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Research Article
Morphological and molecular identification of four new resupinate species of Lyomyces (Hymenochaetales) from southern China
expand article infoJun-Zhu Chen, Chang-Lin Zhao
‡ Southwest Forestry University, Kunming, China
Open Access

Abstract

Four new wood-inhabiting fungal species, Lyomyces bambusinus, L. cremeus, L. macrosporus and L. wuliangshanensis, are proposed based on a combination of morphological and molecular evidence. Lyomyces bambusinus is characterized by resupinate basidiomata with colliculose to tuberculate hymenial surface and broadly ellipsoid, hyaline, slightly thick-walled, smooth basidiospores. Lyomyces cremeus is characterised by resupinate basidiomata with smooth, cream hymenial surface and ellipsoid, hyaline, thin-walled to slightly thick-walled basidiospores. Lyomyces macrosporus is characterized by pruinose basidiomata with reticulate hymenial surface, presence of three kinds of cystidia and larger basidiospores (6.7–8.9 × 4.4–5.4 µm). Lyomyces wuliangshanensis is characterized by coriaceous basidiomata and ellipsoid, hyaline, slightly thick-walled, smooth basidiospores. The phylogenetic analyses based on molecular data of the internal transcribed spacer (ITS) region sequences revealed that the four new species belonged to Lyomyces. Lyomyces bambusinus grouped with L. sambuci. Lyomyces cremeus clade was sister to a clade comprised of L. microfasciculatus. Lyomyces macrosporus was sister to L. allantosporus. Lyomyces wuliangshanensis was closely related to L. mascarensis.

Keywords

Phylogeny, Schizoporaceae, taxonomy, wood-inhabiting fungi, Yunnan Province

Introduction

Lyomyces P. Karst. typified by L. sambuci (Pers.) P. Karst., is a small corticioid genus characterized by resupinate to effused basidiomata with smooth to granular or odontioid hymenophore, a monomitic hyphal system bearing clamp connections, strongly encrusted generative hyphae, the presence of several types of cystidia; clavate to suburniform basidia, and smooth, thin- to slightly thick-walled, cyanophilous basidiospores (Karsten 1881; Bernicchia and Gorjón 2010). The members of Lyomyces grow on dead, still-attached or fallen branches of angiosperms, on dead wooden and herbaceous stems, or occasionally on gymnosperm wood (Yurchenko et al. 2017). Twenty-three species are currently known in Lyomyces worldwide (Rabenhorst 1851; Karsten 1881, 1882; Peck 1903; Bourdot and Galzin 1911; Cunningham 1959, 1963; Wu 1990; Hjortstam and Ryvarden 2009; Yurchenko et al. 2013, 2017; Gafforov et al. 2017; Riebesehl and Langer 2017) and five species were recorded in China (Xiong et al. 2009; Gafforov et al. 2017; Riebesehl and Langer 2017).

Molecular studies on Lyomyces and related genera have been carried out recently (Riebesehl and Langer 2017; Yurchenko et al. 2017; Viner et al. 2018; Riebesehl et al. 2019). Riebesehl and Langer (2017) indicated that Hyphodontia s.l. should be divided into several genera: Hastodontia (Parmasto) Hjortstam & Ryvarden, Hyphodontia J. Erikss, Kneiffiella (Pers.) Gray, Lagarobasidium Jülich, Lyomyces and Xylodon (Pers.) Gray and thus 35 new combinations were proposed, including fourteen Lyomyces species. The clarification of Lyomyces sambuci complex was conducted based on ITS and 28S sequences analyses and four new species of Lyomyces were described (Yurchenko et al. 2017). Viner et al. (2018) studied the taxonomy of Lagarobasidium and Xylodon, and showed that twelve species clustered into Lyomyces clade and then grouped with Xylodon clade. Phylogenetic and morphological studies on Xylodon showed that Xylodon was distinct from Hastodontia, Hyphodontia, Kneiffiella and Lyomyces and the Lyomyces generic species L. sambuci was sister to L. crustosus (Pers.) P. Karst. formed a single lineage with a high support (Riebesehl et al. 2019).

During investigations on wood-inhabiting fungi in southern China, four additional taxa were found, which could not be assigned to any described species in Lyomyces. In this study, the authors expand samplings from previous studies (Gafforov et al. 2017; Riebesehl and Langer 2017) to examine taxonomy and phylogeny of them within Lyomyces, based on the internal transcribed spacer (ITS) regions sequences.

Materials and methods

Morphological studies

The specimens studied have been deposited in the herbarium of Southwest Forestry University (SWFC), Kunming, Yunnan Province, P.R. China. Special color terms follow Petersen (1996). Macromorphological descriptions are based on field notes. Micromorphological data were obtained from the dried specimens and observed under a light microscope following Dai (2010) and Cui et al. (2019). The following abbreviations are used: KOH = 5% potassium hydroxide; CB = cotton blue; CB+ = cyanophilous; IKI = Melzer’s reagent; IKI– = non-amyloid and non-dextrinoid; L = mean spore length (arithmetic average of all spores); W = mean spore width (arithmetic average of all spores); Q = L/W ratio; n (a/b) = number of spores (a) measured from given number (b) of specimens.

DNA extraction and sequencing

CTAB rapid plant genome extraction kit-DN14 (Aidlab Biotechnologies Co., Ltd, Beijing) was used to obtain genomic DNA from dried specimens, according to the manufacturer’s instructions (Han et al. 2016; Song and Cui 2017) . The ITS region was amplified with the primer pair ITS5 and ITS4 (White et al. 1990). The PCR cycling procedure for ITS was as follows: initial denaturation at 95 °C for 3 min, followed by 35 cycles at 94 °C for 40 s, 58 °C for 45 s and 72 °C for 1 min, and a final extension of 72 °C for 10 min followed Shen et al. (2019). The PCR products were purified and directly sequenced at Kunming Tsingke Biological Technology Limited Company, Yunnan Province, P.R.China. All newly generated sequences were deposited in GenBank (Table 1).

Table 1.

List of species, specimens and GenBank accession numbers of sequences used in this study.

Species name Sample no. GenBank accession no. References
ITS
Lyomyces allantosporus KAS-GEL 4933 KY800401 Yurchenko et al. 2017
FR 0249548 KY800397 Yurchenko et al. 2017
Lyomyces bambusinus CLZhao 3675 MN945969 Present study
CLZhao 4808 MN945970 Present study
CLZhao 4831 MN945968 Present study
CLZhao 4840 MN945971 Present study
Lyomyces cremeus CLZhao 2812 MN945973 Present study
CLZhao 4138 MN945974 Present study
CLZhao 8295 MN945972 Present study
Lyomyces crustosus YG-G 39 MF382993 Gafforov et al. 2017
UC 2022841 KP814310 Rosenthal et al. 2017
Lyomyces erastii MA-Fungi 34336 JX857800 Gafforov et al. 2017
YG 022 MF382992 Gafforov et al. 2017
Lyomyces griseliniae KHL 12971 DQ873651 Larsson et al. 2006
Lyomyces juniperi KAS-GEL 4940 DQ340316 Yurchenko et al. 2017
FR 0261086 KY081799 Riebesehl and Langer 2017
Lyomyces macrosporus CLZhao 4516 MN945977 Present study
CLZhao 4531 MN945978 Present study
CLZhao 8605 MN945975 Present study
CLZhao 3951 MN945976 Present study
Lyomyces mascarensis KAS-GEL 4833 KY800399 Yurchenko et al. 2017
KAS-GEL 4908 KY800400 Yurchenko et al. 2017
Lyomyces microfasciculatus CLZhao 4626 MK343568 Present study
CLZhao 5109 MN954311 Present study
TNM F 24757 JN129976 Yurchenko and Wu 2014
Lyomyces organensis MSK 7247 KY800403 Yurchenko et al. 2017
Lyomyces orientalis KAS-GEL 3376 DQ340325 Yurchenko et al. 2017
KAS-GEL 3400 DQ340326 Yurchenko et al. 2017
Lyomyces pruni Ryberg 021018 DQ873624 Larsson et al. 2006
Lyomyces sambuci 80 SAMHYP JX857721 Yurchenko et al. 2017
83 SAMHYP JX857720 Yurchenko et al. 2017
Lyomyces vietnamensis TNM F 9073 JX175044 Yurchenko and Wu 2014
Lyomyces wuliangshanensis CLZhao 4108 MN945980 Present study
CLZhao 4144 MN945981 Present study
CLZhao 4167 MN945979 Present study
CLZhao 4206 MN945982 Present study
CLZhao 4475 MN945983 Present study
Palifer verecundus KHL 12261 DQ873642 Larsson et al. 2006
Xylodon asperus UC 2023169 KP814365 Yurchenko et al. 2017

Phylogenetic analyses

Sequencher 4.6 (GeneCodes, Ann Arbor, MI, USA) was used to assemble and edit the DNA sequence. Sequences were aligned in MAFFT 7 (https://mafft.cbrc.jp/alignment/server/) using the “G-INS-I” strategy and manually adjusted in BioEdit (Hall 1999). The sequence alignment was deposited in TreeBase (submission ID 25382). Sequences of Palifer verecundus (G. Cunn.) Stalpers & P.K. Buchanan and Xylodon asperus (Fr.) Hjortstam & Ryvarden obtained from GenBank were used as outgroups to root trees following Yurchenko et al. (2017) in Fig. 1.

Maximum parsimony (MP), Maximum Likelihood (ML) and Bayesian Inference (BI) analyses were applied to the ITS dataset sequences. Approaches to phylogenetic analyses followed Wu et al. (2018) and Zhu et al. (2019) the tree construction procedure was performed in PAUP* version 4.0b10 (Swofford 2002). 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 1000 random sequence additions. Max-trees were set to 5000, branches of zero length were collapsed and all most-parsimonious trees were saved. Clade robustness was assessed using bootstrap (BT) analysis with 1000 replicates (Felsenstein 1985). Descriptive tree statistics tree length (TL), consistency index (CI), retention index (RI), rescaled consistency index (RC) and homoplasy index (HI) were calculated for each most-parsimonious tree generated.

Sequences were also analyzed using Maximum Likelihood (ML) ML analysis was conducted with RAxML-HPC2 through the Cipres Science Gateway (www.phylo.org; Miller et al. 2009). Branch support (BS) for ML analysis was determined by 1000 bootstrap replicates and evaluated under the gamma model.

MrModeltest 2.3 (Nylander 2004) was used to determine the best-fit evolution model for the data set for Bayesian Inference (BI). Bayesian Inference was performed with MrBayes 3.1.2 with a general time reversible (GTR) model of DNA substitution and a gamma distribution rate variation across sites (Ronquist and Huelsenbeck 2003). Four Markov chains were used in each of 2 runs from random starting trees for 600,000 generations, with trees and parameters sampled every 100 generations. The first quarter of generations were discarded as burn-in. A majority rule consensus tree of all remaining trees and posterior probabilities was calculated. Branches that received bootstrap support for maximum likelihood (BS) ≥75%, maximum parsimony (BP) ≥75%, and Bayesian posterior probabilities (BPP) ≥0.95 were considered significantly supported.

Phylogeny results

The ITS dataset (Fig. 1) included sequences from 39 fungal specimens representing 18 species. The dataset had an aligned length of 608 characters, of which 277 characters were constant and 242 parsimony-informative. MP analysis yielded 8 equally parsimonious trees (TL = 978, CI = 0.523, HI = 0.478, RI = 0.738, RC = 0.385). The best-fit model for ITS alignment estimated and applied in the BI was GTR+I+G. At the end of the BI runs, the average standard deviation of split frequencies was 0.008676. The tree topology obtained by BI and ML was similar to the MP one.

The phylogenetic tree (Fig. 1) demonstrated that all samples grouped into the Lyomyces in the present study. Lyomyces bambusinus grouped with L. sambuci. Lyomyces cremeus formed a monophyletic lineage and then grouped with L. microfasciculatus (Yurchenko & Sheng H. Wu) Riebesehl & Langer. Lyomyces macrosporus was sister to L. allantosporus Riebesehl, Yurchenko & Langer. Lyomyces wuliangshanensis was closely related to L. mascarensis Riebesehl, Yurchenko & Langer.

Figure 1. 

Maximum parsimony strict consensus tree illustrating the phylogeny of four new species and related species in Lyomyces based on ITS sequences. Branches are labelled with maximum likelihood bootstrap equal to or higher than 70%, parsimony bootstrap proportions equal to or higher than 50% and Bayesian posterior probabilities equal to or high than 0.95 respectively.

Taxonomy

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

MycoBank No: 834036
Figs 2, 6

Holotype

China. Yunnan Province: Puer, Zhenyuan County, Heping Town, Ailaoshan National Nature Reserve, on dead bamboo, 11 January 2018, CLZhao 4831 (SWFC).

Etymology

The epithet bambusinus (Lat.): refers to the occurrence on bamboo.

Basidiomata

Annual, resupinate, ceraceous when fresh, becoming brittle and cracking upon drying, up to 20 cm long and 8 cm wide, 100–200 µm thick. Hymenial surface colliculose to tuberculate, white to cream when fresh, turning cream to buff upon drying. Margin narrow, concolorous with hymenial surface.

Hyphal system

Monomitic; generative hyphae with clamp connections, hyaline, thick-walled, branched, 2.5–3.9 µm in diameter, IKI–, cyanophilous; tissues unchanged in KOH. Numerous crystals present among hyphae.

Hymenium

Two kinds of cystidia: 1) capitate, hyaline, thin-walled, 35–55 × 4–7 µm, smooth or slightly encrusted; 2) tapering, hyaline, thin-walled, 40–65 × 4–5.5 µm, smooth or slightly encrusted; cystidioles present, hyaline, thin-walled, 12–17 × 2–3 µm. Basidia clavate, constricted, thin-walled, with four sterigmata and a basal clamp connection, 16.5–35 × 3.5–7 µm.

Spores

Basidiospores broadly ellipsoid, hyaline, slightly thick-walled, smooth, IKI–, cyanophilous, guttulate, (4.5–)4.7–5.9 (–6.2) × (3.4–)3.7–4.6(–4.8) µm, L = 5.31 µm, W = 4.19 µm, Q = 1.23–1.3 (n = 120/4).

Ecology and distribution

On dead bamboo, causing a white rot. China.

Additional specimens examined

China. Yunnan Province: Puer, Jingdong County, Wuliangshan National Nature Reserve, on dead bamboo, 3 October 2017, CLZhao 3675; Zhenyuan County, Heping Town, Ailaoshan National Nature Reserve, on dead bamboo, 11 January 2018, CLZhao 4808, CLZhao 4840 (SWFC).

Figure 2. 

Basidiomata of Lyomyces bambusinus (holotype). Scale bars: 1 cm (A); 5 mm (B).

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

MycoBank No: 834037
Figs 3, 7

Holotype

China. Yunnan Province: Jingdong County, Taizhong Town, Ailaoshan Ecological Station, on fallen branch of angiosperm, 23 August 2018, CLZhao 8295 (SWFC).

Etymology

The epithet cremeus (Lat.): refers to the cream hymenial surface.

Basidiomata

Annual, resupinate, ceraceous when fresh, becoming membranaceous upon drying, up to 13 cm long and 5 cm wide, 50–100 µm thick. Hymenial surface smooth, pale cream when fresh, turn cream upon drying. Margin narrow, white to cream.

Hyphal system

Monomitic; generative hyphae with clamp connections, hyaline, thick-walled, branched, 3–5 µm in diameter, IKI–, cyanophilous; tissues unchanged in KOH. Numerous crystals present among hyphae.

Hymenium

Two kinds of cystidia: 1) capitate, hyaline, thin-walled, 20–40 × 3–5 µm, smooth or slightly encrusted; 2) tapering, hyaline, thin-walled, 18–35 × 3–4.5 µm, smooth or slightly encrusted; cystidioles present, hyaline, thin-walled, 15–20 × 2.5–4 µm. Basidia clavate, with four sterigmata and a basal clamp connection, 9–18.5 × 3–6 µm.

Spores

Basidiospores ellipsoid, hyaline, thin-walled to slightly thick-walled, smooth, IKI–, cyanophilous, guttulate, 4.5–5.6(–5.8) × 3.3–4.3(–4.5) µm, L = 5.01 µm, W = 3.94 µm, Q = 1.25–1.3 (n = 90/3).

Ecology and distribution

Lignicolous, causing a white rot. China.

Additional specimens examined

China. Yunnan Province: Yuxi, Xinping County, Shimenxia Forestry Park, on fallen branch of angiosperm, 21 August 2017, CLZhao 2812; Puer, Jingdong County, Wuliangshan National Nature Reserve, on fallen branch of angiosperm, 5 October 2017, CLZhao 4138 (SWFC).

Figure 3. 

Basidiomata of Lyomyces cremeus (holotype). Scale bars: 1 cm (A); 5 mm (B).

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

MycoBank No: 834038
Figs 4, 8

Holotype

China. Yunnan Province: Puer, Jingdong County, Taizhong Town, Ailaoshan National Nature Reserve, on fallen branch of angiosperm, 24 August 2018, CLZhao 8605 (holotype in SWFC).

Etymology

The epithet macrosporus (Lat.): refers to the larger basidiospores.

Basidiomata

Annual, resupinate, subceraceous when fresh, becoming pruinose upon drying, up to 22 cm long and 3 cm wide, 100–200 µm thick. Hymenial surface reticulate, cream when fresh, turning cream to buff upon drying. Margin narrow, white to buff.

Hyphal system

Monomitic; generative hyphae with clamp connections, hyaline, thick-walled, branched, 2.5–4 µm in diameter, IKI–, cyanophilous; tissues unchanged in KOH. Numerous crystals present among hyphae.

Hymenium

Three kinds of cystidia: 1) capitate, hyaline, thin-walled, 19–35 × 3–7 µm; 2) tapering, hyaline, thin-walled, 13–20 × 2.5–4 µm; 3) moniliform, hyaline, thin-walled, 15–22 × 4.5–6 µm; fusoid cystidioles present, hyaline, thin-walled, 15–20 × 2.5–4 µm. Basidia subclavate to clavate, constricted, hyaline, thin-walled, with four sterigmata and a basal clamp connection, 22.2–38 × 4.5–7 µm.

Spores

Basidiospores ellipsoid, hyaline, slightly thick-walled, smooth, IKI–, cyanophilous, guttulate, (6.4–)6.7–8.9(–9.1) × 4.4–5.4(–5.7) µm, L = 7.84 µm, W = 4.93 µm, Q = 1.48–1.8 (n = 120/4).

Ecology and distribution

Lignicolous, causing a white rot. China.

Additional specimens examined

China. Yunnan Province: Puer, Jingdong County, Taizhong Town, Ailaoshan National Nature Reserve, on fallen branch of angiosperm, 4 October 2017, CLZhao 3951; Wuliangshan National Nature Reserve, on fallen branch of angiosperm, 6 October 2017, CLZhao 4516, CLZhao 4531 (SWFC).

Figure 4. 

Basidiomata of Lyomyces macrosporus (holotype). Scale bars: 1 cm (A); 5 mm (B).

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

MycoBank No: 834039
Figs 5, 9

Holotype

China. Yunnan Province: Puer, Jingdong County, Wuliangshan National Nature Reserve, on fallen branch of angiosperm, 5 October 2017, CLZhao 4167 (SWFC).

Etymology

The epithet wuliangshanensis (Lat.): refers to the locality (Wuliangshan) of the type specimens.

Basidiomata

Annual, resupinate, subcoriaceous when fresh, becoming coriaceous upon drying, up to 15 cm long and 5 cm wide, 50–150 µm thick. Hymenial surface smooth to more or less tuberculate, white to cream when fresh, turning cream to buff upon drying. Margin narrow, concolorous with hymenial surface.

Hyphal system

Monomitic; generative hyphae with clamp connections, hyaline, thick-walled, branched, 2–3 µm in diameter, IKI–, cyanophilous; tissues unchanged in KOH. Numerous crystals present among hyphae.

Hymenium

Two kinds of cystidia: 1) capitate, hyaline, thin-walled, 22–37 × 3–6 µm; 2) tapering, hyaline, thin-walled, 21–35 × 4–6.5 µm; fusoid cystidioles present, hyaline, thin-walled, 16–21 × 2.5–3.5 µm. Basidia clavate, hyaline, thin-walled, with four sterigmata and a basal clamp connection, 12–20 × 3–4.3 µm.

Spores

Basidiospores ellipsoid, hyaline, slightly thick-walled, smooth, IKI–, cyanophilous, guttulate, (3.3–)3.5–5.3(–5.5) × 2.8–4(–4.2) µm, L = 4.3 µm, W = 3.56 µm, Q = 1.22–1.31 (n = 120/4).

Ecology and distribution

Lignicolous, causing a white rot. China.

Additional specimens examined

China. Yunnan Province: Puer, Jingdong County, Wuliangshan National Nature Reserve, on angiosperm trunk, 5 October 2017, CLZhao 4108, CLZhao 4144; on angiosperm stump, 5 October 2017, CLZhao 4206; on fallen branch of angiosperm, 6 October 2017, CLZhao 4475 (SWFC).

Figure 5. 

Basidiomata of Lyomyces wuliangshanensis (holotype). Scale bars: 1 cm (A); 5 mm (B).

Figure 6. 

Microscopic structures of Lyomyces bambusinus (drawn from the holotype) A basidiospores B basidia and basidioles C cystidia D cystidioles E a cross section of basidiomata. Scale bars: 5 μm (A); 10 μm (B–E).

Figure 7. 

Microscopic structures of Lyomyces cremeus (drawn from the holotype) A basidiospores B basidia and basidioles C cystidia D cystidioles E a cross section of basidiomata. Scale bars: 5 μm (A); 10 μm (B–E).

Figure 8. 

Microscopic structures of Lyomyces macrosporus (drawn from the holotype) A basidiospores B basidia and basidioles C cystidia D cystidioles E a cross section of basidiomata. Scale bars: 10 μm (A–E).

Figure 9. 

Microscopic structures of Lyomyces wuliangshanensis (drawn from the holotype) A basidiospores B basidia and basidioles C cystidia D cystidioles E a cross section of basidiomata. Scale bars: 5 μm (A); 10 μm (B–E).

Discussion

Miettinen et al. (2016) analyzed a phylogenetic classification in Phanerochaetaceae (Polyporales, Basidiomycota) and showed that the macromorphology of basidiomata and hymenophore construction did not reflect monophyletic groups. The phylogeny we obtained (Fig. 1) shows that the macromorphological and micromorphological characters are not consistent with monophyletic groups.

In our phylogeny, Lyomyces bambusinus was sisiter to L. sambuci, but morphologically L. sambuci differs from L. bambusinus by having ellipsoid to oblong, narrower basidiospores (4.5–6 × 3–3.5 µm, Yurchenko et al. 2017). Lyomyces cremeus formed a monophyletic lineage with strong supports (100% BS, 100% BP, 1.00 BPP; Fig. 1) and then was sister to a clade comprised of L. microfasciculatus, L. mascarensis, and L. wuliangshanensis. However L. microfasciculatus differs in odontioid hymenophore and presence of minute peg-like fascicles of hyphae (Yurchenko and Wu 2014). Lyomyces macrosporus was sister to L. allantosporus, but morphologically L. allantosporus differs in having porulose hymenophore and suballantoid to allantoid, narrower basidiospores (7–9 × 3–3.8 µm, Yurchenko et al. 2017). Lyomyces wuliangshanensis grouped closely with L. mascarensis, but L. mascarensis differs from L. wuliangshanensis by having thin-walled generative hyphae and presence of capitate cystidia (17–38 × 3.5–6 µm, Yurchenko et al. 2017).

Five Lyomyces species were reported from China prior to this study, Lyomyces albus (Sheng H. Wu) Riebesehl & Langer, L. capitatocystidiatus (H.X. Xiong, Y.C. Dai & Sheng H. Wu) Riebesehl & Langer, L. microfasciculatus, L. sambuci and L. tenuissimus (Yurchenko & Sheng H. Wu) Riebesehl & Langer. Lyomyces albus differs from four new species by its odontioid hymenophore (Riebesehl and Langer 2017); L. capitatocystidiatus by grandinioid hymenophore with arachnoid-farinaceous hymenial surface (Xiong et al. 2009); L. microfasciculatus by minutely odontioid hymenial surface and small emerging fascicles of flexuous hyphae (Yurchenko and Wu 2014); L. tenuissimus by much thinner basidiomata and non-encrusted, subicular hyphae, shorter cystidia and shorter basidia (Yurchenko et al. 2013).

Hyphodontia s.l. is an extensively studied group of Hymenochaetales (Dai 2012; Viner et al. 2018; Riebesehl et al. 2019), but the Chinese species diversity is still not well known, especially in subtropical and tropical areas. The four new Lyomyces species here described are from the subtropics.

Acknowledgements

This research was supported by the National Natural Science Foundation of China (Project No. 31700023), Yunnan Agricultural Foundation Projects (2017FG001-042), and the Key Laboratory of State Forestry Administration for Highly-Efficient Utilization of Forestry Biomass Resources in Southwest China (Southwest Forestry University) (Project No. 2019-KF10).

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