683 MycoKeys MycoKeys 110: 67-92 (2024) DOI: 10.3897/mycokeys.110.133108 Research Article Morphological and molecular identification for four new wood- inhabiting species of Lyomyces (Basidiomycota) from China Qi Yuan'?®, Yunchao Li2®, Yunfei Dai2®, Kunyan Wang”®, Yixuan Wang3, Changlin Zhao'2© 1 The Key Laboratory of Forest Resources Conservation and Utilization in the Southwest Mountains of China Ministry of Education, Key Laboratory of National Forestry and Grassland Administration on Biodiversity Conservation in Southwest China, Yunnan Provincial Key Laboratory for Conservation and Utilization of In-forest Resource, Southwest Forestry University, Kunming 650224, China 2 College of Forestry, Southwest Forestry University, Kunming 650224, China 3 Yunnan Yunzhihuang Health Technology Co., LTD, Kunming 650224, China Corresponding author: Changlin Zhao (fungi@swfu.edu.cn) OPEN Qaccess This article is part of: Exploring the Hidden Fungal Diversity: Biodiversity, Taxonomy, and Phylogeny of Saprobic Fungi Edited by Samantha C. Karunarathna, Danushka Sandaruwan Tennakoon, Ajay Kumar Gautam Academic editor: Samantha C. Karunarathna Received: 25 July 2024 Accepted: 15 October 2024 Published: 30 October 2024 Citation: Yuan Q, Li Y, Dai Y, Wang kK, 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 Copyright: © Qi Yuan et al. This is an open access article distributed under terms of the Creative Commons Attribution License (Attribution 4.0 International - CC BY 4.0). Abstract Fungi are one of the most diverse groups of organisms on Earth, in which the wood-in- habiting fungi play an important role in forest ecosystem processes and functions. Four new wooc-inhabiting fungi, Lyomyces hengduanensis, L. niveomarginatus, L. wumeng- shanensis 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 x 3-4.5 um). Lyomyces niveomarginatus is distinguished by the subceraceous basidiomata with crackled hymenial surface, a monomitic hyphal system and ellipsoid basidiospores (4.5-7 x 3-4 um). Lyomyces wumengshanensis is distinguished by the grandinioid hymenial surface, a monomitic hyphal system and ellipsoid to broad ellip- soid basidiospores (4-6 x 3-5 um). Lyomyces zhaotongensis is unique in the grandin- ioid hymenial surface, a monomitic hyphal system and broadly ellipsoid basidiospores measuring as 2.6-3.5 x 2.5-3 um. 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 phy- logram, 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. niveomargin- atus 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. hengduanen- sis 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 67 Qi Yuan et al.: Four new wood-inhabiting species of Lyomyces (Basidiomycota) from China Schizoporaceae Jilich includes many variations of the fruiting body types with- in 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 represen- tative wood-inhabiting fungal taxa, including diverse hymenophoral morpholo- gies 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-odon- tioid 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 Gor- jon 2010). Based on the MycoBank database (http://www.mycobank.org, ac- cessed 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 current- ly 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 molecu- lar systematics period (Guan et al. 2023; Zhang et al. 2024). The genus Hy- phodontia s.|. 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.|. was ac- cepted (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, Lagaroba- sidium clade, Kneiffiella-Alutaceodontia clade, Xylodon-Lyomyces-Rogersella clade and Xylodon-Schizopora-Palifer clade, were included, based on the phylo- genetical studies for Hyphodontia s.I., in which the genus Lyomyces was nested within the Xylodon-Lyomyces-Rogersella clade (Yurchenko and Wu 2013). The research revealed that Hyphodontia s.|. was divided into six genera, viz., Hast- odontia (Parmasto) Hjortstam & Ryvarden, Hyphodontia J. Erikss., Kneiffiella, Lagarobasidium Julich, 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 MycoKeys 110: 67-92 (2024), DOI: 10.3897/mycokeys.110.133108 68 Qi Yuan et al.: Four new wood-inhabiting species of Lyomyces (Basidiomycota) from China 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 fun- gal diversity of the family Schizoporaceae was analysed, in which six new spe- cies 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. yunna- nensis (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 col- lected from the Honghe, Lincang, Puer, Wenshan and Zhaotong of Yunnan Prov- ince, PR. 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, PR. China. Macromorphological descriptions were based on field notes and photos were captured in the field and lab. Colour terminology follows Petersen (Peters- en 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 aver- age 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, Chi- na) 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 denatur- ation 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 LROR 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 anda 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.nim.nih.gov/genbank/) (Table 1). MycoKeys 110: 67-92 (2024), DOI: 10.3897/mycokeys.110.133108 69 Qi Yuan et al.: Four new wood-inhabiting species of Lyomyces (Basidiomycota) from China Table 1. List of species, specimens, and GenBank accession numbers of sequences used in this study. Species name Fasciodontia brasiliensis F. bugellensis F. bugellensis F, yunnanensis F, yunnanensis Hymenochaete ochromarginata H. rubiginosa Lyomyces albopulverulentus mirlrlrlyer lel irl eye yelper eerie pe ye lr pepe lee yer le pepe lr ere eve lr pepe pe lr pepe le peepee lee le . allantosporus . allantosporus . bambusinus . bambusinus cremeus cremeus . crustosus . crustosus . densiusculus . elaeidicola . elaeidicola erastii erastii . fimbriatus . fimbriatus . fissuratus . fissuratus fumosus . gatesiae . gatesiae . griseliniae . hengduanensis . hengduanensis . hengduanensis . hengduanensis . hengduanensis . juniperi . leptocystidiatus . leptocystidiatus . Macrosporus . Mascarensis . Mascarensis . microfasciculatus . niveomarginatus niveus niveus . ochraceoalbus . ochraceoalbus . ochraceoalbus . orientalis pruni pruni . sambuci . sambuci MycoKeys 110: 67-92 (2024), DOI Specimen No. GenBank accession No. ~GenBankaccessionNo. i ee ee kws-oetaoas ——kvao0aor | rrozisseo = Kvoooss7 | eizheo tes noaszes | etzhao see near | ~clzmaoats# So Mnoasora etzheo e295 nausea tasmyeeas rasa | uecrnazeet | krenato | ~Roverden asi 0x73059 | ~wweaereosr-20rsnowse | ~tweaorenariagrsnoas7 | TASM:YG 022 MF382992 = aaesawve —xas7e00 | —wwot06207 | Ksrsa00 SS a cuznaotas2 wise | euzhaoann wri | euzhaoetas | mwrimaa weamreosisa wrsnoma? | weameosss2 | rsigue | kt 12971 66) oera6s1 —eizaoaaris | —oneeoisa —ciznaosama | oresnsa | Fraasi0gs | kvoer799 ~wezerroeies rszssia | weaorreee2 ress | eazhaoasie | MNOASS7T | kas-cetaans | kvoose | rasceusooe | kveoow00 | euzhaostas nse sero vanoaos = ease | asaoms ceizser | amosre | —ibergoztara (68) | D0s7se28 esr 83SAMHYP : 10.3897/mycokeys.110.133108 References Yurchenko et al. (2020) Yurchenko et al. (2020) Yurchenko et al. (2020) Luo and Zhao (2021) Luo and Zhao (2021) Unpublished He and Li (2013) Guan et al. (2023) Yurchenko et al. (2017) Yurchenko et al. (2017) Chen and Zhao (2020) Chen and Zhao (2020) Chen and Zhao (2020) Chen and Zhao (2020) Gafforov et al. (2017) Unpublished Viner et al. (2022) Wang et al. (2021) Wang et al. (2021) Gafforov et al. (2017) Unpublished Yurchenko et al. (2020) Yurchenko et al. (2020) Luo et al. (2021b) Luo et al. (2021b) Luo et al. (2021b) Wang et al. (2021) Wang et al. (2021) Larsson et al. (2006) Present study Present study Present study Present study Present study Riebesehl and Langer (2017) Wang et al. (2021) Wang et al. (2021) Chen and Zhao (2020) Yurchenko et al. (2020) Yurchenko et al. (2020) Chen and Zhao (2020) Present study Luo et al. (2021b) Luo et al. (2021b) Luo et al. (2021b) Luo et al. (2021b) Yurchenko et al. (2017) Yurchenko et al. (2017) Larsson et al. (2006) Larsson et al. (2006) Yurchenko et al. (2017) Yurchenko et al. (2017) 70 Qi Yuan et al.: Four new wood-inhabiting species of Lyomyces (Basidiomycota) from China Species name . vVietnamensis . wuliangshanensis . wuliangshanensis . wumengshanensis . wumengshanensis . wumengshanensis . wumengshanensis . wumengshanensis mirjelyrier leper lel . wumengshanensis L. wumengshanensis L. yunnanensis L. yunnanensis L. yunnanensis L. zhaotongensis Xylodon afromontanus X. asiaticus X. cystidiatus X. daweishanensis X. daweishanensis X. filicinus X. fissuratus X. fissuratus X. hastifer X. hyphodontinus X. macrosporus X. puerensis X. puerensis X. quercinus X. ramicida X. subflaviporus X. tropicus X. wenshanensis X. xinpingensis GenBank accession No. Specimen No. References ITS nLSU TNM F9073 JX175044 - Yurchenko et al. (2017) CLZhao 4108 MN945980 = Chen and Zhao (2020) CLZhao 4167 MN945979 7 Chen and Zhao (2020) CLZhao 29374 OR803021 PP657613 Present study CLZhao 31486 OR899208 - Present study CLZhao 32705 OR899209 oe Present study CLZhao 32736 OR899210 = Present study CLZhao 32800 OR899211 PP657614 Present study CLZhao 32869 OR899212 = Present study CLZhao 32915 OR899213 PP657615 Present study CLZhao 2463 OP730711 OP730723 Guan et al. (2023) CLZhao 9375 OP730710 = Guan et al. (2023) CLZhao 10041 OP730709 = Guan et al. (2023) CLZhao 32878 PP537950 PP657609 Present study H 7006811 0Q645463 = Yurchenko et al. (2024) CLZhao 10368 OM959479 ; Zhang et al. (2024) FR-0249200 MH880195 MH884896 Riebesehl et al. (2019) CLZhao 18492 OP730719 OP730727 Guan et al. (2023) CLZhao 18446 OP730717 OP730725 Guan et al. (2023) MSK-F 12869 MH880199 NG067836 Riebesehl et al. (2019) CLZhao 7007 OP730713 = Guan et al. (2023) CLZhao 9407 OP730714 = Guan et al. (2023) K(M) 172400 NR166558 = Riebesehl and Langer (2017) KAS-GEL9222 MH880205 MH884903 Riebesehl et al. (2019) CLZhao 10226 MZ663809 MZ663817 Luo et al. (2021a) CLZhao 8142 OP730720 OP730728 Guan et al. (2023) CLZhao 8639 OP730721 OP730729 Guan et al. (2023) Larsson 11076 (GB) KT361633 = Larsson et al. (2004) Spirin 7664 NR138013 rs Unpublished Wu 0809-76 KX857803 = Chen et al. (2017) CLZhao 3351 0L619261 0L619269 Qu et al. (2022) CLZhao 15729 OM338097 OM338104 Luo et al. (2022) CLZhao 11224 MW394662 MW394654 Luo et al. (2022) The sequences were aligned in MAFFT version 7 (Katoh et al. 2019) us- ing 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., re- trieved 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 Xy- lodon 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). Maximum Parsimony (MP), Maximum Likelihood (ML) and Bayesian Infer- ence (BI) analyses were applied to the combined three datasets following a pre- vious 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 parsi- monious trees were saved. Clade robustness was assessed using bootstrap MycoKeys 110: 67-92 (2024), DOI: 10.3897/mycokeys.110.133108 71 Qi Yuan et al.: Four new wood-inhabiting species of Lyomyces (Basidiomycota) from China Xylodon asiaticus C 70/-/- 100/100/1.00 79/ - 10.99 Xylodon puerensis CL Xylodon subflaviporus \ - /52/1.00} - /100/- Xylodon fissuratus CL 99/100/1.00| Xylodon fissuratus” 99/92/1,00 | 85/58/1.00 Xylodon d Xylodon da -Xylodon hyphe Xylodon ous 1100/100/1.00 100/100/1.00 Xylodon macrosporus CLZhao sonoori.oo| Lvomyees hengduan g/100/1.00] 'Lyomyces hengduan a 93/97/1.00 Fasciodontia yunnanensis CLZhao | °) Hone Fasciodontia yunnanensis CLZhao Loo1do/iloo f Hasciodontia bugellensis KAS-FD © ~ 71007 -| Fasciodontia bugellensis MSK-F 7353 Fasciodontia brasiliensis MSK-F 7245a Hymenochaete rubiginosa He 458 i Hymenochaete ochromarginata | t—Y 50 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. (BT) analysis with 1,000 pseudo-replicates (Felsenstein 1985). Descriptive tree statistics - tree length (TL), composite consistency index (Cl), composite re- tention 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 Likeli- hood (ML) in RAxML-HPC2 through the CIPRES Science Gateway (Miller et al. 2012). Branch support (BS) for the ML analysis was determined by 1000 boot- strap pseudo-replicates. MycoKeys 110: 67-92 (2024), DOI: 10.3897/mycokeys.110.133108 72 Qi Yuan et al.: Four new wood-inhabiting species of Lyomyces (Basidiomycota) from China se H ] Loo/1oo/1.00Lvomyces leptocystidiatus LWZ20170818-2 KK y & y otyp € 'Lyomyces leptocystidiatus LWZ20170818-1 Lyomyces griseliniae KHL 12971 (GB) km v ® * Pruinose Loo/100/1.09 Lvontces fimbriatus Wu910620-7 * vy © Smooth 100/100/1.00] 'Lyomyces fimbriatus Wu911204-4 * 100/100/1.00 Lyomyces fumosus CLZhao 8188 * y ® * Tuberculate Lyomyces densiusculus Ryvarden 44818 oe kk y = Odontioid Lyomyces allantosporus KAS-GEL4933 * 100/100/1.007, vomyces allantosporus FR-0249548 % * . aa * Grandinioid Lyomyces macrosporus CLZhao 4516 e . Lyomyces wumengshanensis CLZha * Reticulate 100/100/1.00 Lyomiycés witnenashoneaen ae * Warted iLyomyces wumengshanensis CLZI 100/100/1.00-Lvemyces bambusinus CLZhao 480 * Crackled eeonnod Lyomyces bambusinus CLZhao 4831 xe * 100/1bo/1.00)Lvomyces sambuci KAS-JR7 * * Lyomyces sambuci 83SAMHYP. Lyomyces orientalis GEL3376 ¥ Ellipsoid Lyomyces albopulverulentus CLZhao 21478 be 4 ic 100/100/1.00 ~Lvomyces elaeidicola LWZ20180411-19 ¥ Globose Lyomyces elaeidicola LWZ20180411-20 Ff Kk *& Lyomyces erastii TASM:YG 022 © Ovoid 100/10071-00L_ 7 yomyces erastii 23¢SAMHYP kK . : Loo/100/1.00r Lvomyces wuliangshanensis CLZhao 4108 y Cylindrical Lyomyces wuliangshanensis CLZhao 4167 xe Lyomyces mascarensis KAS-GEL4833 % 109/100/1.001 Tyomyces mascarensis KAS-GEL4908 Lyomyces gatesiae LWZ20180515-3 xB k* FJ 28 ® Allantoid € ® Thin-walled 100/100/1001L yomyces gatesiae LWZ20180515-32 sdioaioor Lvonuces cremeus CLZhao 8295 & Thick-walled Lyomyces cremeus CLZhao 4138 € ors 87 3 49 8 FeO * Lyomyces fissuratus CLZhao 4291 * & Slightly thick-walled 100/100/1.00L-Lyomyces fissuratus CLZhao 4352 Lyomyces microfasciculatus CLZhao 5109 > Lyomyces organensis MSK7247 oe * 100/100/1.007-——Lvomyces pruni GEL2327 Fs * Lyomyces pruni Ryberg 021018 (GB) Lyomyces yunnanensis CLZhao 2463 Lyomyces yunnanensis CLZhao 9375 Lyomyces yunnanensis CLZhao 10041 % ¥ % €€¢e€€¢e€ € €E € € EE EE 100/100/1.00) * -Lyomyces niveus CLZhao 6442 *k 100/100/1.00Lyomyces niveus CLZhao 6431 97/100/0.98 PLyomyces ochraceoalbus CLZhao 4725 * 89/68) - Lyomyces ochraceoalbus CLZhao 4385 Lyomyces crustosus TASM:YG G39 * 97/9970.99L Lyomyces crustosus UC2022841 @€¢¢€¢ o Oo + =a Lyomyces zhaotongensis CLZhao 32878 Lyomyces vietnamensis TNM F9073 Lyomyces niveomarginatus CLZhao 16360 Lyomyces juniperi FR-0261086 71/63/0.96 ro) -Xylodon quercinus Larsson 11076 (GB) Cd ie \\ -Xylodon ramicida Spirin 7664 > : e ye p> 2) oF oe Figure 2. Maximum parsimony strict consensus tree illustrating the phylogeny of the four new species and related spe- cies 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. MrModelTest 2.3 (Nylander 2004) was used to determine the best-fit evolu- tion 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 mil- lion 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 sup- port value (BS) of => 70%, a Maximum Parsimony bootstrap support value (BT) of => 70% or a Bayesian Posterior Probability (BPP) of = 0.95. MycoKeys 110: 67-92 (2024), DOI: 10.3897/mycokeys.110.133108 73 Qi Yuan et al.: Four new wood-inhabiting species of Lyomyces (Basidiomycota) from China 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-unin- formative and 560 were parsimony-informative. Maximum parsimony analysis yielded one equally parsimonious tree (TL = 2,513, Cl = 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-un- informative and 385 were parsimony-informative. Maximum parsimony analy- sis yielded 80 equally parsimonious tree (TL = 1,748, Cl = 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 wumengshan- ensis was retrieved as a sister to L. macrosporus C.L. Zhao. Moreover, Lyomy- ces 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, Fengging 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). Etymology. Hengduanensis (Lat.) refers to the type locality “Hengduan Mountain’. MycoKeys 110: 67-92 (2024), DOI: 10.3897/mycokeys.110.133108 74 Qi Yuan et al.: Four new wood-inhabiting species of Lyomyces (Basidiomycota) from China Figure 3. Basidiomata of Lyomyces hengduanensis (holotype). Scale bars: 1 cm (A); 2 mm (B). MycoKeys 110: 67-92 (2024), DOI: 10.3897/mycokeys.110.133108 75 Qi Yuan et al.: Four new wood-inhabiting species of Lyomyces (Basidiomycota) from China fd Oot A 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 um (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 um 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, co- lourless, thick-walled, branched, 2-3 um in diameter; IKI-, CB-, tissues un- changed 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 x 3-4 um; (2) sub- clavate, colourless, thin-walled, smooth, slightly constricted in the middle to somewhat sinuous, 16-23 x 3-4.5 um; basidia clavate, with 4 sterigmata and a basal clamp connection, 10.5-14 x 3.5-5 um. MycoKeys 110: 67-92 (2024), DOI: 10.3897/mycokeys.110.133108 76 Qi Yuan et al.: Four new wood-inhabiting species of Lyomyces (Basidiomycota) from China Basidiospores ellipsoid, colourless, thin-walled, smooth, with one oil drop, CB-, IKI-, 3.5-6 x 3-4.5 um, L= 4.63 um, W = 3.65 um, 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°92E, 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 Na- tional Nature Reserve, GPS coordinates 23°21'N, 104°10E, altitude 1950 m, on a fallen branch of angiosperm, leg. C.L. Zhao, 26 July 2019, CLZhao 16360 (SWFC). Etymology. Niveomarginatus (Lat.) refers to the niveous margin of basidi- omata. Description. Basidiomata annual, resupinate, adnate, subceraceous, without odour and taste when fresh and up to 7.5 cm long, 2 cm wide, 150 um 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, co- lourless, thin-walled, branched, 1.5-3.5 um in diameter; IKI-, CB-, tissues un- changed in KOH. Numerous crystals present amongst generative hyphae. Cystidia of two types: (1) fusoid, colourless, thin-walled, smooth, 25-29 x 2-3 um; (2) clavate, colourless, thin-walled, smooth, 20-25.5 x 4.5-5.5 um; basidia sub- clavate, with 4 sterigmata and a basal clamp connection, 23-29 x 2.5-3.5 um. Basidiospores ellipsoid, colourless, thin-walled, smooth, with one oil drop, CB-, IKI-, 4.5-7 x (2.5-)3—-4 pm, L = 5.51 um, W = 3.15 um, Q = 1.75 (n = 30/1). Additional specimens examined (paratypes). CHINA * Yunnan Province, Wen- shan, 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 Coun- ty, 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). Etymology. Wumengshanensis (Lat.) refers to the type locality “Wumeng Mountain”. MycoKeys 110: 67-92 (2024), DOI: 10.3897/mycokeys.110.133108 77 Qi Yuan et al.: Four new wood-inhabiting species of Lyomyces (Basidiomycota) from China sal sf waa yf ee , ee ie —— : : (poe aha LAR FS es. Sey asioramels o3 MycoKeys 110: 67-92 (2024), DOI: 10.3897/mycokeys.110.133108 78 Qi Yuan et al.: Four new wood-inhabiting species of Lyomyces (Basidiomycota) from China E 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 um (A-F). 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 um 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, co- lourless, thick-walled, branched, 3-4 um in diameter; IKI-, CB-, tissues un- changed in KOH. Numerous crystals present amongst generative hyphae. Cystidia capitate, colourless, thin-walled, smooth, 24.5-29 x 3-4 um; basid- ia subclavate to barrelled, colourless, with 4 sterigmata and a basal clamp con- nection, 11.5-14 x 5.5-6.5 um. Basidiospores ellipsoid to broad ellipsoid, colourless, thin-walled, smooth, with one oil drop, CB-, IKI-, 4-6 x 3-5 um, L = 5.4 um, W = 4.2 um, Q = 1.28- 1.32 (n = 120/4). Additional specimen examined (paratype). CHINA * Yunnan Province, Zha- otong, Wumeng Mountain National Nature Reserve, 27°72'N, 103°92'E, altitude 1424 m, ona fallen branch of angiosperm, leg. C.L. Zhao, 29 August 2023, CLZ- MycoKeys 110: 67-92 (2024), DOI: 10.3897/mycokeys.110.133108 79 Qi Yuan et al.: Four new wood-inhabiting species of Lyomyces (Basidiomycota) from China + * ae ai ae Figure 7. Basidiomata of Lyomyces wumengshanensis (holotype). Scale bars: 1 cm (A); 2 mm (B). MycoKeys 110: 67-92 (2024), DOI: 10.3897/mycokeys.110.133108 80 Qi Yuan et al.: Four new wood-inhabiting species of Lyomyces (Basidiomycota) from China ABE lee se = 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 um (A-E). hao 31486, CLZhao 32705, CLZhao 32736, CLZhao 32800, CLZhao 32869, CLZ- hao 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 Moun- tain National Nature Reserve, GPS coordinates 27°77'N, 104°29’'E, altitude MycoKeys 110: 67-92 (2024), DOI: 10.3897/mycokeys.110.133108 81 Qi Yuan et al.: Four new wood-inhabiting species of Lyomyces (Basidiomycota) from China Figure 9. Basidiomata of Lyomyces zhaotongensis (holotype). Scale bars: (A) 1 cm; (B) 2 mm. 82 MycoKeys 110: 67-92 (2024), DOI: 10.3897/mycokeys.110.133108 Qi Yuan et al.: Four new wood-inhabiting species of Lyomyces (Basidiomycota) from China A 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 um (A-E). 2900 m, on the fallen branch of angiosperm, leg. C.L. Zhao, 29 August 2023, CLZhao 32878 (SWFC). Etymology. Zhaotongensis (Lat.) refers to the type locality “Zhaotong”. 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, co- lourless, thick-walled, branched, 1.5-2 um in diameter; IKI-, CB-, tissues un- changed in KOH. Numerous crystals present amongst generative hyphae. Cystidia fusoid, colourless, thin-walled, smooth, 16-20.5 x 2.5-3.5 um. Basidia clavate, with 4 sterigmata and a basal clamp connection, 14-16.5 x 2.5-3.5 um. Basidiospores broadly ellipsoid, colourless, thin-walled, smooth, with oil drops, CB-, IKI-, 2.6-3.5 x 2.5-3 um, L = 2.99 um, W = 2.75 um, Q = 1.08 (n = 30/1). Additional specimen examined (paratype). CHINA * Yunnan Province, Zha- otong, 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). MycoKeys 110: 67-92 (2024), DOI: 10.3897/mycokeys.110.133108 83 Qi Yuan et al.: Four new wood-inhabiting species of Lyomyces (Basidiomycota) from China Discussion Many recently new wood-inhabiting fungal taxa have been reported in the sub- tropics 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. lep- tocystidiatus 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) Rie- besehl & 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 morpho- logical 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 Hymenochae- tales (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. zha- otongensis and then closely grouped with L. crustosus, L. ochraceoalbus and L. vietnamensis. Lyomyces niveomarginatus was retrieved as a sister to L. juni- peri. L. wumengshanensis was sister to L. macrosporus. Moreover, L. zhaoton- gensis grouped with L. hengduanensis and then closely clustered with three species: L. crustosus, L. ochraceoalbus and L. vietnamensis. However, mor- phologically, L. zhaotongensis can be delimited from L. hengduanensis by its the grandinioid hymenial surface and longer basidia (14-16.5 x 2.5-3.5 ym); L. crustosus can be separated from L. hengduanensis by its odontioid hyme- nial surface and narrow basidiospores (5—-7.5 x 2.5-3 um) (Lentz and McKay 1976); L. ochraceoalbus differs in L. hengduanensis by having a smooth hyme- nial surface and lacking a cystidium (Luo et al. 2021c); L. vietnamensis differs from L. hengduanensis by its aculeate hymenial surface and narrow basidio- spores (5.8-6.1 x 2.6-2.9 um; 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 x 4-4.5 um; Hjortstam and Ryvarden (2004)); L. macrosporus can be separated from L. wumengshanensis by its reticulate hymenial surface and longer basidiospores (6.7—8.9 x 4.4—5.4 um; Chen and Zhao (2020)); L. crustosus can be delimited from L. zhaoton- gensis by its odontioid hymenial surface and longer basidiospores (5-7.5 x 2.5-3 ym; Lentz and McKay (1976)); L. hengduanensis can be delimited from L. zhaotongensis by its pruinose hymenial surface and shorter basidia (14- 16.5 x 2.5-3.5 um); L. ochraceoalbus differs in L. zhaotongensis by having smooth hymenial surface and longer basidiospores (4-5 x 2.5-3.5 um; Luo et al. (2021c)); L. vietnamensis can be delimited from L. zhaotongensis by its MycoKeys 110: 67-92 (2024), DOI: 10.3897/mycokeys.110.133108 84 Qi Yuan et al.: Four new wood-inhabiting species of Lyomyces (Basidiomycota) from China aculeate hymenial surface and longer basidiospores (5.8-6.1 x 2.6-2.9 um; Yurchenko and Wu (2013)). Morphologically, Lyomyces hengduanensis resembles four taxa viz. L. albopul- verulentus, L. bambusinus, L. mascarensis Riebesehl, Yurch. & Langer and L. yun- nanensis, by the similar ellipsoid basidiospores. However, L. albopulverulentus differs from L. hengduanensis by its larger basidia (24.5—28.5 x 7-9 um) and ba- sidiospores (8-10.5 x 5.5—7 um; Guan et al. (2023)); L. bambusinus can be sep- arated from L. hengduanensis by its colliculose to tuberculate hymenial surface and longer basidia (16.5-35 x 3.5-7 um; Chen and Zhao (2020)); L. mascaren- sis is distinct from L. hengduanensis by having indistinctly colliculose hymenial surface and longer basidia (16-17.5 x 3.5-4.5 tm; Yurchenko et al. (2017)); L. yunnanensis is distinguished from L. hengduanensis by its grandinioid hymenial surface and longer basidia (16.5-27 x 4—5.5 yum; 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 basidio- spores. However, L. albopulverulentus differs from L. niveomarginatus by its pruinose hymenial surface and wider basidia (24.5-28.5 x 7—9 um; Guan et al. (2023)); L. cremeus can be separated from L. niveomarginatus by its smooth hy- menial surface and shorter basidia (9-18.5 x 3-6 um; Chen and Zhao (2020)); L. macrosporus differs from L. niveomarginatus by its reticulate hymenial sur- face and wider basidia (23-29 x 2.5-3.5 um) and wider basidiospores (6.7—8.9 x 4.4-5.4 ym; 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 x 3—4.3 um; Chen and Zhao (2020)); L. yun- nanensis is distinct from L. niveomarginatus by having grandinioid hymenial surface and wider basidia (16.5-27 x 4—5.5 um; Guan et al. (2023)). Morphologically, Lyomyces wumengshanensis resembles L. bambusinus, L. cremeus, L. fumosus, L. fissuratus, L. wuliangshanensis and L. yunnanen- sis by having the capitate cystidia. However, L. bambusinus is distinct from L. wumengshanensis by possessing tapering cystidia (40-65 x 4—5.5 um) and longer basidia (16.5-35 x 3.5-7 um; Chen and Zhao (2020)); L. cremeus differs from L. wumengshanensis by its smooth hymenial surface and possesses taper- ing cystidia (18-35 x 3—4.5 um; Chen and Zhao (2020)); L. fumosus can be sep- arated from L. wumengshanensis by its smooth, smoky grey hymenial surface and narrower basidia (11.5-17.5 x 3-5 um; Luo et al. (2021b)); L. fissuratus can be delimited from L. wumengshanensis by its longer and narrower basidia (14.7- 23.3 x 2.9-4.8 tm; Luo et al. (2021b)); L. wuliangshanensis differs from L. wu- mengshanensis by its smooth to more or less tuberculate hymenial surface and narrower basidia (12-20 x 3-4.3 um; Chen and Zhao (2020)); L. yunnanensis is separated from L. wumengshanensis by the longer basidia (16.5-27 x 4—5.5 um) and possessing fusiform cystidia (18-39 x 4-6 um; Guan et al. (2023)). Morphologically, Lyomyces zhaotongensis reminds L. albopulverulentus, L. cremeus, L. denudatus Viner, L. macrosporus and L. wuliangshanensis by hav- ing the ellipsoid basidiospores. However, L. albopulverulentus can be separat- ed from L. zhaotongensis by its pruinose hymenial surface and larger basidia (24.5-28.5 x 7-9 um) and larger basidiospores (8-10.5 x 5.5-7 um; Guan et al. (2023)); L. cremeus is distinct from L. zhaotongensis by its smooth hyme- nial surface and larger basidiospores (4.5—5.6 x 3.3-4.3 um; Chen and Zhao MycoKeys 110: 67-92 (2024), DOI: 10.3897/mycokeys.110.133108 85 Qi Yuan et al.: Four new wood-inhabiting species of Lyomyces (Basidiomycota) from China (2020)); L. denudatus is separated from L. zhaotongensis by the smooth hyme- nial surface and longer basidiospores (4.8—7 x 2.8—4.2 um; Viner and Miettin- en (2022)); L. macrosporus differs from L. zhaotongensis due to its reticulate hymenial surface and larger basidia (22.2-38 x 4.5-7 um) and larger basid- iospores (6.7-8.9 x 4.4—5.4 um; Chen and Zhao (2020)); L. wuliangshanensis can be delimited from L. zhaotongensis by its smooth to more or less tuber- culate hymenial surface and longer basidiospores (3.5—5.3 x 2.8-4 um; Chen and Zhao (2020)). A morphological comparison amongst four new Lyomyces species and seven similar species are presented in Table 2. 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 Gorjon 2010; Nufiez and Ry- varden 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 recent- ly-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 Table 2. A morphological comparison between four new Lyomyces species and seven similar species in the genus Lyomyces. The bold are new taxa. Species name Lyomyces albopulverulentus Lyomyces bambusinus Lyomyces cremeus Lyomyces denudatus Lyomyces hengduanensis Lyomyces mascarensis Lyomyces niveomarginatus Lyomyces wuliangshanensis Lyomyces wumengshanensis Lyomyces yunnanensis Lyomyces zhaotongensis Hymenial surface Pruinose/ white Colliculose to tuberculate/ cream to buff Smooth/ pale cream Smooth/ cream Pruinose/ cream to slightly buff Smooth / cream or brownish Smooth / cream to slightly buff Tuberculate/ cream to buff Grandinioid/ white to cream Grandinioid/ cream to buff Grandinioid/ cream to buff Generative Cystidia Basidia Basidiospores References hyphae Thick-walled/ | Capitate,37-54x5-9 | Clavate, 24.5-28.5 x | Ellipsoid, (7.5-)8-10.5(- Guan et al. frequently um 7-9 um 11) x (5-)5.5-7 um (2023) branched Thick-walled/ | Capitate, 35-55 x 4-7 Clavate, 16.5-35 x Broadly ellipsoid, Chen and Zhao branched um; tapering, 40-65 x 3.5-7 um (4.5—-)4.7-5.9 (-6.2) x (2020) 4-5.5 um, cystidioles, (3.4-)3.7-4.6(-4.8) um 12-17 x 2-3 um Thick-walled/ | Capitate, 20-40 x 3-5 Clavate,9-18.5x | Ellipsoid, 4.5-5.6(-5.8) x | Chen and Zhao branched um; tapering, 18-35 x 3-6 um 3.3-4.3(-4.5) um (2020) 3-4.5 um Thin-walled | Capitate, (21-)34.9-62 x Suburniform, Ellipsoid, (4.1-)4.8-7 x Viner and to slightly (3.5=)4-5.5(=7) mi 15-21.1(-25) x 2.8-4.2(—4.7) um Miettinen (2022) thick-walled 3.8-5.5 um Thick-walled/ | Fusoid,17.5-25x3-4 | Clavate, 10.5-14 x Ellipsoid, 3.5—6 x Present study branched um; subclavate, 16-23 x 3.5-5 um 3-4.5 um 3-4.5 um Thin-walled Capitate, 17-38 Subcylindrical with Ellipsoid or broadly Yurchenko et al. x 3.5-6(-7) um; one constriction, ellipsoid, (4—)4.5-6 x (2017) submoniliform, 18-22 16-17.5(-19) x (3-)3.3-4 um x 5-5.5 um; tapering, 3.5-4.5(-6) um 25-30 x 3.5-4.5 um Thin-walled, Fusoid, 25-29 x 2-3 | Subclavate, 23-29 x Ellipsoid, 4.5—-7 x Present study branched um; clavate, 20-25.5 x 2.5-3.5 um (2.5-)3-4 um 4.5-5.5 um Thick-walled/ | Capitate, 22-37 x 3-6 Clavate, 12-20 | Ellipsoid, (3.3—)3.5-5.3(— | Chen and Zhao branched um; tapering, 21-35 x 3-4.3 um 5.5) x 2.8-4(-4.2) um (2020) 4-6.5 um Thick-walled/ Capitate, 24.5-29 x Subclavate to Ellipsoid to broad Present study branched 3-4 um barreled, 11.5-14x | ellipsoid, 4-6 x 3-5 um 5:5-6.5 lm Thick-walled, | Tapering, 18-39 4-6 | Clavate, 16.5-27 x Ellipsoid, (4.5-)5-7 x Guan et al. frequently um; capitate, 16-23.5 x 4-5.5 um 3-4.5 um (2023) branched 3-5 um Thick-walled/ Fusoid, 16-20.5 x Clavate, 14-16.5 x | Broadly ellipsoid, 2.6-3.5 | Present study branched 2.5-3.5 um 2.5-3.5 um x 2.5-3 um MycoKeys 110: 67-92 (2024), DOI: 10.3897/mycokeys.110.133108 86 Qi Yuan et al.: Four new wood-inhabiting species of Lyomyces (Basidiomycota) from China 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 (Proj- ect No. 32170004, U2102220), Forestry Innovation Programs of Southwest Forestry Uni- versity (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.; re- sources, C.Z.; writing—original draft preparation, C.Z. and Q.Y.; writing—review and edit- ing, 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/0009-0007-7 734-4142 Kunyan Wang ® hitps://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. References Bernicchia A, Gorjon SP (2010) Fungi Europaei 12: Corticiaceae s.|. Edizioni Candusso, Alassio, Italy, 1008 pp. Bian LS, Wu F, Dai YC (2016) Two new species of Coltricia (Hymenochaetaceae, Ba- sidiomycota) from Southern China based on evidence from morphology and DNA sequence data. 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