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. 2026 Jul 24;137:273–297. doi: 10.3897/mycokeys.137.203862

Three new species of Cantharellales and Hymenochaetales from Pinus massoniana in China

Kai-Yue Luo 1, Yu-Cheng Dai 1, Yuan Yuan 1,✉, Ying-Da Wu 1,2,✉
PMCID: PMC13428250  PMID: 42542819

Abstract

Three new wood-inhabiting fungi species, found on Pinus massoniana, are proposed, based on a combination of morphological features and molecular evidence. Botryobasidium massonianae, belonging to Cantharellales, is characterised by fluffy, pellicular basidiomata, hypochnoid hymenophoral surface, generative hyphae simple septate, basidia with four to six sterigmata and basidiospores measuring 6.3–9.2 × 2.8–3.9 µm. Peniophorella crassibasidiata, belonging to Hymenochaetales, is characterised by soft coriaceous basidiomata, thick-walled generative hyphae, the presence of three types of cystidia, cylindrical basidiospores measuring 8.4–12.4 × 3.1–4.2 µm. Peniophorella variegata, belonging to Hymenochaetales, is characterised by farinaceous basidiomata, smooth hymenophoral surface, three types of cystidia, the presence of hyphidia and basidiospores measuring 7.9–10.8 × 3.9–5.1 µm. Phylogenetic analyses of the combined ITS+nLSU dataset using Maximum Likelihood and Bayesian Inference confirmed that one new species belongs to the genus Botryobasidium (Cantharellales), the other two new species belong to the genus Peniophorella (Hymenochaetales), respectively. Phylogenetically related and morphologically similar species to these three new species are discussed.

Key words: Corticioid fungi, molecular phylogeny, taxonomy, wood-inhabiting fungi

Introduction

Wood-inhabiting fungi, mainly belonging to the class Agaricomycetes (Basidiomycota), colonise various woody substrates and play essential roles in decomposing lignin, cellulose, and hemicellulose, thereby driving matter cycles and carbon storage in forest ecosystems (Wang et al. 2013; Dong et al. 2024a, 2025; Hongsanan et al. 2025; Song et al. 2025; Wijesinghe et al. 2025; Yang et al. 2025; Zhao et al. 2025a). Despite their ecological importance and global distribution, the taxonomy of many wood-inhabiting fungal taxa remains poorly known.

The genus Botryobasidium Donk belongs to the family Botryobasidiaceae Jülich, the order Cantharellales Gäum., and is typified by B. subcoronatum (Höhn. & Litsch.) Donk. The genus is characterised by annual, resupinate basidiomata with smooth, pellicular, hypochnoid, or arachnoid hymenophores; a monomitic hyphal system; generative mostly branched at right angles; basidia with 2–8 sterigmata; smooth or ornamented basidiospores; and causing a white rot (Donk 1964; Langer et al. 2000b; Moncalvo et al. 2006; Xiong et al. 2009; Zhang et al. 2025). Many asexual morph generic names, such as Acladium Link, Allescheriella Henn., Alysidium Kunze, Haplotrichum Link, Neoacladium P.N. Singh & S.K. Singh, Physospora Fr., and Sporocephalium Chevall., are congeneric with Botryobasidium, and were re-combined in Botryobasidium (Stalpers et al. 2021). Recent phylogenetic studies based on ITS data or ITS + nLSU data analyses demonstrated that Botryobasidium forms a well-supported monophyletic group, consistent with previous evidence from its micromorphological and ultrastructural characteristics, consequently, several new species in the genus have been described in China (Li et al. 2025a, 2025b; Zhang et al. 2025). So far, according to the Index Fungorum (www.indexfungorum.org; accessed on July 10, 2026) and MycoBank (https://www.mycobank.org/page/Simple%20names%20search; accessed on July 10, 2026), 120 and 138 taxa were registered in Botryobasidium, respectively, but 97 species are accepted worldwide (Lentz 1967; Jung 1995; Langer et al. 2000a; Hagara 2001; Bernicchia et al. 2010; Saitta et al. 2011; Bates et al. 2017; Kalinina et al. 2020; Ram et al. 2021). Among these accepted species, 24 species were discovered in China (Dong et al. 2024b; Liu et al. 2024; Zhou et al. 2024a; Zhou et al. 2024b; Li et al. 2025a; Wang et al. 2025; Zhang et al. 2025; Yuan et al. 2026).

The genus Peniophorella P. Karst. belongs to the family Peniophorellaceae L.W. Zhou et al., the order Hymenochaetales Oberw., and is typified by P. pubera (Fr.) P. Karst. The genus is characterised by annual, effused, thin basidiomata, white to yellowish, smooth to odontioid hymenophores, a monomitic hyphal system, various cystidia, thin-walled, ellipsoid, cylindrical, or allantoid basidiospores with negative reaction in Melzer’s reagent and Cotton Blue, and causing a white rot (Karsten 1889; Bernicchia and Gorjón 2010). The independence of Peniophorella was reinstated from Hyphoderma Wallr. in hymenochaetoid clade by Larsson (2007a) based on molecular phylogeny, and 19 combinations in Peniophorella were proposed. Subsequent studies by Miettinen and Larsson (2011) and Telleria et al. (2012) confirmed that Peniophorella forms a well-resolved monophyletic group clearly separated from Hyphodermasensu stricto. Although later classified in Hyphodermataceae (Justo et al. 2017), more recent studies have proposed the new family Peniophorellaceae (Wang et al. 2023), which has been accepted in subsequent works. Meanwhile, based on morphological and molecular phylogenetic analyses, several species have been described in the genus (Dong et al. 2024a; Liu et al. 2024; Zhang et al. 2026). So far, according to the Index Fungorum (www.indexfungorum.org; accessed on July 10, 2026) and MycoBank (https://www.mycobank.org/page/Simple%20names%20search; accessed on July 10, 2026), 46 and 49 specific taxa were registered in Peniophorella, respectively, with 46 species accepted worldwide, and 30 species were discovered in China (Larsson 2007b; Duhem 2013; Prasher 2015; Yurchenko et al. 2020; Dong et al. 2024a; Liu et al. 2024; Deng et al. 2025; Li et al. 2025b; Wang et al. 2026; Zhang et al. 2026; Zhu et al. 2026).

A key feature of Hymenochaetales is the high diversity of corticioid fungi, which represent a polyphyletic assemblage based on morphology (Liu et al. 2025; Zhao et al. 2025a, 2025b). In the past, the classification of many corticioid genera relied primarily on morphological traits, which often resulted in unnatural groupings. The advent of molecular phylogenetics has substantially improved our understanding of the systematics within these corticioid taxa (Wu et al. 2022; Yuan et al. 2023; Zhou et al. 2023). Several recently established families—such as Hyphodontiaceae X.Wei Wang & L.W. Zhou, Peniophorellaceae, and Schizoporaceae Jülich—mainly consist of corticioid genera, indicating considerable phylogenetic divergence within this morphological group (Wang et al. 2021, 2023). Consequently, combining morphological evidence with phylogenetic analyses remains essential for the accurate delimitation of these inconspicuous fungi.

Currently, the species diversity of Cantharellales and Hymenochaetales remains largely unexplored, with ongoing studies indicating substantial undiscovered diversity, particularly in biodiverse regions such as East Asia. In the current study, based on ITS + nLSU sequences, one new species belonging to the genus Botryobasidium (Cantharellales) and two new species belonging to the genus Peniophorella (Hymenochaetales) are described and illustrated.

Materials and methods

Morphological studies

Five specimens were collected from natural forests, found on Pinus massoniana from Guangdong, Henan and Jiangxi Province of China, respectively. These are deposited in the Fungarium of the Institute of Microbiology, Beijing Forestry University (BJFC), China. Morphological descriptions are based on field notes and voucher specimens. The microscopic analysis follows Spirin et al. (2017) and Wu et al. (2022). Freehand sections were made from dried basidiomata and mounted in 5% (w/v) potassium hydroxide (KOH) to observe colour changes. Sections were studied at a magnification of up to 1000× using a Nikon Eclipse 80i microscope and phase contrast illumination. Microscopic features and measurements were made from slide preparations stained with Cotton Blue and Melzer’s reagent. To represent the variation in the size of spores, 5% of measurements were excluded from each end of the range and are given in parentheses. In the description: KOH = 5% potassium hydroxide, IKI = Melzer’s reagent, IKI– = neither amyloid nor dextrinoid, CB = Cotton Blue, CB– = acyanophilous in Cotton Blue, L = arithmetic average of spore length, W = arithmetic average of spore width, Q = L/W ratios and n = number of basidiospores measured from a given number of specimens. Colour terms follow Anonymous (1969) and Petersen (1996).

DNA extraction, amplification and sequencing

A CTAB rapid plant genome extraction kit-DN14 (Aidlab Biotechnologies Co., Ltd, Beijing) was used to obtain DNA from dried specimens and to perform the polymerase chain reaction (PCR) according to the manufacturer’s instructions with some modifications (Shen et al. 2019; Sun et al. 2020). The internal transcribed spacer (ITS) and large subunit nuclear ribosomal RNA gene (nLSU) were amplified using the primer pairs ITS5/ITS4 and LR0R/LR7 (White et al. 1990; Hopple and Vilgalys 1999, https://sites.duke.edu/vilgalyslab/rdna_primers_for_fungi/).

PCR amplifications were carried out in 30 μL reaction mixtures containing 15 μL of 2× EasyTaq PCR Super Mix (Beijing Quanshi Gold Biotechnology Co., Ltd., Beijing, China), 12 μL of RNase-free Water, 1 μL of each primer, and 1 μL of genomic DNA. The PCR procedure for ITS was as follows: initial denaturation at 95 °C for 3 min, followed by 34 cycles at 94 °C for 40 s, annealing at 54 °C for 45 s and extension at 72 °C for 1 min, with a final extension of 72 °C for 10 min. The PCR procedure for nLSU was as follows: initial denaturation at 94 °C for 1 min, followed by 34 cycles of denaturation at 94 °C for 30 s, annealing at 50 °C for 1 min and extension at 72 °C for 1.5 min, with a final extension at 72 °C for 10 min. The PCR products were purified and sequenced at the Beijing Genomics Institute (BGI, China), with the same primers. The newlygenerated sequences were deposited in GenBank (Sayers et al. 2024) and listed in Table 1.

Table 1.

Names, specimen numbers, references and corresponding GenBank accession numbers of the taxa used in the phylogenetic analysis of this study. [New species are in bold; * indicates the type material; — indicates no available sequence].

Species Sample No. GenBank Accession No. References
ITS nLSU
Botryobasidium acanthosporum Yuan16326 PP229497 — Zhou et al. 2024a
B. acanthosporum Yuan17989 PP229511 — Zhou et al. 2024a
B. acanthosporum Yuan18083* PP229512 PP218361 Zhou et al. 2024a
B. acanthosporum Yuan18128 PP229517 — Zhou et al. 2024a
B. asperulum RAS552 OR471090 OR470959 Swenie et al. 2024
B. asperulum RAS578 OR471100 OR470964 Swenie et al. 2024
B. aureum RAS571 SV1 OR471098 OR470963 Swenie et al. 2024
B. aureum RAS571 SV2 OR471099 — Swenie et al. 2024
B. bambusinum CLZhao 29916 PQ539057 PQ539060 Dong et al. 2024b
B. bambusinum CLZhao 29936* PQ539058 PQ539061 Dong et al. 2024b
B. bambusinum CLZhao 29938 PQ539059 PQ539062 Dong et al. 2024b
B. botryosum AFTOL-ID 604 DQ267124 FJ176881 Matheny et al. 2006
B. candicans FRDBI 29580226 OR896129 — Unpublished
B. coniferarum LWZ20210928-3* OR557259 OR527282 Liu et al. 2024
B. coniferarum LWZ20171016-15 OR557262 OR527286 Liu et al. 2024
B. conspersum AFTOL-ID 1766 DQ911612 DQ521414 Matheny et al. 2006
B. daweishanense CLZhao 40061 PQ373983 — Zhang et al. 2025
B. daweishanense CLZhao 40062* PQ373984 PQ373977 Zhang et al. 2025
B. gossypirubiginosum CLZhao 26052* OR668924 OR708665 Zhou et al. 2024b
B. gossypirubiginosum Dai 26208 PQ285750 — Li et al. 2025a
B. incanum CLZhao 26697* OR668923 OR708664 Zhou et al. 2024b
B. incanum Dai 25375 PQ285751 PQ28566 Li et al. 2025a
B. indicum Yuan18434 PP209217 PP218365 Li et al. 2025a
B. indicum hr5326 OP806032 — Li et al. 2025a
B. intertextum UC2022959 KP814540 — Rosenthal et al. 2017
B. leave RAS762 OR471128 — Swenie et al. 2024
B. latihyphum Dai 26858* PQ279526 PQ282521 Li et al. 2025a
B. latihyphum Yuan 16496 PP331854 PP218153 Li et al. 2025a
B. leptocystidiatum Yuan17706 PP209200 PP218353 Zhou et al. 2024a
B. leptocystidiatum Yuan17708* PP209197 PP218354 Zhou et al. 2024a
B. massonianae Dai 35993* PZ459029 PZ459034 Present study
B. massonianae Dai 36002 PZ459030 PZ459035 Present study
B. robustius CBS:945.69 MH859491 MH871272 Vu et al. 2019
B. robustius iNaturalist 162067551 PP436446 — Unpublished
B. rubiginosum RAS776 OR471136 — Swenie et al. 2024
B. simile RAS793 OR471146 — Swenie et al. 2024
B. simile RAS794 OR471147 — Swenie et al. 2024
B. subcoronatum RAS620 SV1 OR471110 OR470967 Swenie et al. 2024
B. subincanum LWZ20230417-17b PP959661 PP959649 Wang et al. 2025
B. subincanum LWZ20230417-41a PP959660 — Wang et al. 2025
B. subovalibasidium Yuan16439 PP209199 PP218152 Zhou et al. 2024a
B. subovalibasidium Yuan18179* PP209196 PP218362 Zhou et al. 2024a
B. tubulicystidium DK14_139 OL436769 — Zhou et al. 2024b
B. vagum LWZ20191016-22 PP959659 PP959648 Wang et al. 2025
B. xizangense LWZ20230722-25a* PP959663 PP959650 Wang et al. 2025
B. xizangense LWZ20230722-16a PP959662 — Wang et al. 2025
B. yunnanense CLZhao 24877* OR668925 OR708666 Zhou et al. 2024b
B. zhejiangensis Dai 25056* PQ279530 PQ282525 Li et al. 2025a
B. zhejiangensis Dai 24851 PQ279529 PQ282524 Li et al. 2025a
Lyomyces allantosporus FR 0249548 NR154135 — Yurchenko et al. 2017
L. pruni GEL2327 DQ340312 — Yurchenko et al. 2020
Peniophorella alba Yuan 17692* PV883103 — Zhu et al. 2026
P. albohymenia CLZhao 33187* PQ811412 PQ847496 Deng et al. 2025
P. albohymenia CLZhao 33257 PQ811413 — Deng et al. 2025
P. aspersa TNM F20163 MN062098 MN062143 Yurchenko et al. 2020
P. aspersa TNM F24809* MN062097 MN062142 Yurchenko et al. 2020
P. cremea CLZhao 1606* MT955162 — Xu et al. 2020
P. cremea CLZhao 1719 MT955163 — Xu et al. 2020
P. crystallifera TNM F23666 KX427170 MN062145 Yurchenko et al. 2020
P. crystallifera LWZ 20210626-4a ON063685 ON063885 Yurchenko et al. 2020
P. daweishanensis CLZhao 18600* OR094501 OR449932 Dong et al. 2024a
P. echinocystis KHL 6284 DQ677494 DQ681200 Larsson 2007b
P. euryhypha Dai 25368* PQ726884 PQ726867 Li et al. 2025b
P. fissurata CLZhao 4539 MN864259 OM985775 Guan et al. 2020
P. fissurata CLZhao 9421* MN864260 OM985776 Guan et al. 2020
P. guttulifera CBS 107303 LT603016 LT603001 Kolařík and Vohník 2018
P. guttulifera NH 12012 DQ647501 — Hallenberg et al. 2007
P. odontiiformis MUCL 32673 DQ647498 — Hallenberg et al. 2007
P. odontiiformis SFC20191015-02 OQ996171 OQ996202 Cho et al. 2024
P. olivacea CLZhao 25896* OR094502 OR449933 Dong et al. 2024a
P. pertenuis FCUG 2430 DQ647482 — Hallenberg et al. 2007
P. pertenuis Wu 9606-13 DQ647477 — Hallenberg et al. 2007
P. pinicola Dai 28400* PQ726882 PQ726865 Li et al. 2025b
P. pinicola Dai 28468 PQ726883 PQ726866 Li et al. 2025b
P. pinicola Yuan 1444 PQ726889 PQ726872 Li et al. 2025b
P. praetermissa NH 10986 DQ647462 — Hallenberg et al. 2007
P. praetermissa NH 11192 DQ647461 — Hallenberg et al. 2007
P. praetermissa Yuan 282 PQ726879 PQ726862 Li et al. 2025b
P. praetermissa Yuan 339 PQ726880 PQ726863 Li et al. 2025b
P. pubera CBS 464 86 MH861988 MH873680 Vu et al. 2019
P. pubera LWZ 20210624-16b ON063687 ON063887 Wang et al. 2023
P. punctata CLZhao 33720 PQ811414 PQ847497 Deng et al. 2025
P. punctata CLZhao 33732* PQ811415 PQ847498 Deng et al. 2025
P. reticulata CLZhao 17096 OM985745 OM985783 Unpublished
P. reticulata TNM F22559* NR172776 NG073752 Yurchenko et al. 2020
P. rude LWZ 20171026-7 ON063688 ON063888 Wang et al. 2023
P. sidera LWZ 20180921-15* OQ540892 OQ540850 Wang et al. 2023
P. sidera LWZ 20180922-62 OQ540893 OQ540851 Wang et al. 2023
P. stellata Dai 25341 PQ726876 PQ726859 Li et al. 2025b
P. stellata Dai 25358* PQ726877 PQ726860 Li et al. 2025b
P. stellata Dai 25361 PQ726878 PQ726861 Li et al. 2025b
P. subalbohymenia QYZhang 141* PX270289 PX270293 Zhang et al. 2026
P. subalbohymenia QYZhang 191 PX270290 PX270294 Zhang et al. 2026
P. subglobospora 10628MD HG315519 — Telleria et al. 2013
P. subpraetermissa Wu 9506-27* DQ647493 — Hallenberg et al. 2007
P. subreticulata LWZ 20200812-37a OQ540886 OQ540846 Wang et al. 2023
P. subreticulata LWZ 20200921-49b* OQ540884 OQ540845 Wang et al. 2023
P. tenuissima HMZhou 377* PV475574 PV646276 Wang et al. 2026
P. tongbiguanensis CLZhao 36353* PV475575 PV646277 Wang et al. 2026
P. tsugae NH 7473 — DQ677505 Larsson 2007a
P. yunnanensis CLZhao 4810 MN864263 OM985788 Guan et al. 2020
P. yunnanensis CLZhao 6132 MN864268 OM985789 Guan et al. 2020
P. crassibasidiata Yuan 3208* PZ459031 PZ459036 Present study
P. variegata Yuan 2232* PZ459032 PZ459037 Present study
P. variegata Yuan 2250 PZ459033 PZ459038 Present study
Schizocorticium magnosporum Wu 1510-34 MK405351 MK405337 Wu et al. 2021
S. mediosporum Chen 2456 MK405359 MK405345 Wu et al. 2021
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Sequences generated were aligned manually with additional sequences downloaded from GenBank using AliView version 1.27 (Larsson 2014). The final ITS and nLSU datasets were subsequently aligned using MAFFT v.7 under the E-INS-i strategy with no cost for opening gaps and equal cost for transformations (command line: mafft –genafpair –maxiterate 1000, Katoh and Standley (2013) and visualised in AliView. Alignments were spliced and transformed formats in Mesquite v.3.2. (Maddison and Maddison 2017). Multiple sequence alignments were trimmed by trimal v.1.2 using the -htmlout-gt 0.8 -st option to deal with gaps, when necessary (Capella-Gutierrez et al. 2009).

Phylogenetic analyses

The two-marker DNA multiple sequence alignment (ITS + nLSU) was used to determine the phylogenetic position of the new species. Sequences of Lyomyces allantosporus Riebesehl, Yurchenko & Langer and L. pruni (Lasch) Riebesehl & Langer were chosen as the outgroups (Fig. 1, Spirin et al. 2015; Riebesehl and Langer 2017; Chen and Zhao 2020). Sequences of Schizocorticium magnosporum Sheng H. Wu & C.L. Wei and S. mediosporum Sheng H. Wu & C.L. Wei were chosen as the outgroups (Fig. 2, Cho et al. 2024). The phylogenetic analyses followed the approach of Han et al. (2015) and Zhu et al. (2019). Maximum Likelihood (ML) and Bayesian Inference (BI) analyses were performed, based on the ITS + nLSU datasets.

Figure 1.

Figure 1.

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ML analysis of Botryobasidium, based on dataset of ITS + nLSU. ML bootstrap values equal to or higher than 60% and Bayesian posterior probabilities values equal to or higher than 0.90 are shown. New taxa are in bold, * represents type material.

Figure 2.

Figure 2.

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ML analysis of Peniophorella, based on dataset of ITS + nLSU. ML bootstrap values equal to or higher than 60% and Bayesian posterior probabilities values equal to or higher than 0.90 are shown. New taxa are in bold, * represents type material.

Sequences were analysed using Maximum Likelihood (ML) with RAxML v.8.2.10 (Stamatakis 2014). Branch supports for all parsimony analyses were estimated by performing 1,000 bootstrap replicates with a heuristic search of 10 random-addition replicates for each bootstrap replicate. Bayesian phylogenetic inference and Bayesian Posterior Probabilities (BPP) were computed with MrBayes 3.2.6 with a GTR + I + G model of DNA substitution and a gamma distribution rate variation across sites (Ronquist and Huelsenbeck 2003). Four Markov chains were run for 1.5 million generations (two-marker dataset) until the split deviation frequency value was less than 0.01 and trees were sampled every 100 generations in Fig. 1; for 2 million generations in Fig. 2. The first 25% of the sampled trees were discarded as burn-in and the remaining ones were used to reconstruct a majority rule consensus and calculate Bayesian Posterior Probabilities (BPP) of the clades. All trees were viewed in FigTree v. 1.4.3 (http://tree.bio.ed.ac.uk/software/figtree/). Branches that received bootstrap support for ML (≥ 75% (ML-BS)) and BPP (≥ 0.95 BPP) were considered as significantly supported. The ML bootstrap (ML) ≥ 50% and Bayesian Posterior Probabilities (BPP) ≥ 0.90 are presented on topologies from ML analysis, respectively.

Results

Molecular phylogeny

The combined two-marker dataset (ITS + nLSU, Fig. 1) included sequences from 51 samples representing 30 taxa. The phylogenetic reconstruction performed with Maximum Likelihood (ML) and Bayesian Inference (BI) analyses for the combined dataset showed similar topology and few differences in statistical support. The best model-fit applied in the Bayesian analysis was GTR + I + G, lset nst = 6, rates = invgamma and prset statefreqpr = dirichlet (1, 1, 1, 1). Bayesian analysis and ML analysis resulted in a similar topology to the MP analysis, with an average standard deviation of split frequencies of 0.007700 (BI). The phylogenetic tree inferred from the ITS + nLSU sequences indicated that the new species belonged to Botryobasidium (Fig. 1). In addition, Botryobasidium massonianae grouped together with B. aureum Parmasto, B. coniferarum S.L. Liu & L.W. Zhou and B. indicum (P.N. Singh & S.K. Singh) R. Kirschner & G. Langer with high support (ML = 81%, BPP = 1.00).

The combined two-marker dataset (ITS + nLSU, Fig. 2) included sequences from 55 samples representing 33 taxa. The phylogenetic reconstruction performed with Maximum Likelihood (ML) and Bayesian Inference (BI) analyses for the combined dataset showed similar topology and few differences in statistical support. The best model-fit applied in the Bayesian analysis was GTR + I + G, lset nst = 6, rates = invgamma and prset statefreqpr = dirichlet (1, 1, 1, 1). Bayesian analysis and ML analysis resulted in a similar topology to the MP analysis, with an average standard deviation of split frequencies of 0.008584 (BI). The phylogenetic tree inferred from the ITS + nLSU sequences indicated that the two new species belonged to Peniophorella (Fig. 2). In addition, Peniophorella crassibasidiata formed an independent lineage; P. variegata is related to P. alba L.J. Zhou et al., P. albohymenia Y.L. Deng & C.L. Zhao, P. stellata Y.C. Dai et al., P. subalbohymenia Q.Y. Zhang, and P. tenuissima Lu Wang & C.L. Zhao with high support (ML = 97%, BPP = 1.00).

Based on the ITS and nLSU sequences of the type material, BLAST queries against the authoritative sequences in NCBI yielded alignments, which are shown in Table 2.

Table 2.

Based on the ITS and nLSU sequences of the type material, BLAST queries against the authoritative sequences in NCBI yielded alignments.

Species name Marker Top match species GenBank Accession No. Max score Total score Query cover E-value Identity
Botryobasidium massonianae ITS B. indicum PV191125 1002 1002 99% 0.0 94.74%
nLSU B. aureum OR470963 2370 2370 99% 0.0 98.37%
Peniophorella crassibasidiata ITS P. reticulata NR172776 1064 1064 96% 0.0 96.31%
nLSU P. aspersa OM985770 1797 1797 99% 0.0 99.69%
P. variegata ITS P. subalbohymenia PX270289 1118 1118 99% 0.0 96.35%
nLSU P. subpraetermissa OM985785 2431 2431 99% 0.0 99.4%
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Taxonomy

Botryobasidium massonianae

K.Y. Luo, Yuan Yuan & Y.C. Dai sp. nov.

13338FD3-5060-5669-AEAB-DA8D6E3E8FCE

864541

Figs 3, 4

Figure 3.

Figure 3.

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Basidiomata of Botryobasidium massonianae (Holotype, Dai 35993). Scale bar: 1 cm (A); 0.3 mm (B).

Figure 4.

Figure 4.

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Microscopic structures of Botryobasidium massonianae (Holotype, Dai 35993). a. Basidiospores; b. Basidia and basidioles; c. A section of the basidioma.

Holotype.

China • Henan Province, Xinyang, Shangcheng County, Huangbaishan Forest Park. GPS coordinates: 31.459192°N, 115.370512°E; elevation: 500 m a.s.l. On a fallen branch of Pinus massoniana, 20 September 2025, Dai 35993 (BJFC 057254).

Etymology.

Massonianae (Lat.) refers to the species growing on Pinus massoniana.

Description.

Basidiomata annual, resupinate, effuse, fluffy, pellicular, adherent to the substrate and not easily separated, without odour or taste, up to 6 cm long, 1.5 cm wide, and 200 µm thick at centre. Hymenophoral surface hypochnoid, white when fresh, cream when dry. Sterile margin often indeterminate and not differentiated.

Hyphal structure: Hyphal system monomitic; generative hyphae simple septate, slightly thick-walled with wide lumen, moderately branched mostly with a right angle, more or less straight, loosely interwoven, 5–7.5 μm in diam; IKI–, CB–; tissues unchanged in KOH.

Hymenium: Cystidia and cystidioles absent. Basidia barrel-shaped, thin-walled, with four to six sterigmata and a basal simple septum, 11–16 × 5–10 µm; basidioles in shape similar to basidia, but slightly shorter.

Basidiospores: Fusiform, hyaline, thin-walled, smooth, IKI–, CB–, (5.7–)6.3–9.2(–10.2) × (2.7–)2.8–3.9(–4.3) µm, L = 7.46 µm, W = 3.31 µm, Q = 2.24–2.26 (n = 60/2).

Type of rot.

White rot.

Additional specimen (paratype) examined.

China • Henan Province, Xinyang, Shangcheng County, Huangbaishan Forest Park. GPS coordinates: 31.459192°N, 115.370512°E; elevation: 500 m a.s.l. On a fallen branch of Pinus massoniana, 20 September 2025, Dai 36002 (BJFC 057263).

Peniophorella crassibasidiata

K.Y. Luo, Yuan Yuan & Y.C. Dai sp. nov.

0D622D0E-908F-506C-AB05-F74BC3263983

864542

Figs 5, 6

Figure 5.

Figure 5.

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Basidiomata of Peniophorella crassibasidiata (Holotype, Yuan 3208). Scale bar: 1 cm (A); 0.2 mm (B).

Figure 6.

Figure 6.

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Microscopic structures of Peniophorella crassibasidiata (Holotype, Yuan 3208). a. Basidiospores; b. Basidia and basidioles; c. Stephanocysts; d. Subulate cystidia; e. Fusiform cystidia; f. A section of the basidioma.

Holotype.

China • Guangdong Province, Zhaoqing, Dinghu District, Dinghushan National Nature Reserve. GPS coordinates: 23.159571°N, 112.538344°E; elevation: 51 m a.s.l. On a fallen branch of Pinus massoniana, 8 July 2025, Yuan 3208 (BJFC 062754).

Etymology.

Crassibasidiata (Lat.), crassi- = thick, basidiata = basidia, referring to the thick-walled basidia of this species.

Description.

Basidiomata annual, resupinate, adnate, loosely attached, but not easily separable from substratum, soft coriaceous, without odor and taste when fresh, white when fresh, white to cream upon drying, up to 14 cm long, 1.5 cm wide, and 200 µm thick at center. Hymenophoral surface smooth; margin distinct, thin, white, up to 2 mm long.

Hyphal structure: Hyphal system monomitic; generative hyphae with clamp connections, thick-walled with a wide lumen, occasionally branched, flexuous, loosely interwoven, 2–4 μm in diameter, IKI–, CB–; tissues unchanged in KOH.

Hymenium: Cystidia of three types: (1) stephanocysts infrequent, with some oily contents, hyaline, thin-walled, 9–11 × 6–7 μm; (2) fusiform cystidia frequent, hyaline, thick-walled, 22–25 × 7–9 μm; (3) subulate cystidia frequent, sometimes with oily contents, hyaline, thick-walled, 20–50 × 6–10 μm. Cystidioles absent. Nubbly crystals abundant. Basidia barrel-shaped to subclavate, colorless, thick-walled, smooth, with four sterigmata and a basal clamp connection, 12–22 × 5–9 μm. Basidioles similar in shape to basidia, but smaller.

Basidiospores: cylindrical, hyaline, thin-walled, smooth, occasionally with two oil drops or contents, IKI–, CB–, (8–)8.4–12.4(–12.6) × (3–)3.1–4.2(–4.4) µm, L = 10.21 µm, W = 3.67 µm, Q = 2.78 (n = 30/1).

Type of rot.

White rot.

Peniophorella variegata

K.Y. Luo, Yuan Yuan & Y.C. Dai sp. nov.

30CE719D-A320-5C85-86CA-33E20379D517

864543

Figs 7, 8

Figure 7.

Figure 7.

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Basidiomata of Peniophorella variegata (Holotype, Yuan 2232). Scale bar: 1 cm (A); 0.1 mm (B).

Figure 8.

Figure 8.

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Microscopic structures of Peniophorella variegata (Holotype, Yuan 2232). a. Basidiospores; b. Basidia and basidioles; c. Capitate cystidia; d. Leptocystidia; e. Cystidioles; f. Hyphidia; g. Subulate cystidia; h. A section of the basidioma.

Holotype.

China • Jiangxi Province, Nanchang, Shengshuitang National Forest Park. GPS coordinates: 28.991150°N, 115.533628°E; elevation: 220 m a.s.l. On a fallen branch of Pinus massoniana, 26 April 2025, Yuan 2232 (BJFC 061778).

Etymology.

Variegata (Lat.): refers to the species having variable cystidia.

Description.

Basidiomata annual, resupinate, adnate, loosely attached, not easily separable from substratum, farinaceous, without odour and taste when fresh, white to pale yellowish when fresh and upon drying, up to 7 cm long, 1.5 cm wide, and 300 µm thick. Hymenophoral surface smooth; margin distinct, thin, pale yellowish, up to 2 mm long.

Hyphal structure: Hyphal system monomitic; generative hyphae with clamp connections, thin-walled, moderately branched mostly with a right angle, flexuous, loosely interwoven, 2.5–4 μm in diameter, IKI–, CB–; tissues unchanged in KOH.

Hymenium: Cystidia of three types: (1) capitate cystidia frequent, hyaline, thin-walled, 18.5–24 × 4–7 μm; (2) leptocystidia infrequent, subcylindrical, hyaline, thin-walled, sometimes apically encrusted, 19–31 × 5–7 μm; (3) subulate cystidia infrequent, hyaline, thin-walled, sometimes protruding from the hymenium up to 20 μm, 38–53 × 4–7 μm. Hyphidia frequent, hyaline, thin-walled, 13–16 × 2.5–4 μm. Cystidioles rare, subcylindrical or sphaeropedunculate, hyaline, thin-walled, 11–14 × 6–8 μm. Basidia clavate, more or less flexuous, colorless, thin-walled, smooth, with four sterigmata and a basal clamp connection, occasionally with oily contents, 25–46 × 6–12 μm. Basidioles similar in shape to basidia, but distinctly smaller.

Basidiospores: Short cylindric to oblong-ellipsoid, sometime slightly curved, hyaline, thin-walled, smooth, occasionally with oily contents, IKI–, CB–, (7.3–)7.9–10.8(–12.5) × (3.1–)3.9–5.1(–5.8) µm, L = 9.44 µm, W = 4.50 µm, Q = 2.07–2.13 (n = 60/2).

Type of rot.

White rot.

Additional specimen (paratype) examined.

China • Jiangxi Province, Nanchang, Shengshuitang National Forest Park. GPS coordinates: 28.991150°N, 115.533628°E; elevation: 220 m a.s.l. On a fallen branch of Pinus massoniana, 26 April 2025, Yuan 2250 (BJFC 061796).

Discussion

Cantharellales and Hymenochaetales are two important orders within the class Agaricomycetes (Basidiomycota) that exhibit significant differences in ecological function, wood-inhabiting mechanisms, and phylogenetic relationships. In recent years, with the rapid development of molecular phylogenetic methods and the increasing intensity of global surveys of forest fungal diversity, important progress has been made in understanding the species diversity, taxonomic framework, and evolutionary history of these two orders (Wang et al. 2018; Swenie et al. 2025; Zhao et al. 2025a; Kumar et al. 2026).

Cantharellales is an ecologically highly diverse order, encompassing saprotrophic, mycorrhizal, lichenised, and parasitic nutritional modes (Tuo et al. 2026; Xu et al. 2026). One new species discovered in this study, Botryobasidium massonianae, is a saprotrophic fungus growing on Pinus massoniana.

Hymenochaetales includes a large number of important wood-decaying fungi including some forest pathogens and medicinal fungi (Dai et al. 2000; Wu et al. 2004; Wu et al. 2019; Wu and Dai 2020), and have a worldwide distribution. This order contains corticioid, poroid and hydnoid fungi with different types of basidiomata (Dai et al. 2004; Wu et al. 2022; Zhou et al. 2023; Zhao et al. 2025a). The two species Peniophorella crassibasidiata and P. variegata discovered in this study are both corticioid fungi.

In our phylogeny (Fig. 1), Botryobasidium massonianae grouped together with B. aureum, B. coniferarum and B. indicum with high support (ML = 81%, BPP = 1.00). However, B. aureum is different from B. massonianae by its thin-walled generative hyphae (Parmasto 1965); B. coniferarum is readily distinguished from B. massonianae by longer basidia (16–20 µm vs. 11–16 µm, Liu et al. 2024); B. indicum differs from B. massonianae by its yellow basidiomata (Hyde et al. 2019). Morphologically, B. massonianae, B. acanthosporum L.J. Zhou & H.S. Yuan and B. subovalibasidium L.J. Zhou & H.S. Yuan share simple septate generative hyphae. However, B. acanthosporum is different from B. massonianae by its basidia with 2 sterigmata and subglobose to globose, slightly thick- to thick-walled basidiospores; B. subovalibasidium differs from B. massonianae by its pale yellow to greyish yellow hymenophoral surface (Zhou et al. 2024a).

In our phylogeny (Fig. 2), Peniophorella crassibasidiata formed an independent lineage. Morphologically, P. crassibasidiata, P. pinicola Y.C. Dai et al. and P. stellata share the presence of stephanocysts. However, P. pinicola is different from P. crassibasidiata by its shorter basidiospores (5.7–8 µm vs. 8.4–12.4 µm); P. stellata differs from P. crassibasidiata by its longer basidia (22.5–32 µm vs. 12–22 µm, Li et al. 2025b).

Peniophorella variegata is related to P. alba, P. albohymenia, P. stellata, P. subalbohymenia, and P. tenuissima with high support (ML = 97%, BPP = 1.00). However, P. alba is different from P. variegata by its thick-walled cystidia (Zhu et al. 2026); P. albohymenia is readily distinguished from P. variegata by its absence of hyphidia and cystidioles (Deng et al. 2025); P. stellata differs from P. variegata a (Li et al. 2025b); P. subalbohymenia is different from P. variegata by its shorter basidia (18–25 µm vs. 25–46 µm, Zhang et al. 2026); P. tenuissima is readily distinguished from P. variegata by its wider basidiospores (5.5–7.5 µm vs. 3.9–5.1 µm, Wang et al. 2026). Morphologically, P. variegata, P. olivacea J.H. Dong & C.L. Zhao and P. tongbiguanensis L. Wang & C.L. Zhao share farinaceous basidiomata. However, P. olivacea differs from P. variegata by its presence of halocystidia, the absence of hyphidia, and both shorter basidia (17.5–20 µm vs. 25–46 µm) and basidiospores (6.5–7.5 µm vs. 7.9–10.8 µm, Dong et al. 2024a); P. tongbiguanensis is different from P. variegata by lacking capitate cystidia, hyphidia and cystidioles, and having larger basidiospores (11–13.5 × 5.5–7 µm vs. 7.9–10.8 × 3.9–5.1 µm, Wang et al. 2026).

We collected specimens from Guangdong, Henan, and Jiangxi provinces, representing southern, central, and eastern China, respectively. Notably, new species belonging to these two orders were discovered in all these areas, indicating that the diversity of Cantharellales and Hymenochaetales is more widely distributed across different climatic zones of China. This broader geographic coverage strengthens the significance of our findings and provides a more comprehensive perspective on the national diversity of these fungal groups.

Supplementary Material

XML Treatment for Botryobasidium massonianae
XML Treatment for Peniophorella crassibasidiata
XML Treatment for Peniophorella variegata

Citation

Luo K-Y, Dai Y-C, Yuan Y, Wu Y-D (2026) Three new species of Cantharellales and Hymenochaetales from Pinus massoniana in China. MycoKeys 137: 273–297. https://doi.org/10.3897/mycokeys.137.203862

Contributor Information

Yuan Yuan, Email: yuanyuan1018@bjfu.edu.cn.

Ying-Da Wu, Email: wydbjfu@163.com.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Artificial Intelligence (AI) use

The authors accept full responsibility for the content of the manuscript, including the disclosure of any use of AI.

No AI tools were used in the preparation of this manuscript.

Funding

The research was supported by the National Natural Science Foundation of China (Project Nos. 32370013, U23A20142).

Author contributions

All authors designed the research and contributed to data analysis and interpretation, prepared the samples and drafted the manuscript. All authors have read and agreed to the published version of the manuscript.

Author ORCIDs

K.-Y. Luo https://orcid.org/0000-0001-6145-4864

Y.-C. Dai https://orcid.org/0000-0002-6523-0320

Y. Yuan https://orcid.org/0000-0001-6674-9848

Y.-D. Wu https://orcid.org/0000-0003-1295-4015

Data availability

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

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Associated Data

This section collects any data citations, data availability statements, or supplementary materials included in this article.

Supplementary Materials

XML Treatment for Botryobasidium massonianae
mycokeys.137.203862-treatment1.xml (8.8KB, xml)
XML Treatment for Peniophorella crassibasidiata
mycokeys.137.203862-treatment2.xml (8.1KB, xml)
XML Treatment for Peniophorella variegata
mycokeys.137.203862-treatment3.xml (9.4KB, xml)

Data Availability Statement

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

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