Abstract
Wood-inhabiting fungi serve as primary decomposers of lignocellulose in forest ecosystems, facilitating carbon release and nutrient turnover. These processes are essential for sustaining material cycling, energy flow, and soil health. The genus Hyphoderma, within the Basidiomycota, is a notable group among wood-inhabiting fungi. This study introduces two new resupinate, adnate, wood-inhabiting fungal taxa, Hyphoderma tenuicystidiatum and H. tabuliformis, based on morphological and molecular phylogenetic analyses. Hyphoderma tenuicystidiatum is distinguished by light vinaceous gray fresh basidiomata with a smooth hymenophore, a monomitic hyphal system with clamp connections on generative hyphae, cylindrical cystidia, the presence of fusoid cystidioles, and subcylindrical to cylindrical basidiospores measuring 10.4–14.9 × 5.2–7.1 µm. In contrast, H. tabuliformis exhibits white to cream fresh basidiomata with an arachnoid hymenophore, encrusted generative hyphae, clavate cystidia, and subcylindrical basidiospores measuring 8.7–12.2 × 3.6–4.2 µm. The discovery of two new species in Qinghai enriches the diversity of Hyphoderma and provides additional morphological and molecular data for the genus.
Key words: Hyphodermataceae , phylogeny, taxonomy, wood-rotting fungi
Introduction
Wood-inhabiting fungi grow on various woody materials and constitute an important group of microorganisms (Yuan et al. 2020, 2026; Wu et al. 2022; Liu et al. 2025; Guan et al. 2026). Some of these fungi can decompose and utilize wood, thereby facilitating nutrient cycling in forest ecosystems, and are vital components of these ecosystems (Si et al. 2011; Wang et al. 2023). In addition, certain species, such as medicinal fungi and forest pathogens, have significant economic value (Wu et al. 2019; Yuan et al. 2021; Bhunjun et al. 2024). Although several wood-inhabiting fungi have been described from western China (Dai et al. 2004a; Wu et al. 2017; Yuan et al. 2021), the diversity of this group remains poorly known. Hyphoderma Wallr., a member of the family Hyphodermataceae Jülich (Polyporales, Basidiomycota), was established by Wallroth (1833) and typified by H. setigerum (Fr.) Donk. The genus is characterized by resupinate to effuse-reflexed, adnate, whitish to yellowish basidiomata of ceraceous consistency, a smooth to tuberculate or hydnoid hymenophore, a monomitic (rarely dimitic) hyphal system with clamp connections, the presence or absence of cystidia, suburniform to subcylindrical basidia, and ellipsoid to subglobose, smooth, thin-walled basidiospores (Bernicchia and Gorjón 2010).
The genus Hyphoderma was first introduced by Wallroth (1833) with eight species based on a broad concept that included both resupinate and thelephoroid (clavarioid) forms. After its establishment, the genus was largely neglected until the mid-20th century, when the Dutch mycologist M.A. Donk undertook extensive revisions (Donk 1949, 1957). Donk (1949) selected Hyphoderma spiculosum (syn. Thelephora setigera) as the type, arguing that it was the most prominent element among the taxa listed by Wallroth and also the one most in accordance with the genus description. Subsequently, Donk (1957) provided a more thorough revision of the genus, recognizing 25 species and formally adopting the combination H. setigerum (Fr.) Donk for the type species, thereby confirming H. spiculosum as a later synonym of H. setigerum. Following Donk’s revisions, several taxonomists attempted to subdivide the genus at the sectional level (Parmasto 1968; Eriksson and Ryvarden 1975).
Parmasto (1968) divided Hyphoderma into three sections, namely Hyphoderma, Mutatoderma, and Stephanocystis. Eriksson and Ryvarden (1975) considered the genus to be unclearly delimited and, anticipating a future subdivision, proposed 12 possible natural species groups as a basis for future taxonomic revision. In the 21st century, molecular phylogenetic studies have profoundly reshaped the traditional classification of Hyphoderma (Larsson 2007). As a result, species possessing distinctive structures such as echinocysts were segregated and transferred to a reinstated older genus, Peniophorella. This changed the concept of Hyphoderma to a narrower circumscription.
To date, 137 species of Hyphoderma have been discovered worldwide, among which 45 have been reported from China (Guan and Zhao 2021; Li et al. 2026). During investigations of the diversity of wood-rotting fungi in China, four Hyphoderma-like samples were collected. Phylogenetic analyses based on nuclear ribosomal ITS and nLSU sequences were carried out to confirm their taxonomic status. Morphological and molecular evidence confirmed that the four examined specimens belong to two new Hyphoderma species.
Materials and methods
Morphological studies
The examined specimens are deposited in the Fungarium of the Institute of Microbiology, Beijing Forestry University (BJFC), Beijing, China. Macromorphological characteristics, including the color and thickness of the basidiomata, hymenophoral surface, and sterile margin, were examined under a stereomicroscope (Nikon SMZ 1000, Tokyo, Japan) at 4× magnification. Colors used follow Anonymous (1969) and Petersen (1996). Samples for microscopic examination were stained in Cotton Blue, Melzer’s reagent, 5% potassium hydroxide, and distilled water, and sections or tissues of basidiomata were studied in Cotton Blue at 1000× magnification using a Nikon E80i microscope with phase-contrast illumination (Nikon, Tokyo, Japan). Line drawings were made with the aid of a drawing tube. To show variation in spore sizes, 30 basidiospores were measured from each specimen. Stalks were excluded from basidia measurements, and the hilar appendage was excluded from basidiospore measurements (Yuan and Zhao 2024). Basidiospore measurements are presented as follows: (c) d–e (f) × (g) h–i (j) µm, where c and f represent the extreme minimum and maximum values of basidiospore length, each accounting for 5% of the measurements; d and e represent the minimum and maximum values within the 90% range; basidiospore width values (g–j) are given in the same manner. The following abbreviations were used: L = mean spore length (arithmetical average of all spores), W = mean spore width (arithmetical average of all spores), Q = values of the length/width ratios among the studied specimens, n = a/b (number of spores (a) measured from a given number (b) of specimens), IKI = Melzer’s reagent, IKI− = negative in Melzer’s reagent, KOH = 5% potassium hydroxide, CB = Cotton Blue, CB− = acyanophilous.
DNA extraction, polymerase chain reaction amplification, and sequencing
A CTAB plant genome rapid extraction kit-DN14 (Aidlab Biotechnologies Co., Ltd.) was used for DNA extraction and PCR amplification from dried specimens according to the manufacturer’s instructions, with some modifications (Liu et al. 2023). The ITS region was amplified with the primers ITS5 (5'-GGAAGTAAAAGTCGTAACAAGG-3') and ITS4 (5'-TCCTCCGCTTATTGATATGC-3') (White et al. 1990). The nLSU gene was amplified with the primers LR0R (5'-ACCCGCTGAACTTAAGC-3') and LR7 (5'-TACTACCACCAAGATCT-3') (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, 58 °C for 45 s, and 72 °C for 1 min, and a final extension at 72 °C for 10 min. The PCR procedure for nLSU was as follows: initial denaturation at 94 °C for 1 min, followed by 35 cycles at 94 °C for 30 s, 48 °C for 1 min, and 72 °C for 1.5 min, and a final extension at 72 °C for 10 min. The PCR products were purified and sequenced bidirectionally by Sanger sequencing using the same primers at the Beijing Genomics Institute, Beijing, China. The newly generated sequences were deposited in GenBank. All sequences analyzed in this study are listed in Table 1.
Table 1.
Information on specimens and GenBank accessions used in this study. New sequences are given in bold. Type specimens are indicated with an asterisk, and “-” indicates unavailable data.
| Species | Specimen no. | Locality | GenBank accession no. | |
|---|---|---|---|---|
| ITS | nLSU | |||
| Climacocystis borealis | FD-31 | USA | KP135308 | KP135210 |
| Diplomitoporus crustulinus | FD-137 | USA | KP135299 | KP135211 |
| Hyphoderma alboarachnum | CLZhao 30488* | China | PV470563 | – |
| H. amoenum | USO 286622 | Canada | HE577030 | – |
| H. asianum | CLZhao 18091* | China | OR141726 | PP826262 |
| H. assimile | CBS:125852 | New Zealand | MH863808 | MH875272 |
| H. australosetigerum | MA Fungi 92235* | Chile | MN963764 | – |
| H. australosetigerum | MA Fungi 92240 | Chile | MN963760 | – |
| H. bambusinum | CLZhao 29903* | China | PV469674 | PV819428 |
| H. cinereofuscum | CLZhao 30341* | China | PQ492371 | PQ511128 |
| H. cinereofuscum | CLZhao 30283 | China | PQ492372 | PQ511129 |
| H. cremeoalbum | NH 11538 (GB) | Sweden | DQ677492 | DQ677492 |
| H. crystallinum | CLZhao 9338* | China | MW917161 | MW913414 |
| H. crystallinum | CLZhao 9374 | China | MW917162 | MW913415 |
| H. definitum | NH 12266 (GB) | China | DQ677493 | DQ677493 |
| H. fissuratum | CLZhao 6731 | China | MT791331 | – |
| H. fissuratum | CLZhao 6726* | China | MT791330 | MT791334 |
| H. floccosum | CLZhao 17129* | China | MW301683 | MW293733 |
| H. floccosum | CLZhao 17215 | China | MW301687 | MW293735 |
| H. fulgens | CLZhao 37429* | China | PV829544 | PV810095 |
| H. fulgens | CLZhao 39474 | China | PV829547 | PV810098 |
| H. granuliferum | 5273 | Canada | JN710545 | JN710545 |
| H. grandineum | CLZhao 30046* | China | PV470561 | PV819429 |
| H. grandineum | CLZhao 43328 | China | PV470562 | PV819430 |
| H. guangdongense | CLZhao 12657* | China | PP235513 | PP235514 |
| H. laceratum | CLZhao 34242* | China | PV829552 | PV810101 |
| H. laceratum | CLZhao 34672 | China | PV829553 | PV810103 |
| H. litschaueri | NH 7603 (GB) | Sweden | DQ677496 | DQ677496 |
| H. litschaueri | FP-101740-Sp | USA | KP135295 | KP135219 |
| H. macaronesicum | MA Fungi 90388 | USA | KC984327 | – |
| H. macaronesicum | TFC Mic 15115 | China | HE577011 | – |
| H. marginatum | CLZhao 3404 | China | OM985717 | OM985754 |
| H. medioburiense | NH 10950 (GB) | Spain | DQ677497 | DQ677497 |
| H. membranaceum | CLZhao 5844* | China | MW917167 | MW913420 |
| H. membranaceum | CLZhao 6971 | China | MW917168 | MW913421 |
| H. microporoides | CLZhao 6857* | China | MW917169 | MW913422 |
| H. microporoides | CLZhao 8695 | China | MW917170 | MW913423 |
| H. moniliforme | Wu 0211-42* | China | KC928282 | – |
| H. moniliforme | Wu 0211-46 | China | KC928284 | – |
| H. mopanshanense | CLZhao 6498* | China | MT791329 | MT791333 |
| H. mopanshanense | CLZhao 6449 | China | OM985720 | OM985759 |
| H. nemorale | TNM F3931 | China | KJ885183 | KJ885184 |
| H. nemorale | Wu 9508-14* | China | KC928280 | KC928281 |
| H. niveomarginatum | CLZhao 25078* | China | OR141728 | OR506179 |
| H. nudicephalum | Wu9307_29 | China | AJ534269 | – |
| H. nudicephalum | CLZhao 17839 | China | OM985721 | OM985760 |
| H. obtusiforme | KHL1464 | Spain | JN572909 | – |
| H. obtusiforme | KHL11105 | Spain | JN572910 | – |
| H. occidentale | KHL 8477 (GB) | China | DQ677499 | DQ677499 |
| H. paramacaronesicum | MA Fungi 87736* | USA | KC984399 | – |
| H. paramacaronesicum | MA Fungi 87737 | China | KC984405 | – |
| H. pinicola | Wu 0108-32 | China | KJ885181 | KJ885182 |
| H. pinicola | Wu 0108-36 | China | KC928278 | KC928279 |
| H. prosopidis | ARIZ HHB 8479 | China | HE577029 | – |
| H. puerense | CLZhao 9476* | China | MW443045 | – |
| H. puerense | CLZhao 9583 | China | MW443046 | MW443051 |
| H. qujingense | CLZhao 26018* | China | OR141729 | PP826263 |
| H. setigerum | FCUG 1200 | Sweden | AJ534273 | – |
| H. setigerum | FCUG 1688 | Sweden | AJ534272 | – |
| H. sinense | CLZhao 7963 | China | MW301679 | MW293730 |
| H. sinense | CLZhao 17811* | China | MW301682 | MW293732 |
| H. sordidum | CLZhao 27379 | China | OR141731 | – |
| H. sordidum | CLZhao 27390* | China | OR141732 | OR506180 |
| H. tabuliformis | Dai 35238* | China | PZ745286 | PZ745283 |
| H. tabuliformis | Dai 35239 | China | PZ745287 | – |
| H. tenuicystidiatum | Dai 31215 | China | PZ745288 | PZ745284 |
| H. tenuicystidiatum | Dai 29223* | China | PZ745289 | PZ745285 |
| H. tenuissimum | CLZhao 7221* | China | MW443049 | MW443054 |
| H. tenuissimum | CLZhao 16210 | China | MW443050 | MW443055 |
| H. tongbiguanense | CLZhao 39574 | China | PV470531 | – |
| H. tongbiguanense | CLZhao 39575 | China | PV470532 | – |
| H. transiens | NH 12304 | Sweden | DQ677504 | DQ677504 |
| H. tropicum | CLZhao 17308* | China | OM985727 | OM985768 |
| H. variolosum | CBS 734.91 | China | MH862320 | MH873992 |
| H. variolosum | CBS 735.91 | China | MH862321 | MH873993 |
| H. weishanense | CLZhao 22403* | China | OR141727 | OR506181 |
| H. yingjiangense | CLZhao 37195 | China | PV470533 | – |
| H. yingjiangense | CLZhao 37232 | China | PV470534 | – |
| H. yunnanense | CLZhao 8845* | China | OM985728 | OM985769 |
Phylogenetic analyses
The phylogenetic relationships of Hyphoderma were analyzed based on the ITS + nLSU sequences. Climacocystis borealis (Fr.) Kotl. & Pouzar and Diplomitoporus crustulinus (Bres.) Domański were used as the outgroups. To determine the phylogenetic affinities of the new species, the ITS sequences of the holotype specimens were used as queries in BLAST searches against the GenBank nucleotide database (Sayers et al. 2024), and the top hits with the highest similarity scores were retrieved for comparative analysis. The DNA sequences were aligned using MAFFT v. 7 with the G-INS-i option (Katoh and Standley 2013). Alignments were optimized manually in BioEdit v. 7.0.9 (Hall 1999) and concatenated in Mesquite v. 3.2 (Maddison and Maddison 2017). The partition homogeneity test (PHT; Farris et al. 1994) for the dataset was conducted in PAUP v. 4.0b10 (Swofford 2002) using 1,000 homogeneity replicates. The combined dataset was initially divided into four data blocks (ITS1, 5.8S, ITS2, and nLSU). The optimal partition scheme and substitution models for Bayesian inference (BI) were selected using PartitionFinder (Lanfear et al. 2017; Zhang et al. 2020) under the Akaike information criterion (AIC) within PhyloSuite v. 1.2.3. For maximum likelihood (ML) analysis, the optimal substitution models were automatically determined by ModelFinder (Kalyaanamoorthy et al. 2017), as implemented in IQ-TREE v. 2.2.0 (Minh et al. 2020).
The ML analyses were performed using IQ-TREE v. 2.2.0 (Minh et al. 2020) with the optimal substitution models determined by ModelFinder (Kalyaanamoorthy et al. 2017). Branch support was assessed using ultrafast bootstrap analysis with 5,000 replicates (Hoang et al. 2018). The BI analysis with MrBayes v. 3.2.7 (Ronquist et al. 2012) implemented the Markov chain Monte Carlo (MCMC) technique. Four Markov chains were run in two runs from random starting trees for 6 million generations for ITS + nLSU (Fig. 1), with one tree retained every 100th generation until the average standard deviation of split frequencies was below 0.01. The first 25% of the sampled trees were discarded as burn-in, and the remaining trees were used to reconstruct a majority-rule consensus and to calculate Bayesian posterior probabilities (BPP) for the clades. The sequence alignments and the retrieved topologies were deposited in TreeBASE (http://www.treebase.org), under accession ID 32742. Phylogenetic trees were visualized with FigTree v. 1.4.2 (http://tree.bio.ed.ac.uk/software/figtree), and additional layout was carried out using Adobe Illustrator CS v. 5. Branches that received ML bootstrap support ≥ 50% and BPP ≥ 0.90 were considered significantly supported. ML bootstrap support ≥ 50% and BPP ≥ 0.90 are presented on the topologies from the ML analysis.
Figure 1.

ML analysis of Hyphoderma based on the ITS + nLSU dataset. ML bootstrap values equal to or higher than 50% and Bayesian posterior probability values equal to or greater than 0.90 are shown. New taxa are given in bold.
Results
Phylogenetic analyses
In this study, 134 sequences from 79 fungal samples representing 50 taxa were used to reconstruct phylogenetic trees, including 79 ITS sequences and 55 nLSU sequences. The dataset had an aligned length of 2,040 characters, of which 1,397 (68%) were constant, 116 (6%) were variable and parsimony-uninformative, and 527 (26%) were parsimony-informative. For the BI analysis, the combined dataset was partitioned into three subsets (ITS1 + ITS2, 5.8S, and nLSU) based on the PartitionFinder results. The best-fit models selected under the Akaike information criterion (AIC) were GTR+I+G for ITS1 + ITS2, K80+I+G for 5.8S, and GTR+I+G for nLSU. For the ML analysis in IQ-TREE, the dataset was partitioned into three subsets (ITS1, 5.8S, and ITS2 + nLSU), with the best-fit models being GTR+I+G for ITS1, K80+I+G for 5.8S, and GTR+I+G for ITS2 + nLSU. The phylogenetic tree topology from the Bayesian analysis was similar to that from the ML analysis, so only the ML tree is provided (Fig. 1).
Taxonomy
Hyphoderma tabuliformis
X. Li, Yuan Yuan & Y.C. Dai sp. nov.
27FBFE96-920F-5F97-9B39-E574A4C72B34
864738
Figure 2.

Basidiomata of Hyphoderma tabuliformis (holotype, Dai 35238). Scale bars: 1 cm (A); 1 mm (B).
Figure 3.

Sections of the hymenium of Hyphoderma tabuliformis (holotype, Dai 35238). A–D. Basidiospores; E, F. Basidia; G, J. Basidioles; H, I. Cystidia; K. Generative hyphae. Scale bars: 10 µm (A–K).
Figure 4.

Microscopic structures of Hyphoderma tabuliformis (drawn from the holotype). a. Basidiospores; b. Basidia; c. Basidioles; d. Cystidia; e. Section of basidioma.
Diagnosis.
Hyphoderma tabuliformis is distinguished by membranaceous to ceraceous basidiomata when fresh, arachnoid hymenial surface, clavate cystidia, hyphae encrusted with crystals, and usually with oil drops inside basidiospores.
Holotype.
China • Qinghai Province, Huzhu County, Beishan Forest Park, Langshidang, GPS coordinates 36°53'N, 102°21'E, altitude 2,900 m asl., on a fallen branch of Pinus tabuliformis, 10 August 2025, Dai 35238 (holotype BJFC056499).
Etymology.
Tabuliformis (Lat.): refers to the species growing on Pinus tabuliformis.
Description.
Basidiomata. Annual, resupinate, adnate, without odor and taste, membranaceous to ceraceous when fresh, and up to 7 cm long, 1.6 cm wide, and 100 µm thick at the center. Hymenial surface arachnoid, white to cream when fresh, and pinkish buff when dry. Sterile margin indeterminate, white to cream.
Subicular hyphae. Monomitic, generative hyphae with clamp connections, thin-walled, more or less interwoven, encrusted with crystals, 3.0–4.5 µm in diam., IKI–, CB–; tissues unchanged in KOH.
Cystidia. Clavate, smooth, thin-walled, 48–68 × 4.8–6.6 µm.
Basidia. Clavate, thin-walled, more or less constricted in the middle, usually with oil drops inside, with four sterigmata and a basal clamp connection, 28–40 × 4.0–5.8 µm; basidioles similar to basidia in shape, but smaller.
Basidiospores. Subcylindrical to cylindrical, thin-walled, smooth, usually with oil drops inside, IKI–, CB–, (8.4–)8.7–12.2(–11.9) × (3.5–)3.6–4.2(–4.3) µm, L = 10.24 µm, W = 3.93 µm, Q = 2.57–2.64 (n = 60/2).
Type of rot. White rot.
Additional specimen (paratype) examined.
China • Qinghai Province, Huzhu County, Beishan Forest Park, Langshidang, GPS coordinates 36°53'N, 102°21'E, altitude 2,900 m asl., on a fallen branch of Pinus tabuliformis, 10 August 2025, Dai 35239 (BJFC056500).
Note.
In the phylogenetic tree (Fig. 1), Hyphoderma tabuliformis is closely related to H. medioburiense, although the ITS region of H. tabuliformis differs from that of H. medioburiense by 5.8%. Hyphoderma medioburiense differs from H. tabuliformis by its wider basidia (7–8.5 vs. 4.0–5.8 µm), longer cystidia (70–105 vs. 48–68 µm), and wider basidiospores (4.5–5 vs. 3.6–4.2 µm; Donk 1957). Morphologically, H. tabuliformis is similar to H. alboarachnum and H. prosopidis in sharing an arachnoid hymenial surface and similar hyphal structure. However, H. alboarachnum is distinguished from H. tabuliformis by having smaller basidiospores (5–6 × 2–3 vs. 8.7–12.2 × 3.6–4.2 µm) and lacking cystidia (Li et al. 2026). Hyphoderma prosopidis differs from H. tabuliformis by having wider basidia (8.0–11.0 vs. 4.0–5.8 µm) and wider basidiospores (5.5–8.5 vs. 3.6–4.2 µm; Telleria et al. 2012).
Hyphoderma tenuicystidiatum
X. Li, Yuan Yuan & Y.C. Dai sp. nov.
93473390-34CF-5F37-B745-A29C6E83DD4A
864739
Figure 5.

Basidiomata of Hyphoderma tenuicystidiatum (holotype, Dai 29223). Scale bars: 1 cm (A); 1 mm (B).
Figure 6.

Sections of the hymenium of Hyphoderma tenuicystidiatum (holotype, Dai 29223). A–D. Basidiospores; E, F. Basidia; G, J. Basidioles; H, I. Cystidia; K. Generative hyphae. Scale bars: 10 µm (A–K).
Figure 7.

Microscopic structures of Hyphoderma tenuicystidiatum (drawn from the holotype). a. Basidiospores; b. Basidia; c. Basidioles; d. Cystidia; e. Section of basidioma.
Diagnosis.
Hyphoderma tenuicystidiatum is distinguished by light vinaceous gray hymenial surface when fresh and cream when dry, cylindrical cystidia, fusoid cystidioles and oil drops inside basidiospores.
Holotype.
China • Qinghai Province, Tongren County, Shuangpengxi, GPS coordinates 35°56'N, 102°20'E, altitude 3,068 m asl., on rotten wood of Picea crassifolia, 24 July 2024, Dai 29223 (holotype BJFC049482).
Etymology.
Tenuicystidiatum (Lat.): refers to the species having thin cystidia.
Description.
Basidiomata. Annual, resupinate, adnate, without odor and taste when fresh, membranaceous, up to 5.0 cm long, 3.5 cm wide, and 100 µm thick. Hymenial surface smooth, light vinaceous gray when fresh, cream when dry. Sterile margin indeterminate, light vinaceous gray.
Subicular hyphae. Monomitic, generative hyphae with clamp connections, thin-walled, more or less interwoven, 2.5–3.5 µm diam., CB–, IKI–; tissues unchanged in KOH.
Cystidia. Cylindrical, smooth, thin-walled, occasionally with oil drops inside, 40–55 × 4.3–7.7 µm; cystidioles present, fusoid, smooth, thin-walled, 30–38 × 5.4–6.3 µm.
Basidia. Clavate, occasionally with oil drops inside, with four sterigmata and a basal clamp connection, 25–38 × 6.9–9.0 µm; basidioles similar to basidia in shape, but smaller.
Basidiospores. Subcylindrical to oblong ellipsoid, thin-walled, smooth, with oil drops inside, IKI–, CB–, (9.5–)10.4–14.9(–15.7) × (5.1–)5.2–7.1(–7.2) µm, L = 12.30 µm, W = 6.12 µm, Q = 1.89–2.04 (n = 60/2).
Type of rot. White rot.
Additional specimen (paratype) examined.
China • Qinghai Province, Xunhua County, Mengdatianchi Nature Reserve, GPS coordinates 35°51'N, 102°38'E, altitude 3,000 m asl., on a dead angiosperm tree, 12 October 2024, Dai 31215 (BJFC051474).
Notes.
In the phylogenetic tree (Fig. 1), Hyphoderma tenuicystidiatum and H. tabuliformis formed two well-supported lineages (100% in ML and 1.00 BPP; 100% in ML and 1.00 BPP, respectively). The phylogenetic analyses revealed that H. tenuicystidiatum is related to H. obtusiforme, but the ITS region of H. tenuicystidiatum differs from that of H. obtusiforme by 4.4%. Hyphoderma obtusiforme is distinguished from H. tenuicystidiatum by having wider cystidia (8–10 vs. 4.3–7.7 µm; Eriksson and Ryvarden 1975). Morphologically, H. tenuicystidiatum is highly similar to H. microporoides, H. fissuratum, and H. transiens in having clavate basidia, similar hyphal structure, and a smooth hymenial surface. However, H. microporoides differs from H. tenuicystidiatum by having smaller basidiospores (8.5–10 × 2.5–3.5 vs. 10.4–14.9 × 5.2–7.1 µm) and capitate cystidia (Guan and Zhao 2021). Hyphoderma fissuratum is distinguished from H. tenuicystidiatum by having smaller basidiospores (8.5–10.3 × 3–4 vs. 10.4–14.9 × 5.2–7.1 µm) and lacking cystidia (Ma et al. 2021). Hyphoderma transiens differs from H. tenuicystidiatum by having narrower basidiospores (3.5–4 vs. 5.2–7.1 µm; Parmasto 1968).
Discussion
In this study, morphological and phylogenetic analyses were conducted on Hyphoderma specimens collected from Qinghai Province, and two new species were identified and described (Fig. 1; Table 1), raising the total number of Hyphoderma species known from China from 45 to 47 (Yang et al. 2023; Li et al. 2026). Because the molecular phylogenetic analyses included only a subset of the currently accepted species of Hyphoderma, phylogenetic placement alone cannot exclude the possibility that the newly collected specimens correspond to described species for which molecular data are unavailable. To address this issue, the two new species were compared with morphologically similar accepted species, especially those lacking molecular data, as well as those represented in the phylogenetic analyses (Table 2). The morphological characters compared included basidiomata morphology; cystidia type, shape, and size; basidia shape and size; and basidiospore shape and size. Therefore, although some accepted Hyphoderma species still lack molecular data, the recognition of the new species is supported by an integrated combination of molecular phylogenetic position, morphological diagnosis, comparison with the closest phylogenetic relatives, and comparison with morphologically similar species.
Table 2.
Morphological comparison of the new species with similar taxa of the genus Hyphoderma.
| Species name | Basidiomata | Hyphal system | Cystidia | Basidia | Basidiospores |
|---|---|---|---|---|---|
| Hyphoderma acutatum | Hymenial surface smooth, whitish to ochraceous | Monomitic, thin-walled, generative hyphae with clamp connections | Subulate, thin-walled, 100–170 × 15–20 µm | Suburniform, 25–30 × 5–6 µm, four sterigmata, a basal clamp connection | Ellipsoid, 7–7.5(–8–9) × 4–4.5(–5) µm |
| H. alboarachnum | Hymenial surface arachnoid, white to cream | Monomitic, thin-walled, generative hyphae with clamp connections | Absent | Cylindrical, 23.5–30 × 3.5–4.5 µm, four sterigmata, a basal clamp connection | Oblong–ellipsoid to cylindrical; 5–6 × 2–3 µm. |
| H. ayresii | Hymenial surface smooth, whitish to pale pinkish buff | Monomitic, thin- to slightly thick-walled, generative hyphae with clamp connections | Subulate, thick-walled, encrusted, 60–160 × 15–30 µm | Subutriform to subclavate, 37–47 × 10–12.5 µm, four sterigmata, a basal clamp connection | Ellipsoid, 11.5–13.5 × 7–8 μm |
| H. bicystidiatum | Hymenial surface smooth, yellowish white to pale yellow | Monomitic, thin-walled, generative hyphae with clamp connections | 1) tubular, thin-walled, 82.0–122.0 × 12.3–13.8 µm. 2) clavate to subclavate, thick-walled, 69.0–110.0 × 11.0–13.8 µm | Clavate to subclavate, 13.6–22.3 × 6.2–7.5 µm, four sterigmata, a basal clamp connection | subellipsoid to suballantoid, 6.2–8.8 × 3.0–4.1 µm |
| H. clavatum | Hymenial surface smooth or slightly tuberculate, grayish clay | Monomitic, thin-walled, generative hyphae with clamp connections | Clavate or apically globose, thin-walled or apically slightly thick-walled, 30–60 µm long, 10–15 µm diam for the globose parts | Subclavate, 30–40 × 6.5–8 µm, four sterigmata, a basal clamp connection | Cylindrical, 10–13 × 4.2–5.2 µm |
| H. fissuratum | Hymenial surface smooth, cream | Monomitic, thin-walled, generative hyphae with clamp connections | Absent | Narrowly clavate, 24–28 × 4–5.5 µm, four sterigmata, a basal clamp connection | Cylindrical, 8.5–10.2 × 3–4 µm |
| H. galactinum | Hymenial surface smooth to tuberculate | Monomitic, thin-walled, generative hyphae with clamp connections | Cylindrical, often more or less moniliform with several constrictions, thin to slightly thick-walled, 40–80 × 6–10 µm. | Subclavate to subventricose, 25–35 × 7–8 μm, four sterigmata, a basal clamp connection | Cylindrical to suballantoid, 13–16 × 4–4.5 μm |
| H. hjortstamii | Hymenial surface smooth, yellowish white to pale yellow | Monomitic, thin-walled, generative hyphae with clamp connections | Subcylindrical, thin-walled, 40–55 × 5.6–6.8 µm | Clavate, 17–26 × 5–6.8 µm, four sterigmata, a basal clamp connection | Suballantoid to allantoid, 8–11 × 3.1–4.4 µm |
| H. incrustatum | Hymenial surface smooth to tuberculate, pale orange to greyish orange to greyish red | Monomitic, thin-walled, generative hyphae with clamp connections | 1) tubular, thin-walled, encrusted, 76–90 × 7–8 µm. 2) capitate, encrusted, 33–43 × 4.6–11.7 µm | Clavate, 27–39 × 6.2–8.7 μm, four sterigmata, a basal clamp connection | Ellipsoid to suballantoid, 10.8–16 × 4.3–5.2 μm |
| H. lunasporum | Hymenial surface smooth, white to cream | Monomitic, thin-walled, generative hyphae with clamp connections | Cylindrical to subclavate, thin- to slightly thick-walled, 30–60 × 5–8 µm | Subclavate to suburniform, 20–30 × 5–7 μm, four sterigmata, a basal clamp connection | Allantoid to lunate, 8–12 × 3–4.5 μm |
| H. malenconii | Hymenial surface smooth to tuberculate, ivory yellow to light buff to pale olive–buff | Monomitic, thin- to slightly thick-walled, generative hyphae with clamp connections | Cylindrical, thin-walled, 35–90 × 7–12 µm | Utriform to subutriform, 30–50 × 7–9 µm, four sterigmata, a basal clamp connection | Cylindrical, 12–15.5 × 5–6 μm |
| H. microcystidium | Hymenial surface smooth, whitish | Monomitic, thin- to slightly thick-walled, generative hyphae with clamp connections | 1) Lamprocystidia, slightly thick-walled, encrusted, 17–32 × 7–10.5 µm. 2) Leptocystidia, cylindrical, 20–65 × 8–13 µm | Subutriform, 18–25 × 6.5–7.5 µm, four sterigmata, a basal clamp connection | Narrowly ellipsoid, 10–12.5 × 4–4.8 μm |
| H. microporoides | Hymenial surface smooth, cream to pale buff | Monomitic, thin-walled, generative hyphae with clamp connections | Capitate halocystidia, thin-walled, 18–51 × 4.5–7 µm | Clavate, 18.5–29.5 × 5–7 µm, four sterigmata, a basal clamp connection | Cylindrical to allantoid, 8.5–10 × 2.5–3.5 µm |
| H. rimulosum | Hymenial surface smooth, white to ivory yellow | Monomitic, thin-walled, generative hyphae with clamp connections | Cylindrical, slightly thick-walled, usually encrusted throughout, 30–55 × 4–6 µm | Subclavate, 22–32 × 4.5–6 µm, four sterigmata, a basal clamp connection | Ellipsoid, 6–7 × 3.9–4.1 μm |
| H. subclavatum | Hymenial surface smooth, clay–yellow | Monomitic, thin- to slightly thick-walled, generative hyphae with clamp connections | Clavate, thin-walled, 40–70 × 8–11 µm, apically often with one to several secondary septa. | Suburniform, 40–55 × 7–8 µm, four sterigmata, a basal clamp connection | Cylindrical, 10–12 × 4.2–5.3 µm |
| H. subglobosum | Hymenial surface smooth, grayish yellow to grayish orange | Monomitic, thin-walled, generative hyphae with clamp connections | Clavate, thin-walled, 50.0–69.0 × 8.0–11.8 µm | Clavate to subclavate, 22.0–34.0 × 6.8–10.0 µm with oily contents, four sterigmata, a basal clamp connection | Subglobose to broadly ellipsoid, 11.8–13.1 × 8.0–10.0 µm |
| H. tabuliformis | Hymenial surface arachnoid, white to cream | Monomitic, thin-walled, generative hyphae with clamp connections | Clavate, thin-walled, 48–68 × 4.8–6.6 µm | Clavate, 28–40 × 4.0–5.8 µm, four sterigmata, a basal clamp connection | Subcylindrical to cylindrical, 8.7–12.2 × 3.6–4.2 µm |
| H. tenuicystidiatum | Hymenial surface smooth, light vinaceous gray | Monomitic, thin-walled, generative hyphae with clamp connections | Cylindrical, thin-walled, 40–55 × 4.3–7.7 µm | Clavate, 25–38 × 6.9–9.0 µm, four sterigmata, a basal clamp connection | Subcylindrical to oblong ellipsoid, 10.4–14.9 × 5.2–7.1 µm |
| H. transiens | Hymenial surface odontioid, whitish, capucine buff to light ochraceous buff | Monomitic, thin-walled, generative hyphae with clamp connections | Subcylindrical to cylindrical, thin-walled, 63.0–76.0 × 9.4–11.8 μm | Clavate to subclavate, 20.0–32.0 × 6.2–7.6 μm, four sterigmata, a basal clamp connection | Ellipsoid to suballantoid, 9.6–10.6 × 3.4–4.2 μm |
In terms of ecology and host associations, Hyphoderma is an important genus of wood-decaying fungi, widely distributed worldwide and playing a key role in forest carbon cycling (Dai et al. 2021; Zhou et al. 2023). Most Hyphoderma species grow on hardwoods, whereas a few occur on coniferous wood (Dai et al. 2004b). The two new species differ in host use: H. tenuicystidiatum was found on rotten wood of Picea and dead angiosperm trees, indicating a relatively broad host range, whereas H. tabuliformis was found only on fallen branches of Pinus tabuliformis, appearing to be more specialized. This suggests that different species within the genus may have distinct host preferences and ecological adaptation strategies, potentially reflecting differences in lignin-degrading capacity or niche partitioning, although most species are not strictly selective in their wood substrates (Dai 2010; Yurchenko and Wu 2015; He et al. 2019). Because sampling was restricted to a limited area in Qinghai, additional Hyphoderma species may remain to be discovered in under-surveyed areas, such as the margins of the Qinghai–Tibet Plateau and the boreal coniferous forest zones of northeastern China. In addition, the present study relied solely on ITS and nLSU sequences, mainly because the vast majority of Hyphoderma species currently lack RPB1, RPB2, and TEF1 sequences in public databases, and incorporating these genes would substantially reduce taxonomic coverage. Therefore, future studies should expand sampling across broader geographic and habitat ranges, generate and deposit multilocus sequences of Hyphoderma in public databases, and incorporate these data into future analyses to further investigate the diversity, taxonomy, and evolution of Hyphoderma.
Key to species of Hyphoderma from China
| 1 | Cystidia absent | 2 |
| – | Cystidia present | 5 |
| 2 | Hymenial surface arachnoid | H. alboarachnum |
| – | Hymenial surface smooth | 3 |
| 3 | Hymenial surface cream when fresh | H. fissuratum |
| – | Hymenial surface white when fresh | 4 |
| 4 | Generative hyphae with clamp connection | H. weishanense |
| – | Generative hyphae simple with simple-septa | H. yingjiangense |
| 5 | One type of cystidia present | 6 |
| – | Two types of cystidia present | 28 |
| 6 | Cystidia encrusted with crystals | 7 |
| – | Cystidia not encrusted with crystals | 12 |
| 7 | Basidiospores > 7.5 µm long | 8 |
| – | Basidiospores < 7.5 µm long | 11 |
| 8 | Cystidia > 60 µm long | 9 |
| – | Cystidia < 60 µm long | H. fulgens |
| 9 | Cystidia thick-walled | 10 |
| – | Cystidia thin-walled | H. yunnanense |
| 10 | Cystidia subulate | H. macrosporum |
| – | Cystidia fusiform | H. mopanshanense |
| 11 | Cystidia cylindrical | H. rimulosum |
| – | Cystidia tubular | H. puerense |
| 12 | Basidiospores > 6.5 µm long | 13 |
| – | Basidiospores < 6.5 µm long | 27 |
| 13 | Basidia 2-sterigmata | H. fragilissimum |
| – | Basidia 4-sterigmata | 14 |
| 14 | Halocystidia absent | 15 |
| – | Halocystidia present | 25 |
| 15 | Septate cystidia absent | 16 |
| – | Septate cystidia present | 24 |
| 16 | Cheilocystidia absent | 17 |
| – | Cheilocystidia present | H. marginatum |
| 17 | Hymenial surface grandinioid | H. grandineum |
| – | Hymenial surface smooth | 18 |
| 18 | Basidia > 40 µm long | H. subclavatum |
| – | Basidia < 40 µm long | 19 |
| 19 | Basidiospores > 6 µm wide | 20 |
| – | Basidiospores < 6 µm wide | H. tabuliformis |
| 20 | Basidiospores > 10 µm long | 21 |
| – | Basidiospores < 10 µm long | 23 |
| 21 | Basidiomata membranaceous | 22 |
| – | Basidiomata subceraceous | H. hjortstamii |
| 22 | Basidiospores cylindrical | H. clavatum |
| – | Basidiospores subcylindrical to oblong ellipsoid | H. tenuicystidiatum |
| 23 | Cystidia cylindrical | H. litschaueri |
| – | Cystidia moniliform | H. asianum |
| 24 | Hymenial surface cracked | H. subsetigerum |
| – | Hymenial surface not cracked | H. subpraetermissum |
| 25 | Basidia clavate | 26 |
| – | Basidia barrelled | H. qujingense |
| 26 | Basidiospores > 8.5 µm long | H. microporoides |
| – | Basidiospores < 8.5 µm long | H. punctatum |
| 27 | Basidia clamped at base | H. sordidum |
| – | Basidia simple-septate at base | H. bambusinum |
| 28 | Septate cystidia absent | 29 |
| – | Septate cystidia present | 32 |
| 29 | Basidiospores > 9 µm long | 30 |
| – | Basidiospores < 9 µm long | H. laceratum |
| 30 | Basidia clamped at base | 31 |
| – | Basidia simple-septate at base | H. cinereofuscum |
| 31 | Basidiospores allantoid | H. crystallinum |
| – | Basidiospores cylindrical | H. nitidum |
| 32 | Basidia 2-sterigmata | H. pinicola |
| – | Basidia 4-sterigmata | 33 |
| 33 | Hymenial surface grandinioid | H. setigerum |
| – | Hymenial surface smooth | H. guangdongense |
Supplementary Material
Funding Statement
This study was financed by the National Natural Science Foundation of China (Project Nos. 32370013, 32300013, 32161143013) and the macrofungal diversity of Qinghai Province (Project GHMB-2024-18).
Additional information
Conflict of interest
The authors have declared that no competing interests exist.
Ethical statement
No ethical statement was reported.
Artificial Intelligence (AI) use
The authors accept full responsibility for the content of the manuscript, including the disclosure of any use of AI.
No AI tools were used in the preparation of this manuscript.
Funding
This study was financed by the National Natural Science Foundation of China (Project Nos. 32370013, 32300013, and 32161143013) and the macrofungal diversity of Qinghai Province (Project GHMB-2024-18).
Author contributions
The authors confirm contribution to the paper as follows: conception and design of the study: Li X, Cui YJ; data collection: Li X, Cui YJ, Xing R, Dai YC, Yuan Y, Wu YD; analysis and interpretation of results: Li X, Yuan Y; draft manuscript preparation: Li X. All authors reviewed the results and approved the final version of the manuscript.
Author ORCIDs
X. Li https://orcid.org/0009-0001-1625-8589
Y. Cui https://orcid.org/0009-0000-8706-6771
Y. Dai https://orcid.org/0000-0002-6523-0320
Data availability
All of the data that support the findings of this study are available in the main text.
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Data Availability Statement
All of the data that support the findings of this study are available in the main text.
