNA sequence data are very useful in exploring cryptic taxa and the diversity of corticioid fungi. In the present study, four new species of Phanerochaete are described from these two areas, which improve our knowledge of the diversity of Chinese white rot fungi.

Key to the accepted species of Phanerochaete in China

1 Hymenophore poroid P. inflata
– Hymenophore non-poroid 2
2 Hymenophore grandinioid 3
– Hymenophore smooth to raduloid 5
3 Basidiospores thick-walled P. daliensis
– Basidiospores thin-walled 4
4 Cystidia present P. aculeata
– Cystidia absent P. yunnanensis
5 Hymenophore at first smooth, odontioid to raduloid when mature 6
– Hymenophore smooth, more or less tuberculate 7
6 Distributed in northern China P. cumulodentata
– Distributed in southern China P. magnoliae
7 Rhizomorpha present 8
– Rhizomorpha absent 19
8 Hymenophore purple in KOH 9
– Hymenophore unchanged, or becoming buff, grayish brown, brown, reddish brown, red or black in KOH 10
9 Cystidia without crystal granules P. subrosea
– Cystidia with crystal granules P. citrinorhizomorpha
10 Hyphal cords reddish brown P. citrinosanguinea
– Hyphal cords white, cream, grayish or orange 11
11 Cystidia absent P. citrinoalba
– Cystidia present 12
12 Cystidia obviously encrusted with crystals P. laevis
– Cystidia smooth or sparsely encrusted 13
13 Generative hyphae without clamp connections in subiculum P. subsanguinea
– Generative hyphae with clamp connections in subiculum 14
14 Cystidia thick-walled 15
– Cystidia thin-walled 16
15 Cystidia with septa P. subtropica
– Cystidia without septa P. castanea
16 Hyphal cords turning reddish brown in KOH 17
– Hyphal cords not turning reddish brown in KOH 18
17 Cystidia 30–70 × 4–6 µm; basidiospores 5–6 × 2.5–3 µm P. leptocystidiata
– Cystidia 35–50 × 4–6 µm; basidiospores 4–5 × 2–2.5 µm P. sinensis
18 Basidiomata buff in KOH P. shenghuaii
– Basidiomata darkening in KOH P. rhizomorpha
19 Cystidia absent 20
– Cystidia present 24
20 Hymenophore brown P. porostereoides
– Hymenophore whitish, cream, gray, grayish brown, or yellowish 21
21 Hymenial surface lightly darkening in KOH 22
– Hymenial surface unchanged in KOH 23
22 Basidiomata undetachable from substrate P. pruinosa
– Basidiomata detachable from substrate P. cinerea
23 Basidiospores 4.2–5.2 × 1.8–2.2 µm P. spadicea
– Basidiospores 4.2–5.1 × 2.5–3.3 µm P. brunnea
24 Cystidia obviously encrusted 25
– Cystidia smooth or sparsely encrusted 36
25 Cystidia encrusted with yellow resinous granules 26
– Cystidia encrusted without yellow resinous granules 27
26 Hymenophore brown; quasi-binding hyphae present P. ericina
– Hymenophore lilac pink; quasi-binding hyphae absent P. incarnata
27 On Monocotyledons P. minor
– On Dicotyledons 28
28 Cystidia up to 150 µm long P. velutina
– Cystidia up to 80 µm long 29
29 Cystidia up to 13 µm wide 30
– Cystidia up to 9 µm wide 31
30 Cystidia only apically encrusted; widely distributed in China P. concrescens
– Cystidia encrusted up to one third of the length; distributed only in southern China P. australis
31 Hymenophore yellow to buff 32
– Hymenophore white to cream 34
32 Basidiomata ceraceous; basidiospores > 5.5 µm long P. livescens
– Basidiomata membranaceous; basidiospores < 5.5 µm long 33
33 Hymenophore yellow to yellowish brown; margin determinate P. hymenochaetoides
– Hymenophore cream to light yellow; margin fibrillose P. cystidiata
34 Cystidia thick-walled; basidia up to 70 µm long, 8.5 µm wide P. metuloidea
– Cystidia thin- to slightly thick-walled; basidia up to 50 µm long, 6 µm wide 35
35 Subicular hyphae thin to slightly thick-walled; cystidia subulate P. laevis
– Subicular hyphae thick-walled; cystidia tapering but with obtuse apex P. sordida
36 Cystidia two kinds P. robusta
– Cystidia one kind 37
37 Clamp connections present in subiculum 38
– Clamp connections absent in subiculum 43
38 Basidiospores < 4.6 µm in length 39
– Basidiospores > 4.6 µm in length 41
39 Cystidia < 40 µm in length P. albida
– Cystidia > 40 µm in length 40
40 Basidiomata turning lemon-yellow in KOH P. wuyiensis
– Basidiomata turning greyish green in KOH P. carnosa
41 Cystidia < 40 µm in length P. subcarnosa
– Cystidia > 40 µm in length 42
42 Basidiomata margin fibrillose P. affinis
– Basidiomata margin byssoid P. alpina
43 On bamboo 44
– On wood 45
44 Basidiospores > 6.2 µm in length P. bambucicola
– Basidiospores < 6.2 µm in length P. parmastoi
45 Basidiospores cylindrical 46
– Basidiospores ellipsoid 48
46 Basidiospores > 6 µm in length P. guangdongensis
– Basidiospores < 6 µm in length 47
47 Basidiomata coriaceous P. subtuberculata
– Basidiomata membranaceous P. crystallina
48 Basidiomata coriaceous to soft corky P. hainanensis
– Basidiomata membranaceous 49
49 Basidiospores < 6.5 µm in length P. tongbiguanensis
– Basidiospores > 6.5 µm in length 50
50 Basidiospores with oil-drops P. taiwaniana
– Basidiospores without oil-drops P. stereoides

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

This study was supported by the National Natural Science Foundation of China (Project Nos. 32370013), the Young Elite Scientists Sponsorship Program by CAST (Project No. 2023QNRC001), Zhejiang Key Laboratory of Biological Breeding and Exploitation of Edible and Medicinal Mushrooms, and the Postdoctoral Fellowship Program (Grade C) of China Postdoctoral Science Foundation (GZC20230254).

Author contributions

Conceptualization, Y. Yuan, K.Y. Luo and Y.C. Dai; methodology, Y. Yuan, K.Y. Luo and X. Zhang; software, Y. Yuan, K.Y. Luo and X. Zhang; formal analysis, K.Y. Luo and Y. Yuan; investigation, Y. Yuan, K.Y. Luo and Y.C. Dai; resources, Y. Yuan, Y.C. Dai; writing – original draft preparation, K.Y. Luo; writing – review and editing, Y.C. Dai; visualization, X. Zhang, Y. Yuan; supervision, Y. Yuan, Y.C. Dai; project administration, Y. Yuan and Y.C. Dai; funding acquisition, Y. Yuan. All authors have read and agreed to the published version of the manuscript.

Author ORCIDs

Kai-Yue Luo https://orcid.org/0000-0001-6145-4864

Xin Zhang https://orcid.org/0009-0005-8363-7852

Yu-Cheng Dai https://orcid.org/0000-0002-6523-0320

Yuan Yuan https://orcid.org/0000-0001-6674-9848

Data availability

All of the data that support the findings of this study are available in the main text.

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Kai-Yue Luo and Xin Zhang contributed equally to this work.
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