Five new species of Schizoporaceae (Basidiomycota, Hymenochaetales) from East Asia

Qian-Xin Guan; Jing Huang; Jian Huang; Chang-Lin Zhao

doi:10.3897/mycokeys.96.99327

Research Article

Five new species of Schizoporaceae (Basidiomycota, Hymenochaetales) from East Asia

Qian-Xin Guan^‡, Jing Huang^‡, Jian Huang^§, Chang-Lin Zhao^‡|‡

‡ Southwest Forestry University, Kunming, China

§ Yunnan General Administration of Forestry Seeds and Seedlings, Kunming, China

| Kunming Institute of Botany, Chinese Academy of Science, Kunming, China

Corresponding author: Chang-Lin Zhao ( fungichanglinz@163.com )

Academic editor: Jennifer Luangsa-ard

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: Guan Q-X, Huang J, Huang J, Zhao C-L (2023) Five new species of Schizoporaceae (Basidiomycota, Hymenochaetales) from East Asia. MycoKeys 96: 25-56. https://doi.org/10.3897/mycokeys.96.99327

Abstract

Five new wood-inhabiting fungi, Lyomyces albopulverulentus, L. yunnanensis, Xylodon daweishanensis, X. fissuratus, and X. puerensis spp. nov., are proposed based on a combination of morphological features and molecular evidence. Lyomyces albopulverulentus is characterized by brittle basidiomata, pruinose hymenophore with a white hymenial surface, a monomitic hyphal system with clamped generative hyphae, and ellipsoid basidiospores. Lyomyces yunnanensis is characterized by a grandinioid hymenial surface, the presence of capitate cystidia, and ellipsoid basidiospores. Xylodon daweishanensis is characterized by an odontioid hymenial surface, a monomitic hyphal system with clamped generative hyphae, and broad ellipsoid-to-subglobose basidiospores. Xylodon fissuratus is characterized by a cracking basidiomata with a grandinioid hymenial surface, and ellipsoid basidiospores. Xylodon puerensis is characterized by a poroid hymenophore with an angular or slightly daedaleoid configuration, and ellipsoid-to-broad-ellipsoid basidiospores. Sequences of ITS and nLSU rRNA markers of the studied samples were generated and phylogenetic analyses were performed with the maximum likelihood, maximum parsimony, and Bayesian inference methods. The phylogram based on the ITS+nLSU rDNA gene regions (Fig. 1) included six genera within the families Chaetoporellaceae, Hyphodontiaceae, Hymenochaetaceae, and Schizoporaceae (Hymenochaetales)—Fasciodontia, Hastodontia, Hyphodontia, Kneifiella, Lyomyces, and Xylodon—in which the five new species were grouped into genera Lyomyces and Xylodon. The phylogenetic tree inferred from the ITS sequences highlighted that Lyomyces albopulverulentus formed a monophyletic lineage and was then grouped closely with L. bambusinus, L. orientalis, and L. sambuci; additionally, L. yunnanensis was sister to L. niveus with strong supports. The topology, based on the ITS sequences, revealed that Xylodon daweishanensis was retrieved as a sister to X. hyphodontinus; X. fissuratus was grouped with the four taxa X. montanus, X. subclavatus, X. wenshanensis, and X. xinpingensis; and X. puerensis was clustered with X. flaviporus, X. ovisporus, X. subflaviporus, X. subtropicus, and X. taiwanianus.

Keywords

Biodiversity, China, molecular systematics, taxonomy, wood-inhabiting fungi, Yunnan Province

Introduction

Fungi represent one of the most diverse groups of organisms on the earth, with an indispensable role in the processes and functioning of ecosystems (Hyde 2022). The family Schizoporaceae Jülich includes many variations of the fruiting body types among the order Hymenochaetales Oberw. (Larsson et al. 2006; Wu et al. 2022a), in which it comprises many representative wood-inhabiting fungal taxa, including hydnoid, corticioid, and polyporoid basidiomes with diverse hymenophoral and cystidial morphology (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, b). In addition, the species of Schizoporaceae have been described from different countries, and they cause a white rot (Langer 1994).

The genus Lyomyces P. Karst. is a small corticioid group, typified by L. sambuci (Pers.) P. Karst. 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). The species of Lyomyces are found on fallen angiosperm branches, dead woody or herbaceous stems, or, occasionally, on gymnosperm wood (Yurchenko et al. 2017). Approximately 40 species of Lyomyces are currently known (Rabenhorst 1851; Karsten 1881, 1882; Cunningham 1959, 1963; Wu 1990; Hjortstam and Ryvarden 2009; Xiong et al. 2009; Dai 2010, 2011; Yurchenko et al. 2013, 2017, 2020; Gafforov et al. 2017; Riebesehl and Langer 2017; Chen and Zhao 2020; Luo et al. 2021b, c; Viner et al. 2022). The genus Xylodon (Pers.) Gray is typiﬁed by X. quercinus (Pers.) Gray (Bernicchia and Gorjón 2010). The taxa of this genus grow on rotten gymnosperm or angiosperm trunks and stumps, bamboo and ferns (Girometta et al. 2020; Greslebin and Rajchenberg 2000; Kotiranta and Saarenoksa 2000). They are characterized by resupinate or effuse basidiomata with a smooth, tuberculate, grandinioid, odontioid, coralloid, irpicoid, or poroid hymenophore; a monomitic or dimitic hyphal system with clamped generative hyphae; the presence of different types of cystidia, i.e., utriform or suburniform basidia; and cylindrical-to-ellipsoid-to-globose basidiospores, which can cause a white rot (Gray 1821; Bernicchia and Gorjón 2010). Based on the MycoBank database (http://www.mycobank.org, accessed on 24 December 2022) and the Index Fungorum (http://www.indexfungorum.org, accessed on 24 December 2022), Xylodon has registered 221 speciﬁc and infraspeciﬁc names, and the actual number of the species has reached 100 species (Chevallier 1826; Kuntze 1898; Wu 1990, 2000, 2001, 2006; Hjortstam and Ryvarden 2007, 2009; Xiong et al. 2009, 2010; Bernicchia and Gorjón 2010; Tura et al. 2011; Dai 2012; Lee and Langer 2012; Yurchenko et al. 2013; Yurchenko and Wu 2014; Zhao et al. 2014; Chen et al. 2016; Kan et al. 2017a, b; Riehl and Langer 2017; Wang and Chen 2017; Viner et al. 2018, 2022; Riebesehl et al. 2019; Shi et al. 2019; Dai et al. 2021; Luo et al. 2021a, 2022; Qu and Zhao 2022; Qu et al. 2022).

Classification of the kingdom of fungi has been updated continuously, based on the frequent inclusion of data from DNA sequences in many phylogenetic studies (Wijayawardene et al. 2020). Based on the early embrace of molecular systematics by mycologists, both the discovery and classiﬁcation of fungi, among the more basal branches of the tree, are now coming to light from genomic analyses and environmental DNA surveys that have been conducted (James et al. 2020). Hyphodontia s.l. was indicated to be a polyphyletic group, and Xylodon and Kneiffiella P. Karst. included the most species (Yurchenko and Wu 2016; Riebesehl and Langer 2017; Riebesehl et al. 2019). Given the lack of sequences for a part of the fungal taxa, it is difficult to clearly separate the many genera in this group 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). On the basis of the nuclear DNA sequence data, six well-distinguished clades—Lagarobasidium clade, Kneiffiella-Alutaceodontia clade, Hyphodontia clade, Hastodontia clade, Xylodon-Lyomyces, Rogersella clade, and Xylodon-Schizopora-Palifer clade—were included based on the phylogenetical studies for Hyphodontia s.l., in which the genera Xylodon and Lyomyces were nested within the Xylodon-Lyomyces-Rogersella clade and Xylodon-Schizopora-Palifer clade, respectively (Yurchenko and Wu 2013). Riebesehl and Langer (2017) 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. Yurchenko et al. (2017) clarified the Lyomyces sambuci complex on the basis of the sequences of the internal transcribed spacer (ITS) and the nuclear large subunit (nLSU) ribosomal DNA gene, in which four new species were described. To confirm the taxonomic relationship of Xylodon species, Viner et al. (2018) proposed two genera, Lagarobasidium and Xylodon, which should be synonymous based on molecular data from the ITS and nLSU regions, and in which the three species were combined into Xylodon. Riebesehl et al. (2019) clarified the generic concept and their phylogenetic reconstruction of Lyomyces, in which L. sambuci was sister to L. crustosus (Pers.) P. Karst. Based on a combination of morphological and molecular evidence, the wood-inhabiting fungal diversity within the family Schizoporaceae of the order Hymenochaetales were analyzed, including Lyomyces fissuratus C.L. Zhao, L. fumosus C.L. Zhao, L. niveus C.L. Zhao and L. ochraceoalbus C.L. Zhao. (Luo et al. 2021b, 2021c). Viner et al. (2022) described three new species from Africa as Xylodon angustisporus Viner & Ryvarden, X. dissiliens Viner & Ryvarden, and X. laxiusculus Viner & Ryvarden, based on the morphological descriptions and molecular analyses that they conducted. A phylogenetic and taxonomic study on Xylodon showed that three new species of Xylodon were from southern China; in addition, it was also found that it enriched the fungal diversity of these areas (Qu et al. 2022).

During investigations on the wood-inhabiting fungi in the Yunnan–Guizhou Plateau of China, samples representing five additional species belonging to genera Lyomyces and Xylodon were collected. To clarify the placement and relationships of the five species, we carried out a phylogenetic and taxonomic study on Lyomyces and Xylodon, based on the ITS and nLSU sequences.

Materials and methods

Morphology

The studied specimens were deposited at the Herbarium of Southwest Forestry University (SWFC), Kunming, Yunnan Province, P.R. China. Macromorphological descriptions are based on field notes and photos captured in the field and lab. Color terminology follows Petersen (Petersen 1996). Micromorphological data were obtained from the dried specimens when observed under a light microscope following Dai (2012). 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 CTAB rapid plant genome extraction kit-DN14 (Aidlab Biotechnologies Co., Ltd, Beijing) was used to obtain genomic DNA from the dried specimens following the manufacturer’s instructions (Zhao and Wu 2017). The nuclear ribosomal ITS region was amplified with the primers ITS5 and ITS4 (White et al. 1990). The nuclear ribosomal LSU gene was amplified with the primers LR0R and LR7 (Vilgalys and Hester 1990; Rehner and Samuels 1994). The PCR procedure for ITS and nLSU followed previous study (Zhao and Wu 2017). All newly-generated sequences were deposited in NCBI 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.		Country	References
Species name	Specimen No.	ITS	nLSU	Country	References
Fasciodontia brasiliensis	MSK-F 7245a	MK575201	MK598734	Brazil	Yurchenko et al. 2020
F. bugellensis	KAS-FD 10705a	MK575203	MK598735	France	Yurchenko et al. 2020
F. bugellensis	MSK-F 7353	MK575205	MK598736	Belarus	Yurchenko et al. 2020
F. yunnanensis	CLZhao 6280	MK811275	MZ146327	China	Luo and Zhao 2021
F. yunnanensis	CLZhao 6385	MK811277	–	China	Luo and Zhao 2021
Hastodontia halonata	HHB-17058	MK575207	MK598738	Mexico	Yurchenko et al. 2020
Hymenochaete ochromarginata	He 47	KU978861	JQ279666	China	Unpublished
H. rubiginosa	He 458	JQ279580	–	China	He and Li 2013
Hyphodontia arguta	KHL 11938 (GB)	EU118632	EU118633	Sweden	Larsson 2007
H. pallidula	GEL 2097	DQ340317	DQ340372	Germany	Unpublished
H. zhixiangii	LWZ 20160909-4	KY440396	–	Uzbekistan	Kan et al. 2017
H. zhixiangii	LWZ 20160909-9	KY440398	–	Uzbekistan	Kan et al. 2017
Kneiffiella eucalypticola	LWZ20180515-9	MT319411	MT319143	Australia	Wang et al. 2021
K. palmae	GEL3456	DQ340333	DQ340369	China	Yurchenko et al. 2020
K. subalutacea	GEL2196	DQ340341	DQ340362	Norway	Yurchenko et al. 2020
*Lyomyces albopulverulentus*	CLZhao 21478*	OP730712	OP730724	China	Present study
L. allantosporus	KAS-GEL4933	KY800401	–	France	Yurchenko et al. 2017
L. allantosporus	FR-0249548	KY800397	–	La Réunion	Yurchenko et al. 2017
L. bambusinus	CLZhao 4831	MN945968	–	China	Chen and Zhao 2020
	CLZhao 4808	MN945970	–	China	Chen and Zhao 2020
	CLZhao 4831	MN945968	–	China	Chen and Zhao 2020
L. cremeus	CLZhao 4138	MN945974	–	China	Chen and Zhao 2020
L. cremeus	CLZhao 8295	MN945972	–	China	Chen and Zhao 2020
L. crustosus	TASM:YG G39	MF382993	–	Uzbekistan	Gafforov et al. 2017
L. crustosus	UC2022841	KP814310	–	USA	Unpublished
L. densiusculus	Ryvarden 44818	OK273853	–	Uganda	Viner et al. 2022
L. elaeidicola	LWZ20180411-20	MT319458	–	Malaysia	Wang et al. 2021
L. elaeidicola	LWZ20180411-19	MT319457	–	Malaysia	Wang et al. 2021
L. erastii	TASM:YG 022	MF382992	–	Uzbekistan	Gafforov et al. 2017
L. erastii	23cSAMHYP	JX857800	–	Spain	Unpublished
L. fimbriatus	Wu910620-7	MK575209	–	China	Yurchenko et al. 2020
L. fimbriatus	Wu911204-4	MK575210	–	China	Yurchenko et al. 2020
L. fissuratus	CLZhao 4352	MW713742	–	China	Luo et al. 2021a
L. fissuratus	CLZhao 4291	MW713738	–	China	Luo et al. 2021a
L. fumosus	CLZhao 8188	MW713744	–	China	Luo et al. 2021a
L. gatesiae	LWZ20180515-3	MT319447	–	Australia	Wang et al. 2021
L. gatesiae	LWZ20180515-32	MT319448	–	Australia	Wang et al. 2021
L. griseliniae	KHL 12971 (GB)	DQ873651	–	Costa Rica	Larsson et al. 2006
L. juniperi	FR-0261086	KY081799	–	La Réunion	Riebesehl and Langer 2017
L. leptocystidiatus	LWZ20170818-1	MT326514	–	China	Wang et al. 2021
L. leptocystidiatus	LWZ20170818-2	MT326513	–	China	Wang et al. 2021
L. macrosporus	CLZhao 4516	MN945977	–	China	Chen and Zhao 2020
L. mascarensis	KAS-GEL4833	KY800399	–	La Réunion	Yurchenko et al. 2020
L. mascarensis	KAS-GEL4908	KY800400	–	La Réunion	Yurchenko et al. 2020
L. microfasciculatus	CLZhao 5109	MN954311	–	China	Chen and Zhao 2020
L. niveus	CLZhao 6431	MZ262541	MZ262526	China	Luo et al. 2021b
L. niveus	CLZhao 6442	MZ262542	MZ262527	China	Luo et al. 2021b
L. ochraceoalbus	CLZhao 4385	MZ262535	MZ262521	China	Luo et al. 2021b
L. ochraceoalbus	CLZhao 4725	MZ262536	MZ262522	China	Luo et al. 2021b
L. organensis	MSK7247	KY800403	–	Brazil	Yurchenko et al. 2017
L. orientalis	GEL3376	DQ340325	–	China	Yurchenko et al. 2017
L. pruni	GEL2327	DQ340312	–	Germany	Larsson et al. 2006
L. pruni	Ryberg 021018 (GB)	DQ873624	–	Sweden	Larsson et al. 2006
L. sambuci	KAS-JR7	KY800402	KY795966	Germany	Yurchenko et al. 2017
L. sambuci	83SAMHYP	JX857721	–	USA	Yurchenko et al. 2017
L. vietnamensis	TNM F9073	JX175044	–	Vietnam	Yurchenko et al. 2013
L. wuliangshanensis	CLZhao 4108	MN945980	–	China	Chen and Zhao 2020
L. wuliangshanensis	CLZhao 4167	MN945979	–	China	Chen and Zhao 2020
*L. yunnanensis*	CLZhao 2463	OP730711	OP730723	China	Present study
	CLZhao 9375	OP730710	–	China	Present study
	CLZhao 10041*	OP730709	–	China	Present study
Xylodon acystidiatus	LWZ20180514-9	MT319474	–	Australia	Wang et al. 2021
X. apacheriensis	Wu 0910-58	KX857797	–	China	Chen et al. 2017
X. aspera	KHL 8530	AY463427	–	Sweden	Larsson et al. 2004
X. astrocystidiata	Wu 9211-71	JN129972	–	China	Yurchenko and Wu 2014
X. attenuatus	Spirin 8775	MH324476	–	America	Viner et al. 2018
X. australis	LWZ20180509-8	MT319503	–	China	Wang et al. 2021
X. bambusinus	CLZhao 9174	MW394657	–	China	Ma and Zhao 2021
X. borealis	JS26064	AY463429	–	Norway	Larsson et al. 2004
X. brevisetus	JS17863	AY463428	–	Norway	Larsson et al. 2004
X. crystalliger	LWZ20170816-33	MT319521	–	China	Wang et al. 2021
X. cystidiatus	FR-0249200	MH880195	MH884896	Réunion	Riebesehl et al. 2019
X. damansaraensis	LWZ20180417-23	MT319499	–	Malaysia	Wang et al. 2021
*X. daweishanensis*	CLZhao 18357*	OP730715	–	China	Present study
	CLZhao 18425	OP730716	–	China	Present study
	CLZhao 18446	OP730717	OP730725	China	Present study
	CLZhao 18458	OP730718	OP730726	China	Present study
	CLZhao 18492	OP730719	OP730727	China	Present study
X. detriticus	Zíbarová 30.10.17	MH320793	–	Czech Republic	Viner et al. 2018
X. filicinus	MSK-F 12869	MH880199	NG067836	China	Riebesehl et al. 2019
*X. fissuratus*	CLZhao 9407*	OP730714	–	China	Present study
*X. fissuratus*	CLZhao 7007	OP730713	–	China	Present study
X. flaviporus	FR-0249797	MH880201	–	Réunion	Riebesehl et al. 2019
X. flocculosus	CLZhao 18342	MW980776	–	China	Qu and Zhao 2022
X. follis	FR-0249814	MH880204	–	Réunion	Riebesehl et al. 2019
X. gossypinus	CLZhao 8375	MZ663804	MZ663813	China	Luo et al. 2021
X. grandineus	CLZhao 16075	OM338091	–	China	Luo et al. 2022
X. grandineus	CLZhao 6425	OM338090	–	China	Luo et al. 2022
X. hastifer	K(M) 172400	NR166558	–	USA	Riebesehl and Langer 2017
X. heterocystidiatus	Wei 17-314	MT731753	–	China	Unpublished
X. hyphodontinus	KAS-GEL9222	MH880205	MH884903	Kenya	Riebesehl et al. 2019
X. kunmingensis	TUB-FO 42565	MH880198	–	China	Riebesehl et al. 2019
X. lacerates	CLZhao 9892	OL619258	–	China	Qu et al. 2022
X. lagenicystidiatus	LWZ20180513-16	MT319634	–	Australia	Wang et al. 2021
X. lenis	Wu890714-3	KY081802	–	China	Riebesehl and Langer 2017
X. macrosporus	CLZhao 10226	MZ663809	MZ663817	China	Luo et al. 2021
X. mollissimus	LWZ 20160318-3	KY007517	–	China	Wang et al. 2021
X. montanus	CLZhao 8179	OL619260	–	China	Qu et al. 2022
X. nespori	LWZ20180921-35	MT319655	MT319238	China	Wang et al. 2021
X. niemelaei	LWZ20150707-13	MT319630	–	China	Wang et al. 2021
X. nongravis	GC 1412-22	KX857801	–	China	Chen et al. 2017
X. nothofagi	ICMP 13842	AF145583	–	China	Paulus et al. 2000
X. ovisporus	LWZ20170815-31	MT319666	–	China	Wang et al. 2021
X. papillosa	CBS:114.71	MH860026	–	Netherlands	Vu et al. 2019
X. paradoxus	Dai14983	MT319519	–	China	Wang et al. 2021
X. pruinosus	Spirin 2877	MH332700	–	Estonia	Viner et al. 2018
X. pseudolanatus	CFMR FP-150922	MH880220	–	Belize	Riebesehl et al. 2019
X. pseudotropicus	Dai16167	MT319509	–	China	Wang et al. 2021
*X. puerensis*	CLZhao 8142*	OP730720	OP730728	China	Present study
*X. puerensis*	CLZhao 8639	OP730721	OP730729	China	Present study
X. punctus	CLZhao 17691	OM338092	–	China	Luo et al. 2022
X. quercinus	Larsson 11076 (GB)	KT361633	–	Sweden	Larsson et al. 2004
X. ramicida	Spirin 7664	NR138013	–	usa	Unpublished
X. rhododendricola	LWZ20180513-9	MT319621	–	Australia	Wang et al. 2021
X. rimosissima	Ryberg 021031 (GB)	DQ873627	–	Sweden	Larsson et al. 2006
X. serpentiformis	LWZ20170816-15	MT319673	–	China	Wang et al. 2021
X. sinensis	CLZhao 11120	MZ663811	–	China	Luo et al. 2021
X. spathulatus	LWZ20180804-10	MT319646	–	China	Wang et al. 2021
X. subclavatus	TUB-FO 42167	MH880232	–	China	Riebesehl et al. 2019
X. subflaviporus	Wu 0809-76	KX857803	–	China	Chen et al. 2017
X. subserpentiformis	LWZ20180512-16	MT319486	–	Australia	Wang et al. 2021
X. subtropicus	LWZ20180510-24	MT319541	–	China	Wang et al. 2021
X. taiwanianus	CBS:125875	MH864080	–	Netherlands	Vu et al. 2019
X. tropicus	CLZhao 3351	OL619261	OL619269	China	Qu et al. 2022
X. ussuriensis	KUN 1989	NR166241	–	USA	Unpublished
X. verecundus	KHL 12261 (GB)	DQ873642	–	Sweden	Larsson et al. 2006
X. victoriensis	LWZ20180510-29	MT319487	–	Australia	Wang et al. 2021
X. wenshanensis	CLZhao 10790	OM338095	–	China	Luo et al. 2022
	CLZhao 15729	OM338097	OM338104	China	Luo et al. 2022
X. xinpingensis	CLZhao 11224	MW394662	MW394654	China	Luo et al. 2022
X. yarraensis	LWZ20180510-5	MT319639	–	Australia	Wang et al. 2021
X. yunnanensis	LWZ20180922-47	MT319660	–	China	Wang et al. 2021

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). Each dataset was aligned separately at first and then the ITS and nLSU regions were combined with Mesquite version 3.51. The combined dataset was deposited in TreeBASE (submission ID 29868). Sequences of Hymenochaete ochromarginata P.H.B. Talbot and H. rubiginosa (Dicks.) Lév. retrieved from GenBank were used as an outgroup in the ITS+nLSU analysis (Fig. 1); Sequences of Xylodon quercinus and X. ramicida Spirin & Miettinen retrieved from GenBank were used as an outgroup in the ITS analysis (Fig. 2); Lyomyces bambusinus C.L. Zhao and L. sambuci were selected as outgroup (Fig. 3) as inspired by a previous study (Luo et al. 2021c).

Figure 1.

Maximum parsimony strict consensus tree illustrating the phylogeny of Xylodon, Lyomyces and related genera in the order Hymenochaetales 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 two 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.

Figure 3.

Maximum parsimony strict consensus tree illustrating the phylogeny of the three new species and related species in Xylodon, 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 analysis in PAUP* version 4.0a169 (http://phylosolutions.com/paup-test/) was applied to ITS and the combined ITS+nLSU 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 pseudoreplicates.

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 8 million generations for ITS+nLSU (Fig. 1); 0.5 million generations for ITS (Fig. 2) and 9.5 million generations for ITS (Fig. 3) 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.

Results

Molecular phylogeny

The ITS+nLSU dataset (Fig. 1) comprised sequences from 43 fungal specimens representing 31 taxa. The dataset had an aligned length of 2,100 characters, of which 1,323 characters were constant, 156 were variable and parsimony-uninformative and 621 (35%) were parsimony-informative. Maximum parsimony analysis yielded 1 equally parsimonious tree (TL = 2,867, CI = 0.4423, HI = 0.5577, RI = 0.6488 and RC = 0.2869). 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 = 0.008603 (BI) and the effective sample size (ESS) across the two runs is double the average ESS (avg. ESS) = 1,623. The phylogram based on the ITS+nLSU rDNA gene regions (Fig. 1) include six genera within Schizoporaceae (Hymenochaetales), which are Fasciodontia, Hastodontia, Hyphodontia, Kneifiella, Lyomyces, and Xylodon—in which five new species were grouped into the genera Lyomyces and Xylodon.

The ITS dataset (Fig. 2) comprised sequences from 47 fungal specimens representing 29 taxa. The dataset had an aligned length of 661 characters, of which 316 characters were constant, 53 were variable and parsimony-uninformative and 292 (35%) were parsimony-informative. Maximum parsimony analysis yielded 1 equally parsimonious tree (TL = 1,371, CI = 0.4136, HI = 0.5864, RI = 0.6984 and RC = 0.2888). 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 = 0.006564 (BI) and the effective sample size (ESS) across the two runs is double the average ESS (avg. ESS) = 359. The phylogenetic tree (Fig. 2), inferred from the ITS sequences, highlighted that L. albopulverulentus formed a monophyletic lineage. It was then grouped closely with L. bambusinus, L. orientalis Riebesehl, Yurch. & Langer, and L. sambuci. In addition, L. yunnanensis was found to be the sister to L. niveus with strong supports.

The ITS dataset (Fig. 3) comprised sequences from 72 fungal specimens representing 65 taxa. The dataset had an aligned length of 702 characters, of which 283 characters were constant, 96 were variable and parsimony-uninformative and 323 (35%) were parsimony-informative. Maximum parsimony analysis yielded 5,000 equally parsimonious trees (TL = 2,726, CI = 0.2748, HI = 0.7252, RI = 0.4280 and RC = 0.1176). 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 = 0.02518 (BI) and the effective sample size (ESS) across the two runs is double the average ESS (avg. ESS) = 1,440. The topology (Fig. 3), based on ITS sequences, revealed that X. daweishanensis was retrieved as a sister to X. hyphodontinus (Hjortstam & Ryvarden) Riebesehl, Yurchenko & G. Gruhn. Furthermore, X. fissuratus was grouped with four taxa: X. montanus C.L. Zhao; X. subclavatus (Yurchenko, H.X. Xiong & Sheng H. Wu) Riebesehl, Yurch. & Langer; X. wenshanensis K.Y. Luo & C.L. Zhao; and X. xinpingensis C.L. Zhao & X. Ma. Moreover, X. puerensis was clustered with X. flaviporus (Berk. & M.A. Curtis ex Cooke) Riebesehl & Langer, X. ovisporus (Corner) Riebesehl & Langer, X. subflaviporus C.C. Chen & Sheng H. Wu, X. subtropicus (C.C. Chen & Sheng H. Wu) C.C. Chen & Sheng H. Wu, and X. taiwanianus (Sheng H. Wu) Hjortstam & Ryvarden.

Taxonomy

Lyomyces albopulverulentus C.L. Zhao, sp. nov.

MycoBank No: 846525

Figs 4, 5

Type material

Holotype. China. Yunnan Province, Lijiang, Lashihai Nature Reserve, 26°51'37"N, 100°8'14"E, altitude 2450 m a.s.l., on fallen angiosperm branch, leg. C.L. Zhao, 19 July 2021, CLZhao 21478 (SWFC).

Etymology

Albopulverulentus (Lat.): referring to the white and pruinose hymenial surface.

Description

Basidiomata annual, resupinate, adnate, brittle, without odor or taste when fresh, up to 12 cm long, 1.5 cm wide, and 150 µm thick. Hymenial surface pruinose, white when fresh and drying. Sterile margin indistinct, white, and up to 2 mm wide.

Figure 4.

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

Hyphal system monomitic, generative hyphae with clamp connections, colorless, thick-walled, frequently branched, interwoven, 3.5–5.5 µm in diameter; IKI–, CB–, tissues unchanged in KOH; subhymenial hyphae densely covered by crystals.

Cystidia capitate, colorless, thin-walled, smooth, slightly constricted at the neck, with a globose tip, 37–54 × 5–9 µm; basidia clavate, slightly sinuous, with four sterigmata and a basal clamp connection, 24.5–28.5 × 7–9 µm.

Figure 5.

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

Basidiospores ellipsoid, colorless, thin-walled, smooth, IKI–, CB–, (7.5–)8–10.5(–11) × (5–)5.5–7 µm, L =9.12 µm, W = 6 µm, Q = 1.52 (n = 30/1).

Additional specimen examined

(paratype). China. Yunnan Province, Yuxi, Xinping County, the Ancient Tea Horse Road, 23°57'10"N, 101°30'41"E, altitude 2,600 m a.s.l., on fallen angiosperm branch, leg. C.L. Zhao, 13 January 2018, CLZhao 5234 (SWFC).

Lyomyces yunnanensis C.L. Zhao, sp. nov.

MycoBank No: 846527

Figs 6, 7

Type material

Holotype. China. Yunnan Province, Dali, Nanjian County, Lingbaoshan, 24°46'2"N, 100°30'26"E, altitude 2350 m a.s.l., on fallen angiosperm branch, leg. C.L. Zhao, 9 January 2019, CLZhao 10041 (SWFC).

Etymology

Yunnanensis (Lat.): referring to the locality (Yunnan Province) of the type specimen.

Description

Basidiomata annual, resupinate, adnate, coriaceous when fresh, becoming farinaceous upon drying, without odor or taste when fresh, up to 15 cm long, 2.5 cm wide, and 150 µm thick. Hymenial surface grandinioid, cream to buff when fresh, and buff upon drying. Sterile margin indistinct, buff, and up to 1 mm wide.

Figure 6.

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

Hyphal system monomitic, generative hyphae with clamp connections, colorless, thick-walled, frequently branched, interwoven, 2.5–3 µm in diameter; IKI–, CB–, tissues unchanged in KOH. Numerous crystals present among hyphae.

Cystidia of two types: (1) fusiform, tapering, colorless, thin-walled, 18–39 × 4–6 µm; (2) capitate cystidia, colorless, thin-walled, 16–23.5 × 3–5 µm; fusoid cystidioles present, colorless, thin-walled, 18–25 × 3–6 µm; basidia clavate, slightly sinuous, with four sterigmata and a basal clamp connection, 16.5–27 × 4–5.5 µm.

Figure 7.

Microscopic structures of Lyomyces yunnanensis (holotype) A basidiospores B basidia and basidioles C tapering cystidia D capitate cystidia E fusoid cystidioles F a section of hymenium. Scale bars: 5 µm (A); 10 µm (B–F).

Basidiospores ellipsoid, colorless, thin-walled, smooth, IKI–, CB–, (4.5–)5–7 × 3–4.5 µm, L = 5.72 µm, W = 3.6 µm, Q = 1.54–1.65 (n = 90/3).

Additional specimens examined

(paratypes). China. Yunnan Province, Yuxi, Xinping County, Mopanshan National Forestry Park, 23°55'48"N, 101°59'22"E, altitude 2150 m a.s.l., on fallen angiosperm branch, leg. C.L. Zhao, 19 August 2017, CLZhao 2463 (SWFC); Puer, Jingdong County, the Forest of Pineapple, 24°21'32"N, 100°48'12"E, altitude 2110 m a.s.l., on fallen angiosperm branch, leg. C.L. Zhao, 4 January 2019, CLZhao 9375 (SWFC).

Xylodon daweishanensis C.L. Zhao, sp. nov.

MycoBank No: 846530

Figs 8, 9

Type material

Holotype. China. Yunnan Province, Honghe, Pingbian County, Daweishan National Nature Reserve, 22°53'26"N, 103°35'37"E, altitude 1990 m a.s.l., on angiosperm trunk, leg. C.L. Zhao, 3 August 2019, CLZhao 18357 (SWFC).

Etymology

Daweishanensis (Lat.): referring to the locality (Daweishan) of the type specimen.

Description

Basidiomata annual, resupinate, adnate, without odor or taste when fresh, coriaceous, up to 10 cm long, 5 cm wide, and 150 µm thick. Hymenial surface odontioid, slightly buff when fresh, and buff upon drying. Margin sterile, slightly buff, and 1 mm wide.

Hyphal system monomitic, generative hyphae with clamp connections, colorless, thin to thick-walled, frequently branched, interwoven, 1.5–4 µm in diameter, IKI–, CB–, tissues unchanged in KOH.

Figure 8.

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

Cystidia of two types: (1) capitate cystidia thin-walled, smooth, slightly constricted at the neck, with a globose tip, 11–23.5 × 2.5–5 µm; (2) asterocystidia thin-walled, with the apical part encrusted, 11–26.5 × 2.5–4.5 µm; basidia clavate to subcylindrical, constricted, somewhat sinuous, with four sterigmata and a basal clamp connection, 11–15.5 × 2.5–4 µm.

Figure 9.

Microscopic structures of Xylodon daweishanensis (holotype) A basidiospores B basidia and basidioles C capitate cystidia D asterocystidia E a section of hymenium. Scale bars: 5 µm (A); 10 µm (B–E).

Basidiospores broad ellipsoid to subglobose, colorless, thin-walled, smooth, with oil drops, IKI–, CB–, 3–4 × 2.5–3.5(–4) µm, L = 3.51 µm, W = 3.14 µm, Q = 1.09–1.15 (n = 150/5).

Additional specimens examined

(paratypes). China. Yunnan Province, Honghe, Pingbian County, Daweishan National Nature Reserve, 22°53'26"N, 103°35'37"E, altitude 1990 m a.s.l., on angiosperm trunk, leg. C.L. Zhao, 3 August 2019, CLZhao 18425, CLZhao 18446, CLZhao 18458, and CLZhao 18492 (SWFC).

Xylodon fissuratus C.L. Zhao, sp. nov.

MycoBank No: 846532

Figs 10, 11

Type material

Holotype. China. Yunnan Province, Puer, Jingdong County, the Forest of Pineapple, 24°21'32"N, 100°48'12"E, altitude 2110 m a.s.l., on fallen angiosperm branch, leg. C.L. Zhao, 4 January 2019, CLZhao 9407 (SWFC).

Etymology

Fissuratus (Lat.): referring to the cracking hymenial surface.

Description

Basidiomata annual, resupinate, adnate, coriaceous, without odor or taste when fresh, up to 12 cm long, 2.5 cm wide, and 150 µm thick. Hymenial surface grandinioid, and white when fresh, white to slightly cream on drying, cracking. Sterile margin indistinct, white, and up to 1 mm wide.

Figure 10.

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

Hyphal system monomitic, generative hyphae with clamp connections, colorless, thin-walled, frequently branched, interwoven, 2–3 µm in diameter; IKI–, CB–, tissues unchanged in KOH.

Figure 11.

Microscopic structures of Xylodon fissuratus (holotype) A basidiospores B basidia and basidioles C capitate cystidia D a section of hymenium. Scale bars: 5 µm (A); 10 µm (B–D).

Cystidia capitate, thin-walled, smooth, slightly constricted at the neck, with a globose tip, 11.5–16.5 × 3–4.5 µm; basidia clavate to subcylindrical, slightly constricted in the middle to somewhat sinuous, with four sterigmata and a basal clamp connection, 10.5–16.5 × 2–4 µm.

Basidiospores ellipsoid, colorless, thin-walled, smooth, IKI–, CB–, 4–5 × 3–4 µm, L = 4.44 µm, W = 3.4 µm, Q = 1.3 (n = 30/1).

Additional specimen examined

(paratype). China. Yunnan Province, Chuxiong, Zixishan Forestry Park, 25°01'26"N, 101°24'37"E, altitude 2313 m a.s.l., on fallen angiosperm branch, leg. C.L. Zhao, 1 July 2018, CLZhao 7007 (SWFC).

Xylodon puerensis C.L. Zhao, sp. nov.

MycoBank No: 846533

Figs 12, 13

Type material

Holotype. China. Yunnan Province, Puer, Zhenyuan County, Heping Town, Jinshan Virgin Forest Park, 23°56'21"N, 101°25'32"E, altitude 2240 m a.s.l., on fallen angiosperm branch, leg. C.L. Zhao, 21 August 2018, CLZhao 8142 (SWFC).

Etymology

Puerensis (Lat.): referring to the locality (Yunnan Province) of the type specimen.

Description

Basidiomata annual, resupinate, adnate, coriaceous, without odor or taste when fresh, up to 12 cm long, 5 cm wide, and 200 µm thick. Hymenial surface poroid, pores angular or slightly daedaleoid, 3–6 per mm, and cream when fresh, buff on drying. Sterile margin slightly buff, and up to 1 mm wide.

Figure 12.

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

Hyphal system monomitic, generative hyphae with clamps, colorless, thick-walled, frequently branched, interwoven, 2.5–4.5 µm in diameter; IKI–, CB–, tissues unchanged in KOH.

Cystidia of four types: (1) paraphysoid cystidia colorless, smooth, 12–20.5 × 3–5 µm; (2) astrocystidia colorless, thin-walled, smooth, 9–11 × 3.5–5.5 µm; (3) capitate cystidia, colorless, thin-walled, smooth, embedded, 22–29.5 × 6.5–12 µm; (4) septocystidia, thin-walled, smooth, with the apical part encrusted, 32–51 × 3.5–6 µm; basidia clavate to subcylindrical, slightly sinuous or distinctly sinuous, with four sterigmata and a basal clamp connection, 14.5–20 × 5–7 µm.

Figure 13.

Microscopic structures of Xylodon puerensis (holotype) A basidiospores B basidia and basidioles C paraphysoid cystidia D astrocystidium E capitate cystidia F septocystidium cystidia G a section of hymenium. Scale bars: 5 µm (A); 10 µm (B–G).

Basidiospores ellipsoid to broad ellipsoid, colorless, thin-walled, smooth, with oil drops, IKI–, CB–, (5.5–)6–7 × 4.5–5.5 µm, L = 6.41 µm, W = 5.01 µm, Q = 1.28 (n = 30/1).

Additional specimen examined

(paratype). China. Yunnan Province, Puer, Jingdong County, Taizhong Town, Ailaoshan Ecological Station, 24°29'41"N, 100°56'32"E, altitude 1930 m a.s.l., on angiosperm trunk, leg. C.L. Zhao, 24 August 2018, CLZhao 8639 (SWFC).

Discussion

Many recently described wood-inhabiting fungal taxa have been reported in the subtropics and tropics, including in the genera Lyomyces and Xylodon (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; Luo et al. 2022; Qu and Zhao 2022; Qu et al. 2022; Viner et al. 2022). Prior to this study, the following fourteen Lyomyces species were reported from China: 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 and L. wuliangshanensis 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 five new species in Lyomyces and Xylodon, based on a combination of morphological features and molecular evidence.

Phylogenetically, based on the multiple loci in Hyphodontia s.l., six genera, Fasciodontia, Hastodontia, Hyphodontia, Lyomyces, Kneiffiella, and Xylodon, were divided into four clades in the order Hymenochaetales (Wang et al. 2021). In the present study, based on the ITS+nLSU data (Fig. 1), Lyomyces and Xylodon were grouped with Fasciodontia, Hastodontia, Hyphodontia and Kneiffielle, in which five new species were grouped into the genera Lyomyces and Xylodon. Based on ITS topology (Figs 2, 3), L. albopulverulentus formed a monophyletic lineage, and was then grouped closely with L. bambusinus, L. orientalis, and L. sambuci. In addition, L. yunnanensis was found to be a sister to L. niveus with strong supports. The topology, based on ITS sequences, revealed that X. daweishanensis was retrieved as a sister to X. hyphodontinus. Moreover, X. fissuratus was grouped with the four taxa X. montanus, X. subclavatus, X. wenshanensis, and X. xinpingensis. X. puerensis was clustered with X. flaviporus, X. ovisporus, X. subflaviporus, X. subtropicus, and X. taiwanianus. However, morphologically, L. bambusinus can be delimited from L. albopulverulentus by its colliculose-to-tuberculate hymenial surface, its narrower basidia (16.5–35 × 3.5–7 µm), and its smaller and more broadly ellipsoid basidiospores (4.7–5.9 × 3.7–4.6 µm; Chen and Zhao 2020); Further, L. orientalis can be delimited from L. albopulverulentus by its smooth or slightly tuberculate hymenial surface, and by both its smaller basidia (13–20 × 3.5–4.5 µm) and basidiospores (5–6 × 4–4.5 µm; Yurchenko et al. 2017); L. sambuci can be delimited from L. albopulverulentus by its smooth-to-tuberculate hymenial surface and its smaller basidiospores (4.5–6 × 3.5–4 µm; Bernicchia and Gorjón 2010); L. niveus can be delimited from L. yunnanensis by a smaller basidia (9.5–15.0 × 3.5–5.5 µm) and broadly ellipsoid basidiospores (3.5–5 × 3–4 µm; Luo et al. 2021c). Xylodon hyphodontinus differs from X. daweishanensis by its larger basidiospores (4–5 × 4.5 µm; Hjortstam and Ryvarden 1980). X. montanus could be delimited from X. fissuratus by its smooth hymenial surface and moniliform cystidia (19.5–47.6 × 3.6–7.1 µm; Qu et al. 2022); X. subclavatus differs from X. fissuratus by its larger capitate cystidia (20–25 × 3–4 μm) and wider basidiospores (4–5.5 × 3.5–4; Yurchenko et al. 2013); X. wenshanensis can be delimited from X. fissuratus by its smaller capitate cystidia (6–11 × 3–6.5 µm; Luo et al. 2022); X. xinpingensis can be delimited from X. fissuratus by its reticulate hymenial surface and larger basidia (18.5–33 × 3–6.5 µm; Ma and Zhao 2021). Xylodon flaviporus differs from X. puerensis by its wider basidia (14.5–20 × 5–7 µm) and smaller basidiospores (4.5–5.5 × 3–3.5 µm; Ryvarden 1985); X. ovisporus differs from X. puerensis by its smaller basidiospores (3.5–4.3 × 2.8–3.3 µm; Riebesehl and Langer 2017); X. subflaviporus is distinguishable from X. puerensis by its narrower basidia (8–18 × 4–5 µm) and smaller basidiospores (3.9–4.8 × 2.7–3.5 µm; Chen et al. 2017); X. subtropicus differs from X. puerensis by its smaller basidiospores (5–5.8 × 3.5–4 µm; Chen et al. 2017); X. taiwanianus differs from X. puerensis by its smaller basidiospores (4.5–5.5 × 2.6–3 µm; Wu 2001).

Morphologically, Lyomyces albopulverulentus resembles L. bambusinus, L. cremeus, L. mascarensis Riebesehl, Yurch. & Langer, L. orientalis, and L. wuliangshanensis, by sharing capitate cystidia and ellipsoid basidiospores. However, L. bambusinus differs from L. albopulverulentus by possessing a tapering cystidia (40–65 × 4–5.5 µm) and smaller basidiospores (4.7–5.9 × 3.7–4.6 µm; Chen and Zhao 2020); L. cremeus differs from L. albopulverulentus by its narrower capitate cystidia (20–40 × 3–5 µm), both smaller basidia (9–18.5 × 3–6 µm) and basidiospores (4.5–5.6 × 3.3–4.3 µm; Chen and Zhao 2020); L. mascarensis can be delimited from L. albopulverulentus by smaller capitate cystidia (17–38 × 3.5–6 µm), basidia (16–17.5 × 3.5–4.5 µm) and basidiospores (4.5–6 × 3.3–4 µm; Yurchenko et al. 2017); L. orientalis can be delimited from L. albopulverulentus due to its smaller capitate cystidia (17–38 × 3.5–6 µm), basidia (16–17.5 × 3.5–4.5 µm) and basidiospores (4.5–6 × 3.3–4 µm; Yurchenko et al. 2017); L. wuliangshanensis is different from L. albopulverulentus by smaller capitate cystidia (22–37 × 3–6 µm), basidia (12–20 × 3–4.3 µm) and basidiospores (3.5–5.3 × 2.8–4 µm; Chen and Zhao 2020).

Morphologically, Lyomyces yunnanensis resembles L. bambusinus, L. cremeus, L. fumosus, L. fissuratus and L. wuliangshanensis in both its capitate and tapering cystidia. However, L. bambusinus differs from L. yunnanensis by possessing a larger capitate cystidia (35–55 × 4–7 µm; Chen and Zhao 2020); L. cremeus differs from L. yunnanensis due to its smooth hymenial surface and smaller basidia (9–18.5 × 3–6 µm; Chen and Zhao 2020); L. fissuratus can be delimited from L. yunnanensis by its white-to-cream hymenial surface, and the presence of submoniliform cystidia (15.5–22 × 2.7–4 µm; Luo et al. 2021b); L. fumosus differs from L. yunnanensis due to its smooth hymenial surface, the presence of moniliform cystidia (8.5–22.7 × 2.5–3.7 µm), and its smaller basidia (11.5–17.5 × 3–5 µm; Luo et al. 2021b); L. wuliangshanensis is distinguishable from L. yunnanensis by its larger capitate cystidia (22–37 × 3–6 µm) and its smaller basidiospores (3.5–5.3 × 2.8–4 µm; Chen and Zhao 2020).

Morphologically, Xylodon daweishanensis is similar to X. follis Riebesehl et al., X. grandineus K.Y. Luo & C.L. Zhao, X. laceratus C.L. Zhao, X. macrosporus, X. sinensis C.L. Zhao & K.Y. Luo and X. tropicus C.L. Zhao due to its grandinioid, or odontioid, hymenial surface. However, X. follis differs from X. daweishanensis due to its cream hymenial surface, wider capitate cystidia (17–30 × 4.5–9 µm), and larger, globose to subglobose basidiospores (8–9.5 × 7–8.5 µm; Riebesehl et al. 2019); X. grandineus differs from X. daweishanensis by its subulate cystidia (11–19 × 3–5 µm; Luo et al. 2022); X. laceratus can be delimited from X. daweishanensis by its fusiform cystidia (20.3–26.8 × 5.3–6.4 µm) and its larger basidiospores (3.9–5.3 × 2.6–4.1 µm; Qu et al. 2022); X. macrosporus differs from X. daweishanensis by its cylindrical cystidia (44–79.5 × 3–6 µm), larger basidia (11.5–36 × 5–11 µm) and thick-walled basidiospores (8–10.5 × 7.5–9 µm; Luo et al. 2021a); X. sinensis differs from X. daweishanensis by its fusiform cystidia (10–21 × 3–6 µm), and its buff-to-brown hymenial surface (Luo et al. 2021a); X. tropicus can be delimited from X. daweishanensis by its subglobose, slightly thick-walled basidiospores (Qu et al. 2022).

Xylodon fissuratus resembles X. attenuatus Spirin & Viner, X. borealis (Kotir. & Saaren.) Hjortstam & Ryvarden, X. bresinskyi (Langer) Hjortstam & Ryvarden, X. dimiticus (Jia J. Chen & L.W. Zhou) Riebesehl & E. Langer, X. grandineus and X. vesiculosus Yurchenko et al. by it sharing similar ellipsoid basidiospores. However, X. attenuatus differs from X. fissuratus due to its odontoid hymenial surface, the presence of hyphoid cystidia (17.6–39 × 2.7–4.6 µm) and its larger capitate cystidia (14.2–27.2 × 3.3–4.5 µm; Viner et al. 2018); X. borealis differs from X. fissuratus by its slender hypha-like cystidia (40–70 × 3–5 µm), larger capitate cystidia (20–50 × 4–6 µm) and basidia (15–20 × 4–5 µm; Bernicchia and Gorjón 2010); X. bresinskyi differs from X. fissuratus by its poroid hymenial surface with rudimentary console shaping (Langer 2000); X. dimiticus is distinguishable from X. fissuratus by poroid hymenial surface with angular pores (2–4 per mm; Chen et al. 2016); X. grandineus differs from X. fissuratus due to its subulate cystidia (11–19 × 3–5 µm) and its smaller basidiospores (3–4.5 × 2–3 µm; Luo et al. 2022); X. vesiculosus can be delimited from X. fissuratus by its odontioid hymenial surface and larger basidiospores (5.3–6.3 × 3–4 µm; Riebesehl et al. 2019).

Xylodon puerensis is similar to X. bresinskyi, X. dimiticus, X. hallenbergii (Sheng H. Wu) Hjortstam & Ryvarden, X. poroideoefibulatus (Sheng H. Wu) Hjortstam & Ryvarden, X. reticulatus (C.C. Chen & Sheng H. Wu) C.C. Chen & Sheng H. Wu, X. subtropicus and X. syringae (Langer) Hjortstam & Ryvarden by sharing a similar poroid hymenophore. However, X. bresinskyi can be delimited from X. puerensis by possessing smaller basidiospores (4.5–5.5 × 3–3.5 µm; Langer 2000); X. dimiticus differs from X. puerensis by possessing smaller basidia (9–13 × 4.5–6 µm) and basidiospores (3.8–4.6 × 2.8–3.5 µm; Chen et al. 2016); X. hallenbergii can be delimited from X. fissuratus by its both smaller capitate cystidia (15–23 × 4–5.3 µm) and basidiospores (4.2–5 × 4–4.3 µm; Wu 2001); X. poroideoefibulatus differs from X. puerensis by possessing smaller capitate cystidia (12–23 × 5.5–6.5 µm) and basidiospores (5–5.7 × 4–4.5 µm; Wu 2001); X. reticulatus can be delimited from X. puerensis by possessing smaller basidiospores (5–5.5 × 3.5–4 µm; Wu 1990); X. syringae differs from X. puerensis by its larger basidia (20–32 × 4–5 µm) and suballantoid basidiospores (8–9 × 3–3.5 µm; Hjortstam and Ryvarden 2009).

Acknowledgements

The research was supported by the National Natural Science Foundation of China (Project No. 32170004); the Yunnan Fundamental Research Project (Grant No. 202001AS070043); the High-level Talents Program of Yunnan Province (YNQR-QNRC-2018-111).

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﻿Abstract

Keywords

﻿Introduction

﻿Materials and methods

﻿Morphology

﻿Molecular phylogeny

﻿Results

﻿Molecular phylogeny

﻿Taxonomy

﻿ Lyomyces albopulverulentus C.L. Zhao, sp. nov.

Type material

Etymology

Description

Additional specimen examined

﻿ Lyomyces yunnanensis C.L. Zhao, sp. nov.

Type material

Etymology

Description

Additional specimens examined

﻿ Xylodon daweishanensis C.L. Zhao, sp. nov.

Type material

Etymology

Description

Additional specimens examined

﻿ Xylodon fissuratus C.L. Zhao, sp. nov.

Type material

Etymology

Description

Additional specimen examined

﻿ Xylodon puerensis C.L. Zhao, sp. nov.

Type material

Etymology

Description

Additional specimen examined

﻿Discussion

﻿Acknowledgements

﻿References

Abstract

Introduction

Materials and methods

Morphology

Molecular phylogeny

Results

Molecular phylogeny

Taxonomy

Lyomyces albopulverulentus C.L. Zhao, sp. nov.

Lyomyces yunnanensis C.L. Zhao, sp. nov.

Xylodon daweishanensis C.L. Zhao, sp. nov.

Xylodon fissuratus C.L. Zhao, sp. nov.

Xylodon puerensis C.L. Zhao, sp. nov.

Discussion

Acknowledgements

References