What Are “Lingzhi Wang” or “Zhu Lingzhi”? Notes on Ganoderma (Ganodermataceae, Polyporales) Species Characterized by Diminutive Pilei and Gracile Stipes from Hainan Island, Tropical China

Abstract

Species of Ganoderma (Ganodermataceae, Polyporales) have been extensively utilized in traditional Chinese medicine for over two millennia, owing to their remarkable medicinal properties and diverse chemical constituents. Hainan Island, located in tropical China, harbors a rich diversity of Ganoderma species. Among these, certain varieties referred to as “Lingzhi Wang” or “Zhu Lingzhi” by indigenous communities are distinguished by their diminutive pilei and slender stipes. Despite their traditional recognition, these species have been subject to morphological confusion. In this study, specimens labeled as “Lingzhi Wang” or “Zhu Lingzhi” were subjected to comprehensive morphological examinations and molecular phylogenetic analyses. The findings reveal that the Ganoderma species characterized by small pilei and gracile stipes encompass at least five distinct species. Among these, two are new to science: G. baisuzhenii and G. shennongii. The remaining three species, G. bambusicola, G. flexipes, and G. subflexipes, have been previously described. Taxonomically, G. bambusicola was reported for the first time on the Chinese mainland. This study provides a clearer taxonomic framework for these medicinally significant fungi.

1. Introduction

The family Ganodermataceae (Polyporales, Basidiomycota) is a globally distributed taxonomic group comprising fourteen accepted genera: Amauroderma Murrill, Amaurodermellus Costa-Rezende, Drechsler-Santos & Góes-Neto, Cristataspora Robledo & Costa-Rezende, Foraminispora Robledo, Costa-Rez. & Drechsler-Santos, Furtadoella B.K. Cui & Y.F. Sun, Ganoderma P. Karst., Haddowia Steyaert, Humphreya Steyaert, Magoderna Steyaert, Neoganoderma B.K. Cui & Y.F. Sun, Sanguinoderma Y.F. Sun, D.H. Costa & B.K. Cui, Sinoganoderma B.K. Cui, J.H. Xing & Y.F. Sun, Tomophagus Murrill, and Trachydermella B.K. Cui & Y.F. Sun [1,2]. Among these, the genus Ganoderma, typified by G. lucidum (Curtis) P. Karst., is the most widely recognized. It is characterized by sessile to stipitate basidiomata, double-walled and truncated basidiospores, and its wood-decaying properties [2,3]. According to the Index Fungorum database (http://www.indexfungorum.org/, accessed on 25 January 2025), there are 498 records under Ganoderma, although many of these epithets are synonyms [1,4]. In China, more than 40 species of this genus have been accepted to date [1,2,5,6,7,8,9,10].

For more than a century, the highly valued medicinal fungus “Lingzhi” or “Ruizhi”, widely cultivated and utilized in China, including Hainan Island, has been taxonomically classified as G. lucidum (Curtis) P. Karst., a species first described in Europe. However, recent studies have revealed that the famous “G. lucidum” in China is genetically and taxonomically distinct from the authentic European G. lucidum. This Chinese species has been scientifically named G. lingzhi S.H. Wu, Y. Cao & Y.C. Dai [10,11]. Despite this, Yao et al. and Du et al. have argued that the correct name for the widely recognized Ganoderma species by the public in China should be G. sichuanense J.D. Zhao & X.Q. Zhang, with G. lingzhi considered a later synonym [12,13]. The scientific nomenclature of this fungus remains a contentious issue, with ongoing debates between G. lingzhi and G. sichuanense.

Ganoderma species, including G. lingzhi or G. sichuanense, have been used in traditional Chinese medicine for more than 6800 years [14,15,16]. These fungi are celebrated for their rich array of natural bioactive compounds, which exhibit a broad spectrum of pharmacological properties, including immunoregulatory, anti-tumor, anti-fatigue, gut microbiota regulation, and hepatocyte protection activities [17,18,19]. Subtropical and tropical regions of China, particularly Hainan Island, are biodiversity hotspots for Ganoderma species [20,21]. In these regions, numerous wild Ganoderma species are commercially traded [22], and their fruiting bodies are commonly used by local residents, either soaked in wine or boiled in water, for health preservation and disease treatment [23].

Among the Ganoderma species in Hainan Island, certain varieties are locally known as “Lingzhi Wang” (king of Ganoderma) or “Zhu Lingzhi” (bamboo host of Ganoderma). These species are distinguished by their diminutive pilei and gracile stipes. Due to their high popularity in the region (based on our investigations), they are often subject to morphological confusion, with homonyms (different Ganoderma species sharing the same name) and synonyms (the same Ganoderma species referred to by different names) being common. To address these taxonomic challenges, this study conducted comprehensive field investigations, key informant interviews, morphological examinations, and molecular phylogenetic analyses to accurately identify and clarify the taxonomy of these species.

2. Materials and Methods

2.1. Key Informant Interview Methodology

Key informants were selected based on their expertise and involvement in the collection, trade, or traditional use of Ganoderma species on Hainan Island. The interviews were semi-structured, combining open-ended questions with a predefined interview guide to ensure flexibility and depth. The interview guide covered the following topics: background information, traditional knowledge, harvesting and trade practices, cultural significance, and conservation perspectives.

2.2. Sample Collection and Morphological Studies

Fresh basidiomata of Ganoderma species were collected during the rainfall seasons in southern China, particularly on Hainan Island. Specimens were photographed in situ at their collection sites to document their natural habitat and morphological features. Detailed morphological characteristics, including color, shape, and size, were recorded from fresh samples. The collected specimens were then dried at approximately 60 °C overnight to ensure complete dehydration and were subsequently deposited in the Hainan Biodiversity Science and Technology Museum (FHMU) or the Cryptogamic Herbarium (HKAS) of the Kunming Institute of Botany, Chinese Academy of Sciences. Color descriptions were standardized using the color codes provided by Kornerup and Wanscher [24].

For microscopic analysis, thin sections were prepared freehand using a razor blade and mounted in 5% potassium hydroxide (KOH) solution or Cotton Blue and Melzer’s reagent. These sections were examined under an OLYMPUS CX23 optical microscope to observe and measure microscopic features. All microscopic structures were illustrated freehand based on the observations. The following notations and abbreviations were used in this study: IKI (Melzer’s reagent; IKI– indicates neither amyloid nor dextrinoid reactions); CB (Cotton Blue; CB+ denotes cyanophilous reactions); and basidiospore measurements are expressed as n/m/p, where n = the total number of basidiospores measured from the m basidiomata of p collections. Basidiospore dimensions are presented in the format (a)b–c(d), where b–c represents the range encompassing at least 90% of the measured values, and extreme values (a and d, when present) are given in parentheses. Additional abbreviations include L (mean basidiospore length), W (mean basidiospore width), and Q (quotient of basidiospore length to width, L/W). All measurements were based on a minimum of 20 basidiospores per species.

2.3. Molecular Procedures

Genomic DNA was extracted from dried basidiomata samples using the Magnetic Beads Genomic DNA Extraction Kit (Magen, Guangzhou, China) in accordance with the manufacturer’s protocol. The ITS region, including internal transcribed spacers 1 and 2, along with the 5.8S rDNA, was amplified using the primer pair ITS5/ITS4 [25]. Additionally, the partial RNA polymerase second-largest subunit region (rpb2) was amplified using primers bRPB2-6F/bRPB2-7.1R [26], and the partial translation elongation factor 1-α (tef1) region was amplified using primers EF1-F and EF1-R [27]. All PCR reactions were conducted in 25 μL volumes, containing 13 µL of 2× Taq PCR MasterMix (KANGWEI Company, Guangzhou, China), 2 µL of each primer (10 mM), 2 µL of DNA template, and 8 µL of nuclease-free water. The PCR conditions for all genetic regions included an initial denaturation step at 95 °C for 4 min, followed by 35 cycles of denaturation at 94 °C for 30 s, annealing at specific temperatures (50 °C for ITS, 53 °C for tef1, and 52 °C for rpb2) for 30 s, extension at 72 °C for 120 s, and a final extension at 72 °C for 7 min. PCR products were visualized on 1% (w/v) agarose gels, and those exhibiting a bright single band were purified and sequenced using an ABI 3730xl DNA Analyzer (Guangzhou Branch of BGI, Guangzhou, China) with the same primers used for amplification [28]. The accuracy of the newly obtained sequences was confirmed by comparison with sequences available in GenBank [https://www.ncbi.nlm.nih.gov/genbank/ (accessed on 1 February 2025)]. The sequences were assembled and edited using BioEdit v7.0.9.0 [29] and subsequently deposited in GenBank (Table 1).

2.4. Dataset Assembly

The phylogenetic analyses were conducted using a sequence dataset comprising three loci: ITS, rpb2, and tef1. Foraminispora concentrica (J. Song, Xiao L. He & B.K. Cui) Y.F. Sun & B.K. Cui was selected as the outgroup, which was suggested by Sun et al. [30]. Additionally, other reference taxa for the phylogenetic analysis were retrieved from GenBank and relevant publications, as detailed in Table 1. To assess potential phylogenetic conflicts among the different gene regions, single-gene phylogenetic trees were constructed for ITS, rpb2, and tef1 separately using the maximum likelihood (ML) method. The results indicated no significant conflicts among the gene fragments, supporting their combination for further analysis. The sequences of the three loci were aligned using MUSCLE v. 3.6 [31] and subsequently concatenated into a single dataset using Phyutility v. 2.2 [32] for comprehensive phylogenetic analysis.

Table 1. Taxa, vouchers, locations, and GenBank accession numbers of DNA sequences used in this study.

			GenBank Accession Nos.
Taxon	Voucher	Locality	ITS	rpb2	tef1	Reference
Cristataspora coffeata	FLOR 50933	Brazil	KU315204	—	—	[33]
C. coffeata	Robledo 3183	Brazil	MN077526	—	MN061695	[34]
C. coffeata	Robledo 3182	Brazil	MN077525	—	—	[34]
C. flavipora	Robledo 3288	Argentina	MN077521	—	MN061694	[34]
Foraminispora concentrica	Cui 16238	Yunnan, SW China	MK119816	MK121504	MK121565	[30]
F. concentrica	Cui 16239	Yunnan, SW China	MK119817	MK121506	MK121566	[30]
Ganoderma acaciicola	Cui 16815	Australia	MZ354895	MZ245384	—	[2]
G. acaciicola	Cui 16814	Australia	MZ354894	MZ245383	—	[2]
G. acontextum	JV 0611/21G	Guatemala	KF605667	MG367489	MG367538	[2]
G. acontextum	JV 1208/11J	USA	KF605668	MG367490	MG367540	[2]
G. adspersum	ITA 39	Italy	EF060011	—	—	[35]
G. adspersum	PF263	Italy	JN176908	—	—	Unpublished
G. alpinum	Cui 17467	Yunnan, SW China	MZ354912	—	—	[2]
G. alpinum	Cui 18402	Xizhang, western China	MZ354910	—	—	[2]
G. angustisporum	Cui 13817	Fujian, SE China	MG279170	MG367507	MG367563	[6]
G. angustisporum	Cui 14578	Guangdong, southern China	MG279171	—	MG367564	[6]
G. applanatum	Cui 14062	Jinlin, NE China	MZ354913	MZ358846	MZ221635	[2]
G. applanatum	Cui 14070	Jinlin, NE China	MZ354914	MZ245387	MZ221636	[2]
G. aridicola	Dai 12588	South Africa	KU572491	—	KU572502	[36]
G. aridicola	GanoTK25	Cameroon	JN105707	—	—	[2]
G. artocarpicola	HL173	Yunnan, SW China	ON994239	OP508428	OP508442	[37]
G. artocarpicola	HL188	Yunnan, SW China	ON994240	OP508427	OP508441	[37]
G. australe	DHCR411	Australia	MF436675	—	MF436677	[38]
G. australe	DHCR417	Australia	MF436676	—	MF436678	[38]
G. austroafricanum	CBS138724	South Africa	KM507324	—	—	[39]
G. bambusicola	Wu 1207-151	Taiwan, SE China	MN957781	LC517944	LC517941	[8]
G. baisuzhenii	N.K. Zeng2080 (FHMU2334)	Hainan, southern China	—	PP785032	PV066218	This study
G. baisuzhenii	N.K. Zeng2519 (FHMU7350)	Hainan, southern China	PP663110	PP785031	PV066219	This study
G. bambusicola	N.K. Zeng1892 (FHMU1217) R71	Hainan, southern China	—	—	PP922172	This study
G. bambusicola	N.K. Zeng1892-1 (FHM7610)	Hainan, southern China	—	—	PP922173	This study
G. bambusicola	N.K. Zeng10386 (FHMU8798)	Hainan, southern China	PV052368	—	PV066216	This study
G. bambusicola	N.K. Zeng10387 (FHMU8803)	Hainan, southern China	PV052369	—	—	This study
G. bambusicola	N.K. Zeng10388 (FHMU8791)	Hainan, southern China	PV052370	PV066220	—	This study
G. bambusicola	N.K. Zeng10340 (FHMU7930)	Hainan, southern China	PV052371	PV066221	PV066217	This study
G. boninense	WD 2028	Japan	KJ143905	KJ143964	KJ143924	[40]
G. boninense	WD 2085	Japan	KJ143906	KJ143965	KJ143925	[40]
G. brownii	JV 1105/9J	Australia	MG279159	MG367494	MG367547	[6]
G. brownii	JV 0709/109	—	KF605662	MG367495	MG367548	[2,6]
G. bubalinomarginatum	Dai 20074	Guangxi, southern China	MZ354926	MZ245388	MZ221637	[2]
G. bubalinomarginatum	Dai 20075	Guangxi, southern China	MZ354927	MZ245389	MZ221638	[2]
G. calidophilum	MFLU 19-2174	Yunnan, SW China	MN398337	—	—	[3]
G. calidophilum	H36	Yunnan, SW China	MW750241	MW839003	MW838997	[1]
G. carnosum	JV 8709/8	Czech Republic	KU572493	—	—	[36]
G. carnosum	MJ 21/08	Czech Republic	KU572492	—	—	[36]
G. carocalcareus	DMC 322	Cameroon	EU089969	—	—	[41]
G. carocalcareus	DMC 513	Cameroon	EU089970	—	—	[41]
G. castaneum	Dai 16500	Hainan, southern China	MZ354918	MZ245390	MZ221639	[2]
G. castaneum	Cui 13893	Hainan, southern China	MZ221640	MZ245391	MZ354919	[2]
G. casuarinicola	Dai 16336	Guangdong, southern China	MG279173	MG367508	MG367565	[6]
G. casuarinicola	Dai 16337	Guangdong, southern China	MG279174	MG367509	MG367566	[6]
G. chalceum	URM80457	Brazil	JX310812	—	—	[42]
G. chocoense	QCAM 3123	Ecuador	MH890527	—	—	[43]
G. chuxiongense	Cui 17262	Yunnan, SW China	MZ354907	—	—	[2]
G. cocoicola	Cui 16791	Australia	MZ354984	MZ245393	MZ221643	[2]
G. cocoicola	Cui 16792	Australia	MZ354985	MZ245394	MZ221644	[2]
G. concinnum	Robledo 3192 (FCOS)	—	MN077522	—	—	[34]
G. concinnum	Robledo 3235 (FCOS)	—	MN077523	—	—	[34]
G. cupreum	GanoTK4	Cameroon	JN105701	—	—	Unpublished
G. cupreum	GanoTK7	Cameroon	JN105702	—	—	Unpublished
G. curtisii	CBS 100131	NC, USA	JQ781848	KJ143966	KJ143926	[11,40]
G. curtisii	CBS 100132	NC, USA	JQ781849	KJ143967	KJ143927	[11,40]
G. destructans	CMW43670	South Africa	KR183856	—	—	[39]
G. dianzhongense	L4331	Yunnan, SW China	MW750237	MZ467043	MW838993	[1]
G. dianzhongense	L4969	Yunnan, SW China	MW750240	MZ467044	MW838996	[1]
G. dorsale	MVHC 5701	Uruguay	MN191581	—	—	[44]
G. dorsale	MVHC 5653	Uruguay	MN191578	—	—	[44]
G. dunense	CMW42149	South Africa	MG020248	—	MG020226	[45]
G. dunense	CMW42157	South Africa	MG020255	—	—	[45]
G. ecuadorense	ASL799	Ecuador	KU128524	—	—	[46]
G. ecuadorense	PMC126	Ecuador	KU128525	—	—	[46]
G. eickeri	CMW 49692	South Africa	MH571690	—	MH567287	[47]
G. eickeri	CMW 50325	South Africa	MH571689	—	MH567290	[47]
G. ellipsoideum	GACP14080966	Hainan, southern China	MH106867	—	—	[5]
G. ellipsoideum	GACP14080968	Hainan, southern China	MH106868	—	—	[5]
G. ellipsoideum	GACP14081228	Hainan, southern China	MH106886	—	—	[5]
G. enigmaticum	CMW43669	South Africa	KR183855	—	—	[39]
G. enigmaticum	CBS 139792	South Africa	NR132918	—	—	[39]
G. esculentum	L4935	Yunnan, SW China	MW750242	MW839004	MW838998	[1]
G. esculentum	HL107	Yunnan, SW China	ON994243	OP508424	OP508437	[48]
G. fallax	JV 1009/27	USA	KF605655	—	—	[2]
G. fallax	JV 0709/39	USA	KF605658	—	—	[2]
G. flexipes	Cui 13841	Hainan, southern China	MZ354923	MZ245401	MZ221655	[2]
G. flexipes	Cui 13863	Hainan, southern China	MZ354924	MZ245402	MZ221656	[2]
G. flexipes	GACP14045450	Hainan, southern China	MH106873	—	—	[5]
G. flexipes	Wei5200	China	JN383978	—	—	[49]
G. flexipes	N.K. Zeng2607 (FHMU3352)	Hainan, southern China	PP663099	—	—	This study
G. flexipes	N.K. Zeng2042 (FHMU2329)	Hainan, southern China	PP663094	PP785027	PP911339	This study
G. flexipes	N.K. Zeng2616 (FHMU5678)	Hainan, southern China	PP663101	—	—	This study
G. flexipes	N.K. Zeng2617 (FHMU5663)	Hainan, southern China	PP663096	—	—	This study
G. flexipes	N.K. Zeng2624 (FHMU5681)	Hainan, southern China	PP663103	—	—	This study
G. flexipes	N.K. Zeng2627 (FHMU5659)	Hainan, southern China	PP663098	—	—	This study
G. flexipes	N.K. Zeng2606 (FHMU3360)	Hainan, southern China	PP663100	—	—	This study
G. flexipes	N.K. Zeng2614 (FHMU5672)	Hainan, southern China	PP663104	—	—	This study
G. flexipes	N.K. Zeng210 (FHMU2292)	Hainan, southern China	PP663095	—	—	This study
G. flexipes	N.K. Zeng2618 (FHMU5661)	Hainan, southern China	PP663097	—	—	This study
G. flexipes	N.K. Zeng4561 (FHMU4863)	Hainan, southern China	PP663106	—	PP922159	This study
G. flexipes	N.K. Zeng4595 (FHMU4898)	Hainan, southern China	PP663102	—	PP922158	This study
G. flexipes	N.K. Zeng2087 (FHMU2337)	Hainan, southern China	PP663107	PP785029	PP922162	This study
G. flexipes	N.K. Zeng2085 (FHMU2336)	Hainan, southern China	PP663108	PP785028	PP922161	This study
“G. flexipes”	Wei5491	Hainan, southern China	JQ781850	KJ143968	—	[11,40]
“G. flexipes”	Wei5494	Hainan, southern China	JN383979	—	—	[11]
G. gibbosum	Cui 13940	China	MZ354972	MZ245404	MZ221658	[2]
G. gibbosum	Cui 14338	China	MZ354969	MZ245405	MZ221659	[2]
G. guangxiense	Cui 14453	Guangxi, southern China	MZ354939	MZ245407	MZ221661	[2]
G. guangxiense	Cui 14454	Guangxi, southern China	MZ354941	MZ245408	MZ221662	[2]
G. guixiense	GXU3457	Guangxi, southern China	OQ788244	PP187389	—	[50]
G. guixiense	GXU3709	Guangxi, southern China	OR271986	—	—	[50]
G. hochiminhense	MFLU 19-2224	Vietnam	MN398324	—	MN423176	[3]
G. hochiminhense	MFLU 19-2225	Vietnam	MN396662	—	MN423177	[3]
G. hoehnelianum	GACP14080913	Hainan, southern China	MH106881	—	—	[5]
G. hoehnelianum	MFLU 19-2168	Myanmar	MN396316	MN423123	MN423158	[3]
G. knysnamense	CMW 47755	South Africa	MH571681	—	MH567261	[47]
G. knysnamense	CMW 47756	South Africa	MH571684	—	MH567274	[47]
G. leucocontextum	GDGM 40200	Xizang, western China	KF011548	—	—	[51]
G. lingzhi	Dai 12479	Anhui, central China	JQ781864	JX029979	JX029975	[11]
G. lingzhi	Wu 1006-38	Hubei, eastern China	JQ781858	JX029980	JX029976	[11]
G. lobatum	JV 0402/24	—	KF605677	—	—	Unpublished
G. lobatum	JV 1212/10J	—	KF605676	—	—	Unpublished
G. lucidum	Rivoire 4195	France	KJ143909	KJ143969	—	[40]
G. lucidum	K 175217	UK	KJ143911	KJ143971	KJ143929	[40]
G. magniporum	Dai 19966	Yunnan, SW China	—	MZ345728	MZ221670	[2]
G. martinicense	SWMart08-55	Martinique	KF963256	—	—	[52]
G. mbrekobenum	UMN7-3 GHA	Ghana	KX000896	—	—	[46]
G. mbrekobenum	UMN7-4 GHA	Ghana	KX000898	—	—	[46]
G. meredithiae	UMNFL50	USA	MG654103	—	—	[53]
G. meredithiae	UMNFL64	USA	MG654106	MG754863	—	[53]
G. mexicanum	MUCL 49453 SW17	Martinique	MK531811	MK531836	MK531825	[54]
G. mexicanum	MUCL 55832	Martinique	MK531815	MK531839	MK531829	[54]
G. mirabile	Cui 18271	Malaysia	MZ354958	MZ345729	MZ221672	[2]
G. mirabile	Cui 18283	Malaysia	MZ354959	MZ345730	MZ221673	[2]
G. mizoramense	UMN-MZ4	India	KY643750	—	—	[55]
G. mizoramense	UMN-MZ5	India	KY643751	—	—	[55]
G. multipileum	CWN 04670 (TNM)	Taiwan, SE China	KJ143913	KJ143972	KJ143931	[40]
G. multipileum	Dai 9447 (IFP)	Hainan, southern China	KJ143914	KJ143973	KJ143932	[40]
G. multipileum	MFLU 19-2166	Thailand	MN401406	MN423142	MN423172	[3]
G. mutabile	Yuan 2289	Yunnan, SW China	JN383977	—	—	[50]
G. mutabile	CLZhao 982	Yunnan, SW China	MG231527	—	—	Unpublished
G. myanmarense	MFLU 19-2167	Myanmar	MN396330	—	—	[3]
G. myanmarense	MFLU 19-2169	Myanmar	MN396330	—	—	[3]
G. nasalanense	GACP17060211	Laos	MK345441	—	—	[3]
G. nasalanense	GACP17060212	Laos	MK345442	—	—	[49]
G. neojaponicum	FFPRI WD-1285	Japan	MN957784	—	—	[8]
G. neojaponicum	FFPRI WD-1532	Japan	MN957785	—	—	[8]
G. nitidum	JV 1504/73	Hainan, southern China	MZ354933	—	MZ221681	[2]
G. obscuratum	Lsh88	Yunnan, SW China	ON994237	—	OP508450	[48]
G. obscuratum	Lsh89	Yunnan, SW China	ON994238	—	OP508451	[48]
G. orbiforme	Cui 13880	Hainan, southern China	MG279187	MG367523	MG367577	[6]
G. orbiforme	Cui 13891	China	MZ354953	MZ345736	MZ221682	[2]
G. oregonense	CBS 265.88	USA	JQ781875	KJ143974	KJ143933	[11,40]
G. oregonense	CBS 266.88	USA	JQ781876	KJ143975	—	[11,40]
G. ovisporum	HKAS123193	Guizhou, SW China	MZ519547	MZ547661	—	[9]
G. ovisporum	GACP20071602	Guizhou, SW China	MZ519548	MZ547662	—	[9]
G. parvulum	URM83343	Brazil	JQ618246	—	—	[45]
G. parvulum	URM80765	Brazil	JX310822	—	—	[45]
G. pfeifferi	K(M)120818	UK	AY884185	—	—	Unpublished
G. philippii	MFLU 19-2222	Thailand	MN401410	—	MN423174	[3]
G. philippii	MFLU 19-2223	Thailand	MN401411	—	MN423175	[3]
G. phyllanthicola	L4948	Yunnan, SW China	PP869245	—	—	[37]
G. phyllanthicola	HL308	Yunnan, SW China	PP869246	—		[37]
G. platense	BAFC384	Argentina	AH008109	—	—	[56]
G. platense	BAFC2374	Argentina	AH008110	—	—	[56]
G. podocarpense	QCAM 6422	Ecuador	MF796661	—	—	[55]
G. podocarpense	JV 1504/126	—	MZ354942	MZ345737	MZ221687	[2]
G. polychromum	UMNOR3	USA	MG654204	—	MG754744	[53]
G. polychromum	MS343OR	USA	MG654197	—	MG754743	[53]
G. puerense	Dai 20427	Yunnan, SW China		MZ345738	MZ221688	[2]
G. ramosissium	xsd08085	—	FJ478127	—	—	[2]
G. ramosissium	xsd08032	—	EU918700	—	—	[2]
G. ravenelii	MS187FL	USA	MG654211	MG754865	MG754745	[53]
G. ravenelii	151FL	USA	MG654208	—	—	[53]
G. resinaceum	BCRC 36147	Netherlands	KJ143916	—	KJ143934	[40]
G. resinaceum	BR 4150	France	KJ143915	—	—	[40]
G. ryvardenii	HKAS58053	Cameroon	HM138671	—	—	[57]
G. ryvardenii	HKAS 58054	Cameroon	HM138672	—	—	[57]
G. sanduense	SA18012501	Guizhou, SW China	MK345450	—	—	[58]
G. sanduense	L4906	Yunnan, SW China	ON994251	OP508430	OP508444	[48]
G. sessile	JV 1209/9	USA	KF605629	—	KJ143936	[40]
G. sessile	V 1209/27	USA	KF605630	KJ143976	KJ143937	[40]
G. shanxiense	BJTC FM423	Shanxi, northern China	MK764268	MK783940	MK783937	[59]
G. shanxiense	HSA 539	Shanxi, northern China	MK764269	MK789681	—	[59]
G. shennongii	N.K. Zeng203 (FHMU2290)	Hainan, southern China	PP663109	—	—	This study
G. sichuanense	HMAS 42798	Sichuan, SW China	JQ781877	—	—	[11]
G. sichuanense	Cui 7691	Guangdong, southern China	JQ781878	—	—	[11]
G. sinense	Cui 14526	Guangxi, southern China	MZ354961	MZ345743	MZ221694	[2]
G. sinense	Cui 14461	China	MZ354963	MZ345744	MZ221695	[2]
G. steyaertanum	II-121-1	Indonesia	KJ654427	—	—	[60]
G. steyaertanum	6-WN-15(M)-A	Indonesia	KJ654459	—	—	[60]
G. suae	L4651	Yunnan, SW China	PP869243	PP894784	PP894782	[37]
G. suae	L4817	Yunnan, SW China	PP869244	—	PP894783	[37]
G. subangustisporum	Cui 18592	Yunnan, SW China	MZ354981	—	MZ221697	[2]
G. subangustisporum	Cui 18593	Yunnan, SW China	MZ354982	—	MZ221698	[2]
G. subellipsoideum	Cui 18241	Malaysia	—	—	MZ221701	[2]
G. subellipsoideum	Cui 18325	Malaysia	—	—	MZ221702	[2]
G. subflexipes	HKAS81926-3	Fujian, SE China	PP465553	—	PP922169	[61], this study
G. subflexipes	N.K. Zeng1893-2 (FHMU7611)	Hainan, southern China	—	—	PP922171	This study
G. subflexipes	HKAS80249	Fujian, SE China	—	—	PP922168	This study
G. subflexipes	N.K. Zeng4086 (FHMU3731)	Guangdong, southern China	—	—	PP922165	This study
G. subflexipes	N.K. Zeng1455 (FHMU2320)	Fujian, SE China	PP465552	PP785030	PP922167	[61], this study
G. subflexipes	N.K. Zeng4114 (FHMU5725)	Guangdong, southern China	PP465551	—	PP922164	[61], this study
G. subflexipes	HKAS79603	Guangdong, southern China	PP465550	—	PP922166	[61], this study
G. subflexipes	HKAS81926-1	Hainan, southern China	PP465549	—	PP922163	[61], this study
G. subflexipes	Cui 17257	Guangdong, southern China	MZ354922	MZ245396	MZ221646	[2]
G. sublobatum	Cui 16804	Australia	MZ354973	MZ345747	MZ221704	[2]
G. sublobatum	Cui 16806	Australia	MZ354974	—	MZ221705	[2]
G. thailandicum	HKAS 104640	Thailand	MK848681	MK875831	MK875829	[62]
G. thailandicum	HKAS 104641	Thailand	MK848682	MK875832	MK875830	[62]
G. tongshanense	Cui 17168	Hubei, central China	MZ354975	—	MZ221706	[2]
G. tropicum	BCRC 37122	Taiwan, SE China	EU021457	—	—	[63]
G. tropicum	GACP1408 1518	Hainan, southern China	MH106884	—	—	[5]
G. tsugae	UMNMI20	USA	MG654324		MG754764	[64]
G. tsugae	UMNMI30	USA	MG654326	MG754871	MH025362	[64]
G. tuberculosum	GVL-21	Mexico	MT232639	—	—	[65]
G. tuberculosum	GVL-40	Mexico	MT232634	—	—	[65]
G. weberianum	GanoTK17	Cameroon	JN105705	—	—	[66]
G. weixiensis	YL02	Yunnan, SW China	MK302445	—	MK302443	[7]
G. weixiensis	YL01	Yunnan, SW China	MK302444	—	MK302442	[7]
G. wiiroense	UMN-21-GHA	Ghana	KT952363	—	—	[67]
G. wiiroense	UMN-20-GHA	Ghana	KT952361	—	—	[67]
G. williamsianum	Dai 17790	Singapore	MZ354947	—	—	[2]
G. williamsianum	Dai 16809	Thailand	MG279183	MG367535	MG367588	[6]
G. yunlingense	Cui 16288	Yunnan, SW China	MZ354915	—	MZ221718	[2]
G. yunlingense	Cui 17043	Yunnan, SW China	MZ354916	—	MZ221719	[2]
G. yunnanense	HL45	Yunnan, SW China	ON994235	OP508422	OP508436	[48]
G. yunnanense	L4812	Yunnan, SW China	ON994236	OP508429	OP508443	[48]
G. zonatum	FL-02	USA	KJ143921	KJ143979	KJ143941	[40]
G. zonatum	FL-03	USA	KJ143922	KJ143980	KJ143942	[40]

New sequences are shown in bold. SW: southwestern; NE: northeastern; SE: southeastern.

Table 1. Taxa, vouchers, locations, and GenBank accession numbers of DNA sequences used in this study.

			GenBank Accession Nos.
Taxon	Voucher	Locality	ITS	rpb2	tef1	Reference
Cristataspora coffeata	FLOR 50933	Brazil	KU315204	—	—	[33]
C. coffeata	Robledo 3183	Brazil	MN077526	—	MN061695	[34]
C. coffeata	Robledo 3182	Brazil	MN077525	—	—	[34]
C. flavipora	Robledo 3288	Argentina	MN077521	—	MN061694	[34]
Foraminispora concentrica	Cui 16238	Yunnan, SW China	MK119816	MK121504	MK121565	[30]
F. concentrica	Cui 16239	Yunnan, SW China	MK119817	MK121506	MK121566	[30]
Ganoderma acaciicola	Cui 16815	Australia	MZ354895	MZ245384	—	[2]
G. acaciicola	Cui 16814	Australia	MZ354894	MZ245383	—	[2]
G. acontextum	JV 0611/21G	Guatemala	KF605667	MG367489	MG367538	[2]
G. acontextum	JV 1208/11J	USA	KF605668	MG367490	MG367540	[2]
G. adspersum	ITA 39	Italy	EF060011	—	—	[35]
G. adspersum	PF263	Italy	JN176908	—	—	Unpublished
G. alpinum	Cui 17467	Yunnan, SW China	MZ354912	—	—	[2]
G. alpinum	Cui 18402	Xizhang, western China	MZ354910	—	—	[2]
G. angustisporum	Cui 13817	Fujian, SE China	MG279170	MG367507	MG367563	[6]
G. angustisporum	Cui 14578	Guangdong, southern China	MG279171	—	MG367564	[6]
G. applanatum	Cui 14062	Jinlin, NE China	MZ354913	MZ358846	MZ221635	[2]
G. applanatum	Cui 14070	Jinlin, NE China	MZ354914	MZ245387	MZ221636	[2]
G. aridicola	Dai 12588	South Africa	KU572491	—	KU572502	[36]
G. aridicola	GanoTK25	Cameroon	JN105707	—	—	[2]
G. artocarpicola	HL173	Yunnan, SW China	ON994239	OP508428	OP508442	[37]
G. artocarpicola	HL188	Yunnan, SW China	ON994240	OP508427	OP508441	[37]
G. australe	DHCR411	Australia	MF436675	—	MF436677	[38]
G. australe	DHCR417	Australia	MF436676	—	MF436678	[38]
G. austroafricanum	CBS138724	South Africa	KM507324	—	—	[39]
G. bambusicola	Wu 1207-151	Taiwan, SE China	MN957781	LC517944	LC517941	[8]
G. baisuzhenii	N.K. Zeng2080 (FHMU2334)	Hainan, southern China	—	PP785032	PV066218	This study
G. baisuzhenii	N.K. Zeng2519 (FHMU7350)	Hainan, southern China	PP663110	PP785031	PV066219	This study
G. bambusicola	N.K. Zeng1892 (FHMU1217) R71	Hainan, southern China	—	—	PP922172	This study
G. bambusicola	N.K. Zeng1892-1 (FHM7610)	Hainan, southern China	—	—	PP922173	This study
G. bambusicola	N.K. Zeng10386 (FHMU8798)	Hainan, southern China	PV052368	—	PV066216	This study
G. bambusicola	N.K. Zeng10387 (FHMU8803)	Hainan, southern China	PV052369	—	—	This study
G. bambusicola	N.K. Zeng10388 (FHMU8791)	Hainan, southern China	PV052370	PV066220	—	This study
G. bambusicola	N.K. Zeng10340 (FHMU7930)	Hainan, southern China	PV052371	PV066221	PV066217	This study
G. boninense	WD 2028	Japan	KJ143905	KJ143964	KJ143924	[40]
G. boninense	WD 2085	Japan	KJ143906	KJ143965	KJ143925	[40]
G. brownii	JV 1105/9J	Australia	MG279159	MG367494	MG367547	[6]
G. brownii	JV 0709/109	—	KF605662	MG367495	MG367548	[2,6]
G. bubalinomarginatum	Dai 20074	Guangxi, southern China	MZ354926	MZ245388	MZ221637	[2]
G. bubalinomarginatum	Dai 20075	Guangxi, southern China	MZ354927	MZ245389	MZ221638	[2]
G. calidophilum	MFLU 19-2174	Yunnan, SW China	MN398337	—	—	[3]
G. calidophilum	H36	Yunnan, SW China	MW750241	MW839003	MW838997	[1]
G. carnosum	JV 8709/8	Czech Republic	KU572493	—	—	[36]
G. carnosum	MJ 21/08	Czech Republic	KU572492	—	—	[36]
G. carocalcareus	DMC 322	Cameroon	EU089969	—	—	[41]
G. carocalcareus	DMC 513	Cameroon	EU089970	—	—	[41]
G. castaneum	Dai 16500	Hainan, southern China	MZ354918	MZ245390	MZ221639	[2]
G. castaneum	Cui 13893	Hainan, southern China	MZ221640	MZ245391	MZ354919	[2]
G. casuarinicola	Dai 16336	Guangdong, southern China	MG279173	MG367508	MG367565	[6]
G. casuarinicola	Dai 16337	Guangdong, southern China	MG279174	MG367509	MG367566	[6]
G. chalceum	URM80457	Brazil	JX310812	—	—	[42]
G. chocoense	QCAM 3123	Ecuador	MH890527	—	—	[43]
G. chuxiongense	Cui 17262	Yunnan, SW China	MZ354907	—	—	[2]
G. cocoicola	Cui 16791	Australia	MZ354984	MZ245393	MZ221643	[2]
G. cocoicola	Cui 16792	Australia	MZ354985	MZ245394	MZ221644	[2]
G. concinnum	Robledo 3192 (FCOS)	—	MN077522	—	—	[34]
G. concinnum	Robledo 3235 (FCOS)	—	MN077523	—	—	[34]
G. cupreum	GanoTK4	Cameroon	JN105701	—	—	Unpublished
G. cupreum	GanoTK7	Cameroon	JN105702	—	—	Unpublished
G. curtisii	CBS 100131	NC, USA	JQ781848	KJ143966	KJ143926	[11,40]
G. curtisii	CBS 100132	NC, USA	JQ781849	KJ143967	KJ143927	[11,40]
G. destructans	CMW43670	South Africa	KR183856	—	—	[39]
G. dianzhongense	L4331	Yunnan, SW China	MW750237	MZ467043	MW838993	[1]
G. dianzhongense	L4969	Yunnan, SW China	MW750240	MZ467044	MW838996	[1]
G. dorsale	MVHC 5701	Uruguay	MN191581	—	—	[44]
G. dorsale	MVHC 5653	Uruguay	MN191578	—	—	[44]
G. dunense	CMW42149	South Africa	MG020248	—	MG020226	[45]
G. dunense	CMW42157	South Africa	MG020255	—	—	[45]
G. ecuadorense	ASL799	Ecuador	KU128524	—	—	[46]
G. ecuadorense	PMC126	Ecuador	KU128525	—	—	[46]
G. eickeri	CMW 49692	South Africa	MH571690	—	MH567287	[47]
G. eickeri	CMW 50325	South Africa	MH571689	—	MH567290	[47]
G. ellipsoideum	GACP14080966	Hainan, southern China	MH106867	—	—	[5]
G. ellipsoideum	GACP14080968	Hainan, southern China	MH106868	—	—	[5]
G. ellipsoideum	GACP14081228	Hainan, southern China	MH106886	—	—	[5]
G. enigmaticum	CMW43669	South Africa	KR183855	—	—	[39]
G. enigmaticum	CBS 139792	South Africa	NR132918	—	—	[39]
G. esculentum	L4935	Yunnan, SW China	MW750242	MW839004	MW838998	[1]
G. esculentum	HL107	Yunnan, SW China	ON994243	OP508424	OP508437	[48]
G. fallax	JV 1009/27	USA	KF605655	—	—	[2]
G. fallax	JV 0709/39	USA	KF605658	—	—	[2]
G. flexipes	Cui 13841	Hainan, southern China	MZ354923	MZ245401	MZ221655	[2]
G. flexipes	Cui 13863	Hainan, southern China	MZ354924	MZ245402	MZ221656	[2]
G. flexipes	GACP14045450	Hainan, southern China	MH106873	—	—	[5]
G. flexipes	Wei5200	China	JN383978	—	—	[49]
G. flexipes	N.K. Zeng2607 (FHMU3352)	Hainan, southern China	PP663099	—	—	This study
G. flexipes	N.K. Zeng2042 (FHMU2329)	Hainan, southern China	PP663094	PP785027	PP911339	This study
G. flexipes	N.K. Zeng2616 (FHMU5678)	Hainan, southern China	PP663101	—	—	This study
G. flexipes	N.K. Zeng2617 (FHMU5663)	Hainan, southern China	PP663096	—	—	This study
G. flexipes	N.K. Zeng2624 (FHMU5681)	Hainan, southern China	PP663103	—	—	This study
G. flexipes	N.K. Zeng2627 (FHMU5659)	Hainan, southern China	PP663098	—	—	This study
G. flexipes	N.K. Zeng2606 (FHMU3360)	Hainan, southern China	PP663100	—	—	This study
G. flexipes	N.K. Zeng2614 (FHMU5672)	Hainan, southern China	PP663104	—	—	This study
G. flexipes	N.K. Zeng210 (FHMU2292)	Hainan, southern China	PP663095	—	—	This study
G. flexipes	N.K. Zeng2618 (FHMU5661)	Hainan, southern China	PP663097	—	—	This study
G. flexipes	N.K. Zeng4561 (FHMU4863)	Hainan, southern China	PP663106	—	PP922159	This study
G. flexipes	N.K. Zeng4595 (FHMU4898)	Hainan, southern China	PP663102	—	PP922158	This study
G. flexipes	N.K. Zeng2087 (FHMU2337)	Hainan, southern China	PP663107	PP785029	PP922162	This study
G. flexipes	N.K. Zeng2085 (FHMU2336)	Hainan, southern China	PP663108	PP785028	PP922161	This study
“G. flexipes”	Wei5491	Hainan, southern China	JQ781850	KJ143968	—	[11,40]
“G. flexipes”	Wei5494	Hainan, southern China	JN383979	—	—	[11]
G. gibbosum	Cui 13940	China	MZ354972	MZ245404	MZ221658	[2]
G. gibbosum	Cui 14338	China	MZ354969	MZ245405	MZ221659	[2]
G. guangxiense	Cui 14453	Guangxi, southern China	MZ354939	MZ245407	MZ221661	[2]
G. guangxiense	Cui 14454	Guangxi, southern China	MZ354941	MZ245408	MZ221662	[2]
G. guixiense	GXU3457	Guangxi, southern China	OQ788244	PP187389	—	[50]
G. guixiense	GXU3709	Guangxi, southern China	OR271986	—	—	[50]
G. hochiminhense	MFLU 19-2224	Vietnam	MN398324	—	MN423176	[3]
G. hochiminhense	MFLU 19-2225	Vietnam	MN396662	—	MN423177	[3]
G. hoehnelianum	GACP14080913	Hainan, southern China	MH106881	—	—	[5]
G. hoehnelianum	MFLU 19-2168	Myanmar	MN396316	MN423123	MN423158	[3]
G. knysnamense	CMW 47755	South Africa	MH571681	—	MH567261	[47]
G. knysnamense	CMW 47756	South Africa	MH571684	—	MH567274	[47]
G. leucocontextum	GDGM 40200	Xizang, western China	KF011548	—	—	[51]
G. lingzhi	Dai 12479	Anhui, central China	JQ781864	JX029979	JX029975	[11]
G. lingzhi	Wu 1006-38	Hubei, eastern China	JQ781858	JX029980	JX029976	[11]
G. lobatum	JV 0402/24	—	KF605677	—	—	Unpublished
G. lobatum	JV 1212/10J	—	KF605676	—	—	Unpublished
G. lucidum	Rivoire 4195	France	KJ143909	KJ143969	—	[40]
G. lucidum	K 175217	UK	KJ143911	KJ143971	KJ143929	[40]
G. magniporum	Dai 19966	Yunnan, SW China	—	MZ345728	MZ221670	[2]
G. martinicense	SWMart08-55	Martinique	KF963256	—	—	[52]
G. mbrekobenum	UMN7-3 GHA	Ghana	KX000896	—	—	[46]
G. mbrekobenum	UMN7-4 GHA	Ghana	KX000898	—	—	[46]
G. meredithiae	UMNFL50	USA	MG654103	—	—	[53]
G. meredithiae	UMNFL64	USA	MG654106	MG754863	—	[53]
G. mexicanum	MUCL 49453 SW17	Martinique	MK531811	MK531836	MK531825	[54]
G. mexicanum	MUCL 55832	Martinique	MK531815	MK531839	MK531829	[54]
G. mirabile	Cui 18271	Malaysia	MZ354958	MZ345729	MZ221672	[2]
G. mirabile	Cui 18283	Malaysia	MZ354959	MZ345730	MZ221673	[2]
G. mizoramense	UMN-MZ4	India	KY643750	—	—	[55]
G. mizoramense	UMN-MZ5	India	KY643751	—	—	[55]
G. multipileum	CWN 04670 (TNM)	Taiwan, SE China	KJ143913	KJ143972	KJ143931	[40]
G. multipileum	Dai 9447 (IFP)	Hainan, southern China	KJ143914	KJ143973	KJ143932	[40]
G. multipileum	MFLU 19-2166	Thailand	MN401406	MN423142	MN423172	[3]
G. mutabile	Yuan 2289	Yunnan, SW China	JN383977	—	—	[50]
G. mutabile	CLZhao 982	Yunnan, SW China	MG231527	—	—	Unpublished
G. myanmarense	MFLU 19-2167	Myanmar	MN396330	—	—	[3]
G. myanmarense	MFLU 19-2169	Myanmar	MN396330	—	—	[3]
G. nasalanense	GACP17060211	Laos	MK345441	—	—	[3]
G. nasalanense	GACP17060212	Laos	MK345442	—	—	[49]
G. neojaponicum	FFPRI WD-1285	Japan	MN957784	—	—	[8]
G. neojaponicum	FFPRI WD-1532	Japan	MN957785	—	—	[8]
G. nitidum	JV 1504/73	Hainan, southern China	MZ354933	—	MZ221681	[2]
G. obscuratum	Lsh88	Yunnan, SW China	ON994237	—	OP508450	[48]
G. obscuratum	Lsh89	Yunnan, SW China	ON994238	—	OP508451	[48]
G. orbiforme	Cui 13880	Hainan, southern China	MG279187	MG367523	MG367577	[6]
G. orbiforme	Cui 13891	China	MZ354953	MZ345736	MZ221682	[2]
G. oregonense	CBS 265.88	USA	JQ781875	KJ143974	KJ143933	[11,40]
G. oregonense	CBS 266.88	USA	JQ781876	KJ143975	—	[11,40]
G. ovisporum	HKAS123193	Guizhou, SW China	MZ519547	MZ547661	—	[9]
G. ovisporum	GACP20071602	Guizhou, SW China	MZ519548	MZ547662	—	[9]
G. parvulum	URM83343	Brazil	JQ618246	—	—	[45]
G. parvulum	URM80765	Brazil	JX310822	—	—	[45]
G. pfeifferi	K(M)120818	UK	AY884185	—	—	Unpublished
G. philippii	MFLU 19-2222	Thailand	MN401410	—	MN423174	[3]
G. philippii	MFLU 19-2223	Thailand	MN401411	—	MN423175	[3]
G. phyllanthicola	L4948	Yunnan, SW China	PP869245	—	—	[37]
G. phyllanthicola	HL308	Yunnan, SW China	PP869246	—		[37]
G. platense	BAFC384	Argentina	AH008109	—	—	[56]
G. platense	BAFC2374	Argentina	AH008110	—	—	[56]
G. podocarpense	QCAM 6422	Ecuador	MF796661	—	—	[55]
G. podocarpense	JV 1504/126	—	MZ354942	MZ345737	MZ221687	[2]
G. polychromum	UMNOR3	USA	MG654204	—	MG754744	[53]
G. polychromum	MS343OR	USA	MG654197	—	MG754743	[53]
G. puerense	Dai 20427	Yunnan, SW China		MZ345738	MZ221688	[2]
G. ramosissium	xsd08085	—	FJ478127	—	—	[2]
G. ramosissium	xsd08032	—	EU918700	—	—	[2]
G. ravenelii	MS187FL	USA	MG654211	MG754865	MG754745	[53]
G. ravenelii	151FL	USA	MG654208	—	—	[53]
G. resinaceum	BCRC 36147	Netherlands	KJ143916	—	KJ143934	[40]
G. resinaceum	BR 4150	France	KJ143915	—	—	[40]
G. ryvardenii	HKAS58053	Cameroon	HM138671	—	—	[57]
G. ryvardenii	HKAS 58054	Cameroon	HM138672	—	—	[57]
G. sanduense	SA18012501	Guizhou, SW China	MK345450	—	—	[58]
G. sanduense	L4906	Yunnan, SW China	ON994251	OP508430	OP508444	[48]
G. sessile	JV 1209/9	USA	KF605629	—	KJ143936	[40]
G. sessile	V 1209/27	USA	KF605630	KJ143976	KJ143937	[40]
G. shanxiense	BJTC FM423	Shanxi, northern China	MK764268	MK783940	MK783937	[59]
G. shanxiense	HSA 539	Shanxi, northern China	MK764269	MK789681	—	[59]
G. shennongii	N.K. Zeng203 (FHMU2290)	Hainan, southern China	PP663109	—	—	This study
G. sichuanense	HMAS 42798	Sichuan, SW China	JQ781877	—	—	[11]
G. sichuanense	Cui 7691	Guangdong, southern China	JQ781878	—	—	[11]
G. sinense	Cui 14526	Guangxi, southern China	MZ354961	MZ345743	MZ221694	[2]
G. sinense	Cui 14461	China	MZ354963	MZ345744	MZ221695	[2]
G. steyaertanum	II-121-1	Indonesia	KJ654427	—	—	[60]
G. steyaertanum	6-WN-15(M)-A	Indonesia	KJ654459	—	—	[60]
G. suae	L4651	Yunnan, SW China	PP869243	PP894784	PP894782	[37]
G. suae	L4817	Yunnan, SW China	PP869244	—	PP894783	[37]
G. subangustisporum	Cui 18592	Yunnan, SW China	MZ354981	—	MZ221697	[2]
G. subangustisporum	Cui 18593	Yunnan, SW China	MZ354982	—	MZ221698	[2]
G. subellipsoideum	Cui 18241	Malaysia	—	—	MZ221701	[2]
G. subellipsoideum	Cui 18325	Malaysia	—	—	MZ221702	[2]
G. subflexipes	HKAS81926-3	Fujian, SE China	PP465553	—	PP922169	[61], this study
G. subflexipes	N.K. Zeng1893-2 (FHMU7611)	Hainan, southern China	—	—	PP922171	This study
G. subflexipes	HKAS80249	Fujian, SE China	—	—	PP922168	This study
G. subflexipes	N.K. Zeng4086 (FHMU3731)	Guangdong, southern China	—	—	PP922165	This study
G. subflexipes	N.K. Zeng1455 (FHMU2320)	Fujian, SE China	PP465552	PP785030	PP922167	[61], this study
G. subflexipes	N.K. Zeng4114 (FHMU5725)	Guangdong, southern China	PP465551	—	PP922164	[61], this study
G. subflexipes	HKAS79603	Guangdong, southern China	PP465550	—	PP922166	[61], this study
G. subflexipes	HKAS81926-1	Hainan, southern China	PP465549	—	PP922163	[61], this study
G. subflexipes	Cui 17257	Guangdong, southern China	MZ354922	MZ245396	MZ221646	[2]
G. sublobatum	Cui 16804	Australia	MZ354973	MZ345747	MZ221704	[2]
G. sublobatum	Cui 16806	Australia	MZ354974	—	MZ221705	[2]
G. thailandicum	HKAS 104640	Thailand	MK848681	MK875831	MK875829	[62]
G. thailandicum	HKAS 104641	Thailand	MK848682	MK875832	MK875830	[62]
G. tongshanense	Cui 17168	Hubei, central China	MZ354975	—	MZ221706	[2]
G. tropicum	BCRC 37122	Taiwan, SE China	EU021457	—	—	[63]
G. tropicum	GACP1408 1518	Hainan, southern China	MH106884	—	—	[5]
G. tsugae	UMNMI20	USA	MG654324		MG754764	[64]
G. tsugae	UMNMI30	USA	MG654326	MG754871	MH025362	[64]
G. tuberculosum	GVL-21	Mexico	MT232639	—	—	[65]
G. tuberculosum	GVL-40	Mexico	MT232634	—	—	[65]
G. weberianum	GanoTK17	Cameroon	JN105705	—	—	[66]
G. weixiensis	YL02	Yunnan, SW China	MK302445	—	MK302443	[7]
G. weixiensis	YL01	Yunnan, SW China	MK302444	—	MK302442	[7]
G. wiiroense	UMN-21-GHA	Ghana	KT952363	—	—	[67]
G. wiiroense	UMN-20-GHA	Ghana	KT952361	—	—	[67]
G. williamsianum	Dai 17790	Singapore	MZ354947	—	—	[2]
G. williamsianum	Dai 16809	Thailand	MG279183	MG367535	MG367588	[6]
G. yunlingense	Cui 16288	Yunnan, SW China	MZ354915	—	MZ221718	[2]
G. yunlingense	Cui 17043	Yunnan, SW China	MZ354916	—	MZ221719	[2]
G. yunnanense	HL45	Yunnan, SW China	ON994235	OP508422	OP508436	[48]
G. yunnanense	L4812	Yunnan, SW China	ON994236	OP508429	OP508443	[48]
G. zonatum	FL-02	USA	KJ143921	KJ143979	KJ143941	[40]
G. zonatum	FL-03	USA	KJ143922	KJ143980	KJ143942	[40]

New sequences are shown in bold. SW: southwestern; NE: northeastern; SE: southeastern.

2.5. Phylogenetic Analyses

The phylogenetic tree based on the combined dataset (ITS + rpb2 + tef1) was reconstructed using both maximum likelihood (ML) and Bayesian inference (BI) methods. For the ML analysis, tree generation and bootstrap analyses were conducted using RAxML 7.2.6 [68], with 1000 bootstrap replicates integrated into an ML search. Bayesian analysis was performed using MrBayes 3.1 [69], employing the Markov Chain Monte Carlo (MCMC) technique. The substitution models for the combined dataset were determined using MrModeltest 2.3 [70], with the best-fit models identified as HKY + I + G for ITS and GTR + I + G for both rpb2 and tef1. The Bayesian analysis of the combined nuclear dataset (ITS + rpb2 + tef1) was run for 60 million generations, with trees sampled every 1000 generations. The first 25% of the sampled trees were discarded as burn-in, and a majority consensus tree was constructed from the remaining trees. Bayesian posterior probabilities (PPs) were calculated for the consensus tree. Branches with ML bootstrap values ≥ 70% and Bayesian posterior probabilities (PPs) ≥ 0.95 were considered to have significant support.

3. Results

3.1. Molecular Data

The combined dataset (ITS + rpb2 + tef1) comprised 224 sequences with 2032 nucleotide sites. A phylogram generated using RAxML, displaying branch lengths and support values, is presented in Figure 1. The tree topologies inferred from maximum likelihood (ML) and Bayesian inference (BY) analyses were identical, although slight differences in statistical support were observed (Figure 1). Based on the combined dataset, our newly collected Ganoderma specimens with diminutive pilei and gracile stipes were grouped into five distinct lineages (Figure 1). Lineage 1, with strong statistical support (BS = 97%, PP = 0.99), included four collections of G. flexipes Pat. and 14 newly collected specimens (FHMU2292, FHMU2329, FHMU2336, FHMU2337, FHMU3352, FHMU3360, FHMU4863, FHMU4898, FHMU5659, FHMU5661, FHMU5663, FHMU5672, FHMU5678, and FHMU5681). Lineage 2, also with strong statistical support (BS = 98%, PP = 1.0), consisted of two collections labeled as G. flexipes (Wei5491 and Wei5494) and one newly collected specimen (FHMU2290). Lineage 3, with strong statistical support (BS = 98%, PP = 1.0), included the holotype of G. subflexipes B.K. Cui, J.H. Xing & Y.F. Sun and eight newly collected specimens (FHMU2320, FHMU3731, FHMU5725, FHMU7611, HKAS79603, HKAS80249, HKAS81926-1, and HKAS81926-3). Lineage 4, with robust statistical support (BS = 100%, PP = 1.0), grouped the holotype of G. bambusicola Sheng H. Wu, C.L. Chern & T. Hatt. with six new specimens (FHMU1217, FHMU7610, FHMU7930, FHMU8791, FHMU8798, and FHMU8803). Finally, Lineage 5, also with strong statistical support (BS = 100%, PP = 1.0), comprised two newly collected specimens (FHMU2334 and FHMU7350).

3.2. Taxonomy

Based on morphological examinations and molecular phylogenetic analyses, our new Ganoderma collections from the southern region of China, particularly Hainan Island, were identified as five distinct taxa. Among these, three were recognized as G. bambusicola, G. flexipes, and G. subflexipes, while the remaining two represented novel species. In accordance with local tradition, G. baisuzhenii, G. flexipes, and G. shennongii are commonly known as “Lingzhi Wang”, whereas G. bambusicola and G. subflexipes are referred to as “Zhu Lingzhi”. Detailed morphological descriptions of these five species are provided in the following sections.

Ganoderma baisuzhenii N.K. Zeng, R. Tian & Zhi Q. Liang, sp. nov. (Figure 2 and Figure 3)

MycoBank: MB854442

Etymology: “baisuzhenii” is given in honor of Su-Zhen Bai, an ancient Chinese mythical figure who risked her life in search of Lingzhi to save her husband.

Diagnosis: It differs from the closest species of Ganoderma by a brownish-red to dark brownish-red pileus, a nearly white context, large pores, a pore surface that was initially yellowish, fading to white with age, and relatively large basidiospores, and it grows on decaying hardwood (underground).

Holotype: CHINA. Hainan Province: Changjiang County, Bawangling of Hainan Tropical Rainforest National Park, elev. 850 m, 28 June 2015, N.K. Zeng2080 (FHMU2334).

Description: Basidiomata annual, dorso-laterally stipitate, corky. Pilei up to 2.6 cm diameter and 0.9 cm thick, solitary, sub-reniform to reniform, or flabelliform. Pileal surface brownish-red (8E8) to dark brownish-red (8F8), strongly laccate, glabrous, with concentric furrows and inconspicuously radial rugose; margin obtuse, entire, incurved. Pore surface yellowish when young, then white, turning yellowish-brown (4C5) when injured; pore 2–3 per mm, subcircular, circular or angular; dissepiments slightly thick to moderately thick, entire. Context up to 0.5 cm thick, nearly white (4A3), hard woody. Tube up to 0.7 cm long, brown (5C5), indistinctively stratified. Stipe up to 10 cm long and 0.65 cm diameter, subcylindrical, solid, woody; surface brownish-black (8F7), glabrous to bumpy, laccate.

Hyphal system trimitic; generative hyphae with clamp connections; all hyphae IKI –, CB +; tissues darkening in KOH. Generative hyphae in context 1–3.5 µm diameter, colorless, thin-walled; skeletal hyphae in context 3–6.5 µm diameter, yellowish-brown, thick-walled with a wide to narrow lumen or sub-solid, seldom branched; binding hyphae in context 2.5–4 µm diameter, pale yellow, thin to slight thick-walled, sub-solid to solid, branched and flexuous. Generative hyphae in tubes 2.5–4 µm diameter, colorless, thin-walled; skeletal hyphae in tubes 3–6.5 µm diameter, yellowish-brown, usually thick-walled, solid to sub-solid, rarely branched; binding hyphae in tube 2–4.7 µm diameter, pale yellow, thin to slight thick-walled, sub-solid to solid, branched and flexuous. Pileipellis composed of clamped generative hyphae, thick-walled to sub-solid; apical cells 20–45 × 4–12 μm, clavate, slightly inflated, yellowish-brown, forming a regular palisade. Stipitipellis composed of clamped generative hyphae, thick-walled to sub-solid; apical cells 25–40 × 6–10 μm, clavate, slightly inflated, grayish-yellow, forming a regular palisade. Cystidia and cystidioles absent. Basidioles 15–23 × 10–13.5 μm, clavate, colorless, thin-walled. Basidia not observed. Basidiospores (11–) 11.5–14 (–15) × (7–) 8–10 μm, L = 13.13 μm, W = 9.35 μm, Q = 1.40 (n = 20/1/1, with myxosporium), 9–11 (–11.5) × 6–8 (–8.5) μm, L = 9.9 μm, W = 7.25 μm, Q = 1.37 (n = 20/1/1, without myxosporium), ellipsoid to broadly ellipsoid, pale yellowish-brown to brown, IKI –, CB +, double-walled with distinctly thick walls, exospore wall smooth, endospore wall with dense spinules.

Habitat: Solitary or gregarious, growing on decaying hardwood (often underground) of fagaceous trees, particularly those of the genus Cyclobalanopsis.

Known distribution: Southern China (Hainan Province).

Additional specimen examined: CHINA. Hainan Province: Yinggeling of Hainan Tropical Rainforest National Park, elev. 750 m, 3 August 2015, N.K. Zeng2519 (FHMU7350).

Notes: Ganoderma baisuzhenii, commonly referred to as “Lingzhi Wang” on Hainan Island in tropical China, has been frequently misidentified as G. flexipes (based on our investigations). Ganoderma flexipes can be distinguished from G. baisuzhenii by its richer red pileus, white pore surface, brown pileal context, and smaller basidiospores (see details below). Morphologically and phylogenetically, G. baisuzhenii is closely related to G. magniporum J.D. Zhao & X.Q. Zhang, G. sanduense Hapuar., T.C. Wen & K.D. Hyde, and G. yunnanense Jun He & Shu H. Li. However, these species exhibit distinct characteristics: G. magniporum, originally described in Guangxi in southern China, features a blackish-brown to black pileus, a brown context, and smaller basidiospores measuring 8.7–10.4 × 5.2–7 μm [71]; G. sanduense, first described in Guizhou in southwestern China, is characterized by a reddish-black to brownish-black pileus, smaller pores, and a brown to dark brown context [58]; and G. yunnanense, originally described in Yunnan in southwestern China, has smaller pores (4–6 per mm), a white pore surface, and smaller basidiospores measuring 9–12 × 7–8 μm [48].

Ganoderma bambusicola Sheng H. Wu, C.L. Chern & T. Hatt. Phytotaxa 456(1): 79, 2020 (Figure 4 and Figure 5)

MycoBank: MB835651

Description: Basidiomata annual, dorso-laterally stipitate, woody. Pilei up to 7 cm diameter and 2 mm thick, solitary, applanate, sub-reniform to reniform, or flabelliform. Pileal surface brownish-black (8F7–8F8), reddish-black (10F8) to purplish-brown (12E6–12E7), laccate, glabrous, with conspicuously or obscurely concentric furrows and strongly radial rugose; margin obtuse, entire, slightly incurved. Pore surface white (2B2–2B3), turning yellowish-brown (5D5–5D6) when injured; pores 4–6 per mm, subcircular to circular or angular; dissepiments thin, mostly entire. Context up to 0.3 cm thick, dark brown (5E5–5E8), fibrous to corky. Tubes up to 0.4 cm long, grayish-yellow (5C3–5C4), indistinctively stratified. Stipe up to 15 cm long and 1.5 cm diameter, cylindrical to subcylindrical, solid, woody; surface blackish (7F7), glabrous to bumpy, laccate.

Hyphal system trimitic; generative hyphae with clamp connections; all hyphae IKI –, CB +; tissues darkening in KOH. Generative hyphae in context 2–4 µm diameter, nearly colorless, thin-walled; skeletal hyphae in context 4–7 µm diameter, grayish-yellow, thick-walled with a medium to narrow lumen or sub-solid, rarely branched; binding hyphae in context 3–5 µm diameter, light orange, slightly thick-walled, sub-solid, frequently branched. Generative hyphae in tubes 3–6 μm diameter, colorless, thin-walled; skeletal hyphae in tubes grayish-yellow, thick-walled with a narrow lumen to sub-solid, frequently arboriform and flexuous; binding hyphae in tubes 1.5–3.5 μm diameter, light orange, thick-walled, branched and flexuous. Pileipellis composed of clamped generative hyphae, thick-walled to sub-solid; apical cells 30–64 × 6–14 μm, clavate, slightly inflated, yellowish-brown, forming a regular palisade. Stipitipellis composed of clamped generative hyphae, thick-walled to sub-solid; apical cells 35–50 × 7–15 μm, clavate, slightly inflated, yellowish-brown, forming a regular palisade. Cystidia and cystidioles absent. Basidioles 17–24.5 × 9–13 μm, clavate, colorless, thin-walled. Basidia not observed. Basidiospores 9.5–12.5 × 5–7 μm, L = 11.13 μm, W = 6.32 μm, Q = 1.76 (n = 160/8/7, with myxosporium), 7–11 × 4–5.5 μm, L = 9.03 μm, W = 5.08 μm, Q = 1.78 (n = 160/8/7, without myxosporium), ellipsoid to broadly ellipsoid, pale yellowish-brown to brown, IKI–, CB+, double-walled with distinctly thick walls, exospore wall smooth, endospore wall with dense spinules.

Habitat: Solitary or gregarious, occurring on dead roots of bamboo, particularly those of Dendrocalamus latiflorus Munro.

Known distribution: Southern (Hainan Province) and southeastern (Taiwan Province) China, Laos, and Myanmar [8].

Specimens examined: CHINA. Hainan Province, Haikou City, bought from market, 14 January 2015, N.K. Zeng1892 (FHMU1217); same location and date, N.K. Zeng1892-1 (FHMU7610); Baisha County, Qingsong Town, Edible and Medicinal Fungi Cultivation Base, 2 October 2024, N.K. Zeng10340 (FHMU7930); Baisha County, Nankai Town, elev. 520 m, 7 October 2024, N.K. Zeng10386 (FHMU8798); same location and date, N.K. Zeng10387 (FHMU8803); same location and date, N.K. Zeng10388 (FHMU8791).

Notes: Ganoderma bambusicola was originally described in Taiwan in southeastern China [8]. Taxonomically, this species represents a newly recorded addition to the fungal flora of the Chinese mainland. Commonly referred to as “Zhu Lingzhi” on Hainan Island in tropical China, it has been successfully cultivated (based on our investigations). This species is characterized by a brownish-black, reddish-black to purplish-brown pileus, relatively small pores, a white pore surface, and a dark brown context. It is typically found growing on dead bamboo roots.

Ganoderma flexipes Pat., Bull. Soc. mycol. Fr. 23(2): 75, 1907 (Figure 6 and Figure 7)

≡ Fomes flexipes (Pat.) Sacc. & Traverso, Syll. fung. (Abellini) 19: 710, 1910

≡ Polyporus flexipes (Pat.) Lloyd, Mycol. Writ. 3 (Syn. Stip. Polyporoids): 104, 1912

= Ganoderma atrum J.D. Zhao et al., Acta Microbiol. Sin. 19: 268, 1979

= Ganoderma hainanense J.D. Zhao et al., Acta Microbiol. Sin. 19: 269, 1979

= Ganoderma parviungulatum J.D. Zhao & X.Q. Zhang, Acta Mycol. Sin. 5: 88, 1986

MycoBank: MB249905

Description: Basidiomata annual, dorso-laterally, sometimes centrally stipitate, corky. Pilei solitary, up to 5 cm diameter and 0.6 cm thick, sub-reniform to reniform, subflabellate to flabellate, shell-like or circular. Pileal surface brownish-red (8E5–8E6) to reddish, strongly laccate, glabrous when young, then bumpy, with obvious concentric furrows and slightly radial rugose; margin obtuse, entire, slightly incurved. Pore surface white, turning brownish (5D5–5D6) when injured; pore 3–4 per mm, subcircular, circular or angular; dissepiments slightly thick, entire. Context up to 3 mm thick, upper layer yellowish-brown (5D5–5D6), lower layer brown to dark brown (8E8), fibrous to corky. Tubes pale brown (5B3–5B4), indistinctively stratified, up to 0.6 cm long. Stipe up to 17.5 cm long and 0.6 cm diameter, flattened to subcylindrical, solid, woody; surface brownish-red (8E8) to reddish-black (8F8), glabrous to bumpy, laccate.

Hyphal system trimitic; generative hyphae with clamp connections; all hyphae IKI–, CB+; tissues darkening in KOH. Generative hyphae in context 1–2.5 µm diameter, colorless, thin-walled; skeletal hyphae in context 4–6 µm diameter, yellowish-brown, thick-walled with a wide to narrow lumen or sub-solid, arboriform and flexuous; binding hyphae in context 4–6 µm diameter, colorless, thick-walled, branched and flexuous. Generative hyphae in tubes 1–3 μm diameter, colorless, thin-walled; skeletal hyphae in tubes 2.5–5 μm diameter, colorless, sub-solid, arboriform and flexuous; binding hyphae in tubes 1–3 μm diameter, colorless, thick-walled, branched and flexuous. Pileipellis composed of clamped generative hyphae, thick-walled; apical cells 30–50 × 5–13 µm, clavate, slightly inflated, yellowish-brown or brownish-orange, forming a regular palisade. Stipitipellis composed of clamped generative hyphae, thick-walled to sub-solid; apical cells 32–50 × 5–12 μm, clavate, slightly inflated, pale brown or brownish-orange, forming a regular palisade. Cystidia and cystidioles absent. Basidioles 17.5–23 × 7–11.5 μm, clavate, colorless, thin-walled. Basidia not observed. Basidiospores 8.5–12 × 5.5–7.5 μm, L = 10.15 μm, W = 6.45 μm, Q = 1.57 (n = 80/4/3, with myxosporium); 6.5–9 × 4–6 μm, L = 7.59 μm, W = 5.2 μm, Q = 1.46 (n = 80/4/3, without myxosporium), ellipsoid, not obviously truncated, light yellow (4A5–4A6) to pale yellowish-brown (5C5–5C6), IKI –, CB +, double-walled with moderately thick walls, exospore wall smooth, endospore wall with dense spinules.

Habitat: Solitary or gregarious, occurring on decaying hardwood (often underground) of fagaceous trees, particularly those of Quercus patelliformis Chun.

Known distribution: Southern (Hainan and Guangdong Provinces) [2], southeastern (Taiwan Province) [72], and southwestern (Yunnan Province) China [3], Vietnam, India, Laos, Nepal, Pakistan, and Myanmar [3,58].

Specimens examined: CHINA. Hainan Province: Wuzhishan of Hainan Tropical Rainforest National Park, elev. 1200 m, 30 May 2009, N.K. Zeng210 (FHMU2292); Jianfengling of Hainan Tropical Rainforest National Park, elev. 900 m, 30 June 2015, N.K. Zeng2087 (FHMU2337); same location, 27 June 2018, N.K. Zeng3420 (FHMU2985); same location, 10 August 2020, N.K. Zeng4561 (FHMU4863); same location, 11 August 2020, N.K. Zeng4595 (FHMU4898); Bawangling of Hainan Tropical Rainforest National Park, elev. 950 m, 28 June 2015, N.K. Zeng2042 (FHMU2329); same location and date, N.K. Zeng2085 (FHMU2336); Yinggeling of Hainan Tropical Rainforest National Park, elev. 800 m, 5 August 2015, N.K. Zeng2606 (FHMU3360); same location and date, N.K. Zeng2607 (FHMU3352); Ledong County, Jianfeng Town, 16 November 2015, bought from the market, N.K. Zeng2614 (FHMU5672); same location and date, N.K. Zeng2616 (FHMU5678); same location and date, N.K. Zeng2617 (FHMU5663); same location and date, N.K. Zeng2618 (FHMU5661); same location and date, N.K. Zeng2624 (FHMU5681); same location and date, N.K. Zeng2627 (FHMU5659).

Notes: Ganoderma flexipes was originally described in Vietnam [73] and has since been reported in several other regions, including China, India, Laos, Nepal, Pakistan, and Myanmar [3,58,72,74,75]. In Hainan Island, tropical China, this species is commonly referred to as “Lingzhi Wang”. It is characterized by a brownish-red to reddish pileus, a white pore surface, and a yellowish-brown to dark brown context. It typically grows on the decaying hardwood (often underground) of fagaceous trees.

Ganoderma shennongii N.K. Zeng, R. Tian & Zhi Q. Liang, sp. nov. (Figure 8 and Figure 9)

MycoBank: MB854441

Etymology: “shennongii” is given in honor of our ancestor represented by Shennong who dared to taste hundreds of herbs including Ganoderma spp.

Diagnosis: It differs from the closest species of Ganoderma by a relatively small basidioma, a dark reddish pileus, a white pore surface, relatively small pores, and a yellowish-brown to brown context, and it grows on decaying dead wood (underground).

Holotype: CHINA. Hainan Province, Diaoluoshan of Hainan Tropical Rainforest National Park, elev. 950 m, 28 May 2009, N.K. Zeng203 (FHMU2290).

Description: Basidiomata annual, dorso-laterally stipitate, corky. Pilei up to 1.8 cm diameter and 0.6 cm thick, solitary, sub-reniform to reniform, or subflabellate to flabellate. Pileal surface dark reddish (9E5–9E6), strongly laccate, glabrous, with conspicuously concentric furrows and slightly radial rugose; margin obtuse, entire, incurved. Pore surface white, turning brown (5C5–5C6) when injured; pores 4–5 per mm, subcircular to circular or angular; dissepiments slightly thick to moderately thick, entire. Context up to 0.1 cm thick, yellowish-brown (4C5) to brown (5D7–5D8), fibrous to corky. Tube up to 0.1 cm long, brown (5C5), indistinctively stratified. Stipe up to 12.5 cm long and 4 mm diameter, cylindrical to subcylindrical, solid, fibrous to woody; surface brownish-black (8F7), glabrous to bumpy, laccate.

Hyphal system trimitic; generative hyphae with clamp connections; all hyphae IKI–, CB+; tissues darkening in KOH. Generative hyphae in context 1.2–2 µm diameter, colorless, thin-walled; skeletal hyphae in context 3.8–5 µm diameter, pale yellowish-brown, thick-walled with a wide to narrow lumen or sub-solid, arboriform and flexuous; binding hyphae in context 1.2–3.2 µm diameter, colorless, thick-walled, branched and flexuous. Generative hyphae in tubes 1–2 µm diameter, colorless, thin-walled; skeletal hyphae in tubes 4–6.5 µm diameter, colorless, thick-walled with a wide to narrow lumen or sub-solid, arboriform and flexuous; binding hyphae in tubes 1–3 µm diameter, colorless, thick-walled, branched and flexuous. Pileipellis composed of clamped generative hyphae, thick-walled; apical cells 25–45 × 7–14 μm, clavate, inflated, dark brown, anticlinal, forming a regular palisade. Stipitipellis composed of clamped generative hyphae, thick-walled; apical cells 20–40 × 5–10 μm, clavate, inflated, dark brown, anticlinal, forming a regular palisade. Cystidia and cystidioles absent. Basidioles 20–25 × 9–20 μm, subclavate to clavate, colorless, thin-walled. Basidia not observed. Basidiospores 10–11.5 × 7–8.5 μm, L = 10.44 μm, W = 7.55 μm, Q = 1.38 (n = 20/1/1, with myxosporium), 7.5–9.5 × 5–8 μm, L = 8.17 μm, W = 6.72 μm, Q = 1.12 (n = 20/1/1, without myxosporium), ellipsoid to broadly ellipsoid, pale yellowish-brown, IKI –, CB +, double-walled with slightly thick walls, exospore wall smooth, endospore wall with dense spinules.

Habitat: Solitary or gregarious, occurring on decaying hardwood (often underground) in forests predominantly composed of fagaceous trees.

Known distribution: Southern China (Hainan Province).

Notes: Ganoderma shennongii, commonly referred to as “Lingzhi Wang” on Hainan Island in tropical China, was historically misidentified as G. flexipes [40]. However, G. flexipes can be distinguished by its larger pores and narrower basidiospores, measuring 8.5–12 × 5.5–7.5 μm (see above). In addition to G. flexipes, G. shennongii is morphologically and phylogenetically closely related to G. subflexipes. Nevertheless, G. subflexipes exhibits smaller pores and narrower basidiospores, measuring 8–11.5 × 5–7.5 μm (see below). Furthermore, G. shennongii shares morphological similarities with G. bambusicola; however, the latter is characterized by a more blackish pileus, narrower basidiospores (9.5–12.5 × 5–7 μm), and a specific habitat preference for growing on dead bamboo roots (see above).

Ganoderma subflexipes B.K. Cui, J.H. Xing & Y.F. Sun, Stud. Mycol. 101: 350, 2022 (Figure 10 and Figure 11)

MycoBank: MB839675

Description: Basidiomata annual, eccentric or dorso-laterally stipitate, corky. Pilei up to 5 cm diameter and 0.5 cm thick, solitary, applanate, sub-reniform to reniform, or subflabellate to flabellate. Pileal surface reddish-brown (8D7–8D8) to orangish-brown (6B8), strongly laccate, glabrous when young, then bumpy, with conspicuously concentric furrows and slightly radial rugose; margin obtuse, entire, incurved. Pore surface white, turning yellowish-brown (4C5–4C6) when injured; pore 5–6 per mm, subcircular to circular or angular. Context up to 1.5 cm thick, yellowish-brown (4C5) to brown (5D7–5D8), fibrous to corky. Tubes up to 0.5 cm long, pale yellow (1A3–1A4) to grayish-yellow (4B4), indistinctively stratified. Stipe up to 15 cm long and 7 mm diameter, cylindrical to subcylindrical, solid, fibrous to woody; surface reddish-brown (7E7) to chocolate brown (8E7), glabrous to bumpy, laccate.

Hyphal system trimitic; generative hyphae with clamp connections; all hyphae IKI–, CB+; tissues darkening in KOH. Generative hyphae in context 1–2 µm diameter, colorless, thin-walled; skeletal hyphae in context 2–6 µm in diameter, pale yellowish-brown, thick-walled with a wide to narrow lumen or sub-solid, arboriform and flexuous; binding hyphae in context 1–3.5 µm diameter, colorless, thick-walled, branched and flexuous. Generative hyphae in tubes 1.5–3 µm diameter, colorless, thin-walled; skeletal hyphae in tubes 3–5 µm in diameter, pale yellow, thick-walled with a wide to narrow lumen or sub-solid, arboriform and flexuous; binding hyphae in context 1–3.5 µm diameter, colorless, thick-walled, branched and flexuous. Pileipellis composed of clamped generative hyphae, thick-walled; apical cells 20–50 × 5–12 μm, clavate, inflated, dark brown, anticlinal, forming a regular palisade. Stipitipellis composed of clamped generative hyphae, thick-walled; apical cells 20–40 × 6–12 μm, clavate, inflated, dark brown, forming a regular palisade. Cystidia and cystidioles absent. Basidioles 15–19 × 8.5–13 μm, subclavate to clavate, colorless, thin-walled. Basidia not observed. Basidiospores 8–11.5 (–12) × 5–7.5 (–8) μm, L = 10.27 μm, W = 6.76 μm, Q = 1.52 (n = 120/6/3, with myxosporium), 6–9.5 × 4–7 μm, L = 7.78 μm, W = 5.34 μm, Q = 1.46 (n = 120/6/3, without myxosporium), ellipsoid to broadly ellipsoid, pale yellow, IKI–, CB+, double-walled with slightly thick walls, exospore wall smooth, endospore wall with dense spinules.

Habitat: It is solitary or gregarious and primarily found on dead roots of bamboo, particularly those of Bambusa chungii McClure. Occasionally, it may also occur on decaying hardwood (often underground) in forests dominated by fagaceous trees or in mixed forests where fagaceous trees and Pinus massoniana Lamb. are predominant.

Known distribution: Southern (Hainan and Guangdong Provinces), southeastern (Fujian Province), and eastern China (Jiangxi Province) [2].

Specimens examined: CHINA. Fujian Province: Zhangping City, Xingqiao Town, Chengkou Village, elev. 350 m, 20 August 2013, N.K. Zeng1455 (FHMU2320); Sanming City, Geshikao National Forest Park, elev. 380 m, 8 July 2013, Y.J. Hao969 (HKAS80249); Sanming City, Nature Reserve for Wild Sarcandra glabra, 8 July 2013, T. Guo724 (HKAS81926). Guangdong Province: Fengkai County, Heishiding Nature Reserve, elev. 360 m, 2 June 2013, Q. Cai924 (HKAS79603); Shaoguan City, Danxia National Nature Reserve, elev. 380 m, 5 June 2019, N.K. Zeng4086 (FHMU3731); same collection, 4 June 2019, N.K. Zeng4114 (FHMU5725). Hainan Province: Limushan of Hainan Tropical Rainforest National Park, elev. 650 m, 4 June 2009, N.K. Zeng242 (FHMU2299); Haikou City, 14 January 2015, bought from market, N.K. Zeng1893 (FHMU1218); same location and date, N.K. Zeng 1893-2 (FHMU7611).

Notes: Ganoderma subflexipes, originally described in Guangdong in southern China [2], is commonly referred to as “Zhu Lingzhi” on Hainan Island in tropical China. This species is characterized by a reddish-brown to orangish-brown pileus, relatively small pores, a white pore surface, and a yellowish-brown to brown context. It typically grows on dead bamboo roots, though it is occasionally found on decaying hardwood (often underground).

4. Discussion

The species diversity of Ganoderma has been extensively documented in southern China, particularly on Hainan Island [2]. However, wild populations of Ganoderma species, including those referred to as “Lingzhi Wang” or “Zhu Lingzhi” on Hainan Island, have significantly declined due to overharvesting. Consequently, many species within this genus have been classified as priority protected species in the Hainan Tropical Rainforest National Park [76,77]. Effective conservation of these species necessitates accurate identification. Furthermore, there is an urgent need to reliably distinguish “Lingzhi Wang” or “Zhu Lingzhi”, which are crucial medicinal fungi in Hainan Island, from their close relatives, counterfeits, adulterants, and inferior substitutes to ensure their medicinal efficacy [78]. The present study identifies “Lingzhi Wang” as including G. baisuzhenii, G. flexipes, and G. shennongii, and “Zhu Lingzhi” as including G. bambusicola and G. subflexipes. The precise definition of these species provides essential data for the conservation and medicinal utilization of these Ganoderma species (

Table 2. Main characters of five Ganoderma species called “Lingzhi Wang” or “Zhu Lingzhi” in Hainan Island, tropical China.

Species	Local Name	Pilei Color	Pore	Context	Basidiospore (μm)	Host
Ganoderma baisuzhenii	Lingzhi Wang	Brownish-red to dark brownish-red	2–3 per mm; surface yellowish when young, then white	Nearly white	11.5–14 × 8–10	On decaying hardwood (often underground) of fagaceous trees, particularly those of the genus Cyclobalanopsis
G. bambusicola	Zhu Lingzhi	Brownish-black, reddish-black to purplish-brown	4–6 per mm; surface white	Dark brown	9.5–12.5 × 5–7	On dead roots of bamboo, particularly those of Dendrocalamus latiflorus
G. flexipes	Lingzhi Wang	Brownish-red to reddish	3–4 per mm; surface white	Upper layer yellowish-brown, lower layer brown to dark brown	8.5–12 × 5.5–7.5	On decaying hardwood (often underground) of fagaceous trees, particularly those of Quercus patelliformis
G. shennongii	Lingzhi Wang	Dark reddish	4–5 per mm; surface white	Yellowish-brown to brown	10–11.5 × 7–8.5	On decaying hardwood (often underground) in forests predominantly composed of fagaceous trees
G. subflexipes	Zhu Lingzhi	Reddish-brown to orangish-brown	5–6 per mm; surface white	Yellowish-brown to brown	8–11.5 × 5–7.5	On dead roots of bamboo, particularly those of Bambusa chungii, sometimes on decaying hardwood (often underground) in forests dominated by fagaceous trees, or mixed forests dominated by fagaceous trees and Pinus massoniana

).

In addition to the aforementioned species—namely, G. baisuzhenii, G. bambusicola, G. flexipes, G. shennongii, and G. subflexipes—other taxa such as G. atrum J.D. Zhao, L.W. Hsu & X.Q. Zhang, G. calidophilum J.D. Zhao, L.W. Hsu & X.Q. Zhang, G. hainanense J.D. Zhao, L.W. Hsu & X.Q. Zhang, G. luteomarginatum J.D. Zhao, L.W. Hsu & X.Q. Zhang, and G. parviungulatum J.D. Zhao & X.Q. Zhang, all originally described on Hainan Island in tropical China, are also characterized by their diminutive pilei and gracile stipes [79,80]. Cao et al. [11] proposed that G. atrum, G. hainanense, and G. parviungulatum are synonymous with G. flexipes, a suggestion later supported by Sun et al. [2]. Similarly, G. luteomarginatum was suggested to be synonymous with G. sinense J.D. Zhao, L.W. Hsu & X.Q. Zhang [2]. Although G. calidophilum was initially believed to be synonymous with G. flexipes [11,72], this viewpoint has been challenged in recent studies. The protologue of G. calidophilum does not align well with that of G. flexipes, as the former exhibits denser pores (4–6 per mm) and larger basidiospores, measuring 10–12.1 × 6.2–8.7 μm [79]. Consequently, the taxonomic relationship between G. flexipes and G. calidophilum remains unresolved and warrants further investigation in future studies.

Among Ganoderma species, high phenotypic plasticity at the macroscopic level is a well-documented phenomenon [81]. In a previous study [61], we successfully induced the formation of fruit bodies of G. subflexipes in a greenhouse environment. Notably, the cultivated G. subflexipes exhibited considerable morphological variation, with fruit bodies displaying diverse shapes (Figure 12). Additionally, the pilei of cultivated G. subflexipes were significantly larger, measuring up to 10 cm in length and 8.5 cm in width. These findings align with the observation that the morphological features of G. subflexipes fruit bodies are influenced by growing conditions, consistent with the views of Gilbertson and Ryvarden [82], who emphasized the high variability of fruit bodies in Ganoderma species. It is also noteworthy that the attachment type of the stipe to the pileus in G. flexipes exhibits considerable variation, ranging from lateral (Figure 6a–d) to nearly central (Figure 6e). This morphological variability underscores the challenges of relying solely on macroscopic characteristics for species identification. Therefore, in addition to morphological evaluation, DNA sequence data play a crucial role in the accurate identification of Ganoderma species. Integrating molecular data with traditional morphological approaches is essential for resolving taxonomic ambiguities and ensuring reliable species delineation.

DNA sequence data play a pivotal and indispensable role in resolving taxonomic delimitations within the genus Ganoderma, as evidenced by previous studies [1,2,3,40,51]. To conclusively address the taxonomic ambiguities surrounding “Lingzhi Wang” or “Zhu Lingzhi” specimens, comprehensive multilocus DNA phylogenetic analyses should be prioritized. Our current investigation has identified five candidate species corresponding to these vernacular names, yet this likely represents only a fraction of the taxonomic diversity. Notably, Hainan Island, a tropical biodiversity hotspot in China, harbors remarkable Ganoderma diversity [2,5]. We anticipate that future investigations incorporating advanced molecular techniques will reveal additional taxa characterized by the diagnostic morphological features of diminutive pilei and gracile stipes.

5. Conclusions

The present study reveals that the Ganoderma species characterized by small pilei and gracile stipes on Hainan Island in tropical China comprise at least five distinct species. Among these, two species—G. baisuzhenii and G. shennongii—are newly described. The remaining three species, G. bambusicola, G. flexipes, and G. subflexipes, have been previously documented. Notably, G. bambusicola is reported for the first time on the Chinese mainland. Furthermore, this study clarifies the taxonomic identities of “Lingzhi Wang” and “Zhu Lingzhi”, identifying “Lingzhi Wang” as encompassing G. baisuzhenii, G. flexipes, and G. shennongii, and “Zhu Lingzhi” as including G. bambusicola and G. subflexipes. These findings provide a foundation for the accurate identification, conservation, and sustainable utilization of these medicinally significant fungi.

Key to five Ganoderma species called “Lingzhi Wang” or “Zhu Lingzhi” in Hainan Island, tropical China
1. Host is bamboo	2
1. Host is hardwood (especially Fagaceae trees)	3
2. Pileus color is brownish-black, reddish-black to purplish-brown; basidiospore size 9.5–12.5 × 5–7 μm	G. bambusicola
2. Pileus color is reddish-brown to orangish-brown; basidiospore size 8–11.5 × 5–7.5 μm	G. subflexipes
3. Pileus context is nearly white; pore density up to 3 per mm; basidiospore up to 14 μm in length and 10 μm in width	G. baisuzhenii
3. Pileus context is brown, pore density up to 5 per mm; basidiospore up to 12 μm in length and 8.5 μm in width	4
4. Pileus color is brownish-red to reddish; pore density 3–4 per mm; basidiospore size 8.5–12 × 5.5–7.5 μm	G. flexipes
4. Pileus color is dark reddish; pore density 4–5 per mm; basidiospore size 10–11.5 × 7–8.5 μm	G. shennongii