Abstract
Steccherinaceae is a speciose family of wood-inhabiting fungi containing various hymenophore configurations. Phylogenetic and morphological analyses of Steccherinaceae were carried out. Phylogenies were reconstructed using four genetic loci, including ITS, nLSU, tef1, and mtSSU. A new poroid genus, Bambusiporia, growing on dead bamboo from Yunnan Province in southwest China and typified by Bambusiporia nivea, is illustrated and described. It is characterized by resupinate basidiomata with a white pore surface when fresh, a monomitic hyphal system bearing clamp connections on generative hyphae, and ellipsoid, slightly thick-walled basidiospores. In addition, a new species in Etheirodon, E. lilacinum, is described, and it is characterized by resupinate to effused-reflexed basidiomata with a lilac hydnoid hymenophore when fresh; a dimitic hyphal system bearing clamp connections on generative hyphae; encrusted contextual generative hyphae; the absence of encrusted cystidia; and ellipsoid to broadly ellipsoid basidiospores measuring 4–4.5 × 3–3.8 µm. Moreover, the evolutionary timing of the main clades in Steccherinaceae was revealed based on conserved regions of two nuclear ribosomal genetic markers (ITS + nLSU). The ancestor of Steccherinaceae evolved during the early Cretaceous at 109.27 Myr [95% highest posterior density (HPD) of 81.91–141.87 Mya]. The initial diversification of the Bambusiporia clade occurred during the late Cretaceous with a mean crown age of 83.23 Myr [95% highest posterior density (HPD) of 56.74–112.09 Myr], earlier than other genera in Steccherinaceae. The Etheirodon clade emerged with a mean stem age of 51.98 Myr [95% highest posterior density (HPD) of 34.49–73.82 Myr] and a mean crown age of 19.56 Myr [95% highest posterior density (HPD) of 12.02–29.35 Myr].
Key words: Divergence time estimation, phylogeny, polypore, taxonomy, wood-decaying fungi
Introduction
Polyporales (Basidiomycota) is a major group of wood-inhabiting fungi, and extensive studies in the order have been carried out worldwide (Ryvarden 2015, 2016; Zhou et al. 2016; Ryvarden and Melo 2017; Cui et al. 2019; Ryvarden et al. 2022; Wu et al. 2022; Zhao et al. 2024). Steccherinaceae Parmasto is a family in Polyporales, typified by Steccherinum Gray, and was established by Parmasto (1968). Prior to the onset of the present study, the family included 23 genera, viz., Antella Miettinen, Antrodiella Ryvarden & I. Johans., Atraporiella Ryvarden, Austeria Miettinen, Butyrea Miettinen, Cabalodontia Piątek, Caudicicola Miettinen, M. Kulju & Kotir., Citripora Miettinen, Etheirodon Banker, Flabellophora G. Cunn., Flaviporus Murrill, Frantisekia Spirin & Zmitr., Junghuhnia Corda, Lamelloporus Ryvarden, Loweomyces (Kotl. & Pouzar) Jülich, Metuloidea G. Cunn., Mycorrhaphium Maas Geest., Niemelaea Zmitr., Ezhov & Khimich., Nigroporus Murrill, Rhomboidia C.L. Zhao, Steccherinum, Trullella Zmitr., and Xanthoporus Audet (Justo et al. 2017; Wu et al. 2022; Zhao et al. 2024). Steccherinaceae is characterized by resupinate, effused-reflexed to pileate or stipitate basidiomata with various hymenophores (such as smooth, granular, hydnoid, and poroid), a monomitic to dimitic hyphal system, generative hyphae with clamp connections or simple septa, allantoid, cylindrical, ellipsoid to subglobose basidiospores, and a white rot mode of wood decay (Maas Geesteranus 1971; Westphalen et al. 2021).
The three genera Antrodiella, Junghuhnia, and Steccherinum in Steccherinaceae are shown to be polyphyletic based on phylogenetic analyses (Miettinen et al. 2012; Yuan 2014; Westphalen et al. 2021; Yurchenko et al. 2023). Seven new genera have been segregated from these three genera during the last decade, namely Antella, Austeria, Butyrea, Caudicicola, Citripora, Rhomboidia, and Trullella (Miettinen and Ryvarden 2016; Kotiranta et al. 2017; Xu et al. 2020; Du et al. 2022).
The genus Etheirodon was established by Banker (1902). It has resupinate basidiomata with a fimbriate-rhizomorphic sterile margin, hydnoid to odontoid hymenophore, encrusted cystidia, and cylindrical to ellipsoid basidiospores. Prior to the onset of the present study, the genus contained three species, viz., E. fimbriatus (Pers.) Banker, E. purpureus Westphalen, and E. roseoalbus J.H. Dong & C.L. Zhao (Hjortstam and Ryvarden 2007; Westphalen et al. 2021; Dong et al. 2024). Etheirodon was treated as a synonym of Odontia Fr., a genus that belongs to Thelephoraceae in Thelephorales (Tedersoo et al. 2014).
In the present study, a phylogenetic assessment of Steccherinaceae based on four genetic markers (ITS + nLSU + tef1 + mtSSU) was carried out. Bambusiporia gen. nov. with a new species, B. nivea, is proposed. Furthermore, a new species in Etheirodon, E. lilacinum, is described and illustrated. In addition, the main morphological characteristics of all 24 genera in Steccherinaceae are summarized. The molecular divergence times of Steccherinaceae, including Bambusiporia and Etheirodon, were analyzed based on the combined two-marker dataset (ITS + nLSU) in the present study.
Materials and methods
Morphological studies
The studied specimens are deposited in the Fungarium of the Institute of Microbiology, Beijing Forestry University (BJFC). Morphological descriptions are based on field notes and voucher specimens. The microscopic analysis follows Dai (2010) and Wu et al. (2022). Sections were studied at a magnification of up to 1000× using a Nikon Eclipse 80i microscope and phase contrast illumination. Descriptions of microscopic features and measurements were made from slide preparations stained with Cotton Blue and Melzer’s reagent. Basidiospores were measured from sections cut from the tubes. To represent the variation in the size of spores, 5% of measurements were excluded from each end of the range and are given in parentheses. In the description: KOH = 5% potassium hydroxide, IKI = Melzer’s reagent, IKI– = neither amyloid nor dextrinoid, CB = Cotton Blue, CB– = acyanophilous in Cotton Blue, CB+ = cyanophilous in Cotton Blue, L = arithmetic average of spore length, W = arithmetic average of spore width, Q = L/W ratios, and n = number of basidiospores measured from the given number of specimens. Color terms follow Anonymous (1969) and Petersen (1996).
DNA extraction, amplification, and sequencing
A CTAB rapid plant genome extraction kit-DN14 (Aidlab Biotechnologies Co., Ltd., Beijing) was used to obtain DNA from dried specimens, followed by the polymerase chain reaction (PCR) according to the manufacturer’s instructions with some modifications (Sun et al. 2020; Qin et al. 2025). The internal transcribed spacer regions (ITS), large subunit nuclear ribosomal RNA gene (nLSU), translation elongation factor 1-α gene (tef1), and mitochondrial small subunit rRNA gene (mtSSU) were amplified using the primer pairs ITS5/ITS4, LR0R/LR7, 985F/1567R, and MS1/MS2 (White et al. 1990; Hopple and Vilgalys 1999; Rehner and Buckley 2005) (https://sites.duke.edu/vilgalyslab/rdna_primers_for_fungi/).
The PCR procedure for ITS, tef1, and mtSSU was as follows: initial denaturation at 95 °C for 3 min, followed by 34 cycles at 94 °C for 40 s, annealing at 54 °C for ITS and 56 °C for tef1 and mtSSU for 45 s, and extension at 72 °C for 1 min, with 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 34 cycles of denaturation at 94 °C for 30 s, annealing at 50 °C for 1 min, and extension at 72 °C for 1.5 min, with a final extension at 72 °C for 10 min. The PCR products were purified and sequenced at the Beijing Genomics Institute (BGI), China, with the same primers as used in PCR. Newly generated sequences were deposited in GenBank. All sequences analyzed in this study are listed in Table 1.
Table 1.
A list of species, specimens, and GenBank accession numbers of sequences used in this study.
| Species name | Sample no. | Location | GenBank accession no. | |||
|---|---|---|---|---|---|---|
| ITS | nLSU | mtSSU | tef1 | |||
| Agaricus campestris | LAPAG370 | — | KM657927 | KR006607 | — | — |
| Amylocorticium cebennense | CFMR: HHB-2808 | USA | GU187505 | GU187561 | — | — |
| Antella chinensis | Dai 9019 (holotype) | China | JX110844 | KC485542 | — | — |
| Antella niemelaei | Renvall 3218 | Finland | AF126876 | — | — | — |
| Antrodiella foliaceodentata | X 1238 | Russia | JN710515 | JN710515 | JN710659 | — |
| Antrodiella semisupina | X 242 | Canada | JN710521 | JN710521 | — | — |
| Antrodiella sp. | X 418 | Japan | JN710523 | JN710523 | — | — |
| Aphanobasidium pseudotsugae | CFMR: HHB-822 | USA | GU187509 | GU187567 | — | — |
| Athelia epiphylla | CFMR: FP-100564 | USA | GU187501 | GU187558 | — | — |
| Atraporiella neotropica | Ryvarden 44447 (holotype) | Belize | HQ659221 | HQ659221 | — | — |
| Atraporiella yunnanensis | CLZhao 605 (holotype) | China | MF962483 | MF962486 | — | — |
| Austeria citrea | X 1171 | New Zealand | JN710511 | — | — | — |
| Austeria citrea | PDD 96654 | New Zealand | MK404662 | — | — | — |
| Bambusiporia nivea | Dai 22451 (holotype) | China | PP907129 | PP907121 | — | — |
| Bambusiporia nivea | Dai 22477 | China | PP907130 | PP907122 | — | — |
| Bjerkandera adusta | Dai 14516 | China | MW507097 | MW520204 | — | — |
| Boletopsis leucomelaena | AFTOL-ID 1527 | USA | DQ484064 | DQ154112 | — | — |
| Boletus edulis | HMJAU4637 | — | JN563894 | KF112455 | — | — |
| Bondarzewia tibetica | Yu 56 | China | KT693203 | KT693205 | — | — |
| Butyrea japonica | Li 1648 | China | KC485536 | KC485553 | — | — |
| Butyrea luteoalba | isolate 5403 | Estonia | JN710558 | JN710558 | JN710682 | JN710719 |
| Cabalodontia delicata | MV 370 | — | MT849298 | — | — | — |
| Cabalodontia delicata | SP 512584 (holotype) | Brazil | NR174056 | — | — | — |
| Cabalodontia queletii | CBS 233.56 | France | MH857599 | MH869147 | — | — |
| Callistosporium graminicolor | AFTOL-ID 978 | USA | DQ484065 | AY745702 | — | — |
| Ceriporia allantospora | RLG-10478 (holotype) | USA | KP135039 | — | — | — |
| Ceriporia aurantiocarnescens | Dai 17951 | China | MW491774 | MW491764 | — | — |
| Ceriporia crassa | Dai 22034 (holotype) | China | OQ476823 | OQ476769 | — | — |
| Ceriporia griseoviolascens | Dai 13202 | France | OQ476825 | OQ476771 | — | — |
| Ceriporia hinnulea | Cui 11291 (holotype) | China | OQ476826 | OQ476772 | — | — |
| Ceriporia mellita | BR 4865 | France | KX236485 | KX236485 | — | — |
| Ceriporia sinoviridans | Dai 13621A (holotype) | China | MW491781 | MW491771 | — | — |
| Ceriporia spissa | Dai 19164 | Canada | OQ476845 | OQ476789 | — | — |
| Ceriporia subviridans | Cui 8012 (holotype) | China | KC182774 | — | — | — |
| Ceriporia viridans | Dai 17003 | China | OQ476847 | OQ476790 | — | — |
| Citripora afrocitrina | X 525 | Uganda | JN710507 | JN710507 | JN710655 | JN710710 |
| Citripora bannaensis | X 243 | China | JN710526 | JN710526 | — | — |
| Cotylidia sp. | MB5 | — | AY854079 | AY629317 | — | — |
| Crystallicutis sp. | Dai 6090 | China | JX623934 | JX644066 | — | — |
| Crystallicutis serpens | HHB-15692-Sp | USA | KP135031 | KP135200 | — | — |
| Cymatoderma sp. | OMC 1427 | USA | KY948826 | KY948872 | — | — |
| Etheirodon aff. fimbriatus | HHB-2878-sp | USA | KY948822 | KY948864 | — | — |
| Etheirodon cf. fimbriatus | KUC 20121109-29 | Korea | KJ668456 | KJ668307 | — | — |
| Etheirodon cf. fimbriatus | Dai 24450 | China | PP907136 | PP907128 | — | |
| Etheirodon cf. fimbriatus | CLZhao 13977 | China | — | — | ||
| Etheirodon fimbriatus | KHL 11905 | Sweden | JN710530 | JN710530 | JN710667 | — |
| Etheirodon fimbriatus | HR 97926 | — | MT849299 | — | — | MT833937 |
| Etheirodon fimbriatus | HR 98811 | — | MT849300 | — | — | MT833938 |
| Etheirodon lilacinum | Dai 23571 (holotype) | China | PP907131 | PP907123 | — | — |
| Etheirodon lilacinum | Dai 23574 | China | PP907132 | PP907124 | PX649048 | PX667841 |
| Etheirodon lilacinum | Dai 23131 | China | PP907133 | PP907125 | PX649047 | PX667842 |
| Etheirodon lilacinum | Dai 23568 | China | PP907134 | PP907126 | PX649046 | PX667843 |
| Etheirodon lilacinum | Dai 23140 | China | PP907135 | PP907127 | — | — |
| Etheirodon purpureus | MCW 642/18 (holotype) | Brazil | MT849301 | MT849301 | — | MT833939 |
| Etheirodon roseoalbus | CLZhao 24770 (holotype) | China | OR096187 | OR461452 | — | — |
| Etheirodon roseoalbus | CLZhao 24903 | China | OR096188 | OR461453 | — | — |
| Flabellophora sp.1 | X 1357 | Indonesia | JN710533 | JN710533 | — | — |
| Flabellophora sp.3 | X 1277 | Indonesia | JN710535 | JN710535 | JN710669 | — |
| Flaviporus albus | GXU 5765 (holotype) | China | OQ981991 | OQ981993 | — | — |
| Flaviporus brownii | X 1216 | Ecuador | JN710537 | JN710537 | — | — |
| Flaviporus subundatus | MCW 457/13 | — | KY175005 | — | — | — |
| Flaviporus tenuis | MCW 44213 | — | KY175001 | KY175001 | — | — |
| Frantisekia mentschulensis | AH 1377 | Austria | JN710544 | JN710544 | — | — |
| Frantisekia mentschulensis | BRNM 710170 | Czechia | FJ496670 | FJ496728 | FJ496748 | — |
| Frantisekia ussurii | Wei 3081 | China | KC485527 | KC485545 | — | — |
| Frantisekia ussurii | Dai 8249 | China | KC485526 | — | — | — |
| Gloeophyllum sepiarium | Wilcox-3BB | USA | HM536091 | HM536061 | — | — |
| Gloeoporus dichrous | Dai 23260 | China | OQ476852 | OQ476795 | — | — |
| Gloeoporus pannocinctus | FP 135015 | USA | MG572755 | MG572739 | — | — |
| Hydnochaete duportii | AFTOL-ID 666 | — | DQ404386 | AY635770 | — | — |
| Hydnophanerochaete odontoidea | CWN 00776 | China | LC363487 | GQ470663 | — | — |
| Hyphoderma litschaueri | FP-101740-Sp | USA | KP135295 | KP135219 | — | — |
| Hyphoderma mutatum | HHB-15479-Sp | USA | KP135296 | KP135221 | — | — |
| Hyphoderma praetermissum | AFTOL-ID 518 | — | AY854081 | AY700185 | — | — |
| Hyphoderma setigerum | FD-312 | USA | KP135297 | KP135222 | — | — |
| Hypochnicium karstenii | NH 10924 | Sweden | DQ677510 | DQ677510 | — | — |
| Hypochnicium polonense | NH 12117 | Russia | EU118635 | EU118635 | — | — |
| Jaapia argillacea | CBS: 252.74 | Netherlands | GU187524 | GU187581 | — | — |
| Junghuhnia crustacea | X 262 | Indonesia | JN710553 | JN710553 | JN710678 | — |
| Junghuhnia crustacea | X 1127 | Indonesia | JN710554 | JN710554 | — | — |
| Junghuhnia micropora | Spirin 2652 | Russia | JN710559 | JN710559 | JN710683 | JN710720 |
| Lactarius deceptivus | AFTOL-ID 682 | USA | AY854089 | AY631899 | — | — |
| Lamelloporus americanus | X 670 | Ecuador | JN710567 | JN710567 | — | — |
| Lamelloporus americanus | RLC 779 | Ecuador | OQ871855 | — | — | — |
| Leptosporomyces raunkiaeri | CFMR: HHB-7628 | USA | GU187528 | GU187588 | — | — |
| Loweomyces fractipes | X 1253 | USA | JN710569 | JN710569 | JN710689 | — |
| Loweomyces fractipes | X 1250 | USA | JN710568 | JN710568 | — | — |
| Loweomyces wynneae | X 1215 | Denmark | JN710604 | JN710604 | JN710709 | — |
| Luteochaete subglobosa | GC 1605-4 | China | MZ636995 | MZ637156 | — | — |
| Meripilus albostygius | Kout 1807/15.1 (holotype) | Puerto Rico | OM669892 | OM669976 | — | — |
| Meripilus crataegi | Dai 15497 (holotype) | China | KY131845 | KY131904 | — | — |
| Meripilus eminens | Dai 22472 | China | OM669900 | OM669983 | — | — |
| Meripilus expallescens | Dai 21060 | Belarus | MT840130 | MT840148 | — | — |
| Meripilus furcatus | TAA 150972 (holotype) | Russia | KY131853 | KY131910 | — | — |
| Meripilus giganteus | Cui 9202 | UK | OM669888 | OM669973 | — | — |
| Meripilus neovitreus | JV 1009/59 (holotype) | USA | OM669908 | OM669990 | — | — |
| Meripilus pouzarii | Dai 21043 | Belarus | MT840124 | MT840142 | — | — |
| Meripilus sanguinolentus | JV 1310/11 | Czechia | OM669920 | OM669998 | — | — |
| Meripilus srilankensis | Dai 19535 (holotype) | Sri Lanka | OM669924 | OM670001 | — | — |
| Meripilus sumstinei | Russell 5913 | USA | MN906088 | — | — | — |
| Meripilus tibeticus | Cui 9588 | China | KY131873 | KY131929 | — | — |
| Meripilus vitreus | Dai 12685 | Czechia | MT840115 | MT840133 | — | — |
| Meruliopsis albomellea | Dai 15205 (holotype) | China | KX494574 | KX494578 | — | — |
| Meruliopsis bambusicola | Dai 21944 (holotype) | China | OQ476864 | OQ476806 | — | — |
| Meruliopsis crassitunicata | Dai 10833 | China | JX623935 | JX644064 | — | — |
| Meruliopsis nanlingensis | Dai 13414 | China | OQ476868 | OQ476809 | — | — |
| Meruliopsis tarda | Dai 10226 | China | JX623945 | — | — | — |
| Meruliopsis taxicola | Dai 22625 | China | OL457966 | OL457436 | — | — |
| Metuloidea murashkinskyi | X 449 | Russia | JN710588 | JN710588 | — | — |
| Metuloidea rhinocephala | X 460 | Australia | JN710562 | JN710562 | JN710686 | — |
| Mycorrhaphium adustum | X 8024 | USA | JN710573 | JN710573 | JN710692 | — |
| Mycorrhaphium subadustum | Yuan 12976 (holotype) | China | MW491378 | MW488040 | — | — |
| Neolentinus adhaerens | DAOM 214911 | — | HM536096 | HM536071 | — | — |
| Niemelaea balaenae | H 7002389 | Canada | FJ496669 | FJ496717 | FJ496746 | — |
| Niemelaea consobrina | Rivoire 977 | France | FJ496667 | FJ496716 | — | — |
| Nigroporus stipitatus | X 546 | Cameroon | JN710574 | JN710574 | — | — |
| Nigroporus vinosus | X 839 | USA | JN710575 | JN710575 | — | — |
| Panus fragilis | HHB-11042-Sp | USA | KP135328 | KP135233 | — | — |
| Phanerochaete alnea | FP-151125 | USA | KP135177 | MZ637181 | — | — |
| Phanerochaetella exilis | HHB-6988 | USA | KP135001 | KP135236 | — | — |
| Phanerochaetella xerophila | HHB-8509-Sp | USA | KP134996 | KP135259 | — | — |
| Phlebiopsis gigantea | FCUG 1417 | Norway | MZ637051 | AF141634 | — | — |
| Phlebiporia bubalina | Dai 13168 | China | KC782526 | KC782528 | — | — |
| Podoscypha multizonata | Jahn 751012 | Germany | EU118663 | EU118663 | — | — |
| Podoscypha venustula | LR 40821 | Venezuela | JX109851 | JX109851 | — | — |
| Podoserpula ailaoshanensis | ZJL2015015 | China | KU324484 | KU324487 | — | — |
| Pseudolagarobasidium baiyunshanense | Han 405 (holotype) | China | MT428549 | MT428547 | — | — |
| Radulodon americanus | CFMR-HHB 11240 | USA | JQ070174 | — | — | — |
| Radulodon yunnanensis | Cui 17979 (holotype) | China | OM971917 | OM971898 | — | — |
| Resiniporus pseudogilvescens | Wu 1209-46 | China | KY688203 | MZ637268 | — | — |
| Rhomboidia wuliangshanensis | CLZhao 4406 (holotype) | China | MK860715 | MK860710 | — | — |
| Rhomboidia wuliangshanensis | CLZhao 4411 | China | MK860716 | MK860711 | — | — |
| Russula emeticicolor | FH12253 | Germany | KT934011 | KT933872 | — | — |
| Schizophyllum radiatum | AFTOL-ID-516 | Panama | AY571060 | AY571023 | — | — |
| Scopuloides hydnoides | FP-150473 | USA | KP135355 | KP135284 | — | — |
| Serpula himantioides | MUCL: 30528 | Belgium | GU187545 | GU187600 | — | — |
| Skeletocutis novae-zelandiae | Ryvarden 38641 | New Zealand | JN710582 | JN710582 | — | — |
| Spongipellis quercicola | Cui 10009 (holotype) | China | OM971919 | OM971899 | — | — |
| Spongipellis sibirica | Dai 1723 | China | OM971921 | — | — | — |
| Spongipellis spumeus | BRNM 734877 | Czechia | HQ728283 | HQ729018 | — | — |
| Spongiporus leucospongia | OKM-4335 | USA | KC585395 | KC585228 | — | — |
| Steccherinum autumnale | Spirin 2957 | Russia | JN710549 | JN710549 | JN710675 | JN710716 |
| Steccherinum fragile | Dai 19972 | China | MW364629 | MW364627 | — | — |
| Steccherinum fragile | Dai 20479 (holotype) | China | MW364628 | MW364626 | — | — |
| Steccherinum incrustans | Dai 19442 | China | ON182084 | ON182087 | — | — |
| Steccherinum incrustans | X 1345 | China | JN710550 | JN710550 | — | — |
| Steccherinum juniperi | Dai 23930 | China | OP956076 | — | — | — |
| Steccherinum meridionale | CBS 125887 | New Zealand | MH864086 | MH875544 | — | — |
| Steccherinum nandinae | Dai 21108 | China | MN833678 | MN833680 | — | — |
| Steccherinum nitidum | KHL 11903 | Sweden | JN710560 | JN710560 | JN710684 | JN710721 |
| Steccherinum ochraceum | KHL 11902 | Sweden | JN710590 | JN710590 | — | — |
| Steccherinum polycystidiferum | RP 140 | — | KY174996 | KY174996 | — | — |
| Steccherinum subcollabens | Dai 19345 (holotype) | China | MN871759 | MN877772 | — | — |
| Terana caerulea | FP-104073 | USA | KP134980 | KP135276 | — | — |
| Tomentella sp. | AFTOL-ID 1016 | USA | DQ835998 | DQ835997 | — | — |
| Trullella dentipora | X 200 | Venezuela | JN710512 | JN710512 | — | — |
| Trullella polyporoides | X 510 | Venezuela | JN710602 | JN710602 | — | — |
| Xanthoporus syringae | Gothenburg 1488 | Sweden | JN710607 | JN710607 | — | — |
| Xanthoporus syringae | X 339 | Finland | JN710606 | JN710606 | — | — |
New taxa and newly generated sequences are in bold.
Sequence alignment
Sequences generated from this study were aligned with additional sequences downloaded from GenBank using BioEdit (Hall 1999). The final ITS, nLSU, tef1, and mtSSU datasets were subsequently aligned using MAFFT v.7 under the E-INS-i strategy (command line: mafft –genafpair –maxiterate 1000) (Katoh and Standley 2013) and visualized in BioEdit. Alignments were manually concatenated and processed further in Mesquite v.3.2 (Maddison and Maddison 2017).
Phylogenetic analyses
In this study, one combined matrix was reconstructed for phylogenetic analyses; a four-marker dataset (ITS + nLSU + tef1 + mtSSU) was used to determine the phylogenetic position of the new species. The sequence alignments and the retrieved topologies were deposited in TreeBase (http://www.treebase.org), under accession ID: 32532 (Reviewer access URL: http://purl.org/phylo/treebase/phylows/study/TB2:S32532?x-access-code=74c447e7541163ef3f4cefaf104f2b6&format=html). Sequences of Hyphoderma setigerum (Fr.) Donk and Hyphoderma litschaueri (Burt) J. Erikss. & Å. Strid, obtained from GenBank, were used as the outgroups (Justo et al. 2017). The phylogenetic analyses followed the approach of Han et al. (2016) and Zhu et al. (2019). Maximum likelihood (ML) and Bayesian inference (BI) analyses were performed based on the four-marker dataset. The combined dataset (ITS1 + 5.8S + ITS2 + nLSU + tef1 + mtSSU) was partitioned into six subsets, and the best-fit substitution model for each partition was selected under the Akaike Information Criterion (AIC) using PartitionFinder within PhyloSuite v1.2.3 (Lanfear et al. 2017; Zhang et al. 2020). Then, the models were estimated separately for ITS1, 5.8S, ITS2, nLSU, tef1, and mtSSU, and Bayesian inference (BI) was analyzed using a partitioned, mixed-model run.
Sequences were analyzed using maximum likelihood (ML) with RAxML-HPC v.8.2.12 through the CIPRES Science Gateway (Miller et al. 2010). Branch support (BT) for ML analysis was determined by 1000 bootstrap replicates. Bayesian phylogenetic inference and Bayesian posterior probabilities (BPP) were computed with MrBayes 3.2.6. Four Markov chains were run for 3 million generations until the split deviation frequency value was less than 0.01, and trees were sampled every 100 generations. The first 25% of the sampled trees were discarded as burn-in, and the remaining ones were used to infer a majority rule consensus and calculate Bayesian posterior probabilities (BPP) of the clades. All trees were viewed in FigTree v.1.4.3 (http://tree.bio.ed.ac.uk/software/figtree/). Branches that received bootstrap support ML ≥ 75% and BPP ≥ 0.95 were considered significantly supported. The ML bootstrap supports of ≥ 50% and BPP of ≥ 0.90 are presented on the topology from the ML analysis.
Divergence time estimation
Archaeomarasmius leggetti Hibbett et al. and Quatsinoporites cranhamii Smith et al. were selected as fossil calibrations in the divergence times of Steccherinaceae, including Etheirodon and the new genus Bambusiporia. A. leggetti was recorded at 94–90 Myr (Hibbett et al. 1997) as the representative of the minimum age of Tricholomataceae R. Heim ex Pouzar belonging to the Agaricales. Q. cranhamii, found in marine calcareous concretions on Vancouver Island, was considered to represent the minimum divergence time of the Hymenochaetales at 113 Myr (Smith et al. 2004). Divergence times were estimated using BEAST v2.6.5 (Bouckaert et al. 2014) based on a dataset of ITS + nLSU. The GTR + G + I substitution model was selected as the best-fit model for the two-marker dataset using MrModelTest2-v.2.4 (Nylander 2004). An XML file was executed using BEAUti v2. The clock model was set to an uncorrelated lognormal relaxed clock (Drummond et al. 2006; Lepage et al. 2007). The Yule process speciation was used as the tree prior (Gernhard 2008). For calibration, a gamma distribution prior (scale = 20, shape = 1) was specified for the Agaricales (offset = 90 Myr) and Hymenochaetales (offset = 125 Myr) clades (Sánchez-Ramírez et al. 2014; Zhao et al. 2016, 2017). All the ucld.mean parameters for different genes were set to uniform. Monte Carlo Markov chains were run for 100 million generations, logging states every 1000 generations. The resulting log file was checked for convergence of the chains using Tracer v1.6 (Rambaut et al. 2013; http://tree.bio.ed.ac.uk/software/tracer/). An ultrametric maximum clade credibility (MCC) tree was summarized using TreeAnnotator v2.6.5, discarding 20% of states as burn-in and annotating clades with ≥ 0.8 posterior probability. FigTree v1.4.3 (http://tree.bio.ed.ac.uk/software/figtree/) was used to visualize the resulting tree and to obtain the means and 95% HPD (Drummond and Rambaut 2007). A 95% HPD marks the shortest interval that contains 95% of the values sampled.
Results
Molecular phylogeny
The combined four-marker dataset (ITS + nLSU + tef1 + mtSSU) included sequences from 77 samples representing 55 taxa, and the dataset had an aligned length of 3397 characters. The phylogenetic reconstruction performed with maximum likelihood (ML) and Bayesian inference (BI) analyses on the combined dataset showed a similar topology and only minor differences in statistical support. The substitution model employed for the ML analysis was GTRGAMMA, and all sequences divided into six partitions were Subset1 (ITS1) = 1–335, Subset2 (5.8S) = 336–510, Subset3 (ITS2) = 511–882, Subset4 (nLSU) = 883–2256, Subset5 (tef1) = 2257–2814, and Subset6 (mtSSU) = 2815–3397. The best model fit applied in the Bayesian analysis for each region of the six partitions was ITS1 (GTR+I+G), 5.8S (TVMEF+G), ITS2 (TVM+I+G), nLSU (GTR+I+G), tef1 (TRNEF+G), and mtSSU (GTR+I+G); lset nst for ITS1, 5.8S, ITS2, nLSU, tef1, and mtSSU = 6; rates = invgamma (ITS1, ITS2, nLSU, and mtSSU); gamma (5.8S, tef1); and prset statefreqpr = dirichlet (1, 1, 1, 1). Bayesian analysis resulted in a nearly congruent topology with an average standard deviation of split frequencies = 0.004359, and thus only the ML tree is provided (Fig. 1). The phylogeny (Fig. 1) indicated the taxonomic relationship of genera in Steccherinaceae, and the Bambusiporia clade is phylogenetically close to the Citripora clade without strong support. The new species Etheirodon lilacinum differed from the other three species, E. fimbriatus, E. purpureus, and E. roseoalbus, nested in the Etheirodon clade. The polypore genera Antrodiella, Junghuhnia, and Skeletocutis are polyphyletic within the Steccherinaceae and Incrustoporiaceae, respectively. Three specimens (X418, Spirin 2652, and Ryvarden 38641), identified as Antrodiella sp., Junghuhnia micropora, and Skeletocutis novae-zelandiae, formed an independent clade in the previous study (Miettinen et al. 2012). In our phylogenetic tree, these three specimens are still assigned the original taxon names in accordance with the taxonomic concept of Miettinen et al. (2012).
Figure 1.
ML analysis of Steccherinaceae based on a dataset of ITS + nLSU + tef1 + mtSSU. ML bootstrap values higher than 50% and Bayesian posterior probability values more than 0.90 are shown. New taxa are in bold.
Divergence time estimation
The MCMC tree (Fig. 2) shows that the ancestor of the Steccherinaceae evolved during the early Cretaceous at 109.27 Myr [95% highest posterior density (HPD) of 81.91–141.87 Myr]. The initial diversification of the Etheirodon clade occurred during the Eocene in the Paleogene Period with a mean stem age of 51.98 Myr [95% highest posterior density (HPD) of 34.49–73.82 Myr] and a mean crown age of 19.56 Myr [95% highest posterior density (HPD) of 12.02–29.35 Myr]. The new genus Bambusiporia was estimated at 83.23 Myr, emerging in the late Cretaceous, and the posterior probability is up to 1. However, the divergence time of the Citripora clade emerged with a mean crown age of 44.62 Myr [95% highest posterior density (HPD) of 25–67.98 Myr], which belongs to the Eocene of the Paleogene Period. The international chronostratigraphic chart follows Cohen et al. (2013; updated) (URL: https://stratigraphy.org/chart/).
Figure 2.
Divergence time estimation of Steccherinaceae from Bayesian evolutionary analysis sampling tree based on the conserved regions of two DNA fragments (ITS + nLSU). Posterior probabilities are not less than 0.80, and the mean ages (Myr) of each node are annotated. The 95% highest posterior densities of divergence time estimation are marked by horizontal bars.
Taxonomy
Bambusiporia
Y.C. Dai, Xin Zhang & Chao G. Wang gen. nov.
F70CD51E-EE21-5787-934F-682D922986C0
MB854326
Etymology.
Bambusiporia (Lat.): refers to the genus having resupinate basidiomata and growing on bamboo.
Type species.
Bambusiporia nivea Y.C. Dai & Chao G. Wang, sp. nov.
Description.
Basidiomata tiny, annual, resupinate, detachable, soft when fresh. Poroid hymenophore, white when fresh. Hyphal system monomitic; generative hyphae bearing clamp connections, hyaline, thin- to slightly thick-walled. Cystidia and cystidioles absent. Basidiospores ellipsoid, hyaline, slightly thick-walled, smooth, sometimes with one or two small guttules, IKI−, weakly CB+.
Bambusiporia nivea
Y.C. Dai, Xin Zhang & Chao G. Wang sp. nov.
6C1CCB70-933A-5088-A1F9-810BB54B45E0
MB854327
Figure 3.
Basidiomata of Bambusiporia nivea (holotype, Dai 22451). Scale bar: 1 cm.
Figure 4.
Microscopic structures of Bambusiporia nivea (holotype, Dai 22451). a. Basidiospores; b. Basidia and basidioles; c. Hyphae from subiculum; d. Hyphae from trama.
Holotype.
China • Yunnan Province, Zhaotong, Huanglianhe Forest Park, on dead bamboo, 30 June 2021, Dai 22451 (BJFC037035).
Etymology.
Nivea (Lat.): refers to the species having white pore surface when fresh.
Description.
Basidiomata. Annual, resupinate, separate, soft, without odor or taste when fresh, soft corky when dry, up to 1 cm long, 0.8 cm wide, 2 mm thick at center. Pore surface white when fresh, becoming pinkish buff upon drying; sterile margin very narrow to almost absent; pores round to angular, 4–6 per mm; dissepiments thin, entire. Subiculum thin, cream, corky, up to 0.5 mm thick. Tubes concolorous with pore surface, soft corky, up to 1.5 mm long.
Hyphal structure. Hyphal system monomitic; generative hyphae bearing clamp connections, hyaline, IKI−, CB−; tissues unchanged in KOH.
Subiculum. Generative hyphae slightly thick-walled with a wide lumen, occasionally branched, more or less flexuous, loosely interwoven, 4–7 μm in diam. Irregular crystals present amongst subicular hyphae.
Tubes. Generative hyphae thin- to slightly thick-walled with a wide lumen, occasionally branched, more or less flexuous, 3–5 μm in diam. Cystidia and cystidioles absent. Basidia clavate, with four sterigmata and a basal clamp connection, 15–20 × 6–6.5 μm; basidioles in shape similar to basidia but smaller.
Spores. Basidiospores ellipsoid, hyaline, slightly thick-walled, smooth, sometimes with one or two small guttules, IKI−, weakly CB+, (3.8–)4–4.6(–4.8) × 3–3.7(–3.8) µm, L = 4.19 μm, W = 3.3 μm, Q = 1.26–1.28 (n = 60/2).
Type of rot. White rot.
Additional specimen examined.
China • Yunnan Province, Zhaotong, Huanglianhe Forest Park, on dead bamboo, 30 June 2021, Dai 22477 (BJFC037061).
Notes.
Bambusiporia nivea is characterized by resupinate basidiomata with a white pore surface when fresh, round to angular pores of 4–6 per mm, a monomitic hyphal system with generative hyphae bearing clamp connections, ellipsoid, slightly thick-walled basidiospores measuring 4–4.6 × 3–3.7 µm, and growing on dead bamboo in southwest China.
Etheirodon lilacinum
Y.C. Dai & Chao G. Wang sp. nov.
D0633E10-FB1B-542D-9241-126CD8067065
MB854328
Figure 5.
Basidiomata of Etheirodon lilacinum (holotype, Dai 23568). Scale bar: 1 cm.
Figure 6.
Microscopic structures of Etheirodon lilacinum (holotype, Dai 23568). a. Basidiospores; b. Basidia and basidioles; c. Cystidioles; d. Hyphae from context; e. Hyphae from spines.
Holotype.
China • Xizang Autonomous Region, Linzhi, Bomi County, on fallen trunk of Betula, 26 October 2021, Dai 23568 (BJFC038140).
Etymology.
Lilacinum (Lat.): refers to the species having lilac pore surface when fresh.
Description.
Basidiomata. Annual, resupinate to slightly effused-reflexed with small pilei, soft to fleshy, without odor or taste when fresh, soft corky when dry, up to 20 cm long, 3 cm wide when resupinate. Hymenophore hydnoid, lilac when fresh, dark grayish violet when dry; sterile margin distinct, lilac when fresh, grayish violet when dry, up to 3 mm wide, fimbriate to dentate; spines soft when fresh, soft corky when dry, up to 1 mm long, cylindrical to flattened, 2–4 per mm at base. Context pale grayish violet, soft corky when dry, up to 1.5 mm thick.
Hyphal structure. Hyphal system dimitic; generative hyphae bearing clamp connections; skeletal hyphae IKI−, CB−; tissues becoming pale olivaceous in KOH.
Context. Generative hyphae infrequent, hyaline, thin- to thick-walled with a wide lumen, sometimes covered with fine hyaline crystals, occasionally branched, straight, 3–4 μm in diam.; skeletal hyphae dominant, brownish orange, thick-walled with a narrow lumen to subsolid, smooth, rarely branched, more or less flexuous, loosely interwoven, 3–6 μm in diam.
Spines. Generative hyphae infrequent, hyaline, thin- to slightly thick-walled with a wide lumen, smooth, rarely branched, straight, 3–3.5 μm in diam.; skeletal hyphae dominant, brownish orange, thick-walled with a medium to narrow lumen, smooth, occasionally branched, straight, subparallel along the spines, agglutinated, 3.5–4 μm in diam. Cystidia absent; cystidioles fusoid, hyaline, thin-walled, smooth, 15–20 × 4 μm. Basidia clavate, with four sterigmata and a basal clamp connection, 20–23 × 4.5–5 μm; basidioles in shape similar to basidia, but smaller. Irregular crystals present amongst hymenia.
Spores. Basidiospores ellipsoid to broadly ellipsoid, hyaline, thin-walled, smooth, IKI−, CB−, (3.8–)4–4.5(–5) × (2.8–)3–3.8(–4) µm, L = 4.12 μm, W = 3.2 μm, Q = 1.27–1.3 (n = 90/3).
Type of rot. White rot.
Additional specimens examined.
China • Sichuan Province, Ganzi, Luding County, Hailuogou Forest Park, on fallen branch of Abies, 8 October 2021, Dai 23131 (BJFC037702), Dai 23140 (BJFC037711); Xizang Autonomous Region, Linzhi, Bomi County, on fallen trunk of Betula, 26 October 2021, Dai 23571 (BJFC038143), Dai 23574 (BJFC038146).
Notes.
Etheirodon lilacinum is characterized by resupinate to slightly effused-reflexed basidiomata; a hydnoid and lilac hymenophore when fresh, dark grayish violet when dry; cylindrical to flattened spines of 2–4 per mm at the base; a dimitic hyphal system bearing clamp connections on generative hyphae; thin- to thick-walled generative hyphae sometimes covered with fine hyaline crystals; ellipsoid to broadly ellipsoid basidiospores measuring 4–4.5 × 3–3.8 µm; and growing on angiosperm and gymnosperm wood.
Key to genera within Steccherinaceae
| 1 | Hyphal system monomitic | 2 |
| – | Hyphal system dimitic or dimitic to trimitic | 10 |
| 2 | Hymenophore odontoid, tuberculate or smooth | Cabalodontia |
| – | Hymenophore poroid | 3 |
| 3 | Basidiomata resupinate | 4 |
| – | Basidiomata effused-reflexed, pileate to stipitate | 7 |
| 4 | Pore surface brown | Atraporiella |
| – | Pore surface light-colored | 5 |
| 5 | Ampullaceous septa and gloeocystidia present | Caudicicola |
| – | Ampullaceous septa and gloeocystidia absent | 6 |
| 6 | Tubes shallow, pores large and angular, sterile margin distinct | Niemelaea |
| – | Tubes long, pores small and round to angular, sterile margin absent | Bambusiporia |
| 7 | Hyphae in context swollen | Xanthoporus |
| – | Hyphae in context normal | 8 |
| 8 | Basidiospores thin-walled | Rhomboidia |
| – | Basidiospores thick-walled | 9 |
| 9 | Pilei usually imbricate with a common stipe, pores small | Flabellophora |
| – | Pilei usually solitary with a stipe or not, pores large | Loweomyces |
| 10 | Gloeocystidia present | 11 |
| – | Gloeocystidia absent | 13 |
| 11 | Basidiomata resupinate to pileate | Antrodiella |
| – | Basidiomata completely resupinate | 12 |
| 12 | Pore surface white to yellow | Antella |
| – | Pore surface straw-colored | Butyrea |
| 13 | Hymenophore completely poroid | 14 |
| – | Hymenophore poroid, odontoid, irpicoid, lamellae or corticioid | 19 |
| 14 | Pore surface sulfur-yellow or citric-yellow | 15 |
| – | Pore surface white, reddish, or brownish vinaceous | 16 |
| 15 | Basidiospores subcylindrical | Austeria |
| – | Basidiospores broadly ellipsoid to subglobose | Citripora |
| 16 | Encrusted thick-walled cystidia absent | 17 |
| – | Encrusted thick-walled cystidia present | 18 |
| 17 | Basidiospores allantoid | Nigroporus |
| – | Basidiospores oblong-ellipsoid | Frantisekia |
| 18 | Basidiomata resupinate | Junghuhnia |
| – | Basidiomata pileate | Flaviporus |
| 19 | Sterile margin fimbriate-rhizomorphic | Etheirodon |
| – | Sterile margin smooth | 20 |
| 20 | Encrusted thick-walled cystidia present | 21 |
| – | Encrusted thick-walled cystidia absent | 22 |
| 21 | Hyphal system trimitic, skeletal hyphae rather wide | Metuloidea |
| – | Hyphal system dimitic, skeletal hyphae narrow | Steccherinum |
| 22 | Basidiomata pileate without stipes | Lamelloporus |
| – | Basidiomata pileate with stipes | 23 |
| 23 | Basidiospores cylindrical and curved | Trullella |
| – | Basidiospores cylindrical to broadly ellipsoid | Mycorrhaphium |
Discussion
In the present study, phylogenetic analyses using a four-marker dataset (ITS + nLSU + tef1 + mtSSU) illustrate the phylogeny of genera belonging to Steccherinaceae in Polyporales (Fig. 1). Bambusiporia forms an independent clade and is proposed as a new genus. Recently, divergence time estimation has been applied in fungal taxonomy, especially in genera, families, or higher-ranking taxa, to support fungal systematics (Chen et al. 2015; Varga et al. 2019; Zhao et al. 2023, 2025; Dong et al. 2024). In the present study, divergence time estimation of Bambusiporia was analyzed, and the result supports our proposal for the establishment of the new genus. Multi-gene phylogenetic analysis is a core approach for resolving phylogenetic relationships and delimiting taxonomic units in fungal systematics (Miettinen et al. 2012). Sufficient and matched sequence quantities across all markers can provide accurate resolution of phylogenetic relationships among closely related taxa. However, quantitative disparities among individual sequence fragments will lead to deviations in multi-gene analyses, and sampling more individuals often gives better results than sampling more loci in phylogenetic studies at the genus level (James et al. 2006; Maddison and Knowles 2006). In this study, a total of 77 samples representing 55 taxa were studied in our phylogenetic analysis, including 77 ITS sequences, 66 nLSU sequences, 16 mtSSU sequences, and 12 tef1 sequences obtained. Phylogenetic results from multi-gene (ITS + nLSU + tef1 + mtSSU) and two-gene (ITS + nLSU) analyses showed similar topology.
Bamboos are monocotyledonous woody plants. Fewer studies on bamboo-decaying fungi were reported before the 21st century. However, recently the diversity of these fungi has been investigated, especially in China, and more than 20 new taxa of wood-decaying fungi on bamboos have been described (Dai et al. 2021; Wang et al. 2021; Mao et al. 2023; Zhang et al. 2023; Zhou et al. 2023; Cui et al. 2025). It seems that more unknown taxa of bamboo-decaying fungi will be described after further investigation. The present study proposes a new genus and a new species on bamboos from China.
Bambusiporia is a light-colored polypore that forms a separate, strongly supported clade in Steccherinaceae in our phylogenetic tree. So far, 24 genera accepted in Steccherinaceae have various types of hymenophores (poroid, hydnoid, and corticioid, Table 2). In the dating analyses (Fig. 2), the new genus Bambusiporia was estimated to have emerged in the late Cretaceous with a mean crown age of 83.23 Myr. However, the divergence time of the Citripora clade and the Steccherinum s.s. clade, with mean crown ages of 44.62 Myr and 52.09 Myr, respectively, both occurred during the Eocene in the Paleogene and are posterior to Bambusiporia. So far, the Bambusiporia clade diverged earlier than other genera in Steccherinaceae (Fig. 2). The divergence times of the Poaceae with a crown age of 101 Myr occurred during the early Cretaceous, and the ancestor of the bamboos (Poaceae: Bambusoideae) evolved during the Cretaceous–Paleogene (K–Pg) boundary at 66 Myr (Huang et al. 2022). In contrast, the new genus Bambusiporia was estimated to have diverged at 83.23 Myr in the late Cretaceous, predating the origin of Bambusoideae. Thus, Bambusiporia likely originally decomposed other monocotyledonous plants or gymnosperms and subsequently underwent a host shift to bamboo, a pattern consistent with that of some species such as Meripilus cinereus and M. lineatus, which are known to associate with both bamboos and gymnosperms as hosts (Wang et al. 2025).
Table 2.
The main morphological characteristics of all 24 genera in Steccherinaceae.
| Genera | Type species | Basidiomata | Type of hymenophore | Hyphal system | Shape of basidiospores | References |
|---|---|---|---|---|---|---|
| Antella | A. niemelaei | Resupinate | Poroid to irpicoid | Dimitic | Ellipsoid | Miettinen and Ryvarden (2016) |
| Antrodiella | A. semisupina | Resupinate to pileate | Poroid | Dimitic to trimitic | Cylindrical to ellipsoid | Ryvarden and Gilbertson (1993); Yuan (2014) |
| Atraporiella | A. neotropica | Resupinate | Poroid | Monomitic | Ellipsoid to slightly allantoid | Ryvarden (2007) |
| Austeria | A. citrea | Pileate | Poroid | Dimitic | Subcylindrical | Miettinen and Ryvarden (2016) |
| Bambusiporia | B. nivea | Resupinate | Poroid | Monomitic | Ellipsoid | This study |
| Butyrea | B. luteoalba | Resupinate | Poroid | Dimitic | Cylindrical | Miettinen and Ryvarden (2016) |
| Cabalodontia | C. queletii | Resupinate | Odontoid, tuberculate or smooth | Monomitic | Ellipsoid to subglobose | Piątek (2004) |
| Caudicicola | C. gracilis | Resupinate | Poroid | Monomitic | Broadly ellipsoid to subglobose | Kotiranta et al. (2017) |
| Citripora | C. bannaensis | Effused-reflexed to pileate | Poroid | Dimitic | Broadly ellipsoid to subglobose | Miettinen and Ryvarden (2016) |
| Etheirodon | E. fimbriatus | Resupinate to effused-reflexed | Odontoid to hydnoid | Dimitic | Ellipsoid to broadly ellipsoid | This study |
| Flabellophora | F. superposita | Pileate to stipitate | Poroid | Monomitic | Subglobose to globose | Cunningham (1965) |
| Flaviporus | F. brownii | Resupinate to effused-reflexed | Poroid | Dimitic | Ellipsoid, broadly ellipsoid to subglobose | Ryvarden and Gilbertson (1993); Wei et al. (2023) |
| Frantisekia | F. fissiliformis | Resupinate, effused-reflexed to pileate | Poroid | Dimitic | Oblong-ellipsoid | Spirin and Zmitrovich (2007); Yuan (2014) |
| Junghuhnia | J. crustacea | Resupinate to rarely effused-reflexed | Poroid | Dimitic | Cylindrical to ellipsoid | Ryvarden and Gilbertson (1993) |
| Lamelloporus | L. americanus | Pileate | Lamellae | Dimitic | Ellipsoid | Ryvarden (1987) |
| Loweomyces | L. fractipes | Resupinate, effused-reflexed to stipitate | Poroid | Monomitic | Broadly ellipsoid to subglobose | Kotlába and Pouzar (1976) |
| Metuloidea | M. tawa | Effused-reflexed to pileate | Poroid to hydnoid | Dimitic to trimitic | Ellipsoid to short cylindrical | Cunningham (1965) |
| Mycorrhaphium | M. adustum | Effused-reflexed, pileate to stipitate | Poroid to hydnoid | Dimitic | Cylindrical to ellipsoid | Maas Geesteranus (1971); Westphalen et al. (2019) |
| Niemelaea | N. consobrina | Resupinate | Poroid | Monomitic | Ellipsoid to broadly ellipsoid | Ryvarden and Gilbertson (1993); Zmitrovich et al. (2015) |
| Nigroporus | N. vinosus | Resupinate to pileate | Poroid | Dimitic | Allantoid | Ryvarden and Johansen (1980) |
| Rhomboidia | R. wuliangshanensis | Pileate | Poroid | Monomitic | Broadly ellipsoid | Xu et al. (2020) |
| Steccherinum | S. ochraceum | Resupinate, effused-reflexed to pileate or stipitate | Poroid, odontoid to corticioid | Dimitic | Ellipsoid to subcylindrical | Liu et al. (2023) |
| Trullella | T. dentipora | Pileate to stipitate | Poroid to irpicoid | Dimitic | Cylindrical and curved | Zmitrovich (2018) |
| Xanthoporus | X. peckianus | Pileate to stipitate | Poroid | Monomitic | Ellipsoid to subglobose | Audet (2010) |
Citripora encompasses the two species, C. afrocitrina (Ipulet & Ryvarden) Miettinen & Ryvarden and C. bannaensis Miettinen. It has effused-reflexed to pileate basidiomata with lemon-yellow tints, small pores, a dimitic hyphal system bearing clamp connections on generative hyphae, and tiny, broadly ellipsoid to subglobose basidiospores (Miettinen and Ryvarden 2016). Though Citripora grouped with Bambusiporia in a joint clade, the divergence time of them differs by nearly 40 Myr (Fig. 2). In addition, the Citripora clade is phylogenetically close to the Bambusiporia clade without strong support (Fig. 1). Polypore genera Flabellophora, Nigroporus, Rhomboidia, and Trullella have effused-reflexed to pileate or stipitate basidiomata (Cunningham 1965; Ryvarden and Johansen 1980; Miettinen and Ryvarden 2016; Xu et al. 2020), which differs from Bambusiporia. In addition, Flabellophora and Rhomboidia have broadly ellipsoid to subglobose basidiospores; Nigroporus has grayish-blue, vinaceous-brown to dark brown basidiomata, a dimitic hyphal system, and allantoid basidiospores; Trullella has a dimitic hyphal system and allantoid basidiospores, and these characteristics are also different from Bambusiporia. Mycorrhaphium is a pileate hydnoid genus that encompasses eleven species. Austeria is monotypic and is characterized by pileate basidiomata with sulfur-yellow tints, tiny pores, a dimitic hyphal system, and subcylindrical and thin-walled basidiospores (Miettinen and Ryvarden 2016), while Bambusiporia is a resupinate poroid genus.
Niemelaea features three species, viz., N. balaenae (Niemelä) V. Papp, N. consobrina (Bres.) Zmitr. et al., and N. cremea (Parmasto) Zmitr., Ezhov & Khimich. It has resupinate basidiomata with a light-colored pore surface, a monomitic hyphal system bearing clamp connections on generative hyphae, and ellipsoid to broadly ellipsoid basidiospores (Zmitrovich et al. 2015). These characteristics are similar to Bambusiporia. However, Niemelaea is unrelated to Bambusiporia in the phylogeny (Fig. 1).
Etheirodon fimbriatus (basionym: Odontia fimbriata Pers.) was originally described from France. It has pinkish basidiomata; an odontioid hymenophore, usually with a filamentous to rhizomorphic sterile margin; conical spines of 4–5 per mm at the base; strongly encrusted and thick-walled cystidia; and ellipsoid basidiospores (Eriksson et al. 1984). Etheirodon purpureus was originally described from Brazil. It is very similar to E. fimbriatus, but the former has smaller spines of 7–10 per mm at the base (Westphalen et al. 2021). Etheirodon fimbriatus and E. purpureus differ from the new species E. lilacinum by having an ochraceous to dark ochraceous hymenophore when dry, the presence of encrusted and thick-walled cystidia, and smaller basidiospores (3.5–4 × 2–2.5 µm in E. fimbriatus and 4–4.5 × 2–2.5 µm in E. purpureus vs. 4–4.5 × 3–3.8 µm in E. lilacinum, Westphalen et al. 2021). The two specimens, Dai 24450 and CLZhao 13977, were found in China, and their morphological characteristics are very similar to E. fimbriatus. The phylogenetic analysis suggests a close relatedness to E. fimbriatus. In this study, we treat them as Etheirodon cf. fimbriatus. In addition, specimens KUC 20121109-29 and HHB-2878-sp are treated as “Etheirodon cf. fimbriatus” as well because we did not study their voucher material. Etheirodon diverged in the Miocene with a mean crown age of 19.56 Myr. Etheirodon lilacinum diverged from E. fimbriatus s.l. during the late Miocene with a mean crown age of 6.03 Myr.
Supplementary Material
Citation
Wang C-G, Zhang X, Liu Z-B, Chen J, Sun Y-F, Dai Y-C, Wu Y-D (2026) The new genus Bambusiporia and a new species of Etheirodon in Steccherinaceae (Polyporales, Basidiomycota) from China. MycoKeys 130: 101–126. https://doi.org/10.3897/mycokeys.130.183355
Footnotes
Chao-Ge Wang and Xin Zhang contributed equally to this work.
Contributor Information
Yu-Cheng Dai, Email: yuchengdai@bjfu.edu.cn.
Ying-Da Wu, Email: wydbjfu@163.com.
Additional information
Conflict of interest
The authors have declared that no competing interests exist.
Ethical statement
No ethical statement was reported.
Use of AI
No use of AI was reported.
Funding
The research is supported by the National Natural Science Foundation of China (Project Nos. 32300013, U23A20142) and the Second Tibetan Plateau Scientific Expedition and Research Program (STEP, Grant No. 2019QZKK0503).
Author contributions
Chao-Ge Wang, Yu-Cheng Dai and Ying-Da Wu designed the research and contributed to data analysis and interpretation. Chao-Ge Wang conducted the molecular experiments and analyzed the data. Chao-Ge Wang, Jian Chen and Yi-Fei Sun prepared the samples. Chao-Ge Wang drafted the manuscript. Chao-Ge Wang, Xin Zhang, Zhan-Bo Liu, Yu-Cheng Dai discussed the results and edited the manuscript. All authors contributed to the article and approved the submitted version.
Author ORCIDs
Chao-Ge Wang https://orcid.org/0000-0003-4381-5720
Xin Zhang https://orcid.org/0009-0005-8363-7852
Zhan-Bo Liu https://orcid.org/0000-0002-3894-5398
Jian Chen https://orcid.org/0000-0001-8409-7791
Yi-Fei Sun https://orcid.org/0000-0003-3997-3662
Yu-Cheng Dai https://orcid.org/0000-0002-6523-0320
Ying-Da Wu https://orcid.org/0000-0003-1295-4015
Data availability
All of the data that support the findings of this study are available in the main text or Supplementary Information.
Supplementary materials
Phylogenetic analyses
This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Chao-Ge Wang, Xin Zhang, Zhan-Bo Liu, Jian Chen, Yi-Fei Sun, Yu-Cheng Dai, Ying-Da Wu
Data type
rar
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Associated Data
This section collects any data citations, data availability statements, or supplementary materials included in this article.
Supplementary Materials
Phylogenetic analyses
This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Chao-Ge Wang, Xin Zhang, Zhan-Bo Liu, Jian Chen, Yi-Fei Sun, Yu-Cheng Dai, Ying-Da Wu
Data type
rar
Data Availability Statement
All of the data that support the findings of this study are available in the main text or Supplementary Information.






