Research Article |
Corresponding author: Chang-Lin Zhao ( fungichanglinz@163.com ) Academic editor: Bao-Kai Cui
© 2020 Jun-Zhu Chen, Chang-Lin Zhao.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Chen J-Z, Zhao C-L (2020) Morphological and molecular identification of four new resupinate species of Lyomyces (Hymenochaetales) from southern China. MycoKeys 65: 101-118. https://doi.org/10.3897/mycokeys.65.48660
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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.
Phylogeny, Schizoporaceae, taxonomy, wood-inhabiting fungi, Yunnan Province
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 (
Molecular studies on Lyomyces and related genera have been carried out recently (
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 (
The specimens studied have been deposited in the herbarium of Southwest Forestry University (
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 (
List of species, specimens and GenBank accession numbers of sequences used in this study.
Species name | Sample no. | GenBank accession no. | References |
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ITS | |||
Lyomyces allantosporus | KAS-GEL 4933 | KY800401 |
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FR 0249548 | KY800397 |
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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 |
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UC 2022841 | KP814310 |
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Lyomyces erastii | MA-Fungi 34336 | JX857800 |
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YG 022 | MF382992 |
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Lyomyces griseliniae | KHL 12971 | DQ873651 |
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Lyomyces juniperi | KAS-GEL 4940 | DQ340316 |
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FR 0261086 | KY081799 |
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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 |
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KAS-GEL 4908 | KY800400 |
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Lyomyces microfasciculatus | CLZhao 4626 | MK343568 | Present study |
CLZhao 5109 | MN954311 | Present study | |
TNM F 24757 | JN129976 |
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Lyomyces organensis | MSK 7247 | KY800403 |
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Lyomyces orientalis | KAS-GEL 3376 | DQ340325 |
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KAS-GEL 3400 | DQ340326 |
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Lyomyces pruni | Ryberg 021018 | DQ873624 |
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Lyomyces sambuci | 80 SAMHYP | JX857721 |
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83 SAMHYP | JX857720 |
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Lyomyces vietnamensis | TNM F 9073 | JX175044 |
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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 |
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Xylodon asperus | UC 2023169 | KP814365 |
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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 (
Maximum parsimony (MP), Maximum Likelihood (ML) and Bayesian Inference (BI) analyses were applied to the ITS dataset sequences. Approaches to phylogenetic analyses followed
Sequences were also analyzed using Maximum Likelihood (ML) ML analysis was conducted with RAxML-HPC2 through the Cipres Science Gateway (www.phylo.org;
MrModeltest 2.3 (
The ITS dataset (Fig.
The phylogenetic tree (Fig.
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.
China. Yunnan Province: Puer, Zhenyuan County, Heping Town, Ailaoshan National Nature Reserve, on dead bamboo, 11 January 2018, CLZhao 4831 (
The epithet bambusinus (Lat.): refers to the occurrence on bamboo.
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.
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.
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.
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).
On dead bamboo, causing a white rot. China.
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 (
China. Yunnan Province: Jingdong County, Taizhong Town, Ailaoshan Ecological Station, on fallen branch of angiosperm, 23 August 2018, CLZhao 8295 (
The epithet cremeus (Lat.): refers to the cream hymenial surface.
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.
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.
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.
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).
Lignicolous, causing a white rot. China.
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 (
China. Yunnan Province: Puer, Jingdong County, Taizhong Town, Ailaoshan National Nature Reserve, on fallen branch of angiosperm, 24 August 2018, CLZhao 8605 (holotype in
The epithet macrosporus (Lat.): refers to the larger basidiospores.
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.
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.
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.
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).
Lignicolous, causing a white rot. China.
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 (
China. Yunnan Province: Puer, Jingdong County, Wuliangshan National Nature Reserve, on fallen branch of angiosperm, 5 October 2017, CLZhao 4167 (
The epithet wuliangshanensis (Lat.): refers to the locality (Wuliangshan) of the type specimens.
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.
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.
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.
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).
Lignicolous, causing a white rot. China.
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 (
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,
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 (
Hyphodontia s.l. is an extensively studied group of Hymenochaetales (
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).