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Journal of Fungi
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20 March 2025

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

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1
Ministry of Education Key Laboratory for Ecology of Tropical Islands, Key Laboratory of Tropical Animal and Plant Ecology of Hainan Province, College of Life Sciences, Hainan Normal University, Haikou 571158, China
2
Institute of Medicinal Plant Development, Chinese Academy of Medical Sciences & Peking Union Medical College, Beijing 100193, China
3
Qinghai Provincial Key Laboratory of Tibetan Medicine Pharmacology and Safety Evaluation, Northwest Institute of Plateau Biology, Chinese Academy of Sciences, Xining 810008, China
4
University of Chinese Academy of Sciences, 19(A) Yuquan Road, Beijing 100049, China
J. Fungi2025, 11(3), 237;https://doi.org/10.3390/jof11030237
This article belongs to the Special Issue Edible and Medicinal Macrofungi, 3rd Edition
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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.
Table 1. Taxa, vouchers, locations, and GenBank accession numbers of DNA sequences used in this study.
GenBank Accession Nos.
TaxonVoucherLocalityITSrpb2tef1Reference
Cristataspora coffeataFLOR 50933BrazilKU315204——[33]
C. coffeataRobledo 3183BrazilMN077526—MN061695[34]
C. coffeataRobledo 3182BrazilMN077525——[34]
C. flaviporaRobledo 3288ArgentinaMN077521—MN061694[34]
Foraminispora concentricaCui 16238Yunnan, SW ChinaMK119816MK121504MK121565[30]
F. concentricaCui 16239Yunnan, SW ChinaMK119817MK121506MK121566[30]
Ganoderma acaciicolaCui 16815 AustraliaMZ354895MZ245384—[2]
G. acaciicolaCui 16814AustraliaMZ354894MZ245383—[2]
G. acontextumJV 0611/21G GuatemalaKF605667MG367489MG367538[2]
G. acontextumJV 1208/11JUSAKF605668MG367490MG367540[2]
G. adspersumITA 39ItalyEF060011——[35]
G. adspersumPF263ItalyJN176908——Unpublished
G. alpinumCui 17467 Yunnan, SW ChinaMZ354912——[2]
G. alpinumCui 18402Xizhang, western ChinaMZ354910——[2]
G. angustisporumCui 13817Fujian, SE ChinaMG279170MG367507MG367563[6]
G. angustisporumCui 14578Guangdong, southern ChinaMG279171—MG367564[6]
G. applanatumCui 14062Jinlin, NE ChinaMZ354913MZ358846MZ221635[2]
G. applanatumCui 14070Jinlin, NE ChinaMZ354914MZ245387MZ221636[2]
G. aridicolaDai 12588South AfricaKU572491—KU572502[36]
G. aridicolaGanoTK25CameroonJN105707——[2]
G. artocarpicolaHL173Yunnan, SW ChinaON994239OP508428OP508442[37]
G. artocarpicolaHL188Yunnan, SW ChinaON994240OP508427OP508441[37]
G. australeDHCR411AustraliaMF436675—MF436677[38]
G. australeDHCR417AustraliaMF436676—MF436678[38]
G. austroafricanumCBS138724South AfricaKM507324——[39]
G. bambusicolaWu 1207-151Taiwan, SE ChinaMN957781LC517944LC517941[8]
G. baisuzheniiN.K. Zeng2080
(FHMU2334)		Hainan, southern China—PP785032PV066218This study
G. baisuzheniiN.K. Zeng2519
(FHMU7350)		Hainan, southern ChinaPP663110PP785031PV066219This study
G. bambusicolaN.K. Zeng1892
(FHMU1217) R71		Hainan, southern China——PP922172
This study
G. bambusicolaN.K. Zeng1892-1
(FHM7610)		Hainan, southern China——PP922173
This study
G. bambusicolaN.K. Zeng10386 (FHMU8798)Hainan, southern ChinaPV052368—PV066216This study
G. bambusicolaN.K. Zeng10387 (FHMU8803)Hainan, southern ChinaPV052369——This study
G. bambusicolaN.K. Zeng10388 (FHMU8791)Hainan, southern ChinaPV052370PV066220—This study
G. bambusicolaN.K. Zeng10340 (FHMU7930)Hainan, southern ChinaPV052371PV066221PV066217This study
G. boninenseWD 2028JapanKJ143905KJ143964KJ143924[40]
G. boninenseWD 2085JapanKJ143906KJ143965KJ143925[40]
G. browniiJV 1105/9JAustraliaMG279159MG367494MG367547[6]
G. browniiJV 0709/109—KF605662MG367495MG367548[2,6]
G. bubalinomarginatumDai 20074Guangxi, southern ChinaMZ354926MZ245388MZ221637[2]
G. bubalinomarginatumDai 20075 Guangxi, southern ChinaMZ354927MZ245389MZ221638[2]
G. calidophilumMFLU 19-2174Yunnan, SW ChinaMN398337——[3]
G. calidophilumH36Yunnan, SW ChinaMW750241MW839003MW838997[1]
G. carnosumJV 8709/8Czech Republic KU572493——[36]
G. carnosumMJ 21/08Czech Republic KU572492——[36]
G. carocalcareusDMC 322 CameroonEU089969——[41]
G. carocalcareusDMC 513CameroonEU089970——[41]
G. castaneumDai 16500Hainan, southern ChinaMZ354918MZ245390MZ221639[2]
G. castaneumCui 13893Hainan, southern ChinaMZ221640MZ245391MZ354919[2]
G. casuarinicolaDai 16336 Guangdong, southern ChinaMG279173MG367508MG367565[6]
G. casuarinicolaDai 16337Guangdong, southern ChinaMG279174MG367509MG367566[6]
G. chalceumURM80457BrazilJX310812——[42]
G. chocoenseQCAM 3123 EcuadorMH890527——[43]
G. chuxiongenseCui 17262 Yunnan, SW ChinaMZ354907——[2]
G. cocoicolaCui 16791 AustraliaMZ354984MZ245393MZ221643[2]
G. cocoicolaCui 16792AustraliaMZ354985MZ245394MZ221644[2]
G. concinnumRobledo 3192 (FCOS)—MN077522——[34]
G. concinnumRobledo 3235 (FCOS)—MN077523——[34]
G. cupreumGanoTK4CameroonJN105701——Unpublished
G. cupreumGanoTK7CameroonJN105702——Unpublished
G. curtisiiCBS 100131NC, USAJQ781848KJ143966KJ143926[11,40]
G. curtisiiCBS 100132NC, USAJQ781849KJ143967KJ143927[11,40]
G. destructansCMW43670 South AfricaKR183856——[39]
G. dianzhongenseL4331Yunnan, SW ChinaMW750237MZ467043MW838993[1]
G. dianzhongenseL4969Yunnan, SW ChinaMW750240MZ467044MW838996[1]
G. dorsaleMVHC 5701UruguayMN191581——[44]
G. dorsaleMVHC 5653UruguayMN191578——[44]
G. dunenseCMW42149South AfricaMG020248—MG020226[45]
G. dunenseCMW42157 South AfricaMG020255——[45]
G. ecuadorenseASL799 EcuadorKU128524——[46]
G. ecuadorensePMC126EcuadorKU128525——[46]
G. eickeriCMW 49692South AfricaMH571690—MH567287[47]
G. eickeriCMW 50325South AfricaMH571689—MH567290[47]
G. ellipsoideumGACP14080966Hainan, southern ChinaMH106867——[5]
G. ellipsoideumGACP14080968Hainan, southern ChinaMH106868——[5]
G. ellipsoideumGACP14081228Hainan, southern ChinaMH106886——[5]
G. enigmaticumCMW43669 South AfricaKR183855——[39]
G. enigmaticumCBS 139792 South AfricaNR132918——[39]
G. esculentumL4935Yunnan, SW ChinaMW750242MW839004MW838998[1]
G. esculentumHL107Yunnan, SW ChinaON994243OP508424OP508437[48]
G. fallaxJV 1009/27 USAKF605655——[2]
G. fallaxJV 0709/39USAKF605658——[2]
G. flexipesCui 13841Hainan, southern ChinaMZ354923MZ245401MZ221655[2]
G. flexipesCui 13863Hainan, southern ChinaMZ354924MZ245402MZ221656[2]
G. flexipesGACP14045450Hainan, southern ChinaMH106873——[5]
G. flexipesWei5200 ChinaJN383978——[49]
G. flexipesN.K. Zeng2607
(FHMU3352)		Hainan, southern ChinaPP663099——This study
G. flexipesN.K. Zeng2042 (FHMU2329)Hainan, southern ChinaPP663094PP785027PP911339This study
G. flexipesN.K. Zeng2616 (FHMU5678)Hainan, southern ChinaPP663101——This study
G. flexipesN.K. Zeng2617 (FHMU5663)Hainan, southern ChinaPP663096——This study
G. flexipesN.K. Zeng2624 (FHMU5681)Hainan, southern ChinaPP663103——This study
G. flexipesN.K. Zeng2627 (FHMU5659)Hainan, southern ChinaPP663098——This study
G. flexipesN.K. Zeng2606 (FHMU3360)Hainan, southern ChinaPP663100——This study
G. flexipesN.K. Zeng2614 (FHMU5672)Hainan, southern ChinaPP663104——This study
G. flexipesN.K. Zeng210 (FHMU2292)Hainan, southern ChinaPP663095——This study
G. flexipesN.K. Zeng2618 (FHMU5661)Hainan, southern ChinaPP663097——This study
G. flexipesN.K. Zeng4561 (FHMU4863)Hainan, southern ChinaPP663106—PP922159This study
G. flexipesN.K. Zeng4595 (FHMU4898)Hainan, southern ChinaPP663102—PP922158This study
G. flexipesN.K. Zeng2087 (FHMU2337)Hainan, southern ChinaPP663107PP785029PP922162This study
G. flexipesN.K. Zeng2085 (FHMU2336)Hainan, southern ChinaPP663108PP785028PP922161This study
“G. flexipes”Wei5491Hainan, southern ChinaJQ781850KJ143968—[11,40]
“G. flexipes”Wei5494Hainan, southern ChinaJN383979——[11]
G. gibbosumCui 13940ChinaMZ354972MZ245404MZ221658[2]
G. gibbosumCui 14338ChinaMZ354969MZ245405MZ221659[2]
G. guangxienseCui 14453 Guangxi, southern ChinaMZ354939MZ245407MZ221661[2]
G. guangxienseCui 14454Guangxi, southern ChinaMZ354941MZ245408MZ221662[2]
G. guixienseGXU3457Guangxi, southern ChinaOQ788244PP187389—[50]
G. guixienseGXU3709Guangxi, southern ChinaOR271986——[50]
G. hochiminhenseMFLU 19-2224 VietnamMN398324—MN423176[3]
G. hochiminhenseMFLU 19-2225VietnamMN396662—MN423177[3]
G. hoehnelianumGACP14080913Hainan, southern ChinaMH106881——[5]
G. hoehnelianumMFLU 19-2168MyanmarMN396316MN423123MN423158[3]
G. knysnamenseCMW 47755 South AfricaMH571681—MH567261[47]
G. knysnamenseCMW 47756South AfricaMH571684—MH567274[47]
G. leucocontextumGDGM 40200Xizang, western ChinaKF011548——[51]
G. lingzhiDai 12479Anhui, central ChinaJQ781864JX029979JX029975[11]
G. lingzhiWu 1006-38Hubei, eastern ChinaJQ781858JX029980JX029976[11]
G. lobatumJV 0402/24—KF605677——Unpublished
G. lobatumJV 1212/10J—KF605676——Unpublished
G. lucidumRivoire 4195FranceKJ143909KJ143969—[40]
G. lucidumK 175217UKKJ143911KJ143971KJ143929[40]
G. magniporumDai 19966Yunnan, SW China—MZ345728MZ221670[2]
G. martinicenseSWMart08-55 MartiniqueKF963256——[52]
G. mbrekobenumUMN7-3 GHA GhanaKX000896——[46]
G. mbrekobenumUMN7-4 GHAGhanaKX000898——[46]
G. meredithiaeUMNFL50USAMG654103——[53]
G. meredithiaeUMNFL64USAMG654106MG754863—[53]
G. mexicanumMUCL 49453 SW17MartiniqueMK531811MK531836 MK531825[54]
G. mexicanumMUCL 55832MartiniqueMK531815MK531839 MK531829[54]
G. mirabileCui 18271MalaysiaMZ354958MZ345729MZ221672[2]
G. mirabileCui 18283MalaysiaMZ354959MZ345730MZ221673[2]
G. mizoramenseUMN-MZ4 IndiaKY643750——[55]
G. mizoramenseUMN-MZ5IndiaKY643751——[55]
G. multipileumCWN 04670 (TNM)Taiwan, SE ChinaKJ143913KJ143972KJ143931[40]
G. multipileumDai 9447 (IFP)Hainan, southern ChinaKJ143914KJ143973KJ143932[40]
G. multipileumMFLU 19-2166ThailandMN401406MN423142MN423172[3]
G. mutabileYuan 2289Yunnan, SW ChinaJN383977——[50]
G. mutabileCLZhao 982Yunnan, SW ChinaMG231527——Unpublished
G. myanmarenseMFLU 19-2167MyanmarMN396330——[3]
G. myanmarenseMFLU 19-2169MyanmarMN396330——[3]
G. nasalanenseGACP17060211LaosMK345441——[3]
G. nasalanenseGACP17060212LaosMK345442——[49]
G. neojaponicumFFPRI WD-1285JapanMN957784——[8]
G. neojaponicumFFPRI WD-1532JapanMN957785——[8]
G. nitidumJV 1504/73Hainan, southern ChinaMZ354933—MZ221681[2]
G. obscuratumLsh88Yunnan, SW ChinaON994237—OP508450[48]
G. obscuratumLsh89Yunnan, SW ChinaON994238—OP508451[48]
G. orbiformeCui 13880Hainan, southern ChinaMG279187MG367523MG367577[6]
G. orbiformeCui 13891ChinaMZ354953MZ345736MZ221682[2]
G. oregonenseCBS 265.88USAJQ781875KJ143974KJ143933[11,40]
G. oregonenseCBS 266.88USAJQ781876KJ143975—[11,40]
G. ovisporumHKAS123193Guizhou, SW ChinaMZ519547MZ547661—[9]
G. ovisporumGACP20071602Guizhou, SW ChinaMZ519548MZ547662—[9]
G. parvulumURM83343BrazilJQ618246——[45]
G. parvulumURM80765BrazilJX310822——[45]
G. pfeifferiK(M)120818UKAY884185——Unpublished
G. philippiiMFLU 19-2222ThailandMN401410—MN423174[3]
G. philippiiMFLU 19-2223ThailandMN401411—MN423175[3]
G. phyllanthicolaL4948Yunnan, SW ChinaPP869245——[37]
G. phyllanthicolaHL308Yunnan, SW ChinaPP869246— [37]
G. platenseBAFC384ArgentinaAH008109——[56]
G. platenseBAFC2374ArgentinaAH008110——[56]
G. podocarpenseQCAM 6422 EcuadorMF796661——[55]
G. podocarpenseJV 1504/126—MZ354942MZ345737MZ221687[2]
G. polychromumUMNOR3USAMG654204—MG754744[53]
G. polychromumMS343ORUSAMG654197—MG754743[53]
G. puerenseDai 20427 Yunnan, SW China MZ345738MZ221688[2]
G. ramosissiumxsd08085—FJ478127——[2]
G. ramosissiumxsd08032—EU918700——[2]
G. raveneliiMS187FLUSAMG654211MG754865MG754745[53]
G. ravenelii151FLUSAMG654208——[53]
G. resinaceumBCRC 36147NetherlandsKJ143916—KJ143934[40]
G. resinaceumBR 4150FranceKJ143915——[40]
G. ryvardeniiHKAS58053CameroonHM138671——[57]
G. ryvardeniiHKAS 58054CameroonHM138672——[57]
G. sanduenseSA18012501Guizhou, SW ChinaMK345450——[58]
G. sanduenseL4906Yunnan, SW ChinaON994251OP508430OP508444[48]
G. sessileJV 1209/9USAKF605629—KJ143936[40]
G. sessileV 1209/27USAKF605630KJ143976KJ143937[40]
G. shanxienseBJTC FM423Shanxi, northern ChinaMK764268MK783940MK783937[59]
G. shanxienseHSA 539Shanxi, northern ChinaMK764269MK789681—[59]
G. shennongiiN.K. Zeng203 (FHMU2290)Hainan, southern ChinaPP663109——This study
G. sichuanenseHMAS 42798 Sichuan, SW ChinaJQ781877——[11]
G. sichuanenseCui 7691Guangdong, southern ChinaJQ781878——[11]
G. sinenseCui 14526Guangxi, southern ChinaMZ354961MZ345743MZ221694[2]
G. sinenseCui 14461ChinaMZ354963MZ345744MZ221695[2]
G. steyaertanumII-121-1IndonesiaKJ654427——[60]
G. steyaertanum6-WN-15(M)-AIndonesiaKJ654459——[60]
G. suaeL4651Yunnan, SW ChinaPP869243PP894784PP894782[37]
G. suaeL4817Yunnan, SW ChinaPP869244—PP894783[37]
G. subangustisporumCui 18592 Yunnan, SW ChinaMZ354981—MZ221697[2]
G. subangustisporumCui 18593Yunnan, SW ChinaMZ354982—MZ221698[2]
G. subellipsoideumCui 18241Malaysia——MZ221701[2]
G. subellipsoideumCui 18325 Malaysia——MZ221702[2]
G. subflexipesHKAS81926-3Fujian, SE ChinaPP465553—PP922169[61], this study
G. subflexipesN.K. Zeng1893-2 (FHMU7611)Hainan, southern China——PP922171This study
G. subflexipesHKAS80249Fujian, SE China——PP922168This study
G. subflexipesN.K. Zeng4086
(FHMU3731)		Guangdong, southern China——PP922165This study
G. subflexipesN.K. Zeng1455
(FHMU2320)		Fujian, SE ChinaPP465552PP785030PP922167[61], this study
G. subflexipesN.K. Zeng4114
(FHMU5725)		Guangdong, southern ChinaPP465551—PP922164[61], this study
G. subflexipesHKAS79603Guangdong, southern ChinaPP465550—PP922166[61], this study
G. subflexipesHKAS81926-1Hainan, southern ChinaPP465549—PP922163[61], this study
G. subflexipesCui 17257 Guangdong, southern ChinaMZ354922MZ245396MZ221646[2]
G. sublobatumCui 16804AustraliaMZ354973MZ345747MZ221704[2]
G. sublobatumCui 16806AustraliaMZ354974—MZ221705[2]
G. thailandicumHKAS 104640 ThailandMK848681MK875831MK875829[62]
G. thailandicumHKAS 104641ThailandMK848682MK875832MK875830[62]
G. tongshanenseCui 17168 Hubei, central ChinaMZ354975—MZ221706[2]
G. tropicumBCRC 37122Taiwan, SE ChinaEU021457——[63]
G. tropicumGACP1408 1518Hainan, southern ChinaMH106884——[5]
G. tsugaeUMNMI20USAMG654324 MG754764[64]
G. tsugaeUMNMI30USAMG654326MG754871MH025362[64]
G. tuberculosumGVL-21MexicoMT232639——[65]
G. tuberculosumGVL-40MexicoMT232634——[65]
G. weberianumGanoTK17CameroonJN105705——[66]
G. weixiensisYL02Yunnan, SW ChinaMK302445—MK302443[7]
G. weixiensisYL01 Yunnan, SW ChinaMK302444—MK302442[7]
G. wiiroenseUMN-21-GHAGhanaKT952363——[67]
G. wiiroenseUMN-20-GHAGhanaKT952361——[67]
G. williamsianumDai 17790SingaporeMZ354947——[2]
G. williamsianumDai 16809ThailandMG279183MG367535MG367588[6]
G. yunlingenseCui 16288 Yunnan, SW ChinaMZ354915—MZ221718[2]
G. yunlingenseCui 17043Yunnan, SW ChinaMZ354916—MZ221719[2]
G. yunnanenseHL45Yunnan, SW ChinaON994235OP508422OP508436[48]
G. yunnanenseL4812Yunnan, SW ChinaON994236OP508429OP508443[48]
G. zonatumFL-02USAKJ143921KJ143979KJ143941[40]
G. zonatumFL-03USAKJ143922KJ143980KJ143942[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).
Figure 1. Phylogram for Ganoderma species generated from maximum likelihood analysis of ITS, rpb2, and tef1 sequence dataset using RAxML. BS ≥ 70% and PP ≥ 0.95 are indicated above or below the branches as RAxML BS/PP.

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)
Figure 2. Ganoderma baisuzhenii (a–c) Basidiomata [(a,b) from FHMU2334, holotype; (c) from FHMU7350]. (d) Section of pileus (FHMU2334). Scale bars = 1 cm. Photos by N.K. Zeng.
Figure 3. Microscopic features of Ganoderma baisuzhenii (FHMU 2334, holotype). (a) Basidiospores. (b) Pileipellis. (c) Stipitipellis. (d) Generative hyphae. (e) Skeletal hyphae. (f) Binding hyphae. (g) Basidioles. Scale bars = 10 μm. Drawings by R. Tian.
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)
Figure 4. Ganoderma bambusicola (a–d) Basidiomata [(a,b) from FHMU8798; (c) from FHMU7930, cultivated fruit bodies; (d) from FHMU1217]. (e) Section of pileus (FHMU1217). Scale bars = 1 cm. Photos: (a,b) by Y. Liu and (c–e) by N.K. Zeng.
Figure 5. Microscopic features of Ganoderma bambusicola (FHMU1217). (a) Basidiospores. (b) Pileipellis. (c) Stipitipellis. (d) Generative hyphae. (e) Skeletal hyphae. (f) Binding hyphae. (g) Basidioles. Scale bars = 10 μm. Drawings by R. Tian.
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)
Figure 6. Ganoderma flexipes (a–e) Basidiomata [(a,b) from FHMU2985; (c,d) from FHMU2329; (e) from FHMU2337]. (f) Section of pileus (FHMU2329). Scale bars = 1 cm. Photos by N.K. Zeng.
Figure 7. Microscopic features of Ganoderma flexipes (FHMU2985). (a) Basidiospores. (b) Pileipellis. (c) Stipitipellis. (d) Generative hyphae. (e) Skeletal hyphae. (f) Binding hyphae. (g) Basidioles. Scale bars = 10 μm. Drawings by R. Tian.
≡ 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)
Figure 8. Ganoderma shennongii (FHMU2290, holotype) (a–c) Basidiomata. (d) Section of pileus. Scale bars = 1 cm. Photos by N.K. Zeng.
Figure 9. Microscopic features of Ganoderma shennongii (FHMU2290, holotype). (a) Basidiospores. (b) Pileipellis. (c) Stipitipellis. (d) Generative hyphae. (e) Skeletal hyphae. (f) Binding hyphae. (g) Basidioles. Scale bars = 10 μm. Drawings by R. Tian.
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)
Figure 10. Ganoderma subflexipes (a–e) Basidiomata [(a,b) from FHMU2320; (c,d) from FHMU5725; (e) from FHMU2299]. (f) Section of pileus (FHMU5725). Scale bars = 1 cm. Photos by N.K. Zeng.
Figure 11. Microscopic features of Ganoderma subflexipes (FHMU2320). (a) Basidiospores. (b) Pileipellis. (c) Stipitipellis. (d) Generative hyphae. (e) Skeletal hyphae. (f) Binding hyphae. (g) Basidioles. Scale bars = 10 μm. Drawings by R. Tian.
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).
Table 2. Main characters of five Ganoderma species called “Lingzhi Wang” or “Zhu Lingzhi” in Hainan Island, tropical China.
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.
Figure 12. The varied fruit bodies from cultivated G. subflexipes. Scale bars = 1 cm. Photos by R. Tian.
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 bamboo2
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

Author Contributions

Conceptualization, N.-K.Z. and X.-D.C.; methodology, performing the experiment, and formal analysis, R.T. and H.-Z.Q.; resources, N.-K.Z., Z.-Q.L., X.-Y.Z., X.-D.M., L.X. and T.-C.W.; writing—original draft preparation, R.T. and Q.Z.; writing—review and editing, N.-K.Z. and X.-D.C.; supervision, N.-K.Z.; project administration, N.-K.Z.; funding acquisition, N.-K.Z. All authors have read and agreed to the published version of the manuscript.

Funding

The work was financially supported by the Special Fund Project for Environmental Protection of Hainan Province: Investigation and Assessment of Biodiversity in Changjiang County; the Hainan Institute of National Park, HINP, KY-24ZK02; the Hainan Province Science and Technology Special Fund (ZDYF2023RDYL01); and the Innovation and Entrepreneurship Training Program for College Students in Hainan Province, China (No. S202411658028).

Institutional Review Board Statement

Not applicable.

Data Availability Statement

The datasets presented in this study have been deposited in NCBI GenBank (https://www.ncbi.nlm.nih.gov/genbank/ (accessed on 13 March 2025)) and Mycobank (https://www.mycobank.org/page/Home/MycoBank (accessed on 13 March 2025)).

Acknowledgments

We are grateful to W.Z. Ma, Kunming Institute of Botany, Chinese Academy of Sciences, for providing access to the herbariums to examine the collections of Ganoderma; Y. Liu, Hainan Squirrel Study Abroad Consulting Service Co., Ltd. Haikou, China, for her kind contribution of several specimens and photographs of G. bambusicola; and the forest rangers, Hainan Tropical Rainforest National Park, for their kind help during the field investigations.

Conflicts of Interest

The authors confirm that there are no known conflicts of interest associated with this publication.

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