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Research Article
Morphological and molecular identification for four new wood-inhabiting species of Lyomyces (Basidiomycota) from China
expand article infoQi Yuan‡, Yunchao Li‡, Yunfei Dai‡, Kunyan Wang‡, Yixuan Wang§, Changlin Zhao‡
‡ Southwest Forestry University, Kunming, China
§ Yunnan Yunzhihuang Health Technology Co., LTD, Kunming, China

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

Academic editor: Samantha C. Karunarathna
© 2024 Qi Yuan, Yunchao Li, Yunfei Dai, Kunyan Wang, Yixuan Wang, 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: Yuan Q, Li Y, Dai Y, Wang K, Wang Y, Zhao C (2024) Morphological and molecular identification for four new wood-inhabiting species of Lyomyces (Basidiomycota) from China. MycoKeys 110: 67-92. https://doi.org/10.3897/mycokeys.110.133108
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Abstract

Fungi are one of the most diverse groups of organisms on Earth, in which the wood-inhabiting fungi play an important role in forest ecosystem processes and functions. Four new wood-inhabiting fungi, Lyomyces hengduanensis, L. niveomarginatus, L. wumengshanensis and L. zhaotongensis, are proposed, based on morphological features and molecular evidence. Lyomyces hengduanensis differs in the brittle basidiomata with pruinose hymenial surface, a monomitic hyphal system and ellipsoid basidiospores (3.5–6 × 3–4.5 µm). Lyomyces niveomarginatus is distinguished by the subceraceous basidiomata with crackled hymenial surface, a monomitic hyphal system and ellipsoid basidiospores (4.5–7 × 3–4 µm). Lyomyces wumengshanensis is distinguished by the grandinioid hymenial surface, a monomitic hyphal system and ellipsoid to broad ellipsoid basidiospores (4–6 × 3–5 µm). Lyomyces zhaotongensis is unique in the grandinioid hymenial surface, a monomitic hyphal system and broadly ellipsoid basidiospores measuring as 2.6–3.5 × 2.5–3 µm. Sequences of ITS and nLSU rRNA markers of the studied samples were generated and phylogenetic analyses were performed using the Maximum Likelihood, Maximum Parsimony and Bayesian Inference methods. The phylogram, based on the ITS+nLSU rDNA gene regions, included three genera within the Schizoporaceae viz. Fasciodontia, Lyomyces and Xylodon, in which the four new species were grouped into Lyomyces. The phylogenetic tree inferred from the ITS sequences highlighted that L. hengduanensis group with L. zhaotongensis and then closely grouped with L. crustosus, L. ochraceoalbus, and L. vietnamensis. The new taxon L. niveomarginatus was retrieved as a sister to L. juniperi. The new species L. wumengshanensis was sister to L. macrosporus. The new taxon L. zhaotongensis grouped with L. hengduanensis and then closely grouped with L. crustosus, L. ochraceoalbus and L. vietnamensis.

Key words

Biodiversity, phylogenetic analyses, taxonomy, Yunnan Province

Introduction

Fungi are one of the most diverse groups of organisms on Earth and play an indispensable role in the forest ecosystem processes and functioning (Hyde 2022; Guan et al. 2023; Deng et al. 2024a). The wood-inhabiting fungal family Schizoporaceae Jülich includes many variations of the fruiting body types within the order Hymenochaetales Oberw. (Larsson et al. 2006; Wu et al. 2022a; Guan et al. 2023; Zhang et al. 2024) and it comprises a number of representative wood-inhabiting fungal taxa, including diverse hymenophoral morphologies as hydnoid, corticioid and polyporoid (Yurchenko and Wu 2016; Riebesehl and Langer 2017; Yurchenko et al. 2017; Cui et al. 2019; Riebesehl et al. 2019; Jiang et al. 2021; Wu et al. 2022a, 2022b; Guan et al. 2023; Deng et al. 2024a, b; Zhang et al. 2024). In addition, taxa of the family Schizoporaceae are widely found in different continents, causing white rot (Langer 1994; Luo et al. 2022; Guan et al. 2023; Zhang et al. 2024).

The genus Lyomyces P. Karst. is typified by L. sambuci (Pers.) P. Karst. It is characterised 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). Based on the MycoBank database (http://www.mycobank.org, accessed on 25 April 2024) and the Index Fungorum (http://www.indexfungorum.org, accessed on 25 April 2024), Lyomyces has 55 specific and infraspecific names registered, of which approximately 41 species of Lyomyces are currently known (Rabenhorst 1851; Karsten 1881; Karsten 1882; Cunningham 1959; Cunningham 1963; Wu 1990; Hjortstam and Ryvarden 2009; Xiong et al. 2009; Dai 2010; Dai 2011; Yurchenko and Wu 2013; Gafforov et al. 2017; Riebesehl and Langer 2017; Yurchenko et al. 2017; Chen and Zhao 2020; Yurchenko et al. 2020; Luo et al. 2021b; Luo et al. 2021c; Viner et al. 2022; Guan et al. 2023).

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. 2020). These pioneering research studies into the family Schizoporaceae were just the prelude to the molecular systematics period (Guan et al. 2023; Zhang et al. 2024). The genus Hyphodontia s.l. was indicated to be a polyphyletic group, in which the genera Xylodon (Pers.) Gray and Kneiffiella P. Karst. included the largest number of species (Yurchenko and Wu 2016; Riebesehl and Langer 2017; Riebesehl et al. 2019). Due to the lack of sequences of some wood-inhabiting fungal taxa, it is difficult to clearly distinguish many genera in this family Schizoporaceae using molecular data; therefore, a broad concept of Hyphodontia s.l. was accepted (Yurchenko and Wu 2016; Riebesehl and Langer 2017; Wang and Chen 2017; Riebesehl et al. 2019). Based on the nuclear DNA sequence data, six well-distinguished clades as Hastodontia clade, Hyphodontia clade, Lagarobasidium clade, Kneiffiella-Alutaceodontia clade, Xylodon-Lyomyces-Rogersella clade and Xylodon-Schizopora-Palifer clade, were included, based on the phylogenetical studies for Hyphodontia s.l., in which the genus Lyomyces was nested within the Xylodon-Lyomyces-Rogersella clade (Yurchenko and Wu 2013). The research revealed that Hyphodontia s.l. was divided into six genera, viz., Hastodontia (Parmasto) Hjortstam & Ryvarden, Hyphodontia J. Erikss., Kneiffiella, Lagarobasidium Jülich, Lyomyces and Xylodon, in which 35 new combinations were proposed, including fourteen Lyomyces species (Riebesehl and Langer 2017). On the basis of the sequences of the internal transcribed spacer (ITS) and the nuclear large subunit (nLSU) ribosomal DNA gene, the phylogenetic analysis clarified that the Lyomyces sambuci complex divided into four new species (Yurchenko et al. 2017). Riebesehl et al. (2019) clarified the generic concept and their phylogenetic reconstruction of Lyomyces and the species L. sambuci was sister to L. crustosus (Pers.) P. Karst (Riebesehl et al. 2019). Based on a combination of the morphological and molecular evidence, the fungal diversity of the family Schizoporaceae was analysed, in which six new species were described: L. fissuratus C.L. Zhao, L. fumosus C.L. Zhao, L. niveus C.L. Zhao, L. ochraceoalbus C.L. Zhao, L. albopulverulentus C.L. Zhao and L. yunnanensis (Luo et al. 2021b, 2021c; Guan et al. 2023).

During the investigations of the wood-inhabiting fungi, we collected four new Hymenochaetales taxa from Yunnan Province, China, that could not be assigned to any described species of the order. We present the morphological characteristics and phylogenetic analyses with ITS and nLSU that support the four species in the genus Lyomyces.

Materials and methods

Morphology

Fresh basidiomata of the fungi growing on the angiosperm branch were collected from the Honghe, Lincang, Puer, Wenshan and Zhaotong of Yunnan Province, P.R. China after recording important information (Rathnayaka et al. 2024). Specimens were dried in an electric food dehydrator at 40 °C (Hu et al. 2022), then sealed and stored in an envelope bag and deposited in the Herbarium of the Southwest Forestry University (SWFC), Kunming, Yunnan Province, P.R. China. Macromorphological descriptions were based on field notes and photos were captured in the field and lab. Colour terminology follows Petersen (Petersen 1996). Micromorphological data were obtained from the dried specimens when observed under a light microscope following the previous study (Guan et al. 2023). 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 all spores), W = mean spore width (arithmetic average for all 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).

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 (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 the previous study (Zhao and Wu 2017). All 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.

Species name Specimen No. GenBank accession No. References
ITS nLSU
Fasciodontia brasiliensis MSK-F 7245a MK575201 MK598734 Yurchenko et al. (2020)
F. bugellensis KAS-FD 10705a MK575203 MK598735 Yurchenko et al. (2020)
F. bugellensis MSK-F 7353 MK575205 MK598736 Yurchenko et al. (2020)
F. yunnanensis CLZhao 6280 MK811275 MZ146327 Luo and Zhao (2021)
F. yunnanensis CLZhao 6385 MK811277 — Luo and Zhao (2021)
Hymenochaete ochromarginata He 47 KU978861 JQ279666 Unpublished
H. rubiginosa He 458 JQ279580 — He and Li (2013)
Lyomyces albopulverulentus CLZhao 21478 OP730712 OP730724 Guan et al. (2023)
L. allantosporus KAS-GEL4933 KY800401 — Yurchenko et al. (2017)
L. allantosporus FR-0249548 KY800397 — Yurchenko et al. (2017)
L. bambusinus CLZhao 4831 MN945968 — Chen and Zhao (2020)
L. bambusinus CLZhao 4808 MN945970 — Chen and Zhao (2020)
L. cremeus CLZhao 4138 MN945974 — 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 — Unpublished
L. densiusculus Ryvarden 44818 OK273853 — Viner et al. (2022)
L. elaeidicola LWZ20180411-20 MT319458 — Wang et al. (2021)
L. elaeidicola LWZ20180411-19 MT319457 — Wang et al. (2021)
L. erastii TASM:YG 022 MF382992 — Gafforov et al. (2017)
L. erastii 23cSAMHYP JX857800 — Unpublished
L. fimbriatus Wu910620-7 MK575209 — Yurchenko et al. (2020)
L. fimbriatus Wu911204-4 MK575210 — Yurchenko et al. (2020)
L. fissuratus CLZhao 4352 MW713742 — Luo et al. (2021b)
L. fissuratus CLZhao 4291 MW713738 — Luo et al. (2021b)
L. fumosus CLZhao 8188 MW713744 — Luo et al. (2021b)
L. gatesiae LWZ20180515-3 MT319447 — Wang et al. (2021)
L. gatesiae LWZ20180515-32 MT319448 — Wang et al. (2021)
L. griseliniae KHL 12971 (GB) DQ873651 — Larsson et al. (2006)
L. hengduanensis CLZhao 20627 OR793233 PP657611 Present study
L. hengduanensis CLZhao 25551 OR658999 PP657610 Present study
L. hengduanensis CLZhao 32713 OR899153 — Present study
L. hengduanensis CLZhao 32714 OR899154 — Present study
L. hengduanensis CLZhao 32782 OR899155 PP657612 Present study
L. juniperi FR-0261086 KY081799 — Riebesehl and Langer (2017)
L. leptocystidiatus LWZ20170818-1 MT326514 — Wang et al. (2021)
L. leptocystidiatus LWZ20170818-2 MT326513 — Wang et al. (2021)
L. macrosporus CLZhao 4516 MN945977 — Chen and Zhao (2020)
L. mascarensis KAS-GEL4833 KY800399 — Yurchenko et al. (2020)
L. mascarensis KAS-GEL4908 KY800400 — Yurchenko et al. (2020)
L. microfasciculatus CLZhao 5109 MN954311 — Chen and Zhao (2020)
L. niveomarginatus CLZhao 16360 PP537949 PP657607 Present study
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. ochraceoalbus MSK7247 KY800403 — Yurchenko et al. (2017)
L. orientalis GEL3376 DQ340325 — Yurchenko et al. (2017)
L. pruni GEL2327 DQ340312 — Larsson et al. (2006)
L. pruni Ryberg 021018 (GB) DQ873624 — Larsson et al. (2006)
L. sambuci KAS-JR7 KY800402 KY795966 Yurchenko et al. (2017)
L. sambuci 83SAMHYP JX857721 — Yurchenko et al. (2017)
L. vietnamensis TNM F9073 JX175044 — Yurchenko et al. (2017)
L. wuliangshanensis CLZhao 4108 MN945980 — Chen and Zhao (2020)
L. wuliangshanensis CLZhao 4167 MN945979 — Chen and Zhao (2020)
L. wumengshanensis CLZhao 29374 OR803021 PP657613 Present study
L. wumengshanensis CLZhao 31486 OR899208 — Present study
L. wumengshanensis CLZhao 32705 OR899209 — Present study
L. wumengshanensis CLZhao 32736 OR899210 — Present study
L. wumengshanensis CLZhao 32800 OR899211 PP657614 Present study
L. wumengshanensis CLZhao 32869 OR899212 — Present study
L. wumengshanensis CLZhao 32915 OR899213 PP657615 Present study
L. yunnanensis CLZhao 2463 OP730711 OP730723 Guan et al. (2023)
L. yunnanensis CLZhao 9375 OP730710 — Guan et al. (2023)
L. yunnanensis CLZhao 10041 OP730709 — Guan et al. (2023)
L. zhaotongensis CLZhao 32878 PP537950 PP657609 Present study
Xylodon afromontanus H 7006811 OQ645463 — Yurchenko et al. (2024)
X. asiaticus CLZhao 10368 OM959479 — Zhang et al. (2024)
X. cystidiatus FR-0249200 MH880195 MH884896 Riebesehl et al. (2019)
X. daweishanensis CLZhao 18492 OP730719 OP730727 Guan et al. (2023)
X. daweishanensis CLZhao 18446 OP730717 OP730725 Guan et al. (2023)
X. filicinus MSK-F 12869 MH880199 NG067836 Riebesehl et al. (2019)
X. fissuratus CLZhao 7007 OP730713 — Guan et al. (2023)
X. fissuratus CLZhao 9407 OP730714 — Guan et al. (2023)
X. hastifer K(M) 172400 NR166558 — Riebesehl and Langer (2017)
X. hyphodontinus KAS-GEL9222 MH880205 MH884903 Riebesehl et al. (2019)
X. macrosporus CLZhao 10226 MZ663809 MZ663817 Luo et al. (2021a)
X. puerensis CLZhao 8142 OP730720 OP730728 Guan et al. (2023)
X. puerensis CLZhao 8639 OP730721 OP730729 Guan et al. (2023)
X. quercinus Larsson 11076 (GB) KT361633 — Larsson et al. (2004)
X. ramicida Spirin 7664 NR138013 — Unpublished
X. subflaviporus Wu 0809-76 KX857803 — Chen et al. (2017)
X. tropicus CLZhao 3351 OL619261 OL619269 Qu et al. (2022)
X. wenshanensis CLZhao 15729 OM338097 OM338104 Luo et al. (2022)
X. xinpingensis CLZhao 11224 MW394662 MW394654 Luo et al. (2022)

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 sequence alignments were deposited in figshare (DOI: 10.6084/m9.figshare.27166521). Sequences of Hymenochaete ochromarginata P.H.B. Talbot and Hymenochaete rubiginosa (Dicks.) Lév., retrieved from GenBank, were used as the outgroups in the ITS+nLSU analysis (Fig. 1). The sequence alignments were deposited in figshare (DOI: 10.6084/m9.figshare.27166521). Sequences of Xylodon quercinus (Pers.) Gray and Xylodon ramicida Spirin & Miettinen, retrieved from GenBank, were used as the outgroups in the ITS analysis (Fig. 2) (Guan et al. 2023; Zhang et al. 2024).

Figure 1. 

Maximum Parsimony strict consensus tree illustrating the phylogeny of four new species and related species in Lyomyces within Schizoporaceae, based on ITS+nLSU sequences. Branches are labelled 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 four new species and related species in Lyomyces, based on ITS sequences. Branches are labelled 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 (Zhao and Wu 2017). 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 - 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 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 1.905 million generations for ITS+nLSU (Fig. 1) and 2 million generations for ITS (Fig. 2), with trees and parameters sampled every 1,000 generations. The first quarter of all of the generations were discarded as burn-in. 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.

Results

Molecular phylogeny

The ITS+nLSU dataset (Fig. 1) comprised sequences from 40 fungal specimens representing 29 taxa. The dataset had an aligned length of 2,112 characters, of which 1,298 characters were constant, 254 were variable and parsimony-uninformative and 560 were parsimony-informative. Maximum parsimony analysis yielded one equally parsimonious tree (TL = 2,513, CI = 0.4990, HI = 0.5010, RI = 0.6658 and RC = 0.3322). 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 = 0.009992 (BI) and the effective sample size (ESS) across the two runs is double the average ESS (avg. ESS) = 2078.5. The phylogram, based on the ITS+nLSU rDNA gene regions (Fig. 1), included three genera within Schizoporaceae (Hymenochaetales), which were Fasciodontia, Lyomyces and Xylodon, in which four new species were grouped into the genera Lyomyces.

The ITS dataset (Fig. 2) comprised sequences from 57 fungal specimens representing 33 taxa. The dataset had an aligned length of 696 characters, of which 270 characters were constant, 41 were variable and parsimony-uninformative and 385 were parsimony-informative. Maximum parsimony analysis yielded 80 equally parsimonious tree (TL = 1,748, CI = 0.4027, HI = 0.5973, RI = 0.6935 and RC = 0.2793). The best model of nucleotide evolution for the ITS 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 = 0.014964 (BI) and the effective sample size (ESS) across the two runs is double the average ESS (avg. ESS) = 1,387.5. The phylogenetic tree (Fig. 2), inferred from the ITS sequences, highlighted that L. hengduanensis group with L. zhaotongensis; and then closely grouped with L. crustosus (Pers.) P. Karst., L. ochraceoalbus C.L. Zhao and L. vietnamensis (Yurchenko & Sheng H. Wu) Riebesehl & Langer. Lyomyces niveomarginatus was retrieved as a sister to L. juniperi (Bourdot & Galzin) Riebesehl & Langer. Lyomyces wumengshanensis was retrieved as a sister to L. macrosporus C.L. Zhao. Moreover, Lyomyces zhaotongensis grouped with L. hengduanensis and closely clustered with L. crustosus, L. ochraceoalbus and L. vietnamensis.

Taxonomy

Lyomyces hengduanensis Q. Yuan & C.L. Zhao, sp. nov.

MycoBank No: 853724
Figs 3, 4

Type material

Holotype. China • Yunnan Province, Lincang, Fengqing County, Yaojie Town, GPS coordinates 24°66'N, 100°19'E, altitude 2060 m, on a fallen branch of angiosperm, leg. C.L. Zhao, 22 October 2022, CLZhao 25551 (SWFC).

Figure 3. 

Basidiomata of Lyomyces hengduanensis (holotype). Scale bars: 1 cm (A); 2 mm (B).

Etymology

Hengduanensis (Lat.) refers to the type locality “Hengduan Mountain”.

Figure 4. 

Microscopic structures of Lyomyces hengduanensis (holotype): basidiospores (A), basidia (B), basidioles (C), fusoid cystidia (D), subclavate cystidia (E), a section of hymenium (F). Scale bars: 20 µm (A–F).

Description

Basidiomata annual, resupinate, adnate, brittle, without odour and taste when fresh and up to 3.5 cm long, 1 cm wide, 100 µm thick. Hymenial surface pruinose, white to cream when fresh, to cream to slightly buff upon drying. Sterile margin white to cream and up to 1 mm wide.

Hyphal system monomitic, generative hyphae with clamp connections, colourless, thick-walled, branched, 2–3 µm in diameter; IKI–, CB–, tissues unchanged in KOH. Numerous crystals present amongst generative hyphae.

Cystidia of two types: (1) fusoid, colourless, thin-walled, smooth, slightly constricted in the middle to somewhat sinuous, 17.5–25 × 3–4 µm; (2) subclavate, colourless, thin-walled, smooth, slightly constricted in the middle to somewhat sinuous, 16–23 × 3–4.5 µm; basidia clavate, with 4 sterigmata and a basal clamp connection, 10.5–14 × 3.5–5 µm.

Basidiospores ellipsoid, colourless, thin-walled, smooth, with one oil drop, CB–, IKI–, 3.5–6 × 3–4.5 µm, L = 4.63 µm, W = 3.65 µm, Q = 1.25–1.28 (n = 90/3).

Additional specimens examined

(paratypes). China • Yunnan Province, Zhaotong, Qiaojia County, Yaoshan Town, Yaoshan National Nature Reserve, 26°50'N, 102°59'E, altitude 2500 m, on a fallen branch of angiosperm, leg. C.L. Zhao, 22 August 2020, CLZhao 20627 (SWFC) • Zhaotong, Wumeng Mountain National Nature Reserve, GPS coordinates 27°72'N, 103°92'E, altitude 1424 m, on a fallen branch of angiosperm, leg. C.L. Zhao, 29 August 2023, CLZhao 32713, CLZhao 32714, CLZhao 32782 (SWFC).

Lyomyces niveomarginatus Q. Yuan & C.L. Zhao, sp. nov.

MycoBank No: 853725
Figs 5, 6

Type material

Holotype. China • Yunnan Province, Wenshan, Wenshan National Nature Reserve, GPS coordinates 23°21'N, 104°10'E, altitude 1950 m, on a fallen branch of angiosperm, leg. C.L. Zhao, 26 July 2019, CLZhao 16360 (SWFC).

Figure 5. 

Basidiomata of Lyomyces niveomarginatus (holotype). Scale bars: 1 cm (A); 2 mm (B).

Etymology

Niveomarginatus (Lat.) refers to the niveous margin of basidiomata.

Figure 6. 

Microscopic structures of Lyomyces niveomarginatus (holotype): basidiospores (A), basidia (B), basidioles (C), fusoid cystidia (D), clavate cystidia (E), a section of hymenium (F). Scale bars: 20 µm (A–F).

Description

Basidiomata annual, resupinate, adnate, subceraceous, without odour and taste when fresh and up to 7.5 cm long, 2 cm wide, 150 µm thick. Hymenial surface crackled, white to cream when fresh, to cream to slightly buff upon drying. Sterile margin distinct, whitish and up to 2 mm wide.

Hyphal system monomitic, generative hyphae with clamp connections, colourless, thin-walled, branched, 1.5–3.5 µm in diameter; IKI–, CB–, tissues unchanged in KOH. Numerous crystals present amongst generative hyphae.

Cystidia of two types: (1) fusoid, colourless, thin-walled, smooth, 25–29 × 2–3 µm; (2) clavate, colourless, thin-walled, smooth, 20–25.5 × 4.5–5.5 µm; basidia subclavate, with 4 sterigmata and a basal clamp connection, 23–29 × 2.5–3.5 µm.

Basidiospores ellipsoid, colourless, thin-walled, smooth, with one oil drop, CB–, IKI–, 4.5–7 × (2.5–)3–4 µm, L = 5.51 µm, W = 3.15 µm, Q = 1.75 (n = 30/1).

Additional specimens examined

(paratypes). China • Yunnan Province, Wenshan, Wenshan National Nature Reserve, GPS coordinates 23°21'N, 104°10'E, altitude 1950 m, on a fallen branch of angiosperm, leg. C.L. Zhao, 7 August 2024, CLZhao 40333, CLZhao 40334 (SWFC).

Lyomyces wumengshanensis Q. Yuan & C.L. Zhao, sp. nov.

MycoBank No: 853726
Figs 7, 8

Type material

Holotype. China • Yunnan Province, Zhaotong, Daguan County, Wumeng Mountain National Nature Reserve, GPS coordinates 27°72'N, 103°92'E, altitude 1424 m, on a fallen branch of angiosperm, leg. C.L. Zhao, 3 July 2023, CLZhao 29374 (SWFC).

Figure 7. 

Basidiomata of Lyomyces wumengshanensis (holotype). Scale bars: 1 cm (A); 2 mm (B).

Etymology

Wumengshanensis (Lat.) refers to the type locality “Wumeng Mountain”.

Figure 8. 

Microscopic structures of Lyomyces wumengshanensis (holotype): basidiospores (A), basidia (B), basidioles (C), capitate cystidia (D), a section of hymenium (E). Scale bars: 10 µm (A–E).

Description

Basidiomata annual, resupinate, adnate, coriaceous when fresh, becoming farinaceous upon drying, without odour and taste when fresh and up to 5 cm long, 2 cm wide, 150 µm thick. Hymenial surface grandinioid, white when fresh, to cream upon drying. Sterile margin white and up to 1 mm wide.

Hyphal system monomitic, generative hyphae with clamp connections, colourless, thick-walled, branched, 3–4 µm in diameter; IKI–, CB–, tissues unchanged in KOH. Numerous crystals present amongst generative hyphae.

Cystidia capitate, colourless, thin-walled, smooth, 24.5–29 × 3–4 µm; basidia subclavate to barrelled, colourless, with 4 sterigmata and a basal clamp connection, 11.5–14 × 5.5–6.5 µm.

Basidiospores ellipsoid to broad ellipsoid, colourless, thin-walled, smooth, with one oil drop, CB–, IKI–, 4–6 × 3–5 µm, L = 5.4 µm, W = 4.2 µm, Q = 1.28–1.32 (n = 120/4).

Additional specimen examined

(paratype). China • Yunnan Province, Zhaotong, Wumeng Mountain National Nature Reserve, 27°72'N, 103°92'E, altitude 1424 m, on a fallen branch of angiosperm, leg. C.L. Zhao, 29 August 2023, CLZhao 31486, CLZhao 32705, CLZhao 32736, CLZhao 32800, CLZhao 32869, CLZhao 32915, CLZhao 32933 (SWFC).

Lyomyces zhaotongensis Q. Yuan & C.L. Zhao, sp. nov.

MycoBank No: 853727
Figs 9, 10

Type material

Holotype. China •Yunnan Province, Zhaotong, Wumeng Mountain National Nature Reserve, GPS coordinates 27°77'N, 104°29'E, altitude 2900 m, on the fallen branch of angiosperm, leg. C.L. Zhao, 29 August 2023, CLZhao 32878 (SWFC).

Figure 9. 

Basidiomata of Lyomyces zhaotongensis (holotype). Scale bars: (A) 1 cm; (B) 2 mm.

Etymology

Zhaotongensis (Lat.) refers to the type locality “Zhaotong”.

Figure 10. 

Microscopic structures of Lyomyces zhaotongensis (holotype): basidiospores (A), basidia (B), basidioles (C), fusoid cystidia (D), a section of hymenium (E). Scale bars: 20 µm (A–E).

Description

Basidiomata annual, resupinate, adnate, farinaceous when fresh, becoming coriaceous upon drying and up to 9.5 cm long, 3 cm wide, 30–80 um thick. Hymenial surface grandinioid, cream when fresh and cream to buff upon drying. Sterile margin white to cream and up to 1 mm wide.

Hyphal system monomitic, generative hyphae with clamp connections, colourless, thick-walled, branched, 1.5–2 µm in diameter; IKI–, CB–, tissues unchanged in KOH. Numerous crystals present amongst generative hyphae.

Cystidia fusoid, colourless, thin-walled, smooth, 16–20.5 × 2.5–3.5 µm. Basidia clavate, with 4 sterigmata and a basal clamp connection, 14–16.5 × 2.5–3.5 µm.

Basidiospores broadly ellipsoid, colourless, thin-walled, smooth, with oil drops, CB–, IKI–, 2.6–3.5 × 2.5–3 µm, L = 2.99 µm, W = 2.75 µm, Q = 1.08 (n = 30/1).

Additional specimen examined

(paratype). China • Yunnan Province, Zhaotong, Wumeng Mountain National Nature Reserve, GPS coordinates 27°77'N, 104°29'E, altitude 2900 m, on the fallen branch of angiosperm, leg. C.L. Zhao, 10 August 2024, CLZhao 40335 (SWFC).

Discussion

Many recently new wood-inhabiting fungal taxa have been reported in the subtropics and tropics, including in the genus Lyomyces (Xiong et al. 2009; Chen et al. 2017; Kan et al. 2017a, b; Riebesehl and Langer 2017; Viner et al. 2018; Chen and Zhao 2020; Luo et al. 2021a, b, c, 2022; Qu and Zhao 2022; Qu et al. 2022; Viner et al. 2022; Guan et al. 2023; Deng et al. 2024a, b; Zhang et al. 2024). Prior to this study, the following sixteen Lyomyces species were reported from China as L. albopulverulentus C.L. Zhao, L. albus (Sheng H. Wu) Riebesehl & Langer, L. bambusinus, L. capitatocystidiatus (H.X. Xiong, Y.C. Dai & Sheng H. Wu) Riebesehl & Langer, L. cremeus C.L. Zhao, L. fissuratus, L. fumosus, L. leptocystidiatus Xue W. Wang & L.W. Zhou, L. macrosporus C.L. Zhao & K.Y. Luo, L. microfasciculatus (Yurchenko & Sheng H. Wu) Riebesehl & Langer, L. niveus, L. ochraceoalbus, L. sambuci, L. tenuissimus (Yurchenko & Sheng H. Wu) Riebesehl & Langer, L. wuliangshanensis C.L. Zhao and L. yunnanensis C.L. Zhao (Xiong et al. 2009; Yurchenko et al. 2013; Riebesehl and Langer 2017; Chen and Zhao 2020; Luo et al. 2021b, c; Wang et al. 2021). The present study reports four new species in the genus Lyomyces, based on a combination of morphological features and molecular evidence.

Phylogenetically, based on the multiple loci in Hyphodontia s.l., six genera of Fasciodontia, Hastodontia, Hyphodontia, Lyomyces, Kneiffiella and Xylodon, were divided into four clades in the wood-inhabiting fungal order Hymenochaetales (Wang et al. 2021). In the present study, the phylogram inferred from the ITS+nLSU data, four new species grouped into the genus Lyomyces (Fig. 1). Based on ITS topology (Fig. 2), in which L. hengduanensis group with L. zhaotongensis and then closely grouped with L. crustosus, L. ochraceoalbus and L. vietnamensis. Lyomyces niveomarginatus was retrieved as a sister to L. juniperi. L. wumengshanensis was sister to L. macrosporus. Moreover, L. zhaotongensis grouped with L. hengduanensis and then closely clustered with three species: L. crustosus, L. ochraceoalbus and L. vietnamensis. However, morphologically, L. zhaotongensis can be delimited from L. hengduanensis by its the grandinioid hymenial surface and longer basidia (14–16.5 × 2.5–3.5 µm); L. crustosus can be separated from L. hengduanensis by its odontioid hymenial surface and narrow basidiospores (5–7.5 × 2.5–3 µm) (Lentz and McKay 1976); L. ochraceoalbus differs in L. hengduanensis by having a smooth hymenial surface and lacking a cystidium (Luo et al. 2021c); L. vietnamensis differs from L. hengduanensis by its aculeate hymenial surface and narrow basidiospores (5.8–6.1 × 2.6–2.9 µm; Yurchenko and Wu (2013)). L. juniperi can be delimited from L. niveomarginatus by its smooth hymenial surface with some scattered small granules and wider basidia (15–25 × 4–4.5 µm; Hjortstam and Ryvarden (2004)); L. macrosporus can be separated from L. wumengshanensis by its reticulate hymenial surface and longer basidiospores (6.7–8.9 × 4.4–5.4 µm; Chen and Zhao (2020)); L. crustosus can be delimited from L. zhaotongensis by its odontioid hymenial surface and longer basidiospores (5–7.5 × 2.5–3 µm; Lentz and McKay (1976)); L. hengduanensis can be delimited from L. zhaotongensis by its pruinose hymenial surface and shorter basidia (14–16.5 × 2.5–3.5 µm); L. ochraceoalbus differs in L. zhaotongensis by having smooth hymenial surface and longer basidiospores (4–5 × 2.5–3.5 µm; Luo et al. (2021c)); L. vietnamensis can be delimited from L. zhaotongensis by its aculeate hymenial surface and longer basidiospores (5.8–6.1 × 2.6–2.9 µm; Yurchenko and Wu (2013)).

Morphologically, Lyomyces hengduanensis resembles four taxa viz. L. albopulverulentus, L. bambusinus, L. mascarensis Riebesehl, Yurch. & Langer and L. yunnanensis, by the similar ellipsoid basidiospores. However, L. albopulverulentus differs from L. hengduanensis by its larger basidia (24.5–28.5 × 7–9 µm) and basidiospores (8–10.5 × 5.5–7 µm; Guan et al. (2023)); L. bambusinus can be separated from L. hengduanensis by its colliculose to tuberculate hymenial surface and longer basidia (16.5–35 × 3.5–7 µm; Chen and Zhao (2020)); L. mascarensis is distinct from L. hengduanensis by having indistinctly colliculose hymenial surface and longer basidia (16–17.5 × 3.5–4.5 µm; Yurchenko et al. (2017)); L. yunnanensis is distinguished from L. hengduanensis by its grandinioid hymenial surface and longer basidia (16.5–27 × 4–5.5 µm; Guan et al. (2023)).

Morphologically, Lyomyces niveomarginatus resembles several species viz. L. albopulverulentus, L. cremeus, L. macrosporus, L. wuliangshanensis and L. yunnanensis by the cream to buff hymenial surface and ellipsoid basidiospores. However, L. albopulverulentus differs from L. niveomarginatus by its pruinose hymenial surface and wider basidia (24.5–28.5 × 7–9 µm; Guan et al. (2023)); L. cremeus can be separated from L. niveomarginatus by its smooth hymenial surface and shorter basidia (9–18.5 × 3–6 µm; Chen and Zhao (2020)); L. macrosporus differs from L. niveomarginatus by its reticulate hymenial surface and wider basidia (23–29 × 2.5–3.5 µm) and wider basidiospores (6.7–8.9 × 4.4–5.4 µm; Chen and Zhao (2020)); L. wuliangshanensis can be delimited from L. niveomarginatus by its smooth to more or less tuberculate hymenial surface and shorter basidia (12–20 × 3–4.3 µm; Chen and Zhao (2020)); L. yunnanensis is distinct from L. niveomarginatus by having grandinioid hymenial surface and wider basidia (16.5–27 × 4–5.5 µm; Guan et al. (2023)).

Morphologically, Lyomyces wumengshanensis resembles L. bambusinus, L. cremeus, L. fumosus, L. fissuratus, L. wuliangshanensis and L. yunnanensis by having the capitate cystidia. However, L. bambusinus is distinct from L. wumengshanensis by possessing tapering cystidia (40–65 × 4–5.5 µm) and longer basidia (16.5–35 × 3.5–7 µm; Chen and Zhao (2020)); L. cremeus differs from L. wumengshanensis by its smooth hymenial surface and possesses tapering cystidia (18–35 × 3–4.5 µm; Chen and Zhao (2020)); L. fumosus can be separated from L. wumengshanensis by its smooth, smoky grey hymenial surface and narrower basidia (11.5–17.5 × 3–5 µm; Luo et al. (2021b)); L. fissuratus can be delimited from L. wumengshanensis by its longer and narrower basidia (14.7–23.3 × 2.9–4.8 µm; Luo et al. (2021b)); L. wuliangshanensis differs from L. wumengshanensis by its smooth to more or less tuberculate hymenial surface and narrower basidia (12–20 × 3–4.3 µm; Chen and Zhao (2020)); L. yunnanensis is separated from L. wumengshanensis by the longer basidia (16.5–27 × 4–5.5 µm) and possessing fusiform cystidia (18–39 × 4–6 µm; Guan et al. (2023)).

Morphologically, Lyomyces zhaotongensis reminds L. albopulverulentus, L. cremeus, L. denudatus Viner, L. macrosporus and L. wuliangshanensis by having the ellipsoid basidiospores. However, L. albopulverulentus can be separated from L. zhaotongensis 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. zhaotongensis by its smooth hymenial surface and larger basidiospores (4.5–5.6 × 3.3–4.3 µm; Chen and Zhao (2020)); L. denudatus is separated from L. zhaotongensis by the smooth hymenial surface and longer basidiospores (4.8–7 × 2.8–4.2 µm; Viner and Miettinen (2022)); L. macrosporus differs from L. zhaotongensis due to its reticulate hymenial surface and larger basidia (22.2–38 × 4.5–7 µm) and larger basidiospores (6.7–8.9 × 4.4–5.4 µm; Chen and Zhao (2020)); L. wuliangshanensis can be delimited from L. zhaotongensis by its smooth to more or less tuberculate hymenial surface and longer basidiospores (3.5–5.3 × 2.8–4 µm; Chen and Zhao (2020)). A morphological comparison amongst four new Lyomyces species and seven similar species are presented in Table 2.

Table 2.
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A morphological comparison between four new Lyomyces species and seven similar species in the genus Lyomyces. The bold are new taxa.

Species name Hymenial surface Generative hyphae Cystidia Basidia Basidiospores References
Lyomyces albopulverulentus Pruinose/ white Thick-walled/frequently branched Capitate, 37–54 × 5–9 µm Clavate, 24.5–28.5 × 7–9 µm Ellipsoid, (7.5–)8–10.5(–11) × (5–)5.5–7 µm Guan et al. (2023)
Lyomyces bambusinus Colliculose to tuberculate/ cream to buff Thick-walled/ branched Capitate, 35–55 × 4–7 µm; tapering, 40–65 × 4–5.5 µm, cystidioles, 12–17 × 2–3 µm Clavate, 16.5–35 × 3.5–7 µm Broadly ellipsoid, (4.5–)4.7–5.9 (–6.2) × (3.4–)3.7–4.6(–4.8) µm Chen and Zhao (2020)
Lyomyces cremeus Smooth/ pale cream Thick-walled/ branched Capitate, 20–40 × 3–5 µm; tapering, 18–35 × 3–4.5 µm Clavate, 9–18.5 × 3–6 µm Ellipsoid, 4.5–5.6(–5.8) × 3.3–4.3(–4.5) µm Chen and Zhao (2020)
Lyomyces denudatus Smooth/ cream Thin-walled to slightly thick-walled Capitate, (21–)34.9–62 × (3.5–)4–5.5(–7) μm Suburniform, 15–21.1(–25) × 3.8–5.5 μm Ellipsoid, (4.1–)4.8–7 × 2.8–4.2(–4.7) μm Viner and Miettinen (2022)
Lyomyces hengduanensis Pruinose/ cream to slightly buff Thick-walled/ branched Fusoid, 17.5–25 × 3–4 µm; subclavate, 16–23 × 3–4.5 µm Clavate, 10.5–14 × 3.5–5 µm Ellipsoid, 3.5–6 × 3–4.5 µm Present study
Lyomyces mascarensis Smooth / cream or brownish Thin-walled Capitate, 17–38 × 3.5–6(–7) µm; submoniliform, 18–22 × 5–5.5 µm; tapering, 25–30 × 3.5–4.5 µm Subcylindrical with one constriction, 16–17.5(–19) × 3.5–4.5(–6) µm Ellipsoid or broadly ellipsoid, (4–)4.5–6 × (3–)3.3–4 µm Yurchenko et al. (2017)
Lyomyces niveomarginatus Smooth / cream to slightly buff Thin-walled, branched Fusoid, 25–29 × 2–3 µm; clavate, 20–25.5 × 4.5–5.5 µm Subclavate, 23–29 × 2.5–3.5 µm Ellipsoid, 4.5–7 × (2.5–)3–4 µm Present study
Lyomyces wuliangshanensis Tuberculate/ cream to buff Thick-walled/ branched Capitate, 22–37 × 3–6 µm; tapering, 21–35 × 4–6.5 µm Clavate, 12–20 × 3–4.3 µm Ellipsoid, (3.3–)3.5–5.3(–5.5) × 2.8–4(–4.2) µm Chen and Zhao (2020)
Lyomyces wumengshanensis Grandinioid/ white to cream Thick-walled/ branched Capitate, 24.5–29 × 3–4 µm Subclavate to barreled, 11.5–14 × 5.5–6.5 µm Ellipsoid to broad ellipsoid, 4–6 × 3–5 µm Present study
Lyomyces yunnanensis Grandinioid/ cream to buff Thick-walled, frequently branched Tapering, 18–39 × 4–6 µm; capitate, 16–23.5 × 3–5 µm Clavate, 16.5–27 × 4–5.5 µm Ellipsoid, (4.5–)5–7 × 3–4.5 µm Guan et al. (2023)
Lyomyces zhaotongensis Grandinioid/ cream to buff Thick-walled/ branched Fusoid, 16–20.5 × 2.5–3.5 µm Clavate, 14–16.5 × 2.5–3.5 µm Broadly ellipsoid, 2.6–3.5 × 2.5–3 µm Present study

The Basidiomycota is a major phylum of the kingdom Fungi (He et al. 2019; Wijayawardene et al. 2020; Yuan et al. 2023; He et al. 2024), in which the wood-inhabiting fungi are an extensively studied group of Basidiomycota (Gilbertson and Ryvarden 1987; Bernicchia and Gorjón 2010; Núñez and Ryvarden 2001; Dai 2012; Ryvarden and Melo 2014; Wu et al. 2022b; Zhao et al. 2023; Dong et al. 2024), but the wood-inhabiting fungal diversity is still not well known in China, especially in subtropical and tropical areas, and many recently-described taxa of this ecologically important group were from China (Zhao et al. 2014; Zhao et al. 2015; Zhao et al. 2016; Bian et al. 2016; Ma and Zhao 2019; Guan et al. 2020; Huang and Zhao 2020; Guan et al. 2023; Ji et al. 2023; Liu et al. 2023; Yang et al. 2023; Deng et al. 2024a, b; Yang et al. 2024; Zhang et al. 2024; Zhou et al. 2024). Four new species in the present study are described, based on morphological and molecular phylogenetic analyses, also from the subtropics. This study enriches the wood-inhabiting fungal diversity in China and the world.

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 No. 32170004, U2102220), Forestry Innovation Programs of Southwest Forestry University (Grant No: LXXK-2023Z07), the High-level Talents Program of Yunnan Province (YNQR-QNRC-2018-111).

Author contributions

Conceptualization, C.Z.; methodology, C.Z. and Q.Y.; software, C.Z.; validation, C.Z.and Q.Y.; formal analysis, C.Z. and Q.Y.; investigation, C.Z., Q.Y., Y.D. Y.L., K.W. and Y.W.; resources, C.Z.; writing—original draft preparation, C.Z. and Q.Y.; writing—review and editing, C.Z. and Q.Y.; visualization, C.Z. and Q.Y.; supervision, C.Z.; project administration, C.Z.; funding acquisition, C.Z. All authors have read and agreed to the published version of the manuscript.

Author ORCIDs

Qi Yuan https://orcid.org/0000-0002-6732-1656

Yunchao Li https://orcid.org/0009-0007-3500-3761

Yunfei Dai https://orcid.org/0009-0007-7734-4142

Kunyan Wang https://orcid.org/0009-0004-1678-9963

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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