An Updated Phylogenetic Assessment and Taxonomic Revision of Perenniporia sensu lato (Polyporales, Basidiomycota)

Abstract

Perenniporia is an important genus of Polyporaceae. In its common acceptation, however, the genus is polyphyletic. In this study, phylogenetic analyses on a set of Perenniporia species and related genera were carried out using DNA sequences of multiple loci, including the internal transcribed spacer (ITS) regions, the large subunit nuclear ribosomal RNA gene (nLSU), the small subunit mitochondrial rRNA gene (mtSSU), the translation elongation factor 1-α gene (TEF1) and the b-tubulin gene (TBB1). Based on morphology and phylogeny, 15 new genera, viz., Aurantioporia, Citrinoporia, Cystidioporia, Dendroporia, Luteoperenniporia, Macroporia, Macrosporia, Minoporus, Neoporia, Niveoporia, Rhizoperenniporia, Tropicoporia, Truncatoporia, Vanderbyliella, and Xanthoperenniporia, are proposed; 2 new species, Luteoperenniporia australiensis and Niveoporia subrusseimarginata, are described; and 37 new combinations are proposed. Illustrated descriptions of the new species are provided. Identification keys to Perenniporia and its related genera and keys to the species of these genera are provided.

1. Introduction

Perenniporia Murrill, typified by P. medulla-panis (Jacq.) Donk, is a large and cosmopolitan genus. The genus has been redefined by Decock and Stalpers [1], who designated a neotype. Species in this genus have annual to perennial, resupinate to pileate basidiocarps, a dimitic to trimitic hyphal system with clamped generative hyphae, cyanophilous and variably dextrinoid skeletal hyphae, and thick-walled, ellipsoid to subglobose, truncate or not, cyanophilous, and variably dextrinoid basidiospores [2,3].

In the last few decades, Perenniporia has been intensively studied and the number of species has increased significantly. Many new species were published based on morphological characters [1,4,5,6,7,8,9,10,11,12,13,14] or based on morphological characters and phylogenetic analysis [15,16,17,18,19,20,21,22,23,24,25,26,27,28,29].

These multiple additions considerably enlarged the morphological concept of Perenniporia. A recent trend is to make the morphologically distinct species or morphologically homogeneous alliance separate from Perenniporia. For instance, Perenniporiella Decock & Ryvarden [30] was separated from Perenniporia based on non-truncate basidiospores. This was later confirmed by Robledo et al. [31] by phylogenetic data. Hornodermoporus, Truncospora, and Vanderbylia are usually treated as synonyms of Perenniporia [2,32,33,34], but have been shown by phylogenetic studies to also represent clades distinct from the Perenniporia s. s. clade, each morphologically homogenous [3,17,19,20,26,31].

The current phylogenetic analysis confirmed that Perenniporia is polyphyletic, and some monophyletic clades have been separated from Perenniporia s. l. and recognized as independent genera [3,35,36,37]. Based on such characters as the acyanophily and amyloidity of skeletal hyphae dissolving in KOH, as well as the non-dextrinoidity of basidiospores and the phylogenetic data from the rDNA sequences of ITS and nLSU, P. narymica (Pilát) Pouzar was segregated into Yuchengia B.K. Cui & K.T. Steffen [35]. Wu et al. [36] also separated Perenniporia minutissima (Yasuda) T. Hatt. & Ryvarden from Perenniporia as Perenniporiopsis minutissima (Yasuda) C.L. Zhao based on the phylogenetic analyses, as well as the morphological differences of rigidly osseous basidiocarps when there are dry and large basidiospores. Cui et al. [3] used multiple genes of ITS, nLSU, nSSU, mtSSU, TEF1, TBB1, RPB1, and RPB2 and proposed the separation of Perenniporia hattorii Y.C. Dai and B.K. Cui from Perenniporia into Amylosporia B.K. Cui, C.L. Zhao & Y.C. Dai because of its amyloid skeletal hyphae and basidiospores, and P. subadusta (Z.S. Bi & G.Y. Zheng) Y.C. Dai from Perenniporia into Murinicarpus B.K. Cui & Y.C. Dai on account of its stipitate basidiocarps and cystidia in the hymenium. Chen et al. [37] pointed out that Perenniporia subacida (Peck) Donk formed a well-supported lineage that is distinct from the Perenniporia s. s. clade and proposed that P. subacida should be treated in a new genus named Poriella C.L. Zhao.

Although some monophyletic lineages have been assigned a new genus name (for example, Amylosporia, Murinicarpus, Perenniporiopsis, Poriella, and Yuchengia [3,35,36,37]), the positions of other Perenniporia species are still doubtful, and some species should be separated from Perenniporia. In order to clarify the taxonomy and phylogeny of Perenniporia s. l., reliable specimens and sequences were studied using morphological methods and phylogenetic analyses of ITS, nLSU, mtSSU, TEF1, and TBB1.

2. Materials and Methods

2.1. Morphological Studies

The specimens examined in this study are deposited in the herbaria of the Institute of Microbiology, Beijing Forestry University, China (BJFC) and the Institute of Applied Ecology, Chinese Academy of Sciences, China (IFP). Macro-morphological descriptions were based on field notes. Special color terms followed the Petersen protocol [38]. Micro-morphological data were obtained from dried specimens and observed under a light microscope according to Sun et al. [39] and Liu et al. [40]. Sections were studied at a magnification up to 1000× using a Nikon Eclipse 80i microscope and phase contrast illumination (Nikon, Tokyo). Drawings were made with the aid of a drawing tube. Microscopic features, measurements, and drawings were made from slide preparations stained with Cotton Blue and Melzer’s reagent. Spores were measured from sections cut from the tubes. To represent variation in the size of spores. Five percent of measurements were excluded from each end of the range and given in parentheses. The following abbreviations are used: IKI = Melzer’s reagent, IKI− = non-dextrinoid and inamyloid, KOH = 5% potassium hydroxide, CB = Cotton Blue, CB+ = cyanophilous, L = mean spore length (arithmetic average of all spores), W = mean spore width (arithmetic average of all spores), Q = variation in the L/W ratios between the specimens studied, and n = number of spores (a) measured from given number (b) of specimens.

2.2. DNA Extraction, Amplification and Sequencing

A CTAB rapid plant genome extraction kit-DN14 (Aidlab Biotechnologies Co., Ltd., Beijing, China) was used to extract total genomic DNA from dried specimens and to perform the polymerase chain reaction (PCR) according to the manufacturer’s instructions, with some modifications [3,41]. The ITS regions were amplified with the primer pairs ITS5 and ITS4 [42]. The nLSU regions were amplified with the primer pairs LR0R and LR7 (http://www.biology.duke.edu/fungi/mycolab/primers.htm, accessed on 12 October 2021). The mtSSU regions were amplified with the primer pairs MS1 and MS2 [42]. Part of TEF1 was amplified with the primer pairs EF1-983F and EF1-1567R [43]. TBB1 was amplified with the primer pairs Bt-1a and Bt-1b [44]. The PCR procedures for different DNA sequences were the same as those used by Cui et al. [3]. The PCR products were purified and sequenced at the Beijing Genomics Institute (BGI), China, with the same primers. All newly generated sequences and additional sequences downloaded from GenBank are listed in Table 1.

Table 1.

Samples and GenBank accession numbers of sequences used in this study.

Species	Sample no.	Locality	GenBank Accession No.
			ITS	nLSU	mtSSU	TEF1	TBB1
Abortiporus biennis	FD-319	the United States of America	KP135300	KP135195	—	—	—
Abundisporus pubertatis	Dai 11927	China	KC787569	KC787576	KF051034	KF181128	KF482828
Abundisporus sclerosetosus	MUCL 41438	Singapore	FJ411101	FJ393868	—	—	—
Abundisporus violaceus	Ryvarden 32807	Zimbabwe	KF018127	KF018135	KF051038	KF181132	KF482832
Amylosporia hattorii	Cui 10912	China	KX900675	KX900725	—	KX900852	—
Amylosporia hattorii	Dai 10315	China	JQ861740	JQ861756	KF218290	—	—
Antrodia albida	FP-105979	Unknown	EU232189	EU232272	—	—	—
Antrodia serialis	FP-133692	the United States of America	KC585303	KC585127	—	—	—
Aurantioporia aurantiaca	CBS 125867	French Guiana	MH863779	MH875242	—	—	—
Aurantioporia bambusicola	Cui 11050	China	KX900668	KX900719	KX900771	OK665240	—
Aurantioporia bambusicola	Yuan 3925	China	JQ861736	—	—	—	—
Bjerkandera adusta	HHB-12826-Sp	the United States of America	KP134983	KP135198	—	—	—
Cerrena unicolor	FD-299	the United States of America	KP135304	KP135209	—	—	—
Citrinoporia citrinoalba	Cui 13615	China	MG847215	MG847224	MG847238	MG867708	—
Citrinoporia citrinoalba	Dai 13643	China	KX880622	KX880661	KX880705	—	—
Citrinoporia corticola	Dai 18633	Malaysia	MT117217	MT117222	OK642017	OK665241	—
Citrinoporia corticola	Dai 18641	Malaysia	MT117218	MT117223	—	—	—
Climacocystis borealis	FD-31	the United States of America	KP135308	KP135210	—	—	—
Coriolopsis dendriformis	Cui 6719	China	KC867408	KC867445	—	—	—
Coriolopsis retropicta	Cui 13849	China	MK116481	MK116490	—	—	—
Crassisporus imbricatus	Dai 10788	China	KC867350	KC867425	—	—	—
Crassisporus macroporus	Cui 14468	China	MK116486	MK116495	—	—	—
Cryptoporus sinensis	Cui 17418	China	OK642180	OK642231	—	—	—
Cryptoporus volvatus	CBS 432.48	Canada	MH856424	MH867970	—	—	—
Cystidioporia piceicola	Cui 10460	China	JQ861742	JQ861758	KF218310	KF286316	KF482803
Cystidioporia piceicola	Dai 4181	China	JF706328	JF706336	KF218311	KF286317	KF482802
Daedalea quercina	Dai 12659	Finland	KP171208	KP171230	KR605990	KR610719
Dendroporia cinereofusca	Cui 5280	China	KF568892	KF568894	—	—	—
Dendroporia cinereofusca	Dai 9289	China	KF568893	KF568895	—	—	—
Dichomitus squalens	Cui 9639	China	JQ780407	JQ780426	—	—	—
Dichomitus squalens	CBS 432.34	Poland	MH855594	MH867104	—	—	—
Donkioporia expansa	MUCL 35116	Belgium	FJ411104	FJ393872	—	—	—
Earliella scabrosa	PR1209	Puerto Rico	JN165009	JN164793	—	—	—
Fomitopsis pinicola	Cui 10405	China	KC844852	KC844857	KR605961	KR610690	—
Fragiliporia fragilis	Dai 13080	China	KJ734260	KJ734264	—	—	—
Fragiliporia fragilis	Dai 13559	China	KJ734261	KJ734265	—	—	—
Gelatoporia subvermispora	BRNU 592909	Czech Republic	FJ496694	FJ496706	—	—	—
Grammothele lineata	Cui 6539	China	KX832049	KX832058	—	—	—
Grammothelopsis subtropica	Cui 9035	China	JQ845094	JQ845097	—	—	—
Grammothelopsis subtropica	Cui 9041	China	JQ845096	JQ845099	—	—	—
Haploporus odorus	Dai 11296	China	KU941845	KU941869	—	—	—
Haploporus tuberculosus	15559	Sweden	KU941857	KU941881	—	—	—
Heterobasidion annosum	VL296	Lithuania	JF440572	—	—	—	—
Hornodermoporus latissimus	Dai 12054	China	KX900639	KX900686	KF218297	KF286303	KF482789
Hornodermoporus martius	Cui 7992	China	HQ876603	HQ654114	KF051041	KF181135	KF482835
Irpex lacteus	FD-9	the United States of America	KP135026	KP135224	—	—	—
Jorgewrightia guangdongensis	Cui 9130	China	JQ314373	JQ780428	—	—	—
Jorgewrightia tropica	Cui 13740	China	KY449438	KY449449	—	—	—
Lentinus crinitus	DSH92N43C	Costa Rica	KP283495	KP283523	—	—	—
Luteoperenniporia australiensis	Cui 16742	Australia	OK642220	OK642275	OK642018	OK665242	—
Luteoperenniporia australiensis	Cui 16743	Australia	OK642221	OK642276	OK642019	OK665243	OK665272
Luteoperenniporia bannaensis	Cui 8560	China	JQ291727	JQ291729	KF218280	KF286286	KF482772
Luteoperenniporia bannaensis	Cui 8562	China	JQ291728	JQ291730	KF218281	KF286287	KF482773
Luteoperenniporia mopanshanensis	CLZhao 2404	China	MH784911	MH784915	—	—	—
Luteoperenniporia mopanshanensis	CLZhao 5145	China	MH784912	MH784916	—	—	—
Luteoperenniporia yinggelingensis	Cui 13625	China	MH427960	MH427967	MH427975	MH427996	—
Luteoperenniporia yinggelingensis	Cui 13627	China	MH427961	MH427968	MH427976	MH427997	—
Macroporia lacerata	Cui 7220	China	JX141448	JX141458	KF218295	KF286301	KF482787
Macroporia lacerata	Dai11268	China	JX141449	JX141459	KF218296	KF286302	KF482788
Macroporia macropora	Zhou 280	China	JQ861748	JQ861764	KF494992	KF482765	KF482798
Macroporia macropora	Zhou 407	China	JQ861746	JQ861762	KF494991	KF482767	KF482784
Macroporia subrhizomorpha	LWZ 20190722-36	China	MZ578440	MZ578444	—	—	—
Macroporia subrhizomorpha	LWZ 20190723-10	China	MZ578441	—	—	—	—
Macroporia tibetica	Cui 9457	China	JF706326	JF706332	KF218332	KF286297	KF482782
Macroporia tibetica	Cui 9459	China	JF706327	JF706333	KF218333	KF286296	KF482783
Macrosporia nanlingensis	Cui 7541	China	HQ848479	HQ848488	KF218306	KF286312	—
Macrosporia nanlingensis	Cui 7620	China	HQ848477	HQ848486	KF218307	KF286313	—
Microporellus subadustus	Cui 8459	China	HQ876606	HQ654113	—	—	—
Microporellus subadustus	Dai 13895	China	KX880621	KX880660	—	KX880879	KX880780
Minoporus minor	Dai 9198	China	KF495005	KF495016	KF494994	—	KF494969
Minoporus minor	Cui 5782	China	HQ883475	—	KF218300	—	KF494968
Neoporia bostonensis	CLZhao 144	the United States of America	MG491283	MG491286	—	—	—
Neoporia bostonensis	CLZhao 2854	the United States of America	MG491284	MG491287	—	—	—
Neoporia koreana	KUC20080517-02	Republic of Korea	KJ156308	KJ156300	—	—	—
Neoporia koreana	KUC20080517-15	Republic of Korea	KJ156309	KJ156301	—	—	—
Neoporia rhizomorpha	Dai 7248	China	JF706330	JF706348	KF218315	KF286321	KF482807
Neoporia rhizomorpha	Cui 7507	China	HQ654107	HQ654117	KF218314	KF286320	KF482806
Niveoporia decurrata	Dai 16637	Thailand	KY475566	OP289291	—	OP296862	—
Niveoporia decurrata	Dai 16660	Thailand	KY475567	OP289292	—	OP296863	—
Niveoporia russeimarginata	Yuan 1244	China	JQ861750	JQ861766	KF218316	KF286322	KF482808
Niveoporia russeimarginata	Yuan 1262	China	JQ861751	JQ861767	KF218317	KF286323	KF482809
Niveoporia subrusseimarginata	Cui 16980	China	OK642223	OK642278	—	OK665245	—
Niveoporia subrusseimarginata	Cui 16991	China	OK642224	OK642279	—	OK665246	OK665273
Obba valdiviana	Gates FF503	Australia	HQ659235	HQ659235	—	—	—
Perenniporia eugeissonae	Dai 18600	Malaysia	MT232518	MT232512	—	—	—
Perenniporia eugeissonae	Dai 18605	Malaysia	MT232519	MT232513	—	—	—
Perenniporia hainaniana	Cui 6364	China	JQ861743	JQ861759	KF051044	KF181138	KF482838
Perenniporia hainaniana	Cui 6365	China	JQ861744	JQ861760	KF051045	KF181139	KF482839
Perenniporia medulla-panis	Cui 14515	China	MG847214	MG847223	—	MG867707	MG867711
Perenniporia medulla-panis	MUCL 43250	Norway	FJ411087	FJ393875	—	—	—
Perenniporia straminea	Cui 8718	China	HQ876600	JF706335	KF218318	KF286324	KF482810
Perenniporia straminea	Cui 8858	China	HQ654104	JF706334	KF218319	KF286325	KF482811
Perenniporia substraminea	Cui 10177	China	JQ001852	JQ001844	KF051046	KF181140	KF482840
Perenniporia substraminea	Cui 10191	China	JQ001853	JQ001845	KF051047	KF181141	KF482841
Perenniporia subtephropora	Dai 10962	China	JQ861752	JQ861768	KF218323	KF286329	KF482815
Perenniporia subtephropora	Dai 10964	China	JQ861753	JQ861769	KF218324	KF286330	KF482816
Perenniporia tephropora	Cui 8040	China	JN048763	HQ654118	KF218328	KF286307	KF482793
Perenniporia tephropora	Cui 9029	China	HQ876601	JF706339	KF218327	KF286306	KF482792
Perenniporiella chaquenia	MUCL 47647	Argentina	FJ411083	FJ393855	—	HM467609	—
Perenniporiella micropora	MUCL 43581	Cuba	FJ411086	FJ393858	—	HM467608	—
Perenniporiella pendula	MUCL 46034	Cuba	FJ411081	FJ393853	—	HM467601	—
Perenniporiopsis minutissima	Dai 12457	China	KF495004	KF495014	KF218302	KF286308	KF482794
Perenniporiopsis minutissima	Cui 10979	China	KF495003	KF495013	KF218304	KF286310	KF482796
Phanerochaete carnosa	HHB-9195-Sp	the United States of America	KP135129	KP135242	—	—	—
Phlebia radiata	AFTOL-484	Unknown	AY854087	AF287885	—	—	—
Piloporia sajanensis	HK 17250 (GB)	Russia	JX109853	JX109853	—	—	—
Polyporus tuberaster	Dai 12462	China	KU507580	KU507582	—	—	—
Poriella africana	Cui 8674	China	KF018119	KF018128	KF218276	KF286282	KF482768
Poriella africana	Cui 8676	China	KF018120	KF018129	KF218277	KF286283	KF482769
Poriella ellipsospora	Cui 10276	China	KF018124	KF018132	KF218286	KF286292	KF482778
Poriella ellipsospora	Cui 10284	China	JQ861739	KF018133	KF218285	KF286291	KF482777
Poriella subacida	Cui 10053	China	KF495006	KF495017	KF218321	KF286327	KF482813
Poriella subacida	DLL2010-125	the United States of America	JQ673016	—	—	—	—
Poriella valliculorum	LE 222974	Russia	KM411458	KM411474	—	KM411489	—
Pyrofomes demidoffii	PRM869997	Macedonia	KY940246	KY940259	—	—	—
Rhizoperenniporia japonica	Cui 7047	China	KX900677	KX900727	KF218294	KF286300	KF482786
Rhizoperenniporia japonica	Cui 9181	China	JQ001856	JX141468	KF218293	KF286299	—
Sebipora aquosa	Miettinen 8680.1	Indonesia	HQ659240	HQ659240	—	—	—
Skeletocutis amorpha	Miettinen 11038.1	Finland	FN907913	FN907913	—	—	—
Sparsitubus nelumbiformis	Cui 8497	China	KX880631	KX880670	KX880714	KX880887	KX880786
Sparsitubus nelumbiformis	Cui 6590	China	KX880632	KX880671	KX880715	KX880888	—
Steccherinum laeticolor	FP-102480-sp	the United States of America	KY948823	KY948868	—	—	—
Stereum hirsutum	NBRC 6520	Unknown	AY854063	AF393078	—	—	—
Trametes elegans	FP-105679-Sp	the United States of America	JN164944	JN164799	—	—	—
Trametes hirsuta	RLG-5133-T	the United States of America	JN164941	JN164801	—	—	—
Trametes polyzona	BKW004	Ghana	JN164978	JN164790	—	—	—
Trametes suaveolens	FP-102529-Sp	the United States of America	JN164966	JN164807	—	—	—
Trametes versicolor	FP-135156-Sp	the United States of America	JN164919	JN164809	—	—	—
Tropicoporia aridula	Dai 12396	China	JQ001854	JQ001846	KF218278	KF181158	—
Tropicoporia aridula	Dai 12398	China	JQ001855	JQ001847	KF218279	KF286285	KF482771
Tropicoporia brasiliensis	AN-MA10	Brazil	KX584437	KX619594	—	—	—
Tropicoporia brasiliensis	URM89948	Brazil	KX584436	—	—	—	—
Tropicoporia centrali-africana	URM 82624	Brazil	KX584433	KX619599	—	—	—
Tropicoporia centrali-africana	URM 82568	Brazil	KX584432	KX619598	—	—	—
Tropicoporia vanhulleae	MUCL 46315	Senegal	KP217810	—	—	—	—
Tropicoporia vanhulleae	MUCL 38450	Zimbabwe	KP217811	—	—	—	—
Truncatoporia pyricola	Cui 9149	China	JN048762	JN048782	—	KF286318	KF482804
Truncatoporia pyricola	Dai 10265	China	JN048761	JN048781	—	KF286319	KF482805
Truncatoporia truncatospora	Cui 6987	China	JN048778	HQ654112	KF218334	KF286288	KF482774
Truncatoporia truncatospora	Dai 5125	China	HQ654098	HQ848481	KF218335	KX880880	KF482770
Truncospora macrospora	Cui 8106	China	JX941573	JX941596	KX880763	KX880920	KX880809
Truncospora ochroleuca	Cui 16795	Australia	OK642218	OK642273	OK642015	OK665238	—
Truncospora ochroleuca	JV0610/7B	Belize	KJ410698	—	—	KJ410718	—
Truncospora ohiensis	MUCL 41036	the United States of America	FJ411096	FJ393863	—	—	—
Truncospora ornata	SP6672	Russia	KJ410690	—	—	KJ410712	—
Tyromyces chioneus	FD-4	the United States of America	KP135311	KP135291	—	—	—
Vanderbylia cinnamomea	CLZhao 8952	China	MT372778	MT372788	—	—	—
Vanderbylia delavayi	Dai 6891	China	JQ861738	—	KF218287	KF286293	KF482779
Vanderbylia fraxinea	Cui 8871	China	JF706329	JF706345	KF051050	KF181144	KF482844
Vanderbylia vicina	MUCL 44779	Ethiopia	FJ411095	FJ393862	—	—	—
Vanderbyliella sp.	Knudsen 04-111	China	JQ861737	JQ861755	KF218282	—	KF494963
Vanderbyliella tianmuensis	Cui 2648	China	JX141453	JX141463	KF218329	—	KF494971
Vanderbyliella tianmuensis	Cui 2715	China	JX141454	JX141464	KF218331	—	KF494972
Xanthoperenniporia maackiae	Dai 8929	China	HQ654102	JF706338	KF218299	KF286305	KF482791
Xanthoperenniporia maackiae	Cui 10092	China	KX900680	KX900730	KX900780	KX900856	—
Xanthoperenniporia punctata	Dai 17916	China	MG869686	MG869688	—	—	—
Xanthoperenniporia punctata	Dai 17923	China	MG869687	MG869689	OP293085	OP296864	—
Xanthoperenniporia subcorticola	Cui 1248	China	HQ848472	HQ848482	KF218284	KF286290	KF482776
Xanthoperenniporia subcorticola	Dai 7330	China	HQ654094	HQ654108	KF218283	KF286289	KF482775
Xanthoperenniporia tenuis	Wei 2783	China	JQ001858	JQ001848	KF218325	KF286331	KF482817
Xanthoperenniporia tenuis	Wei 2969	China	JQ001859	JQ001849	KF218326	KF286332	KF482818
Yuchengia kilemariensis	LE 214743	Russia	KM411457	KM411473	—	KM411488	—
Yuchengia narymica	Dai 7050	China	JN048776	JN048795	KF051053	KF181147	KF482836
Yuchengia narymica	Dai 10510	China	HQ654101	JF706346	KF051054	KF181148	KF482833

New sequences are shown in bold.

2.3. Phylogenetic Analyses

Heterobasidion annosum (Fr.) Bref. and Stereum hirsutum (Willd.) Pers. were used as outgroups for the ITS + nLSU analysis [45], Fomitopsis pinicola (Sw.) P. Karst. and Daedalea quercina (L.) Pers. were selected as outgroups for the ITS + nLSU + mtSSU + TEF1 + TBB1 analysis. All sequences were aligned in MAFFT 7 [46] (http://mafft.cbrc.jp/alignment/server/, accessed on 11 June 2022) and manually adjusted in BioEdit [47]. Maximum Likelihood (ML) and Bayesian Inference (BI) phylogenetic analyses were performed as described by Sun et al. [48]. ML studies and Bayesian inference were applied to the ITS + nLSU and ITS + nLSU + mtSSU + TEF1 + TBB1 datasets.

ML studies were conducted with RAxML-HPC through the Cipres Science Gateway (https://www.phylo.org, accessed on 12 June 2022) and involved 1000 ML searches under the GTRGAMMA model. Only the maximum likelihood best tree from all searches was kept. In addition, 1000 rapid bootstrap replicates were run with the GTRCAT model to assess the reliability of the nodes.

BI was performed with MrBayes v3.1.2 (Ronquist and Huelsenbeck, Sweden) [49]. Four Markov chains were run from random starting trees for 6 million generations for ITS + nLSU and for 13 million generations for ITS + nLSU + mtSSU + TFF1 + TBB1, and trees were sampled every 100th generation. The first 25% of trees were discarded as burn-in, and the remaining trees were used to calculate Bayesian posterior probabilities (BPP) of the clades.

Trees were viewed in FigTree v1.4.2 (http://tree.bio.ed.ac.uk/software/figtree/, accessed on 12 June 2022). Branches that received bootstrap support for maximum likelihood (BS) and Bayesian posterior probabilities (BPP) ≥ 50% (BS) and ≥0.90 (BPP) were considered significantly supported. The final concatenated sequence alignment and the retrieved topology were deposited in TreeBase (http://purl.org/phylo/treebase, accessed on 29 November 2022; submission ID: 29932).

3. Results

3.1. Phylogeny

The combined ITS + nLSU dataset included sequences from 159 fungal samples representing 108 taxa. The dataset had an aligned length of 2215 characters, of which 1257 characters were constant, 235 were variable and parsimony-uninformative, and 723 were parsimony-informative. BI analysis generated topologies similar to those of ML analysis, with an average standard deviation of split frequencies at 0.004614 (BI). The topology from the ML analysis with a maximum likelihood bootstrap (BS) ≥ 50% and Bayesian posterior probabilities (BPP) ≥ 0.90 labeled on branches is shown (Figure 1).

Figure 1.

Maximum likelihood tree illustrating the phylogeny of Perenniporia and related species in Polyporales based on the ITS + nLSU dataset. Branches are labeled with maximum likelihood bootstrap support values (≥50%) and Bayesian posterior probabilities (≥0.90).

The combined five-gene (ITS, nLSU, mtSSU, TEF1, TBB1) sequence dataset included sequences from 119 fungal samples representing 70 taxa. The dataset had an aligned length of 3498 characters, of which 2285 characters were constant, 226 were variable and parsimony-uninformative, and 987 were parsimony-informative. BI analysis generated topologies similar to those of ML analysis, with an average standard deviation of split frequencies = 0.007253 (BI). The topology from the ML analysis with a maximum likelihood bootstrap (BS) ≥ 50% and Bayesian posterior probabilities (BPP) ≥ 0.90 labeled on branches is shown (Figure 2).

Figure 2.

Maximum likelihood tree illustrating the phylogeny of Perenniporia and related genera based on the ITS + nLSU + mtSSU + TEF1 + TBB1 dataset. Branches are labeled with maximum likelihood bootstrap support values (≥50%) and Bayesian posterior probabilities (≥0.90).

3.2. Taxonomy

Aurantioporia B.K. Cui & Xing Ji, gen. nov.

MycoBank: MB 847338

Differs from other genera by its resupinate, rhizomorphic basidiocarps with an orange pore surface, a dimitic hyphal system with arboriform skeletal hyphae, tissues becoming violet in KOH, ellipsoid, truncate, and slightly dextrinoid basidiospores.

Type species: Aurantioporia bambusicola (Choeyklin, T. Hatt. & E.B.G. Jones) B.K. Cui & Xing Ji

Etymology: Aurantioporia (Lat.) refers to the orange pore surface of the genus.

Basidiocarps are annual to perennial and resupinate with rhizomorphs. The pore surface is yellow to orange when fresh, grayish orange, and orange-brown to dark orange when dry; pores are round to angular; dissepiments thin, entire. The subiculum is extremely thin and cream to pale orange. The tubes are concolorous with pore surface. The hyphal system is dimitic; generative hyphae with clamp connections; skeletal hyphae arboriform, IKI−, CB+; tissues become violet to dark in KOH. Basidiospores are ellipsoid, truncate, hyaline, thick-walled, smooth, slightly dextrinoid, and CB+.

Notes: In our study, P. aurantiaca (A. David & Rajchenb.) Decock & Ryvarden and P. bambusicola Choeyklin, T. Hatt. & E.B.G. Jones formed a single well-supported clade (82% BS, 1.00 BPP, Figure 1; 89% BS, 1.00 BPP, Figure 2), distant from the Perenniporia s. s. clade. Morphologically, the two species differ from Perenniporia s. s. species by the combination of a rhizomorphic basidiocarp with an orange pore surface, a dimitic hyphal system with arboriform skeletal hyphae, and tissues becoming violet in KOH. Therefore, Aurantioporia gen. nov. is proposed to include Perenniporia aurantiaca and P. bambusicola.

Aurantioporia aurantiaca (A. David & Rajchenb.) B.K. Cui & Xing Ji, comb. nov.

MycoBank: MB 847362

Basionym: Pyrofomes aurantiacus A. David & Rajchenb., Mycotaxon 22(2): 312 (1985).

≡ Perenniporia aurantiaca (A. David & Rajchenb.) Decock & Ryvarden, Mycol. Res. 103(9): 1140 (1999).

For a detailed description of Perenniporia aurantiaca, see David and Rajchenberg [50] and Decock and Ryvarden [4].

Notes: Aurantioporia aurantiaca was originally described in Pyrofomes by David and Rajchenberg [50] and later was transferred to Perenniporia by Decock and Ryvarden [4]. The sequence of Aurantioporia aurantiaca from French Guyana (type locality) fell into the Aurantioporia clade in our phylogeny.

Aurantioporia bambusicola (Choeyklin, T. Hatt. & E.B.G. Jones) B.K. Cui & Xing Ji, comb. nov.

MycoBank: MB 847363

Basionym: Perenniporia bambusicola Choeyklin, T. Hatt. & E.B.G. Jones, Fungal Diversity 36: 122 (2009).

For a detailed description of Perenniporia bambusicola, see Choeyklin et al. [51] and Cui et al. [3].

Notes: Aurantioporia bambusicola was first described in Perenniporia from Thailand [51]. It is characterized by resupinate basidiocarps with an orange pore surface, a dimitic hyphal system with non-dextrinoid skeletal hyphae, tissues becoming violet to dark in KOH, oblong-ellipsoid, and truncate basidiospores. This species only grows on bamboo. Aurantioporia aurantiaca also shares an orange pore surface, but Aurantioporia aurantiaca grows on hardwood trees [51].

Specimen examined: CHINA. Yunnan, Cangyuan County, Banlao, on bamboo, 11 July 2013, Cui 11050 (BJFC).

Key to species of Aurantioporia

• 1. Growing on bamboo; distributed in Southeast Asia ………………………………………………………………………………………………… A. bambusicola

• 1. Growing on other hardwoods; distributed in neotropical areas………………………………………………………………………………………A. aurantiaca

Citrinoporia B.K. Cui & Xing Ji, gen. nov.

MycoBank: MB 847346

Differs from other genera by its slightly cushion shape, yellow pore surface, a dimitic hyphal system with dextrinoid, and cyanophilous shortly arboriform vegetative hyphae and ellipsoid, truncate, thick-walled, dextrinoid, and cyanophilous basidiospores.

Type species:

Etymology: Citrinoporia (Lat.) refers to the yellowish pore surface of the genus.

Basidiocarps are annual to perennial and resupinate. The pore surface is white to yellow; pores are round. Subiculum is cream to buff, corky. Tubes are buff to pale brown and corky to hard corky. The hyphal system is dimitic: generative hyphae with clamp connections; skeletal hyphae arboriform, IKI−, CB+; tissues becoming pale brown to black in KOH. Cystidia is absent, cystidioles are present. Basidiospores are ellipsoid, truncate, hyaline, thick-walled, smooth, dextrinoid, and CB+.

Notes: In our ITS + nLSU and five-gene phylogenetic analyses, P. citrinoalba B.K. Cui, C.L. Zhao & Y.C. Dai and P. corticola (Corner) Decock clustered together and formed a clade distinct from the Perenniporia s. s clade with full support (100% BS, 1.00 BPP, Figure 1; 100% BS, 1.00 BPP, Figure 2). Morphologically, this clade differs from the Perenniporia s. s. by its yellow pore surface and dimitic hyphal system. Thus, the new genus is set up and these two new combinations are proposed. Based on five-gene phylogenetic analysis, Citrinoporia is sister to Aurantioporia as they have the same overall morphology, yellow pore surface, dimitic hyphal system, and ellipsoid, truncate basidiospores, but the former differs in absence of rhizomorphs.

Citrinoporia citrinoalba (B.K. Cui, C.L. Zhao & Y.C. Dai) B.K. Cui & Xing Ji, comb. nov.

MycoBank: MB 847364

Basionym: Perenniporia citrinoalba B.K. Cui, C.L. Zhao & Y.C. Dai, Fungal Diversity 97: 270 (2019)

For a detailed description of Perenniporia citrinoalba, see Cui et al. [3].

Notes: Citrinoporia citrinoalba was newly described in Perenniporia from tropical China [2]. It is characterized by annual and resupinate basidiocarps with white to yellow pore surfaces, a dimitic hyphal system with dextrinoid and cyanophilous skeletal hyphae, tissues becoming pale brown to black in KOH, and broadly ellipsoid and truncate basidiospores.

Specimens examined: CHINA. Hainan, Qiongzhong County, Limushan Forest Park, on fallen trunk of Castanopsis, 15 June 2014, Dai 13643 (holotype, BJFC), on fallen angiosperm trunk, 18 November 2015, Cui 13615 (BJFC).

Citrinoporia corticola (Corner) B.K. Cui & Xing Ji, comb. Nov.

MycoBank: MB 847365

Basionym: Parmastomyces corticola Corner, Beih. Nova Hedwigia 96: 96 (1989).

≡ Perenniporia corticola (Corner) Decock, Mycologia 93(4): 776 (2001).

= Perenniporia dipterocarpicola T. Hatt. & S.S. Lee, Mycologia 91(3): 525 (1999)

Notes: Citrinoporia corticola was originally described in Parmastomyces Kotl. & Pouzar from Malaysia by Corner [52] as having a monomitic hyphal system with simple-septate generative hyphae. Decock [53] studied the type specimens of Parmastomyces corticola and confirmed that this species has a dimitic hyphal system with clamped generative hyphae and transferred the species to Perenniporia. Citrinoporia corticola and Citrinoporia citrinoalba share yellow pore surfaces, dimitic hyphal structures, and truncate basidiospores, but the basidiospores of C. citrinoalba (5.5–6 × 4.7–5.2 µm) [3] are larger than those of Citrinoporia corticola (4.4–5 × 3.4–4 μm) [53].

Specimens examined: MALAYSIA. Selangor, Kota Damansara, Community Forest Reserve, on angiosperm stump, 17 April 2018 Dai 18633, 18641 (BJFC).

Key to species of Citrinoporia

• 1. Basidiospores 4.4–5 μm; growing mainly on trees of Dipterocarpaceae……………………………………………………………………………… C. corticola

• 1. Basidiospores 5.5–6 µm; growing on trees of Fagaceae ……………………………………………………………………………………………… C. citrinoalba

Cystidioporia B.K. Cui & Xing Ji, gen. nov.

MycoBank: MB 847348

Differs from other genera by its resupinate basidiocarps, slightly dextrinoid and cyanophilous skeletal hyphae, presence of thick-walled cystidia, and thick-walled, oblong-ellipsoid, truncate, slightly dextrinoid, and cyanophilous basidiospores.

Type species: Cystidioporia piceicola (Y.C. Dai) B.K. Cui & Xing Ji

Etymology: Cystidioporia (Lat.) refers to resembling Perenniporia but with cystidia.

Basidiocarps are annual to biennial, resupinate, soft corky when fresh, and hard corky when dry. Pore surface is cream to buff when fresh and pale yellowish upon drying. Pores are round and large; dissepiments are thin, entire. Subiculum is yellowish ochraceous and corky. Tubes are yellowish ochraceous or straw yellow and corky. Hyphal system is dimitic to trimitic; generative hyphae with clamp connections; skeletal hyphae is slightly dextrinoid, CB+; tissues unchanged in KOH. Cystidia present, thick-walled, strongly CB+. Basidiospores are ellipsoid, truncate, hyaline, thick-walled, smooth, slightly dextrinoid, and CB+.

Notes: In our present phylogenetic analyses (Figure 1 and Figure 2), two specimens of Perenniporia piceicola Y.C. Dai formed a single clade distant from the Perenniporia s. s. clade. Moreover, this species has thick-walled cystidia, large pores, and basidiospores (pores 2–3 per mm, basidiospores 11–14 × 5.4–7.5 µm) [8] which are different from other species of Perenniporia. Thus, the new genus is set up, and the following combination is proposed.

Cystidioporia piceicola (Y.C. Dai) B.K. Cui & Xing Ji, comb. nov.

MycoBank: MB 847366

Basionym: Perenniporia piceicola Y.C. Dai, Ann. Bot. Fenn. 39(3): 173 (2002).

For a detailed description of Perenniporia piceicola, see Dai et al. [8].

Notes: Cystidioporia piceicola was originally described in Perenniporia by Dai et al. [8]; it is characterized by resupinate basidiocarps, thick-walled cystidia, large pores and basidiospores, and usually grows on Picea and Abies.

Specimens examined: CHINA. Yunnan, Lijiang, Yunshanping, on fallen trunk of Picea likiangensis, 18 June 1999, Dai 3089 (isotype, BJFC), on fallen trunk of Abies, 16 September 2018, Cui 17062 (BJFC), and on fallen trunk of Picea, 16 September 2018, Cui 17069 (BJFC).

Dendroporia B.K. Cui & Xing Ji, gen. nov.

MycoBank: MB 847349

Differs from other genera by annual and resupinate basidiocarps with gray to pale brown pore surface, a dimitic hyphal system with weakly dextrinoid skeletal hyphae, tissues darkening in KOH, presence of dendrohyphidia and large rhomboid crystals, and hyaline to pale yellowish, thick-walled, ellipsoid, truncate, non-dextrinoid, and cyanophilous basidiospores.

Type species: Dendroporia cinereofusca (B.K. Cui & C.L. Zhao) B.K. Cui & Xing Ji

Etymology: Dendroporia (Lat.) refers to the presence of dendrohyphidia.

Basidiocarps are annual, resupinate, adnate, and corky. Pore surface is gray to pale brown. Subiculum is thin and clay buff to pale brown. Tubes are concolorous with pore surface and corky. Hyphal system is dimitic; generative hyphae with clamp connections; skeletal hyphae weakly dextrinoid and CB+; tissues are brown to black in KOH. Dendrohyphidia present at dissepimental edges; cystidia are absent; cystidioles are present. Large rhomboid crystals are present. Basidiospores ellipsoid, truncate, hyaline to pale yellowish, thick-walled, smooth, IKI−, and CB+.

Notes: In our present phylogeny, two specimens of Perenniporia cinereofusca B.K. Cui & C.L. Zhao formed a strongly supported clade distinct from the Perenniporia s. s. clade (Figure 1 and Figure 2). Morphologically, P. cinereofusca differs from species of Perenniporia s. s. by its resupinate basidiocarps with a gray to pale brown pore surface, tissues darkening in KOH, hyaline to pale yellowish, and non-dextrinoid basidiospores. Thus, the new genus Dendroporia is set up and the new combination Dendroporia cinereofusca is proposed.

In the current phylogenetic studies, Dendroporia is related to Tropicoporia and Sparsitubus, but with only weak support, and Tropicoporia differs from Dendroporia by its buff-yellow to grayish orange pore surface, non-dextrinoid skeletal hyphae, and dextrinoid basidiospores. Sparsitubus differs from Dendroporia in having effused reflexed to pileate basidiocarps and non-truncate, ornamented basidiospores [54].

Dendroporia cinereofusca (B.K. Cui & C.L. Zhao) B.K. Cui & Xing Ji, comb. nov.

MycoBank: MB 847367

Basionym: Perenniporia cinereofusca B.K. Cui & C.L. Zhao, Mycoscience 55: 419 (2014).

For a detailed description of Perenniporia cinereofusca, see Zhao et al. [20].

Notes: Dendroporia cinereofusca was first described in Perenniporia from tropical China [20] and is characterized by its resupinate basidiocarps with gray to pale brown pore surfaces, a dimitic hyphal system with weakly dextrinoid skeletal hyphae, tissues becoming brown to black in KOH, the presence of dendrohyphidia, and hyaline to pale yellowish, truncate, and non-dextrinoid basidiospores.

Specimens examined: CHINA. Hainan, Ledong County, Jianfengling Nature Reserve, on fallen angiosperm trunk, 18 November 2007, Dai 9289 (holotype, BJFC); Lingshui County, Diaoluoshan Forest Park, on fallen angiosperm trunk, 20 November 2007, Cui 5280 (paratype, BJFC).

Luteoperenniporia B.K. Cui & Xing Ji, gen. nov.

MycoBank: MB 847350

Differs from other genera by its resupinate basidiocarps with buff-yellow to cinnamon-buff pore surface, a dimitic hyphal system with weak to strong dextrinoid skeletal hyphae, the presence of cystidioles, and thick-walled, ellipsoid, and non-truncate, dextrinoid, and cyanophilous basidiospores.

Type species: Luteoperenniporia bannaensis (B.K. Cui & C.L. Zhao) B.K. Cui & Xing Ji

Etymology: Luteoperenniporia (Lat.) refers to resembling Perenniporia but with a buff-yellow pore surface when dry.

Basidiocarps are annual to perennial and resupinate. Pore surface iscream, buff to pale cinnamon buff when fresh, and becoming buff, buff-yellow to cinnamon-buff upon drying; pores are round to angular; dissepiments thin, entire to lacerate. Subiculum is thin and buff to cinnamon-buff. Tubes are concolorous with pore surface and corky. Hyphal system is dimitic, generative hyphae with clamp connections; skeletal hyphae weakly to strongly dextrinoid, CB+; tissues are unchanged in KOH. Cystidia is absent; cystidioles are present. Basidiospores are ellipsoid, non-truncate, hyaline, thick-walled, smooth, dextrinoid, and CB+.

Notes: In the combined ITS + nLSU and five-gene phylogenetic analyses, the species of Luteoperenniporia formed a single clade with high support (Figure 1 and Figure 2) distant from the Perenniporia s. s. clade. Morphologically, Luteoperenniporia differs from Perenniporia s. s. by its buff, buff-yellow to cinnamon-buff pore surfaces, a dimitic hyphal system with weakly to strongly dextrinoid skeletal hyphae and non-truncate basidiospores. Therefore, three new combinations are proposed in Luteoperenniporia, and the new species is described below.

Luteoperenniporia bannaensis (B.K. Cui & C.L. Zhao) B.K. Cui & Xing Ji, comb. nov.

MycoBank: MB 847368

Basionym: Perenniporia bannaensis B.K. Cui & C.L. Zhao, Fungal Diversity 58: 52 (2013).

For a detailed description of Perenniporia bannaensis, see Zhao et al. [19].

Notes: Luteoperenniporia bannaensis was recently described in Perenniporia from China by Zhao et al. [19]; it is closely related to L. yinggelingensis in morphology and phylogeny; they share annual and resupinate basidiocarps, cream to buff pore surface and a dimitic hyphal system with dextrinoid and cyanophilous skeletal hyphae, and both species are distributed in the tropics. However, L. yinggelingensis is distinguished from L. bannaensis by its larger pores and basidiospores (pores 5–6 per mm, basidiospores 6.2–7.5 × 4.5–5.5 μm) [3].

Specimens examined: CHINA. Yunnan, Xishuangbanna, Mengla County, Wangtianshu Nature Reserve, on fallen angiosperm trunk, 2 November 2009, Cui 8560 (holotype, BJFC), Cui 8562 (paratype, BJFC).

Luteoperenniporia mopanshanensis (C.L. Zhao) B.K. Cui & Xing Ji, comb. nov.

MycoBank: MB 847369

Basionym: Perenniporia mopanshanensis C.L. Zhao, Mycotaxon 134(1): 132 (2019).

For a detailed description of Perenniporia mopanshanensis, see Zhao and Ma [28].

Notes: Luteoperenniporia mopanshanensis was recently described in Perenniporia by Zhao and Ma [28]. L. mopanshanensis and L. bannaensis are both reported from Yunnan Province in southern China. They share resupinate basidiocarps, a dimitic hyphal system with strongly dextrinoid skeletal hyphae, non-truncate, and strongly dextrinoid and similar sized basidiospores. However, L. bannaensis differs by having an annual growth habit and smaller pores (6–8 per mm) [19].

Luteoperenniporia yinggelingensis (B.K. Cui & Y.C. Dai) B.K. Cui & Xing Ji, comb. nov.

MycoBank: MB 847370

Basionym: Perenniporia yinggelingensis B.K. Cui & Y.C. Dai, Fungal Diversity 97: 300 (2019).

For a detailed description of Perenniporia yinggelingensis, see Cui et al. [3].

Notes: Luteoperenniporia yinggelingensis was newly described from a tropical area of China [3]. It is characterized by annual and resupinate basidiocarps with slightly lacerate pores, a distinct sterile margin, and a dimitic hyphal system with weakly dextrinoid skeletal hyphae. Macroscopically, L. yinggelingensis is close to L. mopanshanensis, but the latter species has perennial basidiocarps, larger pores (3–5 per mm), indistinct sterile margins, and strongly dextrinoid skeletal hyphae [28].

Specimens examined: CHINA. Hainan, Baisha County, Yinggeling Nature Reserve, on fallen angiosperm trunk, 17 November 2015, Cui 13625 (holotype, BJFC); on fallen angiosperm branch, 17 June 2016, Cui 13856 (BJFC).

Luteoperenniporia australiensis B.K. Cui & Xing Ji, sp. nov.; Figure 3 and Figure 4

Figure 3.

Basidiocarp of Luteoperenniporia australiensis. Scale bar = 2.0 cm.

Figure 4.

Microscopic structures of Luteoperenniporia australiensis (Holotype). (a). Basidiospores. (b). Basidia and basidioles. (c). Cystidioles. (d). Hyphae from trama. (e). Hyphae from subiculum.

MycoBank: MB 847371

Differs from other species of Luteoperenniporia by its annual to perennial growth habit, resupinate basidiocarps with buff to cinnamon-buff pore surfaces, a dimitic hyphal system with dextrinoid skeletal hyphae, and ellipsoid, non-truncate, dextrinoid basidiospores (6.2–7.5 × 4–5.2 μm).

Holotype: AUSTRALIA. Tasmania, Keogh’s Creek Walk, on fallen trunk of Eucalyptus, 15 May 2018, Cui 16743 (BJFC).

Etymology: australiensis (Lat.) refers to the country where the new species was found.

Fruitbody: Basidiocarps are annual to perennial, resupinate, without odor or taste when fresh, corky to hard corky when dry, and up to 14 cm long, 8.5 cm wide, and 7 mm thick at center. Pore surface is buff-yellow, pinkish buff to pale cinnamon-buff when fresh, and buff, pale yellowish-brown to cinnamon-buff upon drying; pores are round to angular, 4–6 per mm, dissepiments thin, entire to lacerate. Sterile margin is distinct to indistinct, buff, and up to 2 mm wide. Subiculum is thin, buff, and up to 1 mm thick. Tubes are concolorous with pore surface, corky to hard corky when dry, and up to 6 mm long.

Hyphal structure: Hyphal system is dimitic; generative hyphae bearing clamp connections; skeletal hyphae dextrinoid, and CB+; tissues are unchanged in KOH.

Subiculum: Generative hyphae is infrequent, hyaline, thin-walled, occasionally branched, and 1.5–2.5 μm in diameter; skeletal hyphae is dominant, hyaline, thick-walled with a wide to narrow lumen, rarely branched, interwoven, and 2–5 μm in diameter.

Tubes: Generative hyphae is infrequent, hyaline, thin-walled, occasionally branched, and 1–2.4 μm in diameter; skeletal hyphae is dominant, hyaline, thick-walled with a wide to narrow lumen, occasionally branched, interwoven, and 1–3 μm. Cystidia are absent; fusoid cystidioles are present, hyaline, thin-walled, and 15.5–22 × 5.8–8 μm. Basidia clavate, with four sterigmata and a basal clamp connection, 15–24.5 × 6.5–9.7 μm; basidioles are dominant and in shape similar to basidia, but slightly smaller.

Spores: Basidiospores are ellipsoid, non-truncate, hyaline, thick-walled, smooth, dextrinoid, CB+, (6–) 6.2–7.4 (–7.7) × (3.9–) 4–5.2 (–5.4) μm, L = 6.77 μm, W = 4.58 μm, Q = 1.44–1.54 (n = 90/3).

Notes: Luteoperenniporia australiensis is characterized by its annual to perennial and resupinate basidiocarps, buff to cinnamon-buff pore surface, entire to lacerate pores, a dimitic hyphal system with dextrinoid skeletal hyphae, presence of cystidioles, and ellipsoid, non-truncate, dextrinoid, and cyanophilous basidiospores. Luteoperenniporia bannaensis is similar to L. australiensis by sharing a buff-yellow to pinkish buff pore surface, a dimitic hyphal system with dextrinoid, and cyanophilous skeletal hyphae. However, L. bannaensis differs from L. australiensis by its smaller pores (6–8 per mm) and smaller basidiospores (5.2–6 × 4–4.6 μm) [19]. Luteoperenniporia yinggelingensis may be confused with L. australiensis by having resupinate basidiocarps, and similar pore size and basidiospores. However, L. yinggelingensis is distinguished from L. australiensis mainly by its annual growth habit and cream to buff pore surface [3]. Perenniporia subaurantiaca (Rodway & Cleland) P.K. Buchanan & Ryvarden is also described from Tasmania, it is similar to Luteoperenniporia australiensis in its resupinate basidiocarps, dimitic hyphal system with dextrinoid and cyanophilous skeletal hyphae, and presence of cystidioles, but P. subaurantiaca has greyish cream to greyish orange pore surfaces and larger basidiospores (7.2–9.5 × 4.2–5.5 μm) [55]

Additional specimens (paratypes) examined: AUSTRALIA. Victoria, Yarra Ranges National Park, at the base of living Eucalyptus, 9 May 2018, Cui 16524 (BJFC), on fallen trunk of Eucalyptus, 9 May 2018, Cui 16525 (BJFC), 10 May 2018, Cui 16535 (BJFC), on living tree of Eucalyptus, 10 May 2018, Cui 16533, Cui 16534 (BJFC); Tasmania, Keogh’s Creek Walk, on fallen trunk of Eucalyptus, 15 May 2018, Cui 16742 (BJFC).

Key to species of Luteoperenniporia

• 1. Pores 6–8 per mm……………………………………………………………………………………………………………………………………………L. bannaensis

• 1. Pores 3–6 per mm…………………………………………………………………………………………………………………………………………………………2

• 2. Skeletal hyphae unbranched…………………………………………………………………………………………………………………………L. mopanshanensis

• 2. Skeletal hyphae branched ……………………………………………………………………………………………………………………………………………… 3

• 3. Basidiocarps annual, distributed in tropical areas…………………………………………………………………………………………………L. yinggelingensis

• 3. Basidiocarps annual to perennial, distributed in temperate to subtropical areas…………………………………………………………………L. australiensis

Macroporia B.K. Cui & Xing Ji, gen. nov.

MycoBank: MB 847352

Differs from other genera by its annual and resupinate basidiocarps, a dimitic hyphal system with dextrinoid skeletal hyphae, the presence of thin-walled cystidioles, and hyaline, thick-walled, ellipsoid, truncate, dextrinoid, and cyanophilous basidiospores.

Type species: Macroporia macropora (B.K. Cui & C.L. Zhao) B.K. Cui & Xing Ji

Etymology: Macroporia (Lat.) refers to the species with relatively large pores in Perenniporia.

Basidiocarps are annual, resupinate, adnate. Pore surface is white, cream to buff when fresh, and becoming buff, pinkish buff to yellowish buff upon drying; pores are angular; dissepiments are thin, entire to lacerate. Subiculum is thin, cream. Tubes are concolorous with pore surface, corky to hard corky. Hyphal system is dimitic, generative hyphae with clamp connections; skeletal hyphae branched, dextrinoid, CB+; tissues are unchanged in KOH. Cystidia is absent; cystidioles are usually present. Basidiospores are ellipsoid, truncate, hyaline, thick-walled, smooth, dextrinoid, and CB+.

Notes: In our ITS + nLSU and combined five-gene phylogenetic analyses, Perenniporia lacerata B.K. Cui & C.L. Zhao, P. macropora B.K. Cui & C.L. Zhao, P. subrhizomorpha Xue W. Wang, L.W. Zhou & X.M. Tian and P. tibetica B.K. Cui & C.L. Zhao grouped together and formed a well-supported clade (Figure 1 and Figure 2), which was distant from the Perenniporia s. s. clade. Morphologically, species in the clade usually have larger pores than those of Perenniporia s. s.; therefore, the new genus Macroporia is proposed to accommodate the four species.

Macroporia lacerata (B.K. Cui & C.L. Zhao) B.K. Cui & Xing Ji, comb. nov.

MycoBank: MB 847372

Basionym: Perenniporia lacerata B.K. Cui & C.L. Zhao, Mycoscience 54: 232 (2013).

For a detailed description of Perenniporia lacerata, see Zhao and Cui [18].

Notes: Macroporia lacerata was originally described in Perenniporia from China by Zhao and Cui [18]. It is characterized by papery and thin basidiocarps, lacerate pores, a dimitic hyphal system with weakly dextrinoid skeletal hyphae, and ellipsoid, truncate, dextrinoid, and cyanophilous basidiospores.

Specimens examined: CHINA. Henan, Xiuwu County, Yuntaishan Park, on fallen angiosperm trunk, 3 September 2009, Cui 7220 (holotype, BJFC); Neixiang County, Baotianman Nature Reserve, on rotten angiosperm wood, 22 September 2009, Dai 11268 (paratype, BJFC).

Macroporia macropora (B.K. Cui & C.L. Zhao) B.K. Cui & Xing Ji, comb. nov.

MycoBank: MB 847373

Basionym: Perenniporia macropora B.K. Cui & C.L. Zhao, Mycologia 105: 947 (2013).

For a detailed description of Perenniporia macropora, see Zhao and Cui [17].

Notes: Macroporia macropora was first described in Perenniporia from southern China by Zhao and Cui [17]. It is distinguished by large pores (2–3 per mm), a dimitic hyphal system with dextrinoid and branched skeletal hyphae, the presence of dendrohyphidia, and ellipsoid and truncate basidiospores. M. macropora is very close to M. lacerata in the current phylogeny, but M. lacerata differs from M. macropora by its smaller (3–5 per mm) and lacerate pores, the absence of dendrohyphidia, and smaller basidiospores (6.1–7 × 5–5.7 μm) [18].

Specimens examined: CHINA. Guangxi, Ningming County, Nonggang Nature Reserve, on fallen angiosperm branch, 8 July 2007, Zhou 407 (holotype, IFP), 7 July 2007, Zhou 297 (paratype, IFP).

Macroporia subrhizomorpha (Xue W. Wang, L.W. Zhou & X.M. Tian) B.K. Cui & Xing Ji, comb. nov.

MycoBank: MB 847374

Basionym: Perenniporia subrhizomorpha Xue W. Wang, L.W. Zhou & X.M. Tian, Phytotaxa 528(2): 129 (2021).

For a detailed description of Perenniporia subrhizomorpha, see Tian et al. [56].

Notes: Macroporia subrhizomorpha was recently described in Perenniporia as P. subrhizomorpha by Tian et al. [56]. Perenniporia rhizomorpha may be confused with P. subrhizomorpha by sharing cream rhizomorphs and similar pore sizes, but the former species differs by its non-truncate basidiospores [10]. Perenniporia tibetica also has rhizomorphs but differs from P. subrhizomorpha by larger pores (2–3 per mm) [15].

Macroporia tibetica (B.K. Cui & C.L. Zhao) B.K. Cui & Xing Ji, comb. nov.

MycoBank: MB 847375

Basionym: Perenniporia tibetica B.K. Cui & C.L. Zhao, Mycoscience 53: 366 (2012).

For a detailed description of Perenniporia tibetica, see Cui and Zhao [15].

Notes: Macroporia tibetica is characterized by resupinate basidiocarps with white to cream rhizomorphs, a dimitic hyphal system with slightly dextrinoid skeletal hyphae, and ellipsoid, truncate or not, dextrinoid, and cyanophilous basidiospores. M. macropora and M. tibetica share resupinate basidiocarps and similar pores, but the former differs in having dendrohyphidia and lacking rhizomorphs [17].

Specimens examined: CHINA. Xizang, Linzhi County, Tongmai, on fallen angiosperm trunk, 16 September 2010, Cui 9457 (holotype, BJFC), Cui 9459 (paratype, BJFC).

Key to species of Macroporia

• 1. Pores lacerate …………………………………………………………………………………………………………………………………………………  M. lacerata

• 1. Pores entire ………………………………………………………………………………………………………………………………………………………………  2

• 2. Basidiocarps without rhizomorphs……………………………………………………………………………………………………………………… M. macropora

• 2. Basidiocarps with rhizomorphs…………………………………………………………………………………………………………………………………………3

• 3. Pores 2–3 per mm………………………………………………………………………………………………………………………………………………M. tibetica

• 3. Pores 4–6 per mm …………………………………………………………………………………………………………………………………… M. subrhizomorpha

Macrosporia B.K. Cui & Xing Ji, gen. nov.

MycoBank: MB 847353

Differs from other genera by its annual and resupinate basidiocarps, cinnamon-buff pore surface, a trimitic hyphal system with weakly dextrinoid and cyanophilous skeletal hyphae, and hyaline, thick-walled, ellipsoid, truncate, strongly dextrinoid, and cyanophilous basidiospores.

Type species: Macrosporia nanlingensis (B.K. Cui & C.L. Zhao) B.K. Cui & Xing Ji

Etymology: Macrosporia (Lat.) refers to the large basidiospores.

Basidiocarps are annual, resupinate, adnate, corky when fresh, and becoming hard corky upon drying. Pore surface is cream-buff to yellowish buff when fresh, becoming cinnamon-buff upon drying; pores are round; dissepiments are thick, entire. Subiculum is cream to buff. Tubes are concolorous with the pore surface and hard corky. Hyphal system is trimitic; generative hyphae with clamp connections; skeletal and binding hyphae are weakly dextrinoid and CB+. Cystidia is absent; cystidioles are present. Basidiospores areellipsoid, truncate, hyaline, thick-walled, smooth, strongly dextrinoid, and CB+.

Notes: In our study, Perenniporia nanlingensis B.K. Cui & C.L. Zhao formed a single clade that was distant from the Perenniporia s. s. clade. Morphologically, it differs from Perenniporia s. s. by its annual and resupinate basidiocarps with cinnamon-buff pore surfaces and larger basidiospores. Therefore, Macrosporia gen. nov. is proposed to accommodate P. nanlingensis.

Macrosporia nanlingensis (B.K. Cui & C.L. Zhao) B.K. Cui & Xing Ji, comb. nov.

MycoBank: MB 847376

Basionym: Perenniporia nanlingensis B.K. Cui & C.L. Zhao, Mycol. Prog. 11: 556 (2012).

For a detailed description of Perenniporia nanlingensis, see Zhao and Cui [16].

Notes: Macrosporia nanlingensis was first described in Perenniporia from southern China [16]; it is characterized by annual and resupinate basidiocarps, a cinnamon-buff pore surface when dry, a trimitic hyphal system with slightly dextrinoid skeletal hyphae, and ellipsoid, truncate, strongly dextrinoid, and cyanophilous basidiospores that are usually longer than 9 μm.

Specimens examined: CHINA. Guangdong Province, Ruyang County, Nanling Nature Reserve, on dead angiosperm tree, 16 September 2009, Cui 7589 (holotype, BJFC), Cui 7620 (paratype, BJFC).

Minoporus B.K. Cui & Xing Ji, gen. nov.

MycoBank: MB 847354

Differs from other genera by its annual and pileate basidiocarps, cream to pale buff pileal surface when fresh, a dimitic hyphal system with weakly amyloid and cyanophilous skeletal hyphae, and hyaline, thick-walled, ellipsoid, truncate, dextrinoid, and cyanophilous basidiospores.

Type species: Minoporus minor (Y.C. Dai & H.X. Xiong) B.K. Cui & Xing Ji

Etymology: Minoporus (Lat.) refers to the small pilei.

Basidiocarps are annual, pileate, solitary, and soft corky when fresh, becoming hard corky upon drying. Pilei are semicircular to spathulate. Pileal surface is cream to pale buff when fresh, becoming cinnamon-buff when dry. Pore surface is cream when fresh, becoming cinnamon-buff when dry; pores are round. Context is white to cream, corky. Tubes are concolorous with pore surface and hard corky. Hyphal system dimitic; generative hyphae with clamp connections; skeletal hyphae is weakly amyloid and CB+. Cystidia and cystidioles are absent. Basidiospores are ellipsoid, truncate, hyaline, thick-walled, smooth, dextrinoid, and CB+.

Notes: In the ITS + nLSU analysis, two specimens of Perenniporia minor Y.C. Dai & H.X. Xiong formed a highly supported single clade (Figure 1), which is distinct from the Perenniporia s. s. clade and closely related to the Perenniporiella clade. Further phylogeny (Figure 2) inferred from the combined five-gene dataset indicated that the clade of P. minor was distant from the Perenniporia s. s. clade and closely related to the Neoporia clade. However, Neoporia has resupinate basidiocarps, dextrinoid skeletal hyphae, and non-truncate basidiospores; Perenniporiella has dextrinoid skeletal hyphae and non-truncate basidiospores [30]. Perenniporia minor differs from species of Perenniporia s. s. by its annual and pileate basidiocarps with a cream to pale buff pileal surface, a dimitic hyphal system with weakly amyloid skeletal hyphae. Therefore, Minoporus gen. nov. is proposed to accommodate P. minor.

Minoporus minor (Y.C. Dai & H.X. Xiong) B.K. Cui & Xing Ji, comb. nov.

MycoBank: MB 847377

Basionym: Perenniporia minor Y.C. Dai & H.X. Xiong, Mycotaxon 105: 60 (2008).

For a detailed description of Perenniporia minor, see Xiong et al. [11].

Notes: This species was described from northeastern China by Xiong et al. [11] and is characterized by annual and pileate basidiocarps, cream to pale buff when fresh and cinnamon buff when dry pileal surface, a dimitic hyphal system with weakly amyloid skeletal hyphae, and ellipsoid, truncate, dextrinoid, and cyanophilous basidiospores.

Specimens examined: CHINA. Jilin, Antu County, Changbaishan Nature Reserve, Huangsongpu, on fallen branch of Acer, 14 September 2007, Dai 9198 (holotype, IFP); Liaoning, Huanren County, Laotudingzi Nature Reserve, on fallen branch of Quercus, 31 July 2008, Cui 5738 (BJFC).

Neoporia B.K. Cui & Xing Ji, gen. nov.

MycoBank: MB 847355

Differs from other genera by its annual and resupinate basidiocarps with a buff-yellow pore surface, a dimitic hyphal system with dextrinoid and cyanophilous skeletal hyphae, and ellipsoid, non-truncate, dextrinoid, and cyanophilous basidiospores.

Type species: Neoporia rhizomorpha (B.K. Cui, Y.C. Dai & Decock) B.K. Cui & Xing Ji

Etymology: Neoporia (Lat.) refers to the genus resembling Perenniporia.

Basidiocarps are annual, resupinate and corky when dry. Pore surface is cream to buff when fresh, buff-yellow to grayish orange upon drying; pores are round to angular. Subiculum is cream to buff. Tubes are concolorous with the pore surface and corky. Hyphal system is dimitic; generative hyphae with clamp connections; skeletal hyphae dextrinoid, CB+; tissues unchanged in KOH. Basidiospores are ellipsoid, non-truncate, hyaline, thick-walled, smooth, dextrinoid, and CB+.

Notes: In our phylogenetic analyses, three species previously included in Perenniporia, P. bostonensis C.L. Zhao, P. koreana Y. Jang & J.J. Kim, and P. rhizomorpha B.K. Cui, Y.C. Dai & Decock grouped together and formed a highly supported clade (100% BS, 1.00 BPP, Figure 1; 100% BS, 1.00 BPP, Figure 2), which was distant from the Perenniporia s. s. clade. Morphologically, this clade differs from Perenniporia s. s. by its dimitic hyphal system and non-truncate basidiospores. Therefore, Neoporia gen. nov. is proposed to accommodate P. bostonensis, P. koreana and P. rhizomorpha.

In the five-gene phylogenetic analysis, Neoporia is related to Luteoperenniporia, Minoporus, Poriella, Vanderbyliella, and Yuchengia. Luteoperenniporia, Neoporia, Poriella, Vanderbyliella, and Yuchengia all have ellipsoid and non-truncate basidiospores. However, Luteoperenniporia differs from Neoporia in having a perennial growth habit; Poriella differs from Neoporia by having a perennial growth habit, cinnamon to ochraceous pore surfaces, and unbranched skeletal hyphae; Vanderbyliella is different from Neoporia by its pileate basidiocarps; Yuchengia differs from Neoporia in having amyloid skeletal hyphae and non-dextrinoid basidiospores [35]. In addition, Minoporus differs from Neoporia by its pileate basidiocarps, amyloid skeletal hyphae, and truncate basidiospores.

Neoporia bostonensis (C.L. Zhao) B.K. Cui & Xing Ji, comb. nov.

MycoBank: MB 847378

Basionym: Perenniporia bostonensis C.L. Zhao, Phytotaxa 351(1): 67 (2018).

For a detailed description of Perenniporia bostonensis, see Shen et al. [27].

Notes: Neoporia bostonensis was recently described in Perenniporia from North America by Shen et al. [27]. It is characterized by resupinate basidiocarps with cream to buff pore surfaces, a dimitic hyphal system with strongly dextrinoid and unbranched skeletal hyphae, and ovoid to broad ellipsoid, non-truncate, and dextrinoid basidiospores.

Neoporia koreana (Y. Jang & J.J. Kim) B.K. Cui & Xing Ji, comb. nov.

MycoBank: MB 847379

Basionym: Perenniporia koreana Y. Jang & J.J. Kim, Mycotaxon 130(1): 174 (2015).

For a detailed description of Perenniporia bannaensis, see Jang et al. [22].

Notes: Neoporia koreana was originally described from Republic of Korea as Perenniporia koreana by Jang et al. [22]; it has annual and resupinate basidiocarps with grayish orange pore surfaces, dextrinoid skeletal hyphae, and ellipsoid and non-truncate basidiospores. The sequences of Neoporia koreana from type specimens fell into Neoporia in our phylogeny. Thus, P. koreana is transferred to Neoporia. N. koreana is similar to N. bostonensis in resupinate basidiocarps, similar sized pores, dextrinoid skeletal hyphae, and non-truncate basidiospores, but the former has larger basidiospores (6–7 × 3.9–5.2 μm) [22] and the latter has smaller basidiospores (3.5–4.5 × 3–4 μm) [27].

Neoporia rhizomorpha (B.K. Cui, Y.C. Dai & Decock) B.K. Cui & Xing Ji, comb. nov.

MycoBank: MB 847380

Basionym: Perenniporia rhizomorpha B.K. Cui, Y.C. Dai & Decock, Mycotaxon 99: 176 (2007).

For a detailed description of Perenniporia rhizomorpha, see Cui et al. [10].

Notes: Neoporia rhizomorpha was first described in Perenniporia based on morphological characters from China [10]. It is unique in Neoporia due to its resupinate basidiocarps with rhizomorphs.

Specimens examined: CHINA. Anhui, Huangshan, Yellow Mountain, on fallen angiosperm trunk, 13 October 2004, Dai 6165 (holotype, BJFC); Fujian, Wuyishan County, Wuyishan Nature Reserve, on fallen angiosperm branch, 19 October 2005, Cui 7248 (paratype, BJFC).

Key to species of Neoporia

• 1. Basidiocarps with rhizomorphs…………………………………………………………………………………………………………………………N. rhizomorpha

• 1. Basidiocarps without rhizomorphs…………………………………………………………………………………………………………………………………… 2

• 2. Basidiospores 3.5–4.5 × 3–4 μm ………………………………………………………………………………………………………………………… N. bostonensis

• 2. Basidiospores 6–7 × 3.9–5.2 μm………………………………………………………………………………………………………………………………N. koreana

Niveoporia B.K. Cui & Xing Ji, gen. nov.

MycoBank: MB 847356

Differs from other genera by perennial basidiocarps with white pore surface when fresh, distinct rusty red to reddish brown sterile margin, a dimitic hyphal system with dextrinoid and cyanophilous skeletal hyphae, the presence of cystidioles, hyaline, and thick-walled, ellipsoid, and truncate basidiospores.

Type species: Niveoporia russeimarginata (B.K. Cui & C.L. Zhao) B.K. Cui & Xing Ji

Etymology: Niveoporia (Lat.) refers to the white pore surface.

Basidiocarps are perennial, resupinate to pileate, corky to woody hard when dry. Pilei dimidiate to fan shaped. Pileal surface is clay-buff to reddish brown when fresh, grayish brown to umber brown when dry, glabrous, and concentrically sulcate. Pore surface is white when fresh and white to cream upon drying; pores are round. The sterile margin is sometimes distinct, rusty red to reddish brown. Context is buff to fawn. Tubes are cream to pale cinnamon. Hyphal system is dimitic; generative hyphae with clamp connections; skeletal hyphae dextrinoid, CB+; tissues unchanged in KOH. Cystidia is absent; cystidioles are present. Basidiospores are ellipsoid, truncate, hyaline, thick-walled, smooth, dextrinoid or not, and CB+.

Notes: In the phylogenetic analysis of this study, P. decurrata F. Wu & X.H. Ji, P. russeimarginata B.K. Cui & C.L. Zhao and an undescribed species grouped together and formed a well-supported clade (74% BS, 1.00 BPP, Figure 1; 83% BS, 0.97 BPP, Figure 2) which was separated from the Perenniporia s. s. clade. Morphologically, species in the clade usually have pileate basidiocarps and white pore surfaces with reddish brown sterile margins, which are different from the species of Perenniporia s. s. Thus, Niveoporia gen. nov. is proposed to accommodate these species.

Niveoporia decurrata (Corner) B.K. Cui & Xing Ji, comb. nov.

MycoBank: MB 847381

Basionym: Perenniporia decurrata Corner, Beih. Nova Hedwigia 96: 105 (1989)

= Perenniporia chiangraiensis F. Wu & X.H. Ji, Mycosphere 8(8): 1103 (2017).

For a detailed description of Perenniporia decurrata, see Corner [52].

Notes: Perenniporia decurrata was first described from Malaysia [52]. Perenniporia chiangraiensis was recently described from Northern Thailand based on morphological characters and molecular data by Ji et al. [25]. However, these authors overlooked P. decurrata, which is a priority name for this species. The species is characterized by pileate basidiocarps with concentrically sulcate pileal surfaces, white pore surfaces, the presence of dendrohyphidia and cystidioles, and ellipsoid, truncate, thick-walled, and non-dextrinoid basidiospores.

Specimens examined: CHINA. Yunnan, Xishuangbanna, Menglun, on fallen angiosperm trunk, 12 September 2006, Yuan 2334 (IFP). THAILAND. Chiang Rai, Doi Mae Salong, on angiosperm tree root, 22 July 2016, Dai 16637 (BJFC).

Niveoporia russeimarginata (B.K. Cui & C.L. Zhao) B.K. Cui & Xing Ji, comb. nov.

MycoBank: MB 847382

Basionym: Perenniporia russeimarginata B.K. Cui & C.L. Zhao, Mycologia 105(4): 947 (2013).

For a detailed description of Perenniporia russeimarginata, see Zhao and Cui [17].

Notes: Niveoporia russeimarginata was first described in Perenniporia from southern China [17]. Perenniporia alboferruginea Decock, described from Cameroon in Africa, is similar to N. russeimarginata in having resupinate basidiocarps with ferruginous red upper margins and a dimitic hyphal system. However, P. alboferruginea differs by having an annual growth habit, larger pores (5–6 per mm), the absence of cystidioles, and non-dextrinoid basidiospores [57].

Specimens examined: CHINA. Yunnan, Chuxiong, Zixishan Nature Reserve, on fallen angiosperm trunk, 1 August 2005, Yuan 1225 (holotype, IFP), Yuan 1262 (paratype, IFP).

Niveoporia subrusseimarginata B.K. Cui & Xing Ji, sp. nov., Figure 5 and Figure 6

Figure 5.

Basidiocarp of Niveoporia subrusseimarginata (Holotype). Scale bar = 2.0 cm.

Figure 6.

Microscopic structures of Niveoporia subrusseimarginata (Holotype). (a). Basidiospores. (b). Basidia and basidioles. (c). Cystidioles. (d). Hyphae from trama. (e). Hyphae from context.

MycoBank: MB 847383

Differs from other Niveoporia species by its resupinate to pileate basidiocarps with rusty reddish brown sterile margins and pores measuring 5–6 per mm.

Holotype: CHINA. Yunnan, Binchuan County, Jizushan Park, on angiosperm stump, 14 September 2018, Cui 16991 (BJFC).

Etymology: Subrusseimarginata (Lat.) refers to the species resembling Niveoporia russeimarginata.

Fruitbody: Basidiocarps are perennial, resupinate, sometimes pileate, corky, without odor or taste when fresh, becoming woody hard upon drying. Pilei are irregular, projecting up to 2 cm, 5 cm wide, and 3 cm thick at the base; with resupinate up to 9 cm long, 5 cm wide, and 1.3 cm thick at center. Pileal surface is orange-brown to reddish brown when fresh, umber brown when dry, glabrous, and concentrically sulcate. The pore surface is white when fresh, cream upon drying; pores are round to angular, 5–6 per mm, dissepiments thick, entire. Sterile margin is distinct to indistinct, cinnamon to rusty reddish brown, and up to 2 mm wide. Subiculum is buff, thin, up to 0.5 mm thick. Tubes are buff, woody hard when dry, and up to 11.5 mm long.

Hyphal structure: Hyphal system is dimitic; generative hyphae bearing clamp connections; skeletal hyphae is weakly dextrinoid and CB+; tissues are unchanged in KOH.

Subiculum: Generative hyphae are infrequent, hyaline, thin-walled, unbranched, and 1–2.2 μm in diameter; skeletal hyphae are dominant, hyaline, thick-walled, occasionally branched, interwoven, and 1–2.5 μm in diameter.

Tubes: Generative hyphae are infrequent, hyaline, thin-walled, occasionally branched, and 1–2 μm in diameter; skeletal hyphae are dominant, hyaline, thick-walled, moderately branched, and interwoven, 1–2.3 μm. Cystidia are absent; fusoid cystidioles are present, hyaline, thin-walled, and 12.5–14 × 4.8–6 μm. Basidia are clavate, with four sterigmata and a basal clamp connection, 12.5–17.5 × 6.3–9 μm; basidioles are dominant and in shape similar to basidia, but slightly smaller.

Spores: Basidiospores are broadly ellipsoid, truncate or not, hyaline, thick-walled, smooth, weakly dextrinoid, CB+, (4–)4.2–4.8 (–5) × (3–)3.2–3.8(–4) μm, L = 4.48 μm, W = 3.51 μm, Q = 1.27–1.3 (n = 90/3).

Notes: Niveoporia subrusseimarginata is characterized by resupinate to pileate basidiocarps with white to cream pore surface, cinnamon to rust sterile margin, dimitic hyphal system with weakly dextrinoid skeletal hyphae, the presence of cystidioles, and broadly ellipsoid, truncate or not basidiospores.

Additional specimens (paratypes) examined: CHINA. Yunnan, Binchuan County, Jizushan Park, on the stump of Quercus, 14 September 2018, Cui 16973 (BJFC), on fallen angiosperm branch, 14 September 2018, Cui 16980 (BJFC), on angiosperm stump, 14 September 2018, Cui 16988, 16990 (BJFC).

Key to species of Niveoporia

• 1. Sterile margin indistinct …………………………………………………………………………………………………………………………………… N. decurrata

• 1. Sterile margin distinct, reddish brown…………………………………………………………………………………………………………………………………2

• 2. Pores 5–6 per mm………………………………………………………………………………………………………………………………… N. subrusseimarginata

• 2. Pores 6–8 per mm …………………………………………………………………………………………………………………………………… N. russeimarginata

Perenniporia Murrill, Mycologia 34(5): 595 (1942).

MycoBank: MB 18204

Type species: Perenniporia medulla-panis (Jacq.) Donk, Persoonia 5(1): 76 (1967).

Basidiocarps are annual to perennial and resupinate. Pore surface is white to cream when fresh, cream to buff when dry; pores are round and small. Subiculum is thin, cream, and corky. Tubes are concolorous with pore surface and corky. Hyphal system is dimitic to trimitic; generative hyphae with clamp connections; skeletal hyphae are non-dextrinoid to dextrinoid or amyloid, cyanophilous; tissues are unchanged in KOH. Cystidia are absent, cystidioles are present. Basidiospores are ellipsoid, truncate, hyaline, thick-walled, smooth, dextrinoid, and CB+.

Notes: In our phylogenetic analyses, the species of Perenniporia s. l. clustered into several clades, Perenniporia medulla-panis is grouped with P. substraminea B.K. Cui & C.L. Zhao and P. hainaniana B.K. Cui & C.L. Zhao. These three species share similar morphological characteristics and form the core clade of Perenniporia. The above concept of Perenniporia s. s. is determined from P. medulla-panis, P. hainaniana, and P. substraminea.

Specimens examined: Perenniporia hainaniana. China. Hainan, Changjiang County, Bawangling Nature Reserve, on angiosperm stump, 8 May 2009, Cui 6364 (holotype, BJFC). Perenniporia medulla-panis. China. Guangxi, Jinxiu County, Dayaoshan Nature Reserve, on living angiosperm tree, 15 July 2017, Cui 14515 (BJFC). Perenniporia substraminea. China. Zhejiang, Taishun County, Wuyanling Nature Reserve, on angiosperm stump, 22 August 2011, Cui 10177 (holotype, BJFC).

Key to species of Perenniporia s. s.

• 1. Pores > 8 per mm…………………………………………………………………………………………………………………………………………P. substraminea

• 1. Pores < 7 per mm…………………………………………………………………………………………………………………………………………………………2

• 2. Dendrohyphidia present at dissepimental edges………………………………………………………………………………………………………P. hainaniana

• 2. Dendrohyphidia absent at dissepimental edges…………………………………………………………………………………………………… P. medulla-panis

Poriella C.L. Zhao, Agronomy 11(7, no. 1308): 5 (2021).

MycoBank: MB 840061

Type species: Poriella subacida (Peck) C.L. Zhao, Agronomy 11(7, no. 1308): 6 (2021).

Basidiocarps are annual to perennial, resupinate to effused-reflexed, and corky when dry. Pore surface is dingy yellowish, cinnamon to ochraceous; pores are round to angular. Context is thin, cream, buff to pale ochraceous. Tubes are concolorous with pore surface, corky. Hyphal system is dimitic to trimitic; generative hyphae with clamp connections; skeletal hyphae is unbranched, strongly dextrinoid, and cyanophilous. Basidiospores are ellipsoid to subglobose, non-truncate, hyaline, thick-walled, smooth, non-dextrinoid to dextrinoid, and CB+.

Notes: Poriella was newly set up by Chen et al. [37] according to analyses of ITS, nLSU, mtSSU, and TEF1 datasets. The type species Poriella subacida, originally described as Polyporus subacidus Peck [58], was usually treated in Perenniporia in current studies [2,3,59]. In our phylogeny, three species of Perenniporia, P. africana Ipulet & Ryvarden, P. ellipsospora Ryvarden & Gilb., and P. valliculorum Spirin et Zmitr. grouped together with Poriella subacida and formed a highly supported clade (100% BS, 1.00 BPP, Figure 1; 98% BS, 1.00 BPP, Figure 2); morphologically, these three species have resupinate basidiocarps, a dimitic hyphal system with unbranched and strongly dextrinoid skeletal hyphae, and hyaline, thick-walled, and non-truncate basidiospores, which are quite consistent with the concept of Poriella. Therefore, these three species are transferred to Poriella based on molecular data and morphological characters.

Poriella africana (Ipulet & Ryvarden) B.K. Cui & Xing Ji, comb. nov.

MycoBank: MB 847384

Basionym: Perenniporia africana Ipulet & Ryvarden, Syn. Fung. 20: 94 (2005).

For a detailed description of Perenniporia africana, see Ipulet and Ryvarden [60] and Cui et al. [3].

Notes: Poriella africana was originally described from Uganda as Perenniporia africana [60]. It has perennial and resupinate basidiocarps, a dimitic hyphal system with unbranched and strongly dextrinoid skeletal hyphae, and subglobose to broadly ellipsoid and non-truncate basidiospores.

Specimens examined: CHINA. Anhui, She County, Qingliangfeng Nature Reserve, on fallen angiosperm trunk, 14 December 2009, Cui 8674, 8676 (BJFC).

Poriella ellipsospora (Ryvarden & Gilb.) B.K. Cui & Xing Ji, comb. nov.

MycoBank: MB 847385

Basionym: Perenniporia ellipsospora Ryvarden & Gilb., Mycotaxon 19: 140 (1984).

For a detailed description of Perenniporia ellipsospora, see Cui et al. [3].

Notes: Poriella ellipsospora and P. subacida both have resupinate basidiocarps, strongly dextrinoid skeletal hyphae, and ellipsoid and non-truncate basidiospores, but the former has larger pores (3–4 per mm) and dextrinoid basidiospores, and the latter has smaller pores (4–6 per mm) and non-dextrinoid basidiospores [37].

Specimens examined: CHINA. Yunnan, Lanping County, Changyanshan Nature Reserve, on fallen angiosperm trunk, 18 September 2011, Cui 10276, 10284 (BJFC).

Poriella valliculorum (Spirin & Zmitr.) B.K. Cui & Xing Ji, comb. nov.

Mycobank: MB 847386

Basionym: Perenniporia valliculorum Spirin & Zmitr., Folia Cryptogamica Petropolitana (Sankt-Peterburg) 6: 51 (2005).

For a detailed description of Perenniporia valliculorum, see Spirin et al. [9].

Notes: Poriella valliculorum was originally described from Russia as Perenniporia valliculorum by Spirin et al. [9]. It has a pale citric yellow to pale tan pore surface, strongly dextrinoid skeletal hyphae, and non-truncate basidiospores, and these characters fit Poriella well. In addition, the sequence of P. valliculorum from the type specimen fell into Poriella in our phylogeny (Figure 1 and Figure 2). Therefore, Perenniporia valliculorum is transferred to Poriella.

Key to species of Poriella

• 1. Basidiospores non-dextrinoid ………………………………………………………………………………………………………………………………  P. subacida

• 1. Basidiospores dextrinoid…………………………………………………………………………………………………………………………………………………2

• 2. Pores < 5 per mm …………………………………………………………………………………………………………………………………………  P. ellipsospora

• 2. Pores > 5 per mm………………………………………………………………………………………………………………………………………………………… 3

• 3. Basidiocarps perennial…………………………………………………………………………………………………………………………………………P. africana

• 3. Basidiocarps annual to perennial ……………………………………………………………………………………………………………………… P. valliculorum

Rhizoperenniporia B.K. Cui & Xing Ji, gen. nov.

MycoBank: MB 847357

Differs from other genera by its resupinate basidiocarps with rhizomorphs, a dimitic hyphal system with weakly dextrinoid skeletal hyphae, and hyaline, thick-walled, ellipsoid, truncate, dextrinoid, and cyanophilous basidiospores.

Type species: Rhizoperenniporia japonica (Yasuda) B.K. Cui & Xing Ji

Etymology: Rhizoperenniporia (Lat.) refers to resembling Perenniporia but with rhizomorphs.

Basidiocarps are annual to perennial, resupinate, and corky when dry. Rhizomorphs are present. Pore surface is grayish white to pale buff when dry; pores are round; dissepiments thick, entire. Subiculum is thin, cream. Tubes are concolorous with pore surface and corky. Hyphal system is dimitic; generative hyphae with clamp connections; skeletal hyphae are dextrinoid and CB+; tissues are unchanged in KOH. Cystidia are absent; cystidioles are present. Basidiospores are ellipsoid, truncate, hyaline, thick-walled, smooth, dextrinoid, and CB+.

Notes: In our phylogenetic analyses (Figure 1 and Figure 2), two specimens of Perenniporia japonica (Yasuda) T. Hatt. & Ryvarden formed a single clade distant from the Perenniporia s. s. clade. In addition, P. japonica has basidiocarps with rhizomorphs, which are different from other species of Perenniporia s. s. Therefore, the new genus Rhizoperenniporia is proposed to include P. japonica.

Rhizoperenniporia japonica (Yasuda) B.K. Cui & Xing Ji, comb. nov.

MycoBank: MB 847387

Basionym: Trametes japonica Yasuda, Bot. Mag., Tokyo 32: 356 (1918).

= Perenniporia japonica (Yasuda) T. Hatt. & Ryvarden, Mycotaxon 50: 36 (1994).

For a detailed description of Perenniporia japonica, see Núñez and Ryvarden [2] and Cui et al. [3].

Notes: In Perenniporia s. l., P. aurantiaca, P. bambusicola, P. rhizomorpha, P. subrhizomorpha, and P. tibetica also have resupinate basidiocarps with rhizomorphs. However, P. aurantiaca differs from P. japonica by an orange pore surface, arboriform vegetative hyphae, and tissues becoming violet in KOH [4]. Perenniporia bambusicola is distinguished by an orange pore surface turning dark violet to black in KOH, arboriform vegetative hyphae, and growing only on bamboo [51]. Perenniporia rhizomorpha differs by its non-truncate basidiospores [10]. Perenniporia subrhizomorpha differs by the absence of cystidioles and larger basidiospores (5.7–6.5 × 4.3–5.5 μm) [56]. Perenniporia tibetica differs by having larger pores and basidiospores (pores 2–3 per mm, basidiospores 6.7–8.7 × 5.3–6.8 μm) [15].

Specimens examined: CHINA. Shanxi, Yangcheng County, Manghe Nature Reserve, on rotten wood of Vitex, 25 August 2016, Dai 17035 (BJFC); Huguan County, Baquan Gorge, on fallen trunk of Lonicera, 27 August 2016, Dai 17068, 17080 (BJFC).

Tropicoporia B.K. Cui & Xing Ji, gen. nov.

MycoBank: MB 847358

Differs from other genera by its dimitic to trimitic hyphal system with usually non-dextrinoid and inamyloid skeletal hyphae, broadly ellipsoid to subglobose, truncate, dextrinoid, and cyanophilous basidiospores.

Type species: Tropicoporia aridula (B.K. Cui & C.L. Zhao) B.K. Cui & Xing Ji

Etymology: Tropicoporia (Lat.) refers to the distribution of the genus in tropical areas.

Basidiocarps are annual to perennial, mostly resupinate, and pseudopileate to rarely pileate. Pore surface is cream, buff-yellow to grayish orange; pores are round to angular. Hyphal system is dimitic to trimitic; generative hyphae with clamp connections; skeletal hyphae are non-dextrinoid to slightly dextrinoid and CB+. Basidiospores are broadly ellipsoid to subglobose, truncate, hyaline, thick-walled, smooth, dextrinoid, and CB+.

Notes: The phylogenies inferred from the two combined datasets ITS + nLSU and ITS + nLSU + mtSSU + TEF1 + TBB1 showed that Perenniporia aridula B.K. Cui & C.L. Zhao, P. vanhulleae Decock & Ryvarden, P. centrali-africana Decock & Mossebo, and P. brasiliensis C.R.S. de Lira et al. grouped together and formed a clade distinct from the Perenniporia s. s. clade, although the branch support was low (Figure 1 and Figure 2). Morphologically, the four species usually have non-dextrinoid skeletal hyphae and broadly ellipsoid to subglobose basidiospores, which are different from the species of Perenniporia s. s.. Thus, Tropicoporia gen. nov. is proposed to accommodate the four species.

In our study, Tropicoporia was closely related to Sparsitubus and then grouped with Dendroporia with low support (Figure 1 and Figure 2). Morphologically, Sparsitubus differs from Tropicoporia by its effused-reflexed to pileate basidiocarps, and asperulate and non-truncate basidiospores [54]. Dendroporia is distinguished from Tropicoporia by its gray to pale brown pore surface, dextrinoid skeletal hyphae, and non-dextrinoid basidiospores.

Tropicoporia aridula (B.K. Cui & C.L. Zhao) B.K. Cui & Xing Ji, comb. nov.

MycoBank: MB 847388

Basionym: Perenniporia aridula B.K. Cui & C.L. Zhao, Fungal Diversity 58: 48 (2013)

For a detailed description of Perenniporia aridula, see Zhao et al. [19].

Notes: Tropicoporia aridula was first described in Perenniporia by Zhao et al. [19]. It is thus far known only from Southwest China. Tropicoporia vanhulleae is closely related to T. aridula in morphology and phylogeny, but the former has smaller basidiospores (5.5–6.0 × 4.5–5.5 µm) [21].

Specimens examined: CHINA. Yunnan, Yuanjiang County, on fallen angiosperm trunk, 9 June 2011, Dai 12396 (holotype, BJFC), on fallen bamboo, 9 June 2011, Dai 12398 (paratype, BJFC).

Tropicoporia brasiliensis (Lira, A.M.S. Soares, Ryvarden & Gibertoni) B.K. Cui & Xing Ji, comb. nov.

MycoBank: MB 847389

Basionym: Perenniporia brasiliensis Lira, A.M.S. Soares, Ryvarden & Gibertoni, Persoonia 38: 355 (2017).

For a detailed description of Perenniporia brasiliensis, see Lira et al in Crous et al. [23].

Notes: Tropicoporia brasiliensis was originally described from Brazil as Perenniporia brasiliensis by Lira et al (in Crous et al. [23]); it has subglobose to globose and small basidiospores (3–4 × 2–4 μm) [23].

Tropicoporia centrali-africana (Decock & Mossebo) B.K. Cui & Xing Ji, comb. nov.

MycoBank: MB 847390

Basionym: Perenniporia centrali-africana Decock & Mossebo, Systematics and Geography of Plants 71(2): 608 (2002).

For a detailed description of Perenniporia centrali-africana, see Decock and Mossebo [61].

Notes: Tropicoporia centrali-africana was originally described from Cameroon as Perenniporia centrali-africana [61]. Lira et al (in Crous et al. [23]) also reported this species from Brazil. It differs from the other three species by its pileate basidiocarps.

Tropicoporia vanhulleae (Decock & Ryvarden) B.K. Cui & Xing Ji, comb. nov.

MycoBank: MB 847391

Basionym: Perenniporia vanhulleae Decock & Ryvarden, Index Fungorum 234: 1 (2015)

For a detailed description of Perenniporia vanhulleae, see Decock and Ryvarden [21].

Notes: Tropicoporia vanhulleae was originally described from Africa as Perenniporia vanhulleae by Decock and Ryvarden [21]. The sequence of T. vanhulleae from the type specimens fell into Tropicoporia in our phylogeny. Therefore, Perenniporia vanhulleae is transferred to Tropicoporia.

Key to species of Tropicoporia

• 1. Basidiocarps resupinate to pileate………………………………………………………………………………………………………………… T. centrali-africana

• 1. Basidiocarps resupinate…………………………………………………………………………………………………………………………………………………2

• 2. Basidiospores < 5 µm………………………………………………………………………………………………………………………………………T. brasiliensis

• 2. Basidiospores > 5 µm……………………………………………………………………………………………………………………………………………………3

• 3. Basidiospores 6–7 μm………………………………………………………………………………………………………………………………………… T. aridula

• 3. Basidiospores 5.5–6.0 µm………………………………………………………………………………………………………………………………… T. vanhulleae

Truncatoporia B.K. Cui & Xing Ji, gen. nov.

MycoBank: MB 847359

Differs from other genera by its resupinate to pileate basidiocarps, a dimitic to trimitic hyphal system with dextrinoid and cyanophilous skeletal hyphae, and thick-walled, ellipsoid, truncate, and cyanophilous basidiospores.

Type species: Truncatoporia truncatospora (Lloyd) B.K. Cui & Xing Ji

Etymology: Truncatoporia (Lat.) refers to the truncate basidiospores of the genus.

Basidiocarps are annual to perennial, resupinate to pileate, and corky. Pileal surface is brown to ochraceous. Pore surface is buff to pale yellowish buff upon drying; pores are round to angular; dissepiments thin, entire. Context cream buff to pale brown. Tubes are concolorous with pore surface. Hyphal system is dimitic to trimitic; generative hyphae with clamp connections; skeletal hyphae are dextrinoid and CB+; tissues are unchanged in KOH. Basidiospores are ellipsoid, truncate, hyaline, thick-walled, smooth, dextrinoid or not, and CB+.

Notes: In our phylogenetic analyses, two species previously included in Perenniporia, P. pyricola Y.C. Dai & B.K. Cui and P. truncatospora (Lloyd) Ryvarden, grouped together and formed a well-supported clade (78% BS, 1.00 BPP, Figure 1; 98% BS, 1.00 BPP, Figure 2) distinct from the Perenniporia s. s. clade. Therefore, Truncatoporia gen. nov. is proposed to accommodate P. pyricola and P. truncatospora.

Truncatoporia pyricola (Y.C. Dai & B.K. Cui) B.K. Cui & Xing Ji, comb. nov.

MycoBank: MB 847392

Basionym: Perenniporia pyricola Y.C. Dai & B.K. Cui, Mycosystema 29(6): 815 (2010).

For a detailed description of Perenniporia pyricola, see Dai [62].

Notes: Truncatoporia pyricola was first described in Perenniporia from Northeast China [62]. It has a distribution in northern China mainly on Pyrus and Prunus. The species has perennial and resupinate basidiocarps with cream to pale cinnamon, a dimitic hyphal system with dextrinoid and cyanophilous skeletal hyphae, thick-walled and truncate, dextrinoid, and cyanophilous basidiospores.

Specimens examined: CHINA. Liaoning, Anshan, Qianshan Park, on living tree of Pyrus, 2 August 2008, Dai 10265 (holotype, BJFC); Tianjin, Ji County, Panshan Mountain, Living tree of Crataegus, 6 August 2015, Dai 15496 (BJFC), Dai 15498 (BJFC).

Truncatoporia truncatospora (Lloyd) B.K. Cui & Xing Ji, comb. nov.

MycoBank: MB 847393

Basionym: Trametes truncatospora Lloyd, Mycol. Writ. 6: 853 (1919).

= Perenniporia truncatospora (Lloyd) Ryvarden, Acta Mycol. Sin. 5: 228 (1986).

For a detailed description of Perenniporia truncatospora, see Cui et al. [3].

Notes: Truncatoporia truncatospora and T. pyricola both have dextrinoid skeletal hyphae and truncate basidiospores, but the former has pileate basidiocarps, smaller pores (6–8 per mm), and non-dextrinoid basidiospores, and the latter has resupinate basidiocarps, larger pores (3–5 per mm) [62], and dextrinoid basidiospores.

Specimen examined: CHINA. Sichuan, Ganluo County, Shengli, Gaoqiao Village, on living tree of Quercus, 14 September 2019, Cui 17770 (BJFC).

Key to species of Truncatoporia

• 1. Basidiocarps resupinate, 3–5 per mm………………………………………………………………………………………………………………………T. pyricola

• 1. Basidiocarps pileate, 6–8 per mm…………………………………………………………………………………………………………………… T. truncatospora

Vanderbyliella B.K. Cui & Xing Ji, gen. nov.

MycoBank: MB 847360

Differs from other genera by its pileate basidiocarps with an orange brown pileal surface, a dimitic hyphal system with strongly dextrinoid skeletal hyphae, and hyaline, thick-walled, ellipsoid, non-truncate, and cyanophilous basidiospores.

Type species: Vanderbyliella tianmuensis (B.K. Cui & C.L. Zhao) B.K. Cui & Xing Ji

Etymology: Vanderbyliella (Lat.) refers to the morphological similarity to Vanderbylia.

Basidiocarps are annual to perennial, pileate, and hard corky to woody hard when dry. Pileal surface is clay-buff, orange-brown to yellowish brown, glabrous, and concentrically sulcate. Pore surface is buff to pale brown when dry; pores are round to angular; dissepiments thin, entire. Context cream to pale brown, corky to hard corky. Tubes are buff to pale brown, hard corky to woody hard. Hyphal system is dimitic; generative hyphae with clamp connections; skeletal hyphae are strongly dextrinoid and CB+; tissues are unchanged in KOH. Cystidia are absent; cystidioles are present. Basidiospores are ellipsoid, non-truncate, hyaline, thick-walled, smooth, dextrinoid or not, and CB+.

Notes: In our phylogenetic studies, Perenniporia tianmuensis B.K. Cui & C.L. Zhao and an unknown species, grouped together and formed a strongly supported clade (99% BS, 1.00 BPP, Figure 1; 100% BS, 1.00 BPP, Figure 2), which was distant from the Perenniporia s. s. clade. Morphologically, these two species have pileate basidiocarps and non-truncate basidiospores, which are different from species of Perenniporia s. s., so Vanderbyliella gen. nov. is proposed.

Vanderbyliella tianmuensis (B.K. Cui & C.L. Zhao) B.K. Cui & Xing Ji, comb. nov.

MycoBank: MB 847394

Basionym: Perenniporia tianmuensis B.K. Cui & C.L. Zhao, Mycoscience 54: 236 (2013).

For a detailed description of Perenniporia tianmuensis, see Zhao and Cui [18].

Notes: Vanderbyliella tianmuensis was first described in Perenniporia from China by Zhao and Cui [18]; it is characterized by annual and pileate basidiocarps, a dimitic hyphal system with strongly dextrinoid skeletal hyphae, and thick-walled, ellipsoid, non-truncate, dextrinoid, and cyanophilous basidiospores.

Specimens examined: CHINA. Zhejiang, Lin’an County, Tianmushan Nature Reserve, on the basis of dead angiosperm trees, 10 October 2005, Cui 2648 (holotype, BJFC).

Xanthoperenniporia B.K. Cui & Xing Ji, gen. nov.

MycoBank: MB 847361

Differs from other genera by its resupinate basidiocarps with yellow pore surface, weakly dextrinoid, and cyanophilous skeletal hyphae, hyaline, thick-walled, ellipsoid, truncate, and cyanophilous basidiospores.

Type species: Xanthoperenniporia tenuis (Schwein.) B.K. Cui & Xing Ji

Etymology: Xanthoperenniporia (Lat.) refers to resembling Tropicoporia but with a yellow pore surface.

Basidiocarps are annual to perennial, resupinate to reflexed-effused, and corky when dry. Pore surface is cream to yellow when fresh, buff to yellow; pores are round to angular. Subiculum is thin, cream, buff to pale yellowish brown. Tubes are concolorous with pore surface. Hyphal system is dimitic to trimitic, generative hyphae with clamp connections; skeletal hyphae are dextrinoid or weakly dextrinoid and CB+; tissues are unchanged in KOH. Cystidia are absent, cystidioles are usually present. Basidiospores are ellipsoid, truncate, hyaline, thick-walled, smooth, dextrinoid or not, and CB+.

Notes: In the combined five-gene phylogeny, Perenniporia maackiae (Bondartsev & Ljub.) Parmasto, P. punctata Hai J. Li & Jing Si, P. subcorticola Chao G. Wang & F. Wu and P. tenuis (Schwein.) Ryvarden clustered together and formed a single clade with good support (74% BS, 0.96 BPP, Figure 1; 89% BS, 1.00 BPP, Figure 2), which was distant from the Perenniporia s. s. clade. Morphologically, these species differ from Perenniporia s. s. in having a yellow pore surface. Therefore, the new genus Xanthoperenniporia is set up, and four new combinations are proposed.

Xanthoperenniporia maackiae (Bondartsev & Ljub.) B.K. Cui & Xing Ji, comb. nov.

MycoBank: MB 847395

Basionym: Fomitopsis maackiae Bondartsev & Ljub., Botanicheskie Materialy 15: 103 (1962).

= Perenniporia maackiae (Bondartsev & Ljub.) Parmasto, Ann. Bot. fenn. 32(4): 223 (1995).

For a detailed description of Perenniporia maackiae, see Cui et al. [3].

Notes: Xanthoperenniporia maackiae grows mainly on Maackia. Xanthoperenniporia subcorticola is similar to X. maackiae by sharing resupinate basidiocarps with yellow pore surfaces, similar sized pores, and the presence of cystidioles, but X. subcorticola has smaller basidiospores (4.2–5 × 3.5–4.2 µm) [29], and X. maackiae has larger basidiospores (5.4–6.3 × 3.8–5.0 µm).

Specimens examined: CHINA. Heilongjiang, Yichun, Dailing, Liangshui Nature Reserve, on fallen branch of Maackia, 26 August 2014, Cui 11531 (BJFC). Jilin, Antu County, Changbaishan Nature Reserve, on dead tree of Maackia, 11 September 2014, Dai 14780 (BJFC).

Xanthoperenniporia punctata (Hai J. Li & Jing Si) B.K. Cui & Xing Ji, comb. nov.

MycoBank: MB 847396

Basionym: Perenniporia punctata Hai J. Li & Jing Si, Phytotaxa 360(1): 56 (2018).

For a detailed description of Perenniporia punctata, see Li et al. [26].

Notes: Xanthoperenniporia punctata was recently described in Perenniporia from China by Li et al. [26] and is characterized by annual and resupinate basidiocarps with a buff-yellow pore surface, a dimitic hyphal system with slightly dextrinoid skeletal hyphae, broadly ellipsoid to subglobose, and truncate and non-dextrinoid basidiospores.

Specimens examined: CHINA. Hubei, Yichang, Wufeng County, Chaibuxi National Forestry Park, on angiosperm stump, 15 August 2017, Dai 17923 (holotype, BJFC), on rotten wood of Quercus, 14 August 2017, Dai 17916 (paratype, BJFC).

Xanthoperenniporia subcorticola (Chao G. Wang & F. Wu) B.K. Cui & Xing Ji, comb. nov.

MycoBank: MB 847397

Basionym: Perenniporia subcorticola Chao G. Wang & F. Wu, MycoKeys 69: 62 (2020).

For a detailed description of Perenniporia subcorticola, see Wang et al. [29].

Notes: Xanthoperenniporia subcorticola was recently described in Perenniporia as P. subcorticola Chao G. Wang & F. Wu by Wang et al. [29]. It is similar to Perenniporia corticola by having a yellow pore surface, dimitic hyphal system, and truncate and dextrinoid basidiospores of almost the same size, but P. corticola differs from P. subcorticola by having arboriform skeletal hyphae and dendrohyphidia [53].

Specimens examined: CHINA. Fujian, Wuyishan County, Wuyishan Nature Reserve, on rotten wood of Pinus, 21 October 2005, Dai 7330 (holotype, BJFC).

Xanthoperenniporia tenuis (Schwein.) B.K. Cui & Xing Ji, comb. nov.

MycoBank: MB 847398

Basionym: Polyporus tenuis Schwein., Trans. Am. phil. Soc., New Series 4: 159 (1832).

= Perenniporia tenuis (Schwein.) Ryvarden, Norw. J Bot. 20: 9 (1973).

For a detailed description of Perenniporia tenuis, see Cui et al. [3].

Notes: Xanthoperenniporia punctata is similar to X. tenuis in having resupinate basidiocarps, pale yellow to buff-yellow pore surface, and similar sized basidiospores, but the former differs in its smaller pores (6–9 per mm), and the absence of cystidioles and non-dextrinoid basidiospores [26], the latter has larger pores (4–6 per mm), the presence of cystidioles, and dextrinoid basidiospores.

Specimens examined: CHINA. Hubei, Yichang, Wufeng County, Chaibuxi National Forestry Park, on rotten angiosperm stump, 15 August 2017, Dai 17935 (BJFC). Shanxi, Yangcheng County, Manghe Nature Reserve, on fallen trunk of Vitex, 25 August 2016, Dai 17026 (BJFC).

Key to species of Xanthoperenniporia

• 1. Basidiospores non-dextrinoid .……………………………………………………………………………………………………………………………… X. punctata

• 1. Basidiospores dextrinoid …………………………………………………………………………………………………………………………………………………2

• 2. Basidiocarps resupinate to reflexed-effused, growing on Maackia……………………………………………………………………………………… X. maackiae

• 2. Basidiocarps resupinate, growing on other trees………………………………………………………………………………………………………………………3

• 3. Basidiocarps annual, pores 4–6 per mm ……………………………………………………………………………………………………………………… X. tenuis

• 3. Basidiocarps perennial, pores 7–8 per mm………………………………………………………………………………………………………………X. subcorticola

Yuchengia B.K. Cui & K.T. Steffen, Nordic J. Bot. 31(3): 333 (2013).

MycoBank: MB 563490

Type species: Yuchengia narymica (Pilát) B.K. Cui, C.L. Zhao & K.T. Steffen, Nordic J. Bot. 31(3): 333 (2013).

Basidiocarps are annual to perennial, resupinate, corky when fresh, and hard corky when dry. Pore surface is cream, yellowish buff to tan; pores are round to angular, dissepiments thin, entire. Subiculum is cream to pale ochraceous. Tubes are concolorous with pore surface. Hyphal system is dimitic; generative hyphae with clamp connections; skeletal hyphae amyloid or not, acyanophilous or weakly cyanophilous, dissolving in KOH. Cystidia are absent; cystidioles are present. Basidiospores are ellipsoid, truncate or not, hyaline, thick-walled, smooth, IKI−, and CB+.

Notes: The type species Yuchengia narymica was first described as Trametes narymica Pilát [63] and later was transferred to Perenniporia by Pouzar [64]. Zhao et al. [35] proposed Yuchengia to accommodate Perenniporia narymica based on molecular data and morphological characteristics.

Yuchengia kilemariensis (Spirin & Shirokov) B.K. Cui & Xing Ji, comb. nov.

MycoBank: MB 847399

Basionym: Perenniporia kilemariensis Spirin & Shirokov, Folia Cryptogamica Petropolitana (Sankt-Peterburg) 6: 38 (2005).

For a detailed description of Perenniporia kilemariensis, see Spirin et al. [9].

Notes: Yuchengia kilemariensis was originally described from Russia as Perenniporia kilemariensis by Spirin et al. [9]. It has inamyloid skeletal hyphae and truncate basidiospores, which is different from the original descriptions of Yuchengia. However, it has resupinate basidiocarps, a dimitic hyphal system with skeletal hyphae dissolving in KOH, and ellipsoid and non-dextrinoid basidiospores. These characters fit Yuchengia well. Moreover, in ITS + nLSU and five-gene phylogenetic analysis, the sequence of Y. kilemariensis from the type specimen fell into Yuchengia (Figure 1 and Figure 2). Therefore, P. kilemariensis is transferred to Yuchengia.

Key to species of Yuchengia

• 1. Skeletal hyphae amyloid …………………………………………………………………………………………………………………………………… Y. narymica

• 1. Skeletal hyphae inamyloid……………………………………………………………………………………………………………………………… Y. kilemariensis

Microporellus subadustus (Z.S. Bi & G.Y. Zheng) B.K. Cui & Xing Ji, comb. nov.

MycoBank: MB 847400

Basionym: Wrightoporia subadusta Z.S. Bi & G.Y. Zheng, Bull. Bot. Res., Harbin 7(4): 76 (1987).

= Perenniporia subadusta (Z.S. Bi & G.Y. Zheng) Y.C. Dai, Ann. Bot. fenn. 39(3): 180 (2002).

= Murinicarpus subadustus (Z.S. Bi & G.Y. Zheng) B.K. Cui & Y.C. Dai, Fungal Di-versity 97: 255 (2019).

= Perenniporia cystidiata Y.C. Dai, W.N. Chou & Sheng H. Wu, Mycotaxon 83: 209 (2002).

Notes: After studying type materials of Wrightoporia subadusta and Perennipori cystidiata, Cui et al. [3] found that the two represent the same species and separated this species from Perenniporia and proposed Murinicarpus. Murinicarpus has the same characters with Microporellus Murrill, stipitate basidiocarps, dextrinoid skeletal hyphae, non-truncate and non-dextrinoid basidiospores [33], and it is treated as a synonym of Microporellus in the current study.

Key to species of Perenniporia and related genera

• 1. Basidiocarps stipitate………………………………………………………………………………………………………………………………………Microporellus

• 1. Basidiocarps resupinate, effused-reflexed to pileate…………………………………………………………………………………………………………………2

• 2. Basidiospores pale yellowish  …………………………………………………………………………………………………………………………… Abundisporus

• 2. Basidiospores hyaline…………………………………………………………………………………………………………………………………………………… 3

• 3. Skeletal hyphae amyloid…………………………………………………………………………………………………………………………………………………4

• 3. Skeletal hyphae inamyloid………………………………………………………………………………………………………………………………………………6

• 4. Basidiocarps pileate……………………………………………………………………………………………………………………………………………Minoporus

• 4. Basidiocarps resupinate to effused-reflexed ………………………………………………………………………………………………………………………… 5

• 5. Basidiospores amyloid………………………………………………………………………………………………………………………………………Amylosporia

• 5. Basidiospores inamyloid………………………………………………………………………………………………………………………………………Yuchengia

• 6. Basidiocarps with rhizomorphs .……………………………………………………………………………………………………………………………………… 7

• 6. Basidiocarps without rhizomorphs…………………………………………………………………………………………………………………………………… 8

• 7. Pore surface orange to orange-brown  ………………………………………………………………………………………………………………… Aurantioporia

• 7. Pore surface grayish white to pale buff…………………………………………………………………………………………………………… Rhizoperenniporia

• 8. Cystidia present………………………………………………………………………………………………………………………………………………………… 9

• 8. Cystidia absent………………………………………………………………………………………………………………………………………………………… 10

• 9. Basidiocarps pileate……………………………………………………………………………………………………………………………………Hornodermoporus

• 9. Basidiocarps resupinate……………………………………………………………………………………………………………………………………Cystidioporia

• 10. Basidiospores non-truncate………………………………………………………………………………………………………………………………………… 11

• 10. Basidiospores truncate……………………………………………………………………………………………………………………………………………… 15

• 11. Basidiocarps pileate ………………………………………………………………………………………………………………………………………………… 12

• 11. Basidiocarps resupinate .…………………………………………………………………………………………………………………………………………… 13

• 12. Basidiospores obovoid …………………………………………………………………………………………………………………………………… Vanderbylia

• 12. Basidiospores ellipsoid ………………………………………………………………………………………………………………………………… Vanderbyliella

• 13. Basidiocarps annual……………………………………………………………………………………………………………………………………………Neoporia

• 13. Basidiocarps annual to perennial……………………………………………………………………………………………………………………………………14

• 14. Hyphal system dimitic………………………………………………………………………………………………………………………………Luteoperenniporia

• 14. Hyphal system dimitic to trimitic …………………………………………………………………………………………………………………………… Poriella

• 15. Basidiocarps osseous ………………………………………………………………………………………………………………………………… Perenniporiopsis

• 15. Basidiocarps corky to woody hard…………………………………………………………………………………………………………………………………16

• 16. Pore surface white when fresh, usually with reddish brown sterile margin…………………………………………………………………………Niveoporia

• 16. Pore surface cream, buff, yellowish to cinnamon, without reddish brown sterile margin………………………………………………………………… 17

• 17. Basidiocarps resupinate to pileate………………………………………………………………………………………………………………………………… 18

• 17. Basidiocarps resupinate  …………………………………………………………………………………………………………………………………………… 19

• 18. Basidiospores > 9 µm in length ………………………………………………………………………………………………………………………… Truncospora

• 18. Basidiospores < 9 µm in length…………………………………………………………………………………………………………………………Truncatoporia

• 19. Tissues brown to black in KOH …………………………………………………………………………………………………………………………………… 20

• 19. Tissues unchanged in KOH …………………………………………………………………………………………………………………………………………21

• 20. Dendrohyphidia present at dissepiment edges, basidiospores non-dextrinoid……………………………………………………………………Dendroporia

• 20. Dendrohyphidia usually absent at dissepiment edges, basidiospores dextrinoid…………………………………………………………………Citrinoporia

• 21. Pore surface yellow ……………………………………………………………………………………………………………………………… Xanthoperenniporia

• 21. Pore surface usually cream to cinnamon………………………………………………………………………………………………………………………… 22

• 22. Basidiospores ≥ 9 µm in length ………………………………………………………………………………………………………………………… Macrosporia

• 22. Basidiospores ≤ 9 µm in length …………………………………………………………………………………………………………………………………… 23

• 23. Skeletal hyphae usually non-dextrinoid ……………………………………………………………………………………………………………… Tropicoporia

• 23. Skeletal hyphae usually dextrinoid ……………………………………………………………………………………………………………………………… 24

• 24. Basidiospores < 6 µm in length ………………………………………………………………………………………………………………………… Perenniporia

• 24. Basidiospores > 6 µm in length .………………………………………………………………………………………………………………………… Macroporia

4. Discussion

The phylogenetic analyses carried out by Binder et al. [45] placed Perenniporia in the core polyporoid clade, with species of Perenniporia forming more than one lineage. Our phylogenetic analyses confirmed that Perenniporia s. l. is polyphyletic and is nested in the core polyporoid clade.

Abundisporus Ryvarden is closely related to some Perenniporia species. Ryvarden [65] established Abundisporus to accommodate the species with pale yellowish and non-dextrinoid basidiospores, previously accepted in Loweporus J.E. Wright. Our phylogenetic analyses showed that Abundisporus is closely related to Cystidioporia, Macrosporia, and Niveoporia, but distant from the Perenniporia s. s. as previous studies [17,19,31]. Morphologically, Abundisporus is distinguished from these genera by its more or less pinkish basidiocarps, non-truncate, and pale yellowish basidiospores [66].

Truncospora, typified by T. ochroleuca (Berk.) Pilát, was established by Pilát [67]. The genus is characterized by pileate basidiocarps, a dimitic hyphal system with clamped generative hyphae, dextrinoid and cyanophilous skeletal hyphae, and hyaline, thick-walled, truncate, ellipsoid, and strongly dextrinoid basidiospores [68,69]. These characters are the same as those of some species in Perenniporia s. l. This genus has often been treated as a synonym of Perenniporia since its establishment [32,33,34,70]. Decock and Ryvarden [5] considered that Perenniporia detrita (Berk.) Ryvarden, P. ochroleuca (Berk.) Pilát and P. ohiensis (Berk.) Ryvarden formed a morphologically homogeneous group that could be recognized at the genus level. Later phylogeny showed that these three species formed a monophyletic clade that was distinct from Perenniporia s. s. [17,19,31]. Morphologically, Truncospora mainly differs from Perenniporia s. s. by having button-shaped to ungulate basidiocarps, variably dextrinoid skeletal hyphae, and longly ovoid basidiospores [68].

Vanderbylia was established by Reid [71]; species in this genus have pileate and hard corky basidiocarps, a dimitic hyphal system with clamped generative hyphae, dextrinoid and cyanophilous skeletal hyphae, and hyaline, thick-walled, non-truncate, subglobose to obovoid, dextrinoid, and cyanophilous basidiospores [3,17]. It was treated as a synonymy of Perenniporia [32,72]. Recent phylogenetic analyses revealed that Vanderbylia is an independent genus that is distant from Perenniporia s. s. [3,17,18,19]. In our ITS + nLSU phylogeny, Vanderbylia is closely related to Vanderbyliella, but the latter has ellipsoid and dextrinoid or non-dextrinoid basidiospores. Vanderbylia differs from Perenniporia s. s. by its pileate basidiocarps, dextrinoid skeletal hyphae, and non-truncate and subglobose to amygdaliform basidiospores.

Hornodermoporus was established by Teixeira [73] and typified by H. martius (Berk.) Teixeira. The genus is characterized by its pileate basidiocarps with a black crust at the pileal surface, a dimitic hyphal system with strongly dextrinoid and cyanophilous skeletal hyphae, the presence of cystidia, truncate, oblong-ellipsoid, and strongly dextrinoid basidiospores. Hornodermoporus was treated as a synonym of Perenniporia [1,58], but morphologically it differs from Perenniporia s. s. in having pileate basidiocarps with a black crusted pileal surface, dextrinoid skeletal hyphae, the presence of cystidia, and oblong-ellipsoid basidiospores. In our phylogeny, the separation of Hornodermoporus from Perenniporia is supported, as previously reported [3,19,20,36,74].

Amylosporia, Murinicarpus, Perenniporiopsis, Poriella, and Yuchengia have recently been segregated from Perenniporia s. l. These genera were phylogenetically distant from Perenniporia s. s. in our study. Amylosporia differs from Perenniporia s. s. by its amyloid skeletal hyphae and basidiospores [3]. Amylosporia hattorii and Perenniporia amylodextrinoidea Gilb. & Ryvarden both have amyloid skeletal hyphae, but the former has amyloid and larger basidiospores (10–12 × 5.5–7.5 μm) [3], and the latter has dextrinoid and smaller basidiospores (4.5–5.5 × 3–3.5 μm) [33]. Murinicarpus, newly proposed by Cui et al. [3] based on Wrightoporia subadusta = Perenniporia cystidiata, has same characters with Microporellus and is a synonym of Microporellus. It is different from Perenniporia s. s. by its stipitate basidiocarps, dextrinoid skeletal hyphae, presence of thick-walled cystidia, non-truncate and non-dextrinoid basidiospores [3]. Perenniporiopsis is distinguished from Perenniporia s. s. by its basidiocarps waxy when fresh and rigidly osseous when dry, dextrinoid skeletal hyphae, and large basidiospores [36]. Chen et al. [37] described Poriella as having non-dextrinoid basidiospores. Perenniporia africana and P. ellipsospora are congeneric, although they have dextrinoid basidiospores; they are transferred to Poriella. Poriella is distinguished from Perenniporia s. s. by its unbranched, dextrinoid skeletal hyphae, and non-truncate basidiospores. Perenniporia kilemariensis, which has inamyloid skeletal hyphae and truncate basidiospores is transferred to Yuchengia, so the definition of Yuchengia is revised also in this study. Yuchengia differs from Perenniporia s. s. by its non-dextrinoid basidiospores, unbranched “amyloid” skeletal hyphae.

The segregation of Aurantioporia, Citrinoporia, Cystidioporia, Dendroporia, Luteoperenniporia, Macroporia, Macrosporia, Minoporus, Neoporia, Niveoporia, Rhizoperenniporia, Tropicoporia, Truncatoporia, Vanderbyliella, and Xanthoperenniporia from Perenniporia s. l. was well supported by phylogenetic analyses of ITS + nLSU and ITS + nLSU + mtSSU + TEF1 + TBB1. In the ITS + nLSU gene phylogenetic tree, Yuchengia is closely related to Poriella; in the combined five-gene phylogenetic analysis, Yuchengia was related to Minoporus, Neoporia, Poriella, and Vanderbyliella. However, Yuchengia has resupinate basidiocarps, amyloid or not, acyanophilous or weakly cyanophilous skeletal hyphae, and non-dextrinoid basidiospores; Minoporus produces pileate basidiocarps, cyanophilous skeletal hyphae, and truncate and dextrinoid basidiospores; Neoporia has dextrinoid and cyanophilous skeletal hyphae and dextrinoid basidiospores; Poriella has dextrinoid and cyanophilous skeletal hyphae [37]; Vanderbyliella has pileate basidiocarps and dextrinoid and cyanophilous skeletal hyphae.

According to phylogenetic analyses and morphological characters, some species of Perenniporia s. l. have a new generic placement. For four species in the phylogenetic tree, including Perenniporia tephropora (Mont.) Ryvarden, P. subtephropora B.K. Cui & C.L. Zhao, P. eugeissonae P. Du & Chao G. Wang and P. straminea (Bres.) Ryvarden, due to unclear phylogenetic relationships and insufficient morphological difference from Perenniporia s. s. species, their taxonomic positions were remained in Perenniporia s. l.

In summary, we carried out a comprehensive study on Perenniporia s. l., 44 species of Perenniporia s. l. with available sequences. According to phylogenetic evidence and morphological characteristics, 15 new genera were set up, 2 new species were described, and 37 new combinations were proposed. Species lacking reliable molecular sequences were not included in this study.

Author Contributions

Conceptualization, B.-K.C. and X.J.; methodology, B.-K.C., X.J., Y.-F.S., D.-M.W. and N.G.; validation, X.J. and B.-K.C.; formal analysis, X.J.; performing the experiments, X.J.; resources, B.-K.C.; writing—original draft preparation, X.J.; writing—review and editing, B.-K.C.; visualization, X.J.; supervision, B.-K.C.; project administration, B.-K.C.; funding acquisition, B.-K.C. All authors have read and agreed to the published version of the manuscript.

Institutional Review Board Statement

Not applicable.

Informed Consent Statement

Not applicable.

Data Availability Statement

Publicly available datasets were analyzed in this study. These data can be found here: https://www.ncbi.nlm.nih.gov/genbank/, accessed on 24 August 2022; http://purl.org/phylo/treebase, accessed on 29 November 2022; http://www.mycobank.org/, accessed on 24 January 2023.

Conflicts of Interest

The authors declare no conflict of interest.

Funding Statement

The research is supported by the National Natural Science Foundation of China (Nos. 32270010, U2003211), the Scientific and Technological Tackling Plan for the Key Fields of Xinjiang Production and Construction Corps (No. 2021AB004) and Beijing Forestry University Outstanding Young Talent Cultivation Project (No. 2019JQ03016).