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Research Article
Morphological characteristics and phylogenetic analyses revealed four new species (Basidiomycota) in the Yunnan-Guizhou Plateau, China
expand article infoSiyuan He‡, Lu Wang‡, Kaize Shen§, Hongmin Zhou‡
‡ Southwest Forestry University, Kunming, China
§ Zhaotong University, Kunming, China

Corresponding author: Hongmin Zhou ( hongminzhou@foxmail.com )

Academic editor: Samantha C. Karunarathna
© 2025 Siyuan He, Lu Wang, Kaize Shen, Hongmin Zhou.
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: He S, Wang L, Shen K, Zhou H (2025) Morphological characteristics and phylogenetic analyses revealed four new species (Basidiomycota) in the Yunnan-Guizhou Plateau, China. MycoKeys 113: 237-262. https://doi.org/10.3897/mycokeys.113.140932
Open Access

Abstract

Four new fungi, viz. Clavulinopsis wumengshanensis (Clavariaceae, Agaricales), Henningsomyces bambusae (Porotheleaceae, Agaricales), Xenasma bisterigmatae, and X. guttulata (Xenasmataceae, Russulales), from Yunnan Province in China, are proposed, based on a combination of morphological features and molecular evidence. Phylogenetic analyses were conducted using a combined dataset of internal transcribed space and nuclear ribosomal RNA large subunit sequences. The ITS+LSU analysis showed that Clavulinopsis wumengshanensis sp. nov. groups with C. aurantiocinnabarina, Henningsomyces bambusae sp. nov. forms a sister group with H. candidus, Xenasma bisterigmatae sp. nov. is clustered with X. rimicola, and X. guttulata sp. nov. is clustered with X. pruinosum. The morphology and multi-gene phylogenetic analyses confirmed the placement of the four new taxa. Clavulinopsis wumengshanensis is distinguished by buff-yellow to straw-yellow basidiomata, clavate to subcylindrical basidia with four sterigmata, and subglobose basidiospores with several guttules (7–8.5 × 6–7.5 µm); Henningsomyces bambusae is characterized by white to cream basidiomata with short cylindric to tubular colonies, cylindrical to subcylindrical basidia with two sterigmata and globose to subglobose basidiospores (6.5–8.5 × 6.5–8.5 µm); Xenasma bisterigmatae can be characterized by its membranaceous and ash-grey basidiomata, clavate to subcylindrical basidia with two sterigmata and ellipsoid to subglobose basidiospores (10–12.5 × 8–10.5 µm); X. guttulata is distinguished by membranaceous and white to cream basidiomata, clavate to subcylindrical basidia with two sterigmata and ellipsoid to narrowly ellipsoid basidiospores (7–9 × 5.5–7.5 µm).

Key words:

4 new taxa, Agaricales, biodiversity, Russulales, taxonomy, Yunnan province

Introduction

The phylum Basidiomycota R.T. Moore represents one of the major divisions in the fungal tree of life, with global estimates that it encompasses 1.4–4.2 million species in the phylum and the latest estimates of 0.7 to 1 million species, which represent about 28–40% of all fungal diversity (Hyde 2022; He et al. 2024; Liu et al. 2024). The basidiomata of Basidiomycota exhibit complex forms, such as coralloid, corticioid, gilled, hydnoid, poroid, and toothed basidiomata (Bernicchia and Gorjón 2010; He et al. 2019; Zhou et al. 2024a). Traditionally, Basidiomycota is typically characterized by the basidia and basidiospores and some variable morphological characters, such as diverse cellular constructions in hyphal systems and meiosporangia; the basidiomata reflect a profound evolutionary history at the various taxonomic levels within Basidiomycota (Zhao et al. 2016; Hyde et al. 2023; Dong et al. 2024). Nowadays, DNA sequence-based classification and identification have become the standard methodology in fungal taxonomy (Dai et al. 2021; Zhang et al. 2023; Dong et al. 2024; He et al. 2024).

The genus Clavulinopsis Overeem, classified within the family Clavariaceae (Agaricales, Basidiomycota), includes species that are widely distributed. It is characterized by yellow, orange, or creamy white basidiomata, with simple or regularly dichotomously branched, cylindric or fusoid stems; generative hyphae with obtuse, occasionally inflated hyphae with clamp connections; 2–4-spored basidia; and smooth or echinulate basidiospores occasionally with big guttules (Petersen 1996; Knudsen and Vesterholt 2012; Keleş and Kaya 2021; Yan et al. 2023). More than 171 taxon records of the genus currently are listed in the Index Fungorum database (http://www.indexfungorum.org; accessed on 2 November 2024), and approximately 84 species names are legitimately published (Yan et al. 2023). However, the phylogeny of Clavulinopsis is ambiguous due to a lack of molecular evidence and morphological data. Two similar genera, Clavaria and Ramariopsis, are easily confused with Clavulinopsis in clavarioid basidiomata. However, the micromorphology feathers are different between the three genera; viz., Ramariopsis has various basidiomata, but those in Clavaria and Clavulinopsis are simple; clamp connections are present on the basidia and hyphae in Clavulinopsis (Corner 1950, 1970; Petersen and Corner 1968; Yan et al. 2023). Furthermore, the outline of Clavulinopsis was defined by the basidiospore ornamentation, and the related taxa were examined by D.N. Pegler and T.W.K. Young. Applying the molecular phylogenetic methods generated considerable promotion in the generic definition of Clavulinopsis, gradually clarifying the classification boundaries among several closely related genera, particularly Clavaria and Ramariopsis (Furtado et al. 2016; Yan et al. 2023; He et al. 2024).

Henningsomyces Kuntze, a type of cyphelloid fungi first described by Kuntze (1898), belongs to the family Porotheleaceae (Agaricales) (He et al. 2024). Previously, cyphelloid fungi were classified in the family Cyphellaceae based on similar morphology. Subsequently, the name Porotheleaceae (Murrill 1916) was later established for this group (Cooke 1957, 1961). However, the phylogenetic relationship of many cyphelloid fungi remains ambiguous. The genus Henningsomyces, a typical cyphelloid fungus with cylindric basidiomata, is typified by H. candidus (Pers.) Kuntze and is characterized by annual basidiomata consisting of sparse or gregarious tubes, a monomitic hyphae system that typically exhibits both clamp connections and simple septa, an absence of cystidia, and globose to subglobose basidiospores (Wei and Qin 2009; Liu et al. 2023). Among the 50 records of the genus currently listed in the Index Fungorum database (http://www.indexfungorum.org; accessed on 2 November 2024), approximately 22 species names are legitimately published (Liu et al. 2023). Among those studies, Henningsomyces forms a monophyletic lineage and nests into the order Agaricales (Moncalvo et al. 2006, Bodensteiner et al. 2004, Binder et al. 2005, Thorn et al. 2005, Baltazar et al. 2015, Lucas and Dentinger 2015, Moreno et al. 2017). However, the placement of Henningsomyces and related taxa has not yet been substantiated. In recent years, based on morphological examination and molecular phylogenetic analysis, six new species were described in China (Yan et al. 2023). Most mycologists focused on the poroid or corticioid species, but the cyphelloid species bearing cup-, bowl-, or tube-shaped “cyphelloid” hymenophores were rarely reported, such as the genus Henningsomyces bearing the cylindric basidiomata (Wei and Qin 2009).

Xenasma Donk, classified in the family Xenasmataceae (Russulales, Basidiomycota), was introduced in 1957 and is typified by X. rimicola (P. Karst.) Donk (Liberta 1960; Bernicchia and Gorjón 2010). This genus is characterized by the resupinate and smooth basidiomata, a monomitic hyphal system with clamps, generative hyphae, and globose to cylindrical, striate basidiospores (Liberta 1960; Bernicchia and Gorjón 2010). Among the 39 records of the genus currently listed in the Index Fungorum database (http://www.indexfungorum.org; accessed on 2 November 2024), only 11 species names are legitimately published (Bernicchia and Gorjón 2010; He et al. 2024). Xenasma Donk continues to intrigue mycologists due to its unique morphological characteristics and ecological roles (Bernicchia and Gorjón 2010). The genus is primarily found in temperate forest ecosystems, often growing on decaying wood and contributing to wood decomposition (Liu et al. 2024). Recent phylogenetic studies have expanded the understanding of its evolutionary relationships within the Russulales, highlighting the potential for undiscovered species in understudied regions (He et al. 2019, 2024). The morphology of the basidiome and hymenophore, together with habitat, are often regarded as important characters for the taxonomy of the order Russulales, and Xenasmataceae is the only family in which the smooth hymenophore configuration could be found (Larsson and Larsson 2003; Miller et al. 2006; He et al. 2024). In the latest study, there are only two genera in the family Xenasmataceae, namely Xenasma and Xenosperma Oberw, in which the genus Xenasma is a mystery genus, and no new taxon was reported from this genus for nearly half a century (He et al. 2024).

The specimens of the three genera collected in the Yunnan-Guizhou Plateau, China, which could not be assigned to any described species of the order. Therefore, four new species, viz. Clavulinopsis wumengshanensis, Henningsomyces bambusae, Xenasma bisterigmatae, and X. guttulate, are proposed with descriptions, illustrations, and phylogenetic analysis results.

Materials and methods

Morphology

Fresh fruiting bodies of the fungi were collected from Wumengshan National Nature Reserve in Zhaotong of Yunnan Province, China, and the important collection information was recorded (Rathnayaka et al. 2024). Specimens were dried in an electric food dehydrator at 40 °C (Hu et al. 2022), then sealed and stored in an envelope bag and deposited in the herbarium of the Southwest Forestry University (SWFC), Kunming, Yunnan Province, China. Macromorphological descriptions are based on field notes and photos captured in the field and lab. Color terminology follows Petersen (1996). Micromorphological data were obtained from the dried specimens when observed under a light microscope following the previous study (Zhao et al. 2023; Zhou et al. 2024b). The following abbreviations are used: KOH = 5% potassium hydroxide water solution, CB = Cotton Blue, CB– = acyanophilous, IKI = Melzer’s Reagent, IKI– = both inamyloid and indextrinoid, Lm = mean spore length (arithmetic average for all spores), Wm = mean spore width (arithmetic average for all spores), Q = variation in the L/W ratios between the specimens studied, and n = a/b (number of spores (a) measured from given number (b) of specimens).

Molecular phylogeny

The CTAB rapid plant genome extraction kit-DN14 (Aidlab Biotechnologies Co., Ltd., Beijing) was used to obtain DNA from dried specimens, and PCR was performed according to the manufacturer’s instructions with some modifications. ITS locus was amplified using the primer pairs ITS5/ITS4 (White et al. 1990). The nuclear LSU region was amplified with primer pair LR0R and LR7 (Vilgalys and Hester 1990). 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, 54 °C for 45 s, and 72 °C for 1 min, and a final extension at 72 °C for 10 min. The PCR procedure for LSU was as follows: initial denaturation at 94 °C for 1 min, followed by 35 cycles at 94 °C for 30 s, 50 °C for 1 min, and 72 °C for 1.5 min, and a final extension at 72 °C for 10 min. All newly generated sequences were submitted to GenBank and are listed in Table 1.

Table 1.
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Names, voucher numbers, references, and corresponding GenBank accession numbers of the taxa used in the phylogenetic analyses. [* Indicates type materials;—indicates sequence unavailability].

Taxa Locality Voucher no. GenBank accession no. References
ITS 28S
Clavaria apulica Italy AMB 150 MT853065 MT853066 Agnell and Papetti 2020
Clavulinopsis amoena Australia PBM3381 — HQ877702 Hyde et al. 2016
Clavulinopsis aspersa China MHHNU10153 OQ703777 OQ703794 Yan et al. 2023
Clavulinopsis aspersa China MHHNU10342* OQ703778 OQ703795 Yan et al. 2023
Clavulinopsis aurantiaca Brazil URM<BRA>:84212* — KX227749 Hyde et al. 2016
Clavulinopsis aurantiaca Brazil URM<BRA>:84216 KC348464 NG058946 Hyde et al. 2016
Clavulinopsis bicolor China MHHNU10381* OQ703780 OQ703797 Yan et al. 2023
Clavulinopsis bispora China MHHNU11188 OQ703782 OQ703799 Yan et al. 2023
Clavulinopsis bispora China MHHNU11181* OQ703781 OQ703798 Yan et al. 2023
Clavulinopsis corallinorosacea Australia PBM3380 KP257144 HQ877707 Hyde et al. 2016
Clavulinopsis corniculata USA TENN064106 KP257145 HQ877713 Hyde et al. 2016
Clavulinopsis erubescens China MHHNU10290 OQ703784 OQ703801 Yan et al. 2023
Clavulinopsis erubescens China MHHNU8040* OQ703783 OQ703800 Yan et al. 2023
Clavulinopsis fusiformis USA PBM 2804 — EF535273 Hyde et al. 2016
Clavulinopsis fusiformis USA TENN064110 — HQ877717 Hyde et al. 2016
Clavulinopsis gracillima Canada MO 215748 KY706170 — Hay et al. 2019
Clavulinopsis incarnata China MHHNU11331 OQ703788 OQ703805 Yan et al. 2023
Clavulinopsis incarnata China MHHNU11330* OQ703787 OQ703804 Yan et al. 2023
Clavulinopsis miyabeana China ZP-2118 MK427059 — Chen and Zhang 2019
Clavulinopsis sulcata Australia PBM3379 — HQ877709 Hyde et al. 2016
Clavulinopsis sulcata New Zealand PDD78241 — DQ284904 Dentinger and McLaughlin 2006
Clavulinopsis trigonospora China MHHNU9186 OQ703789 OQ703806 Yan et al. 2023
Clavulinopsis trigonospora Italy AMB: 18557* NR176720 NG088120 Franchi and Marchetti 2021
Clavulinopsis tropicalis China MHHNU10721 OQ703792 OQ703809 Yan et al. 2023
Clavulinopsis tropicalis China MHHNU10722* OQ703793 OQ703810 Yan et al. 2023
Clavulinopsis wumengshanensis China CLZhao 29651 PQ408630 PQ408635 Present Study
Clavulinopsis wumengshanensis China CLZhao 29612* PQ408629 PQ408634 Present Study
Clitocybula familia Slovakia BRNM 736053 JF730328 JF730323 Antonín et al. 2011
Clitocybula intervenosa São Tomé BAP 613 SFSU* MH414561 MH385335 Cooper 2018
Clitocybula lacerata Italy AMB 18779 OM422757 OM423633 Consiglio et al. 2022
Clitocybula lacerata Czech Republic PRM 915404 LT854054 LT854030 Antonín et al. 2019
Clitocybula oculus USA PBM 1156 DQ192178 DQ151452 Matheny et al. 2006
Gerronema keralense India CAL 1666* NR159832 NG064531 Latha et al. 2018
Gerronema kuruvense India CAL 1665* NR159831 NG064530 Latha et al. 2018
Gerronema xanthophyllum Czech Republic PRM 924657 LT854023 LT854023 Antonín et al. 2019
Henningsomyces bambusae China CLZhao 33024 PQ408626 — Present study
Henningsomyces bambusae China CLZhao 33085 PQ408627 — Present study
Henningsomyces bambusae China CLZhao 33088* PQ408628 — Present study
Henningsomyces candidus France PB338 AY571044 AY571008 Bodensteiner et al. 2004
Henningsomyces candidus Canada T156 AY571043 — Bodensteiner et al. 2004
Henningsomyces hengduanensis China LWZ 20190807-22b OR557251 OR527277 Liu et al. 2024
Henningsomyces hengduanensis China LWZ 20190807-11b* OR557250 OR527276 Liu et al. 2024
Hydropodia subalpina Italy AMB 18784 OM422761 OM423638 Consiglio et al. 2022
Hydropodia subalpina Italy AMB 18785 OM422762 OM423639 Consiglio et al. 2022
Hydropodia subalpina Turkey OKA TR-K364 MN701620 MN700170 Kaygusuz et al. 2020
Leucoinocybe lenta Italy AMB 18837 OM422765 OM423643 Consiglio et al. 2022
Leucoinocybe taniae Italy AMB 18838 OM422766 OM423644 Consiglio et al. 2022
Leucoinocybe taniae Italy AMB 18839 OM422767 OM423645 Consiglio et al. 2022
Porotheleum albodescendens New Zealand PDD 96321* OL998343 OL998382 Consiglio et al. 2022
Porotheleum domingense Dominican Republic JBSD 131801* OM422768 OM423646 Consiglio et al. 2022
Porotheleum fimbriatum France CBS 465.50 MH856711 — Vu et al. 2019
Porotheleum parvulum Dominican Republic JBSD 131802* OM422783 OM423657 Consiglio et al. 2022
Pseudohydropus commenticius New Zealand PDD 86984* OL998339 OL998379 Consiglio et al. 2022
Pseudohydropus floccipes Czech Republic BRNM 816173 OM422758 OM423634 Consiglio et al. 2022
Pseudohydropus parafunebris New Zealand PDD 87227* JQ694112 — Consiglio et al. 2022
Pterula echo USA AFTOL-ID 711 DQ494693 — Matheny et al. 2006
Radulomyces copelandii China Dai 15061 KU535664 KU535672 Zhao et al. 2016
Radulotubus resupinatus China Cui 8383* KU535660 KU535668 Zhao et al. 2016
Rectipilus afibulatus UK K(M)189533* KT893457 — Lucas and Dentinger 2015
Xenasma bistaminatae China CLZhao 32542* PQ408631 — Present study
Xenasma bistaminatae China CLZhao 32600 PQ408632 — Present study
Xenasma guttulata China CLZhao 32193* PQ408633 — Present study
Xenasma praeteritum USA Alden Dirks:ACD0185 OM009268 — Unpublished
Xenasma pruinosum Japan OTU1299 MT594801 — Unpublished
Xenasma rimicola Australia N.L. Bougher NLB 1449 MT537020 — Unpublished
Xenasma rimicola Australia N.L. Bougher NLB 1571 MT571671 — Unpublished

Sequences generated for this study were aligned, with additional sequences downloaded from GenBank. Sequences were aligned using MAFFT v.7 (https://mafft.cbrc.jp/alignment/server/), adjusting the direction of nucleotide sequences according to the first sequence (accurate enough for most cases) and selecting the G-INS-i iterative refinement method (Katoh et al. 2019). Alignments were manually adjusted to maximize alignment and minimize gaps with BioEdit v.7.0.9 (Hall 1999). A dataset of concatenated ITS and LSU sequences was used to determine the phylogenetic position of the new species. Maximum likelihood (ML) analysis was performed using the CIPRES Science Gateway (Miller et al. 2010) based on the dataset using the RA × ML-HPC BlackBox tool, with the setting RA × ML halt bootstrapping automatically and 0.25 for maximum hours and obtaining the best tree using ML search. Other parameters in ML analysis followed default settings, and statistical support values were obtained using nonparametric bootstrapping with 1,000 replicates. Maximum parsimony (MP) analyses were applied to the combined three datasets following the methods outlined in a previous study (Zhao and Wu 2017), and the tree construction procedure was performed in PAUP* version 4.0b10 (Swofford 2002). Bayesian inference (BI) analysis based on the dataset was performed using MrBayes v.3.2.6 (Ronquist and Huelsenbeck 2003). The best substitution model for the dataset was selected by ModelFinder (Kalyaanamoorthy et al. 2017) using a Bayesian information criterion, and the model was used for Bayesian analysis. Four Markov chains were run from random starting trees. Trees were sampled every 1,000th generation. The first 25% of sampled trees were discarded as burn-in, whereas other trees were used to construct a 50% majority consensus tree and for calculating Bayesian posterior probabilities (BPPs). The bootstrap support for ML is greater than or equal to 70%, and Bayesian posterior probabilities greater than or equal to 0.95 are indicated on the branches in the phylogenetic tree, respectively.

Results

The phylogeny of Clavulinopsis

The dataset included ITS+LSU sequences from 27 samples representing 17 taxa. The datasets had an aligned length of 2,467 characters, of which 1,775 characters are constant, 273 are variable and parsimony-uninformative, and 419 are parsimony-informative. Maximum parsimony analysis yielded 1 equally parsimonious tree (TL = 1359, CI = 0.6799, HI = 0.5829, RI = 0.7129, RC = 0.4847). BI analysis yielded a similar topology to ML analysis, with an average standard deviation of split frequencies of 0.012252; trees were sampled every 1,000th generation, 0.4 million in total. The effective sample size (ESS) for Bayesian analysis across the two runs is double the average ESS (avg ESS) = 201. Branches that received bootstrap support for ML ≥ 70%, MP ≥ 50%, and BI ≥ 0.95 were considered significantly supported, respectively. The ML tree was provided (Fig. 1). The phylogenetic tree (Fig. 1) reveals that the new species Clavulinopsis wumengshanensis is nested into the genus Clavulinopsis and has a close relationship with C. aurantiocinnabarina (Schwein.) Corner with full support (100/100/1.00).

Figure 1. 

Phylogeny of species in Clavulinopsis generated by maximum likelihood based on ITS+LSU sequence data. Branches are labeled with maximum likelihood bootstrap ≥ 70%, a reduced lead value greater than 50%, and Bayesian posterior probabilities ≥ 0.95, respectively.

The phylogeny of Henningsomyces

The dataset included ITS sequences from 31 samples representing 23 taxa. The datasets had an aligned length of 2,375 characters, of which 1,204 characters are constant, 525 are variable and parsimony-uninformative, and 646 are parsimony-informative. Maximum parsimony analysis yielded 1 equally parsimonious tree (TL = 2,997, CI = 0.5706, HI = 0.4294, RI = 0.5875, RC = 0.3352). The BI analysis yielded a similar topology to the ML analysis, with an average standard deviation of split frequencies of 0.007176; trees were sampled every 1,000th generation, 0.4 million in total. And the effective sample size (ESS) for Bayesian analysis across the two runs is double the average ESS (avg ESS) = 349. Branches that received bootstrap support for ML ≥ 70%, MP ≥ 50%, and BI ≥ 0.95 were considered significantly supported, respectively. The ML tree was provided (Fig. 2). The phylogenetic tree (Fig. 2) reveals the new species Henningsomyces bambusae nested into the genus Henningsomyces and has a close relationship with H. candidus (Pers.) Kuntze with full support (100/100/1.00).

Figure 2. 

Phylogeny of species in Henningsomyces generated by maximum likelihood based on ITS sequence data. Branches are labeled with maximum likelihood bootstrap ≥ 70%, a reduced lead value greater than 50%, and Bayesian posterior probabilities ≥ 0.95, respectively.

The phylogeny of Xenasma

The dataset included ITS sequences from nine samples representing seven taxa. The datasets had an aligned length of 944 characters, of which 574 characters are constant, 154 are variable and parsimony-uninformative, and 216 are parsimony-informative. Maximum parsimony analysis yielded 1 equally parsimonious tree (TL = 520, CI = 0.8942, HI = 0.1058, RI = 0.8243, RC = 0.7371). The BI analysis yielded a similar topology to the ML analysis, with an average standard deviation of split frequencies of 0.007297; trees were sampled every 1,000th generation, 0.4 million in total. And the effective sample size (ESS) for Bayesian analysis across the two runs is double the average ESS (avg ESS) = 768.5. Branches that received bootstrap support for ML ≥ 70%, MP ≥ 50%, and BI ≥ 0.95 were considered significantly supported, respectively. The ML tree was provided (Fig. 3). The phylogenetic tree (Fig. 3) reveals the two new species, Xenasma bisterigmatae and X. guttulate, nested into the genus Xenasma. The taxon X. bisterigmatae has a close relationship with X. rimicola (P. Karst.) Donk, while the taxon X. guttulata has a close relationship with X. pruinosum with strong support (97/100/0.96).

Figure 3. 

Phylogeny of species in Xenasma generated by maximum likelihood based on ITS sequence data. Branches are labeled with maximum likelihood bootstrap ≥ 70%, a reduced lead value greater than 50%, and Bayesian posterior probabilities ≥ 0.95, respectively.

The BLAST result of four new species for the closest top 10 taxa and their corresponding parameters are given (Table 2).

Table 2.
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The BLAST result of four new species for the closest top 10 taxa and their corresponding parameters.

Scientific Name Specimens Sequence Number Max score Total score Query cover E value Per. Ident. Acc. len. Accession
Clavulinopsis wumengshanensis CLZhao 29612 PQ408629 1123 1123 100% 0 100% 638 PQ408629
734 734 71% 0 97.24% 431 PQ515870
448 448 44% 6e-121 96.68% 295 PQ453614
911 911 93% 0 95.94% 573 ON943319
972 972 100% 0 95.60% 627 PQ346233
970 970 100% 0 95.59% 626 PQ346239
976 976 100% 0 95.57% 624 PQ346237
966 966 100% 0 95.43% 653 PQ453611
966 966 100% 0 95.43% 632 PQ346229
966 966 100% 0 95.43% 627 PQ346265
Clavulinopsis wumengshanensis CLZhao 29612 PQ408634 1024 1024 100% 0 100.00% 638 PQ408629
896 896 100% 0 95.85% 643 PQ408630
885 885 99% 0 95.99% 627 PQ346233
881 881 100% 0 95.32% 624 PQ346237
880 880 99% 0 95.81% 611 PQ453612
880 880 99% 0 95.81% 653 PQ453611
880 880 99% 0 95.81% 626 PQ346239
880 880 99% 0 95.81% 632 PQ346229
880 880 99% 0 95.81% 627 PQ346265
878 878 99% 0 95.63% 590 PQ453613
Henningsomyces bambusae CLZhao 33088 PQ408628 1282 1282 100% 0 100.00% 694 PQ408628
1253 1253 98% 0 99.85% 694 PQ408626
1223 1223 97% 0 99.26% 697 PQ408627
1081 1081 96% 0 95.71% 701 LC774058
1077 1077 96% 0 95.56% 733 PP849903
1072 1072 96% 0 95.41% 732 MG707601
1059 1059 96% 0 951% 745 MK607599
985 985 89% 0 95.22% 643 AB847016
950 950 89% 0 94.38% 622 AY571057
813 813 90% 0 903% 652 OQ872100
Xenasma bisterigmatae CLZhao 32542 PQ408631 1138 1138 100% 0 100.00% 616 PQ408631
1040 1040 99% 0 97.55% 611 PQ408632
846 846 92% 0 93.58% 634 MT537020
758 676 83% 0 92.83% 676 MT571671
597 597 60% 6e-166 95.48% 411 LR819650
597 597 60% 6e-166 95.48% 411 LR602855
508 508 72% 3e-139 87.25% 636 JF691144
503 503 91% 1e-137 86.96% 675 PQ408633
492 492 59% 3e-134 90.98% 638 OM009268
431 431 60% 6e-116 87.47% 417 LR819360
Xenasma guttulata CLZhao 32193 PQ408633 1247 1247 100% 0 100.00% 675 PQ408633
737 737 91% 0 88.36% 636 JF691144
634 634 91% 5e-177 85.53% 638 OM009268
592 592 55% 3e-164 94.97% 417 LR819360
592 592 55% 3e-164 94.97% 417 LR602599
586 586 55% 1e-162 94.71% 417 LR602541
586 586 55% 1e-162 94.71% 417 LR819292
558 558 55% 3e-154 93.39% 417 LR602585
558 558 55% 3e-154 93.39% 417 LR819344
507 507 47% 1e-138 95.34% 417 MT852420

Taxonomy

Clavulinopsis wumengshanensis S.Y. He., H.M. Zhou & C.L. Zhao, sp. nov.

MycoBank No: 855906
Figs 4, 5

Diagnosis.

Clavulinopsis wumengshanensis differs from C. aurantiocinnabarina by buff-yellow to straw-yellow basidiomata and thick-walled, subglobose, and longer basidiospores (7–8.5 µm vs. 5.6–7.1 µm).

Holotype.

China • Yunnan Province, Zhaotong, Yiliang County, Wumengshan National Nature Reserve, GPS coordinates 27°30'N, 104°12'E, evel. 1710 m asl., on the ground, leg. C.L. Zhao, 12 July 2023, CLZhao 29612 (SWFC).

Etymology.

Wumengshanensis (Lat.) refers to the locality “Wumengshan National Nature Reserve” of the holotype.

Basidiomata.

Basidiomes annual, clavarioid, without odor or taste when fresh, up to 8 cm long, 4 mm wide, and 400–800 µm thick. Fertile part subcylindrical to fusiform, occasionally slightly curved or flexuous and with a distinct longitudinal depression, buff-yellow (4A4) to straw-yellow (3A/B3) when dry. Apex rounded or obtuse acute when mature, concolourous or slightly paler.

Figure 4. 

Basidiomata of Clavulinopsis wumengshanensis (holotype, CLZhao 29612). Scale bars: 1 cm (A); 1 mm (B).

Hyphal structure.

Monomitic, generative hyphae with clamp connections, hyaline, thick-walled, parallel, interwoven, 2–3 µm in diam, some inflated to 13 µm in diam, IKI–, CB–; tissues unchanged in KOH. Cystidia absent. Basidia clavate to subcylindrical, hyaline, thin-walled, with four sterigmata and a basal clamp connection, 35–48.5 × 9–11.5 µm; basidioles in shape similar to basidia, with several guttules.

Figure 5. 

Microscopic structures of Clavulinopsis wumengshanensis (holotype, CLZhao 29612) A basidiospores B basidia and basidioles C a section of basidiomata. Scale bars: 10 µm (A–C).

Spores.

Basidiospores subglobose with a distinct apiculus, hyaline, thick-walled, smooth, with several guttules, IKI–, CB–, 7–8.5(–5) × (5.5–)6–7.5 µm, L = 7.59 µm, W = 6.54 µm, Q = 1.14–1.16 (n = 60/2).

Additional specimen examined

(paratype). China • Yunnan Province, Zhaotong, Yiliang County, Wumengshan National Nature Reserve, GPS coordinates 27°30'N, 104°12'E, evel. 1710 m asl., on the ground, leg. C.L. Zhao, 12 July 2023, CLZhao 29651 (SWFC).

Notes.

Based on the ITS + LSU analysis (Fig. 1), the result showed that the new species Clavulinopsis wumengshanensis is grouped with C. aurantiocinnabarina (Schwein.) Corner. However, C. aurantiocinnabarina differs from C. wumengshanensis by its thin-walled basidiospores and narrower basidia (5.2–7.1 µm vs. 9–11.5 µm, Petersen 1978).

Henningsomyces bambusae S.Y. He, H.M. Zhou & C.L. Zhao, sp. nov.

MycoBank No: 855907
Figs 6, 7

Diagnosis.

Henningsomyces bambusae differs from H. candidus by its globose to subglobose and wider basidiospores (6.5–8.5 µm vs. 4–5 µm).

Holotype.

China • Yunnan Province, Zhaotong, Wumengshan National Nature Reserve, GPS coordinates 27°29'N, 103°55'E, evel. 1900 m asl., on dead bamboo, leg. C.L. Zhao, 18 September 2023, CLZhao 33088 (SWFC).

Etymology.

Bambusae (Lat.) refers to the host genus Bambusa.

Basidiomata.

Basidiomes forming loosely scattered colonies, short cylindric to tubular, sessile, up to 1–1.6 cm in length, 0.5–1 cm diam; external surface white (60) to cream (4A2/3), finely tomentose when dry; inner surface covered by a deeply concave, smooth hymenium.

Figure 6. 

Basidiomata of Henningsomyces bambusae (holotype, CLZhao 33088). Scale bars: 1 cm (A); 1 mm (B).

Hyphal structure.

Monomitic, generative hyphae with clamp connections and simple septa, thin- to thick-walled, IKI–, CB–; tissues unchanged in KOH. External hyphae slightly contorted, widest at the base, slightly tapering towards the obtusely rounded apex, hyaline, thick-walled to solid, occasionally with simple septa, frequently unbranched, dextrinoid, CB–, 3–4 μm diam; tramal hyphae hyaline, thin-walled, with clamp connections, IKI–, 2.5–3 µm diam. Cystidia absent. Basidia cylindrical to subcylindrical, hyaline, thin-walled, with two sterigmata and a basal clamp connection, 19.5–22.5 × 7–8.5 µm; basidioles in shape similar to basidia, but slightly smaller.

Figure 7. 

Microscopic structures of Henningsomyces bambusae (holotype, CLZhao 33088) A basidiospores B basidia and basidioles C a section of basidiomata. Scale bars: 10 µm (A–C).

Spores.

Basidiospores globose to subglobose, hyaline, thin-walled, smooth, with guttules, IKI–, CB–, 6.5–8.5(–9) × (6–)6.5–8.5(–9) µm, Lm = 7.61 µm, Wm = 7.45 µm, Q = 1.02–1.04 (n = 60/2).

Additional specimens examined

(paratypes). China • Yunnan Province, Zhaotong, Wumengshan National Nature Reserve, GPS coordinates 27°29'N, 103°55'E, evel. 1900 m asl., on a dead bamboo, leg. C.L. Zhao, 18 September 2023, CLZhao 33024; CLZhao 33085 (SWFC).

Notes.

Based on the ITS analysis (Fig. 2), the new taxon Henningsomyces bambusae grouped within the genus Henningsomyces and formed a sister group with H. candidus (Pers.) Kuntze. However, H. candidus can be delimited from H. bambusae by its narrower basidiospores (4–5 μm vs. 6–8 μm; Gilbertson and Blackwell 1985).

Xenasma bisterigmatae S.Y. He, H.M. Zhou & C.L. Zhao, sp. nov.

MycoBank No: 855908
Figs 8, 9

Diagnosis.

Xenasma bisterigmatae differs from X. rimicola by its ellipsoid to subglobose and larger basidiospores (8.5–10.5 × 4.5–7 µm vs. 10–12.5 × 8–10.5 µm), two-sterigmata basidia.

Holotype.

China • Yunnan Province, Zhaotong, Wumengshan National Nature Reserve, GPS coordinates 27°29'N, 103°55'E, evel. 1900 m asl., on fallen angiosperm branch, leg. C.L. Zhao, 29 August 2023, CLZhao 32542 (SWFC).

Etymology.

Bisterigmatae (Lat.) refers to two-sterigmata basidia of the holotype.

Basidiomata.

Basidiomes annual, resupinate, smooth, without odor or taste when fresh, up to 6.5 cm long, 2.3 cm wide, and 100 µm thick. Hymenial surface smooth, membranaceous, ash-grey (19C2) when dry; sterile margin indistinct.

Figure 8. 

Basidiomata of Xenasma bisterigmatae (holotype, CLZhao 32542). Scale bars: 1 cm (A); 1 mm (B).

Hyphal structure.

Monomitic, generative hyphae with clamp connections, thin-walled, branched, interwoven, with dense crystal, 3.2–3.5 µm diam, IKI–, CB–; tissues unchanged in KOH. Cystidia abundant, tubular with obtuse apex, with slightly thick walls in the basal part that appears frequently collapsed, often with an apical amorphous globule, 64.5–93 × 5.5–7.5 µm. Cystidioles absent. Basidia clavate to subcylindrical, thin-walled, with two sterigmata and a basal clamp connection, with several guttules, 21.5–28 × 10–12 µm; basidioles in shape similar to basidia, but slightly smaller.

Figure 9. 

Microscopic structures of Xenasma bisterigmatae (holotype, CLZhao 32542) A basidiospores B basidia and basidioles C cystidia D a section of basidiomata. Scale bars: 10 µm (A–C).

Spores.

Basidiospores ellipsoid to subglobose, thick-walled, verrucose, IKI–, CB–, 10–12.5 × 8–10.5(–11) µm, Lm = 11.15 µm, Wm = 9.3 µm, Q = 1.19–1.27 (n = 60/2).

Additional specimens examined

(paratype). China • Yunnan Province, Zhaotong, Wumengshan National Nature Reserve, GPS coordinates 27°29'N, 103°55'E, evel. 1900 m asl., on fallen angiosperm branch, leg. C.L. Zhao, 29 August 2023, CLZhao 32600 (SWFC).

Notes.

Based on the ITS analysis (Fig. 3), the new species Xenasma bisterigmatae was clustered with X. rimicola (P. Karst.) Donk. However, X. rimicola differs from X. bisterigmatae by its four-sterigmata basidia (Cunningham 1963).

Xenasma guttulata S.Y. He, H.M. Zhou & C.L. Zhao, sp. nov.

MycoBank No: 855909
Figs 10, 11

Diagnosis.

Xenasma guttulata differs from X. pruinosum by its membranaceous basidiomata with a white to cream hymenial surface and larger basidiospores (6–7 × 3–4 µm vs. 7–9 × 5.5–7.5 µm).

Holotype.

China • Yunnan Province, Zhaotong, Wumengshan National Nature Reserve, GPS coordinates 27°29'N, 103°55'E, evel. 1900 m asl., on a fallen angiosperm branch, leg. C.L. Zhao, 29 August 2023, CLZhao 32193 (SWFC).

Etymology.

Guttulata (Lat.) refers to the basidiospores with guttules in the holotype.

Basidiomata.

Basidiomes annual, resupinate, closely adnate, without odor or taste when fresh, up to 2.5 cm long, 1.5 cm wide, and 100 µm thick. Hymenial surface smooth, membranaceous, white (60) to cream (4A2/3) when dry; sterile margin indistinct.

Figure 10. 

Basidiomata of Xenasma guttulata (holotype, CLZhao 32193). Scale bars: 1 cm (A); 1 mm (B).

Hyphal structure.

Monomitic, generative hyphae with clamp connections, thin-walled, occasionally branched, interwoven, with dense crystal, 3.3–3.5 µm diam, IKI–, CB–; tissues unchanged in KOH. Cystidia abundant, cystidia tubular with obtuse apex, 75–91.5 × 4–8.5 µm, with slightly thick walls in the basal part that appears frequently collapsed, often with an apical amorphous globule. Cystidioles absent. Basidia clavate to subcylindrical, thin-walled, with two sterigmata and a basal clamp connection, 18.5–24 × 7.5–11 µm; basidioles in shape similar to basidia, but slightly smaller.

Figure 11. 

Microscopic structures of Xenasma guttulata (holotype, CLZhao 32193) A basidiospores B basidia and basidioles C cystidia D a section of basidiomata. Scale bars: 10 µm (A–C).

Spores.

Basidiospores ellipsoid to narrowly ellipsoid, thick-walled, verrucose, with several guttules, IKI–, CB–, 7–9 × 5.5–7.5 µm, Lm = 8.05 µm, Wm = 6.52 µm, Q = 1.23 (n = 30/1).

Notes.

Based on the ITS analysis (Fig. 3), the new species Xenasma guttulata was clustered with X. pruinosum (Pat.) Donk. However, X. pruinosum can be distinguished from X. guttulata by its smaller basidiospores (6–7 × 3–4 µm vs. 7–9 × 5.5–7.5; Donk 1957; Bernicchia and Gorjón 2010).

Discussion

The multilocus (ITS+LSU) analysis (Fig. 1) showed that the new species Clavulinopsis wumengshanensis groups with C. aurantiocinnabarina (Schwein.) Corner. However, C. aurantiocinnabarina differs from C. wumengshanensis by its thin-walled basidiospores and narrower basidia (5.2–7.1 µm vs. 9–11.5 µm) (Petersen 1978). Morphologically, Clavulinopsis fusiformis (Sowerby) Corner and C. incarnata P. Zhang & Jun Yan are similar to C. wumengshanensis by both having smooth, globose to subglobose basidiospores (Keleş and Kaya 2021; Yan et al. 2023). However, C. fusiformis differs in its pale yellow basidiomata, and C. incarnata differs in its pinkish basidiomata (Keleş and Kaya 2021; Yan et al. 2023).

Based on the ITS locus phylogenetic analysis (Fig. 2), the new taxon Henningsomyces bambusae forms a sister group with H. candidus (Pers.) Kuntze. However, H. candidus can be delimited from H. bambusae by its narrower basidiospores (4–5 μm vs. 6–8 μm, Gilbertson and Blackwell 1985). Morphologically, Henningsomyces hengduanensis S.L. Liu & L.W. Zhou and H. minimus (Cooke and W. Phillips) Kuntze are similar to H. bambusae by both having thin-walled, smooth, subglobose basidiospores (Wei and Qin 2009; Liu et al. 2023). However, H. hengduanensis differs in its cream basidiomata and broadly clavate basidia with four sterigmata (Liu et al. 2023); Henningsomyces minimus differs in its shorter (11–15 μm vs. 19.5–22.5 μm), broadly clavate basidia with four sterigmata (Wei and Qin 2009).

Based on the combined ITS locus phylogeny (Fig. 3), X. rimicola (P. Karst.) Donk differs from the new species X. bisterigmatae by its four-sterigmata basida (Cunningham 1963); X. pruinosum (Pat.) Donk can be distinguished from the new species X. guttulata by its smaller basidiospores (6–7 × 3–4 µm vs. 7–9 × 5.5–7.5, Donk 1957; Bernicchia and Gorjón 2010). Morphologically, X. pruinosum is similar to X. bisterigmatae by having tubular cystidia with an apical amorphous globule and slightly thick walls in the basal part (Donk 1957; Bernicchia and Gorjón 2010). However, X. pruinosum differs in its smaller basidiospores (6–7 × 3–4 µm vs. 7–9 × 5.5–7.5 µm; Donk 1957; Bernicchia and Gorjón 2010). Xenasma rimicola is similar to X. guttulata by having tubular cystidia and slightly thick walls in the basal part (Donk 1957). However, X. rimicola differs in its subclavate basidia with four sterigmata (Donk 1957).

Yunnan Province is located in the southwest of China. The climate in Yunnan offers conducive environments for the speciation and diversification of various life forms, and this climatic diversity in Yunnan creates varied landscapes with multiple habitats, resulting in a high species diversity, with over 6,000 recorded fungal species up to now (Wang and Cai 2022; Dong et al. 2024; Wang et al. 2024; Zhou et al. 2024b). Therefore, focusing on the diversity of fungi in the Yunnan-Guizhou Plateau of China is of great significance. Based on the present study, the results not only enrich the species diversity of fungi worldwide but also contribute to the branches of the fungal tree of life.

Acknowledgments

We would like to express our gratitude to Chunqin Zhou (Yunnan Wumeng Mountains National Nature Reserve, Zhaotong, China) for helping us to search fungal specimens.

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 No. 3246006), the Scientific Research Fund of Yunnan Provincial Department of Education (2024J0668), and the Forestry Innovation Programs of Southwest Forestry University (Grant No: LXXK-2023M07).

Author contributions

Data curation: HSY, WL, ZHM. Formal analysis: HSY, WL, SKZ, ZHM. Methodology: HSY, ZHM. Software: HSY, ZHM. Writing—original draft: HSY, WL, SKZ. Writing—review and editing: ZHM.

Author ORCIDs

Siyuan He https://orcid.org/0009-0005-2478-0624

Lu Wang https://orcid.org/0009-0004-6274-5953

Hongmin Zhou https://orcid.org/0000-0002-0724-5815

Data availability

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

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Siyuan He and Lu Wang contributed equally to this work.
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