Skip to main content
NCBI home page
Biodiversity Data Journal logoLink to Biodiversity Data Journal
. 2022 Jul 6;10:e80034. doi: 10.3897/BDJ.10.e80034

Ganodermaovisporum sp. nov. (Polyporales, Polyporaceae) from Southwest China

Hong-De Yang 1,2,3, Yong Ding 1,✉, Ting-Chi Wen 4,2,5,✉, Kalani Kanchana Hapuarachchi 6,2,3,5, De-Ping Wei 3,6,2,5,7
PMCID: PMC9848459  PMID: 36761562

Abstract

Background

Ganoderma is a white-rot fungus with a cosmopolitan distribution and includes several economically important species. This genus has been extensively researched due to its beneficial medicinal properties and chemical constituents with potential nutritional and therapeutic values. Traditionally, species of Ganoderma were identified solely based on morphology; however, recent molecular studies revealed that many morphology-based species are conspecific. Furthermore, some type species are in poor condition, which hinders us from re-examining their taxonomic characteristics and obtaining their molecular data. Therefore, new species and fresh collections with multigene sequences are needed to fill the loopholes and to understand the biological classification system of Ganoderma.

New information

In a survey of Ganoderma in Guizhou Province, southwest China, we found a new species growing on soil and, herein, it was identified by both morphology and phylogenetic evidence. Hence, we propose a new species, Ganodermaovisporum sp. nov. This species is characterised by an annual, stipitate, laccate basidiome, with a red–brown to brownish-black pileus surface and pale white pores, duplex context, clavate pileipellis terminal cells, trimitic hyphal system, ellipsoid basidiospores with dark brown eusporium bearing coarse echinulae and an obtuse turgid appendix. Phylogenetic analyses confirmed that the novel species sisters to G.sandunense with high bootstrap support. Furthermore, the RPB2 sequence of G.sandunense is supplied for the first time. Notably, we re-examined the type specimen of G.sandunense and provide a more precise description of the duplex context, pileipellis terminal cells and basidia. All species collected are described and illustrated with coloured photographs. Moreover, we present an updated phylogeny for Ganoderma, based on nLSU, ITS, RPB2 and TEF1-α DNA sequence data and species relationships and classification are discussed.

Keywords: one new species, Ganoderma , morphology, phylogeny, taxonomy

Introduction

Ganodermataceae Donk is a large family of Polyporales and Ganoderma P. Karst is the most speciose genus in the family (Hapuarachchi et al. 2016a, Hapuarachchi et al. 2017, He et al. 2019). Before the molecular era, Polyporales with double-walled basidiospores with a pigmented endosporium ornamented with columns or ridges and a smooth hyaline exosporium were usually placed in Ganodermataceae (Moncalvo and Ryvarden 1997). This family is comprised of ten genera: Amauroderma Murrill, Amaurodermellus Costa-Rezende, Drechsler-Santos & Góes-Neto, Foraminispora Robledo, Costa-Rezende & Drechsler-Santos, Furtadomyces Leonardo-Silva, Cotrim & Xavier-Santos, Ganoderma P. Karst, Haddowia Steyaert, Humphreya Steyaert, Sanguinoderma Y.F. Sun, D.H. Costa & B.K. Cu,Tomophagus Murrill and Trachyderma Imazeki (Richter et al. 2014, Costa-Rezende et al. 2017, Costa-Rezende et al. 2020, Sun et al. 2020, Leonardo-Silva et al. 2022). However, Ganodermataceae has been treated as a synonym of Polyporaceae (Justo et al. 2017). There have been several discrepancies regarding the treatment of Justo et al. (2017); in particular, the studied collection of Ganodermatoid specimens was insufficient to establish a stable taxonomic and systematic placement in a phylogenetic context because some herbarium materials have been destroyed or cannot be found, lacking molecular and morphological data and the characterised double-walled basidiospores in Ganodermataceae are quite different from those in Polyporaceae (Cui et al. 2019, Costa-Rezende et al. 2020). In this study, we subsequently followed Justo et al. (2017) since the phylogenetic analyses are more convincing and objective than morphological results.

Ganoderma was introduced by Karsten (1881) and typified by G.lucidum (Curtis) P. Karst. (syn. Polyporuslucidus; bas. Boletuslucidus Curtis), a species with stipitate and laccate white-rot Polypore fungi (Karsten 1881, Pegler and Young 1973, Moncalvo and Ryvarden 1997, Keypour et al. 2020). The membership of Ganoderma has been subsequently extended, including species with sessile, non-laccate basidiocarps and pigmented, ellipsoid to ovoid, ornamented, double-walled basidiospores (Murrill 1902, Pegler and Young 1973, Steyaert 1980, Cao and Yuan 2013, Papp 2016). Moncalvo and Ryvarden (1997) accepted 148 Ganoderma species before the molecular era, of which 65% are recognised as only one or some species, but represented different morphology-based species (Ryvarden 2000, Smith and Sivasithamparam 2003, Torres-Torres and Guzmán-Dávalos 2012). Recently, 180 species of Ganoderma were accepted, whereas nearly 500 species are estimated worldwide, of which 60% are awaiting discovery (He et al. 2019, He et al. 2022).

Despite their economic importance, the taxonomy of Ganoderma remains uncertain due to a slew of confusion and misconceptions. During the past several decades, many species of Ganoderma have been delimited, based on the presence of stipe, laccate or non-laccate, the context of pileus and the microscopic characteristics of basidiospores (Chang and Chen 1984, Seo and Kitamot 1998, Wu et al. 2004, Torres-Torres and Guzmán-Dávalos 2012, Zhou et al. 2016, Tchotet-Tchoumi et al. 2019). In general, it is difficult and subjective to identify Ganoderma species solely based on morphological evidence, as their phenotypic traits are sensitive to extrinsic factors, such as illumination, ventilation and humidity (Szedlay et al. 1999, Demoulin 2010, Yang and Feng 2013, Hapuarachchi et al. 2019a). Therefore, morphology-based identification brought Ganoderma into a state of taxonomic chaos (Smith and Sivasithamparam 2003, Coetzee et al. 2015, López-Peña et al. 2019, Náplavová et al. 2020). Compared to morphology, molecular methods have turned out to be more effective in resolving intraspecific relationships with Ganoderma (Yamashita and Hirose 2016, Fryssouli et al. 2020, Gunnels et al. 2020, Jiang et al. 2021, Shen et al. 2021). Phylogenetic markers, such as IGS, nrSSU, ITS, nrLSU, mtSSU, β-TUB, RPB1, RPB2 and TEF1-α sequences, were independently or conjointly used to infer intraspecific relationships within Ganoderma (Cao et al. 2012, Zhou et al. 2015, Xing et al. 2018, Hapuarachchi et al. 2019a, Liu et al. 2019, Ye et al. 2019). In particular, the multilocus phylogeny incorporating sequences from ITS, nrLSU, TEF1-α and RPB2 was applied to give a phylogenetic framework for species delimitation in this genus (Xing et al. 2018, Ye et al. 2019, Tchotet-Tchoumi et al. 2019, Wu et al. 2020, He et al. 2021, Cao et al. 2021). Furthermore, some researchers steered using a combination of morphological, chemotaxonomic and molecular strategies to elevate a steady taxonomy for Ganoderma and resolve taxonomic ambiguities (Richter et al. 2014, Welti et al. 2015).

Ganoderma has a cosmopolitan distribution and most of the species are known from tropical and sub-tropical regions (He et al. 2019). This fungus grows as saprobes or parasites on deciduous and coniferous trees and some of them are considered as plant pathogens that cause basal stem butt rot and root rot (Pinruan et al. 2010, Ding et al. 2020, Mafia et al. 2020, Mohd et al. 2020). Species of Ganoderma play an important role in the nutrient mobilisation process of woody plants. They possess lignocellulose–decomposing enzymes with effective mechanisms for bioenergy production and bioremediation (Coetzee et al. 2015, Kües et al. 2015). In the natural environment, a basidiome has the ability to produce innumerable basidiospores that can be spread by air- or rain-driven and insect vectors (Tuno 1999, Kadowaki et al. 2011, Almaguer et al. 2014, Sadyś et al. 2014). The infection of a plant host by pathogenic Ganoderma species starts with the landing of the basidiospore on the wound trunk or root, followed by germination and colonisation (Rees et al. 2009, Rees et al. 2012, Hushiarian et al. 2013, Ayin et al. 2019). Basal stem rot caused by G.boninense is the main disease that leads to yield losses and death of oil palm, which account for 50% of substantial economic losses to Southeast Asia’s palm oil industry (Hushiarian et al. 2013, Lee and Chang 2016, Midot et al. 2019). Red roots caused by G.philippii are a serious disease of commercial Acaciamangium in Malaysia and India (Glen et al. 2014). Since different Ganoderma species produce different characteristics and pathogenicity, species identification is difficult, which in turn, leads to significant difficulty in disease control (Wong et al. 2012).

Ganoderma was first reported from China by Teng (1934), with four species including G.lucidum and one variety. More than 80 species have been introduced so far and several extensive studies have been carried out to investigate Ganoderma diversity in China, with new species being introduced (Zhao and Zhang 2000, Wu et al. 2004, Dai et al. 2009, Cao et al. 2012, Hapuarachchi et al. 2015, Hapuarachchi et al. 2018c, Wu et al. 2019, Liu et al. 2019). However, the majority of Ganoderma species reported from China have not been subjected to systematic studies (Wang 2012, Hapuarachchi et al. 2016b, Hapuarachchi et al. 2018a, Wang et al. 2019). The objective of the present study is to introduce a novel Ganoderma species, from Guizhou Province, southwest China, with descriptions, colour photographs, illustrations and a multigene phylogeny.

Materials and methods

Ganoderma samples were collected from Sandong Township, Sandu Shuizu Autonomous County, Guizhou Province, China, during the rainy season of July 2020. They were dried and preserved as outlined in Hapuarachchi et al. (2019b). The materials used in this study were deposited at Guizhou University (GACP) and the Herbarium of Kunming Institute of Botany Academia Sinica (HKAS).

Morphological study

Macro-morphological characteristics were described, based on dried material and the photographs provided here. Colour codes (e.g. 8E8) are from Kornerup and Wanscher (1978). Pileus was sectioned with a razor blade and mounted in 5% potassium hydroxide (KOH) solution. Pileipellis, hyphal systems of pileus, basidia and basidiospores were observed and captured using a compound microscope (Leica DM2500) equipped with a camera. Images were measured with Leica Application Suite X (LAS X). In the description section, the number, length, width and length/width ratio of the measured basidiospores are denoted with symbols n, L, W and Q, respectively. The Faces of Fungi number was registered by following Jayasiri et al. (2015).

DNA Extraction, PCR and Sequencing

Genomic DNA was extracted from dried specimens using an HP Fungal DNA Kit (OMEGA, USA) following the protocol of the manufacturer. PCR amplification was performed in a final volume of 50 µl reaction mixture that contained 25 µl 2x BenchTopTM Taq Master Mix (Biomigas), 19 µl distilled water, 2 µl (10 µM) of each primer and 2 µl template DNA. The large subunit ribosomal RNA (LSU), the internal transcribed spacer (ITS), the translation elongation factor (TEF1-α) and the RNA polymerase II second largest subunit (RPB2) were amplified with primer pairs LROR/LR5 (Vilgalys and Hester 1990), ITS5/ITS4 (White et al. 1990), EF1-983F/EF1-1567R (Rehner and Buckley 2017) and RPB2-5f/RPB2-7cR (Liu et al. 1999). PCR amplification reactions were performed with a T100 Thermal Cycler (T100™, Bio-Rad, USA). The procedures used for amplification of ITS were as follows: initial denaturation at 95°C for 3 min, followed by 35 cycles of denaturation at 95°C for 30 s, annealing at 58°C for 30 s, elongation at 72°C for 1 min and a final extension at 72°C for 5 min. The cycling conditions of LSU, TEF1-α and RPB2 consisted of initial denaturation at 95°C for 3 min, followed by 35 cycles of denaturation at 95°C for 30 s, annealing at 56°C for 30 s, elongation at 72°C for 1.3 min and a final extension at 72°C for 10 min. PCR products were verified by 1% agarose gel electrophoresis and sent to Sangon Biotech (Shanghai, China) for purification and sequencing.

Sequence Alignment and Phylogenetic Analysis

The raw sequences generated in this study were assembled with ChromasPro (2.1.8). Megablast analysis was conducted using the assembled ITS and RPB2 sequences as the query to check the closely-related taxa. The taxa used in our phylogenetic analysis were selected, based on megablast results and related publications (Table 1). Alignments were performed using MAFFT v. 7 (http://mafft.cbrc.jp/alignment/server/index.html, Katoh and Standley 2013). The resulting alignments were improved manually when necessary, using BioEdit v. 7.0.5.2 (Hall 1999). The introns in TEF and RPB2 were removed, based on the published CDS sequence in GenBank. The aligned ITS1, 5.8S, ITS2, LSU, TEF1-α and RPB2 sequences were concatenated with SequenceMatrix v.1.7.8 (Vaidya et al. 2011). Maximum Likelihood (ML) analysis was performed using RAxMLHPC2 (Stamatakis 2014) on the CIPRES Science Gateway v. 3.3 (Miller et al. 2010). The phylogenetic tree was inferred from four gene-partition analyses, using the GTRCAT model with 25 categories, with settings that the number of bootstrap replicates to 1,000. PartitionFinder v.2 (Lanfear et al. 2017) was used to estimate the best-fit model of nucleotide evolution, with the dataset subdivided into 10 data partitions (TEF 1st codon positions, TEF 2nd codon positions and TEF 3rd codon positions; RPB2 1st codon positions, RPB2 2nd codon positions and RPB2 3rd codon positions; ITS1; 5.8S; ITS2; LSU) and the following settings: branch lengths = unlinked, models = all, model_selection = AICc and search = greedy. Bayesian Inference (BI) analysis was performed in the CIPRES Science Gateway using MrBayes on XSEDE v. 3.2.7a. The GTR+F+I+G4 (TEF 1st codon positions, TEF 2nd codon positions, RPB2 1st codon positions, RPB2 2nd codon positions, LSU and 5.8S), GTR+F+G4 (TEF 3rd codon positions), GTR+F+G4 (RPB2 2nd codon positions), SYM+G4 (ITS1 and ITS2) were selected as the best model. Two runs of four chains were run until the average standard deviation of split frequencies dropped below 0.01, which occurred after 2,360,000 generations. Tree was sampled every 1000th generation and the chain temperature was decreased to 0.05 to improve convergence. The convergence of the runs was checked using TRACER v.1.6 (Rambaut et al. 2013). The first 25% of the resulting samples were discarded as burn-in and posterior probabilities were calculated from the remaining sampled trees (Larget and Simon 1999). In both ML and BY analyses, Foraminisporaconcentrica (Cui 12644) and Foraminisporayinggelingensis (Cui 13618) were selected as the outgroup (Sun et al. 2020). ML bootstrap values and BY posterior probabilities greater than or equal to 70% and 0.95, respectively, were considered significant support. The phylogenetic tree was visualised with FigTree version 1.4.0 available at http://tree.bio.ed.ac.uk/software/figtree/ (Rambaut 2012).

Table 1.

The species, specimens and GenBank accession numbers of sequences used in this study

Species Voucher Geographic origin GenBank accession numbers References
ITS LSU EF-1 RPB2
G.adspersum SFC20141001-16 Korea KY364251 – KY393284 KY393270 Jargalmaa et al. (2017)
G.adspersum SFC20160115-20 Korea KY364254 – KY393286 KY393272 Jargalmaa et al. (2017)
G.angustisporum Cui 14578 China MG279171 – MG367564 – Xing et al. (2018)
G.angustisporum Cui 13817 (T) China MG279170 – MG367563 MG367507 Xing et al. (2018))
G.applanatum SFC20150930-02 Korea KY364258 – KY393288 KY393274 Jargalmaa et al. (2017)
G.applanatum Wei 5787a China KF495001 – KF494978 – GenBank
G.aridicola Dai 12588 (T) Africa KU572491 – KU572502 – Xing et al. (2016)
G.australe DHCR417 (HUEFS) Australia MF436676 MF436673 MF436678 – Costa-Rezende et al. (2017)
G.australe ZRL20151500 China LT716076 KY418900 KY419088 – Zhao et al. (2017)
G.boninense WD 2085 Japan KJ143906 – KJ143925 KJ143965 Zhou et al. (2015)
G.boninense WD 2028 Japan KJ143905 KU220015 KJ143924 – Zhou et al. (2015)
G.carnosum MUCL 49464 France MG706220 MG706168 MG837838 MG837793 GenBank
G.carnosum GC011ND Slovakia MK415266 MK995647 – Náplavová et al. (2020)
G.carocalcareum DMC 322 (T) Cameroon EU089969 – – – Douanla-Meli and Langer (2009)
G.casuarinicola Dai 16339 China MG279176 – MG367568 MG367511 Xing et al. (2018)
G.casuarinicola Dai 16336 (T) China MG279173 – MG367565 MG367508 Xing et al. (2018)
G.chalceum URM80457 Brazil JX310812 JX310826 – – GenBank
G.concinnum Robledo 3235 – MN077523 MN077557 – – Costa-Rezende et al. (2020)
G.concinnum Robledo 3192 – MN077522 MN077556 – – Costa-Rezende et al. (2020)
G.curtisii CBS 100131 USA JQ781848 – KJ143926 KJ143966 Zhou et al. (2015)
G.curtisii CBS 100132 USA JQ520164 – KJ143927 KJ143967 Zhou et al. (2015)
G.destructans CBS 139793 (T) South Africa NR_132919 NG_058157 – – Coetzee et al. (2015)
G.destructans Dai 16431 South Africa MG279177 – MG367569 MG367512 Xing et al. (2018)
G.dianzhongense L4331(T) China MW750237 – – MZ467043 He et al. (2021)
G.dianzhongense L4737 China MW750238 – – MW839000 He et al. (2021)
G.ecuadorense URM 89449 Brazil MK119828 MK119908 MK121577 MK121535 Sun et al. (2020)
G.ecuadorense URM 89441 Brazil MK119827 MK119907 MK121576 MK121534 Sun et al. (2020)
G.eickeri CMW50325 Africa MH571689 – MH567290 – Tchotet-Tchoumi et al. (2019)
G.eickeri CMW 49692 (T) Africa NR_165524 – – – Tchotet-Tchoumi et al. (2019)
G.ellipsoideum MFLU 19-2221 China MN398339 MN428664 MN423157 – GenBank
G.ellipsoideum GACP 14080966 (T) China NR_160617 – – – Hapuarachchi et al. (2018c)
G.enigmaticum Dai 15971 South Africa KU572487 – KU572497 MG367514 Xing et al. (2016)
G.enigmaticum Dai 15970 South Africa KU572486 – KU572496 MG367513 Xing et al. (2016)
G.esculentum L4935 (T) China MW750242 – – MW839004 He et al. (2021)
G.esculentum L4946 China MW750243 – – – He et al. (2021)
G.flexipes VT17102301 Viet Nam MK345430 MK346830 – – Hapuarachchi et al. (2019b)
G.flexipes Wei5491 China JQ781850 – – KJ143968 Cao et al. (2012)
G.gibbosum SFC20150918-08 Korea KY364271 – KY393291 KY393278 Jargalmaa et al. (2017)
G.gibbosum SFC20150918-03 Korea KY364270 – KY393290 KY393277 Jargalmaa et al. (2017)
G.hoehnelianum Dai 11995 China KU219988 KU220016 MG367550 MG367497 Xing et al. (2018)
G.hoehnelianum Cui 13982 China MG279178 – MG367570 MG367515 Xing et al. (2018)
G.knysnamense CMW 47755 (T) South Africa NR_165523 – MH567261 – Tchotet-Tchoumi et al. (2019)
G.knysnamense CMW49688 Africa MH571683 – MH567266 – Tchotet-Tchoumi et al. (2019)
G.leucocontextum Dai 15601 China KU572485 – KU572495 MG367516 Xing et al. (2016)
G.leucocontextum GDGM 40200 (T) China KM396272 – – – Li et al. (2015)
G.lingzhi Dai12574 (IFP) China KJ143908 – JX029977 JX029981 Zhou et al. (2015)
G.lingzhi Cui9166 China KJ143907 – JX029974 JX029978 Cao et al. (2012)
G.lobatum JV 1008/31 USA KF605671 – MG367553 MG367499 Xing et al. (2018)
G.lobatum JV 1008/32 USA KF605670 – MG367554 MG367500 Xing et al. (2018)
G.lucidum BR 4195 France KJ143909 – – KJ143969 Zhou et al. (2015)
G.lucidum K 175217 Italy KJ143911 – KJ143929 KJ143971 Zhou et al. (2015)
G.lucidum Cui 14405 China MG279182 – MG367574 MG367520 Xing et al. (2018)
G.lucidum CCBAS 707 Europe MG706231 MG706177 MG837846 MG837805 GenBank
G.martinicense UMNTN1 USA MG654178 – MG754738 MG754860 Loyd et al. (2018)
G.martinicense He 2240 USA MG279163 – MG367557 MG367503 Xing et al. (2018)
G.mbrekobenum UMN7-4 GHA Ghana KX000898 KX000899 – – Crous et al. (2016)
G.mbrekobenum UMN7-3 GHA Ghana KX000896 KX000897 – – Crous et al. (2016)
G.meredithiae UMNFL50 USA MG654103 – MG754735 MG754862 Loyd et al. (2018)
G.meredithiae CBS 271.88 (T) USA NR_164435 NG_067432 – – Vu et al. (2019)
G.meredithiae UMNFL64 USA MG654188 – MG754734 MG754861 Loyd et al. (2018)
G.mexicanum MUCL: 55832 Martinique MK531815 – MK531829 MK531839 Cabarroi-Hernández et al. (2019)
G.mexicanum MUCL: 49453 Martinique MK531811 – MK531825 MK531836 Cabarroi-Hernández et al. (2019)
G.mizoramense UMN-MZ5 India KY643751 – – – Crous et al. (2017)
G.mizoramense UMN-MZ4 (T) India KY643750 – – – Crous et al. (2017)
G.multipileum Cui 14373 China MG279184 – MG367575 MG367521 Xing et al. (2018)
G.multipileum Dai 9447 China KJ143914 – KJ143932 KJ143973 Zhou et al. (2015)
G.multiplicatum Dai 12320 China KU572490 – KU572500 – Xing et al. (2016)
G.multiplicatum Dai 13710 China KU572489 – KU572499 – Xing et al. (2016)
G.mutabile Yuan2289 China JN383977 – – – Cao and Yuan (2013)
G.mutabile CLZhao 982 China MG231527 – – – Cao and Yuan (2013)
G.nasalaense LPDR17060212 Laos MK345442 MK346832 – – Hapuarachchi et al. (2019b)
G.nasalaense GACP 17060211 (T) Laos NR_164048 NG_066439 – – Hapuarachchi et al. (2019b)
G.orbiforme Cui 13880 China MG279187 – MG367577 MG367523 Xing et al. (2018)
G.orbiforme Cui 13918 China MG279186 – MG367576 MG367522 Xing et al. (2018)
G.oregonense JV 0108/93 USA KF605620 – MG367558 MG367504 Xing et al. (2018)
G.oregonense CBS 265.88 USA JQ781875 – KJ143933 KJ143974 Zhou et al. (2015)
G.ovisporum HKAS123193 (T) China MZ519547 MZ519545 – MZ547661 This study
G.ovisporum GACP20071602 China MZ519548 MZ519546 – MZ547662 This study
G.perzonatum URM 89437 Brazil MK119830 – MK121579 – Sun et al. (2020)
G.perzonatum SP445990 Brazil KJ792750 – – – GenBank
G.pfeifferi LGAM 336-ACAM DD2118 Greece MG706232 MG706178 MG837847 MG837806 GenBank
G.pfeifferi Dai 12683 Greece MG279165 – MG367560 – Xing et al. (2018)
G.philippii Cui 14444 China MG279189 – MG367579 MG367525 Xing et al. (2018)
G.philippii MFLU 19-2223 Thailand MN401411 MN398327 MN423175 – GenBank
G.podocarpense QCAM6422 Ecuador MF796661 MF796660 – – GenBank
G.polychromum 330OR USA MG654196 – MG754742 – Loyd et al. (2018)
G.polychromum UMNOR3 USA MG654204 – MG754744 – Loyd et al. (2018)
G.ravenelii MS187FL USA MG654211 – MG754745 MG754865 Loyd et al. (2018)
G.ravenelii 150FL USA MG654207 – – – Loyd et al. (2018)
G.resinaceum MUCL: 38956 Netherlands MK554772 – MK554723 MK554747 Cabarroi-Hernández et al. (2019)
G.resinaceum MUCL: 52253 France MK554786 – MK554737 MK554764 Cabarroi-Hernández et al. (2019)
G.ryvardenii GanoTK41 Cameroon JN105699 – – – Kinge et al. (2012)
G.ryvardenii GanoTK43 Cameroon JN105695 – – – Kinge et al. (2012)
G.sandunense GACP18012502 China MK345451 – – MZ547664 Hapuarachchi et al. (2019b)
G.sandunense GACP18012501 (T) China NR_164049 – – MZ547663 Hapuarachchi et al. (2019b)
G.sessile 228DC USA MG654319 – MG754750 MG754869 Loyd et al. (2018)
G.sessile JV 1209/27 USA KF605630 – KJ143937 KJ143976 Zhou et al. (2015)
G.shandongense Dai 15791 China MG279192 – MG367582 MG367528 Xing et al. (2018)
G.shandongense Dai 15787 China MG279191 – MG367581 MG367527 Xing et al. (2018)
G.shanxiense HSA 539 China MK764269 – – MK789681 Liu et al. (2019)
G.shanxiense BJTC FM423 (T) China MK764268 – MK783937 MK783940 Liu et al. (2019)
G.sichuanense CGMCC 5.2175 (T) China NR_152892 – – KC662404 Yao et al. (2013)
G.sinense Cui 13835 China MG279193 – MG367583 MG367530 Xing et al. (2018)
G.sinense Wei 5327 China KF494998 KF495008 KF494976 MG367529 Xing et al. (2018)
G.steyaertanum 6-WN-16(M)-A Indonesia KJ654461 – – – Glen et al. (2014)
G.steyaertanum V-64-3 Indonesia KJ654433 – – – Glen et al. (2014)
G.stipitatum MUCL: 52655 French Guiana MK554770 – MK554717 MK554755 Cabarroi-Hernández et al. (2019)
G.stipitatum MUCL: 43863 Cuba MK554769 – MK554739 MK554745 Cabarroi-Hernández et al. (2019)
G.subamboinense UMNFL100 USA MG654373 – MG754762 – Loyd et al. (2018)
G.subamboinense SPC1 Brazil KU569546 KU570945 – – Bolaños et al. (2016))
G.tenue GTEN24-1 China DQ424977 – – – GenBank
G.tenue GTEN24-2 China DQ424978 – – – GenBank
G.thailandicum HKAS 104641a Thailand MK848682 MK849880 MK875830 MK875832 Luangharn et al. (2019a)
G.thailandicum HKAS 104640a (T) Thailand MK848681 MK849879 MK875829 MK875831 Luangharn et al. (2019a)
G.tropicum Dai 16434 China MG279194 – MG367585 MG367532 Xing et al. (2018)
G.tropicum KUMCC 18–0046a Thailand MH823539 – – MH883621 Luangharn et al. (2019b)
G.tsugae Cui 14112 China MG279196 – MG367587 MG367534 Xing et al. (2018)
G.tsugae Dai 12760 USA KJ143920 – KJ143940 KJ143978 Zhou et al. (2015)
G.tuberculosum UMNFL117 USA MG654359 – MG754771 – Loyd et al. (2018)
G.tuberculosum 233FL USA MG654367 – – MG754873 Loyd et al. (2018)
G.weberianum CBS 219.36 Philippines MH855780 MH867289 MK611974 MK611972 Cabarroi-Hernández et al. (2019)
G.weberianum CBS 128581 Taiwan MH864975 MH876427 MK636693 MK611971 Cabarroi-Hernández et al. (2019)
G.weixiensis HKAS 100649 (T) China NR_166271 NG_067863 MK302442 – Ye et al. (2019)
G.weixiensis HKAS100650 China MK302445 MK302447 MK302443 – Ye et al. (2019)
G.wiiroense UMN-21-GHA Ghana KT952363 KT952364 – – Crous et al. (2015))
G.wiiroense MIN 938704 (T) Ghana NR_158480 NG_064392 – – Crous et al. (2015)
G.williamsianum Dai 16809 Thailand MG279183 – MG367588 MG367535 Xing et al. (2018)
G.williamsianum Wei 5032 China KU219994 KU220024 – – Song et al. (2016)
G.zonatum FL-03 USA KJ143922 – KJ143942 KJ143980 Zhou et al. (2015)
G.zonatum FL-02 USA KJ143921 – KJ143941 KJ143979 Zhou et al. (2015)
Foraminisporaconcentrica Cui 12644 (T) China NR_158325 NG_064396 MK121561 MK121499 Sun et al. (2020)
F.yinggelingensis Cui 13618 (T) China NR_174805 MK119900 MK121570 MK121536 Sun et al. (2020)
Open in a new tab

Taxon treatments

Ganoderma ovisporum

H.D. Yang, T.C. Wen sp. nov.

A07C86F9-799E-59BF-AF73-120508E42B3F

IF558589

FoF 10099

Materials

  1. Type status: Holotype. Occurrence: recordedBy: Hongde Yang; occurrenceID: HKAS123193; Taxon: scientificName: Ganodermaovisporum; kingdom: Fungi; phylum: Basidiomycota; class: Agaricomycetes; order: Polyporales; family: Polyporaceae; genus: Ganoderma; Location: country: China; countryCode: CN; stateProvince: Guizhou; county: Sandu Shuizu Autonomous County; locality: Sandong Township; verbatimElevation: 612 m; verbatimLatitude: 25°70′ N; verbatimLongitude: 107°96′ E; Identification: identifiedBy: Hongde Yang; Event: year: 2020; month: July; day: 16; habitat: Terrestrial; fieldNotes: Rotten wood, in dry dipterocarp forest and in upper mixed deciduous forest and growing up from soil; Record Level: type: HKAS123193; collectionID: SD2020071601

Description

Basidiome annual, stipitate, corky, strongly laccate, becoming lighter when dry. Pileus 3 × 5 cm, up to 0.9 cm thick at the base, applanate, subreniform, upper surface red-brown (8E8) when fresh, becoming brownish-black (6C8) when dry, with slightly concentrically sulcate, radially rugose, irregularly tuberculate bumps and ridges overlying the context. Margin is slightly obtuse, yellow-brown (5D8) or concolorous with the pileus. Pore surface pale white (4A2). Pores nearly round to round, 3–4 per mm, dissepiments thin to slightly thick. Context up to 0.3 cm thick, corky, homogeneous at the periphery, becoming three-layered towards the centre, upper layer creamy-white (6E4), middle layer pale brown (6E4), lower layer brown (6D1), without concentric growth zone, black melanoid band absent. There is a line of independent or confluent, laterally arranged tubes inserted between the upper and middle layers of the context. Tubes up to 0.6 cm long, brownish (6E7). Stipe slightly darker than pileus, lateral, subcylindrical, 4-7 cm long, up to 1 cm in diam. Basidia not observed. Basidiospores (12.5–)13.0–13.5–15.0(–15.5) × (9.0–)9.5–10.0–10.5(–11.5) μm (Qm = 1.3, Q = 1.0–1.7，n = 30, with myxosporium), ellipsoid to broadly ellipsoid, ovoid, brown, double-walled, with a dark brown eusporium bearing coarse echinulae and an obtuse turgid appendix, overlaid by a hyaline, smooth myxosporium. Pileipellis hymeniodermiformic, yellowish-brown, terminal cells clavate, entire, brown (5D6), thick-walled, hollow, 18–29 × 6–11 μm. Hyphal system trimitic, generative hyphae 3.5–6 μm in diam., hyaline, colourless, thin-walled with clamp connections; skeletal hyphae 3–6 μm in diam., thick-walled to nearly solid, sometimes branched; binding hyphae 1.5–3 μm in diam., thick-walled, nearly solid, colourless (Fig. 1).

Figure 1.

Figure 1.

Open in a new tab

Ganodermaovisporum (HKAS123193, holotype). a–b Basidiome; c Pileus; d Pore surface; e Pores; f Sections of pileus; g–i Pileipellis terminal cell; j–m Basidiospores; n Skeletal hyphae; o Generative hyphae; p Binding hyphae. Scale bars: g = 50 µm; h–i = 30 µm; j–m = 10 µm; n = 100 µm; o = 10 µm; p = 100 µm.

Etymology

Referring to the ovoid basidiospores.

Notes

Ganodermaovisporum clusters with G.sandunense in the multigene phylogenetic tree (Fig. 3), the former is similar to the latter by having 98% and 97% homology in ITS and RPB2 sequence data, respectively. These two species are similar in having wide ovoid basidiospores and inhabiting deciduous coniferous mixed forests. However, G.ovisporum differs from G.sandunense in having inconspicuously concentric rings near the pileus margin, lateral stipe and shorter pileipellis terminal cells (18–29 × 6–11 μm), while conspicuously concentric zones and vertically-arranged ridges or grooves, central stipe and longer pileipellis terminal cells (50–95 × 8–13.5 μm) have been observed in the latter. By considering both phylogenetic evidence and morphological observations, we conclude our collection is a new species in Ganoderma.

Ganoderma sandunense

Hapuar., T.C. Wen & K.D. Hyde

EA035F7B-6ED1-584B-80C8-CC61BBA7305C

IF555784

FoF05659

Materials

  1. Type status: Holotype. Occurrence: recordedBy: Ting-Chi Wen; occurrenceID: GACP18012501; Taxon: scientificName: Ganodermasandunense; kingdom: Fungi; phylum: Basidiomycota; class: Agaricomycetes; order: Polyporales; family: Polyporaceae; genus: Ganoderma; Location: country: China; countryCode: CN; stateProvince: Guizhou; county: Sandu Shuizu Autonomous County; verbatimElevation: 590 m; verbatimLatitude: 24°54′N; verbatimLongitude: 107°53′E; Identification: identifiedBy: Kalani Hapuarachchi; Event: year: 2018; month: January; day: 25; habitat: Terrestrial; fieldNotes: Rotten wood, growing up from the soil; Record Level: type: GACP18012501; collectionID: GACP18012501

Description

Basidiome annual, stipitate, corky, strongly laccate. Pileus hemispherical, projecting 8 cm, up to 4 cm wide and 1.5 cm thick. Pileal surface reddish-black (8E8) to brownish-black (6C8), with distinctly concentrically sulcate, vertically-arranged ridges or grooves. Margin obtuse, concolorous with the pileus. Pore surface whitish-yellow (4A2) to light brown (6D4). Pores nearly circular, 3–5 per mm, dissepiments thin. Context up to 0.5 cm thick, inconspicuous triplex, fawn (5C5) to creamy-white (5A1) to dark brown (5E6), without concentric growth zone, black melanoid band absent. There is a line of independent or confluent, laterally-arranged tubes inserted between the upper and middle layer of the context. Tubes up to 1.2 cm long, dark brown (7F8). Stipe slightly darker than pileus, central, subcylindrical, up to 8 cm, 0.5 cm in diam. Basidia broadly ellipsoid, 21–25.5 × 13.5–17.5 μm, with four sterigmata. Basidiospores (12.3–)13.2–13.7–14.2(–15.7) × (9.0–)10–10.3–10.6(–12.5) μm (Qm = 1.3, Q = 1.0–1.7, n = 30, with myxosporium), ellipsoid to broadly ellipsoid, brown (7E5). Pileipellis cells clavate like, entire, brownish-orange (5C5), 50–95 × 8–13.5 μm. Hyphal system trimitic, generative hyphae 4-6 μm in diam., hyaline, colourless, thin-walled with clamp connections; skeletal hyphae 3.5–6 μm in diam., thick-walled to nearly solid, sometimes branched; binding hyphae 1-2 μm in diam., thick-walled, nearly solid, colourless (Fig. 2).

Figure 2.

Figure 2.

Open in a new tab

Ganodermasandunense (GACP18012501, holotype). a Basidiome; b Pore surface; c Sections of pileus; d–e Pileipellis terminal cell; f Basidia; g–j Basidiospores; k Skeletal hyphae and binding hyphae; l Generative hyphae; m Binding hyphae. Scale bars: d–e= 50 μm; f–h = 20 μm; i–j = 10 μm; k = 100 μm; l–m = 50 μm.

Notes

Ganodermasandunense was introduced by Hapuarachchi et al. (2019b) with ITS sequence. In addition, the description of its basidia is absent in their publication. In this study, the holotype of G.sandunense was loaned from Herbarium (GACP) and re-examined. We have refined this species with a more detailed illustration. Furthermore, we provided RPB2 sequence data of this species, which is an important phylogenetic marker used for intraspecific delimitation within Ganoderma.

Identification Keys

Keys to 22 species of laccate Ganoderma species in China

1 Distributed in China with gymnosperms as substrates G.tsugae
– Distributed in China with angiosperms as substrates 2
2 Basidiome sessile 3
– Basidiome stipitate to substipitate 5
3 Pileipellis terminal cells regular, clavate, occasionally with blunt outgrowth and protuberance, context present melanoid bands, basidiospores 8–12 × 3.8–5.2 µm G.angustisporum
– Pileipellis terminal cells are irregular, mainly composed of clavate cells or branched cells with blunt outgrowths in the lateral part or protuberances in the apical 4
4 Melanoid bands absent in the context, concentric growth zones present in the context, basidiospores 9.2–12 × 6.8–8.4 μm G.mutabile
– Melanoid bands present in the context, concentric growth zones absent in the context, basidiospores 8–13.5 × 4.2–6.3 μm G.boninense
5 Distributed in tropical regions 6
– Distributed mainly in temperate regions 8
6 Basidiome notably with a long, lateral stipe, pileus smaller, basidiospores with coarsely echinulate, 8.5–11 × 5–7 μm G.flexipes
– Basidiome stipitate to substipitate, pileus dimidiate, mostly large 7
7 Pileus single or occasionally composed of many small pilei, concentric growth zones present in the context, basidiospores with fine and long echinulate, 8–11.3 × 5–12.8 μm G.multipileum
– Pileus is mostly single, concentric growth zones absent in the context, basidiospores with coarse and short echinulae, 8.5–12.5 × 5.5–7.5 μm G.orbiforme
8 Context nearly homogeneous to homogeneous 9
– Context duplex to triplex 12
9 Pileus context white, pore surface white to cream, basidiospores 9.5–12.5 × 7–9 μm G.leucocontextum
– Pileus context brownish to brown or darker 10
10 Pileipellis terminal cells are mostly irregular, context present melanoid bands and concentric growth zones, basidiospores 10.8–13.1 × 8.3–11 μm G.tropicum
– Pileipellis terminal cells regular, cylindrical to clavate, context absent melanoid bands 11
11 Inhabiting deciduous forests, basidiospores ellipsoid, normally with an orderly arranged echinulae, basidiospores 10.7–12.8 × 7.0–9.0 μm G.sinense
– Inhabiting bamboo forests 12
12 Pileipellis terminal cells 35–65 × 8–16 μm, basidiospores 11–12.5 × 6.5–7.5 μm G.bambusicola
– Pileipellis terminal cells 20–55 × 10–15 μm, basidiospores 8.0–12.5 × 5.0–8.0 μm G.esculentum
13 Chlamydospores present in the context, basidiospores 7.8–10.4 × 5.2–6.4 μm G.weberianum
– Chlamydospores absent in context 14
14 Basidiospores < 8 μm in width and < 12 μm in length 15
– Basidiospores > 8 μm in width and > 9 μm in length 17
15 Basidiome corky, context soft, pores 2–4 per mm, pileipellis terminal cells regular, clavate, 20‒35 × 10‒12 μm, basidiospores 5.7‒8.3 × 2.6‒4.6 μm G.weixiensis
– Basidiome corky to woody, context firm, pores 4–6 per mm, pileipellis terminal cells occasional with outgrowths 16
16 Growing on living trees of Casuarinaequisetifolia, pileipellis terminal cells 40–70 × 5–13 µm, basidiospores 8.3–11.5 × 4.5–7 µm G.casuarinicola
– Growing on deciduous trees, pileipellis terminal cells 20–40 × 7–15 μm, basidiospores 7–9.3 × 4.6–6.8 μm G.lingzhi
17 Basidiospores ellipsoid, with sinuous ridge-like echinulae, 12.3–13.8 × 8.5–9.8 μm G.lucidum
– Basidiospores broadly ellipsoid, with coarse echinulae and an obtuse turgid appendix 18
18 Context brown to dark brown 19
– Context greyish-white to fawn brown 20
19 Pores 4–5 per mm, pileipellis terminal cells 25–30 × 7.5–8.5 μm, basidiospores 11.0–13.0 × 8.0–9.5 μm G.shanxiense
– Pores 5–8 per mm, pileipellis terminal cells 20–45 × 5.5–7.5 μm, basidiospores 9.0–12.5 × 6.5–9.0 μm G.dianzhongense
20 Distributed in Shandong Province, pileipellis terminal cells 17–25 × 4.5–7.5 μm, basidiospores 9‒13 × 6‒9 μm G.shandongense
– Distributed in Guizhou Province 21
21 Basidiome with a central stipe, pileipellis terminal cells 50–95 × 8–13.5 μm, basidiospores 12.3–15.7 × 9.1–12.0 μm G.sandunense
– Basidiome with a lateral stipe, pileipellis terminal cells 18–29 × 6–11 μm, basidiospores 12.5–15.5 × 9.0–11.5 μm G.ovisporum
Open in a new tab

Analysis

Phylogenetic analyses

Eight sequences of ITS, LSU and RPB2 were successfully amplified, but we failed to obtain the TEF1-α sequence from the two specimens HKAS123193 and GACP20071602. The newly-generated sequences and sequences from GenBank represented 132 specimens from 66 species, of which 21 were the type. The combined alignment of sequences comprised 3028 characters of 606, 1020, 809, 593 belonging to TEF1-α, RPB2, ITS and LSU, respectively. The final ML optimisation log-likelihood was -17354.28. The Bayesian Inference stopped at 2915000 generations when the average standard deviation of split frequencies reached 0.009904. The tree topologies derived from ML and BY were identical. Therefore, only the ML tree is shown (Fig. 3). The new species G.ovisporum and G.sandunense formed an individual clade in the phylogenetic tree (Fig. 3).

Figure 3.

Figure 3.

Open in a new tab

Phylogram for Ganoderma generated from Maximum Likelihood analysis of ITS, LSU, TEF1-α and RPB2 sequence data. Bootstrap support values for Maximum Likelihood and maximum parsimony greater than 70% and posterior probabilities of Bayesian Inference ≥ 0.95 are given above branches. Type specimens are marked with letter (T) and new species in this study are indicated in red.

Discussion

In this study, both phylogeny and morphology support G.ovisporum as a new species. Morphologically, it resembles other dark-coloured, laccate, stipitate Ganoderma species. However, it can be distinguished by having larger (12.5–15.5 × 9.0–11.5 μm), wide ovoid, dark brown-pigmented basidiospores. It is mostly similar to G.sandunense in having brownish-black pileus and similarly-sized basidiospores, as well distribution in Guizhou Province (Hapuarachchi et al. 2019b). The former species is distinct from the latter by having a lateral stipe and shorter pileipellis terminal cells (18–29 × 6–11 μm). Phylogenetically, G.ovisporum and G.sandunense are closely related, forming a distinct clade with basal position with strong support.

Ganoderma was extensively researched by the Chinese because it applied to medicine and food, together with the symbolic happiness and immortality culture, those being recognised as long as 2,000 years ago (Hapuarachchi et al. 2016b, Hapuarachchi et al. 2018b, Li et al. 2018, Cui et al. 2019, Du et al. 2021). Chinese taxonomists emphasised the morphological characteristics, such as stipe, pileus, pores, context, pileipellis terminal cells and basidiospores as keys to identity (Zhao 1989, He and Yu 1989, Zhang 1997). Keeping this method, Zhao and Zhang (2000) recorded 76 Ganoderma species from China, providing detailed illustrations. Wu and Dai (2005) identified 77 Ganoderma species with full description and colour photographs. Studies have been implemented to revise the taxonomy of Ganoderma in China by using molecular and morphology methods in the recent decade. The results indicated at least 23 species names are synonyms and confirmed that 24 species are distributed in China, 16 of which possess laccate basidiomes (Wang 2012, Chao 2013, Xing 2019). Since then, six species with laccate basidiomes have been described from China: G.bambusicola, G.dianzhongense, G.esculentum, G.sandunense, G.shanxiense and G.weixinense (Hapuarachchi et al. 2019b, Liu et al. 2019, Ye et al. 2019, Wu et al. 2020, He et al. 2021). Ganoderma taxonomy has undergone tremendous changes since both phenotypic features and phylogeny were used to delineate species (Gottlieb et al. 2000, Hapuarachchi et al. 2018c, Hapuarachchi et al. 2018a, Lin and Yang 2019, Tchotet-Tchoumi et al. 2019, He et al. 2022). Based on the aforementioned characteristics, we have provided a dichotomous key to 22 laccate species, including our new species from China.

Ganoderma could originate from Southeast Asia and later dispersal to the Northern Hemispheres, the Southern Hemispheres and the neotropics before 30 Mya years, during which species radiation and diversification events happened (Moncalvo and Buchanan 2008). Overviewing Ganoderma species worldwide, Imazeki (1939) concluded using subgenera Euganoderma and Elfvingia to accommodate species with laccate and non-laccate characters, respectively. In this study, a phylogenetic analysis was carried out using combined LSU, ITS, TEF1-α and RPB2 sequences from 66 species that included species previously placed in the above two subgenera. The topology of our phylogenetic tree is consistent with the morphology that the laccate species and non-laccate species tend to form groups. It is worth mentioning that the new species G.ovisporum group with the laccate species of G.carnosum, G.dianzhongense, G.leucocontextum, G.lucidum, G.oregonense, G.sandunense, G.shandongense, G.shanxiense, G.tsugae and G.weixiensis had strong support in both ML and Bayesian analyses. Those species were found in only or few ecological niches, except the widely cultivated G.leucocontextum, G.lucidum and G.tsugae (Gottlieb et al. 2000, Moncalvo and Buchanan 2008, Hapuarachchi et al. 2018b, Lin and Yang 2019). Therefore, many Ganoderma species are geographically restricted (He et al. 2022). However, the phylogenetic tree in the case of the laccate species G.pfeifferi and G.mutabile grouped with the non-laccate species G.adspersum, G.australe, G.eickeri, G.ellipsoideum, G.gibbosum, G.knysnamense, G.lobatum, G.podocarpense and G.williamsianum, indicating Euganoderma and Elfvingia are polyphyletic (Gottlieb et al. 2000). However, in fact, they are similar in having a substipitate to sessile basidiome and living as saprobes or parasites (Hapuarachchi et al. 2018c, Tchotet-Tchoumi et al. 2019). Consequently, biogeographic patterns and convergent evolution could explain the population structure and evolution of Ganoderma. Thus, a phylogeography study would help better understand the evolution of Ganoderma.

Supplementary Material

XML Treatment for Ganoderma ovisporum
XML Treatment for Ganoderma sandunense

Acknowledgements

This work was financed by the Science and Technology Foundation of Guizhou Province (KY [2018]039 and No. [2019]2451-3) and by the Open Fund Project of Key Laboratory of Forest Biotechnology in Yunnan, Southwest Forestry University, China (51700201). The authors are very grateful to Professor Xing-Liang Wu for his valuable comments and suggestions.

Contributor Information

Yong Ding, Email: dingyongSWFU@163.com.

Ting-Chi Wen, Email: tingchiwen@yahoo.com.

References

  1. Almaguer M., Rojas-Flores T. I., Rodríguez-Rajo F. J, Aira M. J. Airborne basidiospores of Coprinus and Ganoderma in a Caribbean region. Aerobiologia. 2014;30(2):197–204. doi: 10.1007/s10453-013-9318-y. [DOI] [Google Scholar]
  2. Ayin C. M., Alvarez A. M., Awana C., Schleinzer F. M., Marx B. C., Schlub R. L. Ralstoniasolanacearum, Ganodermaaustrale, and bacterial wetwood as predictors of ironwood tree (Casuarinaequisetifolia) decline in Guam. Australasian Plant Pathology. 2019;48(6):625–636. doi: 10.1007/s13313-019-00666-8. [DOI] [Google Scholar]
  3. Bolaños Ana. Cristina, Bononi Vera. Lúcia. Ramos, Gugliotta Adriana. De. Mello. New records of Ganodermamultiplicatum (Mont.) Pat. (Polyporales, Basidiomycota) from Colombia and its geographic distribution in South America. Check List. 2016;12(4):1–7. doi: 10.15560/12.4.1948. [DOI] [Google Scholar]
  4. Cabarroi-Hernández Milay, Villalobos-Arámbula Alma. Rosa, Torres-Torres Mabel. Gisela, Decock Cony, Guzmán-Dávalos Laura. The Ganodermaweberianum-resinaceum lineage: multilocus phylogenetic analysis and morphology confirm G.mexicanum and G.parvulum in the Neotropics. MycoKeys. 2019;59:95–131. doi: 10.3897/mycokeys.59.33182. [DOI] [PMC free article] [PubMed] [Google Scholar]
  5. Cao Bin, Haelewaters Danny, Schoutteten Nathan, Begerow Dominik, Boekhout Teun, Giachini Admir J., Gorjón Sergio P., Gunde-Cimerman Nina, Hyde Kevin D., Kemler Martin, Li Guo Jie, Liu Dong Mei, Liu Xin Zhan, Nuytinck Jorinde, Papp Viktor, Savchenko Anton, Savchenko Kyryll, Tedersoo Leho, Theelen Bart, Thines Marco, Tomšovský Michal, Toome-Heller Merje, Urón Judith P., Verbeken Annemieke, Vizzini Alfredo, Yurkov Andrey M., Zamora Juan Carlos, Zhao Rui Lin. Delimiting species in Basidiomycota: a review. Fungal Diversity. 2021;109(1):181–237. doi: 10.1007/s13225-021-00479-5. [DOI] [Google Scholar]
  6. Cao Yun, Wu Sheng. Hua, Dai Yu. Cheng. Species clarification of the prize medicinal Ganoderma mushroom “Lingzhi”. Fungal Diversity. 2012;56(1):49–62. doi: 10.1007/s13225-012-0178-5. [DOI] [Google Scholar]
  7. Cao Y., Yuan H. S. Ganodermamutabile sp. nov. from southwestern China based on morphological and molecular data. Mycological Progress. 2013;12(1):121–126. doi: 10.1007/s11557-012-0819-9. [DOI] [Google Scholar]
  8. Chang T. T., Chen T. Ganodermaformosanum sp. nov on formosan sweet gum in Taiwan. Transactions of the British Mycological Society. 1984;82(4):731–73. doi: 10.1016/S0007-1536(84)80119-9. [DOI] [Google Scholar]
  9. Chao Yun. Taxonomy and Phylogeny of Ganoderma in China. University of Chinese Academy of Sciences; Shengyang: 2013. 152. Chinese. [Google Scholar]
  10. Coetzee M. P.A., Marincowitz S., Muthelo V. G., Wingfield M. J. Ganoderma species, including new taxa associated with root rot of the iconic Jacarandamimosifolia in Pretoria, South Africa. IMA Fungus. 2015;6(1):249–256. doi: 10.5598/imafungus.2015.06.01.16. [DOI] [PMC free article] [PubMed] [Google Scholar]
  11. Costa-Rezende D. H., Robledo G. L., Góes-Neto A., Reck M. A., Crespo E., Drechsler-Santos E. R. Morphological reassessment and molecular phylogenetic analyses of Amauroderma s.lat. raised new perspectives in the generic classification of the Ganodermataceae family. Persoonia. 2017;39(1):254–269. doi: 10.3767/persoonia.2017.39.10. [DOI] [PMC free article] [PubMed] [Google Scholar]
  12. Costa-Rezende D. H., Robledo G. L., Drechsler-Santos E. R., Glen M., Gates G., Bonz B. R.M., Popof O. F., Crespo E., Góes-Neto A. Taxonomy and phylogeny of polypores with ganodermatoid basidiospores (Ganodermataceae) Mycological Progress. 2020;19(8):725–741. doi: 10.1007/s11557-020-01589-1. [DOI] [Google Scholar]
  13. Crous P. W., Wingfield M. J., Roux J. J. Le, Richardson D. M., Strasberg D., Shivas R. G., Alvarado P., Edwards J., Moreno G., Sharma R., Sonawane M. S., Tan Y. P., Altés A., Barasubiye T., Barnes C. W., Blanchette R. A., Boertmann D., Bogo A., Carlavilla J. R., Cheewangkoon R., Daniel R., de Beer Z. W., Yáñez-Morales M. de Jesús, Duong T. A., Fernández-Vicente J., Geering A. D.W., Guest D. I., Held B. W., Heykoop M., Hubka V., Ismail A. M., Kajale S. C., Khemmuk W., Kolařík M., Kurli R., Lebeuf R., Lévesque C. A., Lombard L., Magista D., Manjón J. L., Marincowitz S., Mohedano J. M., Nováková A., Oberlies N. H., Otto E. C., Paguigan N. D., Pascoe I. G., Pérez-Butrón J. L., Perrone G., Rahi P., Raja H. A., Rintoul T., Sanhueza R. M.V., Scarlett K., Shouche Y. S., Shuttleworth L. A., Taylor P. W.J., Thorn R. G., Vawdrey L. L., Solano-Vidal R., Voitk A., Wong P. T.W., Wood A. R., Zamora J. C., Groenewald J. Z. Fungal planet description sheets: 371–399. Persoonia. 2015;35(1):264–327. doi: 10.3767/003158515x690269. [DOI] [PMC free article] [PubMed] [Google Scholar]
  14. Crous P. W., Wingfield M. J., Richardson D. M., Leroux J. J., Strasberg D., Edwards J., Roets F., Hubka V., Taylor P. W.J., Heykoop M., Martín M. P., Moreno G., Sutton D. A., Wiederhold N. P., Barnes C. W., Carlavilla J. R., Gené J., Giraldo A., Guarnaccia V., Guarro J., Hernández-Restrepo M., Kolařík M., Manjón J. L., Pascoe I. G., Popov E. S., Sandoval-Denis M., Woudenberg J. H.C., Acharya K., Alexandrova A. V., Alvarado P., Barbosa R. N., Baseia I. G., Blanchette R. A., Boekhout T., Burgess T. I., Cano-Lira J. F., Čmoková A., Dimitrov R. A., Dyakov M. Yu., Dueñas M., Dutta A. K., Esteve-Raventós F., Fedosova A. G., Fournier J., Gamboa P., Gouliamova D. E., Grebenc T., Groenewald M., Hanse B., Hardy G. E.ST.J., Held B. W., Jurjević Ž, Kaewgrajang T., Latha K. P.D., Lombard L., Luangsa-ard J. J., Lysková P., Mallátová N., Manimohan P., Miller A. N., Mirabolfathy M., Morozova O. V., Obodai M., Oliveira N. T., Ordóñez M. E., Otto E. C., Paloi S., Peterson S. W., Phosri C., Roux J., Salazar W. A., Sánchez A., Sarria G. A., Shin H. -D., Silva B. D.B., Silva G. A., Smith M. TH., Souza-Motta C. M., Stchigel A. M., Stoilova-Disheva M. M., Sulzbacher M. A., Telleria M. T., Toapanta C., Traba J. M., Valenzuela-Lopez N., Watling R., Groenewald J. Z. Fungal planet description sheets: 400–468. Persoonia. 2016;36(1):316–458. doi: 10.3767/003158516x692185. [DOI] [PMC free article] [PubMed] [Google Scholar]
  15. Crous P. W., Wingfield M. J., Burgess T. I., Hardy G. E.ST.J., Barber P. A., Alvarado P., Barnes C. W., Buchanan P. K., Heykoop M., Moreno G., Thangavel R., Van der spuy S., Barili A., Barrett S., Cacciola S. O., Cano-Lira J. F., Crane C., Decock C., Gibertoni T. B., Guarro J., Guevara-Suarez M., Hubka V., Kolařík M., Lira C. R.S., Ordoñez M. E., Padamsee M., Ryvarden L., Soares A. M., Stchigel A. M., Sutton D. A., Vizzini A., Weir B. S., Acharya K., Aloi F., Baseia I. G., Blanchette R. A., Bordallo J. J., Bratek Z., Butler T., Cano-Canals J., Carlavilla J. R., Chander J., Cheewangkoon R., Cruz R. H.S.F., Da silva M., Dutta A. K., Ercole E., Escobio V., Esteve-Raventós F., Flores J. A., Gené J., Góis J. S., Haines L., Held B. W., Horta jung M., Hosaka K., Jung T., Jurjević Ž., Kautman V., Kautmanova I., Kiyashko A. A., Kozanek M., Kubátová A., Lafourcade M., La spada F., Latha K. P.D., Madrid H., Malysheva E. F., Manimohan P., Manjón J. L., Martín M. P., Mata M., Merényi Z., Morte A., Nagy I., Normand A. -C., Paloi S., Pattison N., Pawłowska J., Pereira O. L., Petterson M. E., Picillo B., Raj K. N.A. Fungal planet description sheets: 558–624. Persoonia. 2017;38(1):240–384. doi: 10.3767/003158517x698941. [DOI] [PMC free article] [PubMed] [Google Scholar]
  16. Cui B. K., Li H. J., Zhou J. L., Song J., Si J., Yang Z. L., Dai Y. C. Species diversity, taxonomy and phylogeny of Polyporaceae (Basidiomycota) in China. Fungal Diversity. 2019;97(1):137–392. doi: 10.1007/s13225-019-00427-4. [DOI] [Google Scholar]
  17. Dai Yu. Cheng, Yang Zhu. Liang, Cui Bao. Kai, Yu Chang. Jun, Zhou Li. Wei. Species diversity and utilization of medicinal mushrooms and fungi in China (Review) International Journal of Medicinal Mushrooms. 2009;11(3):287–302. doi: 10.1615/intjmedmushr.v11.i3.80. [DOI] [Google Scholar]
  18. Demoulin Vincent. Why conservation of the name Boletusapplanatus should be rejected. Taxon. 2010;59(1):283–286. doi: 10.1002/tax.591026. [DOI] [Google Scholar]
  19. Ding Shun. Ping, Hu Hong. Li, Gu Ji. Dong. Diversity, abundance, and distribution of wood-decay fungi in major parks of Hong Kong. Forests. 2020;11(10):1–16. doi: 10.3390/f11101030. [DOI] [Google Scholar]
  20. Douanla-Meli Clovis, Langer Ewald. Ganodermacarocalcareus sp. nov., with crumbly-friable context parasite to saprobe on Anthocleistanobilis and its phylogenetic relationship in G.resinaceum group. Mycological Progress. 2009;8(2):145–155. doi: 10.1007/s11557-009-0586-4. [DOI] [Google Scholar]
  21. Du Q., Cao Y., Liu C. In: The Lingzhi Mushroom Genome, Compendium of Plant Genomes. 1st. Du Q., editor. Springer, Cham; Beijing: 2021. Lingzhi, An Overview.1-25. [DOI] [Google Scholar]
  22. Fryssouli Vassiliki, Zervakis Georgios, Polemis Elias, Typas Milton A. A global meta-analysis of ITS rDNA sequences from material belonging to the genus Ganoderma (Basidiomycota, Polyporales) including new data from selected taxa. MycoKeys. 2020;75:71–143. doi: 10.3897/mycokeys.75.59872. [DOI] [PMC free article] [PubMed] [Google Scholar]
  23. Glen M., Yuskianti V., Puspitasari D., Francis A., Agustini L., Rimbawanto A., Indrayadi H., Gafur A., Mohammed C. L. Identification of Basidiomycete fungi in Indonesian hardwood plantations by DNA barcoding. Forest Pathology. 2014;44(6):496–508. doi: 10.1111/efp.12146. [DOI] [Google Scholar]
  24. Gottlieb Alexandra M., Ferrer Esther, Wright Jorge E. rDNA analyses as an aid to the taxonomy of species of Ganoderma. Mycological Research. 2000;104(9):1033–1045. doi: 10.1017/s095375620000304x. [DOI] [Google Scholar]
  25. Gunnels Tess, Creswell Matthew, McFerrin Janis, Whittall Justen B. The ITS region provides a reliable DNA barcode for identifying reishi/lingzhi (Ganoderma) from herbal supplements. PLoS One. 2020;15(11):e0236774. doi: 10.1371/journal.pone.0236774. [DOI] [PMC free article] [PubMed] [Google Scholar]
  26. Hall T. A. BioEdit: A user-friendly biological sequence alignment editor andanalysis program for Windows 95/98/NT. Nucleic Acids Symposium Series. 1999;41:95–98. [Google Scholar]
  27. Hapuarachchi K. K., Wen T. C., Deng C. Y., Kang J. C., Hyde K. D. Mycosphere essays 1: Taxonomic confusion in the Ganodermalucidum species complex. Mycosphere. 2015;6(5):542–559. doi: 10.5943/mycosphere/6/5/4. [DOI] [Google Scholar]
  28. Hapuarachchi K. K., Wen T. C., Jeewon R., Wu X. L., Kang J. C. Mycosphere essays 15: Ganodermalucidum - are the beneficial medical properties substantiated? Mycosphere. 2016;7:687–715. doi: 10.5943/mycosphere/7/6/1. [DOI] [Google Scholar]
  29. Hapuarachchi K. K., Wen T. C., Jeewon R., Wu X. L., Kang J. C., Hyde K. D. Mycosphere essays 7. Ganodermalucidum - are the beneficial anti-cancer properties substantiated? Mycosphere. 2016;7(3):305–332. doi: 10.5943/mycosphere/7/3/6. [DOI] [Google Scholar]
  30. Hapuarachchi K. K., Cheng C. R., Wen T. C., Jeewon R., Kakumyan P. Mycosphere essays 20: Therapeutic potential of Ganoderma species: Insights into its use as traditional medicine. Mycosphere. 2017;8(10):1653–1694. doi: 10.5943/mycosphere/8/10/5. [DOI] [Google Scholar]
  31. Hapuarachchi K. K., Karunarathna S. C., Phengsintham P., Kakumyan P., Hyde K. D., Wen T. C. Amauroderma (Ganodermataceae, Polyporales) - bioactive compounds, beneficial properties and two new records from Laos. Asian Journal of Mycology. 2018;1(1):121–136. doi: 10.5943/ajom/1/1/10. [DOI] [Google Scholar]
  32. Hapuarachchi K. K., Elkhateeb W. A., Karunarathna S. C., Cheng C. R., Bandara A. R., Kakumyan P., Hyde K. D., Daba G. M., Wen T. C. Current status of global Ganoderma cultivation, products, industry and market. Mycosphere. 2018;9(5):1025–1052. doi: 10.5943/mycosphere/9/5/6. [DOI] [Google Scholar]
  33. Hapuarachchi K. K., Karunarathna S. C., Rasp&eacute O., De Silva K. H.W.L., Thawthong A., Wu X. L., Kakumyan P., Hyde K. D., Wen T. C. High diversity of Ganoderma and Amauroderma (Ganodermataceae, Polyporales) in Hainan Island, China. Mycosphere. 2018;9(5):931–982. doi: 10.5943/mycosphere/9/5/1. [DOI] [Google Scholar]
  34. Hapuarachchi K. K., Karunarathna S. C., McKenzie E. H.C., Wu X. L., Kakumyan P., Hyde K. D., Wen T. C. High phenotypic plasticity of Ganodermasinense (Ganodermataceae, Polyporales) in China. Asian Journal of Mycology. 2019;2(1):1–47. doi: 10.5943/ajom/2/1/1. [DOI] [Google Scholar]
  35. Hapuarachchi K. K., Karunarathna S. C., Phengsintham P., Yang H. D., Kakumyan P., Hyde K. D., Wen T. C. Ganodermataceae (Polyporales): Diversity in Greater Mekong Subregion countries (China, Laos, Myanmar, Thailand and Vietnam) Mycosphere. 2019;10(1):221–309. doi: 10.5943/mycosphere/10/1/6. [DOI] [Google Scholar]
  36. He Jun, Luo Zong-Long, Tang Song-Ming, Li Yong-Jun, Li Shu-Hong, Su Hong-Yan. Phylogenetic analyses and morphological characters reveal two new species of Ganoderma from Yunnan province, China. MycoKeys. 2021;84:141–162. doi: 10.3897/mycokeys.84.69449. [DOI] [PMC free article] [PubMed] [Google Scholar]
  37. He Mao. Qiang, Zhao Rui. Lin, Hyde Kevin. D., Begerow Dominik, Kemler Martin, Yurkov Andrey, McKenzie Eric. H. C., Raspé Olivier, Kakishima Makoto, Sánchez-Ramírez Santiago, Vellinga Else. C., Halling Roy, Papp Viktor, Zmitrovich Ivan. V., Buyck Bart, Ertz Damien, Wijayawardene Nalin. N., Cui Bao. Kai, Schoutteten Nathan, Liu Xin. Zhan, Li Tai. Hui, Yao Yi. Jian, Zhu Xin. Yu, Liu An. Qi, Li Guo. Jie, Zhang Ming. Zhe, Ling Zhi. Lin, Cao Bin, Antonín Vladimír, Boekhout Teun, da Silva Bianca Denise Barbosa, De Crop Eske, Decock Cony, Dima Bálint, Dutta Arun. Kumar, Fell Jack. W., Geml József, Ghobad-Nejhad Masoomeh, Giachini Admir. J., Gibertoni Tatiana. B., Gorjón Sergio. P., Haelewaters Danny, He Shuang. Hui, Hodkinson Brendan. P., Horak Egon, Hoshino Tamotsu, Justo Alfredo, Lim Young. Woon, Menolli Nelson, Mešić Armin, Moncalvo Jean. Marc, Mueller Gregory. M., Nagy László. G., Nilsson R. Henrik, Noordeloos Machiel, Nuytinck Jorinde, Orihara Takamichi, Ratchadawan Cheewangkoon, Rajchenberg Mario, Silva-Filho Alexandre. G. S., Sulzbacher Marcelo. Aloisio, Tkalčec Zdenko, Valenzuela Ricardo, Verbeken Annemieke, Vizzini Alfredo, Wartchow Felipe, Wei Tie. Zheng, Weiß Michael, Zhao Chang. Lin, Kirk Paul. M. Notes, outline and divergence times of Basidiomycota. Fungal Diversity. 2019;99(1):105–367. doi: 10.1007/s13225-019-00435-4. [DOI] [Google Scholar]
  38. He Mao Qiang, Zhao Rui Lin, Liu Dong Mei, Denchev Teodor T., Begerow Dominik, Yurkov Andrey, Kemler Martin, Millanes Ana M., Wedin Mats, McTaggart A. R., Shivas Roger G., Buyck Bart, Chen Jie, Vizzini Alfredo, Papp Viktor, Zmitrovich Ivan V., Davoodian Naveed, Hyde Kevin D. Species diversity of Basidiomycota. Fungal Diversity. 2022 doi: 10.1007/s13225-021-00497-3. [DOI]
  39. He Shao Chang, Yu Hui Fang. The Family Ganodermataceae from Guizhou province of China. Acta Mycologica Sinca. 1989;8(4):282–288. [Google Scholar]
  40. Hushiarian Roozbeh, Yusof Nor Azah, Dutse Sabo Wada. Detection and control of Ganodermaboninense: strategies and perspectives. Springer Plus. 2013;2(1):1–12. doi: 10.1186/2193-1801-2-555. [DOI] [PMC free article] [PubMed] [Google Scholar]
  41. Imazeki R. Studies on Ganoderma of Nippon. Bulletin of the Tokyo Science Museum. 1939;1:29–52. [Google Scholar]
  42. Jargalmaa S., Eimes J. A., Park M. S., Park J. Y., Oh S. Y., Lim Y. W. Taxonomic evaluation of selected Ganoderma species and database sequence validation. PeerJ. 2017;5:e3596. doi: 10.7717/peerj.3596. [DOI] [PMC free article] [PubMed] [Google Scholar]
  43. Jayasiri Subashini. C., Hyde Kevin. D., Ariyawansa Hiran. A., Bhat Jayarama, Buyck Bart, Cai Lei, Dai Yu. Cheng, Abd-Elsalam Kamel. A., Ertz Damien, Hidayat Iman, Jeewon Rajesh, Jones E. B. Gareth, Bahkali Ali. H., Karunarathna Samantha. C., Liu Jian. Kui, Luangsa-ard J. Jennifer, Lumbsch H. Thorsten, Maharachchikumbura Sajeewa. S. N., McKenzie Eric. H. C., Moncalvo Jean. Marc, Ghobad-Nejhad Masoomeh, Nilsson Henrik, Pang Ka. Lai, Pereira Olinto. L., Phillips Alan J. L., Raspé Olivier, Rollins Adam. W., Romero Andrea. I., Etayo Javier, Selçuk Faruk, Stephenson Steven. L., Suetrong Satinee, Taylor Joanne. E., Tsui Clement K. M., Vizzini Alfredo, Abdel-Wahab Mohamed. A., Wen Ting. Chi, Boonmee Saranyaphat, Dai Dong. Qin, Daranagama Dinushani. A., Dissanayake Asha. J., Ekanayaka Anusha. H., Fryar S. C., Hongsanan Sinang, Jayawardena Ruvishika. S., Li Wen. Jing, Perera Rekhani. H., Phookamsak R., de Silva Nimali. I., Thambugala Kasun. M., Tian Qing, Wijayawardene Nalin. N., Zhao Rui. Lin, Zhao Qi, Kang Ji. Chuan, Promputtha Itthayakorn. The faces of fungi database: fungal names linked with morphology, phylogeny and human impacts. Fungal Diversity. 2015;74(1):3–18. doi: 10.1007/s13225-015-0351-8. [DOI] [Google Scholar]
  44. Jiang Nan, Hu Shuang, Peng Bing, Li Zhen. Hao, Yuan Xiao. Hui, Xiao Shi. Jun, Fu Yong. Ping. Genome of Ganoderma species provides insights into the evolution, conifers substrate utilization, and terpene synthesis for Ganodermatsugae. Frontiers in Microbiology. 2021;12:1–13. doi: 10.3389/fmicb.2021.724451. [DOI] [PMC free article] [PubMed] [Google Scholar]
  45. Justo Alfredo, Miettinen Otto, Floudas Dimitrios, Ortiz-Santana Beatriz, Sjökvist Elisabet, Lindner Daniel, Nakasone Karen, Niemelä Tuomo, Larsson Karl. Henrik, Ryvarden Leif, Hibbett David. S. A revised family-level classification of the Polyporales (Basidiomycota) Fungal Biology. 2017;121(9):798–824. doi: 10.1016/j.funbio.2017.05.010. [DOI] [PubMed] [Google Scholar]
  46. Kadowaki Kohmei, Leschen Richard. A.B., Beggs Jacqueline. R. No evidence for a Ganoderma spore dispersal mutualism in an obligate spore-feeding beetle Zearagytodesmaculifer. Fungal Biology. 2011;115(8):768–774. doi: 10.1016/j.funbio.2011.06.001. [DOI] [PubMed] [Google Scholar]
  47. Karsten P. A. Enumeralio boletinearum et polyporearum fennicarum, systemate novo dispositarum. Revue de Mycologie. 1881;3:16–19. [Google Scholar]
  48. Katoh K., Standley D. M. MAFFT multiple sequence alignment software version 7: Improvements in performance and usability. Molecular Biology and Evolution. 2013;30(4):772–780. doi: 10.1093/molbev/mst010. [DOI] [PMC free article] [PubMed] [Google Scholar]
  49. Keypour Somayeh, Riahi Hossein, Asef Mohammad. Reza, Abdollahzadeh Jafar, borhani Ali, Safaie Naser. The true nature of Ganoderma in Iran: Taxonomy based on ITS and mtSSU rDNA. Forest Pathology. 2020;50(4):e12605. doi: 10.1111/efp.12605. [DOI] [Google Scholar]
  50. Kinge T. R., Mih A. M., Coetzee M. P.A. Phylogenetic relationships among species of Ganoderma (Ganodermataceae, Basidiomycota) from Cameroon. Australian Journal of Botany. 2012;60(6):526–538. doi: 10.1071/BT12011. [DOI] [Google Scholar]
  51. Kornerup A, Wanscher J. H. Methuen Handbook of Colour. 3rd. Methuen; London: 1978. [Google Scholar]
  52. Kües Ursula, Nelson David R., Liu Chang, Yu Guo. Jun, Zhang Jian. Hui, Li Jian. Qin, Wang Xin. Cun, Sun Hui. Genome analysis of medicinal Ganoderma spp. with plant-pathogenic and saprotrophic life-styles. Phytochemistry. 2015;114:18–37. doi: 10.1016/j.phytochem.2014.11.019. [DOI] [PubMed] [Google Scholar]
  53. Lanfear R., Frandsen P. B., Wright A. M. PartitionFinder 2: New methods for selecting partitioned models of evolution for molecular and morphological phylogenetic analyses. Molecular Biology and Evolution. 2017;34:772–773. doi: 10.1093/molbev/msw260. [DOI] [PubMed] [Google Scholar]
  54. Larget B., Simon D. L. Markov chain Monte Carlo algorithms for the Bayesian analysis of phylogenetic trees. Molecular Biology and Evolution. 1999;16(6):750–759. doi: 10.1093/oxfordjournals.molbev.a026160. [DOI] [Google Scholar]
  55. Lee S. S., Chang Y. S. Ganoderma - Jekyll and Hyde mushrooms. Agriculture Science Journal. 2016;2:21–31. [Google Scholar]
  56. Leonardo-Silva Lucas, Cotrim Carlos Filipe Camilo, Xavier-Santos Solange. Furtadomyces nom. nov. (Ganodermataceae, Basidiomycota) with description of F. sumptuosus, a new species of ganodermatoid fungi from Brazil. Mycological Progress. 2022;21(3) doi: 10.1007/s11557-022-01794-0. [DOI] [Google Scholar]
  57. Li Li Feng, Liu Hong Bing, Zhang Quan Wei, Li Zhi Peng, Wong Tin Long, Fung Hau Yee, Zhang Ji Xia, Bai Su Ping, Lu Ai Ping, Han Quan Bin. Comprehensive comparison of polysaccharides from Ganodermalucidum and G.sinense: chemical, antitumor, immunomodulating and gut-microbiota modulatory properties. Scientific Reports. 2018;8(1):1–12. doi: 10.1038/s41598-018-22885-7. [DOI] [PMC free article] [PubMed] [Google Scholar]
  58. Lin Z. B, Yang B. X. Ganoderma and health: Biology, chemistry and industry. Volume 1181. Springer Nature Singapore; Beijing: 2019. [DOI] [Google Scholar]
  59. Li Tai. Hui, Hu Hui. Ping, Deng Wang. Qiu, Wu Sheng. Hua, Wang Dong. Mei, Tsering Tamdrin. Ganodermaleucocontextum, a new member of the G.lucidum complex from southwestern China. Mycoscience. 2015;56(1):81–85. doi: 10.1016/j.myc.2014.03.005. [DOI] [Google Scholar]
  60. Liu HONG, Guo LI. JIE, Li SU. LING, Fan LI. Ganodermashanxiense, a new species from northern China based on morphological and molecular evidence. Phytotaxa. 2019;406(2):129–136. doi: 10.11646/phytotaxa.406.2.4. [DOI] [Google Scholar]
  61. Liu Y. J., Whelen S., Hall B. D. Phylogenetic relationships among ascomycetes: evidence from an RNA polymerse II subunit. Molecular Biology and Evolution. 1999;16(12):1799–1808. doi: 10.1093/oxfordjournals.molbev.a026092. [DOI] [PubMed] [Google Scholar]
  62. López-Peña Damian, Samaniego-Rubiano Crystal, Morales-Estrada Idaly, Gutierrez Aldo, Gaitán-Hernández Rigoberto, Esqueda Martín. Morphological characteristics of wild and cultivated Ganodermasubincrustatum from Sonora, Mexico. Scientia Fungorum. 2019;49:e1213. doi: 10.33885/sf.2019.49.1213. [DOI] [Google Scholar]
  63. Loyd A. L., Barnes C. W., Held B. W., Schink M. J., Smith M. E., Smith J. A., Blanchette R. A. Elucidating "lucidum": Distinguishing the diverse laccate Ganoderma species of the United States. PLOS One. 2018;13(7):1–16. doi: 10.1371/journal.pone.0199738. [DOI] [PMC free article] [PubMed] [Google Scholar]
  64. Luangharn Thatsanee, Karunarathna Samantha. C., Mortimer Peter. E., Hyde Kevin. D., Xu Jian. Chu. Additions to the knowledge of Ganoderma in Thailand: Ganodermacasuarinicola, a new record; and Ganodermathailandicum sp. nov. MycoKeys. 2019;59:47–65. doi: 10.3897/mycokeys.59.36823. [DOI] [PMC free article] [PubMed] [Google Scholar]
  65. Luangharn T., Karunarathna S. C., Mortimer P. E., Hyde K. D., Thongklang N., Xu J. C. A new record of Ganodermatropicum (Basidiomycota, Polyporales) for Thailand and first assessment of optimum conditions for mycelia production. MycoKeys. 2019;51:65–83. doi: 10.3897/mycokeys.51.33513. [DOI] [PMC free article] [PubMed] [Google Scholar]
  66. Mafia M. I., Aminuzzaman F. M., Chowdhury Mohammad. Salahuddin. Mahmood, Tanni Jannatul. Ferdous. Occurrence, diversity and morphology of poroid wood decay by Ganoderma spp. from tropical moist deciduous forest region of Bangladesh. Journal of Agriculture and Natural Resources. 2020;3(2):160–174. doi: 10.3126/janr.v3i2.32498. [DOI] [Google Scholar]
  67. Midot F, Lau S. Y. L, Wong W. C, Tung H. J, Yap M. L, Lo M. L, Jee M. S, Dom S. P, Melling L. Genetic dversity and demographic history of Ganodermaboninense in oil palm plantations of Sarawak, Malaysia inferred from ITS regions. Microorganisms. 2019;7(10):1–17. doi: 10.3390/microorganisms7100464. [DOI] [PMC free article] [PubMed] [Google Scholar]
  68. Miller M. A., Pfeiffer W, Schwartz T. Creating the CIPRES Science Gateway for inference of large phylogenetic trees; Gateway Computing Environments Workshop 2010 (GCE); New Orleans, Louisiana. November; 2010. 1-8. [DOI] [Google Scholar]
  69. Mohd Shukri. I, Izzuddin M. A, Mohd Hefni. R, Idris A. S. Geostatistics of oil palm trees affected by Ganoderma disease in low and high planting density. IOP Conference Series: Earth and Environmental Science. 2020;540(1):1–9. doi: 10.1088/1755-1315/540/1/012065. [DOI] [Google Scholar]
  70. Moncalvo Jean-Marc, Buchanan Peter K. Molecular evidence for long distance dispersal across the Southern Hemisphere in the Ganoderma applanatum-australe species complex (Basidiomycota) Mycological Research. 2008;112(4):425–436. doi: 10.1016/j.mycres.2007.12.001. [DOI] [PubMed] [Google Scholar]
  71. Moncalvo J. M., Ryvarden L. A nomenclatural study of the Ganodermataceae Donk. Synopsis Fungorum. 1997;11:1–114. [Google Scholar]
  72. Murrill William Alphonso. The Polyporaceae of North America. I. The genus Ganoderma. Bulletin of the Torrey Botanical CLub. 1902;29(10):599–608. doi: 10.2307/2478682. [DOI] [Google Scholar]
  73. Náplavová Kateřina, Beck Terézia, Pristaš Peter, Gáperová Svetlana, Šebesta Martin, Piknová Mária, Gáper Ján. Molecular data reveal unrecognized diversity in the European Ganodermaresinaceum. Forests. 2020;11(8):850. doi: 10.3390/f11080850. [DOI] [Google Scholar]
  74. Papp V. The first validly published laccate Ganoderma species from East Asia: G.dimidiatum comb. nov., the correct name for G.japonicum. Studia Botanica Hungarica. 2016;47(2):263–268. doi: 10.17110/studbot.2016.47.2.263. [DOI] [Google Scholar]
  75. Pegler D. N., Young T. W.K. Basidiospore form in the British species of Ganoderma Karst. Kew Bulletin. 1973;28(3):351–364. doi: 10.2307/4108879. [DOI] [Google Scholar]
  76. Pinruan Umpava, Rungjindamai Nattawut, Choeyklin Rattaket, Lumyong Saisamorn, Hyde Kevin. D., Jones E. B. Gareth. Occurrence and diversity of basidiomycetous endophytes from the oil palm, Elaeisguineensis in Thailand. Fungal Diversity. 2010;41(1):71–88. doi: 10.1007/s13225-010-0029-1. [DOI] [Google Scholar]
  77. Rambaut A. FigTree version 1.4.0. http:// tree.bio.ed.ac.uk/software/fgtree/ 2012
  78. Rambaut A, Suchard M. A, Xie D, Drummond A. J. University of Edinburgh; 2013. Tracer v 1.6. [Google Scholar]
  79. Rees R. W., Flood J., Hasan Y., Potter U., Cooper R. M. Basal stem rot of oil palm (Elaeisguineensis); mode of root infection and lower stem invasion by Ganodermaboninense. Plant Pathology. 2009;58(5):982–989. doi: 10.1111/j.1365-3059.2009.02100.x. [DOI] [Google Scholar]
  80. Rees R. W., Flood J., Hasan Y., Wills M. A., Cooper R. M. Ganodermaboninense basidiospores in oil palm plantations: evaluation of their possible role in stem rots of Elaeisguineensis. Plant Pathology. 2012;61(3):567–578. doi: 10.1111/j.1365-3059.2011.02533.x. [DOI] [Google Scholar]
  81. Rehner Stephen A., Buckley Ellen. A Beauveria phylogeny inferred from nuclear ITS and EF1-α sequences: evidence for cryptic diversification and links to Cordycepsteleomorphs. Mycologia. 2017;97(1):84–98. doi: 10.1080/15572536.2006.11832842. [DOI] [PubMed] [Google Scholar]
  82. Richter Christian, Wittstein Kathrin, Kirk Paul. M., Stadler Marc. An assessment of the taxonomy and chemotaxonomy of Ganoderma. Fungal Diversity. 2014;71(1):1–15. doi: 10.1007/s13225-014-0313-6. [DOI] [Google Scholar]
  83. Ryvarden Leif. Studies in Neotropical Polypores 2: A Preliminary Key to Neotropical Species of Ganoderma with a Laccate Pileus. Mycologia. 2000;92(1):180–191. doi: 10.2307/3761462. [DOI] [Google Scholar]
  84. Sadyś M, Skjøth C. A, Kennedy R. Back-trajectories show export of airborne fungal spores (Ganoderma sp.) from forests to agricultural and urban areas in England. Atmospheric Environment. 2014;84:88–99. doi: 10.1016/j.atmosenv.2013.11.015. [DOI] [Google Scholar]
  85. Seo G. S., Kitamot Y. Morphological features and morphogenesis in the Ganodermalucidum complex. Japanese Society of Mushroom Science and Biotechnology. 1998;6:43–54. [Google Scholar]
  86. Shen Shan, Liu Shi. Liang, Jiang Ji. Hang, Zhou Li. Wei. Addressing widespread misidentifications of traditional medicinal mushrooms in Sanghuangporus (Basidiomycota) through ITS barcoding and designation of reference sequences. IMA fungus. 2021;12(10):1-20. doi: 10.1186/s43008-021-00059-x. [DOI] [PMC free article] [PubMed] [Google Scholar]
  87. Smith Brendan. J., Sivasithamparam K. Morphological studies of Ganoderma (Ganodermataceae) from the Australasian and Pacific regions. Australian Systematic Botany. 2003;16(4):487–503. doi: 10.1071/sb02001. [DOI] [Google Scholar]
  88. Song J., Xing J. H., Decock C., He X. L., Cui B. K. Molecular phylogeny and morphology reveal a new species of Amauroderma (Basidiomycota) from China. Phytotaxa. 2016;260(1):47–56. doi: 10.11646/phytotaxa.260.1.5. [DOI] [Google Scholar]
  89. Stamatakis A. RAxML version 8: a tool for phylogenetic analysis and post-analysis of large phylogenies. Bioinformatics. 2014;30:1312–1313. doi: 10.1093/bioinformatics/btu033. [DOI] [PMC free article] [PubMed] [Google Scholar]
  90. Steyaert R. L. Study of some Ganoderma species. http://www.jstor.org/stable/3667780. Bulletin du Jardin Botanique National de Belgique. 1980;50:135–186. doi: 10.2307/3667780. [DOI] [Google Scholar]
  91. Sun Y. F, Costa-Rezende D. H, Xing J. H, Zhou J. L, Zhang B, Gibertoni T. B, Gates G, Glen M, Dai Y. C, Cui B. K. Multi-gene phylogeny and taxonomy of Amauroderma s. lat. (Ganodermataceae) Persoonia. 2020;44:206–239. doi: 10.3767/persoonia.2020.44.08. [DOI] [PMC free article] [PubMed] [Google Scholar]
  92. Szedlay Z., Jakucs E., Boldizsar I., Boka A. Basidiocarp and mycelium morphology of Ganodermalucidum Karst. strains isolated in hungary. Acta Microbiologica et Immunologica Hungarica. 1999;46(1):41–52. doi: 10.1556/amicr.46.1999.1.5. [DOI] [PubMed] [Google Scholar]
  93. Tchotet-Tchoumi James Michel, Coetzee Martin Petrus Albertus, Rajchenberg Mario, Roux Jolanda. Taxonomy and species diversity of Ganoderma species in the Garden Route National Park of South Africa inferred from morphology and multilocus phylogenies. Mycologia. 2019;111(5):730–747. doi: 10.1080/00275514.2019.1635387. [DOI] [PubMed] [Google Scholar]
  94. Torres-Torres Mabel Gisela, Guzmán-Dávalos Laura. The morphology of Ganoderma species with a laccate surface. Mycotaxon. 2012;119(1):201–216. doi: 10.5248/119.201. [DOI] [Google Scholar]
  95. Tuno Nobuko. Insect feeding on spores of a bracket fungus, Elfvingiaapplanata (Pers.) Karst. (Ganodermataceae, Aphyllophorales) Ecological Research. 1999;14(2):97–103. doi: 10.1046/j.1440-1703.1999.00290.x. [DOI] [Google Scholar]
  96. Vaidya Gaurav, Lohman David. J., Meier Rudolf. SequenceMatrix: concatenation software for the fast assembly of multi-gene datasets with character set and codon information. Cladistics. 2011;27(2):171–180. doi: 10.1111/j.1096-0031.2010.00329.x. [DOI] [PubMed] [Google Scholar]
  97. Vilgalys R, Hester M. Rapid genetic identification and mapping of enzymatically amplified ribosomal DNA from several Cryptococcus species. Journal of Bacteriology. 1990;172(8):4238–4246. doi: 10.1128/jb.172.8.4238-4246.1990. [DOI] [PMC free article] [PubMed] [Google Scholar]
  98. Vu D., Groenewald M., de Vries M., Gehrmann T., Stielow B., Eberhardt U., Al-Hatmi A., Groenewald J. Z., Cardinali G., Houbraken J., Boekhout T., Crous P. W., Robert V., Verkley G. J.M. Large-scale generation and analysis of filamentous fungal DNA barcodes boosts coverage for kingdom fungi and reveals thresholds for fungal species and higher taxon delimitation. Studies in Mycology. 2019;92:135–154. doi: 10.1016/j.simyco.2018.05.001. [DOI] [PMC free article] [PubMed] [Google Scholar]
  99. Wang Guang. Yuan, Xu L. L, Yu Hao, Gao Jie, Guo Li. Zhong. Systematic analysis of the lysine succinylome in the model medicinal mushroom Ganodermalucidum. BMC Genomics. 2019;20(1):1–12. doi: 10.1186/s12864-019-5962-0. [DOI] [PMC free article] [PubMed] [Google Scholar]
  100. Wang Xing Cun. Phylogenetic Study on Ganodermataceae Donk. University of Chinese Academy of Sciences; Beijing: 2012. 291. Chinese. [Google Scholar]
  101. Welti Stéphane, Moreau Pierre. Arthur, Decock Cony, Danel Cécile, Duhal Nathalie, Favel Anne, Courtecuisse Régis. Oxygenated lanostane-type triterpenes profiling in laccate Ganoderma chemotaxonomy. Mycological Progress. 2015;14(7):1–15. doi: 10.1007/s11557-015-1066-7. [DOI] [Google Scholar]
  102. White T. J., Bruns T., Lee S., Taylor J. In: PCR Protocols: a guide to methods and applications. Innis M. A., Gelfand D. H., Sninsky J. J., White T. J., editors. Academic Press; 1990. Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics PCR protocols. [DOI] [Google Scholar]
  103. Wong C. L., Bong J. F.C., Idris A. S. Ganoderma species associated with basal stem rot disease of oil palm. American Journal of Applied Sciences. 2012;9(6):879–885. doi: 10.3844/ajassp.2012.879.885. [DOI] [Google Scholar]
  104. Wu Fang, Zhou Li. Wei, Yang Zhu. Liang, Bau Tolgor, Li Tai. Hui, Dai Yu. Cheng. Resource diversity of Chinese macrofungi: edible, medicinal and poisonous species. Fungal Diversity. 2019;98(1):1–76. doi: 10.1007/s13225-019-00432-7. [DOI] [Google Scholar]
  105. Wu SHENG. HUA, Chern CHI. LIANG, Wei CHIA. LING, Chen YU. PING, Akiba MITSUTERU, Hattori TSUTOMU. Ganodermabambusicola sp. nov. (Polyporales, Basidiomycota) from southern Asia. Phytotaxa. 2020;456(1):75–85. doi: 10.11646/phytotaxa.456.1.5. [DOI] [Google Scholar]
  106. Wu X. L., Dai Y. C., Lin Y. H. Study on the Ganodermataceae of China Ⅲ. Guizhou Sciense. 2004;22(4):36–40. [Google Scholar]
  107. Wu X. L., Dai Y. C. Coloured illustrations of Ganodermataceae of China. 1st. Science Press; Beijing: 2005. 228. Chinese. [Google Scholar]
  108. Xing JIA. HUI, Song JIE, Decock CONY, Cui BAO. KAI. Morphological characters and phylogenetic analysis reveal a new species within the Ganodermalucidum complex from South Africa. Phytotaxa. 2016;266(2):115–124. doi: 10.11646/phytotaxa.266.2.5. [DOI] [Google Scholar]
  109. Xing Jia. Hui, Sun Yi. Fei, Han Yu. Li, Cui Bao. Kai, Dai Yu. Cheng. Morphological and molecular identification of two new Ganoderma species on Casuarinaequisetifolia from China. MycoKeys. 2018;34:93–108. doi: 10.3897/mycokeys.34.22593. [DOI] [PMC free article] [PubMed] [Google Scholar]
  110. Xing J. H. Species diversity, taxonomy and phylogeny of Ganoderma. Beijing Forestry University; Beijing: 2019. 1-160. Chinese. [DOI] [Google Scholar]
  111. Yamashita Satoshi, Hirose Dai. Phylogenetic analysis of Ganodermaaustrale complex in a Bornean tropical rainforest and implications for mechanism of coexistence of various phylogenetic types. Fungal Ecology. 2016;24:1–6. doi: 10.1016/j.funeco.2016.04.006. [DOI] [Google Scholar]
  112. Yang Z. L., Feng B. What is the Chinese “Lingzhi”? - a taxonomic mini-review. Mycology. 2013;4(1):1–4. doi: 10.1080/21501203.2013.774299. [DOI] [Google Scholar]
  113. Yao Y. J., Wang X. C., Wang B. Epitypification of Ganodermasichuanense J.D. Zhao & X.Q. Zhang (Ganodermataceae) Taxon. 2013;62(5):1025–1031. doi: 10.12705/625.10. [DOI] [Google Scholar]
  114. Ye L., Karunarathna S. C., Mortimer P. E., Li H., Qiu M. H., Peng X. R., Luangharn T., Li Y. J., I Prom-, Hyde K. D., Xu J. C. Ganodermaweixiensis (Polyporaceae, Basidiomycota), a new member of the G.lucidum complex from Yunnan Province, China. Phytotaxa. 2019;423(2):75–86. doi: 10.11646/phytotaxa.423.2.3. [DOI] [Google Scholar]
  115. Zhang Xiao Qing. Four new records of Ganodermataceae from China. Mycosystema. 1997;16(4):259–26. [Google Scholar]
  116. Zhao J. D. The Ganodermataceae in China. Bibliotheca Mycologica. 1989;132:1–176. [Google Scholar]
  117. Zhao J. D, Zhang X. Q. Flora fungorum cinicorum. 1st. Volume 18, Ganodermataceae. Science Press; Beijing: 2000. 1-204. Chinese. [Google Scholar]
  118. Zhao Rui. Lin, Li Guo. Jie, Sánchez-Ramírez Santiago, Stata Matt, Yang Zhu. Liang, Wu Gang, Dai Yu. Cheng, He Shuang. Hui, Cui Bao. Kai, Zhou Jun. Liang, Wu Fang, He Mao. Qiang, Moncalvo Jean. Marc, Hyde Kevin. D. A six-gene phylogenetic overview of Basidiomycota and allied phyla with estimated divergence times of higher taxa and a phyloproteomics perspective. Fungal Diversity. 2017;84(1):43–74. doi: 10.1007/s13225-017-0381-5. [DOI] [Google Scholar]
  119. Zhou Li. Wei, Cao Yun, Wu Sheng. Hua, Vlasák Josef, Li De. Wei, Li Meng. Jie, Dai Yu. Cheng. Global diversity of the Ganodermalucidum complex (Ganodermataceae, Polyporales) inferred from morphology and multilocus phylogeny. Phytochemistry. 2015;114:7–15. doi: 10.1016/j.phytochem.2014.09.023. [DOI] [PubMed] [Google Scholar]
  120. Zhou Xuan. Wei, Liu Yan, Guo Meng. Yuan, Su Ka. Qi, Zhang Yong. Ming. Species clarification of the widely cultivated Ganoderma in China based on rDNA and FIP gene sequence analysis. International Journal of Agriculture and Biology. 2016;18(5):932–938. doi: 10.17957/ijab/15.0189. [DOI] [Google Scholar]

Associated Data

This section collects any data citations, data availability statements, or supplementary materials included in this article.

Supplementary Materials

XML Treatment for Ganoderma ovisporum
BDJ.10.e80034-treatment1.xml (12.6KB, xml)
XML Treatment for Ganoderma sandunense
BDJ.10.e80034-treatment2.xml (12.4KB, xml)

Articles from Biodiversity Data Journal are provided here courtesy of Pensoft Publishers

RESOURCES