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. 2026 Aug 3;138:57–94. doi: 10.3897/mycokeys.138.204396

Three new wood-inhabiting corticioid fungi from the Qinling Mountains, China

Le-Le Wan 1, Lan-Shuo Zhang 1, Jun-Hong Dong 1, Long Zeng 1, Shun Liu 1,✉, Bao-Kai Cui 1,✉
PMCID: PMC13458435  PMID: 42582694

Abstract

Wood-inhabiting fungi are key decomposers in forest ecosystems, but the diversity of corticioid fungi in the Qinling Mountains remains insufficiently explored. During surveys of macrofungi in Zhashui County and Shangzhou District in the Qinling Mountains, several specimens representing three undescribed taxa of corticioid fungi were collected. Based on detailed morphological examinations and phylogenetic analyses of combined ITS+nLSU datasets by using Maximum Likelihood and Bayesian Inference methods, three new species, viz. Hymenochaete zhashuiensis, Hyphoderma brevicystidiatum and Steccherinum shangzhouense, are described and illustrated. Hymenochaete zhashuiensis is characterized by effused-reflexed basidiomata with a reddish to rust-brown, cracked hymenial surface, abundant dark brown hymenial setae, the presence of dendrohyphidia, and ellipsoid to broadly ellipsoid basidiospores (4–5.6 × 2.8–3.5 µm). Hyphoderma brevicystidiatum is distinguished by resupinate, basidiomata white to cream with hymenial surface, clamped generative hyphae, short cylindrical to subcylindrical cystidia, subclavate to clavate basidia, and ellipsoid to subcylindrical basidiospores (6.2–10 × 2.7–4.2 µm). Steccherinum shangzhouense is characterized by resupinate and membranaceous basidiomata, a white to cream, tuberculate to slightly grandinioid and cracked hymenial surface, simple-septate generative hyphae, the absence of cystidia and skeletocystidia, and broadly ellipsoid to oblong ellipsoid basidiospores (5.2–7.9 × 2.5–5 µm). These findings enrich the wood-inhabiting fungal diversity in the Qinling Mountains and indicate that this region remains underexplored for wood-inhabiting fungi.

Key words: Macrofungi, molecular phylogeny, new taxa, Qinling Mountains, taxonomy

Introduction

Wood-inhabiting fungi are a group of macrofungi primarily composed of polyporoid and corticioid fungi, in which they play a crucial role as decomposers in forest ecosystems, driving wood degradation and the recycling of organic matter (Dong et al. 2024). Among them, polypore fungi usually produce poroid basidiomata and have been well investigated in China (Wu et al. 2022; Xu et al. 2025). While corticioid fungi usually produce resupinate or effused-reflexed basidiomata, their inconspicuous basidiomata often cause these fungi to be overlooked in biodiversity surveys, and 1400 species have been reported in China (Dai et al. 2004b; Yuan et al. 2026). During surveys of macrofungi in Zhashui County and Shangzhou District of the Qinling Mountains, we collected several corticioid fungal specimens that cannot be assigned to any known species. Preliminary morphological observations indicate that these specimens belong to the genera Hymenochaete Lév., Hyphoderma Wallr. and Steccherinum Gray.

The genus Hymenochaete was established by Léveillé (1846), with H. rubiginosa (Dicks.) Lév. as the type species and belongs to Hymenochaetaceae Donk, Hymenochaetales Oberw. (Dai 2010; He and Dai 2012; Parmasto et al. 2014; Liu et al. 2025). The genus exhibits annual to perennial, resupinate, effused-reflexed, pileate or stipitate basidiomata, brownish to reddish brown hymenial surface, monomitic or dimitic hyphal system, simple-septate generative hyphae, hymenial setae or other setal elements, and hyaline to brownish, thin- to thick-walled basidiospores (Parmasto 2001; He and Dai 2012; Liu et al. 2025). Traditional classification discriminated species or sections mainly based on basidiomatal structure and the presence or absence of layers such as the setal layer, context or subiculum, and cortex (Burt 1918; Léger 1998). However, molecular phylogenetic studies have shown that morphological characters alone are insufficient to delimit species within this genus (Wagner and Fischer 2002; Parmasto et al. 2014; Liu et al. 2025). Subsequent molecular phylogenetic studies have clarified the phylogenetic relationships of Hymenochaete and related non-poroid Hymenochaetaceae and revealed extensive cryptic diversity worldwide (Parmasto et al. 2014; Liu et al. 2025; Wang et al. 2026). In China, some species of Hymenochaete have been described based on morphological characteristics and molecular phylogenetic evidence (Dai 2010; He and Dai 2012; He et al. 2017; Liu et al. 2025; Wang et al. 2026). To date, the genus Hymenochaete comprises approximately 265 recognized species, of which 120 have been recorded in China (Liu et al. 2025; Zhao et al. 2025; Wang et al. 2026; Xu et al. 2026; Zhu et al. 2026).

The genus Hyphoderma was established by Wallroth (1833), with H. setigerum (Fr.) Donk as the type species and belongs to Hyphodermataceae Jülich, Polyporales Gäum. (Donk 1957; Larsson 2007; Justo et al. 2017; Li et al. 2026; Yang et al. 2025). The genus exhibits considerable macromorphological diversity, with resupinate to effused-reflexed, ceraceous to membranaceous basidiomata, smooth, tuberculate, grandinioid or odontioid hymenophore, monomitic to dimitic hyphal system with clamped generative hyphae, suburniform to subcylindrical or clavate basidia, and ellipsoid to subglobose basidiospores (Bernicchia and Gorjón 2010; Guan et al. 2021; Duan et al. 2023; Yang et al. 2023). Taxonomically, the generic delimitation of Hyphoderma and its related genera has long been controversial. Traditional classification included a broad assemblage of corticioid fungi in Hyphoderma, but subsequent molecular phylogenetic studies demonstrated that the traditionally circumscribed genus was polyphyletic, leading to the transfer of several species to related genera such as Peniophorella P. Karst. (Larsson 2007; Justo et al. 2017; Li et al. 2026). Subsequent molecular phylogenetic studies have refined the circumscription of Hyphoderma and confirmed the independent phylogenetic position of the core Hyphoderma lineage within Hyphodermataceae (Justo et al. 2017; Yang et al. 2023, 2025; Li et al. 2026). In recent years, multiple new species of Hyphoderma have been described in China based on morphological characteristics and molecular phylogenetic evidence (Guan and Zhao 2021a, 2021b; Ma et al. 2021; Yang et al. 2023, 2025; Su et al. 2024). Currently, the genus Hyphoderma comprises approximately 132 species, of which 54 species are distributed in China (Guan et al. 2021; Li et al. 2026; Yuan et al. 2026).

The genus Steccherinum was established by Gray (1821), with S. ochraceum (Pers.) Gray as the type species and belongs to Steccherinaceae Parmasto, Polyporales (Zhang et al. 2025; Westphalen et al. 2026). The genus exhibits resupinate to effused-reflexed or pileate basidiomata with membranaceous to corky or waxy, and odontioid, hydnoid, grandinioid or poroid hymenophore, monomitic to dimitic hyphal system, thick-walled and encrusted skeletocystidia, and ellipsoid to subglobose basidiospores (Ryvarden 1991; Westphalen et al. 2021, 2026). Taxonomically, the generic delimitation between Steccherinum and its closely related genus Junghuhnia Corda has long been controversial (Westphalen et al. 2018). Traditional classification discriminates these two genera mainly based on hymenophoral morphology, with Junghuhnia characterized by a poroid hymenophore and Steccherinum by an odontioid hymenophore (Du et al. 2020). Subsequent molecular phylogenetic studies have verified that Steccherinum comprised both poroid and odontioid species, further clarifying its independent phylogenetic position within the family Steccherinaceae (Miettinen et al. 2012; Justo et al. 2017; Yuan et al. 2019; Westphalen et al. 2021, 2026). Recently, multiple new Steccherinum species have been described based on morphological characteristics and molecular phylogenetic evidence in China (Dong et al. 2022, 2023; Liu et al. 2023b; Wang et al. 2024; Zhang et al. 2025). The genus Steccherinum currently comprises 91 accepted species worldwide, and 29 Steccherinum taxa have been published or recorded from China (Yuan and Wu 2012; Liu and Dai 2021; Dong et al. 2022; Yuan et al. 2026).

To further clarify the taxonomic status of these specimens, species identifications were performed based on detailed morphological observations and phylogenetic analyses of the combined ITS+nLSU dataset. The results indicate that these specimens represent three new species belonging to Hymenochaete, Hyphoderma, and Steccherinum, respectively.

Materials and methods

Morphological studies

Fresh specimens of Hymenochaete, Hyphoderma and Steccherinum were collected from Zhashui County and Shangzhou District, Shaanxi Province, China. The basidiomata were photographed in situ, and macromorphological characters, substrates and habitats were recorded in the field. The specimens were dried and deposited at the herbarium of the Institute of Microbiology, Beijing Forestry University (BJFC), Beijing, China. Color terms used in macromorphological descriptions follow Kornerup and Wanscher (1967).

Macromorphological descriptions were based on field notes and laboratory observations. Micromorphological characters, measurements and drawings were made from slide preparations mounted in 5% potassium hydroxide (KOH), Cotton Blue (CB) and Melzer’s reagent (IKI), using a Nikon Eclipse 80i microscope (Nikon, Tokyo, Japan). At least 30 basidiospores were measured from each specimen. Sterigmata were excluded from basidial measurements, and the hilar appendage was excluded from basidiospore measurements. The following abbreviations are used: CB = Cotton Blue, CB– = acyanophilous, IKI = Melzer’s reagent, IKI– = neither amyloid nor dextrinoid, KOH = 5% potassium hydroxide, L = mean basidiospore length, W = mean basidiospore width, Q = range of the length/width ratio of basidiospores, Qm = mean length/width ratio of basidiospores, and n = a/b, where a represents the number of measured structures and b represents the number of specimens examined. When sufficient measurements were available, 5% of measurements were excluded from each end of the range and are given in parentheses. The detailed morphological methods followed Sun et al. (2022) and Liu et al. (2023a).

DNA extraction and sequencing

Genomic DNA was extracted from dried specimens using the CTAB rapid plant genome DNA extraction kit DN14 (Aidlab Biotechnologies, Beijing, China), following the manufacturer’s instructions with modifications as described by Song et al. (2025). The internal transcribed spacer region (ITS) was amplified with primers ITS5 and ITS4 (White et al. 1990), and the nuclear large subunit ribosomal RNA gene region (nLSU) was amplified with primers LR0R and LR7 (Hopple Jr and Vilgalys 1999).

The polymerase chain reaction (PCR) program for ITS consisted of an initial denaturation at 95 °C for 3 min, followed by 35 cycles at 94 °C for 40 s, 56 °C for 45 s and 72 °C for 1 min, and a final extension at 72 °C for 10 min. The PCR program for nLSU consisted of an initial denaturation at 94 °C for 1 min, followed by 35 cycles at 94 °C for 30 s, 50 °C for 1 min and 72 °C for 90 s, and a final extension at 72 °C for 10 min. PCR products were sequenced at the Beijing Genomics Institute (BGI), China, using the same primers.

The newly generated sequences were assembled and edited, and then deposited in GenBank. All newly generated sequences and reference sequences used in the phylogenetic analyses are listed in Table 1.

Table 1.

Taxa information and GenBank accession numbers of sequences used in this study.

SPECIES NAME SPECIMEN VOUCHER ITS nLSU REFERENCE
Cabalodontia delicata MCW 693/19 MT849297 MT849297 Westphalen et al. 2021
Diplomitoporus crustulinus FD-137 KP135299 KP135211 Justo et al. 2017
Hydnoporia olivacea Miettinen X3273 MK514610 – Miettinen et al. 2019
H. olivacea Miettinen X3403 MK514612 – Miettinen et al. 2019
Hymenochaete acanthophysata CBS 925.96 MH862623 AF385144 Vu et al. 2019; Wagner and Fischer 2002
H. acerosa He 344 NR_120042 – He and Li 2011
H. adhaerens Spirin 4994 KM017411 – Spirin et al. 2015
H. adhaerens Spirin 6246 KM017412 – Spirin et al. 2015
H. adnata He 560 – KU975527 Liu et al. 2025
H. adnata He 951 – KU975528 Liu et al. 2025
H. adnata He 655 OR287568 OR287643 Liu et al. 2025
H. adusta He 207 JQ279523 KU975497 Nie et al. 2017
H. alpina He 1388 OR287604 OR287674 Liu et al. 2025
H. alpina He 4918 OR287605 OR287675 Liu et al. 2025
H. angustispora Dai 17045 MF370592 MF370598 He et al. 2017
H. angustispora Dai 17049 MF370593 MF370599 He et al. 2017
H. anomala He 1276 OR287531 OR287616 Liu et al. 2025
H. anomala He 592 JQ279566 JQ279650 He and Dai 2012
H. asetosa Dai 10756 JQ279559 JQ279642 He and Dai 2012
H. asiatica He 1270 OR287593 OR287663 Liu et al. 2025
H. asiatica He 5800 OR287594 – Liu et al. 2025
H. atrobrunnea Dai 18810 OR287608 OR287656 Liu et al. 2025
H. atrobrunnea Dai 18821 OR287591 OR287657 Liu et al. 2025
H. attenuata He 28 JQ279526 JQ279633 He and Dai 2012
H. australis TAAM171362 KM017414 HE650990 Spirin et al. 2015; Parmasto et al. 2014
H. austrosinensis He 1009 OR287589 OR287660 Liu et al. 2025
H. austrosinensis He 7078 OR287590 OR287661 Liu et al. 2025
H. bannaensis CLZhao 35721 PQ847494 PQ847499 Deng et al. 2025
H. baishanzuensis Wei 11432 PV883105 – Zhu et al. 2026
H. baishanzuensis Wei 11406 PV883104 – Zhu et al. 2026
H. bambusicola He 4116 KY425674 KY425681 Nie et al. 2017
H. berteroi He 1488 KU975459 KU975498 He et al. 2017
H. biformisetosa He 1445 KF908247 KU975499 Yang and He 2014; He еt al. 2017
H. bispora He 4993 OR287532 OR287617 Liu et al. 2025
H. boddingii MEH-69996 MN030341 MN030347 Rossi et al. 2020
H. boidinii CBS 765.91 MH862335 – Vu et al. 2019
H. borbonica CBS 731.86 MH862026 MH873716 Vu et al. 2019
H. brunnea He 581 OR287582 OR287655 Liu et al. 2025
H. brunnea He 591 OR287581 OR287654 Liu et al. 2025
H. campylopora Cui 7393 JQ279513 JQ279629 He et al. 2017
H. cana He 1305 KF438169 KF438172 He and Li 2014
H. chimonobambusae HMZhou 710 PV475582 PV765708 Wang et al. 2026
H. cervinoidea CBS 736.86 MH862027 – Vu et al. 2019
H. cinerea He 1432 OR287602 OR287672 Liu et al. 2025
H. cinerea He 1483 OR287603 OR287673 Liu et al. 2025
H. cinereoalba CLZhao 10420 OR287566 – Liu et al. 2025
H. cinereoalba CLZhao 4252 OR287567 OR287642 Liu et al. 2025
H. cinereoalba He 6162 OR287559 OR287636 Liu et al. 2025
H. cinnamomea He 2074 KU975460 KU975500 Nie et al. 2017
H. cinnamomea He 755 JQ279548 JQ279658 He and Dai 2012
H. coffeana He 250 OR287533 – Liu et al. 2025
H. colliculosa Dai 16428 MF370596 MF370603 He et al. 2017
H. conchata LWZ 20140728-13 KX258960 – Pan and Zhou 2016
H. conifericola He 779 JQ279538 JQ279641 Liu et al. 2025
H. conifericola He 788 JQ279539 OR287666 Liu et al. 2025
H. contiformis He 1166 KU975461 KU975501 He et al. 2017
H. cruenta He 766 JQ279595 JQ279681 He et al. 2017
H. cruenta SFC20170811_12 MT044423 – Park et al. 2020
H. curtisii USDA FP-103876-Sp – HE650996 Parmasto et al. 2014
H. curtisii He 2061 KU975462 KU975502 Liu et al. 2025
H. cylindrospora Dai 14890 OR287562 – Liu et al. 2025
H. cylindrospora He 773 OR287563 OR287638 Liu et al. 2025
H. damicornis URM 84261 KC348466 – Du et al. 2021
H. denticulata CBS 780.91 MH862336 AF385155 Vu et al. 2019
H. dichotoma Dai 18649 OR287573 OR287647 Liu et al. 2025
H. dichotoma Dai 18779 OR287574 OR287648 Liu et al. 2025
H. dracaenicola Dai 22090 MW559797 MW559802 Du et al. 2021
H. dracaenicola Dai 22096 MW559798 MW559803 Du et al. 2021
H. duportii AFTOL-ID 666 DQ404386 AY635770 Matheny et al. 2006
H. epichlora He 524 OR287534 OR287618 Liu et al. 2025
H. epichlora He 525 JQ279549 JQ279659 He and Dai 2012
H. erastii He 1639 OR287569 OR287644 Liu et al. 2025
H. erastii He 1644 OR287570 – Liu et al. 2025
H. fissurata CLZhao 860 MG231569 – GenBank
H. fissurata He 1193 OR287535 OR287619 Liu et al. 2025
H. flava He 278 OR287586 – Liu et al. 2025
H. flava He 346 OR287587 OR287658 Liu et al. 2025
H. flava He 855 OR287588 OR287659 Liu et al. 2025
H. floridea He 529 JQ279598 KU975505 He and Dai 2012; Liu et al. 2025
H. fuliginosa He 1188 KU975465 KU975506 He et al. 2017
H. fuliginosa He 2077 KU975468 KU975510 Liu et al. 2025
H. fuliginosa UC2023222 KP814513 – GenBank
H. fulva He 640 JQ279565 JQ279648 He and Dai 2012
H. globispora He 911 OR287536 KU975508 Liu et al. 2025
H. granulata He 1472 KU975495 KU975547 Liu et al. 2025
H. hainanensis He 1509 OR287595 OR287664 Liu et al. 2025
H. hainanensis He 1534 OR287596 OR287665 Liu et al. 2025
H. huangshanensis He 432 JQ279533 JQ279671 He and Dai 2012
H. huangshanensis He 441 JQ279535 JQ279669 He and Dai 2012
H. hubeiensis Dai 17959 OR287611 OR287679 Liu et al. 2025
H. hubeiensis He 5084 OR287610 OR287678 Liu et al. 2025
H. hydnoides He 1309 KU975467 KU975509 Liu et al. 2025
H. iliensis Yuan 374 PP769636 – Liu et al. 2025
H. iliensis Yuan 386 PP769637 – Liu et al. 2025
H. innexa He 505 OR287537 OR287620 Liu et al. 2025
H. koeljalgii TAAM 159464 – HE651003 Parmasto et al. 2014
H. legeri He 1021 OR287538 OR287621 Liu et al. 2025
H. legeri He 960 KU975469 KU975511 He et al. 2017
H. leveillei CLZhao 10443 OR287547 – Liu et al. 2025
H. leveillei He 1437 OR287548 OR287628 Liu et al. 2025
H. leveillei He 6202 OR287549 OR287629 Liu et al. 2025
H. lictor He 20140723-2 KU975470 KU975512 Liu et al. 2025
H. liyiriana ARFR260 OR584222 OR569028 Tepin and Singh 2025
H. liyiriana ARFR260a PQ182942 – Tepin and Singh 2025
H. longispora He 217 JQ279537 KU975514 He and Dai 2012; Nie et al. 2017
H. luteomarginata He 1457 OR287614 OR287680 Liu et al. 2025
H. luteomarginata He 1849 OR287615 OR287681 Liu et al. 2025
H. macrochloae ARAN-Fungi 7079 MF990738 MF990743 Crous et al. 2017
H. macrospora Dai 488 KM017413 – Spirin et al. 2015
H. major CLZhao 5136 OR287558 – Liu et al. 2025
H. major He 1942 KU975489 KU975541 Liu et al. 2025
H. megaspora He 302 JQ279553 JQ279660 He and Dai 2012
H. megaspora He 328 JQ279554 – He and Dai 2012
H. membranacea He 1899 KU975491 KU975543 Liu et al. 2025
H. membranacea He 1903 KU975492 KU975544 Liu et al. 2025
H. microcycla CBS 311.39 MH856027 – Vu et al. 2019
H. microcycla Cui 8555 KT283049 JQ279640 Ariyawansa et al. 2015; He and Dai 2012
H. microcycla LWZ 20140719-11 KT283050 – Ariyawansa et al. 2015
H. micropora Cui 8057 JQ279518 – He and Dai 2012
H. minor He 933 JQ279555 JQ279654 He and Dai 2012
H. minuscula He 253 JQ279546 KU975516 He and Dai 2012; Nie et al. 2017
H. moniliformis He 1321 OR287556 OR287634 Liu et al. 2025
H. moniliformis He 916 OR287555 OR287633 Liu et al. 2025
H. montana He 1172 OR287606 OR287676 Liu et al. 2025
H. montana He 1380 OR287607 OR287677 Liu et al. 2025
H. mougeotii CBS 289.54 MH857337 MH868878 Vu et al. 2019
H. murina He 569 JQ716406 JQ716412 He and Li 2013a
H. muroiana He 172 JQ279541 – He and Dai 2012
H. muroiana He 405 JQ279542 KU975517 He and Dai 2012; Nie et al. 2017
H. nanospora CBS 924.96 MH862622 MH874244 Vu et al. 2019
H. nanospora He 475 JQ279531 JQ279672 He and Dai 2012
H. niveomarginata Dai 24611 PP769638 PP769648 Liu et al. 2025
H. niveomarginata He 1611 PP769639 – Liu et al. 2025
H. nothofagicola He 503 JQ279530 JQ279632 He and Dai 2012
H. ochromarginata Cui 8197 JQ279578 – He and Dai 2012
H. ochromarginata He 47 JQ279579 JQ279666 He and Dai 2012
H. odontoides Dai 11635 JQ279563 JQ279647 He and Dai 2012
H. odontoides Cui 24840 PZ458970 PZ466728 This study
H. orientalis He 4601 KY425677 KY425685 Nie et al. 2017
H. parmastoi He 367 JQ780061 – He and Li 2012
H. paucisetigera Cui 7845 JQ279560 JQ279644 He and Dai 2012
H. peroxydata JMB2056 KF371645 KF371648 Baltazar et al. 2014
H. piceae He 1200 OR287572 OR287646 Liu et al. 2025
H. piceae He 1203 OR287571 OR287645 Liu et al. 2025
H. pinnatifida He 2193 KU975472 KU975519 Liu et al. 2025
H. puerensis He 692 OR287612 – Liu et al. 2025
H. puerensis He 708 OR287613 – Liu et al. 2025
H. punctata HMZhou 16 PV475579 PV765709 Wang et al. 2026
H. quercicola He 373 KU975474 KU975521 He et al. 2017
H. quercicola He 377 OR287542 – Liu et al. 2025
H. ramicola He 1703 KU975493 KU975545 Liu et al. 2025
H. ramicola He 1897 KU975494 KU975546 Liu et al. 2025
H. ramicola He 5437 OR287583 – Liu et al. 2025
H. resupinata TU 100039 – HE650988 Parmasto et al. 2014
H. rhabarbarina He 280 JQ279574 KY425688 He and Dai 2012; Nie et al. 2017
H. rhabarbarina LY L527 – HE651007 Parmasto et al. 2014
H. rheicolor He 2192 KU975475 KU975522 Liu et al. 2025
H. rhododendricola He 392 JQ279576 – He and Dai 2012
H. rubiginosa Miettinen X3422 MK757158 – GenBank
H. rubiginosa S58 FJ820546 – Fröhlich-Nowoisky et al. 2009
H. rubrobrunnea He 1982 OR287579 KU975549 Liu et al. 2025
H. rubrobrunnea He 5315 OR287578 – Liu et al. 2025
H. rubrobrunnea He 5346 OR287580 OR287653 Liu et al. 2025
H. rufomarginata He 1489 KU975477 KU975524 He et al. 2017
H. sanguinaria HMZhou 995 PV475583 PV765710 Wang et al. 2026
H. senatoumbrina He 2437 KU975478 KU975525 Liu et al. 2025
H. senatoumbrina He 4940 OR457649 OR452887 Liu et al. 2025
H. separabilis BDC iNaturalist#192495191 PQ509880 – GenBank
H. separabilis He 1479 KU975479 KU975526 Liu et al. 2025
H. separabilis PDD 119545 OL709441 – GenBank
H. separata He 934 OR287544 OR287626 Liu et al. 2025
H. setipora Cui 6301 JQ279515 JQ279639 He and Dai 2012
H. setipora LE-BIN 3006 PQ361605 – GenBank
H. setipora SL1822 OR636140 – GenBank
H. setulohypha He 5570 OR287561 OR287637 Liu et al. 2025
H. setulohypha He 5602 OR287560 – Liu et al. 2025
H. sharmae 66088 MK588753 MK588836 Wang et al. 2019
H. sharmae CAL 1535 KY929017 KY929018 Wang et al. 2019
H. sichuanensis He 1340 OR287577 OR287641 Liu et al. 2025
H. sichuanensis He 1385 OR287585 OR287652 Liu et al. 2025
H. sinensis CLZhao 26040 OR659001 PP425893 Li et al. 2024
H. sinensis CLZhao 26652 PQ060540 – Li et al. 2024
H. spathulata He 685 JQ279591 KU975529 He and Dai 2012; He et al. 2017
H. sphaericola He 303 JQ279599 JQ279684 He and Dai 2012
H. sphaerospora He 715 JQ279594 KU975531 He and Dai 2012; He et al. 2017
H. stereoidea CLZhao 4321 OR287609 – Liu et al. 2025
H. stereoidea He 1661 KU975488 KU975540 Liu et al. 2025
H. stratura HHB-19391 MW740246 – Liu et al. 2025
H. subepichlora He 556 OR287554 OR287632 Liu et al. 2025
H. subepichlora He 559 OR287553 OR287631 Liu et al. 2025
H. subepichlora He 716 PP769642 – Liu et al. 2025
H. subferruginea He 1598 KU975481 – GenBank
H. subfissurata Dai 24358A PP769643 PP769650 Liu et al. 2025
H. subfissurata He 5573 PP769644 PP769651 Liu et al. 2025
H. subinnexa He 1007 OR287598 OR287668 Liu et al. 2025
H. subinnexa He 1213 OR287597 OR287667 Liu et al. 2025
H. subluteobadia He 10 JQ279568 OR287662 Liu et al. 2025
H. subluteobadia He 8 JQ279569 KU975515 He and Dai 2012; Nie et al. 2017
H. subporioides Cui 10163 KT283051 OR287627 Ariyawansa et al. 2015; Liu et al. 2025
H. subrhabarbarina He 737 OR287575 OR287649 Liu et al. 2025
H. subrhabarbarina He 860 OR287576 – Liu et al. 2025
H. subrhabarbarina LY L690 – HE650994 Parmasto et al. 2014
H. tasmanica He 449 JQ279582 JQ279663 He and Dai 2012
H. tasmanica He 455 JQ279583 JQ279664 He and Dai 2012
H. tongbiguanensis He 1552 KF908248 KU975532 Yang and He 2014; Nie et al. 2017
H. tropica He 661 JQ279588 – He and Dai 2012
H. unicolor He 468a JQ279551 JQ279662 He and Dai 2012
H. ustulata He 104 JQ780066 – He and Li 2013b
H. vaginata He 2558 KU975483 KU975535 Liu et al. 2025
H. vaginata He 2599 KU975484 KU975536 Liu et al. 2025
H. variabilis Dai 19779 OR287584 OR287651 Liu et al. 2025
H. variabilis He 1610 OR287592 OR287650 Liu et al. 2025
H. verruculosa Dai 17047 – MF370600 He et al. 2017
H. villosa He 1739 KU975485 KU975537 Liu et al. 2025
H. vitellina CLZhao 17846 OR287550 – Liu et al. 2025
H. vitellina He 5061 OR287551 OR287630 Liu et al. 2025
H. vitellina He 627 OR287552 – Liu et al. 2025
H. vivida He 2040 OR287601 OR287671 Liu et al. 2025
H. vivida He 4986 OR287600 OR287670 Liu et al. 2025
H. vivida He 5288 OR287599 OR287669 Liu et al. 2025
H. xerantica Cui 9209 JQ279519 JQ279635 He and Dai 2012
H. xerantica E060 LC520144 LC520190 Kobayashi et al. 2020
H. yaoshanensis CLZhao 20661 PP356585 – Xu et al. 2026
H. yaoshanensis CLZhao 20626 PP356584 PP785350 Xu et al. 2026
H. yunnanensis He 709 OR287546 – Liu et al. 2025
H. zhashuiensis Cui 24846 PZ458971 PZ466729 This study
Hyphoderma alboarachnum CLZhao 30488 PV470563 – Li et al. 2026
H. amoenum USO 286622 HE577030 – Tellería et al. 2012
H. asianum CLZhao 18091 OR141726 PP826262 Yang et al. 2025
H. assimile CBS:125852 MH863808 MH875272 Vu et al. 2019
H. australosetigerum MA:Fungi:92235 MN963764 – Boonmee et al. 2021
H. australosetigerum MA:Fungi:92240 MN963760 – Boonmee et al. 2021
H. bambusinum CLZhao 29903 PV469674 PV819428 Li et al. 2026
H. brevicystidiatum Cui 24844 PZ458969 PZ466727 This study
H. brevicystidiatum Cui 24841 PZ458968 PZ466726 This study
H. cinereofuscum CLZhao 30283 PQ492372 PQ511129 Li et al. 2025
H. cinereofuscum CLZhao 30341 PQ492371 PQ511128 Li et al. 2025
H. cremeoalbum CLZhao 17007 OM985716 OM985753 Duan et al. 2023
H. cremeoalbum NH 11538 (GB) DQ677492 DQ677492 Larsson 2007
H. crystallinum CLZhao 9338 MW917161 MW913414 Guan and Zhao 2021a
H. crystallinum CLZhao 9374 MW917162 MW913415 Guan and Zhao 2021a
H. definitum NH 12266 (GB) DQ677493 DQ677493 Larsson 2007
H. fissuratum CLZhao 6726 MT791330 MT791334 Ma et al. 2021
H. fissuratum CLZhao 6731 MT791331 – Ma et al. 2021
H. floccosum CLZhao 17129 MW301683 MW293733 Guan and Zhao 2021b
H. floccosum CLZhao 17215 MW301687 MW293735 Guan and Zhao 2021b
H. fragilissimum CLZhao 17245 PV147166 PV185848 Wijesinghe et al. 2025
H. fulgens CLZhao 30254 PV469670 PV819427 Li et al. 2026
H. fulgens CLZhao 36600 PV829546 PV810096 Li et al. 2026
H. fulgens CLZhao 37429 PV829544 PV810095 Li et al. 2026
H. fulgens CLZhao 39474 PV829547 PV810098 Li et al. 2026
H. grandineum CLZhao 30046 PV470561 PV819429 Li et al. 2026
H. grandineum CLZhao 43328 PV470562 PV819430 Li et al. 2026
H. granuliferum 5273 JN710545 JN710545 Yurchenko and Wu 2014a
H. guangdongense CLZhao 12657 PP235513 PP235514 Su et al. 2024
H. incrustatum KHL6685 – AY586668 Yurchenko and Wu 2014a
H. laceratum CLZhao 34242 PV829552 PV810101 Li et al. 2026
H. laceratum CLZhao 34672 PV829553 PV810103 Li et al. 2026
H. laceratum CLZhao 34958 PV829556 PV810106 Li et al. 2026
H. laceratum CLZhao 34961 PV829557 – Li et al. 2026
H. litschaueri FP-101740-Sp KP135295 KP135219 Floudas and Hibbett 2015
H. litschaueri NH 7603 (GB) DQ677496 DQ677496 Larsson 2007
H. macaronesicum MA:Fungi:90388 KC984327 – Tellería et al. 2012
H. macaronesicum TFC:Mic 15115 HE577011 – Yurchenko and Wu 2014b
H. marginatum CLZhao 3404 OM985717 OM985754 Duan et al. 2023
H. medioburiense FD-335 KP135298 KP135220 Floudas and Hibbett 2015
H. membranaceum CLZhao 5844 MW917167 MW913420 Guan and Zhao 2021a
H. membranaceum CLZhao 6971 MW917168 MW913421 Guan and Zhao 2021a
H. microporoides CLZhao 6857 MW917169 MW913422 Guan and Zhao 2021a
H. microporoides CLZhao 8695 MW917170 MW913423 Guan and Zhao 2021a
H. moniliforme Wu 0211-42 KC928282 – Yurchenko and Wu 2015
H. moniliforme Wu 0211-46 KC928284 – Yurchenko and Wu 2015
H. mopanshanense CLZhao 6449 OM985720 OM985759 Duan et al. 2023
H. mopanshanense CLZhao 6498 MT791329 MT791333 Ma et al. 2021
H. nemorale TNM F3931 KJ885183 KJ885184 Yurchenko and Wu 2015
H. nemorale Wu 9508-14 KC928280 KC928281 Yurchenko and Wu 2015
H. nitidum CLZhao 37195 PV470533 – Yuan et al. 2026
H. nitidum CLZhao 37232 PV470534 – Yuan et al. 2026
H. niveomarginatum CLZhao 25078 OR141728 OR506179 Yang et al. 2023
H. nudicephalum CLZhao 17839 OM985721 OM985760 Duan et al. 2023
H. nudicephalum Wu9307_29 AJ534269 – Nilsson et al. 2003
H. obtusiforme KHL11105 JN572910 – Yurchenko and Wu 2014b
H. obtusiforme KHL1464 JN572909 – Yurchenko and Wu 2014b
H. obtusum JS17804 – AY586670 Yurchenko and Wu 2014b
H. occidentale KHL 8477 (GB) DQ677499 DQ677499 Larsson 2007
H. paramacaronesicum MA:Fungi:87736 KC984399 – Martín et al. 2018
H. paramacaronesicum MA:Fungi:87737 KC984405 – Martín et al. 2018
H. pinicola Wu 0108-32 KJ885181 KJ885182 Yurchenko and Wu 2014b
H. pinicola Wu 0108-36 KC928278 KC928279 Yurchenko and Wu 2014b
H. prosopidis ARIZ HHB 8479 HE577029 – Yurchenko and Wu 2015
H. puerense CLZhao 9476 MW443045 – Guan et al. 2021
H. puerense CLZhao 9583 MW443046 MW443051 Guan et al. 2021
H. punctatum CLZhao 33051 PV469675 – Wijesinghe et al. 2025
H. qujingense CLZhao 26018 OR141729 PP826263 Yang et al. 2025
H. roseocremeum NH 10545 – AY586672 Yurchenko and Wu 2014a
H. sibiricum Kotiranta 22801 MF319069 MF318925 GenBank
H. setigerum FCUG 1200 AJ534273 – Nilsson et al. 2003
H. setigerum FCUG 1688 AJ534272 – Nilsson et al. 2003
H. sinense CLZhao 17811 MW301682 MW293732 Guan and Zhao 2021b
H. sinense CLZhao 7963 MW301679 MW293730 Guan and Zhao 2021b
H. sordidum CLZhao 27379 OR141731 – Yang et al. 2023
H. sordidum CLZhao 27390 OR141732 OR506180 Yang et al. 2023
H. subsetigerum HHB11620 GQ409521 – Yurchenko and Wu 2014a
H. subtestaceum CBS:125877 MH864081 MH875539 Yang et al. 2023
H. tenuissimum CLZhao 16210 MW443050 MW443055 Guan et al. 2021
H. tenuissimum CLZhao 7221 MW443049 MW443054 Guan et al. 2021
H. tongbiguanense CLZhao 39575 PV470532 – Yuan et al. 2026
H. tongbiguanense CLZhao 39574 PV470531 – Yuan et al. 2026
H. transiens NH 12304 DQ677504 DQ677504 Larsson 2007
H. tropicum CLZhao 17308 OM985727 OM985768 Duan et al. 2023
H. variolosum CBS:734.91 MH862320 MH873992 Vu et al. 2019
H. variolosum CBS:735.91 MH862321 MH873993 Vu et al. 2019
H. weishanense CLZhao 22403 OR141727 OR506181 Yang et al. 2023
H. yingjiangense CLZhao 37815 PV470536 – Wijesinghe et al. 2025
H. yingjiangense CLZhao 36602 PV470535 – Wijesinghe et al. 2025
H. yunnanense CLZhao 8845 OM985728 OM985769 Duan et al. 2023
Steccherinum aff. nitidum FP-105195-Sp KP135323 KP135227 Floudas and Hibbett 2015
S. amapaense M245 KY977406 KY977405 Hyde et al. 2017
S. austrosinense Dai 17540 MN871755 MN877768 Du et al. 2020
S. austrosinense Dai 17679 MN871756 MN877769 Du et al. 2020
S. autumnale Spirin 2957 JN710549 JN710549 Liu and Dai 2021
S. bononiae AG 1615 PV434851 PV434851 Westphalen et al. 2026
S. bononiae MCW 547/17 PV434850 PV434850 Westphalen et al. 2026
S. bononiae MCW 557/17 PV434854 PV434854 Westphalen et al. 2026
S. bononiae MCW 726/22 PV434852 PV434852 Westphalen et al. 2026
S. bononiae MV446 PV434855 – Westphalen et al. 2026
S. bononiae NR71 PV434853 PV434853 Westphalen et al. 2026
S. bourdotii HR99893 MT849311 – Westphalen et al. 2021
S. bourdotii MT 10/19 MT849312 – Westphalen et al. 2021
S. bourdotii Saarenoksa 10195 JN710584 JN710584 Miettinen et al. 2012
S. ciliolatum Ryvarden 47033 JN710585 JN710585 Miettinen et al. 2012
S. collabens KHL 11848 JN710552 JN710552 Liu and Dai 2021
S. elegantissimum MCW 633/18 PV434856 PV434856 Westphalen et al. 2026
S. elegantissimum MCW 720/21 PV434857 PV434857 Westphalen et al. 2026
S. elegantissimum MCW 721/21 PV434858 – Westphalen et al. 2026
S. farinaceum CLZhao 37844 PV661644 PV661646 Zhang et al. 2025
S. farinaceum CLZhao 39423 PV661645 PV661647 Zhang et al. 2025
S. filiferum BLS M-5230 OP279612 – Yurchenko et al. 2023
S. fimbriatellum Miettinen 2091 JN710555 JN710555 Miettinen et al. 2012
S. fissurutum CLZhao 21803 OP799385 OP799397 Dong et al. 2023
S. fissurutum CLZhao 21841 OP799388 OP799400 Dong et al. 2023
S. formosanum Dai 19345 MN871759 MN877772 Du et al. 2020
S. formosanum TFRI 652 EU232184 EU232268 Chou et al. (unpub.)
S. fragile Dai 19972 MW364629 MW364627 Liu and Dai 2021
S. fragile Dai 20479 MW364628 MW364626 Liu and Dai 2021
S. fragrans MG505 PP838792 PP838793 Crous et al. 2024
S. hirsutum CLZhao 4222 MW290040 MW290054 Dong et al. 2022
S. hirsutum CLZhao 4523 MW290041 MW290055 Dong et al. 2022
S. incrustans Dai 19442 ON182084 ON182087 Liu et al. 2023a
S. juniperi Dai 23931 OP956077 OP956031 Liu et al. 2023a
S. lacerum Niemelä 8246 JN710557 JN710557 Miettinen et al. 2012
S. laeticolor Fp-102480-Sp KY948823 KY948868.1 Justo et al. 2017
S. larssonii MCW 593/17 MT849306 MT849306 Westphalen et al. 2021
S. larssonii MCW 594/17 MT849307 MT849307 Westphalen et al. 2021
S. laxum KHL 12268 JN710577 JN710577 Miettinen et al. 2012
S. lincangense CLZhao 24988 OR096196 OR461455 Dong et al. 2024
S. longiaculeiferum CLZhao 26243 OR096195 – Dong et al. 2024
S. longiaculeiferum CLZhao 26290 OR167202 – Dong et al. 2024
S. meridionalis MR 10466 KY174994 KY174994 Westphalen et al. 2018
S. meridionalis MR 284 KY174992 KY174992 Westphalen et al. 2018
S. molle MCW 568/17 PV434863 PV434863 Westphalen et al. 2026
S. molle MCW 641/18 PV434864 PV434864 Westphalen et al. 2026
S. molle MCW 661/18 PV434865 PV434865 Westphalen et al. 2026
S. molle MCW 687/19 PV434866 PV434866 Westphalen et al. 2026
S. molle MCW 734/22 PV434867 – Westphalen et al. 2026
S. molle MCW 739/23 PV434868 – Westphalen et al. 2026
S. molle MV450 PV434869 – Westphalen et al. 2026
S. nandinae Dai 21107 MN833677 MN833679 Du et al. 2020
S. neonitidum MCW 371/12 KY174990 KY174990 Westphalen et al. 2018
S. neonitidum RP 79 KY174991 KY174991 Westphalen et al. 2018
S. nitidum KHL 11903 JN710560 JN710560 Westphalen et al. 2018
S. nitidum MT 33/12 KY174989 KY174989 Westphalen et al. 2018
S. ochraceum 2060 JN710589 JN710589 Liu and Dai 2021
S. ochraceum KHL11902 JN710590 JN710590 Westphalen et al. 2021
S. perparvulum 524/17 PV434837 PV434837 Westphalen et al. 2026
S. perparvulum 543/17 PV434839 PV434839 Westphalen et al. 2026
S. perparvulum 592/17 PV434847 PV434847 Westphalen et al. 2026
S. perparvulum 659/18 PV434845 PV434845 Westphalen et al. 2026
S. perparvulum 692/19 PV434838 PV434838 Westphalen et al. 2026
S. perparvulum 710/20 PV434841 – Westphalen et al. 2026
S. perparvulum 742/23 PV434846 – Westphalen et al. 2026
S. perparvulum 744/23 PV434843 – Westphalen et al. 2026
S. perparvulum MV728 PV434840 – Westphalen et al. 2026
S. perparvulum MV815 PV434844 PV434844 Westphalen et al. 2026
S. perparvulum NR186 PV434842 – Westphalen et al. 2026
S. polycystidiferum MCW 419/12 KY174995 KY174995 Westphalen et al. 2018
S. polycystidiferum RP 140 KY174996 KY174996 Westphalen et al. 2018
S. pseudozilingianum Kulju 1004 JN710561 JN710561 Miettinen et al. 2012
S. pudorinum KR-M-0093535 OR381577 — Popa et al. 2024
S. puerense CLZhao 3122 MW682341 — Wu et al. 2021a
S. puerense CLZhao 3644 MW682342 MW682338 Wu et al. 2021a
S. puerense Miettinen 13705 JN710592 JN710592 Miettinen et al. 2012
S. resinaceum MCW 540/17 PV434870 PV434870 Westphalen et al. 2026
S. resinaceum MCW 551/17 PV434871 PV434871 Westphalen et al. 2026
S. resinaceum MCW 665/19 PV434872 PV434872 Westphalen et al. 2026
S. resinaceum MCW 679/19 PV434873 PV434873 Westphalen et al. 2026
S. robustius G1195 JN710591 JN710591 Cao et al. 2021
S. rubigimaculatum CLZhao 10638 MW682344 MW682340 Wu et al. 2021a
S. rubigimaculatum CLZhao 4069 MW682343 MW682339 Wu et al. 2021a
S. shangzhouense Cui 24842 PZ458972 PZ466730 This study
S. shangzhouense Cui 24843 PZ458973 PZ466731 This study
S. shangzhouense Cui 24845 PZ458974 PZ466732 This study
S. sp. FD-26 KP135322 KP135289 Floudas and Hibbett 2015
S. sp. 2 Miettinen 9300 JN710593 JN710593 Miettinen et al. 2012
S. sp. 3 Miettinen 14391 JN710594 JN710594 Miettinen et al. 2012
S. sp. 4 Miettinen 13755 JN710596 JN710596 Miettinen et al. 2012
S. straminellum KHL 13849 JN710597 JN710597 Miettinen et al. 2012
S. subcollabens Dai 19344 MN871758 MN877771 Liu and Dai 2021
S. subcollabens Dai 19345 MN871759 MN877772 Liu and Dai 2021
S. subochraceum 730/22 PV434859 – Westphalen et al. 2026
S. subochraceum 746/23 PV434861 PV434861 Westphalen et al. 2026
S. subochraceum 748/23 PV434860 PV434860 Westphalen et al. 2026
S. subochraceum 761/24 PV434862 PV434862 Westphalen et al. 2026
S. subtropicum CLZhao 11059 OP799390 OP799377 Dong et al. 2023
S. subtropicum CLZhao 16901 OP799391 – Dong et al. 2023
S. tenue 5356 JN710598 — Miettinen et al. 2012
S. tenue KHL 12316 JN710598 JN710598 Miettinen et al. 2012
S. tenuispinum LE231603 KM411452 KM411469 Zmitrovich and Kovalenko 2016
S. tenuispinum Miettinen 8065 JN710599 JN710599 Miettinen et al. 2012
S. tenuispinum Spirin 2116 JN710600 JN710600 Miettinen et al. 2012
S. undigerum MCW 426/13 KY174986 KY174986 Westphalen et al. 2018
S. undigerum MCW 436/13 KY174988 KY174988 Westphalen et al. 2018
S. undigerum MCW 472/13 KY174987 KY174987 Westphalen et al. 2018
S. undulatum MCW 743/23 PV434848 PV434848 Westphalen et al. 2026
S. undulatum MCW 760/24 PV434849 PV434849 Westphalen et al. 2026
S. weishanense CLZhao 24911 OR096207 OR461456 Dong et al. 2024
S. wumengshanense CLZhao 23586 OR658995 OR999392 Wang et al. 2024
S. xanthum CLZhao 5030 MW204588 MW204577 Wu et al. 2021b
S. xanthum CLZhao 5032 MW204589 MW204578 Wu et al. 2021b
S. yunnanense CLZhao 1445 MW290042 MW290056 Dong et al. 2022
S. yunnanense CLZhao 2822 MW290043 MW290057 Dong et al. 2022

Phylogenetic analysis

Phylogenetic analyses were conducted separately for Hymenochaete, Hyphoderma and Steccherinum based on the combined ITS+nLSU datasets. Reference sequences were retrieved from GenBank (Table 1). In the phylogenetic analyses, Hydnoporia olivacea (Schwein.) Teixeira was selected as the outgroup for Hymenochaete (Miettinen et al. 2019), Diplomitoporus crustulinus (Bres.) Domański was selected as the outgroup for Hyphoderma (Justo et al. 2017), and Cabalodontia delicata Westph. & Motato-Vásq. was selected as the outgroup for Steccherinum (Westphalen et al. 2021).

Sequence alignments were generated using MAFFT v.7 (Katoh et al. 2019) and manually adjusted in BioEdit v.7.0.9 (Hall 1999). Gaps were treated as missing data. The ITS and nLSU alignments were concatenated using Mesquite v.3.2 (Maddison and Maddison 2017).

Maximum Likelihood (ML) analysis was performed using RAxML-HPC2 v.8.2.3 (Stamatakis 2014). The best-scoring ML tree was searched under the GTRGAMMA model, and branch support was assessed with 1000 rapid bootstrap replicates. Bayesian Inference (BI) was performed using MrBayes v.3.2.6 (Ronquist et al. 2012). The substitution model was set to GTR + I + G, with six substitution types and gamma-distributed rate variation across sites with a proportion of invariable sites (nst = 6, rates = invgamma). Base frequencies were assigned a Dirichlet prior. Five Markov Chain Monte Carlo (MCMC) chains were run for 50 million generations, with trees sampled every 100 generations. The temperature parameter was set to 0.2, and the analysis was allowed to stop when the average standard deviation of split frequencies fell below 0.01 (Ronquist and Huelsenbeck 2003). The first 25% of sampled trees were discarded as burn-in, and the remaining trees were used to generate a 50% majority-rule consensus tree and calculate Bayesian posterior probabilities (BPP).

Phylogenetic trees were visualized using FigTree v.1.4.4 (Rambaut 2018). The ML topology is presented, with ML bootstrap values and Bayesian posterior probabilities indicated at the nodes. Only ML-BS ≥ 75% and BPP ≥ 0.90 are shown. Nodes with ML-BS ≥ 75% and BPP ≥ 0.95 were considered well supported.

Results

Phylogenetic analyses

To clarify the phylogenetic positions of the newly collected specimens, three independent phylogenetic analyses were conducted for Hymenochaete, Hyphoderma and Steccherinum based on combined ITS+nLSU datasets.

The combined ITS+nLSU dataset for Hymenochaete comprised sequences from 225 fungal samples representing 139 species of Hymenochaete and the outgroup species Hydnoporia olivacea. The aligned matrix was 2622 characters in length, including 1115 constant characters, 750 parsimony-uninformative variable characters and 757 parsimony-informative characters. Phylogeny results indicated that specimen Cui 24846 formed a distinct lineage (100% ML, 1.00 BPP; Fig. 1), and exhibited a close phylogenetic affinity to Hymenochaete tasmanica Massee (Fig. 1).

Figure 1.

Figure 1.

Maximum Likelihood tree illustrating the phylogenetic relationships of Hymenochaete inferred from the combined ITS+nLSU dataset. Hydnoporia olivacea was used as the outgroup. Branches are labelled with maximum likelihood bootstrap values and Bayesian posterior probabilities. Only maximum likelihood bootstrap values ≥ 75% and Bayesian posterior probabilities ≥ 0.90 are shown. The new taxon is shown in bold. The type species is indicated by white five-pointed star, and type specimens are marked by *.

The combined two-gene dataset (ITS+nLSU) for Hyphoderma comprised sequences from 91 fungal samples, representing 56 species of Hyphoderma and the outgroup species Diplomitoporus crustulinus. The aligned matrix was 2072 characters in length, of which 1356 were constant, 164 were parsimony-uninformative variable characters and 552 were parsimony-informative. Phylogenetic analyses revealed that specimens Cui 24841 and Cui 24844 formed a well-supported independent clade (98% ML, 0.99 BPP; Fig. 2) and showed a close phylogenetic relationship with Hyphoderma setigerum (Fig. 2). Independent BLAST searches of the ITS and nLSU sequences of Cui 24841 and Cui 24844 were conducted against the NCBI nucleotide database. After exclusion of the query sequence itself, neither gene region yielded close matches to any validly named Hyphoderma species. For ITS sequences, the 20 highest-scoring non-self-hits were assigned to H. subsetigerum Sheng H. Wu, with sequence identities ranging from 97.15% to 98.83%. For the nLSU sequences, the closest non-self matches included H. floccosum C.L. Zhao & Q.X. Guan, H. guangdongense J.Q. Su & C.L. Zhao, H. nudicephalum Gilb. & M. Blackw., H. pinicola Yurchenko & Sheng H. Wu, H. setigerum, H. sordidum Yang Yang & C.L. Zhao, H. subsetigerum and H. tenuissimum C.L. Zhao & Q.X. Guan, with sequence identities ranging from 97.88% to 99.88% (Table 2).

Figure 2.

Figure 2.

Maximum Likelihood tree illustrating the phylogenetic relationships of Hyphoderma inferred from the combined ITS+nLSU dataset. Diplomitoporus crustulinus was used as the outgroup. Branches are labelled with maximum likelihood bootstrap values and Bayesian posterior probabilities. Only maximum likelihood bootstrap values ≥ 75% and Bayesian posterior probabilities ≥ 0.90 are shown. The new taxon is shown in bold. The type species is indicated by white five-pointed star, and type specimens are marked by *.

Table 2.

Independent NCBI BLAST comparisons for ITS and nLSU sequences of Hyphoderma brevicystidiatum (vouchers Cui 24841 and Cui 24844).

Query voucher Marker Query accession Query length (bp) Taxa represented among the top 20 non-self-hits Query cover (%) Perident. (%)
Cui 24841 ITS PZ458968.1 626 Hyphoderma subsetigerum (20/20) 97–100 97.78–98.53
Cui 24841 nLSU PZ466726.1 1395 H. floccosum (4) 95–98 97.88–99.85
H. subsetigerum (2)
H. nudicephalum (5)
H. guangdongense (1)
H. sordidum (1)
H. setigerum (4)
H. tenuissimum (2)
Hyphoderma sp. (1)
Cui 24844 ITS PZ458969.1 635 H. subsetigerum (20/20) 94–99 97.15–98.83
Cui 24844 nLSU PZ466727.1 911 H. pinicola (4) 93–99 98.00–99.88
H. setigerum (1)
H. subsetigerum (4)
H. sp. (1)
H. floccosum (4)
H. guangdongense (1)
H. nudicephalum (5)

The combined ITS+nLSU dataset for Steccherinum included sequences from 116 fungal samples representing 56 species of Steccherinum and the outgroup species Cabalodontia delicata. The aligned matrix was 2016 characters in length, including 1590 constant characters, 125 parsimony-uninformative variable characters and 301 parsimony-informative characters. In the phylogenetic tree, specimens Cui 24842, Cui 24843 and Cui 24845 clustered together and formed an independent lineage within Steccherinum (91% ML, 0.99 BPP; Fig. 3), and exhibited a close phylogenetic affinity to Steccherinum fissurutum J.H. Dong & C.L. Zhao and S. fragrans Ghob.-Nejh. & Langer (Fig. 3).

Figure 3.

Figure 3.

Maximum Likelihood tree illustrating the phylogenetic relationships of Steccherinum inferred from the combined ITS+nLSU dataset. Cabalodontia delicata was used as the outgroup. Branches are labelled with maximum likelihood bootstrap values and Bayesian posterior probabilities. Only maximum likelihood bootstrap values ≥ 75% and Bayesian posterior probabilities ≥ 0.90 are shown. The new taxon is shown in bold. The type species is indicated by white five-pointed star, and type specimens are marked by *.

Taxonomy

Hymenochaete zhashuiensis

L.L. Wan, L.S. Zhang, Shun Liu & B.K. Cui sp. nov.

138FEDB1-9E05-5DA6-9D80-0CE99C98841E

864506

Figs 4, 5

Figure 4.

Figure 4.

Basidiomata of Hymenochaete zhashuiensis (Holotype, Cui 24846). Scale bar: 5 cm.

Figure 5.

Figure 5.

Microscopic structures of Hymenochaete zhashuiensis (drawn from Cui 24846). A. Basidiospores; B. Dendrohyphidia; C. Basidia and basidioles; D. Setae; E. Hymenium. Scale bars: 10 µm (A–E).

Diagnosis.

Hymenochaete zhashuiensis is characterized by resupinate to effused-reflexed basidiomata, a reddish brown to rust-brown, uneven to tuberculate and locally cracked hymenial surface, a monomitic hyphal system with simple-septate, pale yellowish, slightly thick-walled generative hyphae, abundant dark brown, thick-walled hymenial setae (66–89 × 7.8–11 µm), branched dendrohyphidia, and ellipsoid to broadly ellipsoid basidiospores (4–5.6 × 2.8–3.5 µm).

Type.

• China; Shaanxi Province, Zhashui County; 33.807836°N, 108.951155°E; elevation 1184.0 m; on living angiosperm tree; 27 September 2025; Cui 24846 (holotype, BJFC).

Etymology.

“zhashuiensis” (Lat.): referring to Zhashui County, where the type specimen was collected.

Description.

Fruiting body. Basidiomata annual, resupinate to effused-reflexed, broadly attached to the substrate, coriaceous to woody when fresh, becoming hard upon drying, 15–20 cm long, 8–12 cm wide, 0.5–3 mm thick. Reflexed part dark brown to blackish brown, rough, uneven, partly covered by mosses and algae in the field. Hymenial surface reddish brown to rust-brown when fresh, becoming dark reddish brown upon drying, uneven, tuberculate to rugose, locally cracked; margin irregular, thinning out, concolorous to slightly paler than the hymenial surface. Tomentum present; cortex absent; hyphal layer present; setal layer well developed.

Hyphal structure. Hyphal system monomitic; generative hyphae simple-septate, IKI–, CB–; tissues darkening in KOH.

Subiculum. Generative hyphae pale yellowish, slightly thick-walled, frequently branched, interwoven, simple-septate, 1.7–2.3 µm in diam.

Hymenium. Hymenial setae abundant, dark brown, thick-walled, fusiform to subulate, straight to slightly curved, with acute apices, projecting from or embedded in the hymenium, 66–89 × 7.8–11.0 µm. Embedded hymenial setae present; true setal hyphae not observed. Cystidia not observed. Dendrohyphidia present, branched, yellowish brown to brown, thin- to slightly thick-walled. Basidia narrowly clavate to subclavate, with four sterigmata and a simple septum at the base, 14.3–17.2 × 2.9–4.1 µm. Basidioles similar to basidia in shape, but smaller.

Spores. Basidiospores ellipsoid to broadly ellipsoid, hyaline, thin-walled, smooth, IKI–, CB–, 4–5.6 × 2.8–3.5 µm, L = 4.71 µm, W = 3.19 µm, Q = 1.28–1.84 (n = 30/1).

Type of rot.

White rot.

Hyphoderma brevicystidiatum

L.L. Wan, L.S. Zhang, Shun Liu & B.K. Cui sp. nov.

1D1BDD15-6A80-58A4-9688-8F8E229FABA5

864508

Figs 6, 7

Figure 6.

Figure 6.

Basidiomata of Hyphoderma brevicystidiatum (Holotype, Cui 24844). Scale bar: 2 cm.

Figure 7.

Figure 7.

Microscopic structures of Hyphoderma brevicystidiatum (Holotype Cui 24844). A. Basidiospores; B. Basidia and basidioles; C. Cystidia; D. Part of a vertical section of the hymenium. Scale bars: 10 µm (A–D).

Diagnosis.

Hyphoderma brevicystidiatum is characterized by resupinate, white to cream basidiomata, a smooth to slightly grandinioid hymenial surface, a monomitic hyphal system with clamped generative hyphae, cylindrical to subcylindrical cystidia that are mostly smooth and occasionally bear a few crystalline deposits, subclavate to clavate basidia, and ellipsoid to subcylindrical basidiospores (6.2–10 × 2.7–4.2 µm).

Type.

• China; Shaanxi Province, Shangzhou District; 33.721046°N, 110.150691°E; elevation 701.6 m; on fallen branch of Pinus; 22 August 2025; Cui 24844 (holotype, BJFC).

Etymology.

“brevicystidiatum” (Lat.): referring to species having the shorter cystidia.

Description.

Fruiting body. Basidiomata annual, resupinate, closely adnate to the substrate, membranaceous, soft when fresh, becoming brittle upon drying, 15–23 cm long, 2.5–4 cm wide, 0.3–1 mm thick. Hymenial surface smooth to slightly grandinioid, white to cream when fresh, becoming pale gray upon drying, frequently developing slight cracks.

Hyphal structure. Hyphal system monomitic; generative hyphae with clamp connections, IKI–, CB–; tissues unchanged in KOH.

Subiculum. Generative hyphae hyaline, thin-walled, frequently branched, interwoven, 1.8–2.5 µm in diameter.

Hymenium. Cystidia present, cylindrical to subcylindrical, hyaline, thin- to slightly thick-walled, mostly smooth, occasionally with a few crystalline deposits, 24.2–30.5 × 4.2–7.4 µm. Basidia subclavate to clavate, bearing four sterigmata and a basal clamp connection, 12.8–24.2 × 4.8–6.2 µm. Basidioles similar to basidia in shape, but smaller.

Spores. Basidiospores ellipsoid to subcylindrical, hyaline, thin-walled, smooth, IKI–, CB–, (5.2–)6.2–10 (–11.1) × (1.9–)2.7–4.2(–4.4) µm, L = 7.91 µm, W = 3.45 µm, Q = 1.85–2.79 (n = 60/2).

Additional specimen (paratype) examined.

• China; Shaanxi Province, Zhashui County; 33.804981°N, 108.931165°E; elevation 1217.7 m; on fallen angiosperm branch; 16 August 2025; Cui 24841 (BJFC).

Type of rot.

White rot.

Steccherinum shangzhouense

L.L. Wan, L.S. Zhang, Shun Liu & B.K. Cui sp. nov.

D0019657-FC6F-5F1E-8780-20DF0EF2A36D

864510

Figs 8, 9

Figure 8.

Figure 8.

Basidiomata of Steccherinum shangzhouense (Holotype, Cui 24845). Scale bar: 2 cm.

Figure 9.

Figure 9.

Microscopic structures of Steccherinum shangzhouense (drawn from the holotype). A. Basidiospores; B. Basidia and basidioles; C. A section of hymenium. Scale bars: 10 µm (A–C).

Diagnosis.

Steccherinum shangzhouense is characterized by resupinate and membranaceous basidiomata, white to cream, tuberculate to slightly grandinioid and distinctly cracked hymenial surface, monomitic hyphal system with simple-septate generative hyphae, absence of cystidia and skeletocystidia, clavate basidia, and broadly ellipsoid to oblong ellipsoid basidiospores (5.2–7.9 × 2.5–5 µm).

Type.

• China; Shaanxi Province, Shangluo, Shangzhou District; 34.020372°N, 109.983861°E; elevation 852 m; on fallen angiosperm branch; 23 August 2025; Cui 24845 (holotype, BJFC).

Etymology.

The epithet “shangzhouense” (Lat.): referring to Shangzhou District, Shaanxi Province, where the type specimen was collected.

Description.

Fruiting body. Basidiomata annual, resupinate, closely adnate to the substrate, membranaceous, soft when fresh, becoming brittle upon drying, white to cream when fresh, turning pale gray to grayish upon drying, frequently developing conspicuous cracks, 13–18 cm long, 1–2 cm wide, and up to 120 µm thick. Hymenial surface tuberculate to slightly grandinioid, composed of irregular granules, white to cream when fresh, becoming pale gray upon drying, distinctly cracked.

Hyphal structure. Hyphal system monomitic; generative hyphae simple-septate; hyphae IKI–, CB–; tissues unchanged in KOH.

Subiculum. Generative hyphae hyaline, thin-walled, frequently branched, interwoven, 0.8–1.2 µm in diameter.

Hymenium. Cystidia and skeletocystidia not observed. Basidia clavate, with four sterigmata and a simple septum at the base, 15.5–20.1 × 3.5–4.8 µm. Basidioles similar to basidia in shape, but smaller.

Spores. Basidiospores broadly ellipsoid to oblong ellipsoid, hyaline, thin-walled, smooth, occasionally with one oil drop, IKI–, CB–, (4.3–)5.2–7.9(–8.4) × (1.9–)2.5–5(–5.5) µm, L = 6.44 µm, W = 3.6 µm, Q = 1.51–2.09 (n = 90/3).

Additional specimens (paratypes) examined.

• China; Shaanxi Province, Shangzhou District; 33.761057°N, 109.820994°E; elevation 1187 m; on fallen angiosperm trunk; 22 August 2025; Cui 24843 (BJFC). • China; Shaanxi Province, Shangzhou District; 33.760510°N, 109.821477°E; elevation 1206 m; on fallen angiosperm branch; 22 August 2025; Cui 24842 (BJFC).

Type of rot.

White rot.

Discussion

In recent years, numerous new species of wood-inhabiting fungi reported from China have been concentrated in South and Southwest regions (Dai et al. 2004a, 2007, 2009; Yuan and Dai 2008; Zhao et al. 2015; Wu et al. 2016, 2021a, 2021b; Zhang et al. 2023; Zhou et al. 2023), whereas research on the species diversity of corticioid fungi in the Qinling Mountains remains limited. In this study, three new species belonging to the genera Hymenochaete, Hyphoderma and Steccherinum were identified in Zhashui County and Shangzhou District in the eastern Qinling Mountains.

Phylogenetic analysis of Hymenochaete revealed that Cui 24846 clustered with H. tasmanica, forming a well-supported clade (100% ML, 1.00 BPP; Fig. 1). Morphologically, H. tasmanica is similar to H. zhashuiensis by sharing a monomitic hyphal system with simple-septate generative hyphae. However, H. tasmanica differs by having thinner basidiomata (200–800 µm thick), a smooth to slightly tuberculate and usually uncracked hymenial surface, the presence of a cortex, reddish brown setae, larger basidia (19–25 × 3.5–4.5 µm), and slightly narrower basidiospores (4–5.5 × 2.5–3 µm; He and Li 2012). Hymenochaete ochromarginata P.H.B. Talbot and H. rubiginosa (Dicks.) Lév. also produce effused-reflexed basidiomata and abundant hymenial setae, but H. ochromarginata differs from H. zhashuiensis by its yellowish brown to brown pileus with concentrically zonate and sulcate, yellow to ochraceous hymenial margin, shorter hymenial setae (20–56 × 5–10 µm), smaller basidiospores (2–4 × 1.6–2 µm; Vinjusha and Kumar 2024); H. rubiginosa differs from H. zhashuiensis by its perennial, woody hard basidiomata with distinct cortex, subdimitic hyphal system with sclerefied hyphae, usually uncracked hymenium, and narrower, elongate-ellipsoid basidiospores (3.5–5.5 × 2–3 µm; Corfixen and Parmasto 2017). Hymenochaete erastii S.H. He et al. and H. zhashuiensis share the presence of a hyphal layer and setal layer and the absence of a cortex, but the former differs by its strictly resupinate, closely adnate basidiomata, grayish violet to clay buff hymenial surface, crystalline masses in the setal layer, smaller setae (30–46 × 3–7 µm), and narrower oblong-ellipsoid to cylindrical basidiospores (4–5 × 1.7–2 µm; Liu et al. 2025).

In the phylogenetic analyses based on combined ITS and nLSU sequences, two specimens of Hyphoderma brevicystidiatum (Cui 24841 and Cui 24844) formed an independent lineage with strong nodal support (98% ML, 0.99 BPP; Fig. 2). This species clustered together with H. alboarachnum Wen Li & C.L. Zhao, H. floccosum, H. guangdongense, H. paramacaronesicum Tellería et al., H. pinicola and H. setigerum, constituting a highly supported clade (Fig. 2). Morphologically, all these species possess resupinate basidiomata and a monomitic hyphal system bearing clamp connections. Nevertheless, compared to Hyphoderma brevicystidiatum, H. alboarachnum differs by its arachnoid hymenial surface, absence of cystidia and cystidioles, cylindrical basidia (23.5–30 × 3.5–4.5 µm), and smaller basidiospores (5–6 × 2–3 µm; Li et al. 2026); H. floccosum is distinguished by its farinaceous hymenial surface, two types of cystidia, including thick-walled and abundantly encrusted cystidia (60–161 × 5.5–10 µm) and tubular, thin-walled cystidia (37.5–100 × 4–8.5 µm; Guan and Zhao 2021b); H. guangdongense differs by its farinaceous basidiomata, thicker generative hyphae (3.5–6.0 µm), two types of cystidia, including septate cystidia (93–144 × 8–10.4 µm) and tubular cystidia (55–63 × 6.5–10.3 µm), and cylindrical basidiospores (7.4–9 × 3.2–4 µm; Su et al. 2024); H. paramacaronesicum is readily distinguished by its yellowish white to pale orange-yellow basidiomata, cylindrical leptocystidia with several constrictions (70–124 × 8–13 µm), larger basidia (40–48 × 6–9 µm), and larger, ellipsoid basidiospores (12–15 × 5.5–7 µm; Martín et al. 2018); H. pinicola differs by its chalky white, minutely warted to porulose hymenial surface, larger basidia (25–28 × 5–6.5 µm), longer septocystidia (65–180 µm long), and longer, cylindrical to allantoid basidiospores (13–16 × 4–4.5 µm), as well as its occurrence on dead wood of Pinus in Yunnan Province (Yurchenko and Wu 2014a); H. setigerum differs in thick-walled and encrusted cystidia (120–180 × 7.5–10 µm), larger basidia (25–40 × 6.5–7 µm), and subcylindrical basidiospores (9.5–11.5 × 3.5–4.5 µm; Nilsson et al. 2003). Furthermore, additional comparisons were made with four members or closely related segregates of the Hyphoderma setigerum complex. Hyphoderma subsetigerum differs from H. brevicystidiatum in having a grandinioid hymenophore with a whitish to ivory-yellow hymenial surface, thick-walled generative hyphae, narrower basidia (20–30 × 4.5–5.5 µm), and smaller basidiospores (6–8 × 2.8–3.2 µm; Wu 1997); H. bisetigerum is distinguished by predominantly two-sterigmate basidia and thick-walled, heavily encrusted septocystidia, whereas H. brevicystidiatum has four-sterigmate basidia and short, mostly smooth cystidia with only occasional crystalline deposits (Boidin and Gilles 2003); H. nudicephalum differs in its farinaceous to odontioid hymenial surface, thick-walled generative hyphae, and conspicuous capitate cystidia with swollen, non-encrusted apices (Gilbertson and Blackwell 1988); H. tenuissimum is separated from the new species by its tuberculate to minutely grandinioid, slightly buff hymenial surface, thick-walled generative hyphae, and much longer cylindrical cystidia with 4–12 clamped septa and abundant encrustations (50–220 × 6.5–13 µm), and larger, cylindrical basidiospores (7–10.5 × 3–4.5 µm; Guan et al. 2021).

In the phylogenetic tree of the genus Steccherinum, specimens of S. shangzhouense (Cui 24842, Cui 24843 and Cui 24845) formed a distinct independent evolutionary lineage, and clustered with S. fissurutum and S. fragrans to constitute a clade with high statistical support (91% ML, 0.99 BPP; Fig. 3). Nevertheless, these species exhibit conspicuous morphological discrepancies from S. shangzhouense. Specifically, S. fissurutum differs from S. shangzhouense by its subceraceous basidiomata, clamped generative hyphae, presence of numerous strongly encrusted skeletocystidia, and smaller cylindrical basidiospores (4.5–6 × 2.5–3 µm; Dong et al. 2023). Steccherinum fragrans differs by its straw-colored basidiomata with a mild sweet smell, raduloid hymenophore with flattened aculei, pseudodimitic hyphal structure, presence of skeletocystidia, and smaller, broadly ellipsoid, cyanophilous basidiospores (3.5–4 × 2.5–2.8 µm; Crous et al. 2024). During wood-inhabiting corticioid fungi investigations in the Qinling Mountains, we also discovered S. ciliolatum (Berk. & M.A. Curtis) Gilb. & Budington, S. longiaculeiferum J.H. Dong & C.L. Zhao, S. weishanense J.H. Dong & C.L. Zhao and S. straminellum (Bres.) Melo. Compared with S. shangzhouense, S. ciliolatum possesses ceraceous to subceraceous basidiomata, well-developed spines up to 1.5 mm long, dimitic hyphal system with clamped generative hyphae, abundant encrusted cystidia (3.5–8 µm wide), longer basidia (18–22 × 4.5–6 µm), and narrower basidiospores (4.5–5.4 × 1.8–2.7 µm; Maas Geesteranus 1974); S. longiaculeiferum has coriaceous basidiomata, hydnoid hymenial surface with long aculei, dimitic hyphal system with clamped generative hyphae, and smaller, broadly ellipsoid to ellipsoid basidiospores (4–4.3 × 2.5–3 µm; Dong et al. 2024); S. weishanense has coriaceous basidiomata, odontioid hymenial surface with subulate aculei, clamped generative hyphae, two types of cystidia, and smaller ellipsoid basidiospores (4–4.8 × 2.5–3.3 µm; Dong et al. 2024); S. straminellum has an odontioid to hydnoid hymenial surface with spines up to 0.5 mm long, fimbriate margin with hyphal strands, dimitic hyphal system with clamped generative hyphae, frequent encrusted cystidia (7–15 µm wide), and narrower basidiospores (3.5–4.5 × 2–2.2 µm; Ryvarden 2024).

Overall, the present study provides additional evidence that the Qinling Mountains harbor a previously underestimated diversity of wood-inhabiting corticioid fungi. The recognition of three new species, Hymenochaete zhashuiensis, Hyphoderma brevicystidiatum and Steccherinum shangzhouense, is supported by both morphological characters and phylogenetic analyses of the combined ITS+nLSU dataset. Continued surveys in the Qinling Mountains are therefore expected to reveal additional undescribed taxa and will contribute to a more comprehensive understanding of the diversity, taxonomy and biogeography of wood-inhabiting fungi in Qinling Mountains, China.

Supplementary Material

XML Treatment for Hymenochaete zhashuiensis
XML Treatment for Hyphoderma brevicystidiatum
XML Treatment for Steccherinum shangzhouense

Citation

Wan L-L, Zhang L-S, Dong J-H, Zeng L, Liu S, Cui B-K (2026) Three new wood-inhabiting corticioid fungi from the Qinling Mountains, China. MycoKeys 138: 57–94. https://doi.org/10.3897/mycokeys.138.204396

Contributor Information

Shun Liu, Email: liushun2017@bjfu.edu.cn.

Bao-Kai Cui, Email: cuibaokai@bjfu.edu.cn.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Artificial Intelligence (AI) use

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

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

Funding

The research was supported by the National Natural Science Foundation of China (No. 32325001, 32270010), and the Fundamental Research Funds for the Central Universities (QNTD202509).

Author contributions

Conceptualization: Cui BK, Wan LL. Formal analysis: Wan LL, Zhang LS. Funding acquisition: Cui BK. Investigation: Liu S, Wan LL, Dong JH, Zeng L. Methodology: Cui BK. Project administration: Cui BK. Supervision: Cui BK. Validation: Liu S, Cui BK. Writing – original draft: Wan LL. Writing – review and editing: Liu S, Cui BK.

Author ORCIDs

L.-L. Wan https://orcid.org/0009-0002-9625-0962

L.-S. Zhang https://orcid.org/0009-0002-1832-4516

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

L. Zeng https://orcid.org/0009-0004-0124-459X

S. Liu https://orcid.org/0000-0001-9261-4365

B.-K. Cui https://orcid.org/0000-0003-3059-9344

Data availability

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

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Associated Data

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

Supplementary Materials

XML Treatment for Hymenochaete zhashuiensis
XML Treatment for Hyphoderma brevicystidiatum
XML Treatment for Steccherinum shangzhouense

Data Availability Statement

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


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