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Research Article
Morphological and molecular analyses revealed four new wood-inhabiting fungal species (Hymenochaetales, Basidiomycota) from Yunnan
expand article infoYinglian Deng‡, Meng Chen‡, Linfeng Liu‡, Qizhen Li‡, Sicheng Zhang‡, Haisheng Yuan§, Changlin Zhao‡
‡ Southwest Forestry University, Kunming, China
§ Institute of Applied Ecology, Chinese Academy of Sciences, Shenyang, China

Corresponding author: Haisheng Yuan ( hsyuan@iae.ac.cn )

Corresponding author: Changlin Zhao ( fungichanglinz@163.com )

Academic editor: Danushka Sandaruwan Tennakoon
© 2025 Yinglian Deng, Meng Chen, Linfeng Liu, Qizhen Li, Sicheng Zhang, Haisheng Yuan, Changlin Zhao.
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: Deng Y, Chen M, Liu L, Li Q, Zhang S, Yuan H, Zhao C (2025) Morphological and molecular analyses revealed four new wood-inhabiting fungal species (Hymenochaetales, Basidiomycota) from Yunnan. MycoKeys 117: 29-66. https://doi.org/10.3897/mycokeys.117.146236
Open Access

Abstract

Hymenochaetales is one of the fungal orders mainly composed of wood-inhabiting macrofungi within the class Agaricomycetes, Basidiomycota. Four new Hymenochaetales wood-inhabiting fungi, Hymenochaete bannaensis, Lyomyces asiaticus, Peniophorella albohymenia, and P. punctata collected from China are proposed based on morphological characteristics and molecular evidence. H. bannaensis is distinguished by flocculent basidiomata with cinnamon to yellowish brown to rust-brown hymenial surface, generative hyphae with simple septa and broadly ellipsoid to globose basidiospores. L. asiaticus is characterized by the membranaceous basidiomata with white to cream hymenial surface with tuberculate, a monomitic hyphal system with clamped generative hyphae and ellipsoid basidiospores. In addition, P. albohymenia is delimited by membranaceous basidiomata with white hymenial surface, four types of cystidia as stephanocyst, fusiform, cylindrical and capitate, and ellipsoid basidiospores. P. punctata is unique in the membranaceous, punctate basidiomata with white to pale yellow hymenial surface, fusiform cystidia, and allantoid basidiospores. Sequences of ITS and nLSU rRNA markers based on phylogenetic analyses were performed using the Maximum Likelihood, Maximum Parsimony and Bayesian Inference methods.

Key words:

Biodiversity, new taxa, phylogenetic analyses, Southwestern China, taxonomy

Introduction

In terms of taxonomic status, wood inhabiting fungi mainly refer to the classification within the class Agaricomycetes, Basidiomycota (James et al. 2020; Dai et al. 2021; Wijayawardene et al. 2022; Yuan et al. 2023; Zhao et al. 2024; Dai et al. 2025). Most of basidiomycetes are white-rot fungi, which are responsible for degradation of the lignin in the forest ecosystems (Floudas et al. 2012; Osman and El-Nuby 2023). Rot type can be inferred from direct examination of wood substrates or from cultural tests, in which it was classified to three groups, brown rot, soft rot, and white rot fungi, and all species of Hymenochaetales cause white rot (Justo et al. 2017; Wang et al. 2023a).

Hymenochaetales is one of the fungal orders being mainly composed of wood-inhabiting fungi within Agaricomycetes, Basidiomycota (Wang et al. 2023b; Zhou et al. 2023; Dai et al. 2025). This order includes fascinating basidiomycete fungi that occupies a distinct ecological niche, in which there are many variations of the basidiomata types within Hymenochaetales (Larsson et al. 2006; Wu et al. 2019a; Guan et al. 2023). Hymenochaetales comprises a number of representative wood-inhabiting fungal taxa, which includes diverse hymenophoral morphologies as hydnoid, corticioid, and polyporoid (Zhao et al. 2023a; Yuan et al. 2024). Currently, 14 families, 83 genera, and 1205 species, are accepted in Hymenochaetales, and include wood-inhabiting and ectomycorrhizal fungi (Hibbett et al. 2014; Wang et al. 2023a; He et al. 2024). However, the species diversity has been well explored worldwide, therefore more new taxa need to be discovered. Furthermore, the order has various morphological characters for basidiomata, in which all characteristics are also found in the other orders (Polyporales, Russulales) in Agaricomycetes (Wu and Yuan 2020).

On the basis of the frequent inclusion of data from DNA sequences in many phylogenetic studies, the classification of the wood-inhabiting fungi has been updated continuously (Yurchenko et al. 2020a; Mao et al. 2023; Zhang et al. 2023b). Wang et al. (2023a) implemented a multi-locus phylogenetic analysis on the basis of nSSU, ITS, nLSU, mt-SSU, tef1α, rpb1, and rpb2 genes and updated the taxonomic framework of Hymenochaetales. The results have shown that 45 genera were accepted in Hymenochaetaceae, including the genus Hymenochaete (Wang et al. 2023a), and Hymenochaetaceae comprised three major genera Phellinus sensu lato, Hymenochaete and Inonotus sensu lato (Larsen and Cobb-Poulle 1990; Léger 1998; Ryvarden 2005; Wu et al. 2022a). The results showed that the genus Hymenochaete (Hymenochaetaceae) was closely grouped with Hymenochaetopsis S.H. He & Jiao Yang, and they were almost indistinguishable in morphology (Larsson et al. 2006; He and Dai 2012; Baltazar et al. 2014; Parmasto et al. 2014; Yang et al. 2016). According to the anatomical features, the genus has been divided into four sections, Hymenochaete, Fultochaete Escobar ex Léger, Gymnochaete Escobar ex Léger and Paragymnochaete Léger (Léger 1998; He and Dai 2012).

Molecular studies of Hyphoderma Wallr. revealed that the reinstatement of Peniophorella resulted in 19 new combinations in the genus Peniophorella and all species nested in the hymenochaetoid clade (Larsson 2007b). Miettinen et al. (2019) revealed that P. praetermissa (P. Karst.) K.H. Larss. and P. tsugae (Burt) K.H. Larss. grouped together and clustered in Hymenochaetales. Telleria et al. (2012) studied Hyphoderma and its discrimination from closely related taxa and indicated that eight Peniophorella taxa were closely grouped and distinct from Hyphoderma s.s. Justo et al. (2017) revised the family-level classification of the order Polyporales (Basidiomycota) and proposed that 20 species were classified in Peniophorella located in the family Hyphodermataceae (Hymenochaetales). Morphological studies and phylogenetic analyses reported six new taxa of Peniophorella and displayed the taxonomic status for the new taxa within the genus (Guan et al. 2020; Xu et al. 2020; Yurchenko et al. 2020a). Wang et al. (2023a) proposed that the genus Peniophorella bears affinity to a new family Peniophorellaceae.

Due to the lack of sequences of some wood-inhabiting fungal taxa, it is difficult to clearly distinguish many genera in Schizoporaceae, therefore, a broad concept of Hyphodontia s.l. was accepted (Yurchenko and Wu 2014; Riebesehl et al. 2015; Riebesehl and Langer 2017). Based on the nuclear DNA sequence data, six well-distinguished clades as the Hastodontia clade, the Hyphodontia clade, the Lagarobasidium clade, the Kneiffiella-Alutaceodontia clade, the Xylodon-Lyomyces-Rogersella clade and the Xylodon-Schizopora-Palifer clade, were included (Yurchenko and Wu 2014). Inferred from the phylogenetical studies for Hyphodontia s.l., the genus Lyomyces was nested within the Xylodon-Lyomyces-Rogersella clade (Yurchenko and Wu 2014). Riebesehl et al. (2019) clarified the generic concept and their phylogenetic reconstruction of Lyomyces and the species L. sambuci (Pers.) P. Karst. was sister to L. crustosus (Pers.) P. Karst (Riebesehl et al. 2019).

In this study, four taxa from three genera of Hymenochaetales were collected from Yunnan, China and the phylogenetic relationships are studied based on ITS-nLSU gene regions. Therefore, exploring the diversity and taxonomic status of these new taxa using different characters will be significant for wood-inhabiting fungi in Yunnan. Morphology and phylogenetic analyses reveal that these are new to science.

Materials and methods

Sample collection and morphology examination

Fresh basidiomata of the fungi were collected from Xishuangbanna and Zhaotong of Yunnan Province, China, from September 2023 to January 2024. Specimens were dried in an electric food dehydrator at 40 °C, then sealed and stored in an envelope bag and deposited in the herbarium of the Southwest Forestry University (SWFC), Kunming, Yunnan Province, China. Macromorphological descriptions are based on field notes and photos captured in the field and lab. Color terminology was followed as mentioned by Petersen (1996). Micromorphological data were obtained from the dried specimens when observed under a light microscope (Nikon Ni, Tokyo, Japan) following the previous study (Cui et al. 2019; Zhao et al. 2023b; Dong et al. 2024). The following abbreviations are 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 measured spores), W = mean spore width (arithmetic average for measured spores), Q = variation in the L/W ratios between the specimens studied and n = a/b (number of spores (a) measured from given number (b) of specimens). Standardized sampling of microstructures for measurement includes basidiospores (30), basidia, basidioles and cystidia (5), setae and hyphae diameter (10) (Cui et al. 2019; Zhao et al. 2023b; Dong et al. 2024).

Molecular phylogeny

The EZNA HP Fungal DNA Kit (Omega Biotechnologies Co., Ltd., Kunming, China) was used to extract DNA with some modifications from the dried specimens. The nuclear ribosomal ITS region was amplified with primers ITS5 and ITS4 (White et al. 1990). 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 nuclear nLSU region was amplified with primer pair LR0R and LR7 (Vilgalys and Hester 1990; Rehner and Samuels 1994). 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 procedure for ITS and nLSU followed previous study (Zhao and Wu 2017). All of the newly generated sequences were deposited in NCBI GenBank (https://www.ncbi.nlm.nih.gov/genbank/) (Table 1).

Table 1.
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List of species, specimens and GenBank accession numbers of sequences used in this study. [* Indicates type materials].

Species name Sample no. GenBank accessions no. References
ITS nLSU
Basidioradulum mayi LWZ 20180510-18 MN017785 MN017792 Wang et al. (2023a)
B. radula LWZ 20201017-62 ON063684 ON063884 Wang et al. (2023a)
Boletopsis leucomelaena PBM2678 DQ484064 DQ154112 Liu et al. (2022)
Fasciodontia brasiliensis MSK-F 7245a MK575201 MK598734 Yurchenko et al. (2020a)
F. bugellensis KAS-FD 10705a MK575203 MK598735 Yurchenko et al. (2020a)
F. bannaensis MUCL 46950 GU461943 EF429218 Nie et al. (2017)
F. gabonensis MUCL 47576 GU461971 GU461990 Wang et al. (2023a)
F. punctate MUCL 47629 GU461950 GU461982 Nie et al. (2017)
F. rhamnoides LWZ 20180905-15 ON063643 ON063842 Wang et al. (2023a)
Hydnoporia pinicola CLZhao 22505 OR094493 OR449924 Dong et al. (2024)
H. pinicola CLZhao 27154 OR094494 OR449925 Dong et al. (2024)
H. yunnanensis CLZhao 6123 * OR094498 OR449929 Dong et al. (2024)
H. yunnanensis CLZhao 6227 OR094499 OR449930 Dong et al. (2024)
Hymenochaete acerosa He 338 JQ279543 JQ279657 He et al. (2017)
H. adusta He 207 JQ279523 KU975497 He et al. (2017)
H. angustispora Dai 17045 MF370592 MF370598 He et al. (2017)
H. angustispora Dai 17049 MF370593 MF370599 He et al. (2017)
H. anomala He 592 JQ279566 JQ279650 He et al. (2017)
H. asetosa Dai 10756 JQ279559 JQ279642 He et al. (2017)
H. attenuate He 28 JQ279526 JQ279633 He et al. (2017)
H. bambusicola He 4116 KY425674 KY425681 He et al. (2017)
H. berteroi He 1488 KU975459 KU975498 He et al. (2017)
H. biformisetosa He 1445 KF908247 KU975499 Yang and He (2014)
H. boddingii MEH 66068 MN030343 MN030345 Du et al. (2021)
H. boddingii MEH 69996 MN030341 MN030347 Du et al. (2021)
H. borbonica CBS 731.86 MH862026 MH873716 Du et al. (2021)
H. cana He 1305 KF438169 KF438172 He et al. (2017)
H. cinnamomea He 755 JQ279548 JQ279658 He et al. (2017)
H. colliculosa Dai 16427 MF370595 MF370602 He et al. (2017)
H. colliculosa Dai 16428 MF370596 MF370603 He et al. (2017)
H. conchata MEH 70144 MF373838 — Du et al. (2021)
H. contiformis He 1166 KU975461 KU975501 He et al. (2017)
H. cruenta He 766 JQ279595 JQ279681 He et al. (2017)
H. cyclolamellata Cui 7393 JQ279513 JQ279629 He et al. (2017)
H. damicornis URM 84261 KC348466 — Du et al. (2021)
H. damicornis URM 84263 KC348467 — Du et al. (2021)
H. denticulata He 1271 KF438171 KF438174 He et al. (2017)
H. dracaenicola Dai 22090 MW559797 MW559802 Du et al. (2021)
H. dracaenicola Dai 22096 MW559798 MW559803 Du et al. (2021)
H. duportii AFTOL ID666 DQ404386 AY635770 He et al. (2017)
H. epichlora He 525 JQ279549 JQ279659 He and Dai (2012)
H. floridea He 536 JQ279597 JQ279683 He and Dai (2012)
H. fuliginosa He 1188 KU975465 KU975506 Du et al. (2021)
H. fulva He 640 JQ279565 JQ279648 He and Dai (2012)
H. globispora He 911 — KU975508 Du et al. (2021)
H. huangshanensis He 432 JQ279533 JQ279671 He and Dai (2012)
H. innexa He 555 JQ279584 JQ279674 He and Dai (2012)
H. japonica He 245 JQ279590 JQ279680 He and Dai (2012)
H. legeri He 960 KU975469 KU975511 He et al. (2017)
H. longispora He 217 JQ279537 KU975514 He and Dai (2012)
H. luteobadia He 8 JQ279569 KU975515 He and Dai (2012)
H. macrochloae ARAN-fungi 7079 MF990738 MF990743 Du et al. (2021)
H. megaspora He 302 JQ279553 JQ279660 He and Dai (2012)
H. minor He 933 JQ279555 JQ279654 He and Dai (2012)
H. minuscula He 253 JQ279546 KU975516 He and Dai (2012)
H. murina He 569 JQ716406 JQ716412 He et al. (2017)
H. muroiana He 405 JQ279542 KU975517 Du et al. (2021)
H. nanospora He 475 JQ279531 JQ279672 He and Dai (2012)
H. ochromarginata He 47 JQ279579 JQ279666 He and Dai (2012)
H. orientalis He 4601 KY425677 KY425685 He et al. (2017)
H. parmastoi He 277 JQ780064 — He et al. (2017)
H. paucisetigera Cui 7845 JQ279560 JQ279644 He and Dai (2012)
H. quercicola He 373 KU975474 KU975521 He et al. (2017)
H. rhabarbarina He 280 JQ279574 KY425688 He and Dai 2012
H. rheicolor Cui 8317 JQ279529 — Du et al. (2021)
H. rhododendricola He 389 JQ279577 JQ279653 He and Dai (2012)
H. rubiginosa He 1049 JQ716407 JQ279667 Yang et al. (2016)
H. rubiginosa LWZ 20201017-32 ON063655 — Wang et al. (2023a)
H. rufomarginata He 1489 KU975477 KU975524 He et al. (2017)
H. separabilis He 460 JQ279572 JQ279655 He and Dai (2012)
H. setipora Cui 6301 JQ279515 JQ279639 Du et al. (2021)
H. sharmae 66088 MK588753 MK588836 Du et al. (2021)
H. sharmae CAL 1535 KY929017 KY929018 Du et al. (2021)
H. sinensis CLZhao 26040 OR659001 PP425893 Li et al. (2024)
H. spathulata He 685 JQ279591 KU975529 He et al. (2017)
H. sphaericola He 303 JQ279599 JQ279684 Li et al. (2024)
H. sphaericola LWZ 20190808-2b ON063656 ON063855 Li et al. (2024)
H. sphaerospora He 715 JQ279594 KU975531 He et al. (2017)
H. subferruginea Cui 8122 JQ279521 — Du et al. (2021)
H. subferruginea He 1598 KU975481 — Du et al. (2021)
H. tabacina Dai 11635 JQ279563 JQ279647 He and Dai (2012)
H. tasmanica He 449 JQ279582 JQ279663 He et al. (2017)
H. tenuis He 779 JQ279538 JQ279641 Du et al. (2021)
H. tongbiguanensis He 1552 KF908248 KU975532 He et al. (2017)
H. tropica He 574 JQ279587 JQ279675 He et al. (2017)
H. ulmicola He 864 JQ780065 KU975534 He et al. (2017)
H. unicolor He 468a JQ279551 JQ279662 He et al. (2017)
H. verruculosa Dai 17052 MF370594 MF370601 He et al. (2017)
H. villosa He 537 JQ279528 JQ279634 He et al. (2017)
H. xerantica Cui 9209 JQ279519 JQ279635 Du et al. (2021)
H. xerantica LWZ 20190814-13b ON063657 ON063856 Wang et al. (2023a)
H. bannaensis CLZhao 35721 * PQ847494 PQ847499 Present study
H. bannaensis CLZhao 35884 PQ847495 PQ847500 Present study
H. yunnanensis He 1447 KU975486 KU975538 He et al. (2017)
Hyphodontia arguta KHL 11938 (GB) EU118632 EU118633 Larsson (2007a)
H. arguta LWZ 20180905-6 ON063672 ON063871 Wang et al. (2023a)
H. borbonica FR-0219441 KR349240 NG_068856 Riebesehl et al. (2015)
H. pachyspora LWZ20170908-5 MT319426 MT319160 Wang et al. (2023a)
H. pachyspora LWZ20180905-6 MT319425 MT319159 Wang et al. (2021)
H. pallidula He6087 OM100749 OM083983 Unpublished
H. pallidula He6092 OM100750 OM083984 Unpublished
H. wongiae LWZ20180414-16 MT319414 MT319146 Wang et al. (2023a)
H. wongiae LWZ20180417-16 MT319415 MT319147 Wang et al. (2023a)
Lyomyces albopulverulentus CLZhao 21478 OP730712 OP730724 Guan et al. (2023)
L. allantosporus FR-0249548 KY800397 KY795963 Yurchenko et al. (2017)
L. allantosporus KAS-GEL4933 KY800401 KY795965 Yurchenko et al. (2017)
L. asiaticus CLZhao 35703 PQ847492 PQ847501 Present study
L. asiaticus CLZhao 35719 * PQ847493 PQ847502 Present study
L. bambusinus CLZhao 4808 MN945970 — Chen and Zhao (2020)
L. bambusinus CLZhao 4831 MN945968 MW264919 Chen and Zhao (2020)
L. cremeus CLZhao 4138 MN945974 MW264922 Chen and Zhao (2020)
L. cremeus CLZhao 8295 MN945972 — Chen and Zhao (2020)
L. crustosus TASM:YG G39 MF382993 — Gafforov et al. (2017)
L. crustosus UC2022841 KP814310 — Guan et al. (2023)
L. densiusculus Ryvarden 44818 OK273853 OK273853 Viner et al. (2022)
L. elaeidicola LWZ20180411-19 MT319457 MT319190 Wang et al. (2021)
L. elaeidicola LWZ20180411-20 MT319458 NG_153910 Wang et al. (2021)
L. erastii 23cSAMHYP JX857800 — Unpublished
L. erastii TASM:YG 022 MF382992 — Gafforov et al. (2017)
L. fimbriatus Wu910620-7 MK575209 — Yurchenko et al. (2020a)
L. fimbriatus Wu911204-4 MK575210 MK598740 Yurchenko et al. (2020a)
L. fissuratus CLZhao 4291 MW713738 MW713730 Luo et al. (2021b)
L. fissuratus CLZhao 4352 MW713742 MW713732 Luo et al. (2021b)
L. fumosus CLZhao 8188 MW713744 MW713736 Luo et al. (2021b)
L. gatesiae LWZ20180515-3 MT319447 MT319181 Wang et al. (2021)
L. gatesiae LWZ20180515-32 MT319448 MT319182 Wang et al. (2021)
L. griseliniae KHL 12971 (GB) DQ873651 DQ873651 Larsson et al. (2006)
L. hengduanensis CLZhao 20627 OR793233 PP657611 Yuan et al. (2024)
L. hengduanensis CLZhao 25551 OR658999 PP657610 Yuan et al. (2024)
L. incanus CLZhao 22813 OR094480 OR449935 Dong et al. (2024)
L. incanus CLZhao 22900 * OR094481 OR449936 Dong et al. (2024)
L. juniper FR-0261086 KY081799 — Riebesehl and Langer (2017)
L. macrosporus CLZhao 4516 MN945977 MW264920 Chen and Zhao (2020)
L. mascarensis KAS-GEL4833 KY800399 KY795964 Yurchenko et al. (2020a)
L. mascarensis KAS-GEL4908 KY800400 — Yurchenko et al. (2020a)
L. microfasciculatus CLZhao 5109 MN954311 MW264921 Chen and Zhao (2020)
L. niveomarginatus CLZhao 16360 PP537949 PP657607 Yuan et al. (2024)
L. niveus CLZhao 6431 MZ262541 MZ262526 Luo et al. (2021b)
L. niveus CLZhao 6442 MZ262542 MZ262527 Luo et al. (2021b)
L. ochraceoalbus CLZhao 4385 MZ262535 MZ262521 Luo et al. (2021b)
L. ochraceoalbus CLZhao 4725 MZ262536 MZ262522 Luo et al. (2021b)
L. orientalis GEL3376 DQ340325 DQ340351 Yurchenko et al. (2017)
L. pruni GEL2327 DQ340312 DQ340349 Larsson et al. (2006)
L. pruni Ryberg 021018 (GB) DQ873624 — Larsson et al. (2006)
L. sambuci 83SAMHYP JX857721 — Yurchenko et al. (2017)
L. sambuci KAS-JR7 KY800402 KY795966 Yurchenko et al. (2017)
L. vietnamensis He 3260 MW507086 MW507028 Yurchenko et al. (2017)
L. wuliangshanensis He 3498 MW507087 MW507029 Chen and Zhao (2020)
L. wuliangshanensis He 4765 MW507090 MW507032 Chen and Zhao (2020)
L. wumengshanensis CLZhao 29374 OR803021 PP657613 Yuan et al. (2024)
L. wumengshanensis CLZhao 32800 OR899211 PP657614 Yuan et al. (2024)
L. yunnanensis CLZhao 2463 OP730711 OP730723 Guan et al. (2023)
L. yunnanensis CLZhao 9375 OP730710 — Guan et al. (2023)
L. zhaotongensis CLZhao 32878 PP537950 PP657609 Yuan et al. (2024)
Nigrofomes melanoporus JV 1704/39 MF629835 MF629831 Wang et al. (2023a)
N. sinomelanoporus Cui 5277 MF629836 MF629832 Wang et al. (2023a)
Peniophorella albohymenia CLZhao 33187 * PQ811412 PQ847496 Present study
P. albohymenia CLZhao 33257 PQ811413 — Present study
P. aspersa TNM F24809 MN062097 MN062142 Yurchenko et al. (2020b)
P. aspersa TNM F32708 MN062099 MN062144 Yurchenko et al. (2020b)
P. cremea CLZhao 1606 MT955162 — Xu et al. (2020)
P. cremea CLZhao 1719 MT955163 — Xu et al. (2020)
P. crystallifera LWZ 20210626-4a ON063685 ON063885 Wang et al. (2023a)
P. crystallifera TNM F30331 MN062100 MN062147 Yurchenko et al. (2020b)
P. daweishanensis CLZhao 18600 * OR094501 OR449932 Dong et al. (2024)
P. echinocystis KHL 6284 DQ677494 DQ681200 Larsson (2007b)
P. fissurata CLZhao 5848 MN864262 OM985777 Guan et al. (2020)
P. fissurata CLZhao 9421 MN864260 OM985776 Guan et al. (2020)
P. guttulifera CBS 107303 LT603016 LT603001 Kolařík and Vohník (2018)
P. guttulifera NH 12012 (GB) DQ647501 — Hallenberg et al. (2007)
P. odontiiformis TMI 21347 DQ647496 — Hallenberg et al. (2007)
P. odontiiformis TMI 6824 DQ647500 — Hallenberg et al. (2007)
P. olivacea CLZhao 25896 * OR094502 OR449933 Dong et al. (2024)
P. pallida UC 2022844 KP814208 — Rosenthal et al (2017)
P. pallida UC 2022887 KP814201 — Rosenthal et al (2017)
P. pertenuis NH 12429 (GB) DQ647486 — Hallenberg et al. (2007)
P. pertenuis NH 15115 (GB) DQ647487 — Hallenberg et al. (2007)
P. praetermissa NH 10986 (GB) DQ647462 — Hallenberg et al. (2007)
P. praetermissa NH 11192 (GB) DQ647461 — Hallenberg et al. (2007)
P. pubera CBS:464.86 MH861988 MH873680 Vu et al. (2019)
P. pubera LWZ 20210624-16b ON063687 ON063887 Wang et al. (2023a)
P. punctata CLZhao 33720 PQ811414 PQ847497 Present study
P. punctata CLZhao 33732 * PQ811415 PQ847498 Present study
P. reticulate CLZhao 17066 OM985746 OM985783 Dong et al. (2024)
P. reticulate TNM F22559 MN062103 MN062151 Yurchenko et al. (2020b)
P. rude LWZ 20171026-7 ON063688 ON063888 Wang et al. (2023a)
P. subpraetermissa LWZ 20190816-3b ON063689 ON063889 Wang et al. (2023a)
P. subpraetermissa Wu 950627 DQ647493 — Hallenberg et al. (2007)
P. yunnanensis CLZhao 4810 MN864263 OM985788 Guan et al. (2020)
P. yunnanensis CLZhao 6137 MN864266 — Guan et al. (2020)
Phylloporia oreophila LWZ 20190811-27a ON063665 ON063865 Wang et al. (2023a)
P. radiate LWZ 20141122-6 ON063666 ON063866 Wang et al. (2023a)
Porodaedalea himalayensis LWZ 20180903-21 ON063667 ON063867 Wang et al. (2023a)
P. laricis LWZ 20190724-9 ON063668 ON063868 Wang et al. (2023a)
Sanghuangporus quercicola LWZ 20170821-18 ON063669 ON063869 Wang et al. (2023a)
S. weigelae LWZ 20210623-2a ON063671 ON063870 Wang et al. (2023a)
Skvortzovia dabieshanensis LWZ 20201017-55 MW414514 MW414460 Wang et al. (2023a)
S. dabieshanensis LWZ 20210918-15b ON063694 ON063894 Wang et al. (2023a)
S. furfurella KHL 10180 DQ873649 DQ873649 Yu et al. (2021)
S. incana CLZhao 16338 * OR096179 OR449950 Dong et al. (2024)
S. yunnanensis CLZhao 16084 MW472754 MW473473 Wang et al. (2023a)
S. yunnanensis CLZhao 16181 MW472755 MW473474 Dong et al. (2021)
Thelephora ganbajun ZRL20151295 LT716082 KY418908 Liu et al. (2022)
Tubulicrinis glebulosus LWZ 20180903-13 ON063705 — Wang et al. (2023a)
T. pini CLZhao 3679 — OR449951 Dong et al. (2024)
T. pini CLZhao 6881 * OR096210 OR449952 Dong et al. (2024)
T. yunnanensis CLZhao 3418 MT153879 MT153886 He et al. (2020)
T. yunnanensis CLZhao 9717 MT153880 MT153887 He et al. (2020)
Xylodon olivaceobubalinus CLZhao 25164 OR167771 OR449947 Dong et al. (2024)
X. olivaceobubalinus CLZhao 25174 * OR167772 OR449948 Dong et al. (2024)

The sequences were aligned in MAFFT version 7 (Katoh et al. 2019) using the G-INS-i strategy. The alignment was adjusted manually using AliView version 1.27 (Larsson 2014). The combined dataset was deposited in TreeBASE (https://treebase.org/treebase-web/home.html; submission ID 32063). Sequences of Boletopsis leucomelaena (Pers.) Fayod and Thelephora ganbajun M. Zang, retrieved from GenBank were used as the outgroup in the ITS+nLSU analysis (Fig. 1; Dong et al. 2024). Sequences of Fomitiporia bannaensis Y.C. Dai and F. punctata (P. Karst.) Murrill, retrieved from GenBank were used as the outgroup in the ITS+nLSU analysis (Fig. 2; Nie et al. 2017). Sequences of Fasciodontia brasiliensis Yurchenko & Riebesehl and F. bugellensis (Ces.) Yurchenko, Riebesehl & Langer retrieved from GenBank were used as the outgroup in the ITS+nLSU analysis (Fig. 3; Yuan et al. 2024). Sequences of Basidioradulum mayi Xue W. Wang & L.W. Zhou and B. radula (Fr.) Nobles retrieved from GenBank were used as the outgroup in the ITS+nLSU analysis (Fig. 4; Wang et al. 2023a; Dong et al. 2024).

Figure 1. 

Maximum Parsimony strict consensus tree illustrating the phylogeny of the order Hymenochaetales based on ITS+nLSU sequences. Branches are labeled with Maximum Likelihood bootstrap values ≥ 70%, parsimony bootstrap values ≥ 50% and Bayesian posterior probabilities ≥ 0.95, respectively.

Figure 2. 

Maximum Parsimony strict consensus tree illustrating the phylogeny of the two new species and related species in Hymenochaete, based on ITS+nLSU sequences. Branches are labeled with Maximum Likelihood bootstrap values ≥ 70%, parsimony bootstrap values ≥ 50% and Bayesian posterior probabilities ≥ 0.95, respectively.

Figure 3. 

Maximum Parsimony strict consensus tree illustrating the phylogeny of the two new species and related species in Lyomyces, based on ITS+nLSU sequences. Branches are labeled with Maximum Likelihood bootstrap values ≥ 70%, parsimony bootstrap values ≥ 50% and Bayesian posterior probabilities ≥ 0.95, respectively.

Maximum Parsimony (MP), Maximum Likelihood (ML), and Bayesian Inference (BI) analyses were applied to the combined three datasets following a previous study (Dong et al. 2024). All characters were equally weighted and gaps were treated as missing data. Trees were inferred using the heuristic search option with TBR branch swapping and 1,000 random sequence additions. Max-trees were set to 5,000, branches of zero length were collapsed and all parsimonious trees were saved. Clade robustness was assessed using bootstrap (BT) analysis with 1,000 pseudo replicates (Felsenstein 1985). Descriptive tree statistics of tree length (TL), composite consistency index (CI), composite retention index (RI), composite rescaled consistency index (RC), and composite homoplasy index (HI) were calculated for each maximum parsimonious tree generated. The combined dataset was also analysed using Maximum Likelihood (ML) in RAxML-HPC2 through the CIPRES Science Gateway (Miller et al. 2012). Branch support (BS) for the ML analysis was determined by 1000 bootstrap pseudo replicates.

MrModeltest 2.3 (Nylander 2004) was used to determine the best-fit evolution model for each dataset for the purposes of Bayesian inference (BI) which was performed using MrBayes 3.2.7a with a GTR+I+G model of DNA substitution and a gamma distribution rate variation across sites (Ronquist et al. 2012). A total of four Markov chains were run for two runs from random starting trees for 1 million generations for ITS+nLSU (Fig. 1); 0.8 million generations for ITS+nLSU (Figs 2, 3); and 4.4 million generations for ITS+nLSU (Fig. 4) with trees and parameters sampled every 1,000 generations. The first quarter of all of the generations were discarded as burn-ins. A majority rule consensus tree was computed from the remaining trees. Branches were considered as significantly supported if they received a maximum likelihood bootstrap support value (BS) of ≥ 70%, a Maximum Parsimony bootstrap support value (BT) of ≥ 70% or a Bayesian posterior probability (BPP) of ≥ 0.95.

Figure 4. 

Maximum Parsimony strict consensus tree illustrating the phylogeny of the two new species and related species in Peniophorella, based on ITS+nLSU sequences. Branches are labeled with Maximum Likelihood bootstrap values ≥ 70%, parsimony bootstrap values ≥ 50% and Bayesian posterior probabilities ≥ 0.95, respectively.

Results

Molecular phylogeny

The ITS+nLSU dataset (Fig. 1) comprised sequences from 102 fungal specimens representing 63 taxa. The dataset had an aligned length of 2,441 characters, of which 1,233 characters were constant, 204 were variable and parsimony-uninformative and 1,004 (50%) were parsimony-informative. Maximum Parsimony analysis yielded 3 equally parsimonious trees (TL = 7,192, CI = 0.3420, HI = 0.6803, RI = 0.7080 and RC = 0.2263). The best model of nucleotide evolution for the ITS+nLSU dataset estimated and applied in the Bayesian analysis was found to be GTR+I+G. Bayesian analysis and ML analysis resulted in a similar topology as in the MP analysis. The Bayesian analysis had an average standard deviation of split frequencies = 0.008504 (BI). The phylogram based on the ITS+nLSU rDNA gene regions (Fig. 1) included six families within the Hymenochaetales, which were Hymenochaetaceae, Hyphodontiaceae, Peniophorellaceae, Schizoporaceae, Skvortzoviaceae and Tubulicrinaceae, in which the new species Hymenochaete bannaensis was nested in Hymenochaetaceae; two taxa Peniophorella albohymenia and P. punctata were nested in Peniophorellaceae; the species Lyomyces asiaticus was grouped in Schizoporaceae.

The ITS+nLSU dataset (Fig. 2) comprised sequences from 80 fungal specimens representing 71 taxa. The dataset had an aligned length of 2,285 characters, of which 1,479 characters were constant, 199 were variable and parsimony-uninformative and 607 (50%) were parsimony-informative. Maximum Parsimony analysis yielded 3 equally parsimonious trees (TL = 3,749, CI = 0.3420, HI = 0.6580, RI = 0.5462 and RC = 0.1868). The best model of nucleotide evolution for the ITS+nLSU dataset estimated and applied in the Bayesian analysis was found to be GTR+I+G. Bayesian analysis and ML analysis resulted in a similar topology as in the MP analysis. The Bayesian analysis had an average standard deviation of split frequencies = 0.001385 (BI) and the effective sample size (ESS) across the two runs is double the average ESS (avg. ESS) = 365. The phylogram based on the ITS+nLSU rDNA gene regions (Fig. 1) highlighted that the new species Hymenochaete bannaensis was sister to H. muroiana I. Hino & Katum. with strong supports values (100/100/1.00), and then was closely nested with H. colliculosa (Sacc.) Parmasto, H. globispora G.A. Escobar and H. megaspora S.H. He & Hai J. Li.

The ITS+nLSU dataset (Fig. 3) comprised sequences from 51 fungal specimens representing 32 taxa. The dataset had an aligned length of 2,066 characters, of which 1,366 characters were constant, 177 were variable and parsimony-uninformative and 523 (50%) were parsimony-informative. Maximum Parsimony analysis yielded 3 equally parsimonious trees (TL = 2,358, CI = 0.4440, HI = 0.5560, RI = 0.6763 and RC = 0.3003). The best model of nucleotide evolution for the ITS+nLSU dataset estimated and applied in the Bayesian analysis was found to be GTR+I+G. Bayesian analysis and ML analysis resulted in a similar topology as in the MP analysis. The Bayesian analysis had an average standard deviation of split frequencies = 0.006833 (BI) and the effective sample size (ESS) across the two runs is double the average ESS (avg. ESS) = 306. The phylogram based on the ITS+nLSU rDNA gene regions (Fig. 3) indicated that the new species Lyomyces asiaticus was clustered with L. crustosus (Pers.) P. Karst., L. ochraceoalbus C.L. Zhao, L. hengduanensis Q. Yuan & C.L. Zhao and L. zhaotongensis Q. Yuan & C.L. Zhao.

The ITS+nLSU dataset (Fig. 4) comprised sequences from 36 fungal specimens representing 212 taxa. The dataset had an aligned length of 2,016 characters, of which 1,497 characters were constant, 72 were variable and parsimony-uninformative and 447 (50%) were parsimony-informative. Maximum Parsimony analysis yielded 3 equally parsimonious trees (TL = 1,235, CI = 0.6024, HI = 0.3976, RI = 0.7813 and RC = 0.4707). The best model of nucleotide evolution for the ITS+nLSU dataset estimated and applied in the Bayesian analysis was found to be TIM3+I+G. Bayesian analysis and ML analysis resulted in a similar topology as in the MP analysis. The Bayesian analysis had an average standard deviation of split frequencies = 0.009797 (BI) and the effective sample size (ESS) across the two runs is double the average ESS (avg. ESS) = 563. The phylogram based on the ITS+nLSU rDNA gene regions (Fig. 3) revealed that the new species Peniophorella albohymenia was grouped with two taxa as P. olivacea J.H. Dong & C.L. Zhao and P. subpraetermissa (Sheng H. Wu) K.H. Larss. Furthermore, the species P. punctata Y.L. Deng & C.L. Zhao was the sister to P. fissurata C.L. Zhao with strong supports values (100/100/1.00).

Taxonomy

Hymenochaete bannaensis Y.L. Deng & C.L. Zhao, sp. nov.

MycoBank No: 857317
Figs 5, 6, 7

Holotype.

China • Yunnan Province, Xishuangbanna, Wild elephant Valley, GPS coordinates: 22°17'N, 100°85'E, altitude 900 m asl., on the dead bamboo, leg. C.L. Zhao, 25 January 2024, CLZhao 35721 (SWFC).

Etymology.

Bannaensis (Lat.): referring to the locality (banna) of the type specimen.

Diagnosis.

Hymenochaete bannaensis is characterized by the flocculent basidiomata with cinnamon to yellowish brown to rust-brown hymenial surface, generative hyphae with simple septa and broadly ellipsoid to globose basidiospores.

Description.

Basidiomata annual, resupinate, closely adnate, flocculent, without odor or taste when fresh, up to 7 cm long, 3.5 cm wide, and 200 μm thick. Hymenial surface smooth, cinnamon to yellowish brown when fresh, yellowish brown to rust-brown upon drying. Sterile margin yellowish brown, thinning out, up to 2 mm.

Figure 5. 

Basidiomata of Hymenochaete bannaensis (holotype, CLZhao 35721). Scale bars: 1 cm (A); 1 mm (B).

Hyphal system monomitic; generative hyphae with simple-septa, brown, thick-walled, smooth, moderately branched, 2.4–2.9 µm in diameter, IKI–, CB–, tissues darkening in KOH; subhymenial hyphae densely covered by crystals.

Figure 6. 

Microscopic structures of Hymenochaete bannaensis (holotype, CLZhao 35721) A basidiospores B basidia and basidioles C setae D a section of hymenium. Scale bars: 5 µm (A); 10 µm (B–D).

Cystidia and cystidioles absent. Basidia subcylindrical, colorless, thin-walled, with four sterigmata and a basal simple septum, 18.5–23 × 3.5–4 µm; basidioles numerous, in shape similar to basidia but smaller. Setae abundant, distinctly thick-walled, subulate, with an acute tip, yellowish to reddish brown, 67–94 × 7–11 µm, projecting out of the hymenium up to 33–44.5 µm.

Figure 7. 

Sections of hymenium of Hymenochaete bannaensis (holotype, CLZhao 35721). Scale bars: 10 µm (A–C).

Basidiospores broadly ellipsoid to globose, colorless, thin-walled, smooth, always filled with oil drop, CB–, (3.6–)3.7–4(–4.1) × (3.4–)3.5–3.9(–4) µm, L = 3.87 µm, W = 3.67 µm, Q = 1.05–1.06 (n = 60/2).

Another specimen (paratype) examined.

China • Yunnan Province, Xishuangbanna, Wild elephant Valley, GPS coordinates: 22°17'N, 100°85'E, altitude 900 m asl., on the fallen angiosperm branch, leg. C.L. Zhao, 25 January 2024, CLZhao 35884 (SWFC).

Notes.

Phylogenetically, Hymenochaete bannaensis is nested as sister to H. muroiana with strong supports (100/100/1.00) based on ITS+nLSU sequences (Figs 1, 2). Morphologically, H. muroiana is different from H. bannaensis by the brun suie basidiomata, smaller ellipsoid to subcylindrical basidia (8–10 × 3 µm) and narrower ovoid to ellipsoid basidiospores (3.5–4 × 2–2.6 µm; Léger 1998). Hymenochaete acerosa S.H. He & Hai J. Li, H. cystidiata Parmasto and H. sinensis Y.C. Li & C.L. Zhao, are similar to H. bannaensis by thick-walled and acute setae. However, H. acerosa is distinguished from H. bannaensis by its effused and detachable basidiomata, longer setae (85–170 × 5–8 µm), wider clavate basidia (13–21 × 5–8 µm) and larger basidiospores (7–8.5 × 4.8–6 µm; He and Liu 2011). Furthermore, H. cystidiata differs from H. bannaensis by its longer setae (60–100 × 7–10 µm), and subcylindrical basidiospores (4.8–5.8 × 2.5–3.2 µm; Parmasto 2012). Moreover, H. sinensis differs from H. bannaensis by having ceraceous basidiomata, coconut brown to brownish black hymenial surface, shorter barrel-shaped basidia (10–12 × 3.1–3.8 µm) and longer ellipsoid to broadly ellipsoid basidiospores (4–5 × 2.5–3.5 µm; Li et al. 2024).

Lyomyces asiaticus Y.L. Deng & C.L. Zhao, sp. nov.

MycoBank No: 857318
Figs 8, 9, 10

Holotype.

China • Yunnan Province, Xishuangbanna, Wild elephant Valley, GPS coordinates: 22°17'N, 100°85'E, altitude 900 m asl., on the dead bamboo, leg. C.L. Zhao, 25 January 2024, CLZhao 35719 (SWFC).

Etymology.

Asiaticus (Lat.): referring to the continent (Asia) where the species was found.

Diagnosis.

Lyomyces asiaticus is characterized by the membranaceous basidiomata with cream to pale-yellow hymenial surface with tuberculate, a monomitic hyphal system with clamped generative hyphae, and ellipsoid basidiospores 4.6–6.4 × 3–3.9 µm.

Description.

Basidiomata annual, resupinate, closely adnate, membranaceous, without odor or taste when fresh, up to 5 cm long, 1.5 cm wide, and 100 μm thick. Hymenial surface tuberculate, thin, white to cream when fresh, cream upon drying. Sterile margin white to cream, thinning out, up to 2 mm.

Hyphal system monomitic; generative hyphae with clamp connections, colorless, thin-walled, smooth, moderately branched, with some crystals, 2–2.5 µm in diameter, IKI–, CB–, tissues unchanged in KOH; subhymenial hyphae densely covered by crystals.

Figure 8. 

Basidiomata of Lyomyces asiaticus (holotype, CLZhao 35719). Scale bars: 1 cm (A); 1 mm (B)

Hymenium three kinds of cystidia: (1) tapering, colorless, thin-walled, smooth, filled with refractive oil-like matter, 19.5–27 × 1.8–2.3 µm; (2) bottled, colorless, thin-walled, smooth, filled with refractive oil-like matter, 19.3–24.7 × 4.6–5.4 µm; (3) halocystidia, colorless, thin-walled, smooth, filled with refractive oil-like matter, 17–19 × 5.5–7 µm. Basidia cylindrical, with a basal clamp connection and four sterigmata, filled with refractive oil-like matter, 17–21 × 3–7 µm; basidioles numerous, in shape similar to basidia but smaller.

Figure 9. 

Microscopic structures of Lyomyces asiaticus (holotype, CLZhao 35719) A basidiospores B basidia and basidioles C tapering cystidia D bottled cystidia E halocystidia F a section of hymenium. Scale bars: 5μm (A); 10 μm (B–F).

Basidiospores ellipsoid, colorless, thin-walled, smooth, always filled with oil drop, CB–, (4.2–)4.6–6.4(–6.5) × (2.4–)3–3.9(–4) µm, L = 5.63 µm, W = 3.36 µm, Q = 1.62–1.73 (n = 60/2).

Another specimen (paratype) examined.

China • Yunnan Province, Xishuangbanna, Wild elephant Valley, GPS coordinates: 22°17'N, 100°85'E, altitude 900 m asl., on the fallen angiosperm branch, leg. C.L. Zhao, 25 January 2024, CLZhao 35703 (SWFC).

Figure 10. 

Sections of hymenium of Lyomyces asiaticus (holotype, CLZhao 35719). Scale bars: 10 µm (A–C).

Notes.

The phylogenetic tree of ITS+nLSU (Fig. 3) showed that the new species Lyomyces asiaticus was clustered with L. crustosus and L. ochraceoalbus. However, morphologically, L. crustosus can be delimited from L. asiaticus by having effused basidiocarp, clavate basidia (26 × 4.5 µm) and longer ellipsoidal to subcylindrical basidiospores (5–7.5 × 2.5–3 µm; Karsten 1881). L. ochraceoalbus differs from L. asiaticus in its smooth hymenial surface, lacking a cystidium and shorter clavate basidia (11–16.5 × 3.5–5 µm; Luo et al. 2021b). Morphologically, Lyomyces asiaticus reminds L. albopulverulentus C.L. Zhao, L. cremeus, L. denudatus Viner, L. macrosporus C.L. Zhao and L. wuliangshanensis C.L. Zhao by having the ellipsoid basidiospores. However, L. albopulverulentus can be separated from L. asiaticus by its pruinose hymenial surface and larger basidia (24.5–28.5 × 7–9 µm) and larger basidiospores (8–10.5 × 5.5–7 µm; Guan et al. 2023). L. cremeus is distinct from L. asiaticus by its smooth hymenial surface, present two types cystidia as capitate (20–40 × 3–5 µm) and tapering (18–35 × 3–4.5 µm) and thin-walled to slightly thick-walled basidiospores (Chen and Zhao 2020). In addition, L. denudatus is separated from L. asiaticus by the smooth hymenial surface, longer capitate cystidia (34.9–62 × 4–5.5 µm) and suburniform basidia (15–21.1 × 3.8–5.5 µm; Viner and Miettinen 2022). The species L. macrosporus differs from L. asiaticus due to its reticulate hymenial surface and longer basidia (22.2–38 × 4.5–7 µm) and larger basidiospores (6.7–8.9 × 4.4–5.4 µm; Chen and Zhao 2020). Moreover, L. wuliangshanensis can be delimited from L. asiaticus by its smooth to more or less tuberculate hymenial surface and two kinds of cystidia as capitate (22–37 × 3–6 µm) and tapering (21–35 × 4–6.5 µm; Chen and Zhao 2020).

Peniophorella albohymenia Y.L. Deng & C.L. Zhao, sp. nov.

MycoBank No: 857319
Figs 11, 12, 13

Holotype.

China • Yunnan Province, Zhaotong, Xiaocaoba Town, Wumengshan National Nature Reserve, GPS coordinates: 27°33'N, 103°27'E, altitude 2300 m asl., on fallen angiosperm branch, leg. C.L. Zhao, 19 September 2023, CLZhao 33187 (SWFC).

Etymology.

Albohymenia (Lat.): referring to “albus”, the distinctive white hymenium of the type specimen when fresh.

Diagnosis.

Peniophorella albohymenia is characterized by the membranaceous basidiomata with white hymenial surface, four types cystidia as stephanocyst, fusiform, cylindrical and capitate, and ellipsoid basidiospores measuring 9–10.9 × 4.5–5 µm.

Description.

Basidiomata annual, resupinate, adnate, membranaceous, without odor or taste when fresh, up to 4.5 cm long, 1 cm wide, and 0.1 mm thick. Hymenial surface smooth, white when fresh and dry. Sterile margin distinctly, thin, white, up to 1 mm long.

Figure 11. 

Basidiomata of Peniophorella albohymenia (holotype, CLZhao 33187) A basidiomata on the substrate B character hymenophore Scale bars: 1 cm (A); 1 mm (B).

Hyphal system monomitic; generative hyphae with clamp connections, thin-walled, colorless, occasionally branched, 2.5–4 µm in diameter, IKI–, CB–, tissues unchanged in KOH.

Figure 12. 

Microscopic structures of Peniophorella albohymenia (holotype, CLZhao 33187) A basidiospores B basidia and basidioles C stephanocyst D fusiform cystidia E cylindrical cystidia F capitate cystidia G a section of the hymenium. Scale bars: 10 µm (A–G).

Cystidia of four types: (1) stephanocyst, thin-walled, 9–18 × 7–8 µm; (2) fusiform cystidia, smooth, thin-walled, 29–31 × 7–9 µm; (3) cylindrical cystidia, thin-walled, slightly constricted at the neck, the apical part encrusted with asteroid, 27–64 × 5–11 µm; (4) capitate cystidia, thin-walled, the apical part encrusted with asteroid, 22–47 × 5–12 µm. Basidia clavate, with four sterigmata and a basal clamp connection, colorless, thin-walled, 20–28 × 7.5–9 µm; basidioles in shape similar to basidia, but slightly smaller.

Figure 13. 

Sections of hymenium of Peniophorella albohymenia (holotype, CLZhao 33187). Scale bars: 10 µm (A–F).

Basidiospores ellipsoid, colorless, thin-walled, IKI–, CB–, (8.5–)9–10.9(–11.2) × (4.4–)4.5–5(–5.5) µm, L = 9.87 µm, W = 4.93 µm, Q = 1.99–2.02 (n = 60/2).

Another specimen (paratype) examined.

China • Yunnan Province, Zhaotong, Xiaocaoba Town, Wumengshan National Nature Reserve, GPS coordinates: 27°33'N, 103°27'E, altitude 2300 m asl., on fallen angiosperm branch, leg. C.L. Zhao, 19 September 2023, CLZhao 33257 (SWFC).

Notes.

Our results indicate that the new species P. albohymenia was placed within Peniophorella inferring from the dataset of ITS+nLSU (Figs 1, 4). Morphologically, the species P. albohymenia is similar to P. daweishanensis J.H. Dong & C.L. Zhao and P. yunnanensis C.L. Zhao by having capitate cystidia, but P. daweishanensis is distinct from P. albohymenia by its buff to slightly yellowish, reticulate hymenial surface and longer basidia (31.5–38 × 7–9 µm; Dong et al. 2024). P. yunnanensis differs from P. albohymenia due to its grandinioid hymenial surface (Guan et al. 2020).

Peniophorella punctata Y.L. Deng & C.L. Zhao, sp. nov.

MycoBank No: 857320
Figs 14, 15, 16

Holotype.

China • Yunnan Province, Zhaotong, Xiaocaoba Town, Wumengshan National Nature Reserve, GPS coordinates: 27°33'N, 103°27'E, altitude 2300 m asl., on fallen angiosperm branch, leg. C.L. Zhao, 21 September 2023, CLZhao 33732 (SWFC).

Etymology.

Punctata (Lat.): referring to the punctate basidioma of the type specimen.

Diagnosis.

Peniophorella punctata is characterized by the membranaceous, punctate basidiomata with white to pale yellow hymenial surface, fusiform cystidia, and allantoid basidiospores measuring 9.5–12.5 × 4–4.8 µm.

Figure 14. 

Basidiomata of Peniophorella punctata (holotype, CLZhao 33732) A basidiomata on the substrate B character hymenophore. Scale bars: 1 cm (A); 1 mm (B).

Description.

Basidiomata annual, resupinate, adnate, membranaceous, punctate, without odor or taste when fresh, up to 4 cm long, 3 cm wide, and 0.1 mm thick. Hymenial surface smooth, white to pale yellow when fresh, turning to aurantiacus to avellaneus upon drying. Sterile margin distinctly, thin, white, up to 2 mm long.

Figure 15. 

Microscopic structures of Peniophorella punctata (holotype, CLZhao 33732) A basidiospores B basidia and basidioles C fusiform cystidia D a section of the hymenium. Scale bars: 10 µm (A–D).

Hyphal system monomitic, generative hyphae with clamp connections, thin-walled, colorless, occasionally branched, 2.9–4.6 µm in diameter, IKI–, CB–, tissues unchanged in KOH.

Figure 16. 

Sections of hymenium of Peniophorella punctata (holotype, CLZhao 33732). Scale bars: 10 µm (A–C).

Cystidia fusiform, slightly constricted at the neck and forms a long beak, thin-walled, 31–57 × 9–13.5 µm. Basidia clavate, slightly constricted in the middle, with four sterigmata and a basal clamp connection, colorless, thin-walled, 26.3–39.8 × 8.1–9.5 µm; basidioles in shape similar to basidia, but slightly smaller.

Basidiospores cylindrical to allantoid, colorless, thin-walled, IKI–, CB–, (9.1–)9.5–12.5(–12.9) × (3.7–)3.9–4.8(–5.5) µm, L = 10.89 µm, W = 4.30 µm, Q = 2.53–2.54 (n = 60/2).

Another specimen (paratype) examined.

China • Yunnan Province, Zhaotong, Xiaocaoba Town, Wumengshan National Nature Reserve, GPS coordinates: 27°33'N, 103°27'E, altitude 2300 m asl., on fallen angiosperm branch, leg. C.L. Zhao, 21 September 2023, CLZhao 33720 (SWFC).

Notes.

The phylogenetic analysis indicates that species P. punctata was placed within Peniophorella inferring from the dataset of ITS+nLSU. Morphologically, P. allantospora (Sheng H. Wu) K.H. Larss., P. capitulata (Boidin & Gilles) K.H. Larss., P. flagellata (G. Cunn.) K.H. Larss., P. pallida (Bres.) K.H. Larss. and P. praetermissa (P. Karst.) K.H. Larss. are similar to P. punctata based on the smooth hymenophore and allantoid basidiospores. However, P. allantospora differs from P. punctata by its shorter subutriformia to subclavata basidia (17–27 × 7.0–8.5 µm; Larsson 2007b). P. capitulata in its presence of subcylindrica leptocystidia (25–50 × 4–7 µm; Larsson 2007b). P. flagellata is distinct from P. punctata by its cream hymenial surface, obovate or subglobose stephanocysts, narrower basidia (28–40 × 7–8 µm) and shorter basidiospores (8–9 × 5–6 µm; Larsson 2007b). P. pallida can be delimited from P. punctata by its whitish to ochraceous hymenial surface and longer fusiform cystidia (50–70 × 6–8 µm) and smaller basidia (20–25 × 5–6 µm; Larsson 2007b). P. praetermissa can be separated from P. punctata by presence of three types of cystidia as fusiform gloeocystidia (50–100 × 8–12 µm), cylindrical leptocystidia (20–80 × 6–8 µm) and stephanocysts bowl-shaped, and narrower basidia (20–30 × 6–7 µm; Larsson 2007b).

Discussion

Many recently described wood-inhabiting fungal taxa have been reported worldwide, including in the genera Hymenochaete, Lyomyces and Peniophorella (Chen et al. 2017; Kan et al. 2017a, b; Pacheco et al. 2018; Miettinen et al. 2019; Viner and Miettinen 2022; Dong et al. 2024). The Xishuangbanna and Zhaotong are situated in the southwest of China; these areas become focal points for fungal diversity in China due to their complex topography and diverse ecosystems. In our study, four new species were found from Xishuangbanna and Zhaotong, Yunnan Province, China.

The family Hymenochaetaceae is characterized by annual to perennial, brownish basidiomata with a xanthochroic reaction in KOH, poroid or corticioid hymenophore, generative hyphae without clamp connections and setal elements present or absent (Wu et al. 2022a, b). Some forest pathogens species are mainly located in the genus Coniferiporia L.W. Zhou & Y.C. Dai, Onnia P. Karst. and Pyrrhoderma Imazeki (Zhou et al. 2016; Ji et al. 2017). Furthermore, Inonotus obliquus (Fr.) Pilát and Sanghuangporus sanghuang (Sheng H. Wu, T. Hatt. & Y.C. Dai) Sheng H. Wu, L.W. Zhou & Y.C. Dai have therapeutic properties (Wu et al. 2019b; Lv et al. 2023; Zhang et al. 2023a; Ma et al. 2024). Hymenochaetaceae is typed with the genus Hymenochaete Lév., which was erected in 1846 and typified by H. rubiginosa (Dicks.) Lév. The genus Hymenochaete can be identified by the annual to perennial, resupinate, effused-reflexed to pileate basidiomata with smooth, lamellate, tuberculate, poroid or hydnoid hymenophore, hyphal system monomitic or dimitic, presence of setae, and colorless, thin-walled, narrowly cylindrical to globose basidiospores (Léger 1998; Parmasto 2001; He and Dai 2012). There are some characteristics that can be used to circumscribe the genus, such as brown basidiomata darkening in potassium hydroxide, simple-septate generative hyphae, brown setae, and causing a white rot (Léger 1998; Parmasto 2001). In the present study, the new species H. bannaensis is distinguished by flocculent basidiomata with cinnamon to yellowish brown to rust-brown hymenial surface, generative hyphae with simple septa and broadly ellipsoid to globose basidiospores. Phylogenetically, H. bannaensis was close to H. colliculosa, H. globispora, and H. megaspora. However, morphologically, H. colliculosa is distinguished from H. bannaensis by its brown hymenial surface with tuberculate and larger ellipsoid basidiospores measuring 6–7.5 × 4–4.5 µm (Léger 1998). Furthermore, H. globispora differs from H. bannaensis by its deep reddish-brown hymenial surface with granular, longer setae (150 µm) with crystal, wider basidia (15–18 × 4.5–5.5 µm) and globular basidiospores (4 × 5.5 µm; Léger 1990). Moreover, H. megaspora is distinct from H. bannaensis by having effused or effused-reflexed with slightly elevated margins basidiomata (150–600 µm thick), grayish brown to dark gray, silky, tomentose, concentrically sulcate and zonate pileal surface, smooth or tuberculate hymenophore with pale mouse-gray to vinaceous gray, longer setae (90–120 × 8–13 µm), larger clavate basidia (25–32 × 5–8 µm) and larger, broadly ellipsoid basidiospores measuring 7.5–10 × 5–7 µm (He and Liu 2011).

The genus Lyomyces P. Karst. (Schizoporaceae) was established by Karsten (1881) with L. sambuci (Pers.) P. Karst. as the type species. It is characterized by the resupinate-to-effused basidiomata with a smooth to odontioid hymenophore, a monomitic hyphal system with generative hyphae bearing clamp connections, the presence of several types of cystidia and with smooth, thin- to slightly thick-walled basidiospores (Karsten 1881; Bernicchia and Gorjón 2010). Previously, 77 specific and infraspecific names were registered in the Index Fungorum (http://www.indexfungorum.org; accessed on 24 March 2025), of which approximately 65 species of Lyomyces are currently accepted (Cunningham 1963; Dai 2011; Gafforov et al. 2017; Luo et al. 2021a; Yuan et al. 2024; Dai et al. 2025). In this study, the new species Lyomyces asiaticus is characterized by the membranaceous basidiomata with white to cream hymenial surface with tuberculate, a monomitic hyphal system with clamped generative hyphae and ellipsoid basidiospores. The phylogenetic tree of ITS+nLSU (Figs 1, 3) showed that L. asiaticus was grouped closely with L. hengduanensis and L. zhaotongensis. However, morphologically, L. hengduanensis is distinct from L. asiaticus by having pruinose hymenial surface, fusoid cystidia (17.5–25 × 3–4 µm) and subclavate cystidia (16–23 × 3–4.5 µm), and smaller clavate basidia (10.5–14 × 3.5–5 µm; Yuan et al. 2024). L. zhaotongensis is distinguishable from L. asiaticus by its farinaceous basidiomata, fusoid cystidia (16–20.5 × 2.5–3.5 µm), smaller clavate basidia (14–16.5 × 2.5–3.5 µm) and smaller broadly ellipsoid basidiospores measuring 2.6–3.5 × 2.5–3 µm (Yuan et al. 2024).

The family Peniophorellaceae L.W. Zhou et al. was established by Wang et al. (2023a), to include the genus Peniophorella P. Karst, typed with P. pubera (Fr.) P. Karst. Species of the family are characterized by annual, resupinate, effused, adnate, thin, ceraceous basidiomata with smooth to tuberculate, white to yellowish hymenophore, a monomitic hyphal system with clamped generative hyphae thin-walled, and colorless, the presence of different kinds of cystidia, metuloids, gloeocystidia or leptocystidia, echinulate cells usually present, narrowly to broadly clavate basidia with four sterigmata, and hyaline, thin-walled, smooth, ellipsoid, cylindrical or allantoid, inamyloid, acyanophilous basidiospores with oily contents (Wang et al. 2023a). So far, 37 species have been accepted in the genus worldwide (Prasher 2015; Guan et al. 2020; Xu et al. 2020; Dong et al. 2024; Liu et al. 2024). In the present study, two new species have been identified; P. albohymenia is delimited by its membranaceous basidiomata with white hymenial surface, four types cystidia as stephanocyst, fusiform, cylindrical and capitate, and ellipsoid basidiospores. Furthermore, P. punctata is unique in the membranaceous, punctate basidiomata with white to pale yellow hymenial surface, fusiform cystidia, and allantoid basidiospores. In our phylogenies (Figs 1, 4), P. albohymenia was grouped with P. olivacea and P. subpraetermissa with strong supports. Morphologically, P. olivacea can be separated from P. albohymenia by having the ceraceous to farinaceous basidiomata, grayish olivaceous hymenial surface, two types cystidia as fusiform (28.5–39 × 8.5–11 µm) and halocystidia (42–50 × 8–9 µm), smaller basidia (17.5–20 × 6–7 µm) and smaller basidiospores (6.5–7.5 × 3.2–4.5 µm; Dong et al. 2024). P. subpraetermissa is distinct from P. albohymenia by its longer subclavata basidia (20–35 × 6–8 µm) and smaller basidiospores (6.2–7.5 × 4–5 µm; Wu 1997). Furthermore, P. punctata was the sister to P. fissurata with strong supports. However, morphologically, the latter having the ceraceous to corneus basidiomata, smooth to tuberculate and white to cream hymenial surface and larger fusiform cystidia (28–65 × 6.5–15 µm; Guan et al. 2020).

The wood-inhabiting fungi are an extensively studied group of Basidiomycota, and the taxa of Hymenochaetales play a core group in the wood-inhabiting fungi (Dai 2012; Ryvarden and Melo 2014; Liu et al. 2021; Zhou et al. 2022; Deng et al. 2024; Wang et al. 2024), but the species of Hymenochaetales diversity is still not well-known in China, especially in subtropical and tropical areas, and many recently described taxa of this ecologically important Hymenochaetales group were reported from China (Zhao et al. 2014, 2015, 2016; Chen et al. 2020; Guan et al. 2021; Yu et al. 2021; Dong et al. 2024). In the present study, four new species within the order Hymenochaetales are described, based on both morphological and molecular phylogenetic analyses, and are also from the subtropics. This study enriches the wood-inhabiting fungal diversity in China.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

The research was supported by the National Natural Science Foundation of China (Project Nos. 32170004, U2102220), Forestry and Grass Science and Technology Innovation Joint Project of Yunnan Province (Project No. 202404CB090008), and the High-level Talents Program of Yunnan Province (YNQR-QNRC-2018-111), and the Science Foundation of Education Department of Yunnan Province (2025Y0845).

Author contributions

Conceptualization: HY, CZ. Data curation: CZ, HY, QL, MC. Formal analysis: CZ, LL, MC, QL, SZ, YD, HY. Funding acquisition: CZ, HY. Investigation: CZ, HY, YD. Methodology: HY, YD, CZ. Project administration: CZ. Resources: HY, CZ. Software: CZ, HY, MC, QL, SZ, YD, LL. Supervision: HY, CZ, YD. Validation: YD, CZ, HY. Visualization: CZ, HY, YD. Writing - original draft: HY, SZ, YD, CZ, MC, LL. Writing - review and editing: YD, HY, CZ.

Author ORCIDs

Yinglian Deng https://orcid.org/0000-0002-8220-508X

Meng Chen https://orcid.org/0009-0005-9916-8704

Linfeng Liu https://orcid.org/0009-0003-4058-301X

Sicheng Zhang https://orcid.org/0009-0003-6287-9823

Haisheng Yuan https://orcid.org/0000-0001-7056-140X

Changlin Zhao https://orcid.org/0000-0002-8668-1075

Data availability

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

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