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Article

Taxonomy and Phylogeny of Stipitate Stereoid Basidiomycetes from China

by
Jia-Xue Liu
1,2,
Lin-Jiang Zhou
1,2,
Ya-Quan Zhu
1,2,
Hyang Burm Lee
3 and
Hai-Sheng Yuan
1,*
1
CAS Key Laboratory of Forest Ecology and Silviculture, Institute of Applied Ecology, Chinese Academy of Sciences, Shenyang 110016, China
2
University of Chinese Academy of Sciences, Beijing 101408, China
3
Environmental Microbiology Lab, Department of Agricultural Biological Chemistry, College of Agriculture & Life Sciences, Chonnam National University, Gwangju 61186, Republic of Korea
*
Author to whom correspondence should be addressed.
J. Fungi 2026, 12(6), 400; https://doi.org/10.3390/jof12060400
Submission received: 23 March 2026 / Revised: 18 May 2026 / Accepted: 29 May 2026 / Published: 31 May 2026
(This article belongs to the Special Issue Diversity, Phylogeny and Ecology of Forest Fungi, 2nd Edition)
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Abstract

Stipitate stereoid fungi are saprotrophic basidiomycetes characterized by a leathery basidiome, a central-to-lateral stipe and infundibuliform pilei. Although numerous species of stipitate stereoid fungi have been recorded worldwide, understanding of their phylogenetic relationships remains extremely limited, and research on this group of fungi in China is insufficient. In this study, specimens of the three stipitate stereoid genera, namely Podoscypha s. l., Cymatoderma s. l. and Stereopsis s. l., from southern China were investigated. Phylogenetic analyses of the internal transcribed spacer (ITS) regions and the large subunit of the nuclear ribosomal RNA gene (LSU) using maximum likelihood (ML) and Bayesian inference (BI) methods revealed that all three genera are polyphyletic. Consequently, Podoscypha s. s. and Cymatoderma s. s. were delimited, and Cladoderris—previously synonymized with Cymatoderma—was resurrected. Cladoderris is characterized by an imbricate basidiome, tomentose pilei and basidiospores typically shorter than 4 μm in length. Three new species, Podoscypha casiae, Stereopsis buccinata and Cladoderris perennis, were described and illustrated. The morphological distinctions and affinities between the new species and closely related taxa were discussed, the thresholds for the intraspecific and interspecific demarcation within the three genera in this study were provided, and identification keys for the species of each genus were presented.

1. Introduction

Stipitate stereoid fungi are a group of basidiomycetes characterized by the presence of stipes, infundibuliform pilei, a smooth hymenial surface, and smooth basidiospores [1]. This group includes Podoscypha Pat., Cymatoderma Jungh., Cotylidia P. Karst., and Stereopsis D.A. Reid [1,2,3]. These fungi have a global distribution, ranging from tropical to boreal zones, and typically inhabit fallen trunks, stumps and other lignocellulose debris or grow on the ground. As decomposers, they facilitate material and nutrient cycling and play important roles in terrestrial ecosystems.
The genus Podoscypha, with its type species Podoscypha nitidula (Berk.) Pat., was first named and described in 1900 by the French mycologist Pierre Antoine Patouillard [4]. The genus is widely distributed throughout the tropical and temperate regions globally, usually growing on fallen wood or on the ground. In early taxonomic studies, species of this genus were initially classified in the family Polyporaceae, subsequently transferred to Thelephoraceae, and ultimately placed in Podoscyphaceae (Polyporales) based on molecular phylogeny [5,6,7,8,9,10]. According to MycoBank (http://www.MycoBank.org, accessed on 1 May 2026) and Index Fungorum (http://www.indexfungorum.org, accessed on 1 May 2026), the genus comprises 57 legitimate species, and 17 Podoscypha species have been reported or described from China [10,11,12,13,14,15,16,17]. Among them, P. densidisca, P. guangxiensis, P. infundibula, P. lactea, P. petalodes subsp. cystidiata, P. tropica, and P. yunnanensis have been discovered and identified as new species in Guangxi and Yunnan provinces, China [10,13,15].
Cladoderris was established by Persoon in 1826 [18], with C. dendritica as the type. Junghuhn established Cymatoderma in 1840 [19], but Fries [20] later adopted Cladoderris, causing Cymatoderma to be overlooked. This resulted in the misclassification of numerous species until Reid [21,22] confirmed the priority of Cymatoderma and formally synonymized Cladoderris. With the advent of molecular identification techniques and a reevaluation of morphological characteristics, species formerly assigned to Cladoderris have since been transferred to Cymatoderma, Stereopsis, Craterellus, and other genera [2,21,23]. Nevertheless, the taxonomic placement of a small number of species remains uncertain due to the lack of molecular data. The genus Cymatoderma is predominantly found in tropical regions, comprising 15 legitimate species. It has not yielded any new species from China.
The genus Stereopsis, whose type species is Stereopsis radicans (Berk.) D.A. Reid, was originally described and defined in 1965 by the British mycologist David Arthur Reid [2] and is known in most tropical and subtropical regions of the world, usually growing on the ground. With the advancement of molecular systematic studies, it has been classified in the family Stereopsidaceae (Stereopsidales) [24,25]. The genus comprises 22 legitimate species [24,25,26,27], among which one novel species (S. yunnanensis) has been described from China [25].
The genus Cotylidia, typified by Cotylidia undulata (Fr.) P. Karst., was established in 1881 by the mycologist Petter Karsten [28] and recently classified in Rickenellaceae (Hymenochaetales) [1,29,30]. A recent phylogenetic study revealed its polyphyletic status, confirmed Cotylidia sensu stricto, and proposed a new genus, Neocotylidia Jing Si & Hai J. Li [31]. As the taxonomy of Cotylidia has been recently revised, it is not included in the present study.
In recent decades, research on wood-inhabiting fungi in China has made significant progress [32,33,34], yet studies on stipitate stereoid basidiomycetes remain limited. In the present study, dozens of stipitate stereoid fungal specimens from China have been collected, and several undescribed taxa have been identified based on morphological characteristics and phylogenetic analysis of ITS and LSU rDNA sequences. The objectives of this study are to describe the new taxa of stipitate stereoid basidiomycetes from China and to clarify the taxonomic status of Podoscypha and Cymatoderma as well as their intergeneric relationships using multi-gene phylogenetic analyses.

2. Materials and Methods

2.1. Specimen Collection and Morphological Studies

Specimens were collected from Guangzhou South China National Botanical Garden (113°22′2″ E, 23°10′54″ N, altitude: 20–327 m), Shenzhen Fairy Lake Botanical Garden (114°10′32″ E, 22°34′56″ N, altitude: 60–944 m), and Shenzhen Wutong Mountain (114°13′11″ E, 22°35′05″ N, altitude: 692–944 m), Guangdong Province, South China. The regions belong to the South Asian tropical monsoon climate zone. The fresh fruiting bodies of the fungi were dried using an electronic dryer (Evermat, Bjurholm, Sweden) set at temperatures of 40–50 °C. After drying, they were labeled and stored in plastic bags and envelopes. The dried materials studied in this paper are deposited in the herbarium of the Institute of Applied Ecology, Chinese Academy of Sciences (IFP).
The macro-morphological features of the fungal specimens were observed either directly or using a stereomicroscope (Nikon SMZ 1000, Tokyo, Japan). Color terms were defined based on the standards provided by Rayner [35] and Munsell [36]. Micro-morphological data were obtained from dried specimens and observed under a light microscope (Nikon Eclipse E600 microscope, Tokyo, Japan) following the methods described by Mu et al. [37]. Samples for microscopic examination were individually mounted in cotton blue (OriLeaf, Shanghai, China), Melzer’s reagent (OriLeaf, Shanghai, China), and 5% potassium hydroxide (KOH, Damao, Tianjin, China). The range of basidiospores provided in the study excludes the smallest and largest 5% of measurements, denoted in parentheses. The following abbreviations are used in this study: KOH = 5% potassium hydroxide water solution, CB = cotton blue, CB+ = cyanophilous, CB− = acyanophilous, IKI = Melzer’s reagent, IKI− = both inamyloid and indextrinoid, L = mean spore length, W = mean spore width, Q = the range of length-to-width ratios of the basidiospores studied, Qm = mean length-to-width ratios of the basidiospores studied and n for the total number of basidiospores.

2.2. Molecular Analysis

2.2.1. DNA Extraction and PCR

Genomic DNA was extracted from the fungal fruiting bodies using the Rapid Fungi Genomic DNA Isolation Kit, produced by Demeter Biotech Ltd., Beijing, China. The extracted DNA was subsequently used in polymerase chain reaction (PCR) procedures. The PCR amplification system is shown in Table 1. The 2× Taq PCR Mix was purchased from Tiangen Biotech (Beijing) Co., Ltd., Beijing, China.
Two molecular markers were amplified by PCR: the internal transcribed spacer (ITS) region of nuclear ribosomal DNA and the partial large subunit (nLSU) of the nuclear ribosomal RNA gene. The primer pairs used for amplification are listed as follows:
ITS region: ITS1 (5′-CTTGGTCATTTAGAGGAAGTAA-3′) and ITS4 (5′-TCCTCCGCTTATTGATATGC-3′) [38];
nLSU region: LROR (5′-ACCCGCTGAACTTAAGC-3′) and LR7 (5′-TACTACCACCAAGATCT-3′) [39].
The PCR amplification programs for the two markers were set as follows:
ITS region: Initial denaturation at 95 °C for 3 min, followed by 34 cycles of denaturation at 95 °C for 30 s, annealing at 58 °C for 30 s, extension at 72 °C for 60 s, final extension at 72 °C for 5 min, and holding at 14 °C.
LSU region: Initial denaturation at 95 °C for 3 min, followed by 39 cycles of denaturation at 95 °C for 30 s, annealing at 47.2 °C for 30 s, extension at 72 °C for 60 s, final extension at 72 °C for 5 min, and holding at 14 °C.

2.2.2. Sequencing and Sequence Assembly

DNA sequencing was performed at the Beijing Genomics Institute (BGI), and all sequences generated in this study have been submitted to GenBank. The newly obtained sequences were then compared against those in the NCBI GenBank database using the BLAST+ 2.16.0 search tool (https://blast.ncbi.nlm.nih.gov, accessed on 27 September 2024) [40]. Sequence alignments were performed using ClustalX 1.8 [41]. The alignments were manually adjusted to achieve optimal alignment and minimal gaps. Sequence identity and similarity calculations were performed using BioEdit 5.0.6 software [42]. The additional ITS rDNA and nLSU rDNA sequences included in the dataset, which were used to infer phylogenetic relationships, were retrieved from GenBank (http://www.ncbi.nlm.nih.gov/genbank, accessed on 6 August 2025), as detailed in Table 2.

2.3. Phylogenetic Analyses

Phylogenetic relationships were inferred using both maximum likelihood (ML) and Bayesian inference (BI) analyses. The ML analysis was conducted using RAXMLGUI 2.0 [67,68]. The analysis utilized the rapid bootstrap algorithm and the GTR + I + G4 model of nucleotide substitution with 1000 bootstrap replicates.
BI analysis was performed using MrBayes 3.2.7 [69]. The nucleotide substitution model was determined as GTR + I + G based on the Bayesian information criterion (BIC) (Nst = 6, Rates = Invgamma). MCMC analysis was run for 10,000,000 generations, sampling every 4000 generations and printing trees/log-likelihood values every 1000 generations. Two independent runs were performed, each with eight chains. After discarding the initial 25% of trees as burn-in, posterior probabilities were estimated from the remaining trees to assess nodal support. Convergence of the two independent runs was assessed by the average standard deviation of split frequencies (ASDSF), with a value below 0.05 considered indicative of adequate convergence.

2.4. Genetic Distance Analysis and Species Delimitation Method

The genetic distance analysis was conducted in MEGA11 [70]. Specifically, the following parameters were used: the Kimura 2-parameter model was selected as the substitution model [71], and the bootstrap method was set to 10,000 replicates. In addition, rates among sites were assumed to be uniform. Pairwise deletion was applied for missing data treatment.
The visualization of genetic distance analysis was performed in R 4.5.2, utilizing the pheatmap and corrplot packages [72,73,74].
To assess species boundaries within key taxa, we performed Bayesian Poisson Tree Processes (bPTP) analysis following the method of Zhang et al. [75], using the web server (https://species.h-its.org/ptp/, accessed on 1 May 2026). The input phylogenetic tree was the maximum likelihood (ML) tree inferred from the concatenated ITS + nLSU dataset. Analyses were conducted with the following parameters: the tree was set as rooted; the Markov chain Monte Carlo (MCMC) was run for 250,000 generations, with a thinning interval of 100 and a burn-in of 0.15. The “remove outgroup” option was enabled to improve delimitation performance. MCMC convergence was assessed by examining the trace of log-likelihood values. The resulting species delimitation posterior probabilities (PPs) were visualized on the phylogenetic tree to identify putative species-level lineages. According to Malavasi et al. [76], clades with posterior probability (PP) ≥ 0.95 were considered highly well supported.

3. Results

3.1. Molecular Phylogeny

3.1.1. Podoscypha, Cymatoderma and Cladoderris

The dataset of Podoscypha, Cymatoderma, and Cladoderris, based on ITS + nLSU regions, comprises 57 samples, with Steccherinum ochraceum (Pers. ex J.F. Gmel.) Gray designated as the outgroup. The data matrix consists of 1373 constant characters, 318 parsimony-uninformative variable characters, and 509 parsimony-informative sites, resulting in an alignment length of 2200 bases. Both ML and BI analyses were applied to this identical dataset and aligned sequences, generating phylogenetic trees with similar topological structures, as depicted in Figure 1.
In this study (Figure 1), most species of Podoscypha form a well-supported clade (99% ML, 1.00 BPP) together with the type species of the genus, Podoscypha nitidula; this clade is recognized as Podoscypha sensu stricto. Podoscypha gillesii Boidin & Lanq., P. involuta (Klotzsch ex Fr.) Imazeki, P. subinvoluta Jing Si & Hai J. Li, and P. vespillonea (Berk.) Boidin & Lanq. form a distinct clade with full support (100% ML, 1.00 BPP). Due to the unavailability of these specimens, they are provisionally retained in Podoscypha. Three specimens of Podoscypha casiae J.X. Liu & H.S. Yuan (Yuan 19306, Yuan 19320 and Dai 7499) form a highly supported clade (97% ML, 1.00 BPP) and cluster with the clade containing P. tropica Jing Si & Hai J. Li and P. infundibula F.C. Huang & H.F. Zheng with moderate support (1.00 BPP).
In the phylogenetic tree (Figure 1), the type species of the genus, Cymatoderma elegans, and C. caperatum (Berk. & Mont.) D.A. Reid form a highly supported clade (96% ML, 1.00 BPP), which is regarded as Cymatoderma sensu stricto. Cymatoderma dendriticum (Pers.) D.A. Reid and three undescribed samples (Yuan 19310, Yuan 19314 and Yuan 19440) form a fully supported clade (100% ML, 1.00 BPP), prompting the resurrection of the genus Cladoderris to accommodate this lineage.
The bPTP species delimitation analysis showed good MCMC convergence, confirming reliable results. The bPTP analysis based on the ML tree delimited a total of 27 putative species (additional Supporting Information in Supplementary Material S1). Among these, 12 species received relatively high delimitation support (PP ≥ 0.95), including Podoscypha nitidula, P. mellissii, P. gillesii and Cymatoderma pallens, supporting their recognition as independent species. However, sequences from different specimens of the same species, such as P. fulvonitens, P. parvula, and P. involuta, were also split into distinct species units, indicating potential over-splitting by bPTP. By contrast, the PP values of the P. casiae species complex (including P. bolleana, P. brasiliensis, P. bubalina, P. disseminata, P. tropica, and P. infundibula), the C. dendriticum–Cladoderris perennis clade, and C. caperatum ranged from 0.40 to 0.60, indicating low delimitation confidence and ambiguous species boundaries.

3.1.2. Stereopsis

The ITS + nLSU dataset for Stereopsis includes 82 samples, with Protodontia piceicola (Kühner ex Bourdot) G.W. Martin designated as the outgroup. The data matrix comprises 903 constant characters, 124 parsimony-uninformative variable characters, and 1046 parsimony-informative sites, with an alignment length of 2073 bases. Both ML and BI analyses of this dataset generated phylogenetic trees with similar topologies, as shown in Figure 2.
In the phylogenetic tree (Figure 2), Stereopsis yunnanensis Qi Yuan & C.L. Zhao cluster with Stereopsis buccinata J.X. Liu & H.S. Yuan to form a clade with full support (100% ML, 1.00 BPP). Stereopsis globosa (Hjortstam & Ryvarden) Sjökvist, S. radicans, S. yunnanensis, and S. buccinata formed a fully supported clade (100% ML, 1.00 BPP).
The bPTP species delimitation analysis showed good MCMC convergence, confirming reliable results. Species delimitation analysis using bPTP based on the maximum likelihood (ML) phylogenetic tree recovered a total of 39 putative species (additional Supporting Information in Supplementary Material S2). Among these, 15 species received relatively high delimitation support (PP ≥ 0.95); these include Clavulicium delectabile, Sistotrema brinkmannii, S. globosa, S. radicans, and Kavinia himantia. In addition, S. yunnanensis and S. buccinata were delimited as a single species unit with a PP value of 0.90.

3.2. Genetic Distance Analysis

Genetic distances among 57 concatenated ITS and nLSU sequences were calculated under the Kimura 2-parameter model in MEGA 11 and visualized as a heatmap (Figure 3; full matrix in Supplementary Material S3). Distances range from 0 (identical) to 0.43 (highly divergent), with the color gradient transitioning from white (distance < 0.04) to sky blue (>0.30).
As shown in the genetic distance heatmap (Figure 3), interspecific genetic distances within Podoscypha s. s. range from 0.02 to 0.22. In Cymatoderma s. s., interspecific distances are relatively high (0.145–0.313), while intraspecific distances are low (0.024–0.03). Distances between Cymatoderma s. s. and C. pallens range from 0.277 to 0.313. By contrast, interspecific distances within Cladoderris are small (0.015–0.052), with intraspecific distances ranging from 0 to 0.01. Intergeneric distances are as follows: between Podoscypha s. l. and Cladoderris, 0.05–0.353; between Podoscypha s. s. and Cladoderris, 0.063–0.257; between Podoscypha s. l. and Cymatoderma, 0.048–0.38; between Podoscypha s. s. and Cymatoderma s. s., 0.048–0.317; and between Cymatoderma s. s. and Cladoderris, 0.175–0.282. Hence, the minimum intraspecific/interspecific genetic distances for Podoscypha, Cymatoderma, and Cladoderris were 0/0.006, 0.024/0.145, and 0/0.015, respectively. The minimum intergeneric distances between Podoscypha s.s. and the latter two genera were 0.048 and 0.063, respectively, while that between Cymatoderma and Cladoderris was 0.061. In addition, the minimum intergeneric distance between clades 2 and 3 of Podoscypha and Cladoderris was 0.05.

3.3. Taxonomy

3.3.1. Delimitation of the Genera

Based on the results of phylogenetic analysis (Figure 1 and Figure 2), the generic characteristics of Podoscypha s. s. and Cymatoderm s. s. were redefined.
Podoscypha Pat., Essai Tax. Hyménomyc. (Lons-le-Saunier): 70 (1900).
Type species. Podoscypha nitidula.
Description. Basidiome flabellate to infundibuliform, with stipe length exhibiting considerable variation; pileal surface smooth or tomentose, featuring radiating wrinkles or ridges; hymenial surface smooth or with radiating stripes; hyphal structure dimitic, comprising generative hyphae with clamp connections and thick-walled skeletal hyphae; cystidia present; basidia clavate, with a basal clamp; basidiospores ellipsoid to cylindrical, smooth, thin-walled, hyaline, and typically exceeding 3 µm in length.
Distribution. Most of the world except for Northern Asia, Central Asia, and the polar regions.
Substrate. Ground or fallen wood.
Notes. Podoscypha, first described and established in 1900, is a genus within the family Podoscyphaceae. Globally, 57 species of this genus have been recognized to date, predominantly distributed across tropical and temperate regions. In China, 17 species have been recorded, primarily inhabiting tropical to temperate areas.
Cymatoderma Jungh., Tijdschr. Nat. Gesch. Physiol. 7: 290 (1840).
Type species. Cymatoderma elegans.
Description. Basidiome flabellate, pseudo-infundibuliform to infundibuliform, stipitate; pileal surface covered with radiating ridges; hymenial surface with radiating folds or ridges; stipe tomentose; hyphal structure dimitic, hymenium thickened; cuticular layer present, giving rise to the pileal surface tomentum, the hairs that form the tomentum thick-walled except for apices, which may be thin-walled; cystidia present; basidia clavate, with a basal clamp; basidiospores cylindrical to ellipsoid, usually longer than 5 µm.
Distribution. North and South America, Africa, Southeast Asia, and Oceania.
Substrate. Fallen wood.
Notes. The genus Cymatoderma was first described in 1840 and currently belongs to Panaceae. To date, 15 species of this genus are recognized all over the world, predominantly in tropical regions. In China, there are two species, primarily distributed in tropical and subtropical regions.

3.3.2. Resurrected Genus

Cladoderris (Pers.) Berk., 1842, gen. resurr.
Etymology. Cladoderris (Lat.): referring to the dendroid, leathery basidiome.
Type species. Cladoderris dendritica (Pers.) Berk., automatically reinstated.
Description. Basidiome flabellate to infundibuliform, imbricate, stipitate; pileal surface covered by a highly developed felt-like tomentum with indistinct blade ridges beneath it; hymenial surface exhibits prominent ridges; stipe tomentose; hyphal structure dimitic or trimitic; cuticular layer absent; cystidia present; gloeocystidia thin-walled, forming long and undulating bodies; basidia clavate to subcylindrical; basidiospores ellipsoid to subglobose.
Distribution. South America and Africa.
Substrate. Ground or fallen wood.
Notes. Phylogenetic analysis places Cladoderris perennis and Cymatoderma dendriticum within the same clade, distinct from the clade of Cymatoderma s. s. Morphologically, Cymatoderma s. s. and Cladoderris share common traits in having a flabellate to infundibuliform basidiome, a pileus surface with ridges, tomentose stipes, and the presence of cystidia, clavate basidia, and subcylindrical-to-ellipsoidal basidiospores. However, the main difference between Cladoderris and Cymatoderma lies in that Cladoderris species have an imbricate basidiome, lack the cuticular layer, and have basidiospores shorter than 4 μm [2].
Cladoderris dendritica (Pers.) Berk.
Basionym.
Thelephora dendritica Pers., Botanique [5]: 176 (1827).
Cymatoderma dendriticum (Pers.) D.A. Reid, Kew Bull. [13] (3): 523 (1959).
Notes. The holotype of Cladoderris dendritica was collected from Peninsular Malaysia. It is characterized by a lateral stipe, a trimitic hyphal structure, absence of a cuticular layer, absence of cystidia, and broadly ellipsoid-to-subglobose basidiospores (3–4 × 2.5–3 μm) [2].

3.3.3. New Taxa

Cladoderris perennis J.X. Liu & H.S. Yuan, sp. nov. (Figure 4 and Figure 5).
Fungal Names number. FN 572903.
GenBank accession numbers. PQ415521 (ITS) and PQ415509 (nLSU).
Etymology. Perennis (Lat.): referring to perennial habits.
Diagnosis. Similar to Cladoderris dendritica but differs in having a dimitic hyphal system, thick-walled skeletal hyphae, and the presence of cystidia.
Description. Basidiome annual to perennial, stipitate, without taste, umami smell, becoming corky upon drying, 40–90 mm high. Pileus flabellate to reniform, imbricate, projecting up to 7–18 mm, 8–28 mm wide, 0.4–0.9 mm thick at the center, with a thin margin, extending towards the stipe and becoming fleshy; pileal surface golden blonde (5C4), topaz (5C5) and oak brown (5D6) when dry, smooth, with unobvious longitudinal ridges and a developed penniform tomentum; margin of pileal surface dark brown (7F6-7F8), indented. Hymenial surface orange white (5A2) to orange gray (5B2 to 5B4), tomentose, with prominent ridges under the tomentum. Absence of cuticular layer. Stipe lateral or central, soft corky, inflated, great variation in length, up to 70 mm long, sand (4B3) and pale orange (5A3), with yellowish white tomentum. Hyphal system dimitic; generative hyphae with clamps, thin- to slightly thick-walled; skeletal hyphae thick-walled, amyloid in IKI, CB+; tissues unchanged in KOH. Generative hyphae in context with clamps, colorless, thick-walled, frequently branched, subparallel to slightly interwoven, 2.1–5 μm in diam; skeletal hyphae in trama colorless, thick-walled to almost solid, unbranched, 2.8–5.3 μm in diam. Cystidia (leptocystidia) pyriform, clavate to subcylindrical, without apex, frequently present, 18–58 × 2–11 μm, sometimes encrusted; pilo- and caulocystidia absent. Gloeocystidia abundant, thin-walled, forming long and undulating bodies, 32–48 × 3.5–6 μm. Basidia narrowly clavate to subcylindrical, with four sterigmata and a basal clamp, 30–38 × 4.2–6.7 μm; basidioles dominant, in shape similar to basidia, clamps, slightly smaller, thin-walled. Basidiospores subglobose to ellipsoid, colorless, smooth, often monoguttulate, thin-walled, IKI–, CB–, 3.3 ± 0.2 × 3.0 ± 0.1 μm (2.9–4.2 × 2.7–3.2), L = 3.3 μm, W = 3.0 μm, Q = 1.1–1.3, Qm = 1.1, n = 90.
Distribution. Currently known only from China.
Substrate. On the ground.
Holotype. China, Guangdong Province, Shenzhen City, on the ground, 24 Apr. 2024, collected by Yong-Mei Cheng (holotype designated IFP 019974, specimen Yuan 19310).
Additional specimens examined (paratypes). China, Guangdong Province, Shenzhen City, on the ground, 26 April 2024, collected by Li-Fang Peng (paratype designated IFP 019975, specimen Yuan 19314); on the ground, 21 June 2024, collected by Yong-Mei Cheng (paratype designated IFP 019976, specimen Yuan 19440).
Notes. The new species is characterized by the absence of a cuticular layer, an inflated stipe and the presence of cystidia. Cladoderris perennis and C. dendritica form a sister clade, which share robust stipes and tomentose pilei with prominent ridges. The differences between C. dendritica and C. perennis are that C. perennis has a dimitic hyphal system, thick-walled skeletal hyphae, and the presence of cystidia, whereas C. dendritica possesses a trimitic hyphal system with thin-walled skeletal hyphae and lacks cystidia [2]. Furthermore, the pairwise ITS sequence similarity value between the new species (Specimen Yuan 19310) and C. dendritica (Specimen CBS 207.62) was 95.3%.
Podoscypha casiae J.X. Liu & H.S. Yuan, sp. nov. (Figure 6 and Figure 7).
Fungal Names number. FN 572901.
GenBank accession numbers. PQ415526 (ITS) and PQ415513 (nLSU).
Etymology. Casiae (combination of CAS and IAE), commemorating the 70th anniversary of the Institute of Applied Ecology (IAE), Chinese Academy of Sciences (CAS).
Diagnosis. Similar to Podoscypha tropica but differs in tomentose hymenial surface and CB+ skeletal hyphae.
Description. Basidiome annual, stipitate, gregarious, without odor or taste, coriaceous when fresh, becoming hard upon drying, 35–70 mm high. Pileus pseudoinfundibuliform to infundibuliform, projecting up to 13–18 mm long, 10–13 mm wide, 0.2–0.3 mm thick at the center; pileal surface of fresh specimens orange white (5A2) and pale red (7A3), light brown (6D8) when dry, glabrous, with distinct longitudinal stripe and rust brown concentric zones; margin of pileal surface Pompeian yellow (5C6) and clay (5D5) when dry, entire or slightly dentiform. Hymenial surface concolor or paler with pileal surface, very minutely tomentose. Stipe central, slender, cylindrical, up to 50 mm long and 3 mm in diam, golden brown to light brown (5D7–5D8) when dry, with grayish-white tomentum. Hyphal system dimitic; generative hyphae with clamps, thin- to thick-walled; skeletal hyphae thick-walled; IKI–, CB+, tissues unchanged in KOH. Generative hyphae in context dominant, with clamps and simple septs, colorless, thick-walled, moderately branched, subparallel to loosely interwoven, 2.3–3.9 μm in diam; skeletal hyphae in context colorless, thick-walled, unbranched, 3.1–5.4 μm in diam. Cystidia cylindrical, which has no apex, rarely present, thin-walled, 19–34 × 5.2–5.7 μm; pilocystidia absent; caulocystidia present, clavate to subcylindrical, thick-walled, 56–61 × 8.9–10 μm; basidia narrowly clavate to subcylindrical, with four sterigmata and a basal clamp, 9.7–28.5 × 4.0–4.6 μm; basidioles dominant, in shape similar to basidia, clamps, slightly smaller, thin-walled. Basidiospores ellipsoid to broadly ellipsoid, colorless, smooth, often monoguttulate, thin-walled, IKI–, CB–, 4.9 ± 0.67 × 3.3 ± 0.55 μm (4.0–7.0 × 2.4–4.3), L = 4.9 μm, W = 3.3 μm, Q = 1.3–2.0, Qm = 1.5, n = 90.
Distribution. Currently known only from China.
Substrate. On buried wood.
Holotype. China, Guangdong Province, Shenzhen City, on buried wood, 22 Apr. 2024, collected by Zhen-Xing Yao (holotype designated IFP 019977, specimen Yuan 19306).
Additional specimens examined (paratypes). China, Guangdong Province, Shenzhen City, on buried angiosperm wood, 24 May 2006, collected by Yu-Cheng Dai (paratype designated IFP 011924, specimen Dai 7499); on fallen angiosperm trunk, 26 April 2024, collected by Jian-Feng Tan (paratype designated IFP 019978, specimen Yuan 19320).
Notes. The new species is characterized by a glabrous pileal surface, a minutely tomentose hymenial surface and the presence of stipe and cystidia. Podoscypha casiae closely resembles P. tropica and P. infundibula in having caulocystidia and ellipsoid basidiospores. However, P. casiae differs from P. tropica in its tomentose hymenial surface and stipe, thin- to thick-walled generative hyphae, and CB+ skeletal hyphae [15]. P. casiae can be distinguished from P. infundibula in its smooth pileal surface and larger basidiospores (4.9 ± 0.67 × 3.3 ± 0.55 μm) [13]. Moreover, pairwise ITS sequence similarity analysis showed that the new species (Specimen Yuan 19306) had 97.8% similarity with P. tropica (Specimen 140719-19).
Stereopsis buccinata J.X. Liu & H.S. Yuan, sp. nov. (Figure 8 and Figure 9).
Fungal Names number. FN 572925.
GenBank accession numbers. PV190982 (ITS) and PV174382 (nLSU).
Etymology. Buccinata (Lat.): referring to the trumpet-shaped basidiome.
Diagnosis. Similar to Stereopsis yunnanensis but differs in basidiome without odor, tomentose pileal surface and hymenial surface and larger basidiospores.
Description. Basidiome annual, stipitate, solitary, without odor or taste, becoming hard upon drying, 12–41 mm high. Pileus flabelliform or infundibuliform, projecting up to 7–21 mm long, 9–22 mm wide, 0.2–0.4 mm thick at the center; pileal surface of dried specimens cream (4A3) or corn (4B5), tomentose, without radial stripes or zones; margin of pileal surface Somali (7E5) or eye brown (7F6), entire, wavy, with radial folds. Hymenial surface chocolate (6F4) or dark brown (8F4), with a thin and gray–white tomentum. Stipe lateral, bended, flattened, robust, up to 21 mm long and 5 mm in diam, with a white tomentum, concolor or paler with hymenial surface when dry. Hyphal system monomitic; generative hyphae slightly thick-walled, unbranched, IKI–, CB–; tissues unchanged in KOH. Generative hyphae in context colorless, subparallel or loosely interwoven, 1.5–2.2 μm in diam. Cystidia cylindrical, thin-walled, 35–47 × 3.5–5 μm; pilo- and caulocystidia absent. Basidia clavate, with two sterigmata and a basal clamp, 30–40.2 × 3.6–4 μm; basidioles dominant, similar to basidia in shape, generally slightly smaller. Basidiospores subglobose, colorless, smooth, thin-walled, IKI–, CB–, 5.9 ± 0.52 × 5.2 ± 0.45 μm (5.0–7.0 × 4.5–6.0), L = 5.9 μm, W = 5.2 μm, Q = 1.04–1.3, Qm = 1.1, n = 30.
Distribution. Currently known only from China.
Substrate. On the ground.
Holotype. China, Guangdong Province, Guangzhou City, Tianhe District, on the ground, 23 August 2023, collected by Hai-Sheng Yuan (holotype designated IFP 020068, specimen Yuan 18472).
Notes. The new species is characterized by a tomentose pileal surface, a hymenial surface, and basidia with two sterigmata. Stereopsis buccinata clusters with S. yunnanensis and S. radicans, displaying similarities in having a monomitic hyphal system with thick-walled generative hyphae, basidia with two sterigmata, and subglobose basidiospores. However, S. buccinata differs from S. yunnanensis in its basidiome without odor, tomentose pileal surface and hymenial surface, stipe with a white tomentum and larger basidiospores (5.9 ± 0.52 × 5.2 ± 0.45 μm in S. buccinata vs. 4.5–6.5 × 4–5.5 µm in S. yunnanensis) [25]. Stereopsis buccinata is distinguished from S. radicans by unbranched generative hyphae and smaller basidiospores (5.9 ± 0.52 × 5.2 ± 0.45 μm in S. buccinata vs. 6–8 × 5–7.5 µm in S. radicans) [2]. In addition, the pairwise ITS sequence similarity value between the new species and S. yunnanensis (Specimen CLZhao3767) was 94.7%.

4. Discussion

In this study, ITS and LSU regions were amplified from seven specimens of stipitate stereoid fungi collected from subtropical to temperate regions of China. We integrated phylogenetic analysis, genetic distance, bPTP, ITS sequence similarity analysis and morphology to delimit new taxa; consequently, three new species of stipitate stereoid basidiomycetes were proposed, and one genus was resurrected.
Consistent with the results of Sjökvist et al. [1], Wu et al. [10], Huang et al. [13], and Si et al. [15], the phylogenetic analyses in this study indicate that the genus Podoscypha is polyphyletic, forming three distinct lineages. Clade 1 contains the type species P. nitidula and P. casiae, as well as the majority of species belonging to Podoscypha s.s. Clades 2 and 3 include species such as P. involuta, P. subinvoluta, P. gillesii, P. vespillonea, and P. multizonata, all of which exhibit significant genetic differentiation in both tree topology and genetic distances. P. casiae and P. tropica exhibit distinct morphological differences, and P. casiae forms a highly supported monophyletic branch in the phylogenetic tree. The ITS sequence similarity between the two taxa is 97.8%, slightly above the 97% similarity cutoff. However, the bPTP species delimitation analysis provided only weak statistical support (PP 0.50 < 0.95), indicating ambiguous boundaries likely due to limited sampling or barcode resolution. Therefore, given that morphological divergence and phylogenetic independence are unequivocally supported, whereas the uncertainty in molecular evidence stems from methodological limitations, we recognize P. casiae as a new species based on the integration of phylogenetic and morphological evidence. Due to the limited number of specimens, five species within clades 2 and 3 are temporarily retained in the genus Podoscypha pending additional data.
In agreement with the findings of Yuan et al. [25], Stereopsis radicans, S. yunnanensis, and S. globosa cluster into a highly supported monophyletic branch. S. buccinata is also located within this clade and forms a well-supported sister branch with S. yunnanensis. The strong ITS divergence (5.3%, exceeding the 3% threshold), together with phylogenetic and morphological evidence, leads us to recognize S. buccinata as a new species. Although the bPTP analysis grouped S. buccinata and S. yunnanensis into a single putative species with fairly high but not significant posterior probability (PP 0.90), this value falls below our adopted significance threshold (PP ≥ 0.95), and the result is likely influenced by limited sampling or methodological limitations. Due to insufficient sequence data, the generic boundaries and internal relationships of Stereopsis remain unresolved. Additional specimens and molecular data are needed in the future to clarify the generic classification of this genus.
For Cymatoderma s.l., the phylogenetic results are consistent with those of Sjökvist et al. [1] and Huang et al. [13], showing that the genus is polyphyletic. The phylogenetic analysis supports a monophyletic clade corresponding to Cymatoderma s.s., which includes its type species C. elegans; meanwhile, Cladoderris dendritica and C. perennis form a highly supported clade. C. pallens clusters with P. multizonata and Abortiporus biennis (Bull.) Singer in a clade with low support. The ITS sequence similarity between C. perennis and C. dendritica is 95.3%, reflecting a divergence of 4.7%, which surpasses the 3% cutoff typically applied for species differentiation among basidiomycetes. C. perennis also constitutes a separate, strongly supported monophyletic clade in the phylogenetic analysis, and clear morphological distinctions provide additional evidence for its separation. Furthermore, the bPTP analysis yielded only low support (PP 0.60 < 0.95) for grouping C. perennis and C. dendritica as a single species; this outcome points to uncertain species boundaries, potentially attributable to methodological constraints or insufficient sampling. Therefore, we prioritize the robust phylogenetic independence, substantial ITS divergence, and clear morphological distinctions and designate C. perennis as a novel species of Cladoderris. Some species with no available molecular data (C. africanum, C. plicatum, C. sclerotioides, and C. blumei) are provisionally placed within Cymatoderma s.s. based on shared morphological features (a distinct cortex, thick-walled pileal hairs, and basidiospores ≥ 5 μm [2]).
Notably, the minimum interspecific genetic distance within Cymatoderma (0.145) was greater than the minimum intergeneric distances between Cymatoderma and both Podoscypha (0.048) and Cladoderris (0.061), which is consistent with previous studies indicating that Cymatoderma is not monophyletic [1,13]. However, due to the extremely limited number of Cymatoderma specimens and available sequences, there is insufficient evidence to reassess the taxonomic status of this genus at present. Therefore, we have followed the traditional classification scheme for Cymatoderma in this study and focus our discussion on the distinct generic status of Cladoderris. Future studies incorporating more species and population samples of Cymatoderma will be necessary to further test its generic boundaries.
Previous studies have placed most species of Cladoderris within Cymatoderma [21,77,78], but our results show that Cladoderris and Cymatoderma form independent, well-supported clades. The maximum interspecific genetic distance within Cladoderris is 0.052. With the exception of the minimum intergeneric distance between Cladoderris and Podoscypha in clades 2 and 3 (0.05), which is slightly lower than this value, the minimum intergeneric distances between Cladoderris and both Podoscypha s.s. (0.063) and Cymatoderma (0.061) are greater than 0.052. This slight overlap may be due to incomplete lineage sorting, limited sampling of clades 2–3, or the conserved nature of the ITS + LSU regions; nonetheless, the preponderance of evidence (monophyly, morphology, and other intergeneric distances) supports generic separation.
Stipitate stereoid fungi are widely distributed across tropical, subtropical, and temperate zones. The three new species described in this study enrich the diversity of this fungal group in China. Nevertheless, the taxonomy and distribution of related taxa in China still require further extensive sampling and investigation.
  • Keys to the accepted species of Podoscypha, Cymatoderma, Stereopsis and Cladoderris worldwide.
A key to accepted species of Podoscypha
1. Basidiome sessile or without a distinct stipe···································································2
1. Basidiome with a distinct stipe·························································································7
2. Basidiome effused and reflexed························································································3
2. Stipe short and rudimentary·····························································································4
3. Pileocystidia present but scattered·················································Podoscypha caespitosa
3. Pileocystidia absent·················································································P. semiresupinata
4. Pilei surface with radial wrinkles·····················································································5
4. Pilei surface with zones·····································································································6
5. Basidia with 4 sterigmate, 43.5 × 5.5 μm···························································P. aculeata
5. Basidia with 2 or 4 sterigmate, 18–26 × 4–6 μm·········································P. multizonata
6. Hymenial surface ochraceous buff in herbarium material·······························P. corneri
6. Hymenial surface brownish in herbarium material··········································P. gillesii
7. Basidiome infundibuliform or pseudoinfundibuliform················································8
7. Basidiome flabellate, spathulate, or (pseudo)infundibuliform···································19
8. Pileus surface tomentose···································································································9
8. Pileus surface smooth······································································································10
9. Stipe dark brown in herbarium material, with a hyphal disc··················P. infundibula
9. Stipe yellowish-to-pale brown in herbarium material, smooth························P. ursina
10. Hymenial surface tomentose···············································································P. casiae
10. Hymenial surface smooth·····························································································11
11. Hymenial surface with an ashy-gray pruina······························································12
11. Hymenial surface without a pruina·············································································14
12. Pilei surface with minutely radial wrinkles, without zones···················P. brasiliensis
12. Pilei surface with darker concentric zones··································································13
13. The width of basidiospores up to 2.2 μm··················································P. fulvonitens
13. The width of basidiospores range from 2.2 μm to 3.2 μm···························P. ravenelii
14. Stipe lacks a hyphal disc or a conspicuous earth ball at the base·················P. cristata
14. Stipe with a hyphal disc or a conspicuous earth ball at the base······························15
15. Pileocystidia absent, caulocystidia present·································································16
15. Pileo- and caulocystidia absent····················································································17
16. Basidiospores 3.75–4.75 × 2.5–3.2 μm·····························································P. bubalina
16. Basidiospores 2.5 × 2 μm··················································································P. mellissii
17. The length of basidiospores exceed 6 μm························································P. thozetii
17. The length of basidiospores range from 3 μm to 6 μm···············································18
18. Basidiospores up to 2.5 μm broad········································································P. curta
18. Basidiospores range from 3 μm to 4 μm broad··············································P. nitidula
19. Pilei surface smooth·······································································································20
19. Pilei surface tomentose··································································································31
20. Pilei surface dark, blackish brown or black in herbarium material··························21
20. Pilei surface warm brown or chestnut in herbarium material··································22
21. Pileocystidia present·····················································································P. ovalispora
21. Pileocystidia absent···························································································P. moelleri
22. Hymenial surface without a pruina·············································································23
22. Hymenial surface with a gray pruina··········································································26
23. Stipe tomentose, with a hyphal disc·············································································24
23. Stipe without a hyphal disc···························································································25
24. Metuloid cystidia present··················································································P. mellisii
24. Metuloid cystidia absent···················································································P. elegans
25. Basidiospores 4.2–6.1 × 3.3–4.6 μm···································································P. tropica
25. Basidiospores 3.75–4.75 × 2.5–3.2 μm·························································P. glabrescens
26. Stipe without a hyphal disc···························································································27
26. Stipe with a hyphal disc·································································································28
27. Intermediate hyphae present········································································P. venustula
27. Intermediate hyphae absent··············································································P. pusilla
28. Intermediate hyphae present······································································P. tomentipes
28. Intermediate hyphae abesent························································································29
29. Cystidioles thick-walled, up to 52 µm long and 6 µm wide·························P. parvula
29. Cystidioles absent··········································································································30
30. Skeletal hyphae 3–8 μm wide···········································································P. moselei
30. Skeletal hyphae 3–5 μm wide··········································································P. bolleana
31. Pilei surface without zones···························································································32
31. Pilei surface with zones·································································································34
32. Stipe without a hyphal disc···························································P. xanthopusconcinna
32. Stipe with a hyphal disc·································································································33
33. Basidiospores acyanophilous, 3.3–4.3 × 2.6–3 µm·································P. guangxiensis
33. Basidiospores slightly cyanophilous, 4.6–5.6 × 3–3.8 µm································P. lactea
34. Pileus surface with a tobacco-brown pruina········································P. philippinensis
34. Pileus surface without a pruina····················································································35
35. Hymenial surface with a pruina···················································································36
35. Hymenial surface without a pruina·············································································39
36. Stipe with a basal disc····································································································37
36. Stipe without a basal disc······························································································38
37. Pilei surface light gray, stipe light yellow······················································P. involuta
37. Pilei surface brown, stipe maroon······························································P. vespillonea
38. Basidiospores 3.75–5 × 2.5–3.75 µm·······························································P. petalodes
38. Basidiospores 3–3.75 × 2.2–3 µm·····································································P. viridans
39. Hymenial surface with radiating ribs····························································P. replicata
39. Hymenial surface without radiating ribs····································································40
40. Basidiospores cyanophilous·········································································P. densidisca
40. Basidiospores acyanophilous·······················································································41
41. Caulocystidia thin- to thick-walled·························································P. yunnanensis
41. Caulocystidia thick-walled, encrusted with irregular crystals·············P. subinvoluta
 
A key to accepted species of Cymatoderma
1. Basidiome infundibuliform or pseudoinfundibuliform···············································2
1. Basidiome flabellate or pseudoinfundibuliform····························································5
2. Stipe with a hyphal disc······························································Cymatoderma caperatum
2. Stipe without a hyphal disc·······························································································3
3. Hymenial surface with radiating ribs···························································C. africanum
3. Hymenial surface with radial folds··················································································4
4. Basidiospores elliptical, 8–11 × 4–5.5 μm························································C. plicatum
4. Basidiospores elliptical to subcylindrical, 4–5.5 × 2–3 μm·······················C. sclerotioides
5. Pilei surface glabrous····························································································C. blumei
5. Pilei surface tomentose······································································································6
6. Basidiospores elliptical, 6.5–9 × 4–5 μm·····························································C. elegans
6. Basidiospores oval to subglobose, 5–5.8 × 3.8–4.5 μm······································C. pallens
 
A key to accepted species of Stereopsis
1. Pilei surface tomentose······································································································2
1. Pilei surface glabrous or with fibers, wrinkles, ridges···················································6
2. Hymenial surface smooth·································································································3
2. Hymenial surface tomentose or with radial ridges························································5
3. Basidiospores subglobose·································································Stereopsis humphreyi
3. Basidiospores ellipsoid or cylindrical··············································································4
4. Stipe black············································································································S. nigripes
4. Stipe tawny to pale brown···············································································S. straminea
5. Hymenial surface with tiny radial branched ridges······························S. mussooriensis
5. Hymenial surface tomentose, basidiospores 5.2–6.8 × 5–5.7 μm·············S. buccinata
6. Pilei surface glabrous·········································································································7
6. Pilei surface with zones or radial fibers···········································································8
7. Basidiospores ellipsoid···········································································S. pseudocupulata
7. Basidiospores subglobose·····················································································S. albida
8. Cystidia present··················································································································9
8. Cystidia absent·················································································································11
9. Cystidia fusiform·························································································S. yunnanensis
9. Cystidia cylindrical··········································································································10
10. Basidia with 2 sterigmata················································································S. radicans
10. Basidia with 4 sterigmata···················································································S. raphiae
11. Generative hyphae with clamp connections···················································S. hiscens
11. Generative hyphae without clamp connections·························································12
12. Pilei surface with radial wrinkles or ridges·································································13
12. Pilei surface with radial fibers or fibrillose··································································14
13. Stipe surface glabrous················································································S. sparassoides
13. Stipe surface tomentose··············································································S. cartilaginea
14. Hymenial surface smooth·····························································································15
14. Hymenial surface slightly rugulose·············································································16
15. Basidiome infundibuliform, stipe surface tomentose··································S. burtiana
15. Basidiome spathulate or flabelliform··························································S. malaiensis
16. Basidiospores 4.5–6 × 3–4 µm···············································································S. reidii
16. Basidiospores 3–4 × 2.2–2.5 µm·······································································S. vitellina
 
A key to accepted species of Cladoderris
1. Binding hyphae absent·······································································Cladoderris perennis
1. Binding hyphae branched, 2–2.5 μm·····························································C. dendritica

Supplementary Materials

The following supporting information can be downloaded at: https://www.mdpi.com/article/10.3390/jof12060400/s1, Supplementary Material S1: The bPTP species delimitation analysis of the genera Podoscypha, Cymatoderma, and Cladoderris; Supplementary Material S2: The bPTP species delimitation analysis of the genus Stereopsis; Supplementary Material S3: Genetic distance of the genera Podoscypha, Cymatoderma, and Cladoderris based on ITS + nLSU.

Author Contributions

J.-X.L. performed morphological and molecular analysis, made species delimitation, and wrote the manuscript draft. L.-J.Z. provided specimens and some DNA sequences. Y.-Q.Z. provided the software utilization and DNA sequences. H.B.L. revised the manuscript. H.-S.Y. led this study, collected specimens, and revised the manuscript. All authors have read and agreed to the published version of the manuscript.

Funding

This research was financed by the National Natural Science Foundation of China (nos. 32570013 and U2102220) and the Fund of CAS Key Laboratory of Forest Ecology and Silviculture, Institute of Applied Ecology, Chinese Academy of Sciences (no. KLFES-2028).

Institutional Review Board Statement

Not applicable.

Informed Consent Statement

Not applicable.

Data Availability Statement

The sequences from the present study were submitted to the NCBI website (https://www.ncbi.nlm.nih.gov/, accessed on 29 June 2025), and the accession numbers are listed in Table 2.

Acknowledgments

We are grateful to Zhen-Xing Yao, Zhong-Ping Feng, Yu-Cheng Dai, Jian-Feng Tan, Wu-Ping Luo, Yong-Mei Cheng and Li-Fang Peng for collecting specimens.

Conflicts of Interest

The authors have declared that no competing interests exist.

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Figure 1. ML phylogenetic tree of the genera Podoscypha, Cymatoderma, and Cladoderris based on ITS + nLSU. Branches are labeled with maximum likelihood bootstrap values (≥80%) and Bayesian posterior probabilities (≥0.95). Red dotted lines indicate species units supported by bPTP (PP ≥ 0.95). The two new species are shown in bold and marked with red stars. All new taxa and the resurrected genus are highlighted within the blue area.
Figure 1. ML phylogenetic tree of the genera Podoscypha, Cymatoderma, and Cladoderris based on ITS + nLSU. Branches are labeled with maximum likelihood bootstrap values (≥80%) and Bayesian posterior probabilities (≥0.95). Red dotted lines indicate species units supported by bPTP (PP ≥ 0.95). The two new species are shown in bold and marked with red stars. All new taxa and the resurrected genus are highlighted within the blue area.
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Figure 2. ML phylogenetic tree of the genus Stereopsis based on ITS + nLSU. Branches are labeled with maximum likelihood bootstrap values (≥80%) and Bayesian posterior probabilities (≥0.95). Red dotted lines indicate species units supported by bPTP (PP ≥ 0.95). The one new species is in bold and marked with a red star. The new taxa is highlighted within the blue area.
Figure 2. ML phylogenetic tree of the genus Stereopsis based on ITS + nLSU. Branches are labeled with maximum likelihood bootstrap values (≥80%) and Bayesian posterior probabilities (≥0.95). Red dotted lines indicate species units supported by bPTP (PP ≥ 0.95). The one new species is in bold and marked with a red star. The new taxa is highlighted within the blue area.
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Figure 3. Genetic distance heatmap for Podoscypha, Cymatoderma, and Cladoderris based on ITS + nLSU sequences (Kimura 2-parameter model). Colors indicate pairwise genetic distances, ranging from low (blue) to high (yellow), with larger values representing greater sequence divergence and more distant phylogenetic relationships.
Figure 3. Genetic distance heatmap for Podoscypha, Cymatoderma, and Cladoderris based on ITS + nLSU sequences (Kimura 2-parameter model). Colors indicate pairwise genetic distances, ranging from low (blue) to high (yellow), with larger values representing greater sequence divergence and more distant phylogenetic relationships.
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Figure 4. Basidiome of Cladoderris perennis (holotype Yuan 19310, photo by Yong-Mei Cheng).
Figure 4. Basidiome of Cladoderris perennis (holotype Yuan 19310, photo by Yong-Mei Cheng).
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Figure 5. Microscopic structures of Cladoderris perennis (holotype Yuan 19310). (A) Basidiospores; (B) a section of hymenophore with gloeocystidia (granular) and leptocystidia (hyaline); (C) leptocystidia and cystidioles; (D) skeletal hyphae in trama. Scale bars: 5 μm (A); 10 μm (B–D).
Figure 5. Microscopic structures of Cladoderris perennis (holotype Yuan 19310). (A) Basidiospores; (B) a section of hymenophore with gloeocystidia (granular) and leptocystidia (hyaline); (C) leptocystidia and cystidioles; (D) skeletal hyphae in trama. Scale bars: 5 μm (A); 10 μm (B–D).
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Figure 6. Basidiome of Podoscypha casiae (holotype Yuan 19306, photo by Zhen-Xing Yao).
Figure 6. Basidiome of Podoscypha casiae (holotype Yuan 19306, photo by Zhen-Xing Yao).
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Figure 7. Microscopic structures of Podoscypha casiae (holotype Yuan 19306). (A) Basidiospores; (B) a section of hymenophore with basidia and cystidia; (C) caulocystidia. Scale bars: 5 μm (A); 10 μm (B,C).
Figure 7. Microscopic structures of Podoscypha casiae (holotype Yuan 19306). (A) Basidiospores; (B) a section of hymenophore with basidia and cystidia; (C) caulocystidia. Scale bars: 5 μm (A); 10 μm (B,C).
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Figure 8. Basidiome of Stereopsis buccinata (holotype Yuan 18472; photo by Jia-Xue Liu).
Figure 8. Basidiome of Stereopsis buccinata (holotype Yuan 18472; photo by Jia-Xue Liu).
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Figure 9. Microscopic structures of Stereopsis buccinata (holotype Yuan 18472). (A) Basidiospores; (B) a section of hymenophoral trama with cystidia and basidia. Scale bars: 5 μm (A); 10 μm (B).
Figure 9. Microscopic structures of Stereopsis buccinata (holotype Yuan 18472). (A) Basidiospores; (B) a section of hymenophoral trama with cystidia and basidia. Scale bars: 5 μm (A); 10 μm (B).
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Table 1. PCR amplification system.
Table 1. PCR amplification system.
ReagentVolume (μL)
Double distilled water10
2× Taq PCR Mix12.5
Forward primer1
Reverse primer1
DNA1
Table 2. Voucher numbers, geographic origins and GenBank accession numbers used in this study.
Table 2. Voucher numbers, geographic origins and GenBank accession numbers used in this study.
SpeciesSpecimenLocalityITSnLSUReferences
Abortiporus biennisEL65-03 (GB)SwedenJN649325JN649325[1]
A. biennisWei 11890China, JiangxiPQ415519PQ415507Unpublished
Cladoderris perennisYuan 19310 (T)China, ShenzhenPQ415521PQ415509Present study
C. perennisYuan 19314China, GuangzhouPQ415522PQ415510Present study
C. perennisYuan 19440China, ShenzhenPQ415523PQ415511Present study
Clavariadelphus amplusHKAS 76577ChinaMK705851MK704443[43]
C. amplusHKAS 54876 (T)ChinaMK705857MK704444[43]
C. elongatesHMAS 260746ChinaMK705845MK704441[43]
C. elongatesMHHNU 9250ChinaPV463551PV490604[44]
C. ligulaKHL 8560 (GB)SwedenJN649329JN649329[1]
C. ligulaMHHNU 10452ChinaPV463555PV490608[44]
C. pistillarisNAMA 2017-123USAMH979250-Unpublished
C. pistillarisFLAS-F-60521USAMH281842-Unpublished
C. pseudoelongatusMHHNU 32323 (T)ChinaPV463559PV490617[44]
C. pseudoelongatusMHHNU 12123ChinaPV463560PV490618[44]
C. yunnanensisMHHNU 9244ChinaPV463561PV490606[44]
C. yunnanensisHKAS 54849ChinaMK705869MK704453[43]
Clavulicium delectabile1391_PREMIXUSAOQ612508-Unpublished
C. delectabileK(M)236345United KingdomMZ159671-Unpublished
C. globosumKHL12592Costa RicaKC203495KC203495[24]
C. macouniiNorde’n 37145 (GB)SwedenJN649332JN649332[1]
C. macouniiKHL 12129 (GB)SwedenJN649333JN649333[1]
Clavulina castaneipesOSC 108705USAEU669209EU669261Unpublished
C. castaneipesOSC 116725USAEU669210EU669262Unpublished
C. cristataEL7-00 (GB)Finland-AM259213[45]
C. cristataRAS323 SV2USAOR464380OR460872[46]
C. kunmudlutsaTH9206GuyanaHQ680358HQ680358[47]
C. kunmudlutsaTH8932 (T)GuyanaHQ680359HQ680359[47]
C. mahiscolorataFCME 27660MexicoMH542551MN049493[48]
C. mahiscolorataFCME 27662 (T)MexicoMH542554MN049496[48]
C. parvisporaFCME 27657MexicoMH542549MN049491[48]
C. parvisporaFCME 27650 (T)MexicoNR_166245MN049492[48]
C. purpurascensMHHNU 9846ChinaMK564136MK564126[49]
C. purpurascensMHHNU 9848 (T)ChinaMK564137MK564127[49]
Cymatoderma caperatumFLAS-F-61204USAMH211809-Unpublished
C. caperatumMES-3721USAON383384-Unpublished
C. dendriticumCBS 615.73Sri LankaJN649338JN649338[1]
C. dendriticumCBS 207.62CameroonJN649339JN649339[1]
C. elegansDai 17511ChinaON417155-[50]
C. elegansYuan 19343China, HainanPQ415520PQ415508Unpublished
C. pallensCBS 327.66 (T)CameroonJN649342JN649342[1]
Hydnum albomarginatumFJAU66574 (T)ChinaPV329855PV356813[51]
H. albomarginatumFJAU66575ChinaPV329856PV356814[51]
H. fulvostriatumFJAU66566 (T)ChinaPV329849PV356807[51]
H. fulvostriatumFJAU66567ChinaPV329850PV356808[51]
H. jussiiYuan 14008ChinaMW980553MW979539 [52]
H. jussiiYuan 14009ChinaMW980554MW979540[52]
H. pallidocroceumYuan 14023 (T)ChinaMW980568MW979554[52]
H. pallidocroceum Yuan 14017ChinaMW980569MW979555[52]
H. repandumKHL 8552 (GB)SwedenJN649348JN649348[1]
H. repandum H 6003710 (T)Finland NR164553-[53]
H. sphaericumWei 10243 (T)ChinaMW980563MW979549[52]
H. sphaericum Wei 10262ChinaMW980565MW979551[52]
Kavinia alboviridisKM82737United KingdomGQ981505-Unpublished
K. alboviridisEL16-98Estonia-AY463434[54]
K. alboviridisKM141510United KingdomGQ981506-Unpublished
K. altoandinaPSL2302 (T)ChileOP022196-[1]
K. altoandinaPSL3014ChileOP022197-[55]
K. chacoserranaRobledo 2516 (T)ArgentinaMF377531PQ453549Unpublished
K. himantiaLL 36041 (GB)Sweden-AY586682[54]
K. himantiaHMJAU59876ChinaPX411731-Unpublished
K. himantiaLE F-355928RussiaPV017449-Unpublished
Phaeoclavulina abietinaOSC 134649USAJX310378JX287478Unpublished
P. abietinaEL61-03 (GB)SwedenJN649369JN649369[1]
P. bicolorMHHNU10702 (T)ChinaPP809798PP800475[56]
P. bicolorMHHNU10703ChinaPP809799PP800476[56]
P. echinoflavaHKAS 45984 (T)ChinaPP809801PP800478[56]
P. echinoflavaHKAS 45992ChinaPP809800PP800477[56]
P. jilinensisMHHNU9149ChinaPP809802PP800479[56]
P. jilinensisMHHNU9164ChinaPP809803PP800480[56]
P. macrosporaMHHNU9143ChinaPP467359PP493649Unpublished
P. macrosporaMHHNU10399ChinaPP467360PP493650Unpublished
P. minutisporaAMB 18588ItalyMT055969MT053246Unpublished
P. minutisporaAMB 18586 (T)ItalyMT055965MT053243Unpublished
Podoscypha bolleanaCBS 333.66Central African RepublicJN649354JN649354[1]
P. bolleana32032-JQ675332-Unpublished
P. brasiliensisLR 37812 (O)VenezuelaJN649355JN649355[1]
P. bubalina17500-JQ675311-Unpublished
P. casiaeYuan 19306 (T)China, ShenzhenPQ415526PQ415513Present study
P. casiaeDai 7499China, ShenzhenPQ415527PQ415514Present study
P. casiaeYuan 19320China, ShenzhenPQ415528PQ415515Present study
P. cristata8667-JQ675320-Unpublished
P. densidiscaGXU 5784 (T)China, GuangxiOR347558OR347508[13]
P. densidiscaGXU 7524China, GuangxiOR347571OR347572[13]
P. disseminataDMC 232-JQ675326-Unpublished
P. elegansCBS 426.51ArgentinaJN649356JN649356[1]
P. fulvonitens17483-JQ675315-Unpublished
P. fulvonitens18332-JQ675316-Unpublished
P. gillesiiGXU 2176ChinaMG356710MG356793[12]
P. guangxiensisGXU 5849 (T)China, GuangxiOR347573OR347564[13]
P. guangxiensisGXU 6031China, GuangxiOR347565OR347566[13]
P. infundibulaGXU 7837 (T)China, GuangxiOR340036OR340137[13]
P. infundibulaGXU 8829China, GuangxiOR342113OR342114[13]
P. involutaE. Larsson (GB)ThailandJN649357JN649357[1]
P. involutaCBS 113.74Central African RepublicJN649358JN649358[1]
P. lacteaGXU 5952 (T)China, GuangxiOR347570OR347567[13]
P. lacteaGXU 7612China, GuangxiOR347569OR347569[13]
P. mellissiiLR 41658 (O)JamaicaJN649359JN649359[1]
P. multizonataJahn 751012 (GB)GermanyJN649360JN649360[1]
P. multizonata3005GermanyJN710581JN710581[57]
P. nitidula8724-JQ675321-Unpublished
P. parvula32055-JQ675338-Unpublished
P. parvulaDMC 226-JQ675328-Unpublished
P. petalodesYuan 1634ChinaPQ415525-Unpublished
P. petalodes subsp. rosulataCBS 659.84PakistanJN649362JN649362[1]
P. petalodes subsp. rosulataCBS 332.66PakistanJN649363JN649363[1]
P. petalodes subsp. cystidiata140721-15 (T)China: YunnanOQ305833OQ305828[15]
P. petalodes subsp. cystidiata140721-14China: YunnanOQ305832OQ305827[15]
P. raveneliiCBS 664.84USAJN649364JN649364[1]
P. sp.LR43794 (O)Costa RicaJN649365JN649365[1]
P. tropica140719-19 (T)China: YunnanOQ305834OQ305830[15]
P. tropicaYuan 19438China, JiangxiPQ415524PQ415512Unpublished
P. venustulaLR40821 (O)VenezuelaJN649366JN649366[1]
P. venustulaCBS 656.84French GuianaJN649367JN649367[1]
P. vespilloneaCBS 111.74-JN649368JN649368[1]
P. vespilloneaCBS 111.74-MH860836-[58]
P. yunnanensisCLZhao 3963ChinaMK298400MK298404[10]
P. yunnanensisCLZhao 4035 (T)ChinaMK298403MK298407[10]
Protodontia piceicolaKHL 11763 (GB)SwedenDQ873660DQ873660[59]
Sistotrema brinkmanniiNH 11412 (GB)TurkeyAF506473AF506473[60]
S. brinkmanniiKHL14078 (GB)SwedenKF218967KF218967[61]
S. confluensMV-1754SwedenPQ653462PQ653462Unpublished
S. confluensRAS950USAPQ050701PQ050699[62]
S. flavorhizomorphaeTUMH 64409 (T)JapanNR_178118NG_153858[63]
S. flavorhizomorphaeTUMH 64408JapanLC642048LC642066[63]
S. luteovirideHK23176 (H) (T)FinlandKF218963KF218963[61]
S. sinenseCLZhao 24876 (T)ChinaPQ758748PQ758756[64]
S. subconfluensDai 12577 (T)China-JX076810[65]
S. subconfluensDai 12578China-JX076811[65]
S. yunnanenseCLZhao 7357 (T)ChinaON817194ON810362[66]
S. yunnanenseCLZhao 7396ChinaON817195ON810363[66]
Steccherinum ochraceumKHL 11902 (GB)SwedenJQ031130JQ031130[1]
Stereopsis buccinataYuan 18472 (T)China, GuangzhouPV190982PV174382Present study
S. globosaKHL 11228 (GB)Costa RicaJN649330JN649330[1]
S. globosaKHL 12592 (GB)Costa RicaJN649331JN649331[1]
S. radicansPR 5760 (T)Puerto RicoJN649370JN649370[1]
S. radicansTB 8943 (CORT)VenezuelaJN649372JN649372[1]
S. yunnanensisCLZhao 3767 (T)China, YunnanNR198372NG243203[25]
S. yunnanensisYuan 19365China, ShenzhenPV190983PV174383Unpublished
New species are in bold, and type specimens are indicated by (T). Specimen Deposition: (GB) = Herbarium, University of Gothenburg (Sweden); (O) = Oslo Herbarium, Natural History Museum, University of Oslo (Norway).
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Liu, J.-X.; Zhou, L.-J.; Zhu, Y.-Q.; Lee, H.B.; Yuan, H.-S. Taxonomy and Phylogeny of Stipitate Stereoid Basidiomycetes from China. J. Fungi 2026, 12, 400. https://doi.org/10.3390/jof12060400

AMA Style

Liu J-X, Zhou L-J, Zhu Y-Q, Lee HB, Yuan H-S. Taxonomy and Phylogeny of Stipitate Stereoid Basidiomycetes from China. Journal of Fungi. 2026; 12(6):400. https://doi.org/10.3390/jof12060400

Chicago/Turabian Style

Liu, Jia-Xue, Lin-Jiang Zhou, Ya-Quan Zhu, Hyang Burm Lee, and Hai-Sheng Yuan. 2026. "Taxonomy and Phylogeny of Stipitate Stereoid Basidiomycetes from China" Journal of Fungi 12, no. 6: 400. https://doi.org/10.3390/jof12060400

APA Style

Liu, J.-X., Zhou, L.-J., Zhu, Y.-Q., Lee, H. B., & Yuan, H.-S. (2026). Taxonomy and Phylogeny of Stipitate Stereoid Basidiomycetes from China. Journal of Fungi, 12(6), 400. https://doi.org/10.3390/jof12060400

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