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Research Article
Molecular phylogeny and morphology reveal four new wood-inhabiting fungi of Asterostroma and Radulomyces (Basidiomycota) from Southwestern China
expand article infoJunhong Dong‡, Qiaohua Deng‡, Minglan Chen‡, Daxiang Chen§, Chunqin Zhou|, Changlin Zhao‡
‡ Southwest Forestry University, Kunming, China
§ Tongbiguan Provincial Nature Reserve, Mangshi, China
| Management and Conservation Bureau, Yunnan Wumeng Mountain National Nature Reserve, Zhaotong, China
¶ Zhaotong University, Zhaotong, China

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

Academic editor: Samantha C. Karunarathna
© 2025 Junhong Dong, Qiaohua Deng, Minglan Chen, Daxiang Chen, Chunqin Zhou, Changlin Zhao.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation: Dong J, Deng Q, Chen M, Chen D, Zhou C, Zhao C (2025) Molecular phylogeny and morphology reveal four new wood-inhabiting fungi of Asterostroma and Radulomyces (Basidiomycota) from Southwestern China. MycoKeys 112: 35-58. https://doi.org/10.3897/mycokeys.112.137098
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Abstract

In the ecosystem, wood-inhabiting fungi play an indispensable role in wood degradation and the cycle of substances. They are regarded as the “key player” in the process of wood decomposition because of their ability to produce various enzymes that break down woody lignin, cellulose, and hemicellulose. In this study, four new wood-inhabiting fungal species, Asterostroma paramuscicola, Radulomyces bambusinus, R. fissuratus, and R. sinensis, were collected from southwestern China and were proposed based on the morphological and molecular evidence. Asterostroma paramuscicola is characterised by the felted-membranous to pellicular basidiomata with pinkish to slightly salmon-buff, a smooth hymenial surface, a monomitic hyphal system, and generative hyphae bearing simple-septate and subglobose, thin-walled, echinulate basidiospores measuring as 8–8.8 × 7–8 µm. Radulomyces bambusinus is characterised by the resupinate basidiomata with pinkish-white to pink, a tuberculate hymenial surface, a monomitic hyphal system and generative hyphae bearing clamp connections, and subglobose, slightly thick-walled, smooth basidiospores measuring as 6–7.5 × 5.5–7.3 µm. Radulomyces fissuratus is characterised by the coriaceous basidiomata with grey to grey-buff, a tuberculate hymenial surface, a monomitic hyphal system and generative hyphae bearing clamp connections, and globose, slightly thick-walled, smooth basidiospores measuring as 7–9.5 × 6.5–8.5 µm. Radulomyces sinensis is characterised by the coriaceous basidiomata with straw to cinnamon to ocherous, a tuberculate hymenial surface, a monomitic hyphal system and generative hyphae bearing clamp connections, and broadly ellipsoid, slightly thick-walled, smooth basidiospores measuring as 7.5–9 × 6.2–7.5 µm. Sequences of the internal transcribed spacer (ITS) and large subunit (nrLSU) markers of the studied samples were generated, and phylogenetic analyses were performed with maximum likelihood, maximum parsimony, and Bayesian inference methods. Phylogenetic analyses of ITS+nrLSU nuclear RNA gene regions showed that four new species were assigned to the genera Asterostroma and Radulomyces. The phylogenetic tree inferred from the ITS sequences revealed that A. paramuscicola was closely associated with A. macrosporum and A. muscicola. Based on the ITS sequences, the topology showed that Radulomyces bambusinus was retrieved as a sister to R. zixishanensis. The taxon R. fissuratus forms a monophyletic lineage. The other one species, R. sinensis, was closely associated with R. molaris and R. yunnanensis.

Key words

Biodiversity, molecular systematics, new species, taxonomy, wood-decaying fungi, Yunnan Province

Introduction

The term “eukaryote” refers to cell structure and means that an organism’s genetic information is housed inside a structure called a nucleus (Money 2016). Fungi are eukaryotic microorganisms that play key ecological roles as decomposers, mutualists, or pathogens (Hyde et al. 2021). Fungi are a distinct, diverse, and ecologically important branch of the tree of life (James et al. 2020). Classification of the fungi has proven challenging due to the small number of known as compared to estimated species and a lack of sequence data for many extant taxa (Hyde et al. 2021). The phylum Basidiomycota constitutes a major group of the kingdom fungi and is second in species numbers to the phylum Ascomycota (Wijayawardene et al. 2018). Wood-inhabiting fungi have important industrial, medicinal, edible, and economic values, and a small number of them contain toxic metabolites (Niego et al. 2023).

The genus Asterostroma Massee, belonging to the family Peniophoraceae (Russulales, Basidiomycota), is typified by A. apalum (Berk. & Broome) Massee, and it is characterised by the resupinate, effused, brittle, loosely adnate, membranaceous to pellic basidiomata; a cream to ochraceous, smooth hymenial surface; a dimitic hyphal system with simple-septate on generative hyphae and dextrinoid asterosetae; present gloeocystidia; utriform basidia with 4-sterigmata and a basal simple septum; and subglobose to ellipsoid, smooth or tuberculate, amyloid basidiospores (Massee 1889; Bernicchia and Gorjón 2010). Based on the MycoBank database (http://www.mycobank.org, accessed on 9 November 2024) and the Index Fungorum (http://www.indexfungorum.org, accessed on 9 November 2024), the genus Asterostroma has registered 41 specific and infraspecific names, and six have been recorded from China (Liu et al. 2017; Deng et al. 2024; Dong et al. 2024a; Zhou et al. 2024).

The genus Radulomyces M.P. Christ., belonging to the family Radulomycetaceae (Agaricales, Basidiomycota), is typified by R. confluens (Fr.) M.P. Christ., and it is characterised by the resupinate, adnate, effused, ceraceous, hygrophanous basidiomata, smooth, tuberculate, odontioid to raduloid hymenial surface, a monomitic hyphal system with clamp connections on generative hyphae, clavate, sinuous basidia with 4-sterigmata and a basal clamp connection, with abundant oil drops, and ellipsoid to globose, smooth or minutely ornamented (spore dimorphism occurs in some species), slightly thick-walled, acyanophilous basidiospores (Christiansen 1960; Bernicchia and Gorjón 2010). Based on the MycoBank database (http://www.mycobank.org, accessed on 9 November 2024) and the Index Fungorum (http://www.indexfungorum.org, accessed on 9 November 2024), the genus Radulomyces has registered 30 specific and infraspecific names, and three have been found from China (Dong et al. 2024b).

Based on the morphological characteristics of asterosetae, the genus Asterostroma was placed in the family Lachnocladiaceae D.A. Reid (Parmasto 1971; Hallenberg and Eriksson 1985). Later, Asterostroma belonged to the russuloid lineage and was located in the family Peniophoraceae Lotsy in their phylogenetic analysis of ITS+nrLSU (Larsson and Larsson 2003; Miller et al. 2006; Larsson 2007; Liu et al. 2017). According to the morphological characteristics of the basidiospores in Asterostroma (Parmasto 1971; Boidin et al. 1997), this genus was divided into two subgenera: Austroasterostroma Parmasto (smooth and amyloid basidiospores) and Asterostroma ornamented and amyloid basidiospores (Liu et al. 2017; Zhou et al. 2024). Recently, phylogenetic analyses on Asterostroma from China based on ITS+nrLSU sequences and morphology have identified three new species: A. rhizomorpharum H.M. Zhou & C.L. Zhao, A. roseoalbum J.H. Dong & C.L. Zhao and A. yunnanense Y.L. Deng & C.L. Zhao were described (Deng et al. 2024; Dong et al. 2024a; Zhou et al. 2024).

The placement of two genera, Aphanobasidium Jülich and Radulomyces, was previously located in the family Pterulaceae Corner by the phylogenetic reconstructions of corticioid taxa (Larsson et al. 2004; Larsson 2007). Phylogenetically, Radulomyces belonged in the Pterulaceae (Agaricales) and was most closely related to Radulotubus Y.C. Dai, S.H. He & C.L. Zhao and Aphanobasidium (Zhao et al. 2016). These three genera form a strongly supported clade sister to the Pterula-Deflexula-Pterulicium-Merulicium-Coronicium clade based on sequence analyses of ITS+nrLSU (Zhao et al. 2016; Wang et al. 2018). Radulomycetaceae was characterized by the combination of resupinate basidiomes, a monomitic hyphal system, and the absence of cystidia, in which the phylogenetic analyses strongly support the segregation of Radulomycetaceae from Pterulaceae (Leal-Dutra et al. 2020). Based on the phylogenetic and morphological analysis, Leal-Dutra et al. (2020) revealed that no members of the three genera (i.e., Aphanobasidium, Radulomyces, and Radulotubus) within this superclade were pteruloid (i.e., coralloid basidiomes with a dimitic hyphal system) in their morphology. They were distinct from nearly all the other members of Pterulaceae, clearly by morphology and phylogeny, and then consequently Leal-Dutra et al. (2020) proposed the new family Radulomycetaceae to accommodate the three genera. Based on the ITS+nrLSU sequence and morphological characteristics, three new species, R. hydnoides J.H. Dong & C.L. Zhao, R. yunnanensis J.H. Dong & C.L. Zhao, and R. zixishanensis J.H. Dong & C.L. Zhao, were introduced from China (Dong et al. 2024b).

During investigations on wood-inhabiting fungi in southwestern China, some specimens were collected. To clarify the placement and relationships of these specimens, we carried out a phylogenetic and taxonomic study on the genera Asterostroma and Radulomyces based on the ITS and ITS+nrLSU sequences. These specimens were assigned to the genera Asterostroma and Radulomyces. Therefore, four new species, Asterostroma paramuscicola, R. bambusinus, R. fissuratus, and R. sinensis, are proposed, based on the morphological characteristics and phylogenetic analyses.

Materials and methods

Sample collection and herbarium specimen preparation

The fresh fruiting bodies on the dead bamboo and fallen angiosperm branches were collected from Dehong, Lincang, Pu’er, and Zhaotong of Yunnan Province, China. The samples were photographed in situ, and fresh macroscopic details were recorded. Photographs were recorded by a Nikon D7100 camera. All the photos were focus-stacked using Helicon Focus software. Macroscopic details and collection information (Rathnayaka et al. 2024) were taken and transported to a field station where the fruit body was dried on an electronic food dryer at 45 °C (Hu et al. 2022). Once dried, the specimens were sealed in an envelope and zip-lock plastic bags and labeled. The dried specimens were deposited in the herbarium of the Southwest Forestry University (SWFC), Kunming, Yunnan Province, China.

Morphology

The macromorphological descriptions were based on field notes and photos captured in the field and lab. The color terminology follows Petersen (1996). The micromorphological data were obtained from the dried specimens after observation under a light microscope with a magnification of 10 × 100 oil. Sections were mounted in 5% KOH and 2% phloxine B (C20H2Br4Cl4Na2O5), and other reagents were also used, including cotton blue and Melzer’s reagent to observe micromorphology following Wu et al. (2022). To show the variation in spore sizes, 5% of measurements were excluded from each end of the range and shown in parentheses. At least thirty basidiospores from each specimen were measured. Stalks were excluded from basidia measurements, and the hilar appendage was excluded from basidiospore measurements. The following abbreviations are used: KOH = 5% potassium hydroxide water solution, CB– = acyanophilous, IKI– = both inamyloid and non-dextrinoid, 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, Qm represented the average Q of basidiospores measured ± standard deviation, 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, China) was used to extract genomic DNA from the dried specimens according to the manufacturer’s instructions. The nuclear ribosomal internal transcribed spacer (ITS) region was amplified with ITS5 and ITS4 primers (White et al. 1990). The nuclear large subunit (nrLSU) region was amplified with the LR0R and LR7 primer pair (Vilgalys and Hester 1990; Rehner and Samuels 1994). The PCR procedure for ITS was as follows: initial denaturation at 95 °C for 3 min, followed by 35 cycles at 94 °C for 40 s, 58 °C for 45 s, and 72 °C for 1 min, and a final extension of 72 °C for 10 min. The PCR procedure for nrLSU was as follows: initial denaturation at 94 °C for 1 min, followed by 35 cycles at 94 °C for 30 s, 48 °C for 1 min, and 72 °C for 1.5 min, and a final extension of 72 °C for 10 min. The PCR products were purified and sequenced at Kunming Tsingke Biological Technology Limited Company (Yunnan Province, P.R. China). The 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. [New species is shown in bold; * type material].

Species Name Sample No. GenBank Accession No. Country References
ITS nrLSU
Aphanobasidium pseudotsugae HHB-822 GU187509 GU187567 USA Larsson (2007)
Aphanobasidium pseudotsugae UC 2023153 KP814353 AY586696 Sweden Larsson (2007)
Asterostroma andinum He 20120921–17 — KY263874 China Liu et al. (2017)
Asterostroma andinum HHB-8546-sp — AF518600 USA Hibbett and Binder (2002)
Asterostroma bambusicola He 4128 KY263864 — Thailand Liu et al. (2017)
Asterostroma bambusicola He 4132 KY263865 KY263871 Thailand Liu et al. (2017)
Asterostroma cervicolor He 2314 KY263860 KY263868 China Liu et al. (2017)
Asterostroma cervicolor He 4020 KY263859 KY263869 China Liu et al. (2017)
Asterostroma cervicolor KHL 9239 AF506408 AF506408 Puerto Rico Larsson and Larsson (2003)
Asterostroma cervicolor TMI 21362 AB439560 — Japan Larsson and Larsson (2003)
Asterostroma laxum EL 33-99 AF506410 AF506410 Estonia Larsson and Larsson (2003)
Asterostroma macrosporum TMI 25696 AB439544 — Japan Suhara et al. (2010b)
Asterostroma macrosporum TMI 25697 AB439545 — Japan Suhara et al. (2010b)
Asterostroma medium HFRG_EJ220212_2_FRDBI 23891920 OQ133615 — UK Deng et al. (2024)
Asterostroma medium HFRG_EJ210127_2FRDBI 18772203 OL828779 — UK Deng et al. (2024)
Asterostroma muscicola He 4106 KY263861 KY263873 Thailand Liu et al. (2017)
Asterostroma muscicola He 20121104-1 KY263862 KY263872 China Liu et al. (2017)
Asterostroma muscicola TUMH 10017 AB439552 — Japan Suhara et al. (2010b)
Asterostroma ochroleucum HB 9/89 — AF323737 Germany Wagner (2001)
Asterostroma paramuscicola CLZhao 8594 PP392895 PQ306584 China Present study
Asterostroma rhizomorpharum CLZhao 31212 OR672732 OR879302 China Zhou et al. (2024)
Asterostroma rhizomorpharum CLZhao 31216 OR672733 — China Zhou et al. (2024)
Asterostroma vararioides He 4136 KY263866 — Thailand Liu et al. (2017)
Asterostroma vararioides He 4140 KY263867 KY263870 Thailand Liu et al. (2017)
Asterostroma yunnanense CLZhao 22781 OR048809 OR506285 China Deng et al. (2024)
Asterostroma yunnanense CLZhao 22786 OR048811 OR506286 China Deng et al. (2024)
Confertobasidium olivaceoalbum FP 90196 AF511648 AF511648 USA Larsson and Larsson (2003)
Dichostereum durum FG 1985 AF506429 AF506429 France Larsson and Larsson (2003)
Dichostereum effuscatum GG 930915 AF506390 AF506390 France Larsson and Larsson (2003)
Gloiothele lactescens EL 8-98 AF506453 AF506453 Sweden Larsson and Larsson (2003)
Gloiothele lamellosa KHL 11031 AF506454 AF506454 USA Larsson and Larsson (2003)
Merulicium fusisporum Hjm s.n. EU118647 EU118647 Sweden Larsson (2007)
Peniophora cinerea NH 9808/1788 AF506424 AF506424 Spain Larsson and Larsson (2003)
Peniophora incarnata NH 10271/1909 AF506425 AF506425 Denmark Larsson and Larsson (2003)
Pterula echo AFTOL-ID711 DQ494693 AY629315 USA Larsson and Larsson (2003)
Radulomyces bambusinus CLZhao 35383 * PQ306582 PQ306589 China Present study
Radulomyces bambusinus CLZhao 35384 PQ306583 PQ306590 China Present study
Radulomyces confluens Cui 5977 KU535661 KU535669 China Wang et al. (2018)
Radulomyces confluens He 2224 KU535662 KU535670 China Wang et al. (2018)
Radulomyces copelandii Dai 15061 KU535664 KU535672 China Wang et al. (2018)
Radulomyces copelandii Wu 9606-5 KU535663 KU535671 China Wang et al. (2018)
Radulomyces fissuratus CLZhao 29670 * PQ306579 PQ306586 China Present study
Radulomyces fissuratus CLZhao 29695 PQ306580 PQ306587 China Present study
Radulomyces fissuratus CLZhao 29713 PQ306581 PQ306588 China Present study
Radulomyces hydnoides CLZhao 21632 OR096184 OR449914 China Dong et al. (2024b)
Radulomyces hydnoides CLZhao 21668 OR096185 OR449915 China Dong et al. (2024b)
Radulomyces molaris ARAN-Fungi 2003 — MT232311 Spain Olariaga et al. (2020)
Radulomyces molaris ML0499 AY463459 AY586705 Sweden Larsson et al. (2004)
Radulomyces paumanokensis IMG 5985-16 MG050100 MG050110 Spain Wang et al. (2018)
Radulomyces rickii JK 951007 — AY586706 Sweden Larsson et al. (2004)
Radulomyces rickii G1066 — MK278543 Hungary Varga et al. (2019)
Radulomyces sinensis CLZhao 25667 * PQ306578 PQ306585 China Present study
Radulomyces yunnanensis CLZhao 1262 OR096191 OR449917 China Dong et al. (2024b)
Radulomyces yunnanensis CLZhao 7364 OR096192 OR449918 China Dong et al. (2024b)
Radulomyces zixishanensis CLZhao 21127 ON033887 OR449922 China Dong et al. (2024b)
Radulotubus resupinatus Cui 8383 KU535660 KU535668 China Zhao et al. (2016)
Radulotubus resupinatus Cui 8462 KU535657 KU535665 China Zhao et al. (2016)
Scytinostroma portentosum EL 11-99 AF506470 AF506470 Sweden Larsson and Larsson (2003)
Vararia gallica CBS 656.81 — AF323742 France Wagner (2001)
Vararia ochroleuca CBS 465.61 — AF323743 France Wagner (2001)
Vesiculomyces citrinus EL 53-97 AF506486 AF506486 Sweden Larsson and Larsson (2003)

The sequences were aligned in MAFFT v. 7 (Katoh et al. 2019) using the G-INS-i strategy. The alignment was adjusted manually using AliView v. 1.27 (Larsson 2014). The dataset was aligned first, and then the sequences of ITS+nrLSU were combined with Mesquite v. 3.51. The combined ITS+nrLSU sequences and ITS datasets were used to infer the position of the new species and related species. The sequences of Confertobasidium olivaceoalbum (Bourdot & Galzin) Jülich were retrieved from GenBank and used as outgroup taxa in the ITS+nrLSU analysis (Fig. 1) in the family Peniophoraceae; Scytinostroma portentosum (Berk. & M.A. Curtis) Donk was selected as the outgroup taxon for the ITS analysis (Fig. 2) in the genus Asterostroma (Deng et al. 2024). The sequences of Pterula echo D.J. McLaughlin & E.G. McLaughlin and Merulicium fusisporum (Romell) J. Erikss. & Ryvarden were selected as the outgroup taxon for the ITS+nrLSU analysis (Fig. 3) of the family Radulomycetaceae; Radulotubus resupinatus Y.C. Dai, S.H. He & C.L. Zhao was selected as outgroup taxa in the ITS analysis (Fig. 4) in the genus Radulomyces (Zhao et al. 2016).

Figure 1. 

Maximum parsimony strict consensus tree illustrating the phylogeny of Asterostroma and related genera in the family Peniophoraceae based on ITS+nrLSU sequences. Branches are labelled with maximum likelihood bootstrap value ≥ 70%, parsimony bootstrap value ≥ 50%, and Bayesian posterior probabilities ≥ 0.95.

Figure 2. 

Maximum parsimony strict consensus tree illustrating the phylogeny of Asterostroma paramuscicola and related species in the genus Asterostroma based on ITS sequences. Branches are labelled with maximum likelihood bootstrap value ≥ 70%, parsimony bootstrap value ≥ 50%, and Bayesian posterior probabilities ≥ 0.95.

Figure 3. 

Maximum parsimony strict consensus tree illustrating the phylogeny of Radulomyces and related genera in the family Radulomycetaceae based on ITS+nrLSU sequences. Branches are labelled with maximum likelihood bootstrap value ≥ 70%, parsimony bootstrap value ≥ 50%, and Bayesian posterior probabilities ≥ 0.95.

Figure 4. 

Maximum parsimony strict consensus tree illustrating the phylogeny of three new and related species in the genus Radulomyces based on ITS sequences. Branches are labelled with maximum likelihood bootstrap value ≥ 70%, parsimony bootstrap value ≥ 50%, and Bayesian posterior probabilities ≥ 0.95.

Maximum parsimony (MP), maximum likelihood (ML), and Bayesian inference (BI) analyses were applied to the combined three datasets following a previous study (Zhao and Wu 2017), and the tree construction procedure was performed in PAUP* v. 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 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 the homoplasy index (HI) were calculated for each maximum parsimonious tree generated. The multiple sequence alignment was also analysed using Maximum Likelihood (ML) in RAxML-HPC2 on XSEDE v. 8.2.8 with default parameters (Miller et al. 2012). Branch support (BS) for ML analysis was determined by 1,000 bootstrap replicates.

jModelTest v. 2 (Darriba et al. 2012) was used to determine the best-fit evolutionary model for each data set for Bayesian inference (BI), which was performed using MrBayes 3.2.7a (Ronquist et al. 2012). The first one-fourth of all generations was discarded as burn-in. The majority rule consensus tree of all remaining trees was calculated. Branches were considered as significantly supported if they received maximum likelihood bootstrap value (BS) ≥ 70%, maximum parsimony bootstrap value (BT) ≥ 70%, or Bayesian posterior probabilities (BPP) ≥ 0.95.

Results

Molecular phylogeny

The aligned dataset comprised 34 specimens representing 23 species. Four Markov chains were run for two runs from random starting trees, each for one million generations for the combined ITS+nrLSU (Fig. 1) data set with trees and parameters sampled every 1000 generations. The dataset had an aligned length of 2206 characters, of which 1453 characters are constant, 276 are variable and parsimony uninformative, and 477 are informative. Maximum parsimony analysis yielded nine equally parsimonious trees (TL = 1735, CI = 0.6317, HI = 0.3683, RI = 0.6751, and RC = 0.4265). The best model for the ITS+nrLSU dataset, estimated and applied in the Bayesian analysis, was GTR+I+G. Both Bayesian analysis and ML analysis resulted in a similar topology to MP analysis, with an average standard deviation of split frequencies = 0.004237 (BI), and the effective sample size (ESS) for Bayesian analysis across the two runs is double the average ESS (avg. ESS) = 505.

The aligned dataset comprised 20 specimens representing 11 species. Four Markov chains were run for two runs from random starting trees, each for 0.5 million generations for the ITS (Fig. 2) data set with trees and parameters sampled every 1000 generations. The dataset had an aligned length of 620 characters, of which 384 characters are constant, 51 are variable and parsimony uninformative, and 185 are informative. Maximum parsimony analysis yielded one equally parsimonious tree (TL = 412, CI = 0.7694, HI = 0.2306, RI = 0.8450, and RC = 0.6502). The best model for the ITS dataset, estimated and applied in the Bayesian analysis, was HKY+G. Both Bayesian analysis and ML analysis resulted in a similar topology to MP analysis with an average standard deviation of split frequencies = 0.004683 (BI), and the effective sample size (ESS) for Bayesian analysis across the two runs is double the average ESS (avg. ESS) = 435.

The aligned dataset comprised 26 specimens representing 15 species. Four Markov chains were run for two runs from random starting trees, each for one million generations for the combined ITS+nrLSU (Fig. 3) data set with trees and parameters sampled every 1000 generations. The dataset had an aligned length of 2115 characters, of which 1699 characters are constant, 132 are variable and parsimony uninformative, and 284 are informative. Maximum parsimony analysis yielded 143 equally parsimonious trees (TL = 711, CI = 0.7496, HI = 0.2504, RI = 0.7623, and RC = 0.5715). The best model for the ITS+nrLSU dataset, estimated and applied in the Bayesian analysis, was GTR+I+G. Both Bayesian analysis and ML analysis resulted in a similar topology to MP analysis, with an average standard deviation of split frequencies = 0.008051 (BI), and the effective sample size (ESS) for Bayesian analysis across the two runs is double the average ESS (avg. ESS) = 535.5.

The aligned dataset comprised 21 specimens representing 12 species. Four Markov chains were run for two runs from random starting trees, each for 0.5 million generations for the ITS (Fig. 4) data set with trees and parameters sampled every 1,000 generations. The dataset had an aligned length of 671 characters, of which 444 characters are constant, 74 are variable and parsimony uninformative, and 153 are informative. Maximum parsimony analysis yielded ten equally parsimonious trees (TL = 373, CI = 0.7587, HI = 0.2413, RI = 0.8109, and RC = 0.6153). The best model for the ITS dataset, estimated and applied in the Bayesian analysis, was HKY+G. Both Bayesian analysis and ML analysis resulted in a similar topology to MP analysis, with an average standard deviation of split frequencies = 0.006832 (BI), and the effective sample size (ESS) for Bayesian analysis across the two runs is double the average ESS (avg. ESS) = 346.5.

The phylogram, based on the combined ITS+nrLSU sequences (Fig. 1) analysis, showed that the new species, Asterostroma paramuscicola, was assigned to the genus Asterostroma within the family Peniophoraceae. The phylogenetic tree, based on ITS sequences (Fig. 2), revealed that A. paramuscicola was closely associated with A. macrosporum N. Maek. & Suhara. and A. muscicola (Berk. & M.A. Curtis) Massee. The phylogram, based on the combined ITS+nrLSU sequences (Fig. 3) analysis, showed that three new species, Radulomyces bambusinus, R. fissuratus, and R. sinensis, were assigned to the genus Radulomyces within the family Radulomycetaceae. The phylogenetic tree, based on ITS sequences (Fig. 4), revealed that R. bambusinus was retrieved as a sister to R. zixishanensis. The taxon R. fissuratus forms a monophyletic lineage. The other species, R. sinensis, was closely associated with R. molaris (Chaillet ex Fr.) M.P. Christ. and R. yunnanensis J.H. Dong & C.L. Zhao.

Taxonomy

Asterostroma paramuscicola J.H. Dong & C.L. Zhao, sp. nov.

MycoBank No: 855659
Figs 5A, 6, 7

Holotype

China • Yunnan Province, Pu’er, Jingdong County, Taizhong Town, Ailaoshan Ecological Station, GPS coordinates 24°31'N, 101°02'E, altitude 2400 m asl., a dead angiosperm tree, leg. C.L. Zhao, 24 August 2018, CLZhao 8594 (SWFC).

Figure 5. 

Sections of hymenium of Asterostroma and Radulomyces A Asterostroma paramuscicola (holotype, CLZhao 8594) B Radulomyces bambusinus (holotype, CLZhao 35383) C Radulomyces fissuratus (holotype, CLZhao 26970) D Radulomyces sinensis (holotype, CLZhao 25667). 10 × 100 Oil.

Etymology

paramuscicola (Lat.): referring to its close phylogenetic relationship with A. muscicola.

Basidiomata

Annual, resupinate, felted-membranous to pellicular, soft, without odour and taste when fresh, becoming coriaceous upon drying, up to 10 cm long, 5 cm wide, 50–150 µm thick. Hymenial surface smooth, pinkish when fresh, turning to pinkish to slightly salmon-buff upon drying, rhizomorphic. Sterile margin thin, cream to buff, fimbriate, up to 1 mm wide.

Figure 6. 

Basidiomata of Asterostroma paramuscicola in general and detailed views (holotype, CLZhao 8594).

Hyphal system

Dimitic, generative hyphae scattered, simple-septate, colorless, thin-walled, occasionally branched, 2–3.5 µm in diameter, IKI–, CB–, tissues unchanged in KOH. Asterosetae in subiculum abundant, predominant, yellowish brown, thick-walled, regularly star-shaped, 4–6 µm in diameter, weakly dextrinoid, rays up to 75 µm long, with acute tips, CB–, and tissues unchanged in KOH.

Hymenium

Asterohyphidia in hymenium similar to asterosetae in subiculum, but smaller and less regularly shaped, 2–3.5 µm in diameter, rays up to 25 µm long, usually bifurcated at tips. Gloeocystidia subcylindrical to fusiform, thin-walled, with a basal simple septum, 45–83.5 × 9–21.5 µm; cystidioles absent. Basidia subcylindrical, colorless, with four sterigmata and a basal simple septum, 47.5–58 × 7.5–9.5 µm; basidioles dominant, similar to basidia in shape, but slightly smaller.

Figure 7. 

Microscopic structures of Asterostroma paramuscicola (holotype, CLZhao 8594) A basidiospores B basidia C basidioles D gloeocystidia E generative hyphae F asterohyphidia from hymenium G asterosetae from subiculum. Scale bars: 10 µm (A–G).

Basidiospores

Subglobose, with a distinct apiculus, spines conical, 2–3 µm long, colorless, thin-walled, echinulate, amyloid, with one guttula, CB–, (7.5–)8–8.8(–9) × (6.6–)7–8(–8.5) µm, L = 8.39 µm, W = 7.65 µm, Q = 1.02–1.22, Qm = 1.10 ± 0.06 (n = 30/1).

Radulomyces bambusinus J.H. Dong & C.L. Zhao, sp. nov.

MycoBank No: 855660
Figs 5B, 8, 9

Holotype

China • Yunnan Province, Zhaotong, Daguan County, Wumengshan National Nature Reserve, GPS coordinates 27°46'N, 103°52'E, altitude 2200 m asl., on the dead bamboo, leg. C.L. Zhao, 6 November 2023, CLZhao 35383 (SWFC).

Figure 8. 

Basidiomata of Radulomyces bambusinus in general and detailed views A, B CLZhao 35383 (holotype) C, D CLZhao 35384.

Etymology

bambusinus (Lat.): referring to the type species growing on bamboo.

Basidiomata

Annual, resupinate, adnate, soft membranous, without odour or taste, becoming coriaceous upon drying, up to 5 cm long, 2 cm wide, 50–100 µm thick. Hymenial surface tuberculate, cream to pinkish when fresh, turning to pinkish-white to pink upon drying. Sterile margin cream to slightly pinkish, thinning out, up to 1 mm wide.

Figure 9. 

Microscopic structures of Radulomyces bambusinus (holotype, CLZhao 35383) A basidiospores B basidia C basidioles D part of the vertical section of hymenium. Scale bars: 10 µm (A–D).

Hyphal system

Monomitic, generative hyphae with clamp connections, colorless, thin-walled, branched, interwoven, 2.5–3.5 µm in diameter; IKI–, CB–, tissues unchanged in KOH.

Hymenium

Cystidia and cystidioles absent. Basidia clavate to barrelled, with 4 sterigmata and a basal clamp connection, occasionally constricted in the middle, 23.5–40.5 × 7.5–10.5 µm; basidioles dominant, similar to basidia in shape, but slightly smaller.

Basidiospores

Subglobose, slightly thick-walled, smooth, colorless, CB+, (5.7–)6–7.5(–8) × 5.5–7.3(–7.5) µm, L = 6.76 µm, W = 6.43 µm, Q = 1.01–1.18, Qm = 1.05 ± 0.05 (n = 60/2).

Additional specimen (paratype) examined

China • Yunnan Province, Zhaotong, Daguan County, Wumengshan National Nature Reserve, GPS coordinates 27°46'N, 103°52'E, altitude 2200 m asl., on the dead bamboo, leg. C.L. Zhao, 6 November 2023, CLZhao 35384 (SWFC).

Radulomyces fissuratus J.H. Dong & C.L. Zhao, sp. nov.

MycoBank No: 855661
Figs 5C, 10, 11

Holotype

China • Yunnan Province, Dehong, Yingjiang County, Tongbiguan Provincial Nature Reserve, GPS coordinates 23°48'N, 97°38'E, altitude 1000 m asl., on the fallen branch of angiosperm, leg. C.L. Zhao, 17 July 2023, CLZhao 29670 (SWFC).

Figure 10. 

Basidiomata of Radulomyces fissuratus in general and detailed views A, B, CLZhao 29670 (holotype) C, D CLZhao 29713 E, F CLZhao 29695.

Etymology

fissuratus (Lat.): referring to the cracked hymenial surface of the type specimen.

Basidiomata

Annual, resupinate, adnate, membranaceous, without odour or taste, becoming hard coriaceous upon drying, up to 30 cm long, 2 cm wide, 50–150 µm thick. Hymenial surface tuberculate, cream when fresh, turning grey to grey-buff upon drying. Sterile margin cream, thinning out, up to 1 mm wide.

Hyphal system

Monomitic, generative hyphae with clamp connections, colorless, thin-walled, branched, interwoven, 2–3.5 µm in diameter; IKI–, CB–, tissues unchanged in KOH.

Figure 11. 

Microscopic structures of Radulomyces fissuratus (holotype, CLZhao 29670) A basidiospores B basidia C basidioles D part of the vertical section of hymenium. Scale bars: 10 µm (A–D).

Hymenium

Cystidia and cystidioles absent. Basidia clavate to barrelled, with 4 sterigmata and a basal clamp connection, 21.5–32.5 × 8.5–10.5 µm; basidioles dominant, similar to basidia in shape, but slightly smaller.

Basidiospores

Globose, slightly thick-walled, smooth, colorless, CB+, (6.5–)7–9(–9.5) × (6.2–)6.5–8.5(–8.8) µm, L = 8.07 µm, W = 7.76 µm, Q = 1.01–1.06, Qm = 1.04 ± 0.03 (n = 90/3).

Additional specimens (paratypes) examined

China • Yunnan Province, Dehong, Yingjiang County, Tongbiguan Provincial Nature Reserve, GPS coordinates 23°48'N, 97°38'E, altitude 1000 m asl., on the fallen branch of angiosperm, leg. C.L. Zhao, 17 July 2023, CLZhao 29695; CLZhao 29713 (SWFC).

Radulomyces sinensis J.H. Dong & C.L. Zhao, sp. nov.

MycoBank No: 855662
Figs 5D, 12, 13

Holotype

China • Yunnan Province, Lincang, Fengqing County, Yaojie Town, Xingyuan Village, GPS coordinates 24°58'N, 99°92'E, altitude 1600 m asl., on the fallen branch of angiosperm, leg. C.L. Zhao, 22 October 2022, CLZhao 25667 (SWFC).

Figure 12. 

Basidiomata of Radulomyces sinensis in general and detailed views (holotype, CLZhao 25667).

Etymology

sinensis (Lat.): referring to the locality (China) of the type specimen.

Basidiomata

Annual, resupinate, adnate, soft coriaceous, without odour or taste, becoming hard coriaceous upon drying, up to 20 cm long, 2 cm wide, 100–150 µm thick. Hymenial surface tuberculate, buff to slightly straw when fresh, turning to straw to cinnamon to ocherous upon drying. Sterile margin cream to slightly straw, abrupt, up to 1 mm wide.

Figure 13. 

Microscopic structures of Radulomyces sinensis (holotype, CLZhao 25667) A basidiospores B basidia C basidioles D part of the vertical section of hymenium. Scale bars: 10 µm (A–D).

Hyphal system

Monomitic, generative hyphae with clamp connections, colorless, thin-walled, branched, interwoven, 2.5–3.5 µm in diameter; IKI–, CB–, tissues unchanged in KOH.

Hymenium

Cystidia and cystidioles absent. Basidia clavate, with 4 sterigmata and a basal clamp connection, with a median constriction, 35–41.5 × 7.5–9.5 µm; basidioles dominant, similar to basidia in shape, but slightly smaller.

Basidiospores

Broadly ellipsoid, slightly thick-walled, smooth, colorless, some with guttulae, CB+, 7.5–9(–9.5) × (5.8–)6.2–7.5(–7.7) µm, L = 8.42 µm, W = 6.88 µm, Q = 1.10–1.33, Qm = 1.23 ± 0.08 (n = 30/1).

Discussion

In the present study, four new species, Asterostroma paramuscicola, Radulomyces bambusinus, R. fissuratus, and R. sinensis, are described based on phylogenetic analyses and morphological characteristics.

Asterostroma is a monophyletic genus in our phylogenetic analysis with low statistical support, in contrast to the previous study (Liu et al. 2017; Zhou et al. 2024). Seven species with ornamented basidiospores formed the section Asterostroma as A. bambusicola S.L. Liu & S.H. He, A. cervicolor (Berk. & M.A. Curtis) Massee, A. macrosporum, A. medium Bres., A. muscicola, A. ochroleucum, A. paramuscicola, A. rhizomorpharum, and A. yunnanense, while three species with smooth basidiospores formed two clades as the A. andinum Pat. clade (only A. andinum) and sect. Laevispora (A. laxum Bres. and A. vararioides S.L. Liu & S.H. He).

Phylogenetically, the phylogram based on the combined ITS+nrLSU sequences (Fig. 1) analysis showed that the new species Asterostroma paramuscicola was assigned to the genus Asterostroma within the family Peniophoraceae. The phylogenetic tree, based on ITS sequences (Fig. 2), revealed that Asterostroma paramuscicola was closely associated with A. macrosporum and A. muscicola. However, the morphological characteristics of A. macrosporum differ from Asterostroma paramuscicola by having an ochreous to fulvous hymenial surface (Suhara et al. 2010b). The species A. muscicola differentiates from A. paramuscicola by having a salmon hymenial surface, smaller basidia (18–24 × 5–6 µm), and shorter basidiospores (6–8 × 5.5–7.5 µm; Boidin et al. 1997).

The phylogram, based on the combined ITS+nrLSU sequences (Fig. 3) analysis, showed that three new species, Radulomyces bambusinus, R. fissuratus, and R. sinensis, were assigned to the genus Radulomyces within the family Radulomycetaceae. The phylogenetic tree, based on ITS sequences (Fig. 4), revealed that Radulomyces bambusinus was retrieved as a sister to R. zixishanensis. The taxon R. fissuratus forms a monophyletic lineage. The other one species, R. sinensis, was closely associated with R. molaris and R. yunnanensis. However, the morphological characteristics of R. zixishanensis differ from R. bambusinus by having a cream to slightly brown, smooth hymenial surface and ellipsoid, thin-walled, basidiospores (7–8.8 × 5.5–6.8 µm; Dong et al. 2024b). The species R. molaris differs from R. sinensis by having a yellowish to cream, hydnoid hymenial surface (Bernicchia and Gorjón 2010). The species R. yunnanensis can be distinguished from R. sinensis by having a cream, smooth hymenial surface, shorter basidia (24–35 × 7–11 µm), and thin-walled basidiospores (8.2–9.5 × 5.5–7 µm; Dong et al. 2024b).

Asterostroma paramuscicola, A. rhizomorpharum, and A. yunnanense are all described from China. However, A. rhizomorpharum can be distinguished from A. paramuscicola in its cream to buff, cracked hymenial surface, shorter basidia (30–45 × 5.5–8 µm), and smaller basidiospores (5.5–6.8 × 4.6–5.9 µm; Zhou et al. 2024). The species A. yunnanense can be distinguished from A. paramuscicola in its cream to salmon-buff hymenial surface, smaller basidia (31–38 × 4–5 µm), and smaller basidiospores (4.5–6 × 4–5 µm; Deng et al. 2024).

Morphologically, Asterostroma paramuscicola resembles A. bambusicola, A. boninense Suhara & N. Maek., and A. vararioides in sharing a fimbriate basidiomata and subglobose to globose basidiospores. However, A. bambusicola differs from A. paramuscicola by having a brownish yellow, grayish brown, light brown, to brownish-red hymenial surface, shorter basidia (30–45 × 5.5–8 µm; Liu et al. 2017). The species A. boninense differentiates from A. paramuscicola by having the buff to partly ochreous hymenial surface and shorter basidiospores (5.5–7.5 × 5–7.2 µm; Suhara et al. 2010a). The species A. vararioides can be distinguished from A. paramuscicola in its grayish brown, light brown, to dark brown hymenial surface and smooth, smaller basidiospores (6–7 × 5.5–6 µm; Liu et al. 2017).

Radulomyces bambusinus shares similarities with R. yunnanensis and R. zixishanensis in having a cracked hymenial surface. However, R. yunnanensis can be distinguished from R. bambusinus by its smooth hymenial surface, ellipsoid, thin-walled, shorter basidiospores (8.2–9.5 × 5.5–7 µm; Dong et al. 2024b). The species R. zixishanensis differentiates from R. bambusinus by having the smooth hymenial surface, ellipsoid, thin-walled basidiospores (7–8.8 × 5.5–6.8 µm; Dong et al. 2024b).

Radulomyces fissuratus resembles R. copelandii (Pat.) Hjortstam & Spooner, R. hydnoides, and R. paumanokensis J. Horman, Nakasone & B. Ortiz in sharing subglobose to globose, slightly thick-walled basidiospores. However, R. copelandii differs from R. fissuratus due to its white hymenial surface, smaller basidia (29–35 × 6–7 µm), and basidiospores (6.4–7 × 5.4–6.2 µm; Ginns and Millman 2011). The species R. hydnoides can be distinguished from R. fissuratus by its hydnoid hymenial surface and shorter basidia (21–34 × 8.5–12.5 µm; Dong et al. 2024b). The species R. paumanokensis differentiates from R. fissuratus by having a hydnoid hymenial surface, smaller basidia (25–31 × 5–7.5 µm), and shorter basidiospores (5.8–6.9 × 5.2–6.4 µm; Wang et al. 2018).

Radulomyces sinensis shares similarities with R. arborifer Malysheva & Zmitr., R. molaris, and R. zixishanensis in having ellipsoid basidiospores. However, R. arborifer differs from R. sinensis due to its dendroid or coralloid hymenial surface (Malysheva 2006). The species R. molaris can be distinguished from R. sinensis by its hydnoid hymenial surface (Bernicchia and Gorjón 2010). The species R. zixishanensis differentiates from R. sinensis by having the cream to slightly brown, smooth hymenial surface and thin-walled basidiospores (7–8.8 × 5.5–6.8 µm; Dong et al. 2024b).

In recent years, the wood-inhabiting fungi have been an extensively studied group of Basidiomycota, which includes a number of poroid, smooth, grandinoid, odontioid, and hydnoid basidiomata in China (Liu et al. 2023; Mao et al. 2023; Zhao et al. 2023; Dong et al. 2024b; Wang et al. 2024; Zhao et al. 2024). This paper enriches our knowledge of fungal diversity in China. We anticipate that more undescribed wood-inhabiting fungi taxa will be discovered throughout China after extensive collection combined with morphological and molecular analyses.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

The research was supported by the National Natural Science Foundation of China (Project Nos. 32170004, U2102220), the High-level Talents Program of Yunnan Province (YNQR-QNRC-2018-111), the Research Project of Key Laboratory of Forest Disaster Warning and Control in Universities of Yunnan Province (ZKJS-S-202208), and the Research Project of Yunnan Key Laboratory of Gastrodia and Fungal Symbiotic Biology (TMKF2023A03).

Author contributions

Conceptualization, C.Z.; methodology, C.Z. and J.D.; software, C.Z., J.D., Q.D., and M.C.; validation, C.Z. and J.D.; formal analysis, C.Z., J.D., and Q.D.; investigation, C.Z., J.D., D.C., and C.Z.; resources, C.Z. and J.D.; writing—original draft preparation, C.Z., J.D., Q.D., and M.C.; writing—review and editing, C.Z. and J.D.; visualization, C.Z. and J.D.; 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

Junhong Dong https://orcid.org/0000-0001-8740-0805

Qiaohua Deng https://orcid.org/0009-0002-4683-2702

Minglan Chen https://orcid.org/0009-0006-7890-2604

Daxiang Chen https://orcid.org/0009-0001-2309-1991

Chunqin Zhou https://orcid.org/0009-0004-8117-1537

Changlin Zhao https://orcid.org/0000-0002-8668-1075

Data availability

All of the data that support the findings of this study are available in the main text.

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