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Research Article
Phylogenetic analyses and morphological characters reveal two new species of Ganoderma from Yunnan province, China
expand article infoJun He‡, Zong-Long Luo§, Song-Ming Tang|, Yong-Jun Li#, Shu-Hong Li#, Hong-Yan Su‡
‡ Dali University, DaLi, China
§ Dali University, Dali, China
| Mae Fah Luang University, Chiang Rai, Thailand
¶ Institute of Biotechnology and Germplasm Resources, Yunnan Academy of Agricultural Sciences, Kunming, China
# Institute of Biotechnology and Germplasm Resources, Yunnan Academy of Agricultural Sciences, KunMing, China

Corresponding author: Shu-Hong Li ( shuhongfungi@126.com )

Corresponding author: Hong-Yan Su ( suhongyan16@163.com )

Academic editor: Bao-Kai Cui
© 2021 Jun He, Zong-Long Luo, Song-Ming Tang, Yong-Jun Li, Shu-Hong Li, Hong-Yan Su.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation: He J, Luo Z-L, Tang S-M, Li Y-J, Li S-H, Su H-Y (2021) Phylogenetic analyses and morphological characters reveal two new species of Ganoderma from Yunnan province, China. MycoKeys 84: 141-162. https://doi.org/10.3897/mycokeys.84.69449
Open Access

Abstract

Ganoderma dianzhongense sp. nov. and G. esculentum sp. nov. are proposed as two new species based on both phenotypic and genotypic evidences. Ganoderma dianzhongense is characterized by the stipitate basidiomata, laccate and oxblood red pileus, gray white pore surface, duplex context and broadly ellipsoid basidiospores (9.0–12.5 × 6.5–9.0 μm) with coarse interwall pillars. Ganoderma esculentum is characterized by its basidiomata with slender stipe, white pore surface, homogeneous pileus context, and slightly truncate, narrow basidiospores (8.0–12.5 × 5.0–8.0 µm). Phylogenetic analyses were carried out based on the internal transcribed spacer (ITS), translation elongation factor 1-α (TEF1-α) and the second subunit of RNA polymerase II (RPB2) sequence data. The illustrations and descriptions for the new taxa are provided.

Keywords

Ganodermataceae, novel species, phylogeny, taxonomy

Introduction

Ganodermataceae was introduced by Donk (1948) which belongs to Polyporales and the latest studies indicated that it is a monophyletic group (Costa-Rezende et al. 2020). Currently, eleven genera viz. Amauroderma Murril, Amaurodermellus Costa-Rezende, Cristataspora Costa-Rezende, Foraminispora Robledo, Costa-Rezende & Drechsler-Santos, Furtadoa Costa-Rezende, Robledo & Drechsler-Santos, Ganoderma P. Karst., Haddowia Steyaert, Humphreya Steyaert, Magoderna (Murrill) Steyaert, Sanguinoderma Y.F. Sun, D.H. Costa & B.K. Cui and Tomophagus Murrill are accepted in Ganodermataceae and supported by morphology and phylogeny (Steyaert 1972; Furtado 1981; Corner 1983; Zhao and Zhang 2000; Ryvarden 2004; Thametal 2012; Costa-Rezende et al. 2017; Costa-Rezende et al. 2020; Sun et al. 2020).

Ganoderma P. Karst (Ganodermataceae, Polyporales) was introduced to accommodate a laccate and stipitate fungus, Ganoderma lucidum (Curtis) P. Karst (Karsten 1881). Ganoderma is characterized by double-walled basidiospores with inter-wall protuberances (Karsten 1881; Moncalvo and Ryvarden 1997). There are 462 records in the Index Fungorum (http://www.Indexfungorum.org/; accessed date: 7 October 2021) and 506 records in MycoBank (http://www.mycobank.org/; accessed date: 7 October 2021). Ganoderma is one of the most taxonomically scrutinized genera among the Ganodermataceae and even in Polyporales (Richter et al. 2015; Costa-Rezende et al. 2020). Most Ganoderma species are wood decomposers, found in all temperate and tropical regions (Pilotti et al. 2004; Cao et al. 2012; Zhou et al. 2015).

Ganoderma has long been regarded as one of the most important medicinal fungi in the world (Paterson 2006); they have been used as medicine for over two millennia in China (Dai et al. 2009). Several Ganoderma species are known to be prolific sources of highly active bioactive compounds, especially polysaccharides, protein, sterols, and triterpenoids (Ahmadi and Riazipour 2007; Chan et al. 2007). These compounds are known to possess extensive therapeutic properties, such as antioxidant, antitumor, and antiviral agents, and improve sleep function (De Silva et al. 2013).

Species diversity of Ganoderma is abundant in China and more than 30 species have been described (Zhao and Zhang 2000; Wang et al. 2009; Cao et al. 2012; Li et al. 2015; Xing et al. 2016; Hapuarachchi et al. 2018; Liu et al. 2019; He et al. 2019; Wu et al. 2020). Yunnan province is considered as one of the hot-spots for studying biodiversity of polypores, and some new Ganoderma species have been described (Zhao 1989; Wang et Wu 2010; Cao and Yuan 2013).

During our investigation into the diversity of Ganoderma in Yunnan province, several specimens of Ganoderma were collected from central and southern Yunnan. Phylogenetic analysis showed that the seven collections formed two distinct lineages and can be recognized as new species, hence two new species, namely G. dianzhongense and G. esculentum are introduced based on morphology and phylogeny.

Materials and methods

Sample collection

Seven Ganoderma specimens were collected during the rainy season from July 2016 to August 2019 in Yunnan Province of China. The samples were then photographed and transported back to the laboratory where their fresh macroscopic details were described. The specimens were deposited in the herbarium of Kunming Institute of Botany Academia Sinica (KUN-HKAS).

Morphological studies

Macro-morphological characters were described based on fresh material field notes, and the photographs provided here. Color codes are from Kornerup and Wanscher (1978). Micro-morphological data were obtained from the dried specimens and observed by using a microscope following Li et al. (2015). Sections were studied at magnification of up to 1000× using a NiKon E400 microscope and phase contrast illumination. Microscopic features and measurements were made from slide preparations stained with 5% potassium hydroxide (KOH) and 2% Melzer’s reagent. Basidiospore features, hyphal system, color, sizes and shapes were recorded and photographed. Measurements were made using the Image Frame work v.0.9.7 to represent variation in the size of basidiospores, 5% of measurements were excluded from each end of the range and extreme values are given in parentheses.

The following abbreviations are used: IKI = Melzer’s reagent, IKI– = neither amyloid nor dextrinoid, KOH = 5% potassium hydroxide, CB = Cotton Blue, CB+ = Cyanophilous (Xing et al. 2018). The abbreviation for basidiospores measurements (n/m/p) denote “n” basidiospores measured from “m” basidiomata of “p” specimens. Basidiospore dimensions (and “Q” values) are given as (a) b–av–c (d), where “a” represents the minimum, “d’ the biggest, “av” the average “b” and “c” covers a minimum of 90% of the values. “Q”, the length/width ratio of a spore in side view, and “Qm” for the average of all basidiospores ± standard deviation (Wang et al. 2015).

DNA extraction, PCR amplification, and sequencing

Total genomic DNA was extracted from dried pieces of pileus with tubes with modified CTAB protocol Doyle (1987). The genes ITS, TEF1-α and RPB2 were amplified by polymerase chain reaction (PCR) technique. The primers ITS1F / ITS4, TEF1-983 / TEF1-1567, and RPB2-6f / fRPB2-7cR were used to amplify the ITS, TEF1-α, RPB2 region, respectively (White et al. 1990; Liu et al. 1999; Matheny et al. 2007). PCR reactions (25 μL) contained mixture: 2.5 μL PCR reaction buffer, 2.5 μL 0.2% BSA, 2 μL dNTP (2.5 mm), 0.5 μL each of primer, 0.2 μL 5 U/μL Taq DNA polymerase, 1–1.5 μL DNA solution and 16 μL sterilized distilled H2O. The PCR cycling for ITS was as follows: initial denaturation at 94 °C for 5 min, followed by 35 cycles at 94 °C for 30 sec, 53 °C for 30 sec and 72 °C for 50 sec and a final extension of 72 °C for 10 min. The PCR cycling for TEF1-α was as follows: initial denaturation at 94 °C for 5 min, followed by 35 cycles at 94 °C for 30 sec, 55 °C for 30 sec and 72 °C for 50 sec and a final extension of 72 °C for 10 min. The PCR cycling for RPB2 was as follows: initial denaturation at 94 °C for 5 min, followed by 35 cycles at 94 °C for 30 sec, 50 °C for 30 sec and 72 °C for 50 sec and a final extension of 72 °C for 10 min. The PCR products were visualized via UV light after electrophoresis on 1% agarose gels stained with ethidium bromide. Successful PCR products were sent to Sangon Biotech Limited Company (Shanghai, China), using forward PCR primers. When sequences have heterozygous INDELS or ambiguous sites, samples were sequenced bidirectionally to make contigs of the amplified regions or verify the ambiguous sites (Wang et al. 2015). Raw DNA sequences were assembled and edited in Sequencher 4.1.4 and the assembled DNA sequences were deposited in GenBank (Table 1).

Sequencing and sequence alignment

Sequence data of three partial loci Internal transcribed spacer region (ITS), RNA polymerase II subunit 2 (RPB2), and translation elongation factor 1-alpha (TEF1-α) were used in the phylogenetic analyses. Besides the sequences generated from this study, other reference sequences were selected from GenBank for phylogenetic analyses (Table 1). Sequences were aligned using the online version of MAFFT v.7 (http://mafft.cbrc.jp/alignment/server/) (Katoh and Standley 2013) and adjusted using BioEdit v.7.0.9 by hand (Hall 1999) to allow maximum alignment and minimize gaps. Ambiguous regions were excluded from the analyses and gaps were treated as missing data. The phylogeny website tool “ALTER” (Glez-Peña et al. 2010) was used to convert the alignment fasta file to Phylip format for RAxML analysis and AliView and PAUP 4.0b 10 were used to convert the alignment fasta file to a Nexus file for Bayesian analysis (Swofford 2003). Phylogenetic analyses were obtained from Maximum Likelihood (ML) and Bayesian analysis (BI).

Molecular phylogenetic analyses

The maximum likelihood (ML) and Bayesian inference (BI) methods were used to analyze the combined dataset of ITS, TEF1-α and RPB2 sequences. Maximum likelihood analysis was conducted with RAxML-HPC2 on the CIPRES Science Gateway (Miller et al. 2010), involved 100 ML searches; all model parameters were estimated by the program. The ML bootstrap values (ML-BS) were obtained with 1000 rapid bootstrapping replicates. Maximum likelihood bootstrap values (ML) equal to or greater than 70% are given above each node (Figure 1).

Table 1.
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Species, specimens, geographic origin and GenBank accession numbers of sequences used in this study.

Species Voucher/strain Origin GenBank accession numbers Reference
ITS TEF1-α RPB2
Ganoderma aridicola Dai 12588 (Type) South Africa KU572491 KU572502 – Xing et al. 2016
G. adspersum GACP15061220 Thailand MK345425 MK371431 MK371437 Hapuarachchi et al. 2019
MFLU 19-2178 Thailand MN396653 MN423149 MN423114 Luangharn et al. 2021
G. angustisporum Cui 13817(Type) Fujian, China MG279170 MG367563 MG367507 Xing et al. 2018
Cui 14578 Guangdong, China MG279171 MG367564 – Xing et al. 2018
G. austral CMW 47785 South Africa MH571686 MH567276 – Tchoumi et al. 2018
CMW 48146 South Africa MH571685 MH567283 – Tchoumi et al. 2018
G. austroafricanum CBS138724 (Type) South Africa KM507324 – – Coetzee et al. 2015
G. aff. austroafricanum CMW25884 South Africa MH571693 MH567296 – Tchoumi et al. 2019
G. bambusicola Wu 1207-151(Type) Taiwan, China MN957781 LC517941 LC517944 Wu et al. 2020
Wu 1207-152 Taiwan, China MN957782 LC517942 LC517945 Wu et al. 2020
Wu 1207-153 Taiwan, China MN957783 LC517943 LC517946 Wu et al. 2020
G. boninense WD 2028 Japan KJ143905 KJ143924 KJ143964 Zhou et al. 2015
WD 2085 Japan KJ143906 KJ143925 KJ143965 Zhou et al. 2015
G. calidophilum MFLU 19-2174 Yunnan, China MN398337 – – Luangharn et al. 2021
H36 Yunnan, China MW750241* MW838997* MW839003* this study
G. carnosum MJ 21/08 Czech R, Europe KU572492 – – Xing et al. 2016
JV 8709/8 Czech R, Europe KU572493 – – Xing et al. 2016
G. carocalcareus DMC 322 (Type) Cameroon EU089969 – – Douanla and Langer 2009
DMC 513 Cameroon EU089970 – – Douanla and Langer 2009
G. casuarinicola Dai 16336 (Type) Guangdong, China MG279173 MG367565 MG367508 Xing et al. 2018
Dai 16339 Guangdong, China MG279176 MG367568 MG367511 Xing et al. 2018
G. curtisii CBS 100131 NC, USA JQ781848 KJ143926 KJ143966 Zhou et al. 2015
CBS 100132 NC, USA JQ781849 KJ143927 KJ143967 Zhou et al. 2015
G. destructans CBS 139793 (Type) South Africa NR132919 – – Coetzee et al. 2015
Dai 16431 South Africa MG279177 MG367569 MG367512 Xing et al. 2018
G. dunense CMW42157 (Type) South Africa MG020255 MG020227 – Tchoumi et al. 2019
CMW42150 South Africa MG020249 MG020228 – Tchoumi et al. 2019
G. ecuadoriense ASL799 (Type) Ecuador KU128524 – – Crous et al. 2016
PMC126 Ecuador KU128525 – – Crous et al. 2016
G. eickeri CMW 49692 (Type) South Africa MH571690 MH567287 – Tchoumi et al. 2019
CMW 50325 South Africa MH571689 MH567290 – Tchoumi et al. 2019
G. ellipsoideum GACP1408966(Type) Hainan, China MH106867 – – Hapuarachchi et al. 2018
GACP14081215 Hainan, China MH106886 – – Hapuarachchi et al. 2018
G. enigmaticum Dai 15970 Africa KU572486 KU572496 MG367513 Xing et al. 2016
Dai 15971 Africa KU572487 KU572497 MG367514 Xing et al. 2016
G. esculentum L4935 (Type) Yunnan, China MW750242* MW838998* MW839004* this study
L4946 Yunnan, China MW750243* MW838999* – this study
G. flexipes Wei 5494 Hainan, China JN383979 – – Cao and Yuan 2013
MFLU 19-2198 Yunnan, China MN398340 – – Luangharn et al. 2021
G. gibbosum MFLU 19-2176 Thailand MN396311 – MN423118 Luangharn et al. 2021
MFLU 19-2190 Laos MN396310 – MN423117 Luangharn et al. 2021
G. heohnelianum Dai 11995 Yunnan, China KU219988 MG367550 MG367497 Song et al. 2016
Cui 13982 Guangxi, China MG279178 MG367570 MG367515 Xing et al. 2018
G. hochiminhense MFLU 19-2224(Type) Vietnam MN398324 MN423176 – Luangharn et al. 2021
MFLU 19-2225 Vietnam MN396662 MN423177 – Luangharn et al. 2021
G. knysnamense CMW 47755 (Type) South Africa MH571681 MH567261 – Tchoumi et al. 2019
CMW 47756 South Africa MH571684 MH567274 – Tchoumi et al. 2019
G. leucocontextum GDGM 44303 Xizang, China KJ027607 – – Li et al. 2015
GDGM 44305 Xizang, China KJ027609 – – Li et al. 2015
G. lingzhi Cui 9166 China KJ143907 JX029974 JX029978 Cao et al. 2012
Dai 12574 Liaoning, China KJ143908 JX029977 JX029981 Cao et al. 2012
G. lobatum JV 1008/31 USA KF605671 MG367553 MG367499 Xing et al. 2018
JV 1008/32 USA KF605670 MG367554 MG367500 Xing et al. 2018
G. lucidum K 175217 UK KJ143911 KJ143929 KJ143971 Zhou et al. 2015
MT 26/10 Czech Republic KJ143912 KJ143930 – Zhou et al. 2015
G. martinicense 231NC NC, USA MG654182 MG754736 – Loyd et al.2018
246TX TX, USA MG654185 MG754737 MG754858 Loyd et al.2018
G. mbrekobenum UMN7-3 GHA (Type) Ghana KX000896 – – Crous et al. 2016
UMN7-4 GHA Ghana KX000898 – – Crous et al. 2016
G. mexicanum MUCL 49453 SW17 Martinique MK531811 MK531825 MK531836 Cabarroi-Hernández et al. 2019
MUCL 55832 Martinique MK531815 MK531829 MK531839 Cabarroi-Hernández et al. 2019
G. mizoramense UMN-MZ4 (Type) India KY643750 – – Crous et al. 2017
UMN-MZ5 India KY643751 – – Crous et al. 2017
G. multipileum CWN 04670 Taiwan, China KJ143913 KJ143931 KJ143972 Zhou et al. 2015
Dai 9447 Hainan, China KJ143914 – KJ143973 Zhou et al. 2015
G. multiplicatum SPC9 Brazil KU569553 – – Bolaños et al. 2016
URM 83346 Brazil JX310823 – – Bolaños et al. 2016
G. mutabile CLZhao 982 Yunnan, China MG231527 – – GenBank
Yuan 2289(Type) Yunnan, China JN383977 – – Cao and Yuan 2013
G. myanmarense MFLU 19-2167 (Type) Myanmar MN396329 – – Luangharn et al. 2021
MFLU 19-2169 Myanmar MN396330 – – Luangharn et al. 2021
G. nasalanense GACP17060211 (Type) Laos MK345441 – – Hapuarachchi et al. 2019
GACP17060212 Laos MK345442 – – Hapuarachchi et al. 2019
G. neojaponicum FFPRI WD-1285 Tokyo, Japan MN957784 – – Wu et al. 2020
FFPRI WD-1532 Chiba, Japan MN957785 – – Wu et al. 2020
G. orbiforme Cui 13918 Hainan, China MG279186 MG367576 MG367522 Xing et al. 2018
Cui 13880 Hainan, China MG279187 MG367577 MG367523 Xing et al. 2018
G. parvulum MUCL 47096 Cuba MK554783 MK554721 MK554742 Cabarroi-Hernández et al. 2019
MUCL 52655 French Guiana MK554770 MK554717 MK554755 Cabarroi-Hernández et al. 2019
G. philippii Cui 14443 Hainan, China MG279188 MG367578 MG367524 Xing et al. 2018
Cui 14444 Hainan, China MG279189 MG367579 MG367525 Xing et al. 2018
G. resinaceum Rivoire 4150 France, Europe KJ143915 – – Zhou et al. 2015
CBS 19476 Netherlands, Europe KJ143916 KJ143934 – Zhou et al. 2015
G. ryvardenii HKAS 58053 (Type) South Africa HM138670 – – Kinge et al. 2011
HKAS 58054 South Africa HM138671 – – Kinge et al. 2011
G. sessile 111TX TX, USA MG654306 MG754747 MG754866 Loyd et al.2018
113FL FL, USA MG654307 MG754748 MG754867 Loyd et al.2018
G. shanxiense BJTC FM423(Type) Shanxi, China MK764268 MK783937 MK783940 Liu et al. 2019
HSA 539 Shanxi, China MK764269 – MK789681 Liu et al. 2019
G. sichuanense HMAS42798 (Type) Sichuan, China JQ781877 – – Cao et al. 2012
Cui 7691 Guangdong, China JQ781878 – – Cao et al. 2012
G. sinense Wei 5327 Hainan, China KF494998 KF494976 MG367529 Xing et al. 2018
Cui 13835 Hainan, China MG279193 MG367583 MG367530 Xing et al. 2018
G. steyaertanum MEL:2382783 Australia KP012964 – – GenBank
6 WN 20B Indonesia KJ654462 – – Glen et al. 2014
G. thailandicum HKAS 104640 (Type) Thailand MK848681 MK875829 MK875831 Luangharn et al. 2019
HKAS 104641 Thailand MK848682 MK875830 MK875832 Luangharn et al. 2019
G. tropicum He 1232 Guangxi, China KF495000 KF494975 MG367531 Xing et al. 2016
HKAS 97486 Thailand MH823539 – MH883621 Luangharn et al. 2021
G. tsugae UMNMI20 MI, USA MG654324 MG754764 – Loyd et al.2018
UMNMI30 MI, USA MG654326 MH025362 MG754871 Loyd et al.2018
G. tuberculosum GVL-21 Veracruz, Mexico MT232639 – – Espinosa-García et al. 2021
GVL-40 Veracruz, Mexico MT232634 – – Espinosa-García et al. 2021
G. weberianum CBS 128581 Taiwan, China MK603805 MK636693 MK611971 Cabarroi-Hernández et al. 2019
CBS 219.36 Philippines MK603804 MK611974 MK611972 Cabarroi-Hernández et al. 2019
G. wiiroense UMN-21-GHA (Type) Ghana KT952363 – – Crous et al. 2015
UMN-20-GHA Ghana KT952361 – – Crous et al. 2015
G. dianzhongense L4331(Type) Yunnan, China MW750237* MW838993* MZ467043* this study
L4230 Yunnan, China MW750236* MW838992* – this study
L4737 Yunnan, China MW750238* MW838994* MW839000* this study
L4759 Yunnan, China MW750239* MW838995* MW839001* this study
L4969 Yunnan, China MW750240* MW838996* MZ467044* this study
G. zonatum FL-02 FL, USA KJ143921 KJ143941 KJ143979 Zhou et al. 2015
FL-03 FL, USA KJ143922 KJ143942 KJ143980 Zhou et al. 2015
Tomophagus colossus TC-02 Vietnam KJ143923 KJ143943 – Zhou et al. 2015

*Newly generated sequences for this study. Bold font = new species.

Bayesian analysis was performed with MrBayes v3.2 (Ronquist et al. 2012), with the best-fit model of sequence evolution estimated with MrModeltest 2.3 (Nylander et al. 2008) to evaluate posterior probabilities (PP) (Rannala and Yang 1996; Zhaxybayeva and Gogarten 2002) by Markov Chain Monte Carlo (MCMC) sampling. Six simultaneous Markov chains were run for 10,000,000 generations, trees were sampled every 500th generation, and 2,000 trees were obtained. The first 5000 trees, representing the burn-in phase of the analyses, were discarded, while the remaining 1500 trees were used for calculating posterior probabilities in the majority rule consensus tree (the critical value for the topological convergence diagnostic is 0.01).

The phylogenetic tree was visualized with FigTree version 1.4.0 (Rambaut 2012) and made in Adobe Illustrator CS5 (Adobe Systems Inc., USA). Sequences derived in this study were deposited in GenBank (http://www.ncbi.nlm.nih.gov). The final sequence alignments and the phylogenetic trees are available at TreeBase (http://www.treebase.org, accession number: 28875).

Results

Phylogenetic analyses

The dataset composed of ITS, TEF1-α and RPB2 genes, comprising a total of 2092 characters including gaps, ITS (1–656 bp), TEF1-α (657–1192 bp) and RPB2 (1193–2092 bp), including 57 taxa with Tomophagus colossus (Fr.) C.F. Baker as the outgroup taxon (Wang et al. 2009; Cao et al. 2012). Best model for the combined 3-gene dataset estimated and applied in the Bayesian analysis was GTR+I+G, lset nst = 6, rates = invgamma; prset statefreqpr = dirichlet (1,1,1,1). The phylogenetic analysis of ML and BI produce similar topology. The combined dataset analysis of RAxML generates a best-scoring tree (Figure 1), with the final ML optimization likelihood value of -13861.891117. The aligned matrix had 993 distinct alignment patterns, with 38.83% completely undetermined characters or gaps. The base frequency and rate are as follows: A = 0.215319, C = 0.266028, G = 0.260220, T = 0.258433; rate AC = 0.885915, AG = 5.586021, AT = 0.936363, CG = 1.205084, CT = 6.595971, GT = 1.000000; gamma distribution shape: α = 0.246210. Bootstrap support values with a maximum likelihood (ML) greater than 70%, and Bayesian posterior probabilities (BPP) greater than 0.95 are given above the nodes (Figure 1).

Phylogenetic analysis showed that five collections clustered together with high bootstrap support, forming a clade sister to G. shanxiense with strong bootstrap support (ML-BS = 96%, BPP = 1.00, Figure 1). Two other collections clustered with G. aridicola, G. bambusicola, G. casuarinicola, G. calidohilum, G. enigmaticum and G. thailandicum (ML-BS = 100%, BPP = 1.00), but forming as a distinct lineage.

Figure 1. 

Phylogeny of the new Ganoderma species and related taxa based on ITS, TEF1-α and RPB2 sequence data. Branches are labeled with bootstrap values (ML) higher than 70%, and posterior probabilities (BPP) higher than 0.95. The new species are shown in bold red.

Taxonomy

Ganoderma dianzhongense J. He, H.Y. Su & S.H. Li, sp. nov.

Index Fungorum number: 558822
MycoBank No: 841408
Figure 2

Diagnosis

Ganoderma dianzhongense is characterized by its mesopodal basidiomata, oxblood red to violet brown pileus surface, melon seed kernel-shaped and broadly ellipsoid basidiospores.

Holotype

China. Yunnan Province, Kunming City, Luquan County, on the rotten broad-leaved trees, alt. 2480 m, Shu-Hong Li, 8 Sept. 2016, L4331 (HKAS 110005).

Etymology

The epithet ‘dianzhong’ refers to central Yunnan province in Chinese, where the holotype was collected.

Description

Basidiomata annual, stipitate, sub-mesopodal to mesopodal or with the back sides fused, coriaceous to woody. Pileus single, suborbicular to reniform, up to 4.8–13.1 cm diam., 1.1 cm thick, weakly to strongly laccate, glossy and shiny, oxblood red (9E7) to violet brown (11F8), smooth, and covered by a thin hard crust, concentrically zonate or azonate. Margin distinct, slightly obtuse. Stipe 9.0–17.7 × 1.1–1.9 cm, central, cylindrical, strongly laccate, dark red brown (11C8) to purplish (14A8) or almost blackish red-brown (10F4), fibrous to woody. Context up to 0.4 cm thick, duplex; lower layer dark brown (8F8), fibrous, composed of coarse loose fibrils; upper layer putty (4B2); corky to woody, bearing distinct concentric growth zones, without black melanoid band. Tubes woody hard, grayish brown, up to 0.9 cm long, unstratified. Pore 4–6 per mm, round to angular, dissepiments slightly thick, entire; pore surface grey white to lead gray (2D2), turning light buff when dust (5D1).

Hyphal system trimitic. Generative hyphae 2.0–3.5 μm in diameter, colorless, thin-walled, clamp connections present; skeletal hyphae 3.0–6.0 μm in diameter, subthick-walled to solid, non-septate, arboriform with few branches, yellowish to golden-yellow; binding hyphae 1–2.5 μm in diameter, thick-walled, frequently branched, interwoven, hyaline to yellowish, scarce; all the hyphae IKI–, CB+; tissues darkening in KOH.

Pileipellis a crustohymeniderm, cells 20–45 × 5.5–7.5 μm, clavate to cylindrical, entire or rarely with one lateral protuberance, thick-walled, without granulations in the apex, golden-yellow to yellowish-brown, thick-walled, moderately amyloid at maturity.

Basidiospores (80/6/3) (9.0) 10–11.0–12.0 (12.5) × (6.5) 7.0–7.9–8.5 (9.0) μm, Q = (1.12) 1.25–1.55 (1.63), Qm = 1.40±0.09 (including myxosporium); holotype: (40/2/1) 10.0–10.9–12 × 7.0–7.9–8.5 (9.0) μm,Q = (1.20) 1.25–1.52, Qm = 1.39±0.08 (including myxosporium). mostly melon seed-shaped at maturity to broadly ellipsoid, usually with one end tapering and obtuse at maturity, with apical germ pore, yellowish to medium brown, IKI–, CB+, inamyloid; perisporium wrinkled, double-walled, with coarse interwall pillars. Basidia widely clavate to utriform, hyaline, with a clamp connection and four sterigmata, 11–19 ×10–13µm; basidioles pear-shaped to fusiform, 10–15 × 8–12 µm.

Figure 2. 

Ganoderma dianzhongense (HKAS 110005, holotype) A basidiomata B upper surface C cut side of pileus D pore surface E sections of pileipellis (LM) F skeletal hyphae from context (LM) G binging hyphae from tubes (LM) H generative hyphae from tubes (LM) I-J basidia and basidioles (LM) K-L basidiospores (LM) M-N basidiospores (SEM) O-P culture after incubation at 28 °C for 8 days. Scale bars: 20 mm (O, P); 10 µm (E-L); 5 µm (M, N). Photographs Jun He.

Habit

Scattered, during fall, decaying wood of broad-leaved trees including Quercus sp. Currently, only known from central Yunnan province, China.

Additional specimens examined

China. Yunnan province, Shilin County, alt. 2109m, Jun He, 28 Aug., 2019, L4969 (HKAS 112719); Songming County, alt. 2204m, Shu-Hong Li, 8 Jul., 2016, L4230 (HKAS 112716); Wuding County, alt. 2295m, Shu-Hong Li, 24 Jul., 2019, L4737 (HKAS 112717); ibid, alt. 2432m, Jun He, 26 Jul., 2019, L4759 (HKAS 112718).

Ganoderma esculentum J. He & S.H. Li, sp. nov.

Index Fungorum number: 558823
MycoBank No: 841409
Figure 3

Diagnosis

Ganoderma esculentum is characterized by its strongly laccate chocolate brown pileus surface, slender stipe and narrow ellipsoid basidiospores.

Holotype

China. Yunnan Province, Honghe City, Mengzi County, on a decaying wood log, alt. 1370 m, Jun He, 26 Aug., 2019, L4935 (HKAS 110006).

Etymology

The epithet ‘esculentum’ refers to this species named after a food.

Description

Basidiomata annual, stipitate, pleuropodal, laccate, woody-corky. Pileus single, sub-orbicular to reniform to spathulate, up to 2.8–8.0 × 2.0–4.5 cm diam, 0.75 cm thick at the base, slightly convex to applanate; surface glabrous, rugose to radially rugose, strongly laccate, not cracking, with a hard crust, difficult to penetrate with the fingernail; surface brownish-black (6C8) to chocolate brown (6F4), almost homogeneous in the adult. Margin grayish orange(6B5) to concolorous, entire, acute to obtuse, smooth to sulcate. Stipe 10.0–17.5 × 0.5–1.0 cm, dorsally lateral to nearly dorsal, sub-cylindrical, solid, surface smooth, very shiny, dark brown (8F8) almost black, darker than pileus, fibrous to woody. Context up to 0.2 cm thick, composed of coarse loose fibrils, dark brown (8F8), with black melanoid band. Tubes 0.2–0.5 cm long, dark brown, woody hard, unstratified. Pore 5–8 per mm, circular or sub-circular, woody; pore surface white when fresh, darkening to soot brown(5F5) when aging and drying.

Hyphal system trimitic. Generative hyphae 1.5–3.0 μm in diameter, colorless, thin-walled, clamp connections present; skeletal hyphae 3.5–5.5 μm in diameter, thick-walled to solid, non-septate, arboriform or not, non-branched or with a few branches in the distal end, golden brown; binding hyphae 1.0–3.0 μm in diameter, thick-walled, much-branched, arboriform, hyaline to yellowish, scarce; all the hyphae IKI–, CB+; tissues darkening in KOH.

Pileipellis a crustohymeniderm, cells 20–55 × 10–15 μm, narrowly clavate to tubular, generally smooth, slightly thick-walled to thick-walled with a wide lumen, occasionally expanded at the apex, without granulations, entire, yellowish to leather brown, weakly to strongly amyloid.

Basidiospores (40/3/2) (8.0) 9.0–10.6–12.5 × (5.0) 5.5–6.6–7.5 (8.0) μm, Q = (1.15) 1.34–1.62–2.01 (2.06), Qm = 1.62±0.19 (including myxosporium); holotype: (20/2/1) 9.0–10.6–12.5 × (5.0) 5.5–6.5–7.0 (8.0) μm, Q = (1.34) 1.45–1.64–1.83 (2.06), Qm = 1.64±0.15 (including myxosporium). narrow ellipsoid to truncate, slightly visible apical germ pore, brownish orange to light brown, IKI–, CB+, inamyloid; with a brown eusporium bearing fine, overlaid by a hyaline myxosporium, with interwall pillars. Basidia not observed.

Figure 3. 

Ganoderma esculentum holotype (HKAS 110006) A basidiomata B upper surface C lower surface D cut side of pileus E pore surface F sections of pileipellis (LM) G, H skeletal hyphae from context (LM) I binging hyphae from tubes (LM) J generative hyphae from tubes (LM) K–M basidiospores (LM) N, O basidiospores (SEM). Scale bars: 20 µm (H); 10 µm (F, G, I-M); 5 µm (N, O). Photographs Jun He.

Habit

On decaying hardwood trees or bamboo roots, accompanied in humus rich soil with over heavily rotted litter on the ground.

Additional specimens examined

China. Yunnan province, Mengzi City, Xinansuo Town, alt. 1328m, Jun He, 26 Aug., 2019, L4946 (HKAS 112720).

Discussion

Ganodermataceae is a large family of polypores, and has received great attention from mycologists for over many decades. However, species identification and circumscriptions have been unclear and taxonomic segregation of the genera has been controversial because of different viewpoints among mycologists (Moncalvo et al. 1995; Moncalvo and Ryvarden 1997; Costa-Rezende et al. 2020). Ganodermataceae was treated as a synonym of Polyporaceae and classify the genus Ganoderma into Polyporaceae by Justo et al. (2017). Later, Cui et al. (2019) excluded Ganoderma from Polyporaceae, due to Ganoderma having unique double-walled basidiospores. In addition, recent studies have clarified someuncer-tainties of generic delimitation and classification of polypores with ganodermatoid basidiospores, and proved that Ganodermataceae is a monophyletic group (Costa-Rezende et al. 2020). More collections of this family are needed in order to estimate the attributes of this taxon better.

In the phylogenetic inferences, Ganoderma dianzhongense is sister to G. shanxiense, which is known from the northern Shanxi province in China (Figure 1). Morphologically, both species share similar characters of the mesopodal basidiomata, suborbicular to reniform pileus, and broadly ellipsoid basidiospores (Table 2). However, G. shanxiense differs from G. dianzhongense in having a red to reddish-brown pileus surface, wider basidiospores (11.0–13.0 × 8.0–9.5 μm), and narrower skeletal hyphae (2.5–5.0 μm, Liu et al. 2019).

Ganoderma dianzhongense resembles G. sinense and G. orbiforme in having suborbicular pileus (Table 2). However, G. sinense is characterized by wider basidiospores (9.5–13.4 × 7.0–10.2 μm) and slightly longitudinally crested basidiospores (Wang and Wu 2007) and a uniformly brown to dark brown context. Ganoderma orbiforme has a purplish black to light brown pileus, a variably brown context, irregularly digitated pileipellis cells, and ellipsoid to ovoid basidiospores (6.9–10.6 × 3.6–5.7 μm) with fine and short echinulae, and a subtropical to tropical distribution (Wang et al. 2014). Ganoderma orbiforme is also phylogenetically unrelated (Figure 1).

In our multi-locus phylogeny analysis (Figure 1), G. aridicola, G. bambusicola, G. casuarinicola, G. calidohilum, G. enigmaticum, G. mbrekobenum, G. thailandicum and G. esculentum formed a distinct lineage, and was clearly separated from other Ganoderma species. It is easy to distinguish them from the morphological characteristics. Ganoderma bambusicola has a longer pileipellis (35–65 × 8–16 μm) and wider basidiospores than those of G. esculentum (10.0–13.0 × 6.5–8.0 µm, Wu et al. 2020). Ganoderma aridicola can be easily distinguished from G. esculentum by the sessile basidiomata and a fuscous to black pileus surface (Xing et al. 2016). Ganoderma casuarinicola differs from G. esculentum by the latter has smaller basidiospores (8.3–11.5 × 4.5–7.0 µm, Xing et al. 2018), grayish brown longer pores and sectorial to shell-shaped pileus. Ganoderma enigmaticum mainly differs from G. esculentum by its golden yellow pileus surface, narrower basidiospores (8.0–11.0 × 3.5–6.0 µm, Coetzee et al. 2015) and causes root and butt rot of living and dead trees. Ganoderma thailandicum can be distinguished from G. esculentum, by its brownish-red pileus surface without radially rugose, narrowly clavate pileipellis cells with tuberculate and smaller basidiospores (6.8–10.2 × 5.8–7.7 µm, Luangharn et al. 2019). Ganoderma mbrekobenum can be differentiated from G. esculentum by its woody to corky texture when dried, with ovoid basidiospores (25.0–57.0 × 6.0–12.0µm, Crous et al. 2016). Ganoderma calidophilum has a larger diameter binding hypha (2.4–5.2 µm) than G. esculentum (1.0–3.0 μm) and G. calidophilum has larger basidiospores (7.3–14.6 × 5.3–9.6 µm, Zhao et al. 1979; Luangharn et al. 2021) than G. esculentum (including myxosporium).

Table 2.
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Morphological comparison of Ganoderma dianzhongense sp. nov., and G. esculentum sp. nov., with their closest relatives in the combined phylogeny.

Species Shape Context Pileipellis cells Pores Basidiospores (μm) Reference
Ganoderma aridicola sessile dimidiate context corky, fuscous, black melanoid band absent moderately amyloid at maturity, 30–55 × 5–8 μm 6–8 per mm 9.7–11.2 × 7.0–7.8 Xing et al. 2016
G. bambusicola stipitate, reniform to semicircular context fairly homogeneous, brownish,1–2 mm thick clavate or cylindrical, 35–65 × 8–16 μm 5–6 per mm 11.0–12.5 × 6.5––7.5 Wu et al. 2020
G. carnosum laterally to rarely eccentrically stipitate, dimidiate, orbicular to reniform whitish and soft-corky context amyloid elements up to 75 μm from clamp to the ape 3–4 per mm 10.0–13 × 7.0–8.5 Patouillard 1889
G. calidophilum stipitate, round or half-round duplex context, 0.1–0.3 cm thick – 4–6 per mm 10.0–13.0 × 6.2–8.7 Zhao et al. 1979
G. casuarinicola stipitate, sectorial to shell-shaped context corky, black melanoid band absent. moderately amyloid at maturity, 40–70 × 5–13μm 4–6 per mm 8.3–11.5 × 4.5–7.0 Xing et al. 2018
G. dianzhongense stipitate, suborbicular to reniform dark brown context, black melanoid band present amyloid elements, 20–45 × 5.5–7.5 μm 5–8 per mm 9.0–12.5 × 6.5–9.0 this study
G. enigmaticum stipitate globular pileus context soft, dark brown amyloid elements 20–46 × 5.5–9 um 3–5 per mm 8.0–11.0 × 3.5–6.0 Coetzee et al. 2015
G. esculentum stipitate, reniform to spathulate dark brown context, without black melanoid bands weakly to strongly amyloid, 20–55 × 10–15 μm 4–6 per mm 8.0–12.5 × 5.0–8.0 this study
G. kunmingense stipitate, spathulate or half-round context wood color – 4 per mm 7.5–10.5 × 6.0–9.0 Zhao 1989
G. lucidum stipitate to sessile thinner context of white to slightly cream color context amyloid hyphal end cells up to 7–11 μm diam 4–5 per mm 7.7–11.5 × 5.2–8.4 Ryvarden and Gilbertson 1993
G. leucocontextum stipitate, reniform to flabelliform thinner context of white to slightly cream color amyloid elements 30–60 × 8–10 μm 4–6 per mm 9.5–12.5 × 7.0–9.0 Li et al. 2015
G. mbrekobenum stipitate, maroon to liver brown – – 4–6 per mm 8.0–11.5 × 6.0–8.0 Crous et al. 2016
G. neojaponicum stipitate, reniform to suborbicular 0.5 cm thick, duplex brownish orange, clavate like cells 3–5 per mm 9.1–13.5 × 5.7–8.9 Imazeki et al. 1939
G. orbiforme sessile, flabelliform or spathulate context up to 0.4–1.0 cm thick, triplex composed of apically acanthus like branched cells 4–6 per mm 7.1–11.8 × 5.2–7.7 Ryvarden 2000
G. sinense stipitate, dimidiate, suborbicular soft and fibrous, dark brown clavate like cells, dextrinoid 5–6 per mm 9.5–13.8 × 6.9–8.7 Zhao et al. 1979
G. shanxiense stipitate, reniform to dimidiate brown context 25–30 × 7.5–8.5 μm 4–5 per mm 11.0–13.0 × 8.0–9.5 Liu et al. 2019
G. tsugae centrally to laterally stipitate, sub-dimidiate to dimidiate whitish and soft corky context 60–75 × 7–10 μm 4–6 per mm 13.0–15.0 × 7.5–8.5 Murrill 1902
G. thailandicum stipitate, greyish-red to brownish-red context mostly brownish-red to reddish-brown clavate to narrowly clavate, tuberculate 4–8 per mm 6.8–10.2 × 5.8–7.7 Luangharn et al. 2019

Morphologically, G. esculentum resemble G. kunmingense by radially rugose, the pileus and slender stipe (Table 2). However, G. kunmingense has narrower hyphae, tissues not darkening in KOH, and broadly ellipsoid to sub-globose basidiospores (7.5–10.5 × 6.0–9.0 µm, Zhao et al. 1989). In addition, G. esculentum shares also similarities with G. neojaponicum but the latter has a double-layered context with the paler layer near the pileus surface and wider basidiospores than those of G. esculentum (9.1–13.5 × 5.7–8.9 µm, Imazeki et al. 1939; Hapuarachchi et al. 2019).

Acknowledgements

The authors thank Kunming Institute of Botany for providing the experimental platform. Dr Xiang-Hua Wang helped to analyze the molecular data and molecular lab work. We also thank Dr. Dan-Feng Bao and Hong-Wei Shen for their valuable suggestions on this study.

The research was financed by the National Natural Science Foundation of China (Project No. 31970021, 32060006), Yunnan Financial Special Project [YCJ (2020)323, 202102AE090036–05], Yunnan Science Technology Department and Technology Talents and Platform Program “Yunnan Province Technology Innovation Talents Training Objects” (Project ID 2017HB084) and Science and technology innovation and R&D promotion project of Qamdo City.

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Supplementary material

Supplementary material 1 

Phylogenetic sequence dataset

Authors: Jun He

Data type: phylogenetic data

Explanation note: Sequence data of three partial loci internal transcribed spaces region (ITS), RNA polymerase II subunit 2 (RPB2), and translation elongation factor 1-alpha (TEF1-α) were used in the phylogenetic analyses.

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
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