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Species diversity, systematic revision and molecular phylogeny of Ganodermataceae (Polyporales, Basidiomycota) with an emphasis on Chinese collections

May 2022
Studies in Mycology 101(1):287-415

May 2022
101(1):287-415

DOI:10.3114/sim.2022.101.05

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CC BY-NC-ND 3.0

Authors:

Yi-Fei Sun

Beijing Forestry University

Jia-Hui Xing

Guangdong Institute of Microbiology

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Ganodermataceae is one of the main families of macrofungi since species in the family are both ecologically and economically important. The double-walled basidiospores with ornamented endospore walls are the characteristic features of Ganodermataceae . It is a large and complex family; although many studies have focused on Ganodermataceae , the global diversity, geographic distribution, taxonomy and molecular phylogeny of Ganodermataceae still remained incompletely understood. In this work, taxonomic and phylogenetic studies on worldwide species of Ganodermataceae were carried out by morphological examination and molecular phylogenetic analyses inferred from six gene loci including the internal transcribed spacer regions (ITS), the large subunit of nuclear ribosomal RNA gene (nLSU), the second largest subunit of RNA polymerase II gene ( rpb2 ), the translation elongation factor 1-α gene ( tef1 ), the small subunit mitochondrial rRNA gene (mtSSU) and the small subunit nuclear ribosomal RNA gene (nSSU). A total of 1 382 sequences were used in the phylogenetic analyses, of which 817 were newly generated, including 132 sequences of ITS, 139 sequences of nLSU, 83 sequences of rpb2 , 124 sequences of tef1 , 150 sequences of mtSSU and 189 sequences of nSSU. The combined six-gene dataset included sequences from 391 specimens representing 146 taxa from Ganodermataceae . Based on morphological and phylogenetic analyses, 14 genera were confirmed in Ganodermataceae : Amauroderma , Amaurodermellus , Cristataspora , Foraminispora , Furtadoella , Ganoderma , Haddowia , Humphreya , Magoderna , Neoganoderma , Sanguinoderma , Sinoganoderma , Tomophagus and Trachydermella . Among these genera, Neoganoderma gen. nov. is proposed for Ganoderma neurosporum ; Sinoganoderma gen. nov. is proposed for Ganoderma shandongense ; Furtadoella gen. nov. is proposed to include taxa previously belonging to Furtadoa since Furtadoa is a homonym of a plant genus in the Araceae ; Trachydermella gen. nov. is proposed to include Trachyderma tsunodae since Trachyderma is a homonym of a lichen genus in the Pannariaceae . Twenty-three new species, viz., Ganoderma acaciicola , G. acontextum , G. alpinum , G. bubalinomarginatum , G. castaneum , G. chuxiongense , G. cocoicola , G. fallax , G. guangxiense , G. puerense , G. subangustisporum , G. subellipsoideum , G. subflexipes , G. sublobatum , G. tongshanense , G. yunlingense , Haddowia macropora , Sanguinoderma guangdongense , Sa. infundibulare , Sa. longistipitum , Sa. melanocarpum , Sa. microsporum and Sa. tricolor are described. In addition, another 33 known species are also described in detail for comparison. Scanning electron micrographs of basidiospores of 10 genera in Ganodermataceae are provided. A key to the accepted genera of Ganodermataceae and keys to the accepted species of Ganoderma , Haddowia , Humphreya , Magoderna , Sanguinoderma and Tomophagus are also provided. In total, 278 species are accepted as members of Ganodermataceae including 59 species distributed in China.

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287

Studies in Mycology

Species diversity, systematic revision and molecular phylogeny of Ganodermataceae

(Polyporales, Basidiomycota) with an emphasis on Chinese collections

Y.-F. Sun1#, J.-H. Xing1#, X.-L. He2, D.-M. Wu3, C.-G. Song1, S. Liu1, J. Vlasák4, G. Gates5, T.B. Gibertoni6, B.-K. Cui1*

1Institute of Microbiology, School of Ecology and Nature Conservation, Beijing Forestry University, Beijing 100083, China; 2Sichuan Institute of Edible Fungi,

Sichuan Academy of Agricultural Sciences, Chengdu, Sichuan 610066, China; 3Biotechnology Research Institute, Xinjiang Academy of Agricultural and

Reclamation Sciences / Xinjiang Production and Construction Group Key Laboratory of Crop Germplasm Enhancement and Gene Resources Utilization,

Shihezi, Xinjiang 832000, China; 4Biology Centre, Czech Academy of Sciences, Institute of Plant “Molecular Biology, Branišovská 31, CZ-370 05 České

Budějovice, Czech Republic; 5Tasmanian Institute of Agriculture, Private Bag 98, Hobart, Tasmania 7001, Australia; 6Universidade Federal de Pernambuco,

Centro de Biociências, Departamento de Micologia, Av. Avenida da Engenharia, s/n, CEP 50740-600, Recife, Pernambuco, Brasil

#These authors contributed equally to the work

*Corresponding author: Bao-Kai Cui, cuibaokai@bjfu.edu.cn, baokaicui2013@gmail.com

Abstract: Ganodermataceae is one of the main families of macrofungi since species in the family are both ecologically and economically important. The double-

walled basidiospores with ornamented endospore walls are the characteristic features of Ganodermataceae. It is a large and complex family; although many

studies have focused on Ganodermataceae, the global diversity, geographic distribution, taxonomy and molecular phylogeny of Ganodermataceae still remained

incompletely understood. In this work, taxonomic and phylogenetic studies on worldwide species of Ganodermataceae were carried out by morphological

examination and molecular phylogenetic analyses inferred from six gene loci including the internal transcribed spacer regions (ITS), the large subunit of nuclear

ribosomal RNA gene (nLSU), the second largest subunit of RNA polymerase II gene (rpb2), the translation elongation factor 1-α gene (tef1), the small subunit

mitochondrial rRNA gene (mtSSU) and the small subunit nuclear ribosomal RNA gene (nSSU). A total of 1 382 sequences were used in the phylogenetic

analyses, of which 817 were newly generated, including 132 sequences of ITS, 139 sequences of nLSU, 83 sequences of rpb2, 124 sequences of tef1, 150

sequences of mtSSU and 189 sequences of nSSU. The combined six-gene dataset included sequences from 391 specimens representing 146 taxa from

Ganodermataceae. Based on morphological and phylogenetic analyses, 14 genera were conrmed in Ganodermataceae: Amauroderma, Amaurodermellus,

Cristataspora, Foraminispora, Furtadoella, Ganoderma, Haddowia, Humphreya, Magoderna, Neoganoderma, Sanguinoderma, Sinoganoderma, Tomophagus and

Trachydermella. Among these genera, Neoganoderma gen. nov. is proposed for Ganoderma neurosporum; Sinoganoderma gen. nov. is proposed for Ganoderma

shandongense; Furtadoella gen. nov. is proposed to include taxa previously belonging to Furtadoa since Furtadoa is a homonym of a plant genus in the Araceae;

Trachydermella gen. nov. is proposed to include Trachyderma tsunodae since Trachyderma is a homonym of a lichen genus in the Pannariaceae. Twenty-three

new species, viz., Ganoderma acaciicola, G. acontextum, G. alpinum, G. bubalinomarginatum, G. castaneum, G. chuxiongense, G. cocoicola, G. fallax, G.

guangxiense, G. puerense, G. subangustisporum, G. subellipsoideum, G. subexipes, G. sublobatum, G. tongshanense, G. yunlingense, Haddowia macropora,

Sanguinoderma guangdongense, Sa. infundibulare, Sa. longistipitum, Sa. melanocarpum, Sa. microsporum and Sa. tricolor are described. In addition, another

33 known species are also described in detail for comparison. Scanning electron micrographs of basidiospores of 10 genera in Ganodermataceae are provided.

A key to the accepted genera of Ganodermataceae and keys to the accepted species of Ganoderma, Haddowia, Humphreya, Magoderna, Sanguinoderma and

Tomophagus are also provided. In total, 278 species are accepted as members of Ganodermataceae including 59 species distributed in China.

Key words: Ganoderma, macro fungi, medicinal mushrooms, new taxa, phylogeny, ultrastructure, white-rot fungi.

Taxonomic novelties: New genera: Furtadoella B.K. Cui & Y.F. Sun, Neoganoderma B.K. Cui & Y.F. Sun, Sinoganoderma B.K. Cui, J.H. Xing & Y.F. Sun

and Trachydermella B.K. Cui & Y.F. Sun; New species: Ganoderma acaciicola B.K. Cui, J.H. Xing & Y.F. Sun, G. acontextum B.K. Cui, J.H. Xing & Vlasák, G.

alpinum B.K. Cui, J.H. Xing & Y.F. Sun, G. bubalinomarginatum B.K. Cui, J.H. Xing & Y.F. Sun, G. castaneum B.K. Cui, J.H. Xing & Y.F. Sun, G. chuxiongense

B.K. Cui, J.H. Xing & Y.F. Sun, G. cocoicola B.K. Cui, J.H. Xing & Y.F. Sun, G. fallax B.K. Cui, J.H. Xing & Vlasák, G. guangxiense B.K. Cui, J.H. Xing & Y.F.

Sun, G. puerense B.K. Cui, J.H. Xing & Y.F. Sun, G. subangustisporum B.K. Cui, J.H. Xing & Y.F. Sun, G. subellipsoideum B.K. Cui, J.H. Xing & Y.F. Sun, G.

subexipes B.K. Cui, J.H. Xing & Y.F. Sun, G. sublobatum B.K. Cui, J.H. Xing & Y.F. Sun, G. tongshanense B.K. Cui, J.H. Xing & Y.F. Sun, G. yunlingense

B.K. Cui, J.H. Xing & Y.F. Sun, Haddowia macropora B.K. Cui, Vlasák & Y.F. Sun, Sanguinoderma guangdongense B.K. Cui & Y.F. Sun, Sa. infundibulare B.K.

Cui & Y.F. Sun, Sa. longistipitum B.K. Cui & Y.F. Sun, Sa. melanocarpum B.K. Cui & Y.F. Sun, Sa. microsporum B.K. Cui & Y.F. Sun and Sa. tricolor B.K. Cui

& Y.F. Sun; New combinations: Furtadoella biseptata (Costa-Rezende et al.) B.K. Cui & Y.F. Sun, Fu. brasiliensis (Singer) B.K. Cui & Y.F. Sun, Fu. corneri

(Gulaid & Ryvarden) B.K. Cui & Y.F. Sun, Neoganoderma neurosporum (J.S. Furtado) B.K. Cui & Y.F. Sun, Sinoganoderma shandongense (J.D. Zhao & L.W.

Xu) B.K. Cui, J.H. Xing & Y.F. Sun and Trachydermella tsunodae (Yasuda ex Lloyd) B.K. Cui & Y.F. Sun.

available online at www.studiesinmycology.org StudieS in Mycology 101: 287–415 (2022).

Citation: Sun Y-F, Xing J-H, He X-L, Wu D-M, Song C-G, Liu S, Vlasák J, Gates G, Gibertoni TB, Cui B-K (2022). Species diversity, systematic revision and

molecular phylogeny of Ganodermataceae (Polyporales, Basidiomycota) with an emphasis on Chinese collections. Studies in Mycology 101: 287–415. doi:

10.3114/sim.2022.101.05.

Received: 13 December 2021; Accepted: 6 April 2022; Effectively published online: 20 May 2022

Corresponding editor: Robert A. Samson

INTRODUCTION

Ganodermataceae as an important family in the Polyporales has

been researched for many decades due to its high medicinal and

ecological values. As traditional medicine, Ganoderma lingzhi,

G. sinense and Amauroderma rugosum have been used for anti-

cancer treatment, for lowering blood pressure and for improving

immunity (Dai et al. 2009, Cao et al. 2012, Chan et al. 2013,

Zhou et al. 2015, Zhang et al. 2019). Tree pathogens such as G.

boninense can cause a basal stem rot on oil palm trees (Pilotti

2005), and G. philippii can cause a red root rot on Acacia mangium

(Glen et al. 2009). Besides, Ganoderma lucidum and A. rugosum

288

Sun et al.

have been used biotechnologically in the production of biofuel and

degradation of environmental pollutants (Jong et al. 2017, Wang

et al. 2021).

The systematics of the Ganodermataceae have been carried

out for about 100 years. Donk (1948) introduced Ganodermataceae

as a family based on its unique double-walled basidiospores

with obvious ornamentation on the endospore walls, which only

included Ganoderma with a laccate pileal surface and truncated

basidiospores and Amauroderma with globose to ellipsoid

basidiospores without a truncated apex. Murrill (1905) introduced

a genus Tomophagus that included pale-coloured basidiomata

with pale and soft context and truncated basidiospores. In the

past decades, specimens of Ganodermataceae have been

collected from all over the world (Steyaert 1972, Moncalvo &

Ryvarden 1997, Ryvarden 2004b, Hapuarachchi et al. 2019b)

except for the polar region. More genera have been established

by evidence of morphological characters and/or molecular data.

Imazeki (1952) presented Trachyderma as a genus with a eshy

succulent context and truncated basidiospores with spinules.

Steyaert (1972) examined several known species in Ganoderma,

then established Haddowia based on non-truncated basidiospores

with longitudinal ridges partly connected with short transverse

walls on the endospore walls; Humphreya with reticular or erratic

irregularly ridged double walls on truncated basidiospores; and

Magoderna based on anticlinal hyphae in pileipellis and ellipsoid

to ovoid basidiospores without truncated apex. Costa-Rezende

et al. (2017) established Foraminispora according to the hollow

columnar endospore ornamentation, which persist to the exospore

wall sometimes forming holes on the basidiospores, and Furtadoa

based on a monomitic hyphal system in context with clamped and

simple-septate generative hyphae. Sun et al. (2020) studied one

group of Amauroderma with a unique pore surface that changes to

blood red when bruised, and established Sanguinoderma. Costa-

Rezende et al. (2020b) established Amaurodermellus with a dull

pileal surface and ovoid basidiospores, and Cristataspora with

double-walled basidiospores which have endosporic ornamentation

as vertical or transverse ridges.

However, the uncertainty about the family level of

Ganodermataceae should not be ignored. Binder et al. (2013)

carried out the phylogenetic and phylogenomic analyses of

Polyporales based on 356 single-copy genes from 10 genomes

of this order, which showed that Ganoderma was involved

in the ‘core polyporoid clade’. Justo et al. (2017) suggested

Ganodermataceae as a synonym of Polyporaceae based on

ITS, nLSU and rpb1 sequences. The conclusions based on

phylogenetic evidence have certain credibility, while in this study,

Ganodermataceae is still regarded as an independent family based

on its remarkable morphological features to clarify the intergeneric

and interspecic relationships. For the time being, we prefer to

treat Ganodermataceae as an independent family different from

Polyporaceae according to previous specialised studies on the

Ganoderma group (Moncalvo & Ryvarden 1997, Robledo et al.

2015, Zhou et al. 2015, Hapuarachchi et al. 2018a, b, 2019a, b,

Xing et al. 2018, Costa-Rezende et al. 2020a, b, Sun et al. 2020,

Luangharn et al. 2021). The scientic status of Ganodermataceae

within the Polyporales should be considered in morphology,

phylogeny, and even whole genome sequences.

Ongoing taxonomic studies of Ganodermataceae from

Asia, Africa, Europe, Neotropics and North America have been

conducted for a long time with many new species and combinations

continually being reported (Otieno 1968, Steyaert 1972, Moncalvo

& Ryvarden 1997, Ryvarden 2004a, b, Gibertoni et al. 2008, Cao

et al. 2012, Le et al. 2012, Coetzee et al. 2015, Gomes-Silva

et al. 2015, Hapuarachchi et al. 2019b, Sun et al. 2020, Costa-

Rezende et al. 2020b). China has a complex and diverse natural

environment resulting in high species richness, and a total of 130

species of Ganodermataceae have been reported (Zhao & Zhang

2000, Dai 2012, Cao & Yuan 2012, Wang & Wu 2014, Li et al. 2015,

Zhou et al. 2015, Hapuarachchi et al. 2018b, Xing et al. 2018, Ye et

al. 2019, Sun et al. 2020). The great variability in the macroscopic

characters of the basidiomata and the relatively uniform macro-

and micro-morphology of most species in Ganodermataceae have

resulted in many confusions in taxonomy. As of 10 March 2022,

there were 642 records of Ganodermataceae recorded in Index

Fungorum (http://www.indexfungorum.org/), and 698 records in

MycoBank (http://www.mycobank.org/). Nearly half of these records

have been identied as synonyms, especially in Ganoderma and

Amauroderma and it is necessary to assess the validity of these

records.

With the rapid development of molecular techniques in recent

years, DNA sequence data have been widely used in the taxonomic

studies of Ganodermataceae. Moncalvo (1995) used ITS

sequences and the D2 region of nLSU sequences to construct the

relationships among species in Ganoderma, and concluded that

the combined data is useful for intrageneric segregation while the

D2 region is suitable for intergeneric or higher ranks segregation.

Subsequently, ITS and nLSU sequences were often used to identify

species (Cao et al. 2012, Le et al. 2012, de Lima Júnior et al. 2014,

Gomes-Silva et al. 2015, Li et al. 2015). It is worth mentioning

that Fryssouli et al. (2020) carried out a phylogenetic study of

Ganoderma based only on 3 970 ITS sequences obtained from

the GenBank/ENA/DDBJ database which evaluated the accuracy

of sequences and showed that Ganoderma can be divided into ve

main lineages. However, for the complex groups in Ganoderma

or for the higher rank classication of Ganodermataceae, most

researchers use multi-gene datasets to construct phylogenetic

trees (Zhou et al. 2015, Costa-Rezende et al. 2017, 2020b, Justo

et al. 2017, Cabarroi-Hernández et al. 2019, Hapuarachchi et

al. 2019b, Luangharn et al. 2020, Sun et al. 2020). At present,

eight genes have been applied to the phylogenetic analyses in

Ganodermataceae, viz., the internal transcribed spacer regions

(ITS), the large subunit of nuclear ribosomal RNA gene (nLSU),

the largest subunit of RNA polymerase II gene (rpb1), the second

largest subunit of RNA polymerase II gene (rpb2), the translation

elongation factor 1-α gene (tef1), the β-tubulin gene (tub), the small

subunit mitochondrial rRNA gene (mtSSU) and the small subunit

nuclear ribosomal RNA gene (nSSU). According to the records in

GenBank (https://www.ncbi.nlm.nih.gov/) as of 21 April 2021, 150

801 items were found by searching ‘Ganodermataceae’ directly, but

only about 65 000 items among them were identied as species of

Ganodermataceae. The number of sequences is considerable, but

repetitive sequences of the same species or specimens, inaccurate

identication and low quality of sequences make it necessary to

select only the reliable molecular data for phylogenetic analyses.

In this study, the specimens collected from all over the world

were studied by macromorphological and microscopic examinations

together with ultrastructural observations and phylogenetic

analyses based on six gene loci (ITS, nLSU, rpb2, tef1, mtSSU and

nSSU). A total of 146 species in Ganodermataceae with available

DNA sequences were involved in the phylogenetic analyses.

Based on morphological characters and phylogenetic evidence,

14 genera were conrmed within Ganodermataceae, Furtadoella

gen. nov., Neoganoderma gen. nov., Sinoganoderma gen. nov.

and Trachydermella gen. nov. were proposed as new genera;

289www.studiesinmycology.org

Ganodermataceae from China

278 species were conrmed in Ganodermataceae including

23 new species which are listed in Table 2. The ultrastructural

features observed under SEM of basidiospores of 10 genera in

Ganodermataceae were described and photographed. In total, 56

species and nine genera are described and illustrated here. A key

to accepted genera of Ganodermataceae and keys to accepted

species of Ganoderma, Haddowia, Humphreya, Magoderna,

Sanguinoderma, Tomophagus are also provided.

MATERIALS AND METHODS

Morphological studies

The studied specimens are deposited at the fungaria of the

Institute of Microbiology, Beijing Forestry University (BJFC,

Beijing, China), the Institute of Applied Ecology, Chinese Academy

of Sciences (IFP, Shenyang, China), the private fungarium of J.

Vlasák of Czech Republic (JV) and the Universidade Federal de

Pernambuco, Brazil (URM). Macro-morphological descriptions

of the new taxa (or selected taxa) were based on eld notes

and fungarium specimens. Special colour terms followed

Petersen (1996). Micro-morphological data were obtained from

dried specimens and observed under a compound microscope

following Cui et al. (2019) and Sun et al. (2020). Sections

were studied at a magnication up to 1 000× using Nikon E80i

microscope and phase contrast illumination (Nikon, Tokyo,

Japan). Line drawings were made with the aid of a drawing tube.

Ultrastructure of basidiospores was observed with Scanning

Electron Microscopy (SEM) using a Field Emission Scanning

Electron Microscope (FESEM) Hitachi SU-8010 (Hitachi, Ltd,

Tokyo, Japan) at Beijing Forestry University, China (BJFU).

Microscopic features, measurements and drawings were made

from slide preparations stained with Cotton Blue and Melzer’s

reagent. Spores were measured from sections cut from the tubes.

To represent the variation in the size of the basidiospores, 5 % of

measurements were excluded from each end of the range, and

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, L = mean spore length (arithmetic average of all

spores), W = mean spore width (arithmetic average of all spores),

Q = variation in the L/W ratios between the specimens studied, n

(a/b) = number of spores: (a) measured from given number, (b)

of specimens.

DNA extraction, amplication and sequencing

A cetyl trimethylammonium bromide (CTAB) rapid plant genome

extraction kit-DN14 (Aidlab Biotechnologies Co., Ltd, Beijing, China)

and a FH plant DNA kit II (Demeter Biotech Co., Ltd., Beijing, China)

were used to extract total genomic DNA from dried specimens and

to perform the polymerase chain reaction (PCR) according to the

manufacturer’s instructions with some modications (Xing et al.

2018, Sun et al. 2020). The ITS regions were amplied with primer

pairs ITS5 and ITS4 (White et al. 1990). The nLSU regions were

amplied with primer pairs LR0R and LR7, and the primer LR5 was

used sometimes as an alternative to LR7 (Vilgalys & Hester 1990).

The rpb2 regions were amplied with primer pairs fRPB2-5F and

fRPB2-7CR (Liu et al. 1999). The tef1 regions were amplied with

primer pairs EF1-983F and EF1-1567R (Rehner & Buckley 2005).

The mtSSU regions were amplied with primer pairs MS1 and MS2

(White et al. 1990). The nSSU regions were amplied with primer

pairs PNS1 and NS41 (White et al. 1990).

The PCR cycling schedule for ITS, tef1 and mtSSU included

an initial denaturation at 95 °C for 3 min, followed by 35 cycles

at 94 °C for 40 s, 54 °C for ITS and mtSSU, 55 °C for tef1 for

45 s, 72 °C for 1 min, and a nal extension at 72 °C for 10 min.

The PCR cycling schedule for nLSU and nSSU included an initial

denaturation at 94 °C for 1 min, followed by 35 cycles at 94 °C for

30 s, 50 °C for nLSU and 53 °C for nSSU for 1 min, 72 °C for 1.5

min, and a nal extension at 72 °C for 10 min. The PCR cycling

schedule for rpb2 included an initial denaturation at 94 °C for 2

min, followed by 10 cycles at 94 °C for 40 s, 60 °C for 40 s and

72 °C for 2 min, then followed by 37 cycles at 94 °C for 45 s, 55 °C

for 1.5 min and 72 °C for 2 min, and a nal extension of 72 °C

for 10 min. The PCR products were puried and sequenced at the

Beijing Genomics Institute (BGI), China, with the same primers. All

sequences analysed in this study were deposited at GenBank and

are listed in Table 1.

Phylogenetic analyses

The sequences generated in this study and retrieved from

GenBank were combined with ITS, nLSU, rpb2, tef1, mtSSU and

nSSU. Perenniporia subtephropora was selected as the outgroup

(Xing 2019). Phylogenetic analyses used in this study followed the

approach of Song & Cui (2017) and Shen et al. (2019). All sequences

of ITS, nLSU, rpb2, tef1, mtSSU and nSSU were respectively

aligned in MAFFT v. 7 (Katoh & Standley 2013, https://mafft.cbrc.jp/

alignment/server/) and manually adjusted in BioEdit v. 7.0.9. (Hall

1999). Alignments were spliced in Mesquite v. 3.2. (Maddison &

Maddison 2017). The partition homogeneity test (PHT) (Farris et

al. 1994) of the six-gene dataset was tested by PAUP v. 4.0b10

(Swofford 2002) under 1 000 homogeneity replicates. The best-t

evolutionary model was selected by hierarchical likelihood ratio

tests (hLRT) and Akaike information criterion (AIC) in MrModeltest

v. 2.3 (Nylander 2004) after scoring 24 models of evolution by

PAUP v. 4.0b10.

The Maximum Likelihood (ML) and Bayesian Inference (BI)

analyses were performed based on the combined dataset. Each

gene of ITS, nLSU, rpb2, tef1, mtSSU and nSSU was used to

perform ML analyses respectively. The ML analyses were performed

in RAxML-HPC v. 8.2.3 (Stamatakis 2014) and involved 1 000 ML

searches under the GTRGAMMA model, and only the Maximum

Likelihood best tree from all searches was provided. In addition,

1 000 rapid bootstrap replicates were run with the GTRCAT model

to assess ML bootstrap values of the nodes. Bayesian Inference

was calculated using MrBayes v. 3.1.2 (Ronquist & Huelsenbeck

2003) with four Markov chains, starting trees for 80 M generations

until the split deviation frequency < 0.01, and trees were sampled

every 100 generations. The rst 25 % of the sampled trees

were discarded as burn-in and the remaining ones were used

to reconstruct a majority rule consensus and calculate Bayesian

Posterior Probabilities (BPP) of the clades.

All trees were viewed in FigTree v. 1.4.2 (http://tree.bio.ed.ac.

uk/software/gtree/). The ML bootstrap support values ≥ 50 %

and Bayesian Posterior Probabilities ≥ 0.95 were presented on

topologies from ML analyses respectively. The nal alignments and

the retrieved topologies were deposited in TreeBASE (http:// www.

treebase.org), under accession ID: 27788 (http://purl.org/phylo/

treebase/phylows/study/TB2:S27788).

290

Sun et al.

Table 1. Taxa information and GenBank accession numbers of the sequences used in this study. Species in bold are new species or new combinations.

Species Voucher

GenBank accession No.

ReferencesITS nLSU rpb2 tef1 mtSSU nSSU

Amauroderma aurantiacum FLOR 52205 KR816510 KU315205 – – – – Costa-Rezende et al. (2016)

DHCR540 MF409961 MF409953 – – – – Costa-Rezende et al. (2017)

A. calcigenum FLOR 52315 KR816514 – – – – – Costa-Rezende et al. (2016)

URM 89213 MK119792 MK119870 MK121484 MK121541 MZ352778aMZ355137aSun et al. (2020), this study

JV 1808/51 MZ354865aMZ354997aMZ245374aMZ221624aMZ352779aMZ355138aThis study

URM83864 JX982565 – – – – – Gomes-Silva et al. (2015)

URM86847 KT006601 – – – – – Gomes-Silva et al. (2015)

URM 89566 MZ354866aMZ355111aMZ245375aMZ221625aMZ352780aMZ355146aThis study

A. calcitum FLOR 50931/DHCR538 KR816528 KU315207 – – – – Costa-Rezende et al. (2016)

FLOR 52230 (TYPE) KR816529 – – – – – Costa-Rezende et al. (2016)

A. camerarium FLOR 52169 KR816523 – – – – – Costa-Rezende et al. (2016)

FLOR 52216 KR816509 – – – – – Costa-Rezende et al. (2016)

A. cf. schomburgkii URM 89271 MK119802 MK119880 MK121495 MK121552 – MZ355289aSun et al. (2020), this study

URM 89272 MK119803 MK119881 MK121496 MK121553 – MZ355280aSun et al. (2020), this study

JV 1908/39 MZ354989aMZ354999aMZ245376aMZ221626a– MZ355147aThis study

A. elegantissimum Dai 17431 MK119794 MK119872 MK121493 MK121543 – MZ355288aSun et al. (2020), this study

URM 83822 MK119795 MK119873 MK121494 MK121544 MZ352784aMZ355308aSun et al. (2020), this study

A. exile URM82794 JX310845 – – – – – Gomes-Silva et al. (2015)

URM 89226 MK119796 MK119874 – MK121545 – MZ355249aSun et al. (2020), this study

A. oriformum URM83250 (TYPE) JX310846 – – – – – Gomes-Silva et al. (2015)

A. intermedium GAS910 MF409959 – – – – – Costa-Rezende et al. (2017)

FLOR 52248 KR816527 KU315209 – – – – Costa-Rezende et al. (2016)

JV 1312/E14-J MZ354868aMZ355096a– MZ221627a– MZ355151aThis study

A. laccatostipitatum HFSL ACGS7 KT006602 – – – – – Gomes-Silva et al. (2015)

URM 89240 MK119797 MK119875 MK121489 MK121546 MZ352781aMZ355262aSun et al. (2020), this study

A. omphalodes DHCR499/501 MF409956 MF409951 – MF421238 – – Costa-Rezende et al. (2017)

DHCR500 MF409957 MF409952 – MF421239 – – Costa-Rezende et al. (2017)

JV 1909/23-J MZ354991aMZ355000aMZ245377aMZ221628aMZ352785aMZ355250aThis study

A. partitum URM82882 JX310852 – – – – – Gomes-Silva et al. (2015)

URM83039 JX310853 – – – – – Gomes-Silva et al. (2015)

A. praetervisum URM 89233 MK119801 MK119879 MK121486 MK121551 MZ352926aMZ355315aSun et al. (2020), this study

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Ganodermataceae from China

Table 1. (Continued).

Species Voucher

GenBank accession No.

ReferencesITS nLSU rpb2 tef1 mtSSU nSSU

JV 1467/40 MZ354867a– – – – – This study

A. pseudoboletum FLOR 52318 KR816516 – – – – – Costa-Rezende et al. (2016)

A. robledoi FLOR 52249 KR816511 – – – – – Costa-Rezende et al. (2016)

URM84230 KC348461 – – – – – Unpublished

URM 87687 MK119800 MK119878 MK121487 MK121550 MZ352786aMZ355290aSun et al. (2020), this study

A. schomburgkii JV 1908/9 MZ354990a– – MZ221629a– MZ355152aThis study

URM 89225 MK119805 MK119883 MK121498 MK121555 – MZ355324aSun et al. (2020), this study

URM83228 JX310848 KT006621 – – – – Gomes-Silva et al. (2015)

Amauroderma sp.URM 89239 MZ354869aMZ355112aMZ245378aMZ221630aMZ352927aMZ355253aThis study

A. subsessile URM 89293 MK119806 MK119885 MK121490 MK121556 MZ352782aMZ355319aSun et al. (2020), this study

URM 89294 MK119807 MK119886 MK121491 MK121557 MZ352783aMZ355317aSun et al. (2020), this study

Amaurodermellus ovisporum DHCR127 (FLOR) MN077530 – – – – – Costa-Rezende et al. (2020b)

DHCR546 (HUEFS) MN077528 – – – – – Costa-Rezende et al. (2020b)

DHCR547 (HUEFS) MN077527 MN077553 – – – – Costa-Rezende et al. (2020b)

DHCR539 (HUEFS) MN077529 – – – – – Costa-Rezende et al. (2020b)

Cristataspora coffeata FLOR 50933 KU315204 – – – – – Costa-Rezende et al. (2020b)

1504/50 MZ354891a– – MZ221631a– – This study

Robledo 3183 (FCOS) MN077526 MN077560 – MN061695 – – Costa-Rezende et al. (2020b)

Robledo 3182 (FCOS) MN077525 MN077559 – – – – Costa-Rezende et al. (2020b)

C. avipora G299 MN077521 MN077555 – MN061694 – – Costa-Rezende et al. (2020b)

Foraminispora austrosinensis Cui 16425 MK119809 MK119888 – MK121559 MZ352835aMZ355257aSun et al. (2020), this study

Cui 14318 MK119810 MK119889 – MK121560 MZ352848aMZ355309aSun et al. (2020), this study

Fo. concentrica Cui 12644 (TYPE) MK119812 MK119891 MK121499 MK121561 MZ352839aMZ355310aSun et al. (2020), this study

Cui 16238 MK119816 MK119895 – MK121565 MZ352840aMZ355279aSun et al. (2020), this study

Cui 17141 MZ354892aMZ355001aMZ245379a– MZ352837aMZ355311aThis study

Cui 12648 MK119815 MK119894 MK121501 MK121564 MZ352849aMZ355314aSun et al. (2020), this study

Fo. rugosa DHCR512 MF409960 – – MF421240 – – Costa-Rezende et al. (2017)

DHCR560 MF409963 MF409955 – MF421241 – – Costa-Rezende et al. (2017)

URM 86888 MZ354945aMZ355131aMZ245380a– MZ352921aMZ355277aThis study

JV 1608/889-ND MZ354946a– – – MZ352919a– This study

Fo. yinggelingensis Cui 13618 (TYPE) MK119821 MK119900 MK121536 MK121570 MZ352838aMZ355275aSun et al. (2020), this study

292

Sun et al.

Table 1. (Continued).

Species Voucher

GenBank accession No.

ReferencesITS nLSU rpb2 tef1 mtSSU nSSU

Cui 13630 MK119822 MK119901 – – MZ352836aMZ355276aSun et al. (2020), this study

Fo. yunnanensis Cui 7974 KJ531653 KU220013 – – MZ352841a– Li & Yuan (2015), this study

Yuan 2253 KJ531655 – – – – – Li & Yuan (2015)

Furtadoella biseptata FLOR 50932 (TYPE) KU315196 KU315206 – – – – Costa-Rezende et al. (2016)

Fu. brasiliensis URM83578 JX310841 – – – – – Gomes-Silva et al. (2015)

TBG58 JX982569 – – – – – Gomes-Silva et al. (2015)

JV 1909/75 MZ354994aMZ355100aMZ245381aMZ221632aMZ352922aMZ355214aThis study

Ganoderma acaciicola Cui 16813 MZ354893aMZ355003aMZ245382a– – MZ355221aThis study

Cui 16814 MZ354894aMZ355004aMZ245383a– – MZ355219aThis study

Cui 16815 (TYPE) MZ354895aMZ355005aMZ245384a– – MZ355282aThis study

G. acontextum JV 0611/21G (TYPE) KF605667 – MG367489 MG367538 – – This study

JV 1208/11J KF605668 – MG367490 MG367540 – – This study

JV 1407/64 MG279151 – MG367491 MG367539 – – This study

G. adspersum Dai 13191 MG279153 – MG367492 MG367541 – MZ355157aXing et al. (2018), this study

HSBU-200894 MG279154 – – MG367542 – MZ355158aXing et al. (2018), this study

G. alpinum Cui 18402 MZ354910a– – – – – This study

Cui 17325 MZ354911a– – – – – This study

Cui 17467 (TYPE) MZ354912a– – – – – This study

G. angustisporum Cui 13817 (TYPE) MG279170 MZ355090aMG367507 MG367563 MZ352850aMZ355166aXing et al. (2018), this study

Dai 19603 MZ354978aMZ355047aMZ245385aMZ221633aMZ352856aMZ355207aThis study

Cui 18240 MZ354979aMZ355074aMZ245386aMZ221634aMZ352851aMZ355246aThis study

G. applanatum Cui 14062 MZ354913aMZ355076aMZ358846aMZ221635aMZ352857aMZ355202aThis study

Cui 14070 MZ354914aMZ355079aMZ245387aMZ221636a– MZ355203aThis study

G. aridicola Dai 12588 (TYPE) KU572491 – – KU572502 MZ352842aMZ355195aXing et al. (2016), this study

GanoTK01 JN105707 – – – JN105734 – Unpublished

GanoTK25 JN105708 – – – JN105719 – Unpublished

G. australe DHCR411 MF436675 MF436672 – MF436677 – – Costa-Rezende et al. (2017)

DHCR417 MF436676 MF436673 – MF436678 – – Costa-Rezende et al. (2017)

G. austroafricanum CBS 138724 KM507324 KM507325 MK611970 – – – Coetzee et al. (2015)

G. boninense WD 2085 KJ143906 – KJ143965 KJ143925 – – Zhou et al. (2015)

WD 2028 KJ143905 KU220015 KJ143964 KJ143924 – – Zhou et al. (2015)

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Ganodermataceae from China

Table 1. (Continued).

Species Voucher

GenBank accession No.

ReferencesITS nLSU rpb2 tef1 mtSSU nSSU

G. brownii JV 1105/9J MG279159 – MG367494 MG367547 – – Xing et al. (2018)

JV 0709/109 KF605662 – MG367495 MG367548 – – Unpublished

G. bubalinomarginatum Dai 20074 MZ354926aMZ355010aMZ245388aMZ221637aMZ352881aMZ355312aThis study

Dai 20075 (TYPE) MZ354927aMZ355040aMZ245389aMZ221638aMZ352882aMZ355224aThis study

G. calidophilum MFLU 19-2174 MN398337 – – – – – Luangharn et al. (2021)

MFLU 19-2219 MN398338 – – – – – Luangharn et al. (2021)

G. carnosum MUCL 49464 MG706220 MG706168 – – – – Unpublished

LGAM 1642 MG706217 MG706165 – – – – Unpublished

G. carocalcareum DMC 322 (TYPE) EU089969 – – – – – Douanla-Meli & Langer (2009)

DMC 513 EU089970 – – – EU089971 – Douanla-Meli & Langer (2009)

G. castaneum Dai 16500 MZ354918a– MZ245390aMZ221639aMZ352914aMZ355173aThis study

Cui 13893 MZ354919aMZ355013aMZ245391aMZ221640aMZ352915aMZ355185aThis study

Dai 13710 KU572489 MZ355045a– KU572499 MZ352917aMZ355229aXing et al. (2016), this study

Cui 17283 (TYPE) MZ354920a– – – MZ352916aMZ355230aThis study

G. casuarinicola Dai 16336 (TYPE) MG279173 MZ355103aMG367508 MG367565 MZ352843aMZ355297aXing et al. (2018), this study

Dai 16337 MG279174 MZ355104aMG367509 MG367566 MZ352844aMZ355196aXing et al. (2018), this study

Dai 19678 MZ354995aMZ355105a– MZ221641aMZ352845aMZ355204aThis study

Dai 19470 MZ354996a– MZ245392aMZ221642a– MZ355142aThis study

G. chalceum URM80457 JX310812 JX310826 – – – – De Lima Júnior et al. (2014)

G. chocoense QCAM 3123 (TYPE) MH890527 – – – – – Crous et al. (2018)

G. chuxiongense Cui 17262 (TYPE) MZ354907a– – – MZ352925aMZ355316aThis study

G. cocoicola Cui 16791 (TYPE) MZ354984aMZ355091aMZ245393aMZ221643a– MZ355321aThis study

Cui 16792 MZ354985aMZ355092aMZ245394aMZ221644a– MZ355278aThis study

G. concinnum Robledo 3192 MN077522 MN077556 – – – – Costa-Rezende et al. (2020b)

Robledo 3235 MN077523 MN077557 – – – – Costa-Rezende et al. (2020b)

G. cupreum GanoTK4 JN105701 – – – JN105732 – Unpublished

GanoTK7 JN105702 – – – JN105730 – Unpublished

G. curtisii CBS 100131 JQ781848 – KJ143966 KJ143926 – – Cao et al. (2012), Zhou et al. (2015)

CBS 100132 JQ781849 – KJ143967 KJ143927 – – Cao et al. (2012), Zhou et al. (2015)

G. destructans CMW43670 (TYPE) KR183856 – – – – – Coetzee et al. (2015)

Dai 16431 MG279177 – – – – – Xing et al. (2018)

294

Sun et al.

Table 1. (Continued).

Species Voucher

GenBank accession No.

ReferencesITS nLSU rpb2 tef1 mtSSU nSSU

G. dunense CMW42149 MG020248 – – MG020226 – – Tchotet Tchoumi et al. (2018)

CMW42157 (TYPE) MG020255 – – MG020227 – – Tchotet Tchoumi et al. (2018)

G. ecuadorense Dai 17397 MZ354950aMZ355019aMZ245398aMZ221648a– MZ355180aThis study

Dai 17418 MZ354951aMZ355020aMZ245399aMZ221649a– MZ355181aThis study

JV 1808/85 MZ354952aMZ355075a– MZ221650a– MZ355247aThis study

G. eickeri CMW50325 MH571689 – – MH567290 – – Tchotet Tchoumi et al. (2019)

CMW49692 (TYPE) MH571690 – – MH567287 – – Tchotet Tchoumi et al. (2019)

Dai 12595 MZ354964aMZ355035a– MZ221651a– MZ355159aThis study

Dai 12598 MZ354965aMZ355036a– MZ221652a– MZ355160aThis study

G. ellipsoideum MFLU 19-2221 MN398339 – – – – – Luangharn et al. (2021)

CMW 14080966 (TYPE) MH106867 – – – – – Hapuarachchi et al. (2018b)

Dai 19683 MZ354970aMZ355018a– MZ221653aMZ352893aMZ355217aThis study

Dai 20544 MZ354971aMZ355033aMZ245400aMZ221654aMZ352895aMZ355205aThis study

G. enigmaticum CMW43669 (TYPE) KR183855 KR183859 – – – – Coetzee et al. (2015)

Dai 15970 KU572486 MZ355106aMG367513 KU572496 MZ352846aMZ355197aXing et al. (2016, 2018), this study

Dai 15971 KU572487 MZ355107aMG367514 KU572497 MZ352847aMZ355198aXing et al. (2016, 2018), this study

G. fallax JV 1009/27 (TYPE) KF605655 – – – – – This study

JV 0709/39 KF605658 – – – – – This study

JV 0509/93K KF605653 – – – – – This study

G. exipes Cui 13841 MZ354923aMZ355063aMZ245401aMZ221655aMZ352905aMZ355177aThis study

Cui 13863 MZ354924aMZ355064aMZ245402aMZ221656a– MZ355178aThis study

Dai 20461 MZ354925aMZ355065aMZ245403aMZ221657a– MZ355153aThis study

G. fornicatum BCRC35374 JX840349 – – – – – Wang et al. (2014)

TNM-F0010592 JX840347 – – – – – Wang et al. (2014)

G. gibbosum KUMCC17-0003 MH035681 MH553157 – – – – Luangharn et al. (2020)

KUMCC17-0005 MH035682 MH553158 – – – – Luangharn et al. (2020)

Cui 13940 MZ354972aMZ355021aMZ245404aMZ221658aMZ352894aMZ355161aThis study

Cui 14338 MZ354969aMZ355014aMZ245405aMZ221659aMZ352876aMZ355162aThis study

Cui 17769 MZ354967a– – – – – This study

Cui 17780 MZ354968a– – – – – This study

Cui 17254 MZ354966aMZ355115aMZ245406aMZ221660aMZ352877aMZ355286aThis study

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Ganodermataceae from China

Table 1. (Continued).

Species Voucher

GenBank accession No.

ReferencesITS nLSU rpb2 tef1 mtSSU nSSU

G. guangxiense Cui 14453 (TYPE) MZ354939aMZ355037aMZ245407aMZ221661aMZ352896aMZ355163aThis study

Cui 14454 MZ354941aMZ355039aMZ245408aMZ221662aMZ352897aMZ355164aThis study

Cui 14508 MZ354940aMZ355038a– MZ221663aMZ352865aMZ355240aThis study

G. hochiminhense MFLU 19-2224 (TYPE) MN398324 MN396390 – MN423176 – – Luangharn et al. (2021)

MFLU 19-2225 MN396662 MN396391 – MN423177 – – Luangharn et al. (2021)

Cui 18229 MZ354986aMZ355094aMZ245409aMZ221664a– MZ355283aThis study

Dai 18488 MZ354987aMZ355093aMZ245410aMZ221665a– MZ355218aThis study

G. hoehnelianum Cui 13904 MZ354935aMZ355135aMZ245411aMZ221666aMZ352888aMZ355169aThis study

Cui 13982 MG279178 MZ355071aMG367515 MG367570 – MZ355170aXing et al. (2018), this study

Dai 20783 – MZ355002aMZ245412aMZ221667aMZ352892aMZ355255aThis study

G. japonicum AS569 AY593864 – – – – – Wang & Yao (2005)

Gja-1 GU213475 – – – – – Unpublished

G. knysnamense CMW47755 (TYPE) MH571681 – – MH56726 – – Tchotet Tchoumi et al. (2019)

CMW49688 MH571683 – – MH567266 – – Tchotet Tchoumi et al. (2019)

G. leucocontextum Dai 15601 KU572485 MZ355049aMG367516 KU572495 MZ352899aMZ355318aXing et al. (2018), this study

GDGM 40200 (TYPE) KF011548 – – – – – Li et al. (2015)

G. lingzhi Wu 1006-38 (TYPE) JQ781858 – JX029980 JX029976 – – Cao et al. (2012)

Cui 9166 KJ143907 – JX029978 JX029974 JX029987 – Zhou et al. (2015)

Dai 20895 MZ354904aMZ355006aMZ245413aMZ221668a– – This study

Cui 18161 MZ354905a– – – – – This study

Cui 18167 MZ354906a– – – – – This study

G. lobatum JV 1008/31 KF605671 – MG367499 MG367553 – – Xing et al. (2018)

JV 1008/32 KF605670 – MG367500 MG367554 – – Xing et al. (2018)

G. lucidum Cui 14404 MG279181 MZ355051aMG367519 MG367573 MZ352858aMZ355191aXing et al. (2018), this study

Cui 14405 MG279182 MZ355089aMG367520 MG367574 – MZ355194aXing et al. (2018), this study

K 175217 KJ143911 – KJ143971 KJ143929 – – Zhou et al. (2015)

MT 26/10 KJ143912 – – KJ143930 – – Zhou et al. (2015)

Dai 20017 MZ354937aMZ355050a– MZ221669a– MZ355256aThis study

G. magniporum Zhou 439 MZ354936aMZ355097a– – MZ352863a– This study

Dai 19966 – MZ355098aMZ345728aMZ221670a– MZ355223aThis study

G. martinicense LIP SW-Mart08-55 (TYPE) KF963256 – – – – – Unpublished

296

Sun et al.

Table 1. (Continued).

Species Voucher

GenBank accession No.

ReferencesITS nLSU rpb2 tef1 mtSSU nSSU

He 2240 MG279163 – MG367503 MG367557 – – Xing et al. (2018)

G. mastoporum K15-86 MF680427 – – – – – Unpublished

TNM-F0018835 JX840351 – – – – – Wang et al. (2014)

G. mbrekobenum UMN7-3 GHA (TYPE) KX000896 KX000897 – – – – Crous et al. (2016)

UMN7-4 GHA KX000898 KX000899 – – – – Crous et al. (2016)

G. meredithiae UMNFL50 MG654103 – – – – – Loyd et al. (2018)

UMNFL64 MG654106 – MG754863 – – – Loyd et al. (2018)

G. mexicanum MUCL 49453 MK531811 – MK531836 MK531825 – – Cabarroi-Hernández et al. (2019)

MUCL 55832 MK531815 – MK531839 MK531829 – – Cabarroi-Hernández et al. (2019)

G. mirabile Cui 16408 – MZ355066a– MZ221671a– MZ355227aThis study

Cui 18271 MZ354958aMZ355067aMZ345729aMZ221672aMZ352860aMZ355231aThis study

Cui 18283 MZ354959aMZ355069aMZ345730aMZ221673aMZ352861aMZ355248aThis study

Cui 18237 MZ354960aMZ355068aMZ345731aMZ221674aMZ352862aMZ355243aThis study

G. mizoramense UMN-MZ4 (TYPE) KY643750 – – – – – Crous et al. (2017a)

UMN-MZ5 KY643751 KY747490 – – – – Crous et al. (2017a)

JZ8 MG437336 – – – – – Unpublished

G. multipileum Cui 13597 MZ354899aMZ355043aMZ345732aMZ221675aMZ352866aMZ355174aThis study

Dai 17569 MZ354896aMZ355007aMZ345733aMZ221676aMZ352867aMZ355175aThis study

Dai 19690 MZ354897aMZ355008a– MZ221677aMZ352868aMZ355215aThis study

Dai 19691 MZ354898aMZ355041a– – MZ352869a– This study

G. multiplicatum SPC9 KU569553 KU570951 – – – – Bolaños et al. (2016)

CC8 KU569515 KU570915 – – – – Bolaños et al. (2016)

URM83346 JX310823 JX310837 – – – – De Lima Júnior et al. (2014)

Dai 17395 MZ354903a– MZ345734aMZ221678aMZ352870aMZ355209aThis study

G. mutabile Yuan 2289 (TYPE) JN383977 – – – – – Cao & Yuan (2012)

Cui 17189 MZ354976a– – MZ221679a– – This study

Dai 20414 MZ354977aMZ355110aMZ345735aMZ221680aMZ352864aMZ355292aThis study

G. myanmarense MFLU 19-2167 (TYPE) MN396330 MN428672 – – – – Luangharn et al. (2021)

MFLU 19-2211/2169 MN396329 MN398325 – – – – Luangharn et al. (2021)

G. nasalanense CACP17060211 (TYPE) MK345441 MK346831 – – – MK346842 Hapuarachchi et al. (2019b)

CACP17060212 MK345442 MK346832 – – – MK346843 Hapuarachchi et al. (2019b)

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Ganodermataceae from China

Table 1. (Continued).

Species Voucher

GenBank accession No.

ReferencesITS nLSU rpb2 tef1 mtSSU nSSU

G. neojaponicum AS5.541 (TYPE) AY593866 – – – – – Wang & Yao (2005)

36073 AY335163 – – – – – Unpublished

G. nitidum JV 1504/73 MZ354933a– – MZ221681aMZ352883a– This study

G. orbiforme Cui 13880 MG279187 MZ355016aMG367523 MG367577 MZ352908aMZ355188aXing et al. (2018), this study

Cui 13891 MZ354953aMZ355017aMZ345736aMZ221682aMZ352910aMZ355167aThis study

Cui 18301 MZ354954aMZ355070a– MZ221683aMZ352911aMZ355232aThis study

Cui 18302 MZ354955aMZ355072a– MZ221684aMZ352912aMZ355233aThis study

Cui 18317 MZ354956aMZ355059a– MZ221685aMZ352909aMZ355285aThis study

Cui 18326 MZ354957aMZ355062a– MZ221686aMZ352913aMZ355244aThis study

G. oregonense CBS 266.88 JQ781876 – KJ143975 – – – Cao et al. (2012), Zhou et al. (2015)

CBS 265.88 JQ781875 – KJ143974 KJ143933 – – Cao et al. (2012), Zhou et al. (2015)

G. parvulum URM83345 JX310820 JX310834 – – – – De Lima Júnior et al. (2014)

URM83344 JX310819 JX310833 – – – – De Lima Júnior et al. (2014)

G. pfeifferi Dai 12153 MG279164 MZ355109a– MG367559 – MZ355168aXing et al. (2018), this study

Dai 12683 MG279165 MZ355108a– MG367560 – – Xing et al. (2018), this study

CBS 221.48 – MH867868 – – – – Unpublished

G. philippii Cui 14443 MG279188 MZ355023aMG367524 MG367578 MZ352871aMZ355186aXing et al. (2018), this study

Cui 14444 MG279189 MZ355022aMG367525 MG367579 – MZ355187aXing et al. (2018), this study

G. platense BAFC384 AH008109 – – – – – Gottlieb et al. (2000)

BAFC2374 AH008110 – – – – – Gottlieb et al. (2000)

G. podocarpense QCAM 6422 (TYPE) MF796661 MF796660 – – – – Crous et al. (2017b)

JV 1504/126 MZ354942a– MZ345737aMZ221687a– – This study

G. polychromum UMNOR3 MG654204 – – MG754744 – – Loyd et al. (2018)

MS343OR MG654197 – – MG754743 – – Loyd et al. (2018)

G. puerense Dai 20427 (TYPE) – MZ355012aMZ345738aMZ221688aMZ352884aMZ355241aThis study

G. ravenelii MS187FL MG654211 – MG754865 MG754745 – – Loyd et al. (2018)

151FL MG654208 – – – – – Loyd et al. (2018)

G. resinaceum MS1211 MT397406 – – MT415669 – – Náplavová et al. (2020)

MS1212 MT397407 – – MT415670 – – Náplavová et al. (2020)

LGAM 462 MG706250 MG706196 MG837821 MG837858 – – Unpublished

LGAM 448 MG706249 MG706195 MG837820 MG837857 – – Unpublished

298

Sun et al.

Table 1. (Continued).

Species Voucher

GenBank accession No.

ReferencesITS nLSU rpb2 tef1 mtSSU nSSU

G. ryvardenii HKAS 58053 (TYPE) HM138671 – – – – – Kinge & Mih (2011)

HKAS 58054 HM138672 – – – – – Kinge & Mih (2011)

HKAS 58055 HM138670 – – – – – Kinge & Mih (2011)

G. sanduense SA18012501 (TYPE) MK345450 – – – – – Hapuarachchi et al. (2019b)

SA18012502 MK345451 – – – – – Hapuarachchi et al. (2019b)

G. sessile Dai 16403 MZ354934aMZ355015aMZ345739aMZ221689aMZ352907aMZ355184aThis study

JV 1209/27 KF605630 – KJ143976 KJ143937 – – Zhou et al. (2015)

G. shanxiense HSA 539 MK764269 – MK789681 – – – Liu et al. (2019)

BJTC FM423 (TYPE) MK764268 – MK783940 MK783937 – – Liu et al. (2019)

Cui 14565 MZ354908a– – MZ221690a– MZ355245aThis study

Dai 18921 MZ354909aMZ355044aMZ345740aMZ221691a– MZ355320aThis study

G. sichuanense HMAS42798 (TYPE) JQ781877 – – – – – Cao et al. (2012)

Cui 16343 MZ354928aMZ355011aMZ345741aMZ221692aMZ352885aMZ355171aThis study

Dai 19651 MZ354929aMZ355031aMZ345742aMZ221693aMZ352889aMZ355208aThis study

G. sinense Cui 14526 MZ354961aMZ355056aMZ345743aMZ221694a– MZ355189aThis study

Cui 14461 MZ354963aMZ355057aMZ345744aMZ221695a– MZ355190aThis study

Dai 20079 MZ354962aMZ355058aMZ345745aMZ221696a– MZ355284aThis study

Ganoderma sp.N.1 MH806359 – – – – – Le et al. (2018)

N.3 MH806363 – – – – – Le et al. (2018)

G. steyaertianum 6-WN-15(M)-A KJ654459 – – – – – Unpublished

6-WN-16(M)-A KJ654461 – – – – – Unpublished

G. stipitatum THC 16 KC884264 – – – – – Unpublished

G. subangustisporum Cui 18592 (TYPE) MZ354981aMZ355027a– MZ221697aMZ352854a– This study

Cui 18593 MZ354982aMZ355028a– MZ221698aMZ352852a– This study

Cui 18596 MZ354983aMZ355029a– MZ221699aMZ352853a– This study

Cui 18597 MZ354980aMZ355025aMZ345746aMZ221700aMZ352855aMZ355216aThis study

G. subellipsoideum Cui 18241 – MZ355132a– MZ221701aMZ352878a– This study

Cui 18325 (TYPE) – MZ355134a– MZ221702a– MZ355295aThis study

Cui 18327 – MZ355133a– MZ221703aMZ352859aMZ355296aThis study

G. subexipes Cui 17247 MZ354921aMZ355128aMZ245395aMZ221645a– MZ355140aThis study

Cui 17257 (TYPE) MZ354922aMZ355129aMZ245396aMZ221646a– MZ355220aThis study

299www.studiesinmycology.org

Ganodermataceae from China

Table 1. (Continued).

Species Voucher

GenBank accession No.

ReferencesITS nLSU rpb2 tef1 mtSSU nSSU

Cui 17258 – MZ355130aMZ245397aMZ221647a– MZ355143aThis study

G. sublobatum Cui 16804 (TYPE) MZ354973a– MZ345747aMZ221704aMZ352879aMZ355293aThis study

Cui 16806 MZ354974aMZ355034a– MZ221705aMZ352918aMZ355165aThis study

G. thailandicum HKAS 104640 (TYPE) MK848681 MK849879 MK875831 MK875829 – – Luangharn et al. (2019)

HKAS 104641 MK848682 MK849880 MK875832 MK875830 – – Luangharn et al. (2019)

G. tongshanense Cui 17168 (TYPE) MZ354975aMZ355024a– MZ221706a– MZ355206aThis study

G. tornatum URM82776 JQ514110 JX310800 – – – – Unpublished

TBG01AM2009 JQ514108 JX310808 – – – – Unpublished

G. tropicum Dai 16434 MG279194 MZ355026aMG367532 MG367585 MZ352872aMZ355176aXing et al. (2018), this study

Dai 19679 MZ354900aMZ355009aMZ358825aMZ221707aMZ352873aMZ355222aThis study

Dai 20029 MZ354902a– MZ358826aMZ221708aMZ352880aMZ355225aThis study

Dai 19491 MZ354901a– MZ358827aMZ221709aMZ352874aMZ355141aThis study

G. tsugae Cui 14110 MG279195 – MG367533 MG367586 MZ352903aMZ355192aXing et al. (2018), this study

Cui 14112 MG279196 – MG367534 MG367587 MZ352904aMZ355193aXing et al. (2018), this study

G. tuberculosum Dai 17412 MZ354943a– – – MZ352906aMZ355199aThis study

JV 1607/62 MZ354944aMZ355087a– MZ221710aMZ352875aMZ355294aThis study

G. weberianum Cui 16359 – MZ355116aMZ358828aMZ221711aMZ352886aMZ355172aThis study

CBS 219.36 MK603804 MH867289 MK611972 MK611974 – – Cabarroi-Hernández et al. (2019)

CBS 128581 MK603805 MH876427 MK611971 MK636693 – – Cabarroi-Hernández et al. (2019)

Dai 19673 MZ354930aMZ355032aMZ358829aMZ221712aMZ352890aMZ355210aThis study

Dai 19682 MZ354932aMZ355042aMZ358830aMZ221713aMZ352887aMZ355213aThis study

Dai 19689 MZ354931aMZ355046a– MZ221714aMZ352891aMZ355226aThis study

G. weixiense HKAS 100649 (TYPE) MK302444 MK302446 – MK302442 – – Ye et al. (2019)

HKAS 100650 MK302445 MK302447 – MK302443 – – Ye et al. (2019)

G. wiiroense UMN-20-GHA KT952361 KT952362 – – – – Crous et al. (2015)

MIN 938704 (TYPE) KT952363 KT952364 – – – – Crous et al. (2015)

G. williamsianum Dai 17790 MZ354947aMZ355060a– MZ221715a– MZ355182aThis study

Dai 16809 MG279183 MZ355030aMG367535 MG367588 – MZ355183aXing et al. (2018), this study

Dai 20553 MZ354948aMZ355061aMZ358831aMZ221716a– MZ355242aThis study

Dai 19611 MZ354949aMZ355048aMZ358832aMZ221717a– – This study

G. yunlingense Cui 16288 (TYPE) MZ354915aMZ355077a– MZ221718a– MZ355179aThis study

300

Sun et al.

Table 1. (Continued).

Species Voucher

GenBank accession No.

ReferencesITS nLSU rpb2 tef1 mtSSU nSSU

Cui 17043 MZ354916aMZ355078a– MZ221719a– MZ355228aThis study

Cui 17958 MZ354917a– – – – – This study

G. zonatum FL-03 KJ143922 – KJ143980 KJ143942 – – Zhou et al. (2015)

FL-02 KJ143921 – KJ143979 KJ143941 – – Zhou et al. (2015)

Haddowia longipes LPDR17072708 MK345423 MK346828 – – – MK346836 Hapuarachchi et al. (2019b)

LPDR17072709 MK345424 MK346829 – – – MK346837 Hapuarachchi et al. (2019b)

Ha. macropora JV 1908/46 (TYPE) MZ354870aMZ354998aMZ358847aMZ221720aMZ352923aMZ355251aThis study

Magoderna subresinosum Dai 18626 MK119823 MK119902 MK121507 MK121571 MZ352831aMZ355211aSun et al. (2020), this study

Cui 18262 MZ354871aMZ355088a– – MZ352832aMZ355258aThis study

Cui 18280 MZ354872aMZ355095a– MZ221721aMZ352833aMZ355304aThis study

Neoganoderma neurospora DHCR559 MN077531 – – – – – Costa-Rezende et al. (2020b)

GAS1013 MN077532 – – – – – Costa-Rezende et al. (2020b)

Sanguinoderma bataanense Cui 6285 MK119831 MK119910 MK121537 MK121580 MZ352793aMZ355238aSun et al. (2020), this study

Dai 10746 MK119832 MK119911 MK121511 MK121581 MZ352801aMZ355267aSun et al. (2020), this study

Zhou 153 KJ531657 – – – – – Li & Yuan (2015)

Dai 7862 KJ531658 – – – – – Li & Yuan (2015)

Sa. elmerianum Cui 8940 MK119833 MK119912 – – MZ352812aMZ355305aSun et al. (2020), this study

HMAS 133187 MK119834 MK119913 – – MZ352824aMZ355234aSun et al. (2020), this study

Cui 18234 MZ354873aMZ355080a– MZ221722aMZ352814aMZ355236aThis study

Dai 20503 MZ354874aMZ355081a– MZ221723aMZ352813aMZ355154aThis study

Dai 20634 MZ354875aMZ355082a– MZ221724aMZ352821aMZ355148aThis study

Sa. avovirens Cui 16935 (TYPE) – MK119914 MK121532 MK121582 MZ352811aMZ355254aSun et al. (2020), this study

Sa. guangdongense Dai 16724 MZ354876aMZ355117aMZ358833aMZ221725aMZ352815aMZ355271aThis study

Cui 17259 (TYPE) MZ354877aMZ355123aMZ358834aMZ221726aMZ352816aMZ355139aThis study

Dai 20419 MZ354890aMZ355083aMZ358835aMZ221727aMZ352818aMZ355155aThis study

Cui 17240 – MZ355124aMZ358836aMZ221728aMZ352817aMZ355287aThis study

Sa. infundibulare Dai 18148 MK119846 MK119925 MK121528 MK121596 MZ352789aMZ355259aSun et al. (2020), this study

Dai 18149 MK119847 MK119926 MK121529 MK121597 MZ352790aMZ355239aSun et al. (2020), this study

Cui 17238 OM780277 – MZ358837aMZ221729aMZ352800aMZ355149aThis study

Cui 17248 (TYPE) MZ354880aMZ355125a– MZ221730aMZ352787aMZ355150aThis study

Dai 18151 MK119848 – MK121530 MK121598 MZ352788aMZ355274aSun et al. (2020), this study

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Ganodermataceae from China

Table 1. (Continued).

Species Voucher

GenBank accession No.

ReferencesITS nLSU rpb2 tef1 mtSSU nSSU

Cui 17256 MZ354885a– MZ358838aMZ221731aMZ352791aMZ355144aThis study

URM 450213 MK119849 MK119927 – – MZ352792aMZ355252aSun et al. (2020), this study

Sa. laceratum Cui 8155 (TYPE) MK119851 MK119928 – – MZ352810a– Sun et al. (2020), this study

A5 MG383652 – – – – – Unpublished

Sa. longistipitum Dai 20696 (TYPE) MZ354881aMZ355084a– MZ221732aMZ352822aMZ355145aThis study

Cui 13903 MZ354882aMZ355114aMZ358839aMZ221733aMZ352809aMZ355301aThis study

Dai 13891 MZ354886aMZ355126a– – MZ352834aMZ355325aThis study

Dai 16635 MZ354883aMZ355120aMZ358840aMZ221734aMZ352802aMZ355260aThis study

Sa. melanocarpum Dai 18512 MZ354888aMZ355118a– MZ221735aMZ352794aMZ355313aThis study

Dai 18603 (TYPE) MZ354889aMZ355113aMZ358841aMZ221736aMZ352796aMZ355281aThis study

Sa. microporum Cui 13851 (TYPE) MK119854 MK119933 MK121512 MK121602 MZ352797aMZ355270aSun et al. (2020), this study

Cui 18270 – MZ355086a– – MZ352808aMZ355235aThis study

Cui 14022 MK119856 MK119935 MK121515 MK121604 MZ352798aMZ355298aSun et al. (2020), this study

Sa. microsporum Dai 16726 (TYPE) – MZ355119a– MZ221737aMZ352795aMZ355272aThis study

Cui 13901 MZ354879aMZ355121a– MZ221738aMZ352803aMZ355299aThis study

Cui 13897 MZ354878aMZ355127a– MZ221739aMZ352804aMZ355300aThis study

Sa. perplexum Cui 6496 KJ531650 KU220001 MK121538 MK121583 MZ352825aMZ355263aLi & Yuan (2015), this study

Cui 6554 MK119835 MK119915 MK121540 MK121585 MZ352826aMZ355264aSun et al. (2020), this study

Wei 5562 KJ531652 – – – – – Li & Yuan (2015)

Dai 10811 KJ531651 KU220002 MK121539 MK121584 MZ352827aMZ355302aLi & Yuan (2015), this study

Sa. reniforme Cui 16511 (TYPE) MK119850 MK119929 MK121531 MK121599 – MZ355322aSun et al. (2020), this study

Sa. rude Cui 16592 MK119836 MK119916 MK121521 MK121586 MZ352924aMZ355307aSun et al. (2020), this study

DHCR457 MN077517 MN077551 – MN061693 – – Costa-Rezende et al. (2020b)

MEL 2317411 MK119842 – MK121524 MK121592 MZ352819aMZ355306aSun et al. (2020), this study

Sa. rugosum Cui 8795 MK119843 MK119922 MK121516 MK121593 MZ352799aMZ355266aSun et al. (2020), this study

Cui 9011 KJ531664 KU220010 MK121517 KU572504 MZ352805aMZ355237aLi & Yuan (2015), this study

Cui 9066 MZ354884aMZ355122a– MZ221740aMZ352806aMZ355268aThis study

Dai 20582 MZ354887aMZ355085aMZ358842aMZ221741aMZ352823aMZ355156aThis study

Cui 9012 KJ531665 KU220011 MK121518 KU572503 MZ352807aMZ355269aLi & Yuan (2015), this study

Sa. sinuosum MEL 2366586 MK119852 MK119930 MK121527 MK121600 MZ352920aMZ355261aSun et al. (2020), this study

MEL 2341763 (TYPE) MK119853 MK119931 MK121525 MK121601 MZ352820aMZ355291aSun et al. (2020), this study

302

Sun et al.

Table 1. (Continued).

Species Voucher

GenBank accession No.

ReferencesITS nLSU rpb2 tef1 mtSSU nSSU

Sa. tricolor Cui 18292 (TYPE) – MZ355101a– MZ221742aMZ352828aMZ355273aThis study

Cui 18242 MZ354992aMZ355099aMZ358843aMZ221743aMZ352829aMZ355303aThis study

Dai 18574 MZ354993aMZ355102aMZ358844aMZ221744aMZ352830aMZ355265aThis study

Sinoganoderma shandongense Dai 15785 MG279190 MZ355052aMG367526 MG367580 MZ352900aMZ355200aXing et al. (2018), this study

Dai 15787 MG279191 MZ355053aMG367527 MG367581 MZ352901aMZ355201aXing et al. (2018), this study

Dai 15791 MG279192 MZ355054aMG367528 MG367582 MZ352902aMZ355323aXing et al. (2018), this study

Dai 20243 – MZ355055a– MZ221745a– – This study

Dai 20244 MZ354938aMZ355073a– MZ221746a– – This study

xsd08032 EU918700 – – – – – Unpublished

xsd08085 FJ478127 – – – – – Unpublished

Tomophagus cattienensis CT119 JN184398 – – – – – Le et al. (2012)

CT99 (TYPE) JN184397 – – – – – Le et al. (2012)

Dai 18487 MZ354988a– MZ358845aMZ221747aMZ352898aMZ355212aThis study

To. colossus URM80450 JX310825 JX310839 – – – – De Lima Júnior et al. (2014)

URM83330 JQ618247 JX310811 – – – – De Lima Júnior et al. (2014)

Trachydermella tsunodae GR363 FJ154773 – – – – – Unpublished

WD2034 AB588989 AB368069 AB368127 – – – Sotome et al. (2011)

Perenniporia subtephropora Dai 10962 (TYPE) JQ861752 JQ861768 KX880850 KF286329 KF218323 – Zhao & Cui (2013)

Dai 10964 JQ861753 JQ861769 KX880851 KF286330 KF218324 – Zhao & Cui (2013)

a Newly generated sequences for this study.

303www.studiesinmycology.org

Ganodermataceae from China

RESULTS

Molecular phylogeny

In this study, 1 382 sequences derived from six gene loci (ITS, nLSU,

rpb2, tef1, mtSSU and nSSU) were used to reconstruct phylogenetic

trees of Ganodermataceae, including 374 sequences of ITS, 242

sequences of nLSU, 173 sequences of rpb2, 242 sequences of

tef1, 158 sequences of mtSSU and 193 sequences of nSSU. The

combined six-gene dataset (ITS + nLSU + rpb2 + tef1 + mtSSU +

nSSU) included sequences from 391 specimens representing 146

taxa from Ganodermataceae and Perenniporia subtephropora as

the outgroup. The partition homogeneity test indicated all the six

different genes displayed a congruent phylogenetic signal (P value

= 1.00). The best-t evolutionary models selected by MrModeltest

v. 2.3 for each region of the six genes were GTR + I + G (ITS1), K80

(5.8S), HKY + I + G (ITS2), GTR + I + G (nLSU), K80 + I + G (rpb2

introns), K80 + G (rpb2 1st codon), GTR + I + G (rpb2 2nd codon),

GTR + I + G (tef1 introns), HKY + I + G (tef1 1st codon), SYM + I

+ G (tef1 2nd codon), SYM + I + G (tef1 3rd codon), GTR + I + G

(mtSSU) and GTR + I + G (nSSU). These models were applied in

Bayesian analyses for the combined dataset.

The combined six-gene dataset has an aligned length of 5 172

total characters including gaps, of which 3 780 are constant, 197 are

variable and parsimony-uninformative, and 1 195 are parsimony-

informative. The average standard deviation of split frequencies in

the Bayesian analyses reached 0.008329. The calculated values

based on the combined six-gene dataset are shown in Fig. 1.

Thirteen clades were obtained in the phylogenetic analyses of

Ganodermataceae: Amauroderma clade (100 % ML, 1.00 BPP),

Amaurodermellus clade (100 % ML, 1.00 BPP), Cristataspora

clade (100 % ML, 1.00 BPP), Foraminispora clade (99 % ML,

1.00 BPP), Furtadoella gen. nov. clade (100 % ML, 1.00 BPP),

Ganoderma clade (58 % ML), Haddowia clade (85 % ML, 0.99

BPP), Magoderna clade (100 % ML, 1.00 BPP), Neoganoderma

gen. nov. clade (100 % ML, 1.00 BPP), Sanguinoderma clade

(88 % ML, 0.98 BPP), Sinoganoderma gen. nov. clade (100 %

ML, 1.00 BPP), Tomophagus clade (100 % ML, 1.00 BPP) and

Trachydermella gen. nov. clade (100 % ML, 1.00 BPP)

The Ganoderma clade is composed of 95 taxa including 16

new species. All taxa in this clade were divided into two groups

according to laccate or dull pileal surface, and 10 subclades are

separated by this feature: subclade I-laccate/dull (84 % ML, 1.00

BPP), subclade II-laccate (100 % ML, 1.00 BPP), subclade III-

laccate (100 % ML, 1.00 BPP), subclade IV-dull (100 % ML, 1.00

BPP), subclade V-laccate/dull (93 % ML, 1.00 BPP), subclade

VI-dull (98 % ML, 1.00 BPP), subclade VII-laccate/dull (75 % ML,

0.99 BPP), subclade VIII, subclade IX (100 % ML, 1.00 BPP) and

subclade X (99 % ML, 1.00 BPP), these subclades were shown in

Fig. 1.

The phylogenetic topologies of Ganodermataceae based on

ITS, nLSU, rpb2, tef1, mtSSU and nSSU sequences respectively

with ML bootstrap support values ≥ 50 % are shown in Figs 2–7.

Besides, including Perenniporia subtephropora as outgroup, there

were 146 taxa included in the ITS dataset, 107 taxa included in

the nLSU dataset, 87 taxa included in the rpb2 dataset, 102 taxa

included in the tef1 dataset, 70 taxa included in the mtSSU dataset,

and 81 taxa included in the nSSU dataset.

Taxonomy

Ganodermataceae Donk, Bull. Bot. Gdns Buitenz. 17: 474. 1948.

Fig. 8. MycoBank MB 80782.

Type genus: Ganoderma P. Karst.

Description: Basidiomata annual to perennial, sessile to stipitate,

pileate, eshy to woody hard. Pilei variable in shape and colour, with

or without laccate surface. Hyphal system dimitic to trimitic, rarely

monomitic in context; generative hyphae mostly bearing clamp

connections, rarely simple-septate. Basidiospores subglobose to

ovoid or reniform, truncated or not, double-walled and slightly to

distinctly thick-walled with varied ornamentation.

Notes: In this study, 12 genera of Ganodermataceae:

Amauroderma, Amaurodermellus, Cristataspora, Foraminispora,

Furtadoella, Ganoderma, Haddowia, Humphreya, Magoderna,

Sanguinoderma, Tomophagus, Trachydermella and two new

genera: Neoganoderma and Sinoganoderma were conrmed

based on morphological and molecular studies. Humphreya was

not included in the phylogenetic analyses since there are no

available specimens to obtain sequences, but it is treated as an

independent genus within Ganodermataceae based on its unique

basidiospore ornamentation.

Key to accepted genera of Ganodermataceae

1a. Colour of fresh pore surface becoming blood red when bruised .................................................................................... Sanguinoderma

1b. Colour of fresh pore surface darkening or unchanged when bruised .................................................................................................... 2

2a. Basidiospores non-truncated ................................................................................................................................................................. 3

2b. Basidiospores truncated ......................................................................................................................................................................... 8

3a. Hyphal system monomitic in context, generative hyphae clamped to simple-septate ........................................................... Furtadoella

3b. Hyphal system di-trimitic in context, generative hyphae clamped .......................................................................................................... 4

4a. Exospore wall incomplete and smooth, endospore wall with two longitudinal crests and transverse membranes .................. Haddowia

4b. Exospore wall complete and uneven to foveolate or verrucose, endospore wall with obvious spinules ................................................ 5

5a. Endospore wall with hollow spinules which persist until exospore wall forming holes ...................................................... Foraminispora

5b. Endospore wall with solid spinules, exospore wall verrucose ................................................................................................................ 6

6a. Basidiospores globose to oblong ...................................................................................................................................... Amauroderma

6b. Basidiospores ellipsoid to ovoid ............................................................................................................................................................. 7

304

Sun et al.

Amauroderma Murrill, Bull. Torrey Bot. Club 32: 366. 1905.

MycoBank MB 17052.

Type species: Amauroderma schomburgkii (Mont. & Berk.) Torrend.

For a detailed description of Amauroderma, see Costa-Rezende et

al. (2016) and Sun et al. (2020).

Notes: The Amauroderma clade is composed of species from the

Neotropics. According to Costa-Rezende et al. (2020a), 24 species

of Amauroderma have been recorded from the Neotropics, 16

species with available DNA sequences were included in the current

phylogenetic analyses. Besides these species, this genus contains

40 taxa which have been recorded from Africa, Southeast Asia and

Central America, and the sequences of these taxa are not available.

Until now, 58 species (Table 2) can be recognised in Amauroderma

based on previous literature (Furtado 1967b, Steyaert 1972,

Corner 1983, Henao-M 1997, Moncalvo & Ryvarden 1997, Gulaid

& Ryvarden 1998, Aime et al. 2003, Ryvarden 2004b, Gomes-Silva

et al. 2015, Ryvarden 2020).

Amaurodermellus Costa-Rezende et al., Mycol. Prog. 19: 727.

2020. MycoBank MB 833561.

Type species: Amaurodermellus ovisporum (Gomes-Silva et al.)

Costa-Rezende et al.

For a detailed description of Amaurodermellus, see Costa-Rezende

et al. (2020b).

Notes: Amaurodermellus was established by Costa-Rezende et

al. (2020b) with type species, Amaurodermellus ovisporum. It can

be distinguished from the other genera in Ganodermataceae by

ovoid basidiospores with inconspicuous spinules on the endospore

wall. Several species in Ganoderma also have ovoid basidiospores

such as G. sichuanense, but Amaurodermellus ovisporum has a

dark dull pileal surface and non-truncated basidiospores which

is similar to Amauroderma. In this study, the taxonomic status of

Amaurodermellus was further conrmed by multi-gene based

phylogenetic analyses (Fig. 1).

Cristataspora Robledo & Costa-Rezende, Mycol. Prog. 19: 733.

2020. MycoBank MB 833558.

Type species: Cristataspora coffeata (Berk.) Robledo et al.

For a detailed description of Cristataspora, see Costa-Rezende et

al. (2020b).

Notes: Cristataspora coffeata as the only species in Cristataspora

was previously placed in Humphreya due to the reticulate or

disjointed crests on the endospore wall (Steyaert 1972). Costa-

Rezende et al. (2020b) examined the specimens of C. coffeata

collected from neotropical areas, and the vertical or transverse

ridges on the endospore wall observed under SEM rendered it

distinct from Humphreya. Therefore, Cristataspora was established

as a new genus based on its different basidiospores and

independent clade in the phylogenetic analysis (Fig. 1).

Foraminispora Robledo et al., Persoonia 39: 258. 2017. MycoBank

MB 819015.

Type species: Foraminispora rugosa (Berk.) Costa-Rezende et al.

For a detailed description of Foraminispora, see Costa-Rezende et

al. (2017) and Sun et al. (2020).

Notes: Foraminispora is characterised by the unique ultrastructure

of its basidiospores, which shows an uneven exospore wall with

holes caused by hollow and columnar spinules on the endospore

wall. In this study, Foraminispora is recognised as an independent

clade including ve species with high support (Fig. 1). Species of

this genus were reported from East Asia and Neotropics, and the

descriptions of these species can be found in Costa-Rezende et al.

(2017) and Sun et al. (2020).

Furtadoella B.K. Cui & Y.F. Sun, gen. nov. MycoBank MB 840977.

Diagnosis: Differs from other genera by its soft basidiomata, white

context, monomitic hyphal system in context, with both clamped

and simple-septate generative hyphae.

7a. Basidiomata woody hard, with short stipe or sessile .............................................................................................................. Magoderna

7b. Basidiomata corky, with long stipe ............................................................................................................................... Amaurodermellus

8a. Basidiomata soft to eshy when fresh .................................................................................................................................................... 9

8b. Basidiomata soft corky to woody hard when fresh ............................................................................................................................... 10

9a. Hyphal system dimitic, generative hyphae branched .......................................................................................................... Tomophagus

9b. Hyphal system trimitic, generative hyphae unbranched .................................................................................................. Trachydermella

10a. Endospore wall with spiny ornamentation .......................................................................................................................................... 11

10b. Endospore wall with ridged ornamentation ........................................................................................................................................ 12

11a. Pore dissepiments thin, context cream; exospore wall uneven to foveolate ................................................................ Sinoganoderma

11b. Pore dissepiments thick, context pale white to dark brown; exospore wall verrucose to vermicular ................................... Ganoderma

12a. Basidiomata sessile to subsessile; basidiospores inconspicuously truncated .............................................................. Neoganoderma

12b. Basidiomata stipitate; basidiospores conspicuously truncated .......................................................................................................... 13

13a. Context white; endospore wall with vertical or transverse ridges ..................................................................................... Cristataspora

13b. Context honey; endospore wall with reticular ridges ........................................................................................................... Humphreya

305www.studiesinmycology.org

Ganodermataceae from China

Ganoderma eickeri Dai 12595

Ganoderma acontextum sp. nov. JV 0611/21G (TYPE)

Ganoderma casuarinicola Dai 19678

Ganoderma gibbosum Cui 14338

Ganoderma angustisporum Dai 19603

Ganoderma fornicatum TNM-F0010592

Ganoderma nasalanense CACP17060211 (TYPE)

Ganoderma fornicatum BCRC35374

Ganoderma fallax sp. nov. JV 0509/93K

Ganoderma alpinum sp. nov. Cui 18402

Ganoderma podocarpense JV 1504/126

Ganoderma knysnamense CMW47755 (TYPE)

Ganoderma subangustisporum sp. nov. Cui 18592 (TYPE)

Ganoderma sinense Cui 14526

Ganoderma angustisporum Cui 18240

Ganoderma subangustisporum sp. nov. Cui 18597

Ganoderma calidophilum MFLU 19-2219

Ganoderma mutabile Dai 20414

Ganoderma williamsianum Dai 16809

Ganoderma hochiminhense Dai 18488

Ganoderma dunense CMW42157 (TYPE)

Ganoderma enigmaticum Dai 15971

Ganoderma yunlingense sp. nov. Cui 17958

Ganoderma ryvardenii HKAS 58053 (TYPE)

Ganoderma lobatum JV 1008/32

Ganoderma mirabile Cui 18283

Ganoderma guangxiense sp. nov. Cui 14508

Ganoderma knysnamense CMW49688

Ganoderma australe DHCR417

Ganoderma orbiforme Cui 13880

Ganoderma tornatum TBG01AM2009

Ganoderma gibbosum Cui 17254

Ganoderma orbiforme Cui 18326

Ganoderma mirabile Cui 18271

Ganoderma mastoporum TNM-F0018835

Ganoderma subangustisporum sp. nov. Cui 18593

Ganoderma subellipsoideum sp. nov. Cui 18241

Ganoderma thailandicum HKAS 104641

Ganoderma mbrekobenum UMN7-4GHA

Ganoderma adspersum Dai 13191

Ganoderma gibbosum KUMCC17-0005

Ganoderma adspersum HSBU-200894

Ganoderma destructans CMW43670 (TYPE)

Ganoderma casuarinicola Dai 16337

Ganoderma cocoicola sp. nov. Cui 16792

Ganoderma thailandicum HKAS 104640 (TYPE)

Ganoderma sinense Dai 20079

Ganoderma sinense Cui 14461

Ganoderma ecuadorense JV 1808/85

Ganoderma subellipsoideum sp. nov. Cui 18327

Ganoderma guangxiense sp. nov. Cui 14453 (TYPE)

Ganoderma eickeri CMW50325

Ganoderma alpinum sp. nov. Cui 17325

Ganoderma japonicum Gja-1

Ganoderma williamsianum Dai 19611

Ganoderma enigmaticum Dai 15970

Ganoderma ryvardenii HKAS 58054

Ganoderma gibbosum Cui 17769

Ganoderma cupreum GanoTK7

Ganoderma enigmaticum CMW43669 (TYPE)

Ganoderma australe DHCR411

Ganoderma chocoense QCAM 3123 (TYPE)

Ganoderma cocoicola sp. nov. Cui 16791 (TYPE)

Ganoderma tongshanense sp. nov. Cui 17168 (TYPE)

Ganoderma acontextum sp. nov. JV 1407/64

Ganoderma gibbosum KUMCC17-0003

Ganoderma sublobatum sp. nov. Cui 16806

Ganoderma boninense WD 2085

Ganoderma hochiminhense Cui 18229

Ganoderma casuarinicola Dai 19470

Ganoderma calidophilum MFLU 19-2174

Ganoderma orbiforme Cui 18302

Ganoderma nasalanense CACP17060212

Ganoderma orbiforme Cui 18301

Ganoderma neojaponicum 36073

Ganoderma neojaponicum AS5.541 (TYPE)

Ganoderma ellipsoideum Dai 19683

Ganoderma zonatum FL-02

Ganoderma mirabile Cui 16408

Ganoderma subellipsoideum sp. nov. Cui 18325 (TYPE)

Ganoderma eickeri CMW49692 (TYPE)

Ganoderma orbiforme Cui 18317

Ganoderma ellipsoideum Dai 20544

Ganoderma aridicola GanoTK25

Ganoderma fallax sp. nov. JV 0709/39

Ganoderma eickeri Dai 12598

Ganoderma zonatum FL-03

Ganoderma acontextum sp. nov. JV 1208/11J

Ganoderma gibbosum Cui 17780

Ganoderma williamsianum Dai 20553

Ganoderma aridicola Dai 12588 (TYPE)

Ganoderma cupreum GanoTK4

Ganoderma applanatum Cui 14062

Ganoderma pfeifferi CBS 221.48

Ganoderma orbiforme Cui 13891

Ganoderma yunlingense sp. nov. Cui 17043

Ganoderma aridicola GanoTK01

Ganoderma hochiminhense MFLU 19-2224

Ganoderma yunlingense sp. nov. Cui 16288 (TYPE)

Ganoderma ellipsoideum CMW 14080966 (TYPE)

Ganoderma pfeifferi Dai 12153

Ganoderma boninense WD 2028

Ganoderma alpinum sp. nov. Cui 17467 (TYPE)

Ganoderma casuarinicola Dai 16336 (TYPE)

Ganoderma mutabile Cui 17189

Ganoderma mirabile Cui 18237

Ganoderma pfeifferi Dai 12683

Ganoderma mbrekobenum UMN7-3GHA (TYPE)

Ganoderma japonicum AS569

Ganoderma tornatum URM82776

Ganoderma ecuadorense Dai 17397

Ganoderma mastoporum K15-86

Ganoderma brownii JV 1105/9J

Ganoderma mutabile Yuan 2289 (TYPE)

Ganoderma ellipsoideum MFLU 19-2221

Ganoderma gibbosum Cui 13940

Ganoderma guangxiense sp. nov. Cui 14454

Ganoderma destructans Dai 16431

Ganoderma dunense CMW42149

Ganoderma sublobatum sp. nov. Cui 16804 (TYPE)

Ganoderma hochiminhense MFLU 19-2225

Ganoderma podocarpense QCAM 6422 (TYPE)

Ganoderma ryvardenii HKAS 58055

Ganoderma fallax sp. nov. JV 1009/27 (TYPE)

Ganoderma williamsianum Dai 17790

Ganoderma subangustisporum sp. nov. Cui 18596

Ganoderma applanatum Cui 14070

Ganoderma angustisporum Cui 13817 (TYPE)

Ganoderma ecuadorense Dai 17418

Ganoderma brownii JV 0709/109

Ganoderma lobatum JV 1008/31

70/-

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52/-

100/1.00

Subc

lade VI-dull

Ganoderma

dull Subclade I-laccate/dull

Subclade II-laccate

Subclade III-laccate

Subclade IV-dull

laccate

Subclade V-laccate/dull

Fig. 1. Maximum Likelihood analyses of Ganodermataceae based on dataset of ITS + nLSU + rpb2 + tef1 + mtSSU + nSSU. Maximum Likelihood bootstrap

values higher than 50 % and Bayesian posterior probabilities values more than 0.95 are shown. New species are in bold. Ganoderma clade is divided by

laccate or dull pileal surface.

306

Sun et al.

Fig. 1. (Continued).

Ganoderma weberianum Dai 19689

Ganoderma concinnum Robledo 3192

Cristataspora coffeata Robledo 3183

Foraminispora concentrica Cui 12648

Ganoderma ravenelii 151FL

Ganoderma oregonense CBS 265.88

Ganoderma curtisii CBS 100132

Ganoderma shanxiense Dai 18921

Ganoderma tropicum Dai 20029

Cristataspora coffeata 1504/50

Ganoderma resinaceum LGAM 448

Ganoderma platense BAFC384

Ganoderma parvulum URM83345

Ganoderma shanxiense Cui 14565

Ganoderma multipileum Dai 19690

Ganoderma flexipes Cui 13863

Ganoderma ravenelii MS187FL

Ganoderma tropicum Dai 16434

Ganoderma platense BAFC2374

Ganoderma mizoramense UMN-MZ5

Ganoderma bubalinomarginatum sp. nov. Dai 20075 (TYPE)

Ganoderma tropicum Dai 19491

Foraminispora rugosa DHCR512

Ganoderma multiplicatum URM83346

Foraminispora concentrica Cui 17141

Ganoderma myanmarense MFLU 19-2167

Ganoderma chalceum URM80457

Ganoderma carnosum LGAM 1642

Ganoderma curtisii CBS 100131

Ganoderma subflexipes sp. nov. Cui 17247

Ganoderma weberianum CBS 219.36

Ganoderma lingzhi Cui 9166

Ganoderma parvulum URM83344

Foraminispora rugosa DHCR560

Foraminispora yunnanensis Cui 7974

Ganoderma multipileum Dai 19691

Ganoderma leucocontextum Dai15601

Ganoderma sp. N.3

Ganoderma sanduense SA18012502

Ganoderma hoehnelianum Cui 13982

Ganoderma myanmarense MFLU 19-2211/2169

Ganoderma castaneum sp. nov. Cui 17283 (TYPE)

Foraminispora austrosinensis Cui 14318

Ganoderma castaneum sp. nov. Cui 13893

Ganoderma lingzhi Dai 20895

Ganoderma lucidum Dai 20017

Ganoderma magniporum Dai 19966

Ganoderma resinaceum MS1212

Ganoderma lingzhi Cui 18167

Cristataspora flavipora G299

Ganoderma wiiroense MIN 938704 (TYPE)

Ganoderma multiplicatum SPC9

Ganoderma flexipes Dai 20461

Ganoderma lucidum Cui 14405

Ganoderma bubalinomarginatum sp. nov. Dai 20074

Ganoderma meredithae UMNFL64

Ganoderma lucidum Cui 14404

Ganoderma acaciicola sp. nov.Cui 16813

Foraminispora rugosa URM86888

Foraminispora concentrica Cui 12644 (TYPE)

Ganoderma multipileum Cui 13597

Ganoderma subflexipes sp. nov. Cui 17258

Ganoderma hoehnelianum Dai 20783

Ganoderma steyaertianum 6-WN-15(M)-A

Ganoderma multiplicatum CC8

Ganoderma subflexipes sp. nov. Cui 17257 (TYPE)

Ganoderma stipitatum THC 16

Ganoderma weberianum CBS 128581

Ganoderma sp. N.1

Foraminispora concentrica Cui 16238

Ganoderma castaneum sp. nov. Dai 16500

Foraminispora yinggelingensis Cui 13618 (TYPE)

Ganoderma chuxiongense sp. nov. Cui 17262 (TYPE)

Ganoderma austroafricanum CBS 138724

Ganoderma carnosum MUCL 49464

Ganoderma philippii Cui 14443

Ganoderma steyaertianum 6-WN-16(M)-A

Ganoderma shanxiense BJTC FM423 (TYPE)

Ganoderma resinaceum MS1211

Ganoderma tuberculosum Dai 17412

Ganoderma carocalcareum DMC 322 (TYPE)

Ganoderma weixiense HKAS 100649 (TYPE)

Ganoderma acaciicola sp. nov. Cui 16814

Ganoderma oregonense CBS 266.88

Ganoderma sichuanense Cui 16343

Cristataspora coffeata Robledo 3182

Ganoderma resinaceum LGAM 462

Ganoderma polychromum UMNOR3

Ganoderma sessile JV 1209/27

Ganoderma tsugae Cui 14110

Ganoderma flexipes Cui 13841

Ganoderma sichuanense Dai 19651

Ganoderma castaneum sp. nov. Dai 13710

Ganoderma hoehnelianum Cui 13904

Ganoderma weberianum Cui 16359

Ganoderma acaciicola sp. nov. Cui 16815 (TYPE)

Ganoderma sichuanense HMAS42798 (TYPE)

Ganoderma tropicum Dai 19679

Ganoderma magniporum Zhou 439

Foraminispora yinggelingensis Cui 13630

Ganoderma mizoramense JZ8

Ganoderma lucidum K 175217

Ganoderma tsugae Cui 14112

Ganoderma weberianum Dai 19682

Ganoderma sanduense SA18012501 (TYPE)

Ganoderma mizoramense UMN-MZ4 (TYPE)

Ganoderma wiiroense UMN-20-GHA

Ganoderma lingzhi Wu 1006-38 (TYPE)

Ganoderma weixiense HKAS 100650

Ganoderma polychromum MS343OR

Ganoderma lingzhi Cui 18161

Ganoderma tuberculosum JV 1607/62

Ganoderma weberianum Dai 19673

Ganoderma carocalcareum DMC 513

Ganoderma mexicanum MUCL 55832

Ganoderma lucidum MT 26/10

Ganoderma leucocontextum GDGM 40200 (TYPE)

Foraminispora austrosinensis Cui 16425

Foraminispora rugosa JV 1608/889-ND

Cristataspora coffeata FLOR 50933

Ganoderma martinicense LIP SW-Mart08-55 (TYPE)

Foraminispora yunnanensis Yuan 2253

Ganoderma multipileum Dai 17569

Ganoderma multiplicatum Dai 17395

Ganoderma concinnum Robledo 3235

Ganoderma sessile Dai 16403

Ganoderma mexicanum MUCL 49453

Ganoderma puerense sp. nov. Dai 20427 (TYPE)

Ganoderma martinicense He 2240

Ganoderma philippii Cui 14444

Ganoderma shanxiense HSA 539

Ganoderma meredithae UMNFL50

Ganoderma nitidum JV 1504/73

98/1.00

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86/0.99

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52/-

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58/-

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100/1.00

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100/1.00

Cristataspora

Foraminispora

dull

Subc

lade VII-laccate/dull

Subc

lade VIII-laccate

Subc

lade IX-laccate

Subc

lade X-laccate

307www.studiesinmycology.org

Ganodermataceae from China

Fig. 1. (Continued).

0.03

Amauroderma cf. schomburgkii URM89271

Sanguinoderma guangdongense sp. nov. Cui 17259 (TYPE)

Sanguinoderma elmerianum Dai 20634

Magoderna subresinosum Dai 18626

Amauroderma sp. URM89239

Amauroderma schomburgkii URM89225

Sanguinoderma melanocarpum sp. nov. Dai 18603 (TYPE)

Sanguinoderma perplexum Wei 5562

Amauroderma partitum URM83039

Perenniporia subtephropora Dai10962 (TYPE)

Amauroderma intermedium GAS910

Tomophagus cattienensis CT119

Sinoganoderma shandongense comb. nov. Dai 15787

Tomophagus colossus URM80450

Amauroderma praetervisum JV 1407/40

Sanguinoderma microporum Cui 13851 (TYPE)

Neoganoderma neurosporum comb. nov. GAS1013

Trachydermella tsunodae comb. nov. WD2034

Tomophagus cattienensis Dai 18487

Sanguinoderma microporum Cui 14022

Amauroderma subsessile URM89293

Sanguinoderma melanocarpum sp. nov. Dai 18512

Amauroderma elegantissimum Dai 17431

Sanguinoderma bataaense Cui 6285

Amaurodermellus ovisporum DHCR546

Sanguinoderma infundibulare sp. nov. Cui 17248

Sanguinoderma elmerianum HMAS 133187

Amauroderma floriformum URM83250 (TYPE)

Sinoganoderma shandongense comb. nov. Dai 20244

Amauroderma aurantiacum FLOR 52205

Sanguinoderma tricolor sp. nov. Cui 18242

Amauroderma aurantiacum DHCR540

Amauroderma robledoi URM84230

Sanguinoderma infundibulare sp. nov. Dai 18151

Amauroderma calcigenum URM86847

Sanguinoderma bataaense Zhou 153

Sanguinoderma rugosum Cui 8795

Amauroderma omphalodes JV 1909/23-J

Amauroderma camerarium FLOR 52216

Amaurodermellus ovisporum DHCR539

Sanguinoderma tricolor sp. nov. Cui 18292 (TYPE)

Sanguinoderma longistipitum sp. nov. Dai 20696 (TYPE)

Amauroderma omphalodes DHCR500

Sanguinoderma flavovirens Cui 16935 (TYPE)

Amauroderma schomburgkii JV 1908/9

Sanguinoderma elmerianum Cui 18234

Sanguinoderma infundibulare sp. nov. Dai 18148

Sanguinoderma elmerianum Dai 20503

Amauroderma calcigenum URM83864

Sanguinoderma elmerianum Cui 8940

Sanguinoderma rude MEL 2317411

Sanguinoderma rude DHCR457

Haddowia longipes LPDR17072708

Sanguinoderma laceratum A5

Sinoganoderma shandongense comb. nov. xsd08085

Sanguinoderma bataaense Dai 10746

Amauroderma camerarium FLOR 52169

Sanguinoderma microsporum sp. nov. Cui 13901

Amauroderma intermedium JV 1312/E14-J

Sanguinoderma guangdongense sp. nov. Dai 16724

Sanguinoderma sinuosum MEL 2341763 (TYPE)

Sanguinoderma rugosum Cui 9011

Amauroderma calcigenum FLOR 52315

Sanguinoderma tricolor sp. nov. Dai 18574

Sanguinoderma infundibulare sp. nov. Cui 17238

Furtadoella brasiliensis comb. nov. TBG58

Sanguinoderma perplexum Dai 10811

Amauroderma calcitum FLOR 50931

Furtadoella brasiliensis comb. nov. JV 1909/75

Sanguinoderma longistipitum sp. nov. Dai 13891

Amauroderma calcigenum URM89213

Sanguinoderma microporum Cui 18270

Amaurodermellus ovisporum DHCR547

Amauroderma laccatostipitatum URM89240

Sanguinoderma perplexum Cui 6554

Amauroderma calcigenum URM89566

Sanguinoderma rugosum Cui 9012

Perenniporia subtephropora Dai10964

Sanguinoderma microsporum sp. nov. Dai 16726 (TYPE)

Sanguinoderma rugosum Cui 9066

Sanguinoderma guangdongense sp. nov. Cui 17240

Amauroderma calcigenum JV 1808/51

Amauroderma laccatostipitatum HFSL ACGS7

Sanguinoderma rude Cui 16592

Amauroderma calcitum FLOR 52230 (TYPE)

Sanguinoderma infundibulare sp. nov. URM 450213

Sinoganoderma shandongense comb. nov. Dai 15785

Amauroderma robledoi FLOR 52249

Sinoganoderma shandongense comb. nov. xsd08032

Sanguinoderma bataaense Dai 7862

Amauroderma cf. schomburgkii JV 1908/39

Sanguinoderma sinuosum MEL 2366586

Sanguinoderma reniforme Cui 16511 (TYPE)

Amauroderma exile URM89226

Amauroderma cf. schomburgkii URM89272

Sanguinoderma perplexum Cui 6496

Magoderna subresinosum Cui 18280

Sanguinoderma microsporum sp. nov. Cui 13897

Haddowia macropora sp. nov. JV 1908/46 (TYPE)

Neoganoderma neurosporum comb. nov. DHCR559

Sinoganoderma shandongense comb. nov. Dai 15791

Amauroderma omphalodes DHCR499

Amauroderma partitum URM82882

Sanguinoderma longistipitum sp. nov. Dai 16635

Sanguinoderma rugosum Dai 20582

Furtadoella brasiliensis comb. nov. URM83578

Amauroderma elegantissimum URM83822

Haddowia longipes LPDR17072709

Sanguinoderma guangdongense sp. nov. Dai 20419

Sanguinoderma longistipitum sp. nov. Cui 13903

Amaurodermellus ovisporum DHCR127

Amauroderma praetervisum URM89233

Amauroderma schomburgkii URM83228

Tomophagus colossus URM83330

Sanguinoderma laceratum Cui 8155 (TYPE)

Sinoganoderma shandongense comb. nov. Dai 20243

Amauroderma exile URM82794

Sanguinoderma infundibulare sp. nov. Cui 17256

Trachydermella tsunodae comb. nov. GR363

Amauroderma intermedium FLOR 52248

Sanguinoderma infundibulare sp. nov. Dai 18149 (TYPE)

Amauroderma pseudoboletum FLOR 52318

Magoderna subresinosum Cui 18262

Amauroderma robledoi URM87687

Tomophagus cattienensis CT99 (TYPE)

Amauroderma subsessile URM89294

Furtadoella biseptata comb. nov. FLOR50932 (TYPE)

100/1.00

99/1.00

100/1.00

96/1.00

80/0.99

100/1.00

95/1.00

50/-

100/1.00

81/0.99

99/-

100/1.00

85/-

89/0.99

100/1.00

92/1.00

70/-

86/-

100/1.00

95/1.00

94/1.00

96/0.99

91/0.99

96/1.00

97/1.00

68/-

100/1.00

94/1.00

98/1.00

52/0.99

87/1.00

100/1.00

78/1.00

83/0.99

86/1.00

80/0.97

100/1.00

100/0.99

89/0.98

94/1.00

100/1.00

99/1.00

85/0.99

78/0.99

88/0.98

100/1.00

85/0.99

58/-

60/-

51/-

76/-

Sinoganoderma gen. nov.

Amauroderma

Trachydermella gen. nov.

Furtadoella gen. nov.

Sanguinoderma

Amaurodermellus

100/1.00

Tomophagus

Haddowia

Magoderna

Neoganoderma gen. nov.

308

Sun et al.

Ganoderma adspersum Dai 13191

Ganoderma zonatum FL-03

Ganoderma hochiminhense Dai 18488

Ganoderma mutabile Dai 20414

Ganoderma casuarinicola Dai 19470

Ganoderma australe DHCR417

Amauroderma aurantiacum DHCR540

Ganoderma gibbosum Cui 17769

Amaurodermellus ovisporum DHCR546

Ganoderma acontextum sp. nov. JV 1407/64

Ganoderma mbrekobenum UMN7-3GHA (TYPE)

Ganoderma brownii JV 1105/9J

Amaurodermellus ovisporum DHCR547

Amauroderma calcigenum URM86847

Trachydermella tsunodae comb. nov. GR363

Ganoderma yunlingense sp. nov. Cui 16288 (TYPE)

Ganoderma ryvardenii HKAS 58054

Ganoderma ellipsoideum MFLU 19-2221

Amauroderma pseudoboletum FLOR 52318

Ganoderma subangustisporum sp. nov. Cui 18592

Ganoderma acontextum sp. nov. JV 1208/11J

Ganoderma mirabile Cui 18237

Ganoderma podocarpense QCAM 6422 (TYPE)

Ganoderma orbiforme Cui 13891

Ganoderma gibbosum Cui 17254

Ganoderma boninense WD 2028

Ganoderma orbiforme Cui 13880

Ganoderma hochiminhense MFLU 19-2225

Ganoderma brownii JV 0709/109

Ganoderma fallax sp. nov. JV 0709/39

Ganoderma applanatum Cui 14062

Ganoderma neojaponicum AS5.541 (TYPE)

Amauroderma calcigenum JV 1808/51

Ganoderma mastoporum TNM-F001883

Ganoderma australe DHCR411

Ganoderma sublobatum sp. nov. Cui 16806

Ganoderma mirabile Cui 18283

Ganoderma pfeifferi Dai 12683

Ganoderma nasalanense CACP17060211 (TYPE)

Ganoderma gibbosum Cui 13940

Ganoderma aridicola Dai 12588 (TYPE)

Ganoderma enigmaticum Dai 15970

Ganoderma cocoicola sp. nov. Cui 16791 (TYPE)

Ganoderma mutabile Yuan 2289 (TYPE)

Amauroderma praetervisum JV 1407/40

Ganoderma cupreum GanoTK7

Ganoderma mirabile Cui 18271

Amaurodermellus ovisporum DHCR127

Ganoderma calidophilum MFLU 19-2219

Ganoderma alpinum sp. nov. Cui 18402

Ganoderma fallax sp. nov. JV 0509/93K

Ganoderma subangustisporum sp. nov. Cui 18593

Ganoderma casuarinicola Dai 16337

Ganoderma ecuadorense JV 1808/85

Ganoderma japonicum Gja-1

Ganoderma fallax sp. nov. JV 1009/27 (TYPE)

Ganoderma orbiforme Cui 18301

Ganoderma nasalanense CACP17060212

Ganoderma gibbosum KUMCC17-0005

Ganoderma enigmaticum CMW43669 (TYPE)

Ganoderma sinense Cui 14526

Ganoderma lobatum JV 1008/31

Ganoderma ellipsoideum Dai 19683

Ganoderma orbiforme Cui 18302

Ganoderma eickeri CMW49692 (TYPE)

Ganoderma gibbosum KUMCC17-0003

Ganoderma angustisporum Cui 18240

Ganoderma alpinum sp. nov. Cui 17467 (TYPE)

Ganoderma gibbosum Cui 14338

Trachydermella tsunodae comb. nov. WD2034

Ganoderma ryvardenii HKAS 58055

Ganoderma japonicum AS569

Ganoderma aridicola GanoTK01

Ganoderma chocoense QCAM 3123 (TYPE)

Ganoderma guangxiense sp. nov. Cui 14508

Ganoderma sinense Dai 20079

Ganoderma eickeri Dai 12595

Ganoderma tornatum TBG01AM2009

Amaurodermellus ovisporum DHCR539

Ganoderma enigmaticum Dai 15971

Ganoderma gibbosum Cui 17780

Ganoderma ryvardenii HKAS 58053 (TYPE)

Ganoderma adspersum HSBU-200894

Ganoderma ellipsoideum Dai 20544

Ganoderma ecuadorense Dai 17418

Ganoderma pfeifferi Dai 12153

Amauroderma calcigenum URM89566

Ganoderma subangustisporum sp. nov. Cui 18597

Ganoderma thailandicum HKAS 104640 (TYPE)

Ganoderma zonatum FL-02

Ganoderma cupreum GanoTK4

Ganoderma angustisporum Dai 19603

Ganoderma knysnamense CMW47755 (TYPE)

Ganoderma thailandicum HKAS 104641

Ganoderma alpinum sp. nov. Cui 17325

Amauroderma calcigenum URM83864

Ganoderma casuarinicola Dai 19678

Ganoderma cocoicola sp. nov. Cui 16792

Ganoderma tornatum URM82776

Ganoderma hochiminhense MFLU 19-2224

Ganoderma applanatum Cui 14070

Ganoderma fornicatum BCRC35374

Ganoderma ecuadorense Dai 17397

Ganoderma guangxiense sp. nov. Cui 14454

Ganoderma guangxiense sp. nov. Cui 14453 (TYPE)

Ganoderma lobatum JV 1008/32

Ganoderma fornicatum TNM-F0010592

Ganoderma orbiforme Cui 18317

Ganoderma yunlingense sp. nov. Cui 17958

Ganoderma ellipsoideum CMW 14080966 (TYPE)

Ganoderma calidophilum MFLU 19-2174

Ganoderma tongshanense sp. nov. Cui 17168 (TYPE)

Amauroderma robledoi URM87687

Ganoderma aridicola GanoTK25

Ganoderma subangustisporum sp. nov. Cui 18596

Ganoderma neojaponicum 36073

Amauroderma robledoi FLOR 52249

Ganoderma boninense WD 2085

Ganoderma eickeri CMW50325

Ganoderma hochiminhense Cui 18229

Ganoderma mutabile Cui 17189

Ganoderma sinense Cui 14461

Ganoderma podocarpense JV 1504/126

Amauroderma robledoi URM84230

Amauroderma aurantiacum FLOR 52205

Ganoderma casuarinicola Dai 16336 (TYPE)

Ganoderma orbiforme Cui 18326

Ganoderma yunlingense sp. nov. Cui 17043

Amauroderma praetervisum URM89233

Ganoderma angustisporum Cui 13817 (TYPE)

Ganoderma mastoporum K15-86

Ganoderma acontextum sp. nov. JV 0611/21G (TYPE)

Ganoderma eickeri Dai 12598

Ganoderma mbrekobenum UMN7-4GHA

Ganoderma knysnamense CMW49688

Ganoderma sublobatum sp. nov. Cui 16804 (TYPE)

100

73 100

100

Fig. 2. Maximum Likelihood analyses of Ganodermataceae based on dataset of ITS. Maximum Likelihood bootstrap values higher than 50 % are shown.

New species are in bold.

309www.studiesinmycology.org

Ganodermataceae from China

Fig. 2. (Continued).

Amauroderma elegantissimum URM83822

Ganoderma wiiroense UMN-20-GHA

Ganoderma platense BAFC2374

Ganoderma destructans Dai 16431

Ganoderma flexipes Cui 13841

Amauroderma omphalodes DHCR499

Ganoderma stipitatum THC 16

Ganoderma multipileum Cui 13597

Ganoderma wiiroense MIN 938704 (TYPE)

Ganoderma concinnum Robledo 3192

Ganoderma mexicanum MUCL 49453

Ganoderma weberianum Dai 19689

Ganoderma weberianum Dai 19682

Amauroderma intermedium GAS910

Amauroderma calcitum FLOR 50931

Ganoderma myanmarense MFLU 19-2211/2169

Ganoderma lingzhi Wu 1006-38 (TYPE)

Amauroderma intermedium FLOR 52248

Amauroderma cf. schomburgkii URM89272

Amauroderma calcitum FLOR 52230 (TYPE)

Ganoderma mizoramense JZ8

Ganoderma sichuanense Cui 16343

Ganoderma multipileum Dai 17569

Ganoderma austroafricanum CBS 138724

Amauroderma calcigenum URM89213

Amauroderma omphalodes JV 1909/23-J

Amauroderma camerarium FLOR 52169

Ganoderma parvulum URM83345

Ganoderma dunense CMW42149

Amauroderma laccatostipitatum URM89240

Ganoderma tuberculosum Dai 17412

Amauroderma subsessile URM89294

Amauroderma exile URM82794

Ganoderma ravenelii MS187FL

Ganoderma mizoramense UMN-MZ4 (TYPE)

Amauroderma sp. URM89239

Amauroderma cf. schomburgkii JV 1908/39

Ganoderma mizoramense UMN-MZ5

Ganoderma bubalinomarginatum sp. nov. Dai 20075 (TYPE)

Amauroderma elegantissimum Dai 17431

Amauroderma cf. schomburgkii URM89271

Ganoderma castaneum sp. nov.Cui 17283 (TYPE)

Ganoderma curtisii CBS 100132

Ganoderma philippii Cui 14443

Ganoderma hoehnelianum Cui 13982

Ganoderma multipileum Dai 19691

Ganoderma polychromum UMNOR3

Ganoderma multiplicatum SPC9

Ganoderma lingzhi Cui 18161

Ganoderma hoehnelianum Cui 13904

Ganoderma tropicum Dai 19679

Ganoderma bubalinomarginatum sp. nov. Dai 20074

Ganoderma acaciicola sp. nov.Cui 16815 (TYPE)

Ganoderma resinaceum LGAM 448

Ganoderma sessile Dai 16403

Ganoderma subflexipes sp. nov.Cui 17257 (TYPE)

Ganoderma ravenelii 151FL

Ganoderma tropicum Dai 16434

Ganoderma weberianum Dai 19673

Ganoderma martinicense He 2240

Ganoderma meredithae UMNFL64

Ganoderma nitidum JV 1504/73

Ganoderma philippii Cui 14444

Ganoderma carocalcareum DMC 513

Ganoderma subflexipes sp. nov.Cui 17247

Ganoderma carocalcareum DMC 322 (TYPE)

Ganoderma lingzhi Cui 9166

Amauroderma calcigenum FLOR 52315

Ganoderma tropicum Dai 19491

Ganoderma resinaceum LGAM 462

Amauroderma omphalodes DHCR500

Amauroderma intermedium JV 1312/E14-J

Ganoderma castaneum sp. nov.Dai 13710

Ganoderma steyaertianum 6-WN-15(M)-A

Ganoderma meredithae UMNFL50

Ganoderma sessile JV 1209/27

Ganoderma platense BAFC384

Ganoderma multiplicatum URM83346

Ganoderma mexicanum MUCL 55832

Ganoderma flexipes Dai 20461

Ganoderma weberianum CBS 128581

Amauroderma floriformum URM83250 (TYPE)

Ganoderma multiplicatum Dai 17395

Ganoderma lingzhi Dai 20895

Amauroderma schomburgkii URM83228

Ganoderma curtisii CBS 100131

Amauroderma laccatostipitatum HFSL ACGS7

Ganoderma resinaceum MS1211

Ganoderma acaciicola sp. nov.Cui 16814

Ganoderma multipileum Dai 19690

Ganoderma castaneum sp. nov.Dai 16500

Amauroderma subsessile URM89293

Ganoderma myanmarense MFLU 19-2167

Ganoderma tuberculosum JV 1607/62

Ganoderma steyaertianum 6-WN-16(M)-A

Ganoderma lingzhi Cui 18167

Amauroderma schomburgkii URM89225

Amauroderma exile URM89226

Ganoderma destructans CMW43670 (TYPE)

Ganoderma sichuanense HMAS42798 (TYPE)

Ganoderma dunense CMW42157 (TYPE)

Ganoderma acaciicola sp. nov.Cui 16813

Ganoderma sichuanense Dai 19651

Ganoderma flexipes Cui 13863

Ganoderma martinicense LIP SW-Mart08-55 (TYPE)

Ganoderma weberianum CBS 219.36

Ganoderma parvulum URM83344

Ganoderma resinaceum MS1212

Amauroderma camerarium FLOR 52216

Amauroderma schomburgkii JV 1908/9

Ganoderma tropicum Dai 20029

Ganoderma castaneum sp. nov.Cui 13893

Ganoderma concinnum Robledo 3235

Amauroderma partitum URM82882

Amauroderma partitum URM83039

Ganoderma multiplicatum CC8

Ganoderma polychromum MS343OR

Ganoderma chalceum URM80457

100

97 100

100

57 100

100

310

Sun et al.

Fig. 2. (Continued).

0.03

Sanguinoderma guangdongense sp. nov. Cui 17259 (TYPE)

Furtadoella brasiliensis comb. nov. JV 1909/75

Ganoderma shanxiense BJTC FM423 (TYPE)

Sanguinoderma bataaense Dai 7862

Sanguinoderma bataaense Cui 6285

Ganoderma lucidum Cui 14405

Ganoderma williamsianum Dai 19611

Ganoderma lucidum K 175217

Perenniporia subtephropora Dai10964

Sanguinoderma microsporum sp. nov. Cui 13897

Sanguinoderma rude Cui 16592

Sanguinoderma infundibulare sp. nov. Dai 18149 (TYPE)

Ganoderma lucidum Dai 20017

Foraminispora rugosa JV 1608/889-ND

Cristataspora coffeata Robledo 3182

Sanguinoderma rugosum Dai 20582

Sanguinoderma infundibulare sp. nov. URM 450213

Foraminispora yunnanensis Yuan 2253

Tomophagus cattienensis Dai 18487

Sanguinoderma sinuosum MEL 2366586

Ganoderma magniporum Zhou 439

Sinoganoderma shandongense comb. nov. Dai 20244

Ganoderma shanxiense Cui 14565

Cristataspora coffeata 1504/50

Sanguinoderma perplexum Cui 6496

Tomophagus cattienensis CT119

Ganoderma williamsianum Dai 16809

Haddowia longipes LPDR17072709

Sanguinoderma guangdongense sp. nov. Dai 20419

Sanguinoderma melanocarpum sp. nov. Dai 18603 (TYPE)

Cristataspora flavipora G299

Sanguinoderma laceratum Cui 8155 (TYPE)

Sanguinoderma bataaense Dai 10746

Ganoderma lucidum MT 26/10

Magoderna subresinosum Cui 18262

Ganoderma tsugae Cui 14110

Haddowia macropora sp. nov. JV 1908/46 (TYPE)

Sanguinoderma microsporum sp. nov. Cui 13901

Ganoderma oregonense CBS 266.88

Magoderna subresinosum Dai 18626

Ganoderma carnosum LGAM 1642

Sanguinoderma laceratum A5

Sanguinoderma rugosum Cui 9012

Foraminispora concentrica Cui 12644 (TYPE)

Cristataspora coffeata Robledo 3183

Sanguinoderma perplexum Cui 6554

Ganoderma sanduense SA18012502

Ganoderma sanduense SA18012501 (TYPE)

Sanguinoderma longistipitum sp. nov. Dai 20696 (TYPE)

Magoderna subresinosum Cui 18280

Cristataspora coffeata FLOR 50933

Foraminispora rugosa URM86888

Perenniporia subtephropora Dai10962 (TYPE)

Sanguinoderma perplexum Wei 5562

Sanguinoderma infundibulare sp. nov. Cui 17248

Sanguinoderma elmerianum Cui 8940

Ganoderma sp. N.3

Sanguinoderma longistipitum sp. nov. Cui 13903

Tomophagus cattienensis CT99 (TYPE)

Furtadoella biseptata comb. nov. FLOR50932 (TYPE)

Foraminispora rugosa DHCR512

Tomophagus colossus URM80450

Foraminispora yunnanensis Cui 7974

Ganoderma tsugae Cui 14112

Sanguinoderma infundibulare sp. nov. Cui 17238

Sinoganoderma shandongense comb. nov. xsd08032

Sanguinoderma elmerianum Cui 18234

Sinoganoderma shandongense comb. nov. Dai 15785

Foraminispora rugosa DHCR560

Ganoderma leucocontextum GDGM 40200 (TYPE)

Sanguinoderma rude DHCR457

Sanguinoderma rugosum Cui 9011

Sanguinoderma infundibulare sp. nov. Cui 17256

Sanguinoderma microporum Cui 14022

Ganoderma williamsianum Dai 17790

Ganoderma shanxiense Dai 18921

Ganoderma weixiense HKAS 100650

Sanguinoderma microporum Cui 13851 (TYPE)

Foraminispora yinggelingensis Cui 13630

Sanguinoderma rude MEL 2317411

Ganoderma weixiense HKAS 100649 (TYPE)

Sanguinoderma rugosum Cui 8795

Sanguinoderma rugosum Cui 9066

Ganoderma shanxiense HSA 539

Foraminispora yinggelingensis Cui 13618 (TYPE)

Foraminispora concentrica Cui 12648

Ganoderma leucocontextum Dai15601

Foraminispora austrosinensis Cui 16425

Ganoderma oregonense CBS 265.88

Foraminispora austrosinensis Cui 14318

Haddowia longipes LPDR17072708

Tomophagus colossus URM83330

Sanguinoderma guangdongense sp. nov. Dai 16724

Ganoderma sp. N.1

Sanguinoderma reniforme Cui 16511 (TYPE)

Sanguinoderma bataaense Zhou 153

Sinoganoderma shandongense comb. nov. Dai 15787

Neoganoderma neurosporum comb. nov. DHCR559

Sanguinoderma perplexum Dai 10811

Sanguinoderma longistipitum sp. nov. Dai 13891

Sanguinoderma elmerianum HMAS 133187

Ganoderma williamsianum Dai 20553

Sanguinoderma tricolor sp. nov. Cui 18242

Foraminispora concentrica Cui 17141

Sanguinoderma longistipitum sp. nov. Dai 16635

Sanguinoderma melanocarpum sp. nov. Dai 18512

Neoganoderma neurosporum comb. nov. GAS1013

Sanguinoderma tricolor sp. nov. Dai 18574

Sinoganoderma shandongense comb. nov. Dai 15791

Sanguinoderma elmerianum Dai 20503

Ganoderma lucidum Cui 14404

Furtadoella brasiliensis comb. nov. TBG58

Sinoganoderma shandongense comb. nov. xsd08085

Sanguinoderma elmerianum Dai 20634

Furtadoella brasiliensis comb. nov. URM83578

Sanguinoderma sinuosum MEL 2341763 (TYPE)

Ganoderma dianzhongense sp. nov. Cui 17262 (TYPE)

Sanguinoderma infundibulare sp. nov. Dai 18148

Foraminispora concentrica Cui 16238

Ganoderma carnosum MUCL 49464

Sanguinoderma infundibulare sp. nov. Dai 18151

100

77 100

100

311www.studiesinmycology.org

Ganodermataceae from China

Fig. 3. Maximum Likelihood analyses of Ganodermataceae based on dataset of nLSU. Maximum Likelihood bootstrap values higher than 50 % are shown.

New species are in bold.

Ganoderma hoehnelianum Cui 13982

Ganoderma subangustisporum sp. nov. Cui 18592 (TYPE)

Ganoderma casuarinicola Dai 16337

Ganoderma myanmarense MFLU 19-2167

Ganoderma mbrekobenum UMN7-3GHA (TYPE)

Ganoderma tornatum TBG01AM2009

Ganoderma subflexipes sp. nov. Cui 17247

Ganoderma nasalanense CACP17060211 (TYPE)

Ganoderma australe DHCR411

Ganoderma resinaceum LGAM 448

Ganoderma guangxiense sp. nov. Cui 14508

Ganoderma flexipes Dai 20461

Ganoderma guangxiense sp. nov. Cui 14454

Sanguinoderma laceratum Cui 8155 (TYPE)

Ganoderma bubalinomarginatum sp. nov. Dai 20074

Ganoderma pfeifferi Dai 12153

Ganoderma williamsianum Dai 20553

Ganoderma tropicum Dai 19679

Ganoderma subellipsoideum sp. nov. Cui 18327

Ganoderma gibbosum KUMCC17-0005

Ganoderma weberianum Dai 19689

Ganoderma sinense Cui 14461

Ganoderma castaneum sp. nov. Dai 13710

Ganoderma enigmaticum CMW43669 (TYPE)

Ganoderma bubalinomarginatum sp. nov. Dai 20075 (TYPE)

Ganoderma subangustisporum sp. nov. Cui 18596

Ganoderma mirabile Cui 18283

Ganoderma gibbosum Cui 17254

Ganoderma weberianum Dai 19682

Ganoderma tropicum Dai 16434

Ganoderma flexipes Cui 13863

Ganoderma acaciicola sp. nov. Cui 16813

Ganoderma nasalanense CACP17060212

Ganoderma ecuadorense JV 1808/85

Ganoderma ecuadorense Dai 17418

Ganoderma enigmaticum Dai 15970

Ganoderma subflexipes sp. nov. Cui 17258

Ganoderma hoehnelianum Cui 13904

Ganoderma acaciicola sp. nov. Cui 16814

Ganoderma guangxiense sp. nov. Cui 14453 (TYPE)

Ganoderma tongshanense sp. nov. Cui 17168 (TYPE)

Ganoderma mirabile Cui 18271

Ganoderma angustisporum Cui 13817 (TYPE)

Ganoderma tornatum URM82776

Ganoderma williamsianum Dai 16809

Ganoderma multiplicatum CC8

Ganoderma gibbosum KUMCC17-0003

Ganoderma subangustisporum sp. nov. Cui 18593

Ganoderma multiplicatum URM83346

Ganoderma acaciicola sp. nov. Cui 16815 (TYPE)

Ganoderma boninense WD 2028

Ganoderma ellipsoideum Dai 19683

Ganoderma sichuanense Cui 16343

Ganoderma multipileum Dai 19690

Ganoderma sessile Dai 16403

Ganoderma sinense Cui 14526

Ganoderma multiplicatum SPC9

Ganoderma hochiminhense Cui 18229

Ganoderma cocoicola sp. nov. Cui 16791 (TYPE)

Ganoderma orbiforme Cui 18301

Ganoderma hochiminhense MFLU 19-2224

Ganoderma pfeifferi Dai 12683

Ganoderma multipileum Cui 13597

Ganoderma ecuadorense Dai 17397

Ganoderma subellipsoideum sp. nov. Cui 18325 (TYPE)

Ganoderma multipileum Dai 19691

Ganoderma subellipsoideum sp. nov. Cui 18241

Ganoderma hochiminhense Dai 18488

Ganoderma australe DHCR417

Ganoderma gibbosum Cui 13940

Ganoderma hochiminhense MFLU 19-2225

Ganoderma thailandicum HKAS 104641

Ganoderma puerense sp. nov. Dai 20427 (TYPE)

Ganoderma wiiroense UMN-20-GHA

Ganoderma subangustisporum sp. nov. Cui 18597

Ganoderma ellipsoideum Dai 20544

Ganoderma flexipes Cui 13841

Ganoderma weberianum CBS 219.36

Ganoderma sichuanense Dai 19651

Ganoderma orbiforme Cui 13880

Ganoderma williamsianum Dai 17790

Ganoderma thailandicum HKAS 104640 (TYPE)

Ganoderma shanxiense Dai 18921

Ganoderma orbiforme Cui 13891

Ganoderma orbiforme Cui 18326

Ganoderma myanmarense MFLU 19-2211/2169

Ganoderma hoehnelianum Dai 20783

Ganoderma eickeri Dai 12595

Ganoderma casuarinicola Dai 16336 (TYPE)

Ganoderma wiiroense MIN 938704 (TYPE)

Ganoderma casuarinicola Dai 19678

Ganoderma castaneum sp. nov. Cui 13893

Ganoderma sublobatum sp. nov. Cui 16806

Ganoderma mutabile Dai 20414

Ganoderma weberianum Dai 19673

Ganoderma parvulum URM83344

Ganoderma parvulum URM83345

Ganoderma sinense Dai 20079

Ganoderma lingzhi Dai 20895

Ganoderma orbiforme Cui 18302

Ganoderma philippii Cui 14444

Ganoderma angustisporum Dai 19603

Ganoderma mirabile Cui 18237

Ganoderma williamsianum Dai 19611

Ganoderma mirabile Cui 16408

Ganoderma resinaceum LGAM 462

Ganoderma weberianum CBS 128581

Ganoderma philippii Cui 14443

Ganoderma mbrekobenum UMN7-4GHA

Ganoderma cocoicola sp. nov. Cui 16792

Ganoderma enigmaticum Dai 15971

Ganoderma subflexipes sp. nov. Cui 17257 (TYPE)

Ganoderma multipileum Dai 17569

Ganoderma pfeifferi CBS 221.48

Ganoderma eickeri Dai 12598

Ganoderma mizoramense UMN-MZ5

Ganoderma orbiforme Cui 18317

Ganoderma austroafricanum CBS 138724

Ganoderma weberianum Cui 16359

Ganoderma angustisporum Cui 18240

Ganoderma gibbosum Cui 14338

100

312

Sun et al.

Fig. 3. (Continued).

0.009

Amauroderma subsessile URM89294

Sanguinoderma elmerianum Dai 20503

Ganoderma applanatum Cui 14070

Amauroderma schomburgkii URM83228

Sanguinoderma tricolor sp. nov. Cui 18292 (TYPE)

Foraminispora concentrica Cui 17141

Sanguinoderma longistipitum sp. nov. Dai 20696 (TYPE)

Sanguinoderma longistipitum sp. nov. Dai 13891

Furtadoella biseptata comb. nov. FLOR50932 (TYPE)

Amauroderma aurantiacum FLOR 52205

Sanguinoderma elmerianum Cui 8940

Trachydermella tsunodae comb. nov. WD2034

Sanguinoderma rude Cui 16592

Sanguinoderma melanocarpum sp. nov. Dai 18603 (TYPE)

Sanguinoderma elmerianum HMAS 133187

Haddowia longipes LPDR17072708

Cristataspora coffeata Robledo 3183

Ganoderma podocarpense QCAM6422

Amauroderma sp. URM89239

Amauroderma omphalodes JV 1909/23-J

Amauroderma schomburgkii URM89225

Sanguinoderma elmerianum Dai 20634

Ganoderma applanatum Cui 14062

Sanguinoderma rude DHCR457

Amauroderma subsessile URM89293

Amauroderma calcitum FLOR 50931

Sinoganoderma shandongense comb. nov. Dai 15791

Amauroderma laccatostipitatum URM89240

Ganoderma yunlingense sp. nov. Cui 17043

Magoderna subresinosum Cui 18280

Ganoderma concinnum Robledo 3235

Sanguinoderma microsporum sp. nov. Cui 13897

Ganoderma magniporum Dai 19966

Ganoderma carnosum LGAM 1642

Furtadoella brasiliensis comb. nov. JV 1909/75

Sanguinoderma infundibulare sp. nov. Dai 18148

Cristataspora flavipora G299

Sanguinoderma rugosum Dai 20582

Ganoderma lucidum Cui 14405

Sanguinoderma infundibulare sp. nov. Dai 18149 (TYPE)

Ganoderma magniporum Zhou 439

Ganoderma chalceum URM80457

Sanguinoderma longistipitum sp. nov. Dai 16635

Tomophagus colossus URM80450

Cristataspora coffeata Robledo 3182

Sanguinoderma infundibulare sp. nov. URM 450213

Foraminispora concentrica Cui 12644 (TYPE)

Sinoganoderma shandongense comb. nov. Dai 15787

Amauroderma cf. schomburgkii JV 1908/39

Sanguinoderma reniforme Cui 16511 (TYPE)

Foraminispora rugosa DHCR560

Sanguinoderma elmerianum Cui 18234

Foraminispora yinggelingensis Cui 13630

Sanguinoderma infundibulare sp. nov. Cui 17248

Foraminispora austrosinensis Cui 16425

Amauroderma praetervisum URM89233

Amauroderma cf. schomburgkii URM89271

Magoderna subresinosum Dai 18626

Sanguinoderma microsporum sp. nov. Cui 13901

Amauroderma intermedium JV 1312/E14-J

Amauroderma aurantiacum DHCR540

Sanguinoderma guangdongense sp. nov. Dai 16724

Sanguinoderma longistipitum sp. nov. Cui 13903

Amauroderma intermedium FLOR 52248

Sanguinoderma rugosum Cui 8795

Sinoganoderma shandongense comb. nov. Dai 20244

Ganoderma tuberculosum JV 1607/62

Amauroderma elegantissimum Dai 17431

Amauroderma omphalodes DHCR499

Foraminispora concentrica Cui 12648

Sanguinoderma bataaense Cui 6285

Ganoderma leucocontextum Dai15601

Amauroderma elegantissimum URM83822

Sanguinoderma rugosum Cui 9011

Perenniporia subtephropora Dai10962 (TYPE)

Sanguinoderma tricolor sp. nov. Cui 18242

Haddowia longipes LPDR17072709

Sanguinoderma guangdongense sp. nov. Cui 17240

Amauroderma calcigenum JV 1808/51

Sanguinoderma microporum Cui 14022

Ganoderma weixiense HKAS 100649 (TYPE)

Tomophagus colossus URM83330

Amauroderma calcigenum URM89213

Amauroderma robledoi URM87687

Sanguinoderma sinuosum MEL 2341763 (TYPE)

Sanguinoderma perplexum Dai 10811

Sanguinoderma bataaense Dai 10746

Foraminispora austrosinensis Cui 14318

Sanguinoderma perplexum Cui 6496

Perenniporia subtephropora Dai10964

Ganoderma carnosum MUCL 49464

Sanguinoderma microporum Cui 13851 (TYPE)

Foraminispora rugosa URM86888

Ganoderma yunlingense sp. nov. Cui 16288 (TYPE)

Ganoderma concinnum Robledo 3192

Foraminispora yinggelingensis Cui 13618 (TYPE)

Foraminispora yunnanensis Cui 7974

Amauroderma exile URM89226

Haddowia macropora sp. nov. JV 1908/46 (TYPE)

Amaurodermellus ovisporum DHCR547

Sanguinoderma microporum Cui 18270

Sanguinoderma melanocarpum sp. nov. Dai 18512

Sanguinoderma rugosum Cui 9012

Ganoderma lucidum Cui 14404

Sanguinoderma flavovirens Cui 16935 (TYPE)

Sanguinoderma sinuosum MEL 2366586

Amauroderma cf. schomburgkii URM89272

Amauroderma calcigenum URM89566

Sanguinoderma guangdongense sp. nov. Cui 17259 (TYPE)

Amauroderma omphalodes DHCR500

Sanguinoderma rugosum Cui 9066

Sanguinoderma microsporum sp. nov. Dai 16726 (TYPE)

Sanguinoderma guangdongense sp. nov. Dai 20419

Sinoganoderma shandongense comb. nov. Dai 20243

Foraminispora concentrica Cui 16238

Sanguinoderma perplexum Cui 6554

Ganoderma lucidum Dai 20017

Sanguinoderma tricolor sp. nov. Dai 18574

Sinoganoderma shandongense comb. nov. Dai 15785

Magoderna subresinosum Cui 18262

Ganoderma weixiense HKAS 100650

100

80 100

78 74

72 89

100

313www.studiesinmycology.org

Ganodermataceae from China

Fig. 4. Maximum Likelihood analyses of Ganodermataceae based on dataset of rpb2. Maximum Likelihood bootstrap values higher than 50 % are shown.

New species are in bold.

Ganoderma ellipsoideum Dai 20544

Ganoderma gibbosum Cui 14338

Ganoderma meredithae UMNFL64

Ganoderma tropicum Dai 19679

Ganoderma mirabile Cui 18271

Ganoderma orbiforme Cui 13891

Ganoderma shanxiense Dai 18921

Ganoderma ecuadorense Dai 17397

Ganoderma subflexipes sp. nov.Cui 17258

Ganoderma brownii JV 0709/109

Ganoderma boninense WD 2028

Ganoderma boninense WD 2085

Ganoderma multiplicatum Dai 17395

Ganoderma lingzhi Dai 20895

Ganoderma thailandicum HKAS 104641

Ganoderma casuarinicola Dai 16337

Ganoderma brownii JV 1105/9J

Ganoderma tropicum Dai 20029

Ganoderma acontextum sp. nov. JV 0611/21G (TYPE)

Ganoderma williamsianum Dai 16809

Ganoderma subangustisporum sp. nov. Cui 18597

Ganoderma shanxiense BJTC FM423 (TYPE)

Ganoderma zonatum FL-02

Ganoderma thailandicum HKAS 104640 (TYPE)

Ganoderma williamsianum Dai 20553

Ganoderma sinense Cui 14526

Ganoderma applanatum Cui 14062

Ganoderma sinense Cui 14461

Ganoderma gibbosum Cui 17254

Ganoderma lingzhi Wu 1006-38 (TYPE)

Ganoderma martinicense He 2240

Ganoderma mirabile Cui 18283

Ganoderma shanxiense HSA 539

Ganoderma lobatum JV 1008/31

Ganoderma acontextum sp. nov. JV 1407/64

Ganoderma castaneum sp. nov. Dai 16500

Ganoderma orbiforme Cui 13880

Ganoderma acaciicola sp. nov. Cui 16813

Ganoderma adspersum Dai 13191

Ganoderma guangxiense sp. nov. Cui 14453 (TYPE)

Ganoderma flexipes Cui 13841

Ganoderma acaciicola sp. nov. Cui 16814

Ganoderma acontextum sp. nov. JV 1208/11J

Ganoderma tropicum Dai 19491

Ganoderma cocoicola sp. nov. Cui 16791 (TYPE)

Ganoderma ecuadorense Dai 17418

Ganoderma angustisporum Dai 19603

Ganoderma enigmaticum Dai 15970

Ganoderma hochiminhense Cui 18229

Ganoderma zonatum FL-03

Ganoderma multipileum Dai 17569

Ganoderma sinense Dai 20079

Ganoderma casuarinicola Dai 16336 (TYPE)

Ganoderma castaneum sp. nov. Cui 13893

Ganoderma subflexipes sp. nov. Cui 17257 (TYPE)

Ganoderma philippii Cui 14444

Ganoderma mutabile Dai 20414

Ganoderma gibbosum Cui 13940

Ganoderma lobatum JV 1008/32

Ganoderma acaciicola sp. nov. Cui 16815 (TYPE)

Ganoderma curtisii CBS 100132

Ganoderma mirabile Cui 18237

Ganoderma sublobatum sp. nov. Cui 16804 (TYPE)

Ganoderma applanatum Cui 14070

Ganoderma angustisporum Cui 18240

Ganoderma flexipes Dai 20461

Ganoderma williamsianum Dai 19611

Ganoderma casuarinicola Dai 19470

Ganoderma subflexipes sp. nov. Cui 17247

Ganoderma curtisii CBS 100131

Ganoderma angustisporum Cui 13817 (TYPE)

Ganoderma tropicum Dai 16434

Ganoderma cocoicola sp. nov. Cui 16792

Ganoderma enigmaticum Dai 15971

Ganoderma ravenelii MS187FL

Ganoderma podocarpense JV 1504/126

Ganoderma hochiminhense Dai 18488

Ganoderma flexipes Cui 13863

Ganoderma philippii Cui 14443

Ganoderma lingzhi Cui 9166

Ganoderma guangxiense sp. nov. Cui 14454

Ganoderma multipileum Cui 13597

100

100 100

100

100 100

100

94 99

100

314

Sun et al.

Fig. 4. (Continued).

0.03

Ganoderma weberianum Dai 19682

Ganoderma bubalinomarginatum sp. nov. Dai 20074

Sanguinoderma flavovirens Cui 16935 (TYPE)

Haddowia macropora sp. nov. JV 1908/46 (TYPE)

Ganoderma tsugae Cui 14112

Ganoderma weberianum CBS 219.36

Sanguinoderma sinuosum MEL 2366586

Perenniporia subtephropora Dai10964

Ganoderma austroafricanum CBS 138724

Foraminispora concentrica Cui 12644 (TYPE)

Sanguinoderma rude Cui 16592

Sanguinoderma microporum Cui 13851 (TYPE)

Amauroderma subsessile URM89294

Amauroderma cf. schomburgkii URM89271

Foraminispora concentrica Cui 17141

Ganoderma sessile Dai 16403

Sinoganoderma shandongense comb. nov. Dai 15791

Ganoderma hoehnelianum Cui 13982

Sanguinoderma infundibulare sp. nov. Cui 17238

Sanguinoderma rude MEL 2317411

Trachydermella tsunodae comb. nov. WD2034

Sanguinoderma infundibulare sp. nov. Dai 18151

Ganoderma magniporum Dai 19966

Magoderna subresinosum Dai 18626

Ganoderma resinaceum LGAM 462

Foraminispora concentrica Cui 12648

Sanguinoderma perplexum Cui 6496

Ganoderma weberianum Cui 16359

Ganoderma sichuanense Dai 19651

Sanguinoderma longistipitum sp. nov. Dai 16635

Ganoderma oregonense CBS 266.88

Ganoderma lucidum Cui 14404

Ganoderma weberianum Dai 19673

Tomophagus cattienensis Dai 18487

Amauroderma sp. URM89239

Ganoderma sichuanense Cui 16343

Ganoderma oregonense CBS 265.88

Sanguinoderma microporum Cui 14022

Sanguinoderma sinuosum MEL 2341763 (TYPE)

Amauroderma cf. schomburgkii JV 1908/39

Ganoderma mexicanum MUCL 49453

Sanguinoderma perplexum Dai 10811

Ganoderma weberianum CBS 128581

Ganoderma mexicanum MUCL 55832

Ganoderma bubalinomarginatum sp. nov. Dai 20075 (TYPE)

Sanguinoderma bataaense Cui 6285

Sanguinoderma guangdongense sp. nov. Cui 17259 (TYPE)

Amauroderma praetervisum URM89233

Ganoderma hoehnelianum Dai 20783

Ganoderma leucocontextum Dai15601

Sanguinoderma bataaense Dai 10746

Sanguinoderma perplexum Cui 6554

Ganoderma sessile JV 1209/27

Foraminispora rugosa URM86888

Ganoderma puerense sp. nov. Dai 20427 (TYPE)

Sanguinoderma rugosum Cui 9012

Amauroderma elegantissimum Dai 17431

Furtadoella brasiliensis comb. nov. JV 1909/75

Sanguinoderma reniforme Cui 16511 (TYPE)

Amauroderma calcigenum URM89213

Ganoderma lucidum Cui 14405

Sanguinoderma guangdongense sp. nov. Dai 16724

Sanguinoderma infundibulare sp. nov. Dai 18149 (TYPE)

Amauroderma subsessile URM89293

Sanguinoderma longistipitum sp. nov. Cui 13903

Amauroderma calcigenum URM89566

Foraminispora yinggelingensis Cui 13618 (TYPE)

Ganoderma lucidum K 175217

Amauroderma laccatostipitatum URM89240

Sanguinoderma guangdongense sp. nov.Cui 17240

Sinoganoderma shandongense comb. nov. Dai 15785

Amauroderma omphalodes JV 1909/23-J

Sanguinoderma rugosum Cui 9011

Amauroderma calcigenum JV 1808/51

Sinoganoderma shandongense comb. nov. Dai 15787

Sanguinoderma rugosum Cui 8795

Sanguinoderma infundibulare sp. nov. Cui 17256

Sanguinoderma tricolor sp. nov. Cui 18242

Ganoderma resinaceum LGAM 448

Sanguinoderma melanocarpum sp. nov. Dai 18603 (TYPE)

Amauroderma cf. schomburgkii URM89272

Amauroderma schomburgkii URM89225

Sanguinoderma tricolor sp. nov. Dai 18574

Perenniporia subtephropora Dai10962 (TYPE)

Sanguinoderma rugosum Dai 20582

Amauroderma elegantissimum URM83822

Ganoderma hoehnelianum Cui 13904

Sanguinoderma guangdongense sp. nov. Dai 20419

Amauroderma robledoi URM87687

Ganoderma tsugae Cui 14110

Sanguinoderma infundibulare sp. nov. Dai 18148

100

100 100

100

92 99

100

315www.studiesinmycology.org

Ganodermataceae from China

Fig. 5. Maximum Likelihood analyses of Ganodermataceae based on dataset of tef1. Maximum Likelihood bootstrap values higher than 50 % are shown.

New species are in bold.

Ganoderma subangustisporum sp. nov. Cui 18593

Ganoderma brownii JV 0709/109

Ganoderma boninense WD 2085

Ganoderma tropicum Dai 20029

Ganoderma knysnamense CMW47755 (TYPE)

Ganoderma casuarinicola Dai 16336 (TYPE)

Ganoderma zonatum FL-02

Ganoderma flexipes Dai 20461

Ganoderma subellipsoideum sp. nov. Cui 18325 (TYPE)

Ganoderma boninense WD 2028

Ganoderma curtisii CBS 100131

Ganoderma orbiforme Cui 18326

Ganoderma tropicum Dai 16434

Ganoderma flexipes Cui 13841

Ganoderma orbiforme Cui 13880

Ganoderma multiplicatum Dai 17395

Ganoderma lingzhi Dai 20895

Ganoderma casuarinicola Dai 16337

Ganoderma mirabile Cui 18283

Ganoderma multipileum Dai 17569

Ganoderma australe DHCR411

Ganoderma orbiforme Cui 18302

Ganoderma podocarpense JV 1504/126

Ganoderma tropicum Dai 19491

Ganoderma weberianum Dai 19682

Ganoderma pfeifferi Dai 12153

Ganoderma guangxiense sp. nov. Cui 14508

Ganoderma williamsianum Dai 17790

Ganoderma mirabile Cui 18237

Ganoderma pfeifferi Dai 12683

Ganoderma hochiminhense Cui 18229

Ganoderma ecuadorense Dai 17418

Ganoderma thailandicum HKAS 104641

Ganoderma eickeri Dai 12598

Ganoderma ecuadorense JV 1808/85

Ganoderma subellipsoideum sp. nov. Cui 18241

Ganoderma casuarinicola Dai 19470

Ganoderma tuberculosum JV 1607/62

Ganoderma lobatum JV 1008/31

Ganoderma thailandicum HKAS 104640 (TYPE)

Ganoderma cocoicola sp. nov. Cui 16792

Ganoderma subangustisporum sp. nov. Cui 18592 (TYPE)

Ganoderma weberianum Dai 19673

Ganoderma castaneum sp. nov. Dai 16500

Ganoderma casuarinicola Dai 19678

Ganoderma flexipes Cui 13863

Ganoderma ecuadorense Dai 17397

Ganoderma adspersum Dai 13191

Ganoderma applanatum Cui 14062

Ganoderma aridicola Dai 12588 (TYPE)

Ganoderma dunense CMW42149

Ganoderma acontextum sp. nov. JV 0611/21G (TYPE)

Ganoderma ellipsoideum Dai 20544

Ganoderma sinense Cui 14461

Ganoderma yunlingense sp. nov. Cui 17043

Ganoderma tropicum Dai 19679

Ganoderma gibbosum Cui 13940

Ganoderma subflexipes sp. nov. Cui 17257 (TYPE)

Ganoderma adspersum HSBU-200894

Ganoderma ravenelii MS187FL

Ganoderma knysnamense CMW49688

Ganoderma orbiforme Cui 18301

Ganoderma applanatum Cui 14070

Ganoderma multipileum Cui 13597

Ganoderma curtisii CBS 100132

Ganoderma weberianum Cui 16359

Ganoderma multipileum Dai 19690

Ganoderma weberianum Dai 19689

Ganoderma williamsianum Dai 19611

Ganoderma acontextum sp. nov.JV 1407/64

Ganoderma hochiminhense Dai 18488

Ganoderma australe DHCR417

Ganoderma guangxiense sp. nov. Cui 14454

Ganoderma acontextum sp. nov. JV 1208/11J

Ganoderma gibbosum Cui 14338

Ganoderma eickeri Dai 12595

Ganoderma guangxiense sp. nov. Cui 14453 (TYPE)

Ganoderma subflexipes sp. nov. Cui 17258

Ganoderma lingzhi Cui 9166

Ganoderma subangustisporum sp. nov. Cui 18596

Ganoderma angustisporum Cui 13817 (TYPE)

Ganoderma mutabile Cui 17189

Ganoderma philippii Cui 14443

Ganoderma sublobatum sp. nov. Cui 16806

Ganoderma martinicense He 2240

Ganoderma castaneum sp. nov. Dai 13710

Ganoderma mirabile Cui 18271

Ganoderma weberianum CBS 219.36

Ganoderma sinense Dai 20079

Ganoderma sichuanense Dai 19651

Ganoderma gibbosum Cui 17254

Ganoderma angustisporum Cui 18240

Ganoderma sinense Cui 14526

Ganoderma subangustisporum sp. nov. Cui 18597

Ganoderma eickeri CMW49692 (TYPE)

Ganoderma philippii Cui 14444

Ganoderma subellipsoideum sp. nov. Cui 18327

Ganoderma cocoicola sp. nov. Cui 16791 (TYPE)

Ganoderma enigmaticum Dai 15970

Ganoderma sublobatum sp. nov. Cui 16804 (TYPE)

Ganoderma tongshanense sp. nov. Cui 17168 (TYPE)

Ganoderma lingzhi Wu 1006-38 (TYPE)

Ganoderma castaneum sp. nov. Cui 13893

Ganoderma weberianum CBS 128581

Ganoderma mirabile Cui 16408

Ganoderma angustisporum Dai 19603

Ganoderma hochiminhense MFLU 19-2225

Ganoderma zonatum FL-03

Ganoderma eickeri CMW50325

Ganoderma orbiforme Cui 13891

Ganoderma hochiminhense MFLU 19-2224

Ganoderma yunlingense sp. nov. Cui 16288 (TYPE)

Ganoderma dunense CMW42157 (TYPE)

Ganoderma mutabile Dai 20414

Ganoderma brownii JV 1105/9J

Ganoderma subflexipes sp. nov. Cui 17247

Ganoderma orbiforme Cui 18317

Ganoderma williamsianum Dai 16809

Ganoderma enigmaticum Dai 15971

Ganoderma sichuanense Cui 16343

Ganoderma lobatum JV 1008/32

Ganoderma williamsianum Dai 20553

Ganoderma ellipsoideum Dai 19683

100

76 93

316

Sun et al.

Fig. 5. (Continued).

0.04

Sanguinoderma guangdongense sp. nov. Cui 17240

Magoderna subresinosum Dai 18626

Amauroderma cf. schomburgkii URM89272

Foraminispora concentrica Cui 12644 (TYPE)

Sanguinoderma rugosum Dai 20582

Sanguinoderma elmerianum Dai 20503

Cristataspora flavipora G299

Sanguinoderma infundibulare sp. nov. Dai 18148

Sinoganoderma shandongense comb. nov. Dai 20244

Ganoderma resinaceum LGAM 462

Sinoganoderma shandongense comb. nov. Dai 15791

Sanguinoderma bataaense Dai 10746

Sinoganoderma shandongense comb. nov. Dai 15787

Sanguinoderma melanocarpum sp. nov. Dai 18603 (TYPE)

Sanguinoderma perplexum Cui 6554

Ganoderma bubalinomarginatum sp. nov. Dai 20075 (TYPE)

Amauroderma elegantissimum URM83822

Sanguinoderma microporum Cui 14022

Ganoderma lucidum K 175217

Sanguinoderma infundibulare sp. nov. Cui 17238

Amauroderma schomburgkii JV 1908/9

Ganoderma lucidum Cui 14405

Furtadoella brasiliensis comb. nov. JV 1909/75

Sanguinoderma elmerianum Cui 18234

Sanguinoderma microsporum sp. nov. Cui 13901

Amauroderma subsessile URM89294

Ganoderma lucidum Cui 14404

Sanguinoderma elmerianum Dai 20634

Sanguinoderma perplexum Dai 10811

Ganoderma polychromum MS343OR

Sanguinoderma bataaense Cui 6285

Ganoderma resinaceum MS1212

Sanguinoderma infundibulare sp. nov. Dai 18151

Amauroderma laccatostipitatum URM89240

Ganoderma nitidum JV 1504/73

Tomophagus cattienensis Dai 18487

Amauroderma omphalodes JV 1909/23-J

Sanguinoderma rugosum Cui 8795

Ganoderma weixiense HKAS 100649 (TYPE)

Sinoganoderma shandongense comb. nov. Dai 20243

Amauroderma calcigenum JV 1808/51

Sanguinoderma rugosum Cui 9066

Foraminispora yinggelingensis Cui 13618 (TYPE)

Amauroderma elegantissimum Dai 17431

Ganoderma leucocontextum Dai15601

Cristataspora coffeata Robledo 3183

Sanguinoderma microsporum sp. nov. Dai 16726 (TYPE)

Ganoderma shanxiense Cui 14565

Sanguinoderma sinuosum MEL 2341763 (TYPE)

Foraminispora austrosinensis Cui 14318

Sanguinoderma tricolor sp. nov. Cui 18292 (TYPE)

Sanguinoderma infundibulare sp. nov. Dai 18149 (TYPE)

Ganoderma bubalinomarginatum sp. nov. Dai 20074

Sanguinoderma rude MEL 2317411

Sanguinoderma rude DHCR457

Amauroderma praetervisum URM89233

Ganoderma hoehnelianum Cui 13982

Sanguinoderma infundibulare sp. nov. Cui 17256

Sanguinoderma reniforme Cui 16511 (TYPE)

Amauroderma exile URM89226

Sanguinoderma microporum Cui 13851 (TYPE)

Sanguinoderma longistipitum sp. nov. Dai 20696 (TYPE)

Ganoderma resinaceum LGAM 448

Sanguinoderma rude Cui 16592

Perenniporia subtephropora Dai10964

Sinoganoderma shandongense comb. nov. Dai 15785

Ganoderma magniporum Dai 19966

Foraminispora rugosa DHCR560

Sanguinoderma sinuosum MEL 2366586

Sanguinoderma flavovirens Cui 16935 (TYPE)

Ganoderma sessile Dai 16403

Sanguinoderma guangdongense sp. nov. Dai 20419

Sanguinoderma perplexum Cui 6496

Ganoderma oregonense CBS 265.88

Sanguinoderma rugosum Cui 9012

Foraminispora concentrica Cui 16238

Amauroderma schomburgkii URM89225

Magoderna subresinosum Cui 18280

Ganoderma weixiense HKAS 100650

Amauroderma robledoi URM87687

Haddowia macropora sp. nov. JV 1908/46 (TYPE)

Ganoderma hoehnelianum Dai 20783

Foraminispora concentrica Cui 12648

Ganoderma mexicanum MUCL 55832

Amauroderma omphalodes DHCR500

Perenniporia subtephropora Dai10962 (TYPE)

Amauroderma omphalodes DHCR499

Ganoderma lucidum Dai 20017

Ganoderma tsugae Cui 14112

Sanguinoderma longistipitum sp. nov. Dai 16635

Sanguinoderma longistipitum sp. nov. Cui 13903

Sanguinoderma tricolor sp. nov. Dai 18574

Ganoderma shanxiense BJTC FM423 (TYPE)

Ganoderma shanxiense Dai 18921

Sanguinoderma infundibulare sp. nov. Cui 17248

Amauroderma calcigenum URM89566

Ganoderma puerense sp. nov. Dai 20427 (TYPE)

Sanguinoderma microsporum sp. nov. Cui 13897

Amauroderma subsessile URM89293

Amauroderma sp. URM89239

Amauroderma calcigenum URM89213

Cristataspora coffeata 1504/50

Amauroderma cf. schomburgkii URM89271

Sanguinoderma guangdongense sp. nov. Cui 17259 (TYPE)

Amauroderma cf. schomburgkii JV 1908/39

Sanguinoderma rugosum Cui 9011

Sanguinoderma melanocarpum sp. nov. Dai 18512

Ganoderma hoehnelianum Cui 13904

Ganoderma sessile JV 1209/27

Foraminispora austrosinensis Cui 16425

Ganoderma lucidum MT 26/10

Amauroderma intermedium JV 1312/E14-J

Ganoderma resinaceum MS1211

Ganoderma polychromum UMNOR3

Ganoderma tsugae Cui 14110

Sanguinoderma guangdongense sp. nov. Dai 16724

Foraminispora rugosa DHCR512

Ganoderma mexicanum MUCL 49453

Sanguinoderma tricolor sp. nov. Cui 18242

100

9956

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Ganodermataceae from China

Fig. 6. Maximum Likelihood analyses of Ganodermataceae based on dataset of mtSSU. Maximum Likelihood bootstrap values higher than 50 % are shown.

New species are in bold.

Ganoderma multipileum Dai 19690

Ganoderma sessile Dai 16403

Ganoderma orbiforme Cui 18317

Ganoderma multipileum Cui 13597

Ganoderma subangustisporum sp. nov. Cui 18593

Ganoderma aridicola GanoTK25

Foraminispora yinggelingensis Cui 13618 (TYPE)

Ganoderma multiplicatum Dai 17395

Ganoderma tropicum Dai 19491

Ganoderma sublobatum sp. nov.Cui 16806

Ganoderma lingzhi Cui 9166

Ganoderma sichuanense Dai 19651

Ganoderma tuberculosum Dai 17412

Ganoderma weberianum Dai 19689

Ganoderma bubalinomarginatum sp. nov. Dai 20074

Ganoderma lucidum Cui 14404

Ganoderma subangustisporum sp. nov. Cui 18597

Ganoderma puerense sp. nov. Dai 20427 (TYPE)

Ganoderma castaneum sp. nov. Dai 13710

Ganoderma subangustisporum sp. nov. Cui 18592 (TYPE)

Ganoderma hoehnelianum Cui 13904

Ganoderma cupreum GanoTK4

Foraminispora rugosa JV 1608/889-ND

Ganoderma orbiforme Cui 13891

Ganoderma guangxiense sp. nov. Cui 14508

Ganoderma carocalcareum DMC 513

Ganoderma casuarinicola Dai 16336 (TYPE)

Ganoderma applanatum Cui 14062

Ganoderma sublobatum sp. nov. Cui 16804 (TYPE)

Ganoderma gibbosum Cui 13940

Ganoderma chuxiongense sp. nov. Cui 17262 (TYPE)

Ganoderma bubalinomarginatum sp. nov. Dai 20075 (TYPE)

Ganoderma casuarinicola Dai 19678

Ganoderma tropicum Dai 19679

Ganoderma guangxiense sp. nov. Cui 14453 (TYPE)

Ganoderma angustisporum Dai 19603

Ganoderma enigmaticum Dai 15971

Ganoderma mirabile Cui 18271

Ganoderma multipileum Dai 19691

Ganoderma gibbosum Cui 14338

Ganoderma philippii Cui 14443

Ganoderma orbiforme Cui 18326

Sinoganoderma shandongense comb. nov. Dai 15791

Foraminispora concentrica Cui 12644 (TYPE)

Ganoderma aridicola GanoTK01

Ganoderma ellipsoideum Dai 20544

Ganoderma magniporum Zhou 439

Ganoderma leucocontextum Dai15601

Ganoderma flexipes Cui 13841

Ganoderma castaneum sp. nov. Dai 16500

Ganoderma gibbosum Cui 17254

Ganoderma cupreum GanoTK7

Ganoderma orbiforme Cui 18301

Ganoderma castaneum sp. nov. Cui 13893

Foraminispora rugosa URM86888

Ganoderma subellipsoideum sp. nov. Cui 18327

Sinoganoderma shandongense comb. nov. Dai 15787

Ganoderma weberianum Dai 19682

Ganoderma aridicola Dai 12588 (TYPE)

Ganoderma tsugae Cui 14112

Ganoderma mirabile Cui 18237

Ganoderma sichuanense Cui 16343

Foraminispora concentrica Cui 12648

Foraminispora austrosinensis Cui 16425

Ganoderma nitidum JV 1504/73

Sinoganoderma shandongense comb. nov. Dai 15785

Ganoderma weberianum Cui 16359

Foraminispora yinggelingensis Cui 13630

Ganoderma tropicum Dai 16434

Ganoderma multipileum Dai 17569

Ganoderma tropicum Dai 20029

Ganoderma casuarinicola Dai 16337

Ganoderma orbiforme Cui 18302

Ganoderma mutabile Dai 20414

Ganoderma weberianum Dai 19673

Ganoderma orbiforme Cui 13880

Ganoderma ellipsoideum Dai 19683

Ganoderma subellipsoideum sp. nov. Cui 18241

Foraminispora concentrica Cui 17141

Foraminispora austrosinensis Cui 14318

Ganoderma hoehnelianum Dai 20783

Ganoderma angustisporum Cui 13817 (TYPE)

Ganoderma enigmaticum Dai 15970

Ganoderma mirabile Cui 18283

Ganoderma subangustisporum sp. nov. Cui 18596

Ganoderma guangxiense sp. nov. Cui 14454

Foraminispora concentrica Cui 16238

Foraminispora yunnanensis Cui 7974

Ganoderma angustisporum Cui 18240

Ganoderma castaneum sp. nov. Cui 17283 (TYPE)

Tomophagus cattienensis Dai 18487

Ganoderma tuberculosum JV 1607/62

Ganoderma tsugae Cui 14110

318

Sun et al.

Etymology: furtadoella (Lat.), refers to the Dr João Salvador Furtado

who contributed signicantly to the taxonomy of Ganodermataceae.

Type species: Furtadoella biseptata (Costa-Rezende et al.) B.K.

Cui & Y.F. Sun.

Description: Basidiomata annual, stipitate, soft to corky. Pilei

solitary, orbicular to abelliform or infundibuliform. Pileal surface

yellowish brown to greyish brown, dull, glabrous to tomentose,

obviously concentrically zonate. Pore surface white to straw colour;

pores circular to angular; dissepiments thin to thick, entire to

lacerate. Context white to pale brown, with dark resinous lines, soft

corky. Hyphal system dimitic in trama and monomitic in context;

context composed of clamped to simple-septate generative

hyphae, thin- to slightly thick-walled; tubes composed of clamped

generative hyphae and arboriform skeletal hyphae. Basidiospores

subglobose to ellipsoid, colourless, double-walled with verrucose to

reticulate exospore wall, IKI–.

Notes: Furtadoa is an illegitimate name as it is a homonym of one

genus in Araceae and was renamed as Furtadoella in this study.

Furtadoella was described from the Neotropics comprising three

species in Costa-Rezende et al. (2017). In this study one specimen

collected in French Guiana supported the views of Costa-

Rezende et al. (2017) and Sun et al. (2020) in the morphological

and phylogenetic analyses. Under SEM, the ornamentation of

basidiospores in Furtadoella (Fig. 8C) was obviously shown to

have a verrucose to reticulate exospore wall which is similar with

the ultrastructural features of Amauroderma and Trachydermella.

However, Furtadoella can be distinguished from other genera in

Ganodermataceae by its soft basidiomata, a monomitic hyphal

structure in context and non-truncated basidiospores.

Fig. 6. (Continued).

0.006

Sanguinoderma rugosum Cui 9012

Sanguinoderma melanocarpum sp. nov. Dai 18512

Magoderna subresinosum Dai 18626

Amauroderma subsessile URM89293

Amauroderma robledoi URM87687

Sanguinoderma microsporum sp. nov. Cui 13897

Amauroderma omphalodes JV 1909/23-J

Amauroderma laccatostipitatum URM89240

Sanguinoderma tricolor sp. nov. Dai 18574

Sanguinoderma perplexum Dai 10811

Sanguinoderma elmerianum HMAS 133187

Sanguinoderma elmerianum Cui 8940

Sanguinoderma guangdongense sp. nov. Dai 16724

Sanguinoderma perplexum Cui 6554

Sanguinoderma rude MEL 2317411

Sanguinoderma microporum Cui 13851 (TYPE)

Sanguinoderma infundibulare sp. nov. Cui 17248

Sanguinoderma melanocarpum sp. nov. Dai 18603 (TYPE)

Sanguinoderma bataaense Dai 10746

Sanguinoderma rugosum Dai 20582

Sanguinoderma infundibulare sp. nov. Dai 18149 (TYPE)

Perenniporia subtephropora Dai10964

Sanguinoderma microsporum sp. nov. Dai 16726 (TYPE)

Sanguinoderma flavovirens Cui 16935 (TYPE)

Sanguinoderma longistipitum sp. nov. Cui 13903

Furtadoella brasiliensis comb. nov.

JV 1909/75

Sanguinoderma microporum Cui 18270

Sanguinoderma laceratum Cui 8155 (TYPE)

Sanguinoderma longistipitum sp. nov. Dai 16635

Sanguinoderma tricolor sp. nov. Cui 18242

Amauroderma praetervisum URM89233

Amauroderma elegantissimum URM83822

Sanguinoderma longistipitum sp. nov. Dai 20696 (TYPE)

Magoderna subresinosum Cui 18262

Sanguinoderma rugosum Cui 9011

Sanguinoderma perplexum Cui 6496

Sanguinoderma infundibulare sp. nov. Dai 18151

Sanguinoderma rugosum Cui 9066

Sanguinoderma elmerianum Dai 20634

Perenniporia subtephropora Dai10962 (TYPE)

Sanguinoderma infundibulare sp. nov. Cui 17256

Sanguinoderma infundibulare sp. nov. Dai 18148

Amauroderma subsessile URM89294

Sanguinoderma guangdongense sp. nov. Dai 20419

Sanguinoderma infundibulare sp. nov. Cui 17238

Sanguinoderma elmerianum Cui 18234

Haddowia macropora sp. nov. JV 1908/46 (TYPE)

Magoderna subresinosum Cui 18280

Sanguinoderma infundibulare sp. nov. URM 450213

Sanguinoderma sinuosum MEL 2341763 (TYPE)

Amauroderma calcigenum URM89566

Amauroderma sp. URM89239

Amauroderma calcigenum URM89213

Sanguinoderma rugosum Cui 8795

Sanguinoderma guangdongense sp. nov. Cui 17240

Sanguinoderma microporum Cui 14022

Sanguinoderma tricolor sp. nov. Cui 18292 (TYPE)

Sanguinoderma rude Cui 16592

Sanguinoderma longistipitum sp. nov. Dai 13891

Sanguinoderma guangdongense sp. nov. Cui 17259 (TYPE)

Sanguinoderma sinuosum MEL 2366586

Sanguinoderma bataaense Cui 6285

Amauroderma calcigenum JV 1808/51

Sanguinoderma elmerianum Dai 20503

Sanguinoderma microsporum sp. nov. Cui 13901

100

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Ganodermataceae from China

Fig. 7. Maximum Likelihood ML analyses of Ganodermataceae based on dataset of nSSU. Maximum Likelihood bootstrap values higher than 50 % are

shown. New species are in bold.

Sanguinoderma elmerianum Cui 18234

Haddowia macropora sp. nov. JV 1908/46 (TYPE)

Sanguinoderma infundibulare sp. nov. Cui 17238

Sinoganoderma shandongense comb. nov. Dai 15785

Sanguinoderma longistipitum sp. nov. Dai 13891

Sanguinoderma guangdongense sp. nov. Dai 16724

Sanguinoderma perplexum Dai 10811

Ganoderma gibbosum Cui 17254

Ganoderma shanxiense Dai 18921

Magoderna subresinosum Dai 18626

Foraminispora yinggelingensis Cui 13618 (TYPE)

Sanguinoderma longistipitum sp. nov. Dai 16635

Amauroderma calcigenum URM89213

Ganoderma multipileum Dai 17569

Foraminispora austrosinensis Cui 16425

Amauroderma sp. URM89239

Sanguinoderma elmerianum Cui 8940

Sanguinoderma rugosum Cui 8795

Amauroderma cf. schomburgkii URM89271

Sanguinoderma rugosum Cui 9066

Sanguinoderma elmerianum HMAS 133187

Sanguinoderma perplexum Cui 6496

Sanguinoderma rude MEL 2317411

Sanguinoderma flavovirens Cui 16935 (TYPE)

Ganoderma flexipes Dai 20461

Sanguinoderma microporum Cui 13851

Sanguinoderma microsporum sp. nov. Cui 13901

Amauroderma calcigenum JV 1808/51

Foraminispora concentrica Cui 12644 (TYPE)

Amauroderma omphalodes JV 1909/23-J

Sanguinoderma infundibulare sp. nov. Dai 18148

Ganoderma flexipes Cui 13841

Sanguinoderma perplexum Cui 6554

Sanguinoderma tricolor sp. nov. Dai 18574

Sanguinoderma bataaense Cui 6285

Amauroderma schomburgkii URM89225

Amauroderma intermedium JV 1312/E14-J

Sanguinoderma elmerianum Dai 20503

Ganoderma subflexipes sp. nov.Cui 17258

Sanguinoderma rugosum Dai 20582

Sanguinoderma microsporum sp. nov. Dai 16726 (TYPE)

Sanguinoderma guangdongense sp. nov. Cui 17240

Amauroderma laccatostipitatum URM89240

Sanguinoderma rugosum Cui 9012

Ganoderma subflexipes sp. nov. Cui 17257 (TYPE)

Amauroderma cf. schomburgkii URM89272

Amauroderma elegantissimum URM83822

Tomophagus cattienensis Dai 18487

Sanguinoderma elmerianum Dai 20634

Sanguinoderma microsporum sp. nov. Cui 13897

Sanguinoderma infundibulare sp. nov. Dai 18151

Sanguinoderma infundibulare sp. nov. Cui 17248

Foraminispora yinggelingensis Cui 13630

Foraminispora austrosinensis Cui 14318

Sinoganoderma shandongense comb. nov. Dai 15791

Sanguinoderma guangdongense sp. nov. Dai 20419

Amauroderma cf. schomburgkii JV 1908/39

Amauroderma praetervisum URM89233

Amauroderma subsessile URM89293

Sinoganoderma shandongense comb. nov. Dai 15787

Sanguinoderma sinuosum MEL 2341763 (TYPE)

Foraminispora concentrica Cui 12648

Foraminispora concentrica Cui 17141

Magoderna subresinosum Cui 18262

Amauroderma elegantissimum Dai 17431

Sanguinoderma tricolor sp. nov. Cui 18292 (TYPE)

Magoderna subresinosum Cui 18280

Ganoderma castaneum sp. nov. Dai 16500

Sanguinoderma infundibulare sp. nov. URM 450213

Sanguinoderma longistipitum sp. nov. Dai 20696 (TYPE)

Sanguinoderma infundibulare sp. nov. Dai 18149 (TYPE)

Sanguinoderma rude Cui 16592

Sanguinoderma tricolor sp. nov. Cui 18242

Sanguinoderma bataaense Dai 10746

Sanguinoderma infundibulare sp. nov. Cui 17256

Sanguinoderma melanocarpum sp. nov. Dai 18512

Sanguinoderma sinuosum MEL 2366586

Ganoderma multipileum Cui 13597

Amauroderma robledoi URM87687

Ganoderma tropicum Dai 19679

Amauroderma subsessile URM89294

Amauroderma calcigenum URM89566

Ganoderma subflexipes sp. nov. Cui 17247

Ganoderma acaciicola sp. nov. Cui 16814

Sanguinoderma reniforme Cui 16511 (TYPE)

Sanguinoderma guangdongense sp. nov. Cui 17259 (TYPE)

Amauroderma schomburgkii JV 1908/9

Sanguinoderma melanocarpum sp. nov. Dai 18603 (TYPE)

Sanguinoderma microporum Cui 14022

Sanguinoderma microporum Cui 18270

Ganoderma tropicum Dai 19491

Furtadoella brasiliensis comb. nov. JV 1909/75

Sanguinoderma rugosum Cui 9011

Foraminispora concentrica Cui 16238

Amauroderma exile URM89226

Sanguinoderma longistipitum sp. nov. Cui 13903

Ganoderma flexipes Cui 13863

6171

320

Sun et al.

Fig. 7. (Continued).

0.007

Ganoderma sessile Dai 16403

Ganoderma subellipsoideum sp. nov. Cui 18327

Ganoderma williamsianum Dai 16809

Ganoderma guangxiense sp. nov. Cui 14508

Ganoderma hoehnelianum Cui 13904

Ganoderma hochiminhense Cui 18229

Ganoderma sinense Cui 14461

Ganoderma angustisporum Dai 19603

Ganoderma aridicola Dai 12588 (TYPE)

Ganoderma bubalinomarginatum sp. nov. Dai 20074

Ganoderma castaneum sp. nov. Cui 17283 (TYPE)

Ganoderma subellipsoideum sp. nov. Cui 18325 (TYPE)

Ganoderma ellipsoideum Dai 20544

Ganoderma eickeri Dai 12598

Ganoderma guangxiense sp. nov. Cui 14453 (TYPE)

Ganoderma angustisporum Cui 18240

Ganoderma enigmaticum Dai 15971

Ganoderma applanatum Cui 14070

Ganoderma tongshanense sp. nov. Cui 17168 (TYPE)

Ganoderma casuarinicola Dai 16336 (TYPE)

Ganoderma lucidum Dai 20017

Ganoderma weberianum Dai 19673

Ganoderma enigmaticum Dai 15970

Ganoderma tropicum Dai 16434

Ganoderma hoehnelianum Cui 13982

Ganoderma orbiforme Cui 18317

Ganoderma bubalinomarginatum sp. nov. Dai 20075 (TYPE)

Ganoderma orbiforme Cui 18301

Ganoderma tuberculosum Dai 17412

Ganoderma sublobatum sp. nov. Cui 16804 (TYPE)

Ganoderma hoehnelianum Dai 20783

Ganoderma yunlingense sp. nov. Cui 17043

Ganoderma sichuanense Dai 19651

Haddowia longipes LPDR17072708

Ganoderma sichuanense Cui 16343

Haddowia longipes LPDR17072709

Ganoderma ecuadorense Dai 17418

Ganoderma tsugae Cui 14112

Ganoderma orbiforme Cui 18302

Foraminispora rugosa URM86888

Ganoderma leucocontextum Dai15601

Ganoderma weberianum Dai 19689

Ganoderma cocoicola sp. nov. Cui 16792

Ganoderma yunlingense sp. nov. Cui 16288 (TYPE)

Ganoderma acaciicola sp. nov. Cui 16815 (TYPE)

Ganoderma sinense Cui 14526

Ganoderma casuarinicola Dai 19678

Ganoderma shanxiense Cui 14565

Ganoderma pfeifferi Dai 12153

Ganoderma cocoicola sp. nov. Cui 16791 (TYPE)

Ganoderma sublobatum sp. nov. Cui 16806

Ganoderma mirabile Cui 18237

Ganoderma gibbosum Cui 13940

Ganoderma tropicum Dai 20029

Ganoderma puerense sp. nov. Dai 20427 (TYPE)

Ganoderma gibbosum Cui 14338

Ganoderma ecuadorense Dai 17397

Ganoderma subangustisporum sp. nov. Cui 18597

Ganoderma tuberculosum JV 1607/62

Ganoderma guangxiense sp. nov. Cui 14454

Ganoderma orbiforme Cui 13891

Ganoderma orbiforme Cui 13880

Ganoderma hochiminhense Dai 18488

Ganoderma adspersum HSBU-200894

Ganoderma ecuadorense JV 1808/85

Ganoderma philippii Cui 14444

Ganoderma casuarinicola Dai 19470

Ganoderma casuarinicola Dai 16337

Ganoderma weberianum Dai 19682

Ganoderma angustisporum Cui 13817 (TYPE)

Ganoderma adspersum Dai 13191

Ganoderma castaneum sp. nov. Dai 13710

Ganoderma multipileum Dai 19690

Ganoderma mirabile Cui 16408

Ganoderma lucidum Cui 14404

Ganoderma williamsianum Dai 20553

Ganoderma multiplicatum Dai 17395

Ganoderma mirabile Cui 18283

Ganoderma acaciicola sp. nov. Cui 16813

Ganoderma eickeri Dai 12595

Ganoderma castaneum sp. nov. Cui 13893

Ganoderma mutabile Dai 20414

Ganoderma williamsianum Dai 17790

Ganoderma weberianum Cui 16359

Ganoderma chuxiongense sp. nov. Cui 17262 (TYPE)

Ganoderma lucidum Cui 14405

Ganoderma ellipsoideum Dai 19683

Ganoderma applanatum Cui 14062

Ganoderma orbiforme Cui 18326

Ganoderma mirabile Cui 18271

Ganoderma nasalanense CACP17060211 (TYPE)

Ganoderma philippii Cui 14443

Ganoderma tsugae Cui 14110

Ganoderma nasalanense CACP17060212

Ganoderma sinense Dai 20079

Ganoderma magniporum Dai 19966

100

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Ganodermataceae from China

Fig. 8. Scanning Electron Micrograph (SEM) of basidiospores of 10 genera in Ganodermataceae. A. Amauroderma schomburgkii (JV 1908/9). B.

Foraminispora rugosa (JV 1608/889-ND). C. Furtadoella brasiliensis (JV 1909/75). D. Ganoderma lucidum (Cui 14405). E. Haddowia macropora (JV

1908/46). F. Magoderna subresinosum (Cui 18280). G. Sanguinoderma rude (MEL 2150776). H. Sinoganoderma shandongense (Dai 20244). I. Tomophagus

cattienensis (Dai 18487). J. Trachydermella tsunodae (Dai 3221c). Scale bars = 2 μm.

322

Sun et al.

Table 2. The list of conrmed species in Ganodermataceae. Species in bold occur in China.

Genus Species Type locality Sequences References

Amauroderma (58) A. africanum Liberia – Ryvarden (2004b)

A. albocontextum Cameroon – Ryvarden (2020)

A. albostipitatum Brazil – Gomes-Silva et al. (2015)

A. andinum Venezuela – Ryvarden (2004b)

A. argenteofulvum Zimbabwe – Moncalvo & Ryvarden (1997)

A. aurantiacum Brazil √Gibertoni et al. (2008)

A. boleticeum Venezuela – Ryvarden (2004a)

A. buloloi Papua New Guinea – Moncalvo & Ryvarden (1997)

A. calcigenum Brazil √Ryvarden (2004a)

A. calcitum Brazil √TCosta-Rezende et al. (2016)

A. camerarium Brazil √Ryvarden (2004a)

A. coltricioides Guyana – Aime et al. (2003)

A. congregatum Malaysia – Corner (1983)

A. conicum Madagascar – Moncalvo & Ryvarden (1997)

A. conjunctum Africa – Moncalvo & Ryvarden (1997)

A. deviatum Ecuador – Ryvarden (2004a)

A. ealaense Zaire – Moncalvo & Ryvarden (1997)

A. elegantissimum Venezuela √Ryvarden (2004a)

A. exile Brazil √Ryvarden (2004a)

A. faculum Colombia – Henao-M (1997)

A. abellatum Guyana – Aime et al. (2007)

A. oriformum Brazil √TGomes-Silva et al. (2015)

A. fuscatum Uganda – Moncalvo & Ryvarden (1997)

A. fuscoporia Zimbabwe – Moncalvo & Ryvarden (1997)

A. grandisporum Burundi – Gulaid & Ryvarden (1998)

A. insulare Pacic: New Caledonia – Moncalvo & Ryvarden (1997)

A. intermedium Brazil √Ryvarden (2004a)

A. kwiluense Zaire – Ryvarden (1974)

A. laccatostipitatum Brazil √Gomes-Silva et al. (2015)

A. leptopus New Guinea – Furtado (1967b)

A. leucosporum Singapore – Corner (1983)

A. malesianum Malaysia – Corner (1983)

A. minuta Zimbabwe – Ryvarden (2018)

A. nigrum Cameroon – Moncalvo & Ryvarden (1997)

A. oblongisporum Angola – Campacci & Gugliotta (2009)

A. omphalodes Brazil √Ryvarden (2004a)

A. parasiticum Singapore – Corner (1983)

A. partitum Brazil √Gomes-Silva et al. (2010)

A. picipes Brazil – Gomes-Silva & Gibertoni (2012)

A. praetervisum Brazil √Ryvarden (2004a)

A. preussii Cameroon – Steyaert (1972)

A. pseudoboletus Paraguay √Ryvarden (2004a)

A. pudens India – Moncalvo & Ryvarden (1997)

A. renidens Brazil – Furtado (1967b), Ryvarden (2004a)

A. robledoi Brazil √Costa-Rezende et al. (2020a)

A. ryvardenii Zambia – Ryvarden (2020)

A. salisburiense Zimbabwe – Moncalvo & Ryvarden (1997)

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Ganodermataceae from China

Table 2. (Continued).

Genus Species Type locality Sequences References

A. schomburgkii Guyana √Ryvarden (2004a)

A. secedens Malaysia: Pahang – Corner (1983)

A. sericatum Nigeria – Moncalvo & Ryvarden (1997)

A. sessile Brazil – Gomes-Silva et al. (2015)

A. solomonense Solomon Islands – Corner (1983)

A. subrugosum Samoa Islands – Moncalvo & Ryvarden (1997)

A. subsessile Brazil √Gomes-Silva et al. (2015)

A. tapetellum Colombia – Henao-M (1997)

A. trichodermatum Brazil – Robledo et al. (2015)

A. unilaterum Brazil – Ryvarden (2004a)

A. velutina Cameroon – Ryvarden (2020)

Amaurodermellus (1) Amaurodermellus ovisporum Brazil √Gomes-Silva et al. (2015), Costa-

Rezende et al. (2020b)

Cristataspora (2) C. coffeata St. Vincent √Costa-Rezende et al. (2020b)

C. avipora Jamaica √Costa-Rezende et al. (2020b)

Foraminispora (5) Fo. austrosinensis China: Hainan √Zhao et al. (1984), Sun et al. (2020)

Fo. concentrica China: Sichuan √TSong et al. (2016), Sun et al. (2020)

Fo. rugosa Brazil √Costa-Rezende et al. (2017)

Fo. yinggelingensis China: Hainan √TSun et al. (2020)

Fo. yunnanensis China: Yunnan √Zhao & Zhang (1986b), Sun et al. (2020)

Furtadoella (3) Fu. biseptata comb. nov. Brazil √TCosta-Rezende et al. (2017)

Fu. brasiliensis comb. nov. Brazil √Costa-Rezende et al. (2017)

Fu. corneri comb. nov. Brazil – Costa-Rezende et al. (2017)

Ganoderma (181) G. acaciicola sp. nov. Australia √TThis study

G. acontextum sp. nov. Guatemala √TThis study

G. adspersum Slovenia: Vinkovce √Steyaert (1972), this study

G. aetii Indonesia: Kalimantan – Zmitrovich (2018)

G. ahmadii Pakistan: Sialkot – Steyaert (1972)

G. alluaudii Kenya: Nairobi – Ryvarden (1983)

G. alpinum sp. nov. China: Yunnan √TThis study

G. amazonense Brazil: Para State – Furtado (1967a)

G. angustisporum China: Fujian √TXing et al. (2018)

G. applanatum Europe √Patouillard (1887)

G. aridicola South Africa: Durban √TXing et al. (2016)

G. aureolum Angola: Tchivinguiro – Moncalvo & Ryvarden (1997)

G. australe Pacic island √ Ryvarden (2004a)

G. austroafricanum South Africa: Gauteng √Crous et al. (2014)

G. barretoi Brazil: Madeira – Moncalvo & Ryvarden (1997)

G. baudonii Central African Republic – Moncalvo & Ryvarden (1997)

G. bilobum – – –

G. boninense Japan: Bonin Islands √Ryvarden (1983)

G. brownii USA: California √Steyaert (1972)

G. bruggemanii Indonesia: Java – Steyaert (1972)

G. bubalinomarginatum sp.

nov.

China: Guangxi √TThis study

G. calidophilum China: Hainan √Cao (2013), this study

G. capense South Africa – Teng (1963)

G. carnosum France: Pyrenees √Moncalvo & Ryvarden (1997)

324

Sun et al.

Table 2. (Continued).

Genus Species Type locality Sequences References

G. carocalcareum Cameroon √TDouanla-Meli & Langer (2009)

G. castaneum sp. nov. China: Hainan √TThis study

G. casuarinicola China: Guangdong √TXing et al. (2018)

G. cervinum Papua New Guinea – Moncalvo & Ryvarden (1997)

G. chalceum Sierra Leone: Kenema √Steyaert (1967)

G. chocoense Ecuador: Esmeraldas √TCrous et al. (2018)

G. chonoides Zaire: Shaba – Moncalvo & Ryvarden (1997)

G. chuxiongense sp. nov. China: Yunnan √TThis study

G. cinnamomea Cameroon – Ryvarden (2020)

G. citriporum Venezuela: Yutaje – Ryvarden (2004a)

G. cocoicola sp. nov. Australia √TThis study

G. concinnum Colombia: Choco State √Ryvarden (2000)

G. corrugatum Zaire: Kasai – Steyaert (1961)

G. cupreum Guinea √Moncalvo & Ryvarden (1997), this study

G. curranii Philippines: Luzon – Murrill (1908a)

G. curtisii USA: South Carolina √Murrill (1908b)

G. dejongii Indonesia: Java – Steyaert (1972)

G. destructans South Africa: Gauteng √TCoetzee et al. (2015)

G. dianzhongense China: Yunnan √THe et al. (2021)

G. dimidiatum Japan – Papp (2016)

G. donkii Indonesia: West Java – Steyaert (1972)

G. dorsale Brazil – Moncalvo & Ryvarden (1997)

G. dubio-cochlear Madagascar – Moncalvo & Ryvarden (1997)

G. dunense South Africa: Western

Cape

√TTchotet Tchoumi et al. (2018)

G. dussii Guadeloupe – Moncalvo & Ryvarden (1997)

G. ecuadorense Ecuador: Orellana √TCrous et al. (2016)

G. eickeri South Africa √TTchotet Tchoumi et al. (2019)

G. elegantum Ecuador: Yasuni National

Park

– Ryvarden (2004a)

G. ellipsoideum China: Hainan √THapuarachchi et al. (2018b)

G. endochrum Uganda: Entebbe – Moncalvo & Ryvarden (1997)

G. enigmaticum South Africa: Gauteng √TCoetzee et al. (2015)

G. esculentum China: Yunnan √THe et al. (2021)

G. fallax sp. nov. USA √TThis study

G. fassii Congo: Ubangi – Steyaert (1961)

G. fassioides Congo: Yangambi – Steyaert (1961)

G. ci Tunisia: Gafsa – Moncalvo & Ryvarden (1997)

G. exipes Vietnam: Tonkin √Steyaert (1972)

G. fuscum Zaire: Shaba – Moncalvo & Ryvarden (1997)

G. gabonensis Gabon – Decock & Ryvarden (2020)

G. ghesquierei Zaire: Lukoleka – Moncalvo & Ryvarden (1997)

G. gibbosum Indonesia: Java √Moncalvo & Ryvarden (1997), this study

G. gilletii Zaire: Moanda – Moncalvo & Ryvarden (1997)

G. guangxiense sp. nov. China: Guangxi √TThis study

G. guianensis French Guiana – Ryvarden (2004a)

G. hildebrandii Comores Islands – Moncalvo & Ryvarden (1997)

G. hinnuleum Zaire: Yangambi – Moncalvo & Ryvarden (1997)

325www.studiesinmycology.org

Ganodermataceae from China

Table 2. (Continued).

Genus Species Type locality Sequences References

G. hochiminhense Vietnam √TLuangharn et al. (2021)

G. hoehnelianum Indonesia: Java √Luangharn et al. (2021)

G. hoploides Congo: Virunga National

Park

– Steyaert (1961)

G. impolitum Malaysia: Pahang – Moncalvo & Ryvarden (1997)

G. insulare Seychelles – Ryvarden (2020)

G. knysnamense South Africa √TTchotet Tchoumi et al. (2019)

G. kosteri The Netherlands: Gouda – Steyaert (1972)

G. lamaoense Philippines: Lamao – Steyaert (1972)

G. leucocontextum China: Tibet √TLi et al. (2015)

G. leucocreas Zaire: Loango – Moncalvo & Ryvarden (1997)

G. leytense Philippines: Leyte – Steyaert (1972)

G. lingua Indonesia: Java – Moncalvo & Ryvarden (1997)

G. lingzhi China: Hubei √TCao et al. (2012)

G. lobatoideum Guyana – Steyaert (1980)

G. lobatum USA: North Carolina √Steyaert (1980)

G. lobenense Cameroon – Kinge & Mih (2014)

G. longistipitatum Venezuela – Ryvarden (2000)

G. lucidum England: London √Steyaert (1972)

G. luteicinctum Singapore – Foroutan & Vaidya (2007)

G. magniporum China: Guangxi √This study

G. mangiferae Tahiti – Moncalvo & Ryvarden (1997)

G. manoutchehrii Iran: Ramsar – Steyaert (1972)

G. martinicense Martinique √TWelti & Courtecuisse (2010)

G. mbrekobenum Ghana √TCrous et al. (2016)

G. megalosporum Kenya: Nairobi – Moncalvo & Ryvarden (1997)

G. melanophaeum Zaire: Shaba – Moncalvo & Ryvarden (1997)

G. mexicanum Mexico √Torres-Torres & Guzmán-Dávalos (2012)

G. miniatocinctum Malaysia: Banting – Steyaert (1967)

G. mirabile Malaysia: Pahang √Steyaert (1972)

G. mizoramense India: Mizoram √TCrous et al. (2017a)

G. multicornum Venezuela – Ryvarden (2000)

G. multipileum China: Taiwan √Wang et al. (2009)

G. multiplicatum French Guiana √Steyaert (1980), Ryvarden (2000)

G. mutabile China: Yunnan √TCao & Yuan (2012)

G. myanmarense Myanmar √TLuangharn et al. (2021)

G. namutambalaense Uganda – Moncalvo & Ryvarden (1997)

G. nasalaense Laos √THapuarachchi et al. (2019b)

G. neogibbosum Martinica insula – Welti & Courtecuisse (2010)

G. neojaponicum Japan: Tokyo √TThis study

G. nitidum Honduras: Puerto Sierra √Moncalvo & Ryvarden (1997), this study

G. ochrolaccatum Philippines: Manila – Moncalvo & Ryvarden (1997)

G. oerstedii USA: Puerto Rico √Moncalvo & Ryvarden (1997)

G. orbiforme Guinea √Ryvarden (2000)

G. oregonense USA: Oregon √Murrill (1908b)

G. ostracodes Vietnam: Tonkin – Moncalvo & Ryvarden (1997)

G. parvigibbosum Martinique – Welti & Courtecuisse (2010)

326

Sun et al.

Table 2. (Continued).

Genus Species Type locality Sequences References

G. parvulum Nicaragua √Ryvarden (2004a), this study

G. petchii Sri Lanka: Hakgala – Steyaert (1972)

G. pfeifferi Germany √Foroutan & Vaidya (2007)

G. philippii Myanmar: Mergui √Steyaert (1972)

G. piceum Malaysia – Ryvarden (2015)

G. platense Argentina √Moncalvo & Ryvarden (1997), this study

G. podocarpense Ecuador √TCrous et al. (2017b)

G. polychromum USA: California √Moncalvo & Ryvarden (1997), this study

G. puerense sp. nov. China: Yunnan √TThis study

G. puglisii Italy: Potenza – Steyaert (1972)

G. pulchella – – Bresadola (1912)

G. pygmoideum Brazil – Moncalvo & Ryvarden (1997)

G. ramosissimum China: Yunnan √Zhao (1989a), this study

G. ravenelii USA: South Carolina √Steyaert (1980)

G. resinaceum France: Blois √Ryvarden (2000), Ryvarden (2004a)

G. reticulatosporum Zimbabwe: Harare – Moncalvo & Ryvarden (1997)

G. rhacodes – – Patouillard (1914)

G. rothwellii Zimbabwe – Steyaert (1980)

G. rufoalbum Venezuela – Moncalvo & Ryvarden (1997)

G. ryvardenii Cameroon √TKinge & Mih (2011)

G. sanduense China: Guizhou √THapuarachchi et al. (2019b)

G. sarasinii New Caledonia: Yate – Steyaert (1961)

G. sculpturatum Madagascar – Moncalvo & Ryvarden (1997)

G. septatum Zaire: Kivu – Moncalvo & Ryvarden (1997)

G. sessile USA: New York √Steyaert (1972), this study

G. sessiliforme Mexico √Torres-Torres & Guzmán-Dávalos (2012)

G. shanxiense China: Shanxi √TLiu et al. (2019)

G. sichuanense China: Sichuan √TZhao et al. (1983)

G. silveirae Brazil: Madeire – Moncalvo & Ryvarden (1997)

G. sinense China: Hainan √Zhao et al. (1979)

G. soyeri Zaire: Shaba – Steyaert (1961)

G. sp. Vietnam √Le et al. (2018)

G. steyaertianum Indonesia: Tirtaganga √Smith & Sivasithamparam (2003)

G. stipitatum Nicaragua √Murrill (1908b)

G. subangustisporum sp. nov. China: Yunnan √TThis study

G. subellipsoideum sp. nov. Malaysia √TThis study

G. subexipes sp. nov. China: Guangdong √TThis study

G. sublobatum sp. nov. Australia √TThis study

G. sublucidum Zaire: Eala – Moncalvo & Ryvarden (1997)

G. substipitata – – Bresadola (1915)

G. subumbraculum Japan – Moncalvo & Ryvarden (1997)

G. testaceum Brazil – Moncalvo & Ryvarden (1997)

G. thailandicum Thailand √TLuangharn et al. (2019)

G. tongshanense sp. nov. China: Hubei √TThis study

G. tornatum Mariana Island √Moncalvo & Ryvarden (1997), this study

G. torosum Thailand: Nakhawn

Strithamarat

– Moncalvo & Ryvarden (1997)

327www.studiesinmycology.org

Ganodermataceae from China

Table 2. (Continued).

Genus Species Type locality Sequences References

G. trengganuense Malaysia: Trengganu – Foroutan & Vaidya (2007)

G. tropicum Indonesia: Java √Steyaert (1972)

G. trulla Indonesia: Java – Moncalvo & Ryvarden (1997)

G. trulliforme Indonesia: Java – Moncalvo & Ryvarden (1997)

G. tsugae USA: New York √Murrill (1902)

G. tuberculosum Belize √Murrill (1908b)

G. turbinatum Uganda: Kabale – Ipulet & Ryvarden (2005)

G. umbrinum Indonesia: Java – Moncalvo & Ryvarden (1997)

G. valesiacum Switzerland: Valais – Moncalvo & Ryvarden (1997)

G. vanheurnii Indonesia: Java – Steyaert (1972)

G. vanmeelii Zaire: Shaba – Steyaert (1961)

G. vivianimercedianum Mexico – Torres-Torres (2008)

G. weberianum Samoa Islands √Steyaert (1972)

G. weixiense China: Yunnan √TYe et al. (2019)

G. wiiroense Ghana √TCrous et al. (2015)

G. williamsianum Philippines: Luzon √Murrill (1907)

G. xylonoides Zaire: Bongabo – Steyaert (1961)

G. yunlingense sp. nov. China: Yunnan √TThis study

G. zonatum USA: Florida √Murrill (1902)

Haddowia (2) Ha. longipes French Guyana √Steyaert (1972)

Ha. macropora sp. nov. French Guyana √TThis study

Humphreya (3) Hu. eminii Tanzania – Moncalvo & Ryvarden (1997)

Hu. endertii Indonesia – Steyaert (1972)

Hu. lloydii – – Steyaert (1972)

Magoderna (2) M. infundibuliforme Uganda – Steyaert (1972)

M. subresinosum Philippines: Luzon √Steyaert (1972)

Neoganoderma gen. nov.

(1)

N. neurosporum comb. nov. Panama √Furtado (1967a), Ryvarden (2004a)

Sanguinoderma (16) Sa. bataanense Philippines: Luzon √Murrill (1908a), Sun et al. (2020)

Sa. elmerianum Philippines: Luzon √Murrill (1907), Sun et al. (2020)

Sa. avovirens Zambia √TSun et al. (2020)

Sa. guangdongense sp. nov. China: Guangdong √TThis study

Sa. infundibulare sp. nov. China: Guangdong √TThis study

Sa. laceratum China: Yunnan √TSun et al. (2020)

Sa. longistipitum sp. nov. China: Yunnan √TThis study

Sa. melanocarpum sp. nov. Malaysia √TThis study

Sa. microporum China: Hainan √TSun et al. (2020)

Sa. microsporum sp. nov. Thailand √TThis study

Sa. perplexum Malaysia √Corner (1983), Sun et al. (2020)

Sa. reniforme Zambia √TSun et al. (2020)

Sa. rude Australia: Tasmania √Sun et al. (2020)

Sa. rugosum Indonesia: Java √Sun et al. (2020)

Sa. sinuosum Australia: Queensland √TSun et al. (2020)

Sa. tricolor sp. nov. Malaysia √TThis study

Sinoganoderma gen. nov.

(1)

Si. shandongense comb. nov. China: Shandong √This study

328

Sun et al.

Table 2. (Continued).

Genus Species Type locality Sequences References

Tomophagus (2) To. cattienensis Vietnam √TLe et al. (2012)

To. colossus Costa Rica √Le et al. (2012)

Trachydermella (1) Tr. tsunodae comb. nov. Japan √Imazeki (1952)

T Sequences from type specimens.

Furtadoella biseptata (Costa-Rezende et al.) B.K. Cui & Y.F. Sun,

comb. nov. MycoBank MB 843287.

Basionym: Furtadoa biseptata Costa-Rezende et al., Persoonia 39:

265. 2017.

Notes: Furtadoa biseptata was described as a new species by its

simple septate generative hyphae in the context. However, due to

the illegality of Furtadoa, this species was transferred to Furtadoella

as a new combination in this study. The description of Fu. biseptata

can be found in Costa-Rezende et al. (2017).

Furtadoella brasiliensis (Singer) B.K. Cui & Y.F. Sun, comb. nov.

MycoBank MB 843289.

Basionym: Scutiger brasiliensis Singer, Beih. Nova Hedwigia 77:

22. 1983.

Notes: Costa-Rezende et al. (2017) transferred Scutiger brasiliensis

to Furtadoa based on its similar morphological characters. But the

name Furatadoa is illegitimate, and therefore S. brasiliensis is

placed in Furtadoella. The description of Fu. brasiliensis can be

found in Coelho et al. (2007).

Furtadoella corneri (Gulaid & Ryvarden) B.K. Cui & Y.F. Sun,

comb. nov. MycoBank MB 843290.

Basionym: Amauroderma corneri Gulaid & Ryvarden, Mycol. Helv.

10: 28. 1998.

Notes: This species was rstly described from Brazil, and it was

recombined to Furtadoa by its monomitic hyphal system in context

(Costa-Rezende et al. 2017). In this study, Amauroderma corneri

was treated as a new combination in Furtadoella due to the illegality

of Furtadoa. The detailed description of Fu. corneri can be found in

Gulaid & Ryvarden (1998).

Ganoderma P. Karst., Revue Mycol., Toulouse 3: 17. 1881.

MycoBank MB 17639.

Type species: Ganoderma lucidum (Curtis) P. Karst.

Description: Basidiomata annual to perennial, sessile or subsessile

to stipitate. Pilei solitary or imbricate, of variable shape. Pileal

surface pale brown, reddish brown to almost black, dull to

laccate, with variable ornamentation. Context homogeneous or

heterogeneous. Tubes stratied or not. Hyphal system trimitic,

Fig. 9. Basidiomata of Ganoderma acaciicola.

329www.studiesinmycology.org

Ganodermataceae from China

Fig. 10. Microscopic structures of Ganoderma acaciicola (drawn from Cui 16815). A. Basidiospores. B. Apical cells from cuticle. C. Basidia and basidioles.

D. Hyphae from trama. E. Hyphae from context. Scale bars = 10 μm.

generative hyphae usually with clamp connections. Basidiospores

subglobose to ellipsoid or ovoid, truncated, double-walled with thick

walls, exospore wall semi-reticulate, endospore wall smooth or with

conspicuous spinules.

Notes: In this study, 95 species of Ganoderma were included to

construct the phylogenetic tree, and they formed an independent

clade with proper support (Fig. 1). Based on morphological

characters and phylogenetic evidence, 16 new species are described

and illustrated. In addition, 28 known species are also described, and

a key to conrmed species of Ganoderma in China is provided.

Ganoderma acaciicola B.K. Cui, J.H. Xing & Y.F. Sun, sp. nov.

MycoBank MB 839670. Figs 9, 10.

Diagnosis: Differs from other species in the genus by its sessile

and concrescent basidiomata with reddish brown and laccate pileal

surface, homogeneous context, non-stratied tubes, cream to buff

pore surface unchanging when bruised, broadly ellipsoid to ovoid

basidiospores with truncated apex.

Etymology: acaciicola (Lat.), refers to this species growing on

Acacia.

330

Sun et al.

Typus: Australia, Queensland, Cairns, on stump of Acacia, 18 May 2018,

Cui 16815 (holotype BJFC030114).

Additional materials examined: Australia, Queensland, Cairns, on stump of

Acacia, 18 May 2018, Cui 16813 (BJFC030112), Cui 16814 (BJFC030113);

on root of living Acacia, 18 May 2018, Cui 16817 (BJFC030116).

Description: Basidiomata annual, sessile or subsessile and broadly

attached, usually concrescent, hard corky to woody hard. Pilei sub-

circular to abelliform, up to 16.5 cm diam and 3 cm thick. Pileal

surface rusty orange brown to reddish brown, laccate, glabrous, pileal

margin distinct, cream buff; margin obtuse, entire, irregularly wavy.

Pore surface cream to buff when fresh, unchanging when bruised,

pale straw yellow when dry; pores circular to angular, 4–6 per mm;

dissepiments moderately thick, entire. Context cinnamon brown to

dark brown, homogeneous, with black melanoid lines, hard corky,

up to 2 cm thick. Tubes yellowish brown to greyish brown, non-

stratied, up to 1 cm long. Hyphal system trimitic; generative hyphae

with clamp connections; all hyphae IKI –, CB +; tissues darkening in

KOH. Generative hyphae in context colourless, thin-walled, 2–4 μm

diam; skeletal hyphae in context dark brown, thick-walled with a wide

to narrow lumen or sub-solid, arboriform and exuous, 2–6 μm diam;

binding hyphae in context colourless, thick-walled, branched and

exuous, up to 1 µm diam. Generative hyphae in tubes colourless, thin-

walled, 2–3 μm diam; skeletal hyphae in tubes pale yellowish brown,

thick-walled with a wide to narrow lumen or sub-solid, arboriform and

exuous, 2–4 μm diam; binding hyphae in tubes colourless, thick-

walled, branched and exuous, up to 1 μm diam. Pileipellis composed

of clamped generative hyphae, thick-walled to sub-solid, apical cells

clavate, inated and exuous, pale yellow to golden yellow, about

25–38 × 6–10 μm, forming a regular palisade. Cystidia and cystidioles

absent. Basidia barrel-shaped, colourless, thin-walled, 15–20 × 9–12

µm; basidioles clavate, colourless, thin-walled, 15–20 × 6–10 μm.

Basidiospores broadly ellipsoid to ovoid, truncated, yellowish brown,

IKI –, CB +, double-walled with distinctly thick walls, exospore wall

smooth, endospore wall with dense spinules, (9–)9.2–11.3(–11.6) ×

(5–)5.6–7 μm, L = 10.12 μm, W = 6.29 μm, Q = 1.61 (n = 60/2, with

the turgid vesicular appendix excluded); (10.5–)10.7–11.8(–12.1) ×

(5.5–)5.8–7 μm, L = 11.12 μm, W = 6.31 μm, Q = 1.72–1.81 (n = 60/2,

with the turgid vesicular appendix included).

Notes: Ganoderma acaciicola was collected from Australia on

Acacia. It can be characterised by concrescent basidiomata without

a stipe, rusty orange brown to reddish brown and laccate pileal

surface, pore surface unchanging when bruised. In the phylogenetic

analyses, G. acaciicola is closely related to G. mizoramense

which was described from Mizoram, India; however, its stipitate

basidiomata with irregular pileal surface, ellipsoid basidiospores in

larger size (10–12.5 × 6–9 μm, Crous et al. 2017a) differentiate it

from G. acaciicola.

Ganoderma acontextum B.K. Cui, J.H. Xing & Vlasák, sp. nov.

MycoBank MB 805754. Figs 11, 12.

Diagnosis: Differs from other species in the genus by its ungulate

pilei with non-laccate pileal surface, heterogeneous and thin

context, non-stratied tubes, almond-shaped basidiospores without

spinules on the endospore wall.

Etymology: acontextum (Lat.), refers to the basidiomata having

extremely thin context.

Typus: Guatemala, San Mateo, on angiosperm tree, 22 Nov. 2006

(holotype JV 0611/21G).

Fig. 11. Basidiomata of Ganoderma acontextum.

331www.studiesinmycology.org

Ganodermataceae from China

Additional materials examined: USA, Virginia, Woodbridge, Mason Neck

State Park, on Quercus, 11 Aug. 2012, JV 1208/11J, JV 1407/64 (JV).

Diagnosis: Basidiomata perennial, sessile, woody hard. Pilei solitary,

ungulate to columnar at maturity, up to 6 cm diam and 4 cm thick.

Pileal surface reddish brown to dark brown, dull, glabrous, with dense

concentric furrows; margin obtuse, entire, wavy. Pore surface white

when fresh, turning darker when bruised, clay-buff to dark brown

when dry; pores circular, 4–6 per mm; dissepiments thick, entire.

Context dark brown, heterogeneous, composed of a strikingly light

brown to ochre zone under pileal crust, followed by black melanoid

lines, and dark brown context above the tubes, corky, thin, up to

3 mm thick altogether. Tubes dark brown, non-stratied, up to 4

cm long. Hyphal system trimitic; generative hyphae with clamp

connections; skeletal hyphae occasionally with simple septa; all

hyphae IKI –, CB +; tissues darkening in KOH. Generative hyphae

in context colourless, thin-walled, 2–3 µm diam; skeletal hyphae in

context yellow to brown, thick-walled with a wide to narrow lumen or

sub-solid, arboriform and exuous, 2–5 µm diam; binding hyphae in

context colourless, thick-walled, branched and exuous, 1.2–2.5 μm

diam. Generative hyphae in tubes colourless, thin-walled, 2.2–2.8

μm diam; skeletal hyphae in tubes pale brown to brown, thick-walled

with a wide to narrow lumen or sub-solid, arboriform and exuous,

2.3–4 μm diam; binding hyphae in tubes colourless, thick-walled,

branched and exuous, 1–2.5 μm diam. Cystidia and cystidioles

absent. Basidia broadly clavate, colourless, thin-walled, 20–30 ×

7–11 μm; basidioles in shape like the basidia, colourless, thin-walled,

14–25 × 6–10 μm. Basidiospores almond-shaped, not obviously

Fig. 12. Microscopic structures of Ganoderma acontextum (drawn from JV 0611/21G). A. Basidiospores. B. Basidia and basidioles. C. Hyphae from trama.

D. Hyphae from context. Scale bars = 10 μm.

332

Sun et al.

truncated, golden-brown, IKI –, CB +, double-walled with slightly

thick walls, exospore wall smooth, endospore wall without spinules,

(9–)9.8–10.2(–11) × (4–)5.5–6.3(–6.5) μm, L = 9.89 μm, W = 5.73

μm, Q = 1.73 (n = 30/1, with the turgid vesicular appendix excluded).

Notes: Ganoderma acontextum was collected from Central America

and the USA belonging to the non-laccate group. It has distinctive

features such as ungulate pilei with densely concentrically furrowed

pileal surface, heterogeneous and thin context, non-stratied tubes,

almond-shaped basidiospores without an obviously truncated apex,

and no spinules on the endospore wall.

Ganoderma alpinum B.K. Cui, J.H. Xing & Y.F. Sun, sp. nov.

MycoBank MB 839671. Figs 13, 14.

Diagnosis: Differs from other species in the genus by its perennial

and sessile basidiomata, pale brown to greyish brown pileal surface

with concentric furrows and radial wrinkles, cracked margin,

homogeneous context and non-stratied tubes.

Etymology: alpinum (Lat.), refers to this species being collected

from an alpine area.

Typus: China, Yunnan, Shangri-La, Daxueshan, on stump of Populus, 12

Aug. 2019, Cui 17467 (holotype BJFC034326).

Additional materials examined: China, Sichuan, Yajiang County, on stump

of Pinus, 8 Aug. 2019, Cui 17325 (BJFC034183); Xizang, Chayu County,

on stump of Cupressus, 10 Sep. 2020, Cui 18402 (BJFC035263).

Description: Basidiomata perennial, sessile, woody hard. Pilei

abelliform to shell-shaped, applanate, up to 15 cm diam and 4 cm

thick. Pileal surface pale brown to greyish brown, dull, glabrous, with

concentric furrows and radial wrinkles; margin subacute to obtuse,

entire, slightly wavy, cracked when dry. Pore surface white to cream

when fresh, turning darker when bruised, clay buff to dark brown

when dry; pores circular, 5–7 per mm; dissepiments slightly thick,

entire. Context cinnamon brown to dark brown, homogeneous,

with black melanoid lines, hard corky and brous, up to 2 cm thick.

Tubes yellowish brown to dark brown, non-stratied, up to 2 cm long.

Hyphal system trimitic; generative hyphae with clamp connections;

all hyphae IKI –, CB +; tissues darkening in KOH. Generative hyphae

in context colourless, thin-walled, 3–4 µm diam; skeletal hyphae

in context pale yellowish brown, thick-walled with a wide to narrow

lumen or sub-solid, arboriform and exuous, 3–6 µm diam; binding

hyphae in context colourless, thick-walled, branched and exuous,

up to 2 µm diam. Generative hyphae in tubes colourless, thin-walled,

2–4 μm diam; skeletal hyphae in tubes pale brown, thick-walled with

a wide to narrow lumen or sub-solid, arboriform and exuous, 3–5

μm diam; binding hyphae in tubes colourless, thick-walled, branched

and exuous, up to 2 μm diam. Pileipellis composed of clamped

generative hyphae, thick-walled, apical cells clavate, slightly inated

and exuous, yellowish brown, about 20–30 × 5–8 μm, forming a

regular palisade. Cystidia and cystidioles absent. Basidia barrel-

shaped, colourless, thin-walled, 20–30 × 11–16 µm; basidioles

clavate, colourless, thin-walled, 12–18 × 6–10 μm. Basidiospores

broadly ellipsoid to ovoid, truncated, yellowish brown, IKI –, CB +,

double-walled with distinctly thick walls, exospore wall smooth,

endospore wall with dense spinules, (6.2–)6.3–7.6(–7.8) × (4–)4.1–

5.4(–5.5) μm, L = 6.97 μm, W = 4.83 μm, Q = 1.43–1.45 (n = 60/2,

Fig. 13. Basidiomata of Ganoderma alpinum.

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Ganodermataceae from China

with the turgid vesicular appendix excluded); (7.7–)7.8–9.2(–9.4) ×

(4–)4.1–5.4(–5.8) μm, L = 8.37 μm, W = 4.85 μm, Q = 1.71–1.74 (n =

60/2, with the turgid vesicular appendix included).

Notes: Ganoderma alpinum was collected from high altitude areas

of southwestern China. It is hard to distinguish G. alpinum from G.

applanatum on morphology, however, G. alpinum can be separated

from G. applanatum by phylogenetic analyses and ecological

distribution.

Ganoderma bubalinomarginatum B.K. Cui, J.H. Xing & Y.F. Sun,

sp. nov. MycoBank MB 839672. Figs 15, 16.

Fig. 14. Microscopic structures of Ganoderma alpinum (drawn from Cui 17467). A. Basidiospores. B. Apical cells from cuticle. C. Basidia and basidioles. D.

Hyphae from trama. E. Hyphae from context. Scale bars = 10 μm.

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Sun et al.

Diagnosis: Differs from other species in the genus by its pale-

coloured basidiomata without stipe, laccate pileal surface with buff

margin, homogeneous context and non-stratied tubes.

Etymology: bubalinomarginatum (Lat.), refers to the pilei with buff

margin.

Typus: China, Guangxi, Nanning, Guangxi Academy of Forestry, on stump

of Castanopsis, 4 Jul. 2019, Dai 20075 (holotype BJFC031749).

Additional material examined: China, Guangxi, Nanning, Guangxi

Academy of Forestry, on living tree of Phoebe, 4 Jul. 2019, Dai 20074

(BJFC031748).

Description: Basidiomata annual, sessile and broadly attached,

usually concrescent, hard corky. Pilei solitary, abelliform to

shell-shaped, up to 7.5 cm diam and 6 mm thick. Pileal surface

reddish brown at the base, yellowish brown at the centre, buff at

the margin, laccate, glabrous, with wide concentric furrows and

slightly radial wrinkles; margin obtuse, entire, wavy when dry.

Pore surface white to greyish white when fresh, turning darker

when bruised, pale wood brown to greyish brown when dry;

pores circular to angular, 5–6 per mm; dissepiments moderately

thick, entire. Context straw yellow, homogeneous, without black

melanoid lines, hard corky, up to 4 mm thick. Tubes pale brown,

non-stratied, up to 3 mm long. Hyphal system trimitic; generative

hyphae with clamp connections; all hyphae IKI –, CB +; tissues

darkening in KOH. Generative hyphae in context colourless, thin-

walled, 2–4 μm diam; skeletal hyphae in context pale yellow,

thick-walled with a wide to narrow lumen or sub-solid, arboriform

and exuous, 2–5 μm diam; binding hyphae in context colourless,

thick-walled, rarely branched and exuous, up to 2 µm diam.

Generative hyphae in tubes colourless, thin-walled, 2–3 μm

diam; skeletal hyphae in tubes pale yellow, thick-walled with a

wide to narrow lumen or sub-solid, arboriform and exuous, 2–4

μm diam; binding hyphae in tubes colourless, thick-walled, rarely

branched and exuous, up to 1.5 μm diam. Pileipellis composed

of clamped generative hyphae, thick-walled to sub-solid, apical

cells clavate, inated and exuous, pale golden yellow, about 28–

42 × 9–11 μm, forming a regular palisade. Cystidia and cystidioles

absent. Basidia broadly clavate, colourless, thin-walled, 15–22 ×

7–11 μm; basidioles in shape like the basidia, colourless, thin-

walled, 12–20 × 5–9 μm. Basidiospores broadly ellipsoid to ovoid,

truncated, pale yellowish brown, IKI –, CB +, with oily drop,

double-walled with slightly thick walls, exospore wall smooth,

endospore wall with dense spinules, (6.2–)6.5–7.4(–7.6) × (4.2–)

4.5–5.3(–5.8) μm, L = 6.91 μm, W = 4.87 μm, Q = 1.41–1.43 (n =

60/2, with the turgid vesicular appendix excluded); (7–)7.2–8.3(–

8.8) × (4.3–)4.5–5.6(–5.8) μm, L = 7.82 μm, W = 4.96 μm, Q =

1.57–1.59 (n = 60/2, with the turgid vesicular appendix included).

Notes: Ganoderma bubalinomarginatum has pale-coloured

basidiomata without a stipe which differentiates it from the

species described from Guangxi Autonomous Region: G.

daiqingshanense, G. guinanense and G. magniporum. Ganoderma

Fig. 15. Basidiomata of Ganoderma bubalinomarginatum.

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Ganodermataceae from China

bubalinomarginatum is like G. sessile, described from America: it

shares connate and sessile basidiomata, reddish brown to yellowish

brown pileal surface, absence of black melanoid lines, white pore

surface when fresh, homogeneous context and non-stratied

tubes. However, G. sessile differs by the white, thin and acute pileal

margin, larger pores (3–5 per mm), and larger basidiospores (9–11

× 6–8 μm, Murrill 1902).

Ganoderma castaneum B.K. Cui, J.H. Xing & Y.F. Sun, sp. nov.

MycoBank MB 839673. Figs 17, 18.

Diagnosis: Differs from other species in the genus by its chestnut

brown and laccate pileal surface with wide concentric ridges,

heterogeneous context, and broadly ellipsoid basidiospores with

smooth endospore wall.

Fig. 16. Microscopic structures of Ganoderma bubalinomarginatum (drawn from Dai 20075). A. Basidiospores. B. Apical cells from cuticle. C. Basidia and

basidioles. D. Hyphae from trama. E. Hyphae from context. Scale bars = 10 μm.

336

Sun et al.

Etymology: castaneum (Lat.), refers to the reddish brown pileal

surface like a chestnut.

Typus: China, Hainan, Ledong County, Jianfengling Nature Reserve, on

stump of angiosperm tree, 3 Jul. 2019, Cui 17283 (holotype BJFC034139).

Additional materials examined: China, Hainan, Ledong County,

Jianfengling Nature Reserve, on stump of angiosperm tree, 17 Jun. 2014,

Dai 13710 (BJFC017447); on fallen branch of angiosperm tree, 19 Jun.

2016, Cui 13893 (BJFC028759); Wuzhishan, Wuzhishan Forest Park,

on stump of Acacia, 11 Jun. 2016, Dai 16500 (BJFC022616), Dai 16501

(BJFC022617).

Description: Basidiomata annual, sessile, broadly attached, hard

corky to woody hard. Pilei solitary, abelliform, applanate, up to 8.5

cm diam and 2 cm thick. Pileal surface reddish brown like chestnut,

laccate, glabrous, with wide concentric ridges and slightly radial

wrinkles; margin obtuse, entire, wavy. Pore surface white to cream

when fresh, turning darker when bruised, buff to pale straw yellow

when dry; pores circular, 4–6 per mm; dissepiments moderately

thick, entire. Context heterogeneous, the upper layer pale straw

yellow, the lower layer cinnamon brown to dark brown, with black

melanoid lines, hard corky, up to 1.6 cm thick. Tubes pale greyish

brown, non-stratied, up to 5 mm long. Hyphal system trimitic;

generative hyphae with clamp connections; all hyphae IKI –, CB +;

tissues darkening in KOH. Generative hyphae in context colourless,

thin-walled, 2–4 μm diam; skeletal hyphae in context pale yellowish

brown, thick-walled with a wide to narrow lumen or sub-solid,

arboriform and exuous, 3–6 µm diam; binding hyphae in context

colourless, thick-walled, branched and exuous, up to 2 μm diam.

Generative hyphae in tubes colourless, thin-walled, 2–4 μm diam;

skeletal hyphae in tubes pale yellowish brown, thick-walled with a

wide to narrow lumen or sub-solid, arboriform and exuous, 2–6 μm

diam; binding hyphae in tubes colourless, thick-walled, branched

and exuous, up to 1 μm diam. Pileipellis composed of clamped

generative hyphae, slightly thick-walled, apical cells clavate,

exuous, pale yellow, about 25–40 × 3–5 μm, forming a regular

palisade. Cystidia and cystidioles absent. Basidia barrel-shaped,

colourless, thin-walled, 14–21 × 7–12 μm; basidioles broadly

clavate, colourless, thin-walled, 9–12 × 5–10 μm. Basidiospores

broadly ellipsoid, not obviously truncated, golden yellow, IKI –,

slightly CB +, double-walled with distinctly thick walls, exospore

and endospore walls smooth, (6.2–)6.7–8.3(–8.5) × (4.2–)4.8–6(–

6.3) μm, L = 7.42 µm, W = 5.43 μm, Q = 1.37 (n = 60/1, with the

turgid vesicular appendix excluded).

Notes: Ganoderma castaneum was collected from a tropical

rainforest of Hainan Province. When compared with the species

in the checklist of Ganoderma reported from Hainan Island by

Hapuarachchi et al. (2018b), G. castaneum has distinguished

features such as chestnut-coloured and laccate pileal surface with

wide concentric ridges, heterogeneous context, broadly ellipsoid

and basidiospores not obviously truncated with smooth endospore

walls.

Ganoderma chuxiongense B.K. Cui, J.H. Xing & Y.F. Sun, sp.

nov. MycoBank MB 840397. Figs 19, 20.

Diagnosis: Differs from other species in the genus by its thin

basidiomata, dimidiate and lobate pileus with reddish brown and

laccate pileal surface, pale light-yellow pore surface.

Fig. 17. Basidiomata of Ganoderma castaneum.

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Ganodermataceae from China

Etymology: chuxiongense (Lat.), refers to the holotype of this

species being found in Chuxiong City of Yunnan Province.

Typus: China, Yunnan, Chuxiong, Xishan Park, on stump of angiosperm

tree, 25 Aug. 2018, Cui 17262 (holotype BJFC034120).

Description: Basidiomata annual, laterally stipitate, corky. Pilei

solitary, abelliform, dimidiate and lobate, up to 6 cm diam and

4 mm thick. Pileal surface reddish brown when fresh becoming

dark-red when dry, laccate, glabrous, with concentric bands and

slightly radial rugose; margin pale yellow, acute to obtuse, entire.

Pore surface pale light-yellow when fresh, turning darker when

bruised, buff to pale straw yellow when dry; pores circular, 4–6

per mm; dissepiments slightly thick, entire. Context dark wood

brown, not obviously stratied, without black melanoid lines, soft

corky, up to 2 mm thick. Tubes pale to dark wood brown, non-

Fig. 18. Microscopic structures of Ganoderma castaneum (drawn from Cui 17283). A. Basidiospores. B. Apical cells from cuticle. C. Basidia and basidioles.

D. Hyphae from trama. E. Hyphae from context. Scale bars = 10 μm.

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Sun et al.

stratied, up to 2 mm long. Stipe concolorous with pileal surface,

cylindrical and solid, up to 7 cm long and 6 mm diam. Hyphal

system trimitic; generative hyphae with clamp connections; all

hyphae IKI –, CB +; tissues darkening in KOH. Generative hyphae

in context colourless, thin-walled, 2–3 µm diam; skeletal hyphae

in context pale yellow, thick-walled with a wide to narrow lumen or

sub-solid, arboriform and exuous, 2–4 µm diam; binding hyphae

in context colourless, thick-walled, branched and exuous, up to

2 μm diam. Generative hyphae in tubes colourless, thin-walled,

2–3 μm diam; skeletal hyphae in tubes pale yellow, thick-walled

with a wide to narrow lumen or sub-solid, arboriform and exuous,

2–4 μm diam; binding hyphae in tubes colourless, thick-walled,

branched and exuous, up to 1.5 μm diam. Pileipellis composed

of clamped generative hyphae, slightly thick-walled, apical cells

clavate, slightly inated and exuous, golden yellow, about 23–30

× 6–11 μm, forming a regular palisade. Cystidia and cystidioles

absent. Basidia barrel-shaped, colourless, thin-walled, 15–20 ×

10–12 μm; basidioles broadly clavate, colourless, thin-walled,

13–18 × 7–11 μm. Basidiospores broadly ellipsoid to ovoid,

truncated, pale yellowish brown, IKI –, CB +, double-walled with

distinctly thick walls, exospore wall smooth, endospore wall with

dense spinules, (7.9–)8–9(–9.2) × (6–)6.2–7(–7.1) μm, L = 8.45

μm, W = 6.58 μm, Q = 1.28 (n = 60/1, with the turgid vesicular

appendix excluded); (9.5–)10–11.3(–11.5) × (6.2–)6.5–7.3 μm, L

= 10.35 μm, W = 6.88 μm, Q = 1.5 (n = 60/1, with the turgid

vesicular appendix included).

Notes: Ganoderma chuxiongense is characterised by its stipitate

basidiomata with lobate pilei, reddish brown pileal surface and

pale-yellow pore surface when fresh. Ganoderma kunmingense

described from Yunnan Province is like G. chuxiongense in the

thin basidiomata with laccate and reddish brown pileal surface,

margin entire or incised, but it has no concentric bands on the pileal

surface, a cream pore surface, and broadly ellipsoid to subglobose

basidiospores without obvious spinules on the endospore wall

(Zhao 1989a).

Ganoderma cocoicola B.K. Cui, J.H. Xing & Y.F. Sun, sp. nov.

MycoBank MB 839674. Figs 21, 22.

Diagnosis: Differs from other species in the genus by its small and

hard basidiomata without stipe, dark and laccate pileal surface,

homogeneous context, non-stratied tubes, oblong-ellipsoid and

truncated basidiospores.

Etymology: cocoicola (Lat.), refers to this species growing on

Cocos.

Typus: Australia, Queensland, Cairns, Cairns Botanical Garden, on stump

of Cocos, 17 May 2018, Cui 16791 (holotype BJFC030090).

Additional material examined: Australia, Queensland, Cairns, Cairns

Botanical Garden, on stump of Cocos, 17 May 2018, Cui 16792

(BJFC030091).

Fig. 19. Basidiomata of Ganoderma chuxiongense.

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Ganodermataceae from China

Description: Basidiomata annual to perennial, sessile or sometimes

sub-stipitate, hard corky to woody hard. Pilei solitary, abelliform to

ungulate, up to 3.5 cm diam and 1.5 cm thick. Pileal surface dark-

red to near black, laccate, glabrous, with concentric furrows and

slightly radial wrinkles; margin obtuse, entire. Pore surface white

when fresh, turning darker when bruised, straw yellow to pale brown

when dry; pores circular to angular, 4–6 per mm; dissepiments

distinctly thick, entire. Context dark brown to cinnamon brown,

homogeneous, without black melanoid lines, hard corky, up to 3

mm thick. Tubes dark grey to greyish brown, non-stratied, up to

1.2 cm long. Hyphal system trimitic; generative hyphae with clamp

connections; all hyphae IKI –, CB +; tissues darkening in KOH.

Generative hyphae in context colourless, thin-walled, 2–3 μm

diam; skeletal hyphae in context pale yellowish brown, thick-walled

Fig. 20. Microscopic structures of Ganoderma chuxiongense (drawn from Cui 17262). A. Basidiospores. B. Apical cells from cuticle. C. Basidia and basidioles.

D. Hyphae from trama. E. Hyphae from context. Scale bars = 10 μm.

340

Sun et al.

with a wide to narrow lumen or sub-solid, frequently arboriform and

exuous, 2.5–5 μm diam; binding hyphae in context colourless,

thick-walled, branched and exuous, up to 2 μm diam. Generative

hyphae in tubes colourless, thin-walled, 2–3 μm diam; skeletal

hyphae in tubes pale yellowish brown, thick-walled with a wide

to narrow lumen or sub-solid, frequently arboriform and exuous,

3–5 μm diam; binding hyphae in tubes colourless, thick-walled,

branched and exuous, up to 1.5 μm diam. Pileipellis composed of

clamped generative hyphae, thick-walled to sub-solid, apical cells

clavate, exuous, yellowish brown, about 27–35 × 5–7 μm, forming

a regular palisade. Cystidia and cystidioles absent. Basidia broadly

clavate, colourless, thin-walled, 16–22 × 7–10 µm; basidioles in

shape like the basidia, colourless, thin-walled, 13–20 × 7–10 μm.

Basidiospores oblong-ellipsoid, truncated, pale yellowish brown,

IKI –, CB +, double-walled with moderately thick walls, exospore

wall smooth, endospore wall with dense spinules, 9–9.8(–10) ×

4.2–5.4(–5.6) μm, L = 9.42 µm, W = 4.77 μm, Q = 1.91–2.04 (n =

60/2, with the turgid vesicular appendix excluded); (9.6–)9.7–10.7(–

10.8) × (4.2–)4.4–5.4(–5.5) μm, L = 10.15 µm, W = 4.83 µm, Q =

2.08–2.12 (n = 60/2, with the turgid vesicular appendix included).

Notes: Ganoderma cocoicola was collected from Australia on

Cocos. It is characterised by small and hard basidiomata without

a stipe, blackish brown and laccate pileal surface, homogeneous

context, non-stratied tubes, oblong-ellipsoid and truncated

basidiospores. In the phylogenetic analyses, G. cocoicola

clustered with G. zonatum, G. ryvardenii, G. boninense, and G.

hochiminhense with high support (Fig. 1); all these species grow

on palm trees (Patouillard 1889, Murrill 1902, Kinge & Mih 2011,

Zhou et al. 2015), except G. hochiminhense which grows on Areca

(Luangharn et al. 2021).

Ganoderma fallax B.K. Cui, J.H. Xing & Vlasák, sp. nov. MycoBank

MB 839677. Figs 23, 24.

Diagnosis: Differs from other species in the genus by its non-laccate

pileal surface with faintly concentric furrows, stratied tubes, and

basidiospores not obviously truncated with dense spinules on the

endospore wall.

Etymology: fallax (Lat.), refers to this species being easily confused

morphologically with other non-laccate species.

Typus: USA, Pennsylvania (holotype JV 1009/27).

Materials examined: USA, Arizona, JV 1209/60J (JV); New Jersey, JV

0109/B1-J (JV); Pennsylvania, JV 0709/39 (JV); Tennessee, JV 0509/93K,

JV 1410/14J (JV).

Description: Basidiomata perennial, sessile, hard corky to woody

hard. Pilei solitary, abelliform to ungulate or shell-shaped, up to

11 cm diam and 3 cm thick. Pileal surface yellowish brown to dark

brown, dull, glabrous, with faint concentric furrows; margin acute,

entire, slightly wavy. Pore surface white to pale brown when fresh,

Fig. 21. Basidiomata of Ganoderma cocoicola.

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Ganodermataceae from China

turning darker when bruised, straw yellow when dry; pores circular,

4–6 per mm; dissepiments slightly thick, entire. Context yellowish

brown to dark brown, homogeneous, with black melanoid lines,

corky, up to 7 mm thick. Tubes concolorous with context, stratied,

up to 1.2 cm long. Hyphal system trimitic; generative hyphae with

clamp connections; all hyphae IKI –, CB +; tissues darkening in

KOH. Generative hyphae in context colourless, thin-walled, 2.5–5.5

µm diam; skeletal hyphae in context pale yellowish brown, thick-

walled with narrow lumen or sub-solid, arboriform and exuous,

3–6 μm diam; binding hyphae in context colourless, thick-walled,

branched and exuous, 2–3 μm diam. Generative hyphae in tubes

colourless, thin-walled, 2–3.5 μm diam; skeletal hyphae in tubes

Fig. 22. Microscopic structures of Ganoderma cocoicola (drawn from Cui 16791). A. Basidiospores. B. Apical cells from cuticle. C. Basidia and basidioles.

D. Hyphae from trama. E. Hyphae from context. Scale bars = 10 μm.

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Sun et al.

pale brown, thick-walled with narrow lumen or sub-solid, arboriform

and exuous, 2–5 μm diam; binding hyphae in tubes colourless,

thick-walled, branched and exuous, 2–2.5 μm diam. Cystidia and

cystidioles absent. Basidia barrel-shaped to clavate, colourless,

thin-walled, 25–30 × 14–21 μm; basidioles in shape like the basidia,

colourless, thin-walled, 16–17 × 9–18 μm. Basidiospores ovoid, not

obviously truncated, pale yellowish brown, IKI –, CB +, double-

walled with distinctly thick walls, exospore wall smooth, endospore

wall with dense spinules, (8.5–)9–10(–10.5) × 6–7.5 µm, L = 9.25

μm, W = 6.56 μm, Q = 1.37 (n = 30/1, with the turgid vesicular

appendix included).

Notes: Ganoderma fallax may be confused with the non-laccate

species G. adspersum, which has large basidiomata with a

yellowish brown pileal surface, homogeneous context and

stratied tubes, but G. adspersum has larger and more obviously

truncated basidiospores, and lacks concentric ornamentation on

the pileal surface. In the phylogenetic analyses, G. fallax formed

an independent lineage then clustered with G. adspersum (Fig. 1).

Ganoderma guangxiense B.K. Cui, J.H. Xing & Y.F. Sun, sp. nov.

MycoBank MB 839678. Figs 25, 26.

Diagnosis: Differs from other species in the genus by its sessile

basidiomata, greyish brown to near black pileal surface with obvious

concentric furrows, cracked when dry, homogeneous context, non-

stratied tubes, ellipsoid to ovoid and truncated basidiospores.

Etymology: guangxiense (Lat.), refers to the holotype of this

species located at Guangxi.

Typus: China, Guangxi, Tianlin County, Cenwanglaoshan Nature

Reserve, on stump of angiosperm tree, 8 Jul. 2017, Cui 14453 (holotype

BJFC029321).

Additional materials examined: China, Guangxi, Tianlin County,

Cenwanglaoshan Nature Reserve, on stump of angiosperm tree, 8 Jul.

2017, Cui 14454 (BJFC029322), Cui 14455 (BJFC029323); Jinxiu County,

Dayaoshan Nature Reserve, on fallen trunk of angiosperm tree, 15 Jul.

2017, Cui 14500 (BJFC029369), Cui 14508 (BJFC029377).

Description: Basidiomata annual, sessile and broadly attached, hard

corky to woody hard. Pilei solitary, abelliform to shell-shaped, up to

11 cm diam and 4.3 cm thick. Pileal surface greyish brown to near

black when fresh, dull, glabrous, with obvious concentric furrows,

cracked when dry; margin obtuse, entire. Pore surface cream when

Fig. 23. Basidiomata of Ganoderma fallax.

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Ganodermataceae from China

Fig. 24. Microscopic structures of Ganoderma fallax (drawn from JV 1009/27). A. Basidiospores. B. Basidia and basidioles. C. Hyphae from trama. D.

Hyphae from context. Scale bars = 10 μm.

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fresh, turning darker when bruised, dark straw yellow to pale brown

when dry; pores circular, 5–7 per mm; dissepiments moderately thick,

entire. Context dark brown to cinnamon brown, homogeneous, with

black melanoid lines, hard corky, up to 3 cm thick. Tubes slightly paler

than context, non-stratied, up to 2 cm long. Hyphal system trimitic;

generative hyphae with clamp connections; all hyphae IKI –, CB +;

tissues darkening in KOH. Generative hyphae in context colourless,

thin-walled, 2–3 µm diam; skeletal hyphae in context reddish brown,

thick-walled with a wide to narrow lumen or sub-solid, arboriform and

exuous, 2–6 μm diam; binding hyphae in context colourless, thick-

walled, branched and exuous, up to 2 µm diam. Generative hyphae

in tubes colourless, thin-walled, 2–3 μm diam; skeletal hyphae in

tubes reddish brown, thick-walled with a wide to narrow lumen or

sub-solid, arboriform and exuous, 2–4 μm diam; binding hyphae in

tubes colourless, thick-walled, branched and exuous, up to 2 μm

diam. Pileipellis composed of clamped generative hyphae, thick-

walled to sub-solid, apical cells clavate, inated and exuous, golden

yellow, about 28–37 × 5–8 μm, forming a regular palisade. Cystidia

and cystidioles absent. Basidia barrel-shaped, colourless, thin-

walled, 15–20 × 7–11 μm; basidioles broadly clavate, colourless, thin-

walled, 12–15 × 7–9 μm. Basidiospores ellipsoid to ovoid, truncated,

yellowish brown, IKI –, CB +, double-walled with moderately thick

walls, exospore wall smooth, endospore wall with dense spinules,

(7.5–)7.8–8.8(–8.9) × (4.9–)5–6(–6.5) μm, L = 8.23 μm, W = 5.49 μm,

Q = 1.47–1.53 (n = 60/2, with the turgid vesicular appendix excluded);

8.7–9.8(–10) × (4.9–)5–6(–6.1) μm, L = 9.23 μm, W = 5.52 µm, Q =

1.67–1.68 (n = 60/2, with the turgid vesicular appendix included).

Notes: Morphologically, Ganoderma guangxiense is very similar

to G. australe and they are not easy to separate. However,

phylogenetically, G. guangxiense and G. australe are divided into

two independent lineages with good support (Fig. 1). Ganoderma

daiqingshanense also has sessile and hard basidiomata with

blackish pilei, but it differs from G. guangxiense by having a

heterogeneous context, larger pores (4–5 per mm) and irregular

palisade of pileipellis structure (Zhao 1989a).

Ganoderma puerense B.K. Cui, J.H. Xing & Y.F. Sun, sp. nov.

MycoBank MB 839679. Figs 27, 28.

Diagnosis: Differs from other species in the genus by its woody

hard and thin basidiomata, yellowish brown pileal surface with

dense concentric black bands or furrows, and a margin lacerated

like petals.

Etymology: puerense (Lat.), refers to this species being collected

from Puer City of Yunnan Province.

Typus: China, Yunnan, Puer, Puer Forest Park, on living tree of

Cinnamomum, 17 Aug. 2019, Dai 20427 (holotype BJFC032095).

Description: Basidiomata annual, sessile or with short stipe,

usually growing together, imbricate, woody hard. Pilei solitary,

sub-orbicular to abelliform, applanate, up to 7.5 cm diam and

8 mm thick. Pileal surface yellowish brown to dark brown, dull,

Fig. 25. Basidiomata of Ganoderma guangxiense.

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glabrous, with dense concentric black bands or furrows and

slightly radial wrinkles; margin acute to obtuse, lacerated like

petals. Pore surface white when fresh, turning darker when

bruised, buff to pale brown when dry; pores circular to angular,

5–7 per mm; dissepiments moderately thick, entire. Context straw

yellow to dark brown, homogeneous, with black melanoid lines,

hard corky, up to 3 mm thick. Tubes dark brown, non-stratied,

up to 5 mm long. Stipe concolorous with pileal surface, cylindrical

Fig. 26. Microscopic structures of Ganoderma guangxiense (drawn from Cui 14453). A. Basidiospores. B. Apical cells from cuticle. C. Basidia and basidioles.

D. Hyphae from trama. E. Hyphae from context. Scale bars = 10 μm.

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Sun et al.

and solid, up to 1.5 cm long and 3 mm diam. Hyphal system

trimitic; generative hyphae with clamp connections; all hyphae

IKI –, CB +; tissues darkening in KOH. Generative hyphae in

context colourless, thin-walled, 2–3 μm diam; skeletal hyphae in

context pale yellowish brown, thick-walled with a wide to narrow

lumen or sub-solid, frequently arboriform and exuous, 2–5 μm

diam; binding hyphae in context colourless, thick-walled, rarely

branched and exuous, 1–2 μm diam. Generative hyphae in

tubes colourless, thin-walled, 2–3 μm diam; skeletal hyphae in

tubes pale yellowish brown, thick-walled with a wide to narrow

lumen or sub-solid, arboriform and exuous, 2–4 μm diam;

binding hyphae in tubes colourless, thick-walled, rarely branched

and exuous, 1–2 μm diam. Pileipellis composed of clamped

generative hyphae, thick-walled to sub-solid, apical cells clavate,

faintly inated and exuous, yellowish brown, about 25–40 × 5–7

μm, forming a regular palisade. Cystidia and cystidioles absent.

Basidia barrel-shaped, colourless, thin-walled, 12–20 × 7–10 µm;

basidioles clavate to fusiform, colourless, thin-walled, 11–18 ×

7–10 μm. Basidiospores ellipsoid to ovoid, truncated, yellowish

brown, IKI –, CB +, double-walled with slightly thick walls,

exospore wall smooth, endospore wall with dense spinules, (7–)

7.2–8.5(–8.8) × (4.8–)5–6(–6.4) μm, L = 7.83 μm, W = 5.31 μm,

Q = 1.47 (n = 60/1, with the turgid vesicular appendix excluded);

(8.5–)8.8–9.5(–9.8) × (4.8–)5–6.2(–6.8) μm, L = 9.1 μm, W =

5.52 μm, Q = 1.65 (n = 60/1, with the turgid vesicular appendix

included).

Notes: Ganoderma bicharacteristicum also has a dull pileal surface,

homogeneous context and was described from Yunnan Province,

but G. bicharacteristicum can be distinguished from G. puerense

by its stipitate basidiomata with black pileal surface, wood brown

pore surface and subglobose basidiospores (6.3–9 × 5.6–8.7 μm,

Zhang 1994).

Ganoderma subangustisporum B.K. Cui, J.H. Xing & Y.F. Sun,

sp. nov. MycoBank MB 839680. Figs 29, 30.

Diagnosis: Differs from other species in the genus by its dark-

red to near black pileal surface with concentric bands and radial

rugosities made by dark and continuous spots, margin acute and

thin, and ellipsoid basidiospores with truncated apex.

Etymology: subangustisporum (Lat.), refers to this species being

closely related to Ganoderma angustisporum.

Typus: China, Yunnan, Pingbian County, Daweishan National Nature

Reserve, on stump of angiosperm tree, 3 Aug. 2019, Cui 18592 (holotype

BJFC035453).

Materials examined: China, Yunnan, Pingbian County, Daweishan National

Nature Reserve, on stump of angiosperm tree, 3 Aug. 2019, Cui 18593

(BJFC035454), Cui 18594 (BJFC035455), Cui 18595 (BJFC035456), Cui

18596 (BJFC035457), Cui 18597 (BJFC035458).

Fig. 27. Basidiomata of Ganoderma puerense.

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Ganodermataceae from China

Description: Basidiomata annual, sessile or with short and lateral

stipe, hard corky. Pilei solitary, sub-circular to abelliform, up to

8 cm diam and 1.6 cm thick. Pileal surface dark reddish brown,

laccate, glabrous, with concentric bands and radial rugose made

by dark and continuous spots; margin acute, thin, entire. Pore

surface white when fresh, turning darker when bruised, buff to

pale straw yellow when dry; pores circular to angular, 4–6 per

mm; dissepiments slightly thick, entire. Context cinnamon brown,

homogeneous, with pale melanoid lines, hard corky, up to 9 mm

thick. Tubes pale brown, non-stratied, up to 9 mm long. Stipe

Fig. 28. Microscopic structures of Ganoderma puerense (drawn from Dai 20427). A. Basidiospores. B. Apical cells from cuticle. C. Basidia and basidioles.

D. Hyphae from trama. E. Hyphae from context. Scale bars = 10 μm.

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Sun et al.

concolourous with pileal surface, cylindrical and solid, up to 1 cm

long and 1.5 cm diam. Hyphal system trimitic; generative hyphae

with clamp connections; all hyphae IKI –, CB +; tissues darkening

in KOH. Generative hyphae in context colourless, thin-walled, 3–4

μm diam; skeletal hyphae in context yellowish brown, thick-walled

with a wide to narrow lumen or sub-solid, arboriform and exuous,

3–5 μm diam; binding hyphae in context colourless, thick-walled,

branched and exuous, up to 2 µm diam. Generative hyphae in

tubes colourless, thin-walled, 3–4 μm diam; skeletal hyphae in

tubes yellowish brown, thick-walled with a wide to narrow lumen or

sub-solid, arboriform and exuous, 2–5 μm diam; binding hyphae

in tubes colourless, thick-walled, branched and exuous, up to

2 μm diam. Pileipellis composed of clamped generative hyphae,

thick-walled to sub-solid, apical cells clavate, inated and exuous,

dark yellowish brown to reddish brown, about 32–50 × 7–12

μm, forming a regular palisade. Cystidia and cystidioles absent.

Basidia barrel-shaped, colourless, thin-walled, 13–19 × 10–13

μm; basidioles broadly clavate, colourless, thin-walled, 13–18 ×

6–12 μm. Basidiospores broadly ellipsoid to ellipsoid, truncated,

yellowish brown, IKI –, CB +, double-walled with distinctly thick

walls, exospore wall smooth, endospore wall with dense spinules,

10–11.8(–12.1) × (5–)5.1–7(–7.3) μm, L = 10.88 μm, W = 6.13

μm, Q = 1.72–1.84 (n = 60/2, with the turgid vesicular appendix

excluded); (10.3–)10.7–12.8(–13.1) × (5–)5.2–7(–7.3) μm, L =

11.72 μm, W = 6.02 μm, Q = 1.93–1.97 (n = 60/2, with the turgid

vesicular appendix included).

Notes: Ganoderma subangustisporum clusters with G.

angustisporum with high support in the phylogenetic analyses

(Fig. 1). Morphologically, G. angustisporum differs from G.

subangustisporum by having large basidiomata with lacerated

margin, darker pileal surface with concentric furrows, and narrower

basidiospores (9–11.3 × 4–5.2 µm, Xing et al. 2018).

Ganoderma subellipsoideum B.K. Cui, J.H. Xing & Y.F. Sun, sp.

nov. MycoBank MB 839681. Figs 31, 32.

Diagnosis: Differs from other species in the genus by its subsessile

and hard basidiomata, dark yellowish brown to blackish pileal

surface with dense concentric furrows, small pores and stratied

tubes, and ellipsoid basidiospores with truncated apex.

Etymology: subellipsoideum (Lat.), refers to this species being

closely related to Ganoderma ellipsoideum.

Typus: Malaysia, Kuala Lumpur, Ecological Forest Park, on fallen trunk of

angiosperm tree, 8 Dec. 2019, Cui 18325 (holotype BJFC035184).

Additional materials examined: Malaysia, Selangor, Kota Damansara,

National Forest Reserve, on fallen trunk of angiosperm tree, 6 Dec. 2019,

Cui 18241 (BJFC035100); Kuala Lumpur, Ecological Forest Park, on fallen

trunk of angiosperm tree, 8 Dec. 2019, Cui 18327(BJFC035186).

Fig. 29. Basidiomata of Ganoderma subangustisporum.

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Ganodermataceae from China

Description: Basidiomata annual, sessile or with short and lateral

stipe, woody hard. Pilei solitary, sub-circular to abelliform,

applanate, up to 9.5 cm diam and 2 cm thick. Pileal surface dark

yellowish brown to near black when fresh becoming yellowish

brown or greyish brown when dry, slightly laccate when fresh

becoming dull when dry, glabrous, with dense concentric furrows;

margin obtuse, entire. Pore surface white when fresh, turning

darker when bruised, cream to buff when dry; pores circular, 6–8

per mm; dissepiments moderately thick, entire. Context cinnamon

brown, homogeneous, with black melanoid lines, hard corky, up to

1 cm thick. Tubes dark brown, stratied by a layer of context, up

to 9 mm long. Stipe concolourous with pileal surface, cylindrical

and solid, up to 1.2 cm long and 1.3 cm diam. Hyphal system

trimitic; generative hyphae with clamp connections; all hyphae IKI

–, CB +; tissues darkening in KOH. Generative hyphae in context

colourless, thin-walled, 2–4 μm diam; skeletal hyphae in context

Fig. 30. Microscopic structures of Ganoderma subangustisporum (drawn from Cui 18592). A. Basidiospores. B. Apical cells from cuticle. C. Basidia and

basidioles. D. Hyphae from trama. E. Hyphae from context. Scale bars = 10 μm.

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Sun et al.

pale brown, thick-walled with a wide to narrow lumen or sub-solid,

arboriform and exuous, 2–5 μm diam; binding hyphae in context

colourless, thick-walled, branched and exuous, up to 1.5 μm diam.

Generative hyphae in tubes colourless, thin-walled, 2–3 μm diam;

skeletal hyphae in tubes yellowish brown, thick-walled with a wide

to narrow lumen or sub-solid, arboriform and exuous, 2–4 μm

diam; binding hyphae in tubes colourless, thick-walled, branched

and exuous, up to 1 μm diam. Pileipellis composed of clamped

generative hyphae, thick-walled, apical cells clavate, exuous,

pale yellowish brown, about 30–40 × 3–5 μm, forming a regular

palisade. Cystidia and cystidioles absent. Basidia barrel-shaped,

colourless, thin-walled, 12–17 × 9–11 µm; basidioles broadly

clavate, colourless, thin-walled, 11–15 × 6–10 μm. Basidiospores

ellipsoid, truncated, yellowish brown, IKI –, CB +, double-walled

with moderately thick walls, exospore wall smooth, endospore wall

with dense spinules, 6–6.9(–7) × 4–5.3(–5.5) μm, L = 6.46 μm,

W = 4.62 μm, Q = 1.36–1.44 (n = 60/2, with the turgid vesicular

appendix excluded); (8–)8.2–9.6(–10) × (4–)4.2–5.2(–6) μm, L =

8.89 μm, W = 4.69 μm, Q = 1.89–1.9 (n = 60/2, with the turgid

vesicular appendix included).

Notes: In the phylogenetic analyses, G. subellipsoideum is closely

related to G. ellipsoideum. Ganoderma ellipsoideum was described

from Hainan Province and it has a yellowish brown pileal surface

with alternating brownish orange to yellowish brown zones,

heterogeneous context and non-stratied tubes, and ellipsoid

basidiospores of smaller size (6.1–7.3 × 3.7–4.6 μm, Hapuarachchi

et al. 2018b) which make it different from G. subellipsoideum.

Ganoderma subexipes B.K. Cui, J.H. Xing & Y.F. Sun, sp. nov.

MycoBank MB 839675. Figs 33, 34.

Diagnosis: Differs from other species in the genus by its small and

dorso-laterally stipitate basidiomata, dark yellow to orange-brown

and laccate pileal surface, wavy margin, and obviously truncated

basidiospores.

Etymology: subexipes (Lat.), refers to this species being closely

related to Ganoderma exipes.

Typus: China, Guangdong, Renhua County, Danxiashan Nature Reserve,

on stump of angiosperm tree, 4 Jun. 2019, Cui 17257 (holotype

BJFC034115).

Additional materials examined: China, Guangdong, Renhua County,

Danxiashan Nature Reserve, on root of angiosperm tree, 4 Jun. 2019,

Cui 17247 (BJFC034105); on stump of angiosperm tree, 4 Jun. 2019,

Cui 17258 (BJFC034116); Jiangxi, Shangrao, Daomaoshan Park, on

the ground of Cyclobalanopsis jenseniana, 30 Aug. 2021, Dai 23665

(BJFC038237).

Description: Basidiomata annual, dorso-laterally stipitate, hard

corky. Pilei solitary, abelliform to shell-shaped, up to 3 cm diam

and 1 cm thick. Pileal surface dark yellow to orange-brown, laccate,

glabrous, with concentric bands and slightly radial rugose; margin

obtuse, entire, wavy when fresh, incurved when dry. Pore surface

buff to straw yellow when dry; pores circular to angular, 5–7 per

mm; dissepiments slightly thick to moderately thick, entire. Context

Fig. 31. Basidiomata of Ganoderma subellipsoideum.

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yellowish brown, homogeneous, without black melanoid lines, soft

corky, up to 1.6 mm thick. Tubes buff, non-stratied, up to 7 mm

long. Stipe dark-red, cylindrical and solid, sometimes budding

without pilei, up to 14cm long and 6 mm diam. Hyphal system

trimitic; generative hyphae with clamp connections; all hyphae IKI

–, CB +; tissues darkening in KOH. Generative hyphae in context

colourless, thin-walled, 2–3 μm diam; skeletal hyphae in context

pale yellow, thick-walled with a wide to narrow lumen or sub-solid,

arboriform and exuous, 2–5 μm diam; binding hyphae in context

colourless, thick-walled, branched and exuous, up to 2 μm diam.

Generative hyphae in tubes colourless, thin-walled, 2–3 μm diam;

skeletal hyphae in tubes pale yellow, thick-walled with a wide to

Fig. 32. Microscopic structures of Ganoderma subellipsoideum (drawn from Cui 18325). A. Basidiospores. B. Apical cells from cuticle. C. Basidia and

basidioles. D. Hyphae from trama. E. Hyphae from context. Scale bars = 10 μm.

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Sun et al.

narrow lumen or sub-solid, arboriform and exuous, 2–4 μm

diam; binding hyphae in tubes colourless, thick-walled, branched

and exuous, up to 2 μm diam. Pileipellis composed of clamped

generative hyphae, thick-walled to sub-solid, apical cells clavate,

slightly inated and exuous, golden yellow, about 27–35 × 4–8

μm, forming a regular palisade. Cystidia and cystidioles absent.

Basidia barrel-shaped, colourless, thin-walled, 15–22 × 9–11 µm;

basidioles broadly clavate, colourless, thin-walled, 14–18 × 7–10

μm. Basidiospores ellipsoid to ovoid, truncated, yellowish brown,

IKI –, CB +, double-walled with distinctly thick walls, exospore wall

smooth, endospore wall with dense spinules, (6.3–)6.7–9.8(–10.1)

× (4–)4.2–6.5(–7) μm, L = 8.2 μm, W = 5.37 µm, Q = 1.53–1.65

(n = 60/2, with the turgid vesicular appendix excluded); (7.6–)7.8–

11(–11.5) × (4–)4.2–6.6(–6.8) μm, L = 9.25 μm, W = 5.46 μm, Q =

1.64–1.74 (n = 60/2, with the turgid vesicular appendix included).

Notes: Ganoderma subexipes is distinguished by its small and

dorso-laterally stipitate basidiomata, dark yellow to orange-brown

pileal surface, wavy margin, and obviously truncated basidiospores.

Ganoderma exipes is sister to G. subexipes in phylogenetic tree

(Fig. 1), that species also has small and dorso-laterally stipitate

basidiomata with a laccate pileal surface, but it differs from G.

subexipes by its reddish brown pileal surface, round and smooth

margin, and basidiospores not obviously truncated (Steyaert 1972,

Ryvarden 1983, Wang & Wu 2014).

Ganoderma sublobatum B.K. Cui, J.H. Xing & Y.F. Sun, sp. nov.

MycoBank MB 839682. Figs 35, 36.

Diagnosis: Differs from other species in the genus by its annual

basidiomata, non-laccate pileal surface with shallow concentric

furrows and radial rugose, homogeneous context, non-stratied

tubes and ellipsoid basidiospores with truncated apex.

Etymology: sublobatum (Lat.), refers to this species being closely

related to Ganoderma lobatum.

Typus: Australia, Queensland, Cairns, Mount Whiteld Conservation

Park, on stump of angiosperm tree, 18 May 2018, Cui 16804 (holotype

BJFC030103).

Additional materials examined: Australia, Queensland, Cairns, Mount

Whiteld Conservation Park, on stump of angiosperm tree, 18 May 2018,

Cui 16805 (BJFC030104), Cui 16806 (BJFC030105).

Description: Basidiomata annual, sessile or subsessile, hard corky

to woody hard. Pilei solitary, abelliform to reniform or shell-shaped,

applanate, up to 8 cm diam and 7 cm thick. Pileal surface pale brown

to dark brown, dull, glabrous, with shallowly concentric furrows and

radial rugose; margin obtuse, entire. Pore surface white to pale

brown when fresh, turning darker when bruised, pale grey when

Fig. 33. Basidiomata of Ganoderma subexipes.

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dry; pores circular, 5–6 per mm; dissepiments moderately thick,

entire. Context dark brown, homogeneous, with black melanoid

lines, corky, up to 3 mm thick. Tubes dark brown, non-stratied,

up to 7 mm long. Hyphal system trimitic; generative hyphae with

clamp connections; all hyphae IKI –, CB +; tissues darkening in

KOH. Generative hyphae in context colourless, thin-walled, 2.5–5

μm diam; skeletal hyphae in context pale brown, thick-walled

with narrow lumen or sub-solid, arboriform and exuous, 3.5–6.5

μm diam; binding hyphae in context colourless, thick-walled,

branched and exuous, 2–3 μm diam. Generative hyphae in tubes

colourless, thin-walled, 2–3.5 μm diam; skeletal hyphae in tubes

pale brown, thick-walled with narrow lumen or sub-solid, arboriform

and exuous, 2–4.5 μm diam; binding hyphae in tubes colourless,

thick-walled, branched and exuous, 1.2–2.5 μm diam. Pileipellis

composed of clamped generative hyphae, thin to slightly thick-

walled, apical cells clavate, faintly constricted with obvious branch,

Fig. 34. Microscopic structures of Ganoderma subexipes (drawn from Cui 17257). A. Basidiospores. B. Apical cells from cuticle. C. Basidia and basidioles.

D. Hyphae from trama. E. Hyphae from context. Scale bars = 10 μm.

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Sun et al.

dark brown, about 30–40 × 3–6 μm, forming an untidy palisade.

Cystidia and cystidioles absent. Basidia barrel-shaped, colourless,

thin-walled, 15–25 × 10–17 μm; basidioles clavate, colourless, thin-

walled, 16–22 × 7–14 μm. Basidiospores ellipsoid, truncated, pale

yellowish brown, IKI –, CB +, double-walled with distinctly thick

walls, exospore wall smooth, endospore wall with dense spinules,

(7.5–)9–11(–11.5) × (6–)6.5–8(–8.5) μm, L = 9.67 μm, W = 7.3 μm,

Q = 1.32 (n = 30/1, with the turgid vesicular appendix excluded);

(10.5–)11–12.5(–13) × 6.5–8.5(–9) μm, L = 11.41 μm, W = 7.25 μm,

Q = 1.57 (n = 30/1, with the turgid vesicular appendix included).

Notes: Ganoderma sublobatum was collected from Queensland in

Australia, it is closely related to G. lobatum. However, G. lobatum

has perennial basidiomata with darker pilei, white pore surface, and

smaller basidiospores (7–8.6 × 5–5.5 µm) which are different from

G. sublobatum.

Ganoderma tongshanense B.K. Cui, J.H. Xing & Y.F. Sun, sp.

nov. MycoBank MB 839683. Figs 37, 38.

Diagnosis: Differs from other species in the genus by its pale

yellowish brown pileal surface, large pores, homogeneous context,

non-stratied tubes, and ellipsoid and truncated basidiospores.

Etymology: tongshanense (Lat.), refers to the holotype of this

species being found at Tongshan County of Hubei Province.

Typus: China, Hubei, Tongshan County, Jiugongshan National Park,

on fallen trunk of angiosperm tree, 20 Oct. 2018, Cui 17168 (holotype

BJFC030468).

Description: Basidiomata annual, sessile and broadly attached,

hard corky. Pilei solitary, abelliform, up to 7 cm diam and 2.5 cm

thick. Pileal surface clay buff to pale yellowish brown, dull, glabrous,

with concentric furrows and slightly radial wrinkles; margin obtuse,

entire. Pore surface greyish white when fresh, turning darker when

bruised, pale brown when dry; pores circular, shallow, 3–4 per

mm; dissepiments moderately thick, entire. Context dark brown,

homogeneous, without black melanoid lines, hard corky, up to 1.6

cm thick. Tubes slightly paler than context, non-stratied, stuffed

with white mycelium, up to 9 mm long. Hyphal system trimitic;

generative hyphae with clamp connections; all hyphae IKI –, CB +;

tissues darkening in KOH. Generative hyphae in context colourless,

thin-walled, 2–4 μm diam; skeletal hyphae in context dark brown,

thick-walled with a wide to narrow lumen or sub-solid, arboriform

and exuous, 2–5 μm diam; binding hyphae in context colourless,

thick-walled, branched and exuous, up to 2 µm diam. Generative

hyphae in tubes colourless, thin-walled, 2–4 μm diam; skeletal

hyphae in tubes dark brown, thick-walled with a wide to narrow

lumen or sub-solid, arboriform and exuous, 2–5 μm diam; binding

hyphae in tubes colourless, thick-walled, branched and exuous,

up to 2 μm diam. Pileipellis composed of clamped generative

hyphae, thick-walled to sub-solid, apical cells clavate, inated and

Fig. 35. Basidiomata of Ganoderma sublobatum.

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Ganodermataceae from China

exuous, pale yellowish brown, about 30–45 × 6–8 μm, forming a

regular palisade. Cystidia and cystidioles absent. Basidia clavate,

colourless, thin-walled, 10–13 × 3–5 µm; basidioles in shape like

the basidia, colourless, thin-walled, 8–11 × 4–6 μm. Basidiospores

ellipsoid, truncated, pale yellowish brown, IKI –, CB +, double-

walled with distinctly thick walls, exospore wall smooth, endospore

wall with dense spinules, (8–)8.2–9.8(–10) × (5–)5.2–6.2(–6.5) μm,

L = 8.91 μm, W = 5.62 μm, Q = 1.59 (n = 60/1, with the turgid

vesicular appendix excluded); (9.2–)9.4–10.5(–11) × (5–)5.2–6.1(–

6.3) μm, L = 9.81 µm, W = 5.65 μm, Q = 1.74 (n = 60/1, with the

turgid vesicular appendix included).

Notes: Morphologically, G. tongshanense may be confused with G.

australe which is widely distributed in South China. They share similar

macro-morphological characters, but the latter has perennial and

larger basidiomata and stratied tubes (Patouillard 1889, Ryvarden

& Johansen 1980, Corner 1983). In the phylogenetic analyses, G.

tongshanense and G. australe are distinct from each other (Fig. 1).

Fig. 36. Microscopic structures of Ganoderma sublobatum (drawn from Cui 16804). A. Basidiospores. B. Basidia and basidioles. C. Hyphae from trama. D.

Hyphae from context. Scale bars = 10 μm.

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Sun et al.

Ganoderma yunlingense B.K. Cui, J.H. Xing & Y.F. Sun, sp. nov.

MycoBank MB 839684. Figs 39, 40.

Diagnosis: Differs from other species in the genus by its non-laccate

pileal surface with concentrically irregular ridges, extremely thin

context, stratied tubes, and ovoid to almond-shaped basidiospores

without spinules on the endospore wall.

Etymology: yunlingense (Lat.), refers to this species being collected

from the Yunling Mountains of Yunnan Province.

Typus: China, Yunnan, Lanping County, Luoguqing, on fallen trunk of

Quercus, 9 Sep. 2017, Cui 16288 (holotype BJFC029587).

Additional materials examined: China, Yunnan, Lijiang, Yulongxueshan,

on stump of Quercus, 16 Sep. 2018, Cui 17043 (BJFC030342), Cui 17060

(BJFC030359); Lanping County, Luoguqing, on fallen trunk of Quercus

semecarpifolia, 18 Sep. 2018, Cui 17161 (BJFC030461).

Description: Basidiomata perennial, sessile, hard corky to woody

hard. Pilei solitary, abelliform to shell-shaped, occasionally

dimidiate, up to 11.5 cm diam and 10 cm thick. Pileal surface

greyish brown to grey, dull, glabrous, with concentrically irregular

ridges; margin acute, entire, wavy and sometimes lacerated

as petals. Pore surface white when fresh, turning darker when

bruised, straw yellow when dry; pores circular to angular, 4–6 per

mm; dissepiments moderately thick, entire. Context extremely thin,

without black melanoid lines. Tubes brown to dark brown, distinctly

stratied, up to 5 cm long. Hyphal system trimitic; generative

hyphae with clamp connections; all hyphae IKI –, CB +; tissues

darkening in KOH. Generative hyphae in context colourless,

thin-walled, 2–3.5 µm diam; skeletal hyphae in context yellowish

brown, thick-walled with narrow lumen or sub-solid, arboriform and

exuous, 3–5.5 µm diam; binding hyphae in context colourless,

thick-walled, branched and exuous, 1.5–3 µm diam. Generative

hyphae in tubes colourless, thin-walled, 2–3 μm diam; skeletal

hyphae in tubes yellowish brown, thick-walled with narrow lumen or

sub-solid, arboriform and exuous, 2.5–5 μm diam; binding hyphae

in tubes colourless, thick-walled, branched and exuous, 1.5–3 μm

diam. Pileipellis composed of clamped generative hyphae, thick-

walled, apical cells clavate, exuous, golden yellow to pale brown,

about 25–40 × 4–7 μm, forming a regular palisade. Cystidia and

cystidioles absent. Basidia barrel-shaped, colourless, thin-walled,

15–20 × 8–13 µm; basidioles clavate, colourless, thin-walled,

13–17 × 6–10 μm. Basidiospores ovoid to almond-shaped, not

obviously truncated, pale yellowish brown, IKI –, CB +, double-

walled with slightly thick walls, exospore wall smooth, endospore

wall without spinules, (7.5–)8–10(–10.5) × (4.3–)4.8–5.5(–6) µm, L

= 8.93 µm, W = 5.16 µm, Q = 1.70–1.73 (n = 60/2, with the turgid

vesicular appendix excluded).

Fig. 37. Basidiomata of Ganoderma tongshanense.

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Ganodermataceae from China

Notes: Ganoderma yunlingense is characterised by its non-laccate

pileal surface with concentrically irregular ridges, extremely thin

context, stratied and long tubes, basidiospores not obviously

truncated, without spinules on the endospore wall. It may be

confused with G. acontextum because of the dull pileal surface,

thin context and basidiospores not obviously truncated without any

ornamentation; however, G. acontextum was collected from the

United States of America and has reddish brown pileal surface with

dense concentric furrows, and non-stratied tubes.

Fig. 38. Microscopic structures of Ganoderma tongshanense (drawn from Cui 17168). A. Basidiospores. B. Apical cells from cuticle. C. Basidia and

basidioles. D. Hyphae from trama. E. Hyphae from context. Scale bars = 10 μm.

358

Sun et al.

Notes on accepted species of Ganoderma recorded

from China

Ganoderma ahmadii Steyaert, Persoonia 7: 91. 1972. MycoBank

MB 314303.

For a detailed description of Ganoderma ahmadii, see Steyaert

(1972) and Zhao (1989b).

Notes: Ganoderma ahmadii was described from Pakistan by

Steyaert (1972). Zhao (1989b) examined an authentic specimen

collected from Pakistan, but the description regarding the reddish

brown pileal surface and homogeneous context is different to the

original description. Wang (2005) studied one specimen of G.

ahmadii collected from Sichuan Province in China and the ITS1-

ITS2 sequences are similar to the records in GenBank. So, G.

ahmadii was conrmed in Sichuan Province in China based on

morphological and phylogenetic analyses.

Ganoderma angustisporum J.H. Xing et al., MycoKeys 34: 98.

2018. MycoBank MB 823320.

For a detailed description of Ganoderma angustisporum, see Xing

et al. (2018).

Notes: Ganoderma angustisporum was recently described from

China and classied in Ganoderma according to its narrow and

truncated basidiospores in Ganoderma (Xing et al. 2018). It is also

found in Malaysia, Sri Lanka and Thailand.

Ganoderma applanatum (Pers.) Pat., Hyménomyc. Eur. (Paris):

143. 1887. MycoBank MB 119872. Figs 41, 42.

Basionym: Boletus applanatus Pers., Observ. Mycol. (Lipsiae) 2:

2. 1800.

Description: Basidiomata perennial, sessile and broadly attached,

hard corky to woody hard. Pilei solitary or imbricate, variable,

applanate, abelliform to shell-like or ungulate, up to 23 cm

diam and 7 cm thick. Pileal surface pale brown to dark brown,

dull, glabrous, with shallow to deep concentric furrows; margin

obtuse, entire and wavy. Pore surface white to pale brown when

fresh, turning darker when bruised, greyish white to straw yellow

when dry; pores circular to angular, 4–7 per mm; dissepiments

moderately thick, mostly entire. Context yellowish brown to dark

brown, homogeneous, with black melanoid lines when mature,

corky, up to 3 cm thick. Tubes concolorous with context, stratied

by a layer of context, up to 4 cm long. Hyphal system trimitic;

generative hyphae with clamp connections; all hyphae IKI –,

CB +; tissues darkening in KOH. Generative hyphae in context

colourless, thin-walled, 2.5–7 µm diam; skeletal hyphae in context

pale yellowish brown, thick-walled with a wide to narrow lumen or

sub-solid, frequently arboriform and exuous, 3.5–6.5 µm diam;

binding hyphae in context colourless, thick-walled, branched and

exuous, 2–3 µm diam. Generative hyphae in tubes colourless,

thin-walled, 2–3.5 μm diam; skeletal hyphae in tubes pale brown

to dark brown, thick-walled with a wide to narrow lumen or sub-

solid, frequently arboriform and exuous, 2–4.5 μm diam; binding

hyphae in tubes colourless, thick-walled, branched and exuous,

1–2.5 μm diam. Cystidia, cystidioles, basidia and basidioles

Fig. 39. Basidiomata of Ganoderma yunlingense.

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Ganodermataceae from China

absent. Basidiospores ellipsoid, truncated, yellowish to pale brown,

IKI –, CB +, double-walled with moderately thick walls, exospore

wall smooth, endospore wall with dense spinules, (5–)5.5–7(–8) ×

(4–)4.1–5.2(–6) µm, L = 6.25 μm, W = 4.56 μm, Q = 1.37 (n=30/1,

with the turgid vesicular appendix included).

Additional materials examined: China, Jilin, Jiaohe, Hongyegu Park, on

stump of angiosperm tree, 1 Aug. 2016, Cui 14062 (BJFC028930); on

stump of Betula, 1 Aug. 2016, Cui 14070 (BJFC028938); Fusong County,

Changbaishan Nature Reserve, on fallen trunk of angiosperm tree, 3 Aug.

2016, Cui 14121 (BJFC028989).

Notes: Ganoderma applanatum was rstly described from

Europe and has a Holarctic distribution (Steyaert 1972, Moncalvo

& Ryvarden 1997, Hapuarachchi et al. 2019b). Ganoderma

applanatum is similar with G. australe based on perennial and

sessile basidiomata, non-laccate and pale pileal surface, brown to

dark brown context, but the latter has larger basidiospores (7–12

× 5–8 µm, Ryvarden 2004b). Besides, G. applanatum and G.

australe can be separated in the phylogenetic analyses (Fig. 1).

Ganoderma australe (Fr.) Pat., Bull. Soc. Mycol. Fr. 5: 65. 1889.

MycoBank MB 100745.

Basionym: Polyporus australis Fr., Elench. Fung. (Greifswald) 1:

108. 1828.

Synonyms: Ganoderma triangulum J.D. Zhao & L.W. Hsu, Acta

Mycol. Sin. 3: 18. 1984.

Ganoderma ungulatum J.D. Zhao & X.Q. Zhang, Acta Mycol. Sin.

3: 19. 1984.

Ganoderma bawanglingense J.D. Zhao & X.Q. Zhang, Acta Mycol.

Sin. 6: 205. 1987.

Ganoderma mirivelutinum J.D. Zhao, Acta Mycol. Sin. 7: 206. 1987.

For a detailed description of G. australe, see Ryvarden (2004b) and

Hapuarachchi et al. (2018b).

Fig. 40. Microscopic structures of Ganoderma yunlingense (drawn from Cui 16288). A. Basidiospores. B. Basidia and basidioles. C. Hyphae from trama.

D. Hyphae from context. Scale bars = 10 μm.

360

Sun et al.

Notes: Ganoderma australe was described from an island in the

Pacic Ocean, but the type specimen has been lost and the only

specimen deposited at the Royal Botanic Gardens Kew was from

Europe which is inconsistent with its tropical distribution (Moncalvo

& Ryvarden 1997). Cao (2013) suggested that G. bawanglingense,

G. mirivelutinum, G. triangulum and G. ungulatum should be

regarded as the synonyms of G. applanatum, but their tropical

distribution and morphological similarity of G. australe support that

they should be treated as the synonyms of G. australe. Ganoderma

australe may be confused with G. adspersum, G. applanatum and

G. gibbosum in morphology. Previously, many specimens collected

from South China were identied as “Ganoderma australe”. Two

specimens from Australia are recognised as G. australe by Costa-

Rezende et al. (2017), and most specimens collected from South

China clustered with G. gibbosum (Fig. 1). Whether there is true G.

australe in China is unknown and more detailed studies about the

G. australe complex should be done.

Ganoderma boninense Pat., Bull. Soc. Mycol. Fr. 5: 72. 1889.

MycoBank MB 100062.

For a detailed description of G. boninense, see Steyaert (1967) and

Ryvarden (1983).

Notes: Ganoderma boninense was described from the Bonin

Islands in Japan, and it has been recognised as the main pathogen

of oil palm trees causing a basal stem rot (Pilotti 2005). Ganoderma

boninense can be distinguished by its reddish and shiny pileal

surface, irregular apical cells with swellings and protuberance of

pileipellis, and oblong ellipsoid basidiospores (Ryvarden 1983).

Cao (2013) compared the specimens collected from Hainan

Province and Japan, and they showed very similar morphological

characters and ITS sequences.

Ganoderma calidophilum J.D. Zhao et al., Acta Microbiol. Sin. 19:

270. 1979. MycoBank MB 314307.

For a detailed description of G. calidophilum, see Zhao et al. (1979)

and Luangharn et al. (2021).

Notes: Zhao et al. (1979) described Ganoderma calidophilum from

Hainan Province based on its small basidiomata with heterogenous

context, whitish pore surface and large basidiospores (10–12.1 ×

6.2–8.7 μm). The sequences of G. calidophilum used in this study

were downloaded from GenBank, and showed to be distinct from

G. exipes (Fig. 1), which has been considered as a doubtful

synonym (Wang & Wu 2014).

Ganoderma casuarinicola J.H. Xing et al., MycoKeys 34: 100.

2018. MycoBank MB 823321.

For a detailed description of G. casuarinicola, see Xing et al. (2018)

and Luangharn et al. (2019).

Fig. 41. Basidiomata of Ganoderma applanatum.

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Ganodermataceae from China

Fig. 42. Microscopic structures of Ganoderma applanatum (drawn from He 2139). A. Basidiospores. B. Hyphae from trama. C. Hyphae from context. Scale

bars = 10 μm.

362

Sun et al.

Notes: Ganoderma casuarinicola was collected on a living tree of

Casuarina equisetifolia from Guangdong Province in China by Xing

et al. (2018). Luangharn et al. (2019) reported G. casuarinicola,

where it was found on Pinus kesiya stump, as a new record

from Thailand. There are some differences such as applanate

to dimidiate pilei, longer tubes and larger basidiospores when

comparing these specimens with the type of G. casuarinicola (Dai

16336). Geographical and climatic divergences may be the reason

for the intraspecic differences (Boddy et al. 2014).

Ganoderma ellipsoideum Hapuar. et al., Mycosphere 9: 951.

2018. MycoBank MB 554384.

For a detailed description of G. ellipsoideum, see Hapuarachchi et

al. (2018b).

Notes: Ganoderma ellipsoideum is distinguished by its ellipsoid

spores (6.1–7.3 × 3.7–4.6 μm) with distinct spinules on the

endospore wall. Hapuarachchi et al. (2018b) stated that G.

ellipsoideum was known only from the type locality of Hainan

Province in China. In this study, several specimens collected from

Yunnan and Guangdong provinces showed similar morphological

features and close phylogenetic relationships with G. ellipsoideum.

Ganoderma exipes Pat., Bull. Soc. Mycol. Fr. 23: 75. 1907.

MycoBank MB 249905. Figs 43, 44.

Synonyms: Ganoderma atrum J.D. Zhao et al., Acta Microbiol. Sin.

19: 268. 1979.

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

269. 1979.

Ganoderma parviungulatum J.D. Zhao & X.Q. Zhang, Acta Mycol.

Sin. 5: 88. 1986.

Description: Basidiomata annual, dorso-laterally stipitate, hard

corky to woody hard. Pilei solitary, variable, abelliform to shell-like

or circular, up to 5 cm diam and 1.5 cm thick. Pileal surface dark

brown to reddish brown, strongly laccate, glabrous, with obvious

concentric furrows and slightly radial rugose; margin obtuse, entire,

slightly incurved. Pore surface white when fresh, turning darker

when bruised, light buff when dry; pores circular to angular, 4–6 per

mm; dissepiments slightly thick, entire. Context heterogeneous, the

upper layer pale yellowish brown, the lower layer dark brown, with

black melanoid lines, corky, up to 3 mm thick. Tubes dark brown,

non-stratied, up to 1.2 cm long. Stipe reddish brown to purplish

black, attened to cylindrical, up to 25 cm long and 6 mm diam.

Hyphal system trimitic; generative hyphae with clamp connections;

all hyphae IKI –, CB +; tissues darkening in KOH. Generative

hyphae in context colourless, thin-walled, 1.5–3.5 μm diam;

skeletal hyphae in context pale yellowish brown, thick-walled with

a narrow lumen or sub-solid, arboriform and exuous, 2.5–6 μm

diam; binding hyphae in context colourless, thick-walled, branched

and exuous, 1–2 μm diam. Generative hyphae in tubes colourless,

thin-walled, occasionally branched, 2–3 μm diam; skeletal hyphae

in tubes pale brown to dark brown, thick-walled with a narrow

lumen to sub-solid, frequently arboriform and exuous, 2–4 μm

diam; binding hyphae in tubes colourless, thick-walled, branched

and exuous, 0.5–1 μm diam. Pileipellis composed of clamped

generative hyphae, thick-walled to sub-solid, apical cells clavate,

slightly inated, pale brown to yellowish brown, 30–45 × 5–10

μm, forming a regular palisade. Cystidia and cystidioles absent.

Basidia barrel-shaped, colourless, thin-walled, 23–30 × 12–17 µm;

basidioles in shape like the basidia, colourless, thin-walled, 17–23

× 8–13 μm. Basidiospores ellipsoid, not obviously truncated, pale

yellowish brown, IKI –, CB +, double-walled with moderately thick

walls, exospore wall smooth, endospore walls with dense spinules,

(6.5–)7–9(–9.5) × (4–)4.8–5.5(–6) μm, L = 8.17 μm, W = 5.23 μm,

Q = 1.56 (n = 30/1, with the turgid vesicular appendix excluded).

Materials examined: China, Guangdong, Guangzhou, campus of Sun

Yat-Sen University, on stump of angiosperm tree, 20 Jul. 2012, Cui 17209

(BJFC030563); Hainan, Qiongzhong County, Limushan Forest Park, on

ground of angiosperm forest, 16 Jun. 2016, Cui 13841 (BJFC028707);

Changjiang County, Bawangling Nature Reserve, on ground of angiosperm

forest, 18 Jun. 2016, Cui 13863 (BJFC028729); Yunnan, Puer, Puer Forest

Park, on rotten angiosperm wood, 17 Aug. 2019, Dai 20461 (BJFC032129).

Notes: Ganoderma exipes was rst described from Vietnam by

Patouillard (1907), and many researchers have conducted detailed

studies on it (Steyaert 1972, Ryvarden 1983, Hapuarachchi et

al. 2019b). Cao (2013) considered G. atrum, G. hainanense

Fig. 43. Basidiomata of Ganoderma exipes.

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Ganodermataceae from China

Fig. 44. Microscopic structures of Ganoderma exipes (drawn from Cui 13882). A. Basidiospores. B. Apical cells from cuticle. C. Basidia and basidioles. D.

Hyphae from trama. E. Hyphae from context. Scale bars = 10 μm.

364

Sun et al.

and G. parviungulatum as synonyms of G. exipes based on

comprehensive observations of the holotype specimens.

Ganoderma gibbosum (Blume & T. Nees) Pat., Ann. Jard. Bot.

Buitenzorg, suppl. 1: 114. 1897. MycoBank MB 250058.

Basionym: Polyporus gibbosus Blume & T. Nees, Nova Acta Phys.-

Med. Acad. Caes. Leop.-Carol. Nat. Cur. 13: 19. 1826.

For a detailed description of G. gibbosum, see Luangharn et al.

(2020).

Notes: Ganoderma gibbosum was rst described from Java,

however, the type specimen was lost (Moncalvo & Ryvarden, 1997).

In previous studies, G. gibbosum was regarded as a synonym of G.

applanatum or classied in the G. applanatum–australe complex

based on the non-laccate basidiomata (Zhao 1989b, Moncalvo &

Ryvarden 1997). Luangharn et al. (2020) conducted a study on the

taxonomy of G. gibbosum collected from Kunming, China, and the

results showed that samples of G. gibbosum from Asia and South

America formed two different lineages. In this study, specimens of

G. gibbosum collected from Guangdong, Guangxi and Sichuan

provinces clustered together and formed a well-supported lineage

(Fig. 1).

Ganoderma hoehnelianum Bres., Annls Mycol. 10: 502. 1912.

MycoBank MB 243431. Figs 45, 46.

Synonym: Ganoderma shangsiense J.D. Zhao, Acta Mycol. Sin. 7:

17. 1988.

Description: Basidiomata perennial, sessile or broadly attached to

laterally stipitate, hard corky to woody hard. Pilei solitary, variable,

abelliform or shell-like to reniform, applanate, up to 10 cm diam

and 2.2 cm thick. Pileal surface yellowish brown to dark brown, dull,

glabrous, with obvious concentric furrows; margin acute to obtuse,

entire. Pore surface white when fresh, turning darker when bruised,

straw yellow to pale yellowish brown when dry; pores circular to

angular, 3–6 per mm; dissepiments moderately thick, entire. Context

heterogeneous, the upper layer pale yellowish brown, the lower layer

dark brown, with black melanoid lines, corky, up to 1 cm thick. Tubes

light buff to greyish brown, stratied, up to 1.2 cm long. Stipe dark

brown, attened, up to 3 cm long and 6 mm diam. Hyphal system

trimitic; generative hyphae with clamp connections; all hyphae IKI

–, CB +; tissues darkening in KOH. Generative hyphae in context

absent; skeletal hyphae in context pale yellowish brown, thick-walled

with a wide to narrow lumen or sub-solid, frequently arboriform and

exuous, 2–4 µm diam; binding hyphae in context colourless, thick-

walled, branched and exuous, 1–2.5 µm diam. Generative hyphae

in tubes colourless, thin-walled, 2.3–3.5 μm diam; skeletal hyphae in

tubes pale brown to dark brown, thick-walled with a narrow lumen or

sub-solid, frequently arboriform and exuous, 2–5 μm diam; binding

hyphae in tubes colourless, thick-walled, branched and exuous,

1.5–3 μm diam. Cystidia and cystidioles absent. Basidia absent;

basidioles barrel-shaped, colourless, thin-walled, 21–30 ×9–13 μm.

Basidiospores subglobose, not obviously truncated, pale yellowish

brown, IKI –, CB +, double-walled with distinctly thick walls, exospore

walls smooth, endospore walls with dense spinules, (6.5–)7–8.2(–9)

× (–5.5)5.8–7.8(–8) µm, L = 7.43 μm, W = 6.73 μm, Q = 1.1 (n =

30/1).

Fig. 45. Basidiomata of Ganoderma hoehnelianum.

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Ganodermataceae from China

Fig. 46. Microscopic structures of Ganoderma hoehnelianum (drawn from Dai 16166). A. Basidiospores. B. Basidioles. C. Hyphae from trama. D. Hyphae

from context. Scale bars = 10 μm.

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Sun et al.

Materials examined: China, Hainan, Ledong County, Jianfengling

Nature Reserve, on stump of angiosperm tree, 19 Jun. 2016, Cui 13904

(BJFC028770); Guangxi, Shangsi County, Shiwandashan Forest Park, on

dead tree of angiosperm, 6 Jul. 2016, Cui 13982 (BJFC028850); Nanning,

Guangxi Academy of Forestry, on living tree of angiosperm, 21 Aug. 2019,

Dai 20783 (BJFC032450); Yunnan, Mengla County, Wangtianshu Park, 20

Jul. 2014, Dai 13915 (BJFC017645).

Notes: Ganoderma hoehnelianum was described by Bresadola

from Java. It has typically ganodermoid macro-morphological

features, but the obviously amaurodermoid basidiospores make

G. hoehnelianum different from other species. In the phylogenetic

analyses, G. hoehnelianum grouped in the Ganoderma clade,

and formed an independent lineage with high support (Fig. 1).

Wang & Wu (2010) and Cao (2013) regarded G. shangsiense as

a synonym of G. hoehnelianum after the studies on the holotype of

G. shangsiense.

Ganoderma leucocontextum T.H. Li et al., Mycoscience 56: 82.

2015. MycoBank MB 804187.

For a detailed description of G. leucocontextum, see Li et al. (2015).

Notes: Ganoderma leucocontextum as a member of the G. lucidum

complex shares reddish brown and laccate pilei and truncated

basidiospores, but it is distinguished by the white context Li et

al. (2015). In the current study, G. leucocontextum grouped with

G. weixiense (Fig. 1). According to the comparison between G.

leucocontextum and G. weixiense in Ye et al. (2019), the size of

pores (4–6 per mm vs 2–4 per mm) and basidiospores (9.5–12.5 ×

7–9 μm vs 6–8 × 3–4 μm) are the main differences.

Ganoderma lingzhi Sheng H. Wu et al., Fungal Divers. 56: 54.

2012. MycoBank MB 564240.

For a detailed description of G. lingzhi, see Cao et al. (2012).

Notes: Cao et al. (2012) revised the taxonomic status of the widely

cultivated “Ganoderma lucidum” in China and described it as a

new species called G. lingzhi based on geographical distribution,

morphological features and phylogenetic analyses. Ganoderma

lingzhi is widely distributed in temperate and subtropical areas of

China, and it has also been reported from Korea and Laos (Kim et

al. 2001, Hapuarachchi et al. 2019b).

Ganoderma lucidum (Curtis) P. Karst., Revue Mycol., Toulouse 3:

17. 1881. MycoBank MB 148413. Figs 47, 48.

Basionym: Boletus lucidus Curtis, Fl. Londin. 1: 72. 1781.

Synonym: Ganoderma cantharelloideum M.H. Liu, Acta Mycol. Sin.

8: 279. 1989.

Description: Basidiomata annual, laterally stipitate, hard corky

to woody hard. Pilei solitary, variable, abelliform or shell-like to

circular, up to 11 cm diam and 3 cm thick. Pileal surface yellowish

brown to reddish brown, laccate, glabrous, with concentric furrows;

margin acute to obtuse, entire, slightly wavy. Pore surface white

when fresh, turning darker when bruised, pale yellow to straw

yellow when dry; pores circular, 4–6 per mm; dissepiments slightly

thick, mostly entire. Context heterogeneous, the upper layer cream

to buff, the lower layer clay-buff, without black melanoid lines, soft

corky, up to 1.8 cm thick. Tubes pale brown, non-stratied, up to

1.2 cm long. Stipe reddish brown to purplish black, attened to

cylindrical, up to 12 cm long and 1.5 cm diam. Hyphal system

trimitic; generative hyphae with clamp connections; all hyphae IKI

–, CB +; tissues darkening in KOH. Generative hyphae in context

colourless, thin-walled, 2.5–4 µm diam; skeletal hyphae in context

pale yellowish brown, thick-walled with a narrow lumen or sub-

solid, frequently arboriform and exuous, 3–10 µm diam; binding

hyphae in context colourless, thick-walled, branched and exuous,

2–3 µm diam. Generative hyphae in tubes colourless, thin-walled,

2–3 μm diam; skeletal hyphae in tubes pale brown to dark brown,

thick-walled with a narrow lumen or sub-solid, frequently arboriform

and exuous, 2–8 μm diam; binding hyphae in tubes colourless,

thick-walled, branched and exuous, 1–2.5 μm diam. Pileipellis

composed of clamped generative hyphae, thick-walled to sub-solid,

apical cells clavate, strongly inated and exuous, pale yellowish

brown, about 30–50 × 8–16 µm, forming a regular palisade.

Cystidia and cystidioles absent. Basidia absent; basidioles barrel-

shaped, colourless, thin-walled, 20–25 ×9–12 µm. Basidiospores

ellipsoid, truncated, pale yellowish brown, IKI –, CB +, double-

walled with moderately to distinctly thick walls, exospore walls

smooth, endospore walls with dense spinules, (7–)7.5–9.5(–10) ×

(5–)5.5–7(–7.5) µm, L = 8.52 μm, W = 6.2 μm, Q = 1.37 (n = 30/1,

with the turgid vesicular appendix excluded); (8–)9–11(–11.5) ×

(5–)5.5–7(–8) µm, L = 10.05 μm, W = 6.45 μm, Q = 1.56 (n = 30/1,

with the turgid vesicular appendix included).

Materials examined: China, Sichuan, Qingchuan County, Qingxi Town, on

stump of Quercus, 31 Oct. 2016, Cui 14404 (BJFC029272), Cui 14405

(BJFC029273), Cui 14406 (BJFC029274); Yunnan, Kunming, Xiaoshao

Forest Farm, on rotten angiosperm wood, 1 Jul. 2019, Dai 20017

(BJFC031691).

Notes: Ganoderma lucidum was rstly described from London in

the UK. It is the type species of Ganoderma and has the typical

ganodermoid characters, such as stipitate basidiomata with

laccate pileal surface, and truncated basidiospores with spinules

on the endospore walls (Steyaert 1972, Ryvarden 2004b). Cao

(2013) suggested that G. cantharelloideum should be treated as

a synonym of G. lucidum according to the original description

and observations on the holotype of G. cantharelloideum. In the

current phylogenetic study, the G. lucidum lineage consisted of

specimens collected from the UK, Czech Republic and China with

high divergence (Figs 1, 2). The divergence in G. lucidum groups

also appeared in phylogenetic analyses inferred from single or

multiple genes by Zhou et al. (2015), Hapuarachchi et al. (2018b),

Liu et al. (2019) and Luangharn et al. (2021). This may be caused

by geographic separation. An essential systemic study of the G.

lucidum lineage should be conducted based on more specimens

from different regions and more gene markers.

Ganoderma magniporum J.D. Zhao & X.Q. Zhang, Acta Mycol.

Sin. 3: 15. 1984. MycoBank MB 124473.

For a detailed description of G.a magniporum, see Zhao et al.

(1984).

Notes: Ganoderma magniporum was described from Guangxi of

South China based on its small and nearly black pilei, large pores

(2–2.5 per mm), and ellipsoid basidiospores (8.7–10.4 × 5.2–7 μm,

Zhao et al. 1984). Cao (2013) observed the type specimen (HMAS

42696) of G. magniporum, and stated it was immature and had

smaller basidiospores than another specimen (Zhou 439) collected

from Guangxi. In the current study, one specimen collected from

Yunnan grouped with G. magniporum.

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Ganodermataceae from China

Ganoderma multipileum Ding Hou, Quarterly Journal of the

Taiwan Museum 3: 101. 1950. MycoBank MB 344109.

Synonym: Ganoderma chenghaiense J.D. Zhao, Acta Mycol. Sin.

8: 31. 1989.

For a detailed description of Ganoderma multipileum, see Wang

et al. (2009).

Notes: Ganoderma multipileum was described from Taiwan (China)

based on its imbricate basidiomata with yellowish brown to reddish

brown pileal surface, small pores (6–8 per mm), corky context, and

ellipsoid to ovoid basidiospores (Wang et al. 2009). Cao (2013)

studied the holotype of G. multipileum and G. chenghaiense, and

suggested that the latter should be a synonym of G. multipileum

based on the morphological characters. As more specimens were

collected, the distribution of G. multipileum became widespread,

found in South China, Laos and India. In the phylogenetic analysis,

one specimen collected from the type locality was included and

G. multipileum formed a stable lineage with good support (Fig. 1).

Ganoderma mutabile Y. Cao & H.S. Yuan, Mycol. Prog. 12: 122.

2013. MycoBank MB 563047. Figs 49, 50.

Description: Basidiomata perennial, sessile, hard corky to woody

hard. Pilei solitary, abelliform to shell-like, up to 18 cm diam

and 7 cm thick. Pileal surface reddish brown to purplish brown,

laccate, glabrous, with concentric furrows; margin obtuse, entire,

slightly wavy. Pore surface white when fresh, turning darker when

bruised, dark brown when dry; pores circular to angular, 4–5

per mm; dissepiments moderately thick, entire. Context brown,

homogeneous, with black melanoid lines, hard corky, up to 3 cm

thick. Tubes brown, stratied, up to 4 cm long. Hyphal system

trimitic; generative hyphae with clamp connections; all hyphae IKI

–, CB +; tissues darkening in KOH. Generative hyphae in context

colourless, thin-walled, 2–3.5 µm diam; skeletal hyphae in context

pale brown, thick-walled with a narrow lumen or sub-solid, frequently

arboriform and exuous, 2–5.5 µm diam; binding hyphae in context

colourless, thick-walled, branched and exuous, 1–2.5 µm diam.

Generative hyphae in tubes colourless, thin-walled, 2–3 μm diam;

skeletal hyphae in tubes pale brown, thick-walled with a narrow

lumen or sub-solid, frequently arboriform and exuous, 2–4.5 μm

diam; binding hyphae in tubes colourless, thick-walled, branched

and exuous, 1–2.5 μm diam. Pileipellis composed of clamped

generative hyphae, thick-walled to sub-solid, apical cells mostly

with lateral outgrowths and protuberances, branched and exuous,

yellowish brown, about 40–65 × 4–9 µm, forming an irregular

palisade. Cystidia, cystidioles, basidia and basidioles absent.

Basidiospores broadly ellipsoid, truncated, pale yellowish brown,

IKI –, CB +, double-walled with distinctly thick walls, exospore walls

smooth, endospore walls with dense spinules, (7.5–)8–11(–11.5)

× 5.5–7(–7.5) µm, L = 9.1 μm, W = 6.25 μm, Q = 1.46 (n = 30/1,

with the turgid vesicular appendix excluded); (8.5–)9–12.5(–13) ×

(5.5–)6–7.5(–8) µm, L = 10.57 μm, W = 6.83 μm, Q = 1.55 (n =

30/1, with the turgid vesicular appendix included).

Fig. 47. Basidiomata of Ganoderma lucidum.

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Sun et al.

Materials examined: China, Yunnan, Chuxiong, Zixishan Forest Park, on

living tree of angiosperm, 8 Sep. 2006, holotype Yuan 2289 (IFP); Xinping

County, Mopanshan Forest Park, on ground of mixed forest, 16 Aug. 2019,

Dai 20414 (BJFC032082); Xizang, on angiosperm wood, Dec. 2018, Cui

17189 (BJFC030489).

Notes: Ganoderma mutabile was described from Yunnan in China

based on the irregular apical cells of the pileipellis which are

strongly exuous and frequently branched Cao & Yuan (2012).

Only one specimen was included in the original description, and

in this study additional specimens were collected from Yunnan and

Xizang.

Fig. 48. Microscopic structures of Ganoderma lucidum (drawn from Dai 15805). A. Basidiospores. B. Apical cells from cuticle. C. Basidioles. D. Hyphae from

trama. E. Hyphae from context. Scale bars = 10 μm.

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Ganodermataceae from China

Ganoderma orbiforme (Fr.) Ryvarden, Mycologia 92: 187. 2000.

MycoBank MB 464692.

Basionym: Polyporus orbiformis Fr., Epicr. Syst. Mycol. (Upsaliae):

463. 1838.

Synonyms: Ganoderma fornicatum (Fr.) Pat., Bull. Soc. Mycol. Fr.

5: 71. 1889.

Ganoderma mastoporum (Lév.) Pat., Bull. Soc. Mycol. Fr. 5: 75.

1889.

Ganoderma subtornatum Murrill, Bull. Torrey Bot. Club 34: 477.

1907.

Ganoderma pygmoideum Steyaert, Bull. Jard. Bot. État Brux. 32:

103. 1962.

Ganoderma crebrostriatum J.D. Zhao & L.W. Hsu, Acta Mycol. Sin.

2: 161. 1983.

Ganoderma densizonatum J.D. Zhao & X.Q. Zhang, Acta. Mycol.

Sin. 5: 86. 1986.

Ganoderma limushanense J.D. Zhao & X.Q. Zhang, Acta. Mycol.

Sin. 5: 219. 1986.

Ganoderma diaoluoshanense J.D. Zhao & X.Q. Zhang, Acta Mycol.

Sin. 6: 1. 1987.

For a detailed description of G. orbiforme, see Ryvarden (2000)

and Wang et al. (2014).

Notes: Polyporus orbiformis was combined as Ganoderma orbiforme

by Ryvarden (2000), while G. boninense and G. pygmoideum were

considered as synonyms of G. orbiforme by Ryvarden (2000).

The treatment of G. boninense as a synonym of G. orbiforme is

not accepted by most other mycologists (Pilotti 2005). Wang et

al. (2014) claried the taxonomic status of G. orbiforme based

on morphological and molecular data. Ganoderma cupreum, G.

densizonatum, G. fornicatum, G. limushanense, G. mastoporum

and G. subtornatum were treated as synonyms of G. orbiforme by

Wang et al. (2014). Cao (2013) regarded G. crebrostriatum and

G. diaoluoshanense as synonyms of G. mastoporum based on the

observation of holotype specimens, while G. crebrostriatum and

G. diaoluoshanense should be regarded as the synonyms of G.

orbiforme according to the revision of G. mastoporum by Wang

et al. (2014). In this study, two specimens of G. cupreum from

Cameroon were included in the phylogenetic analyses, and they

formed an independent lineage which is distinct from G. orbiforme

(Fig. 1). Thus, G. cupreum should be treated as an independent

species.

Ganoderma philippii (Bres. et Henn.) Bres., Iconogr. Mycol. 21:

1014. 1932. MycoBank MB 314321. Figs 51, 52.

Basionym: Fomes philippii Bres. & Henn. ex Sacc., Syll. Fung.

(Abellini) 9: 180. 1891.

Description: Basidiomata annual or perennial, sessile and broadly

attached, sometimes growing together, hard corky. Pilei solitary,

variable, abelliform to circular, applanate, up to 26 cm diam and 1.6

cm thick. Pileal surface pale brown to purplish black, dull, glabrous,

with dense concentric zones; margin acute to obtuse, wavy like

petal. Pore surface white when fresh, turning darker when bruised,

pale brown when dry; pores circular to angular, 5–6 per mm;

Fig. 49. Basidiomata of Ganoderma mutabile.

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Sun et al.

dissepiments thin, mostly entire. Context brown, homogeneous,

with black melanoid lines, hard corky, up to 1.4 cm thick. Tubes

yellowish brown, non-stratied, up to 2 mm long. Hyphal system

trimitic; generative hyphae with clamp connections; all hyphae IKI

–, CB +; tissues darkening in KOH. Generative hyphae in context

colourless, thin-walled, 2–3 µm diam; skeletal hyphae in context

pale yellowish brown, thick-walled with a narrow lumen or sub-

solid, arboriform and exuous, 3–5 µm diam; binding hyphae in

Fig. 50. Microscopic structures of Ganoderma mutabile (drawn from Cui 17189). A. Basidiospores. B. Apical cells from cuticle. C. Hyphae from trama. D.

Hyphae from context. Scale bars = 10 μm.

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Ganodermataceae from China

context colourless, thick-walled, branched and exuous, 1–2 µm

diam. Generative hyphae in tubes colourless, thin-walled, 1–3 μm

diam; skeletal hyphae in tubes pale brown to dark brown, thick-

walled with a narrow lumen or sub-solid, frequently arboriform and

exuous, 2–5 μm diam; binding in tubes colourless, thick-walled,

branched and exuous, 1–2 μm diam. Cystidia, cystidioles, basidia

and basidioles absent. Basidiospores obovoid, not obviously

truncated, with obvious vesicular appendix, pale yellowish brown,

IKI –, CB +, double-walled with slightly thick walls, exospore walls

smooth, endospore walls with dense spinules, 5–6 × 3–4 µm, L =

5.63 μm, W = 3.38 μm, Q = 1.67 (n = 30/1, with the turgid vesicular

appendix excluded); 6–8 × 3–4 µm, L = 7.32 μm, W = 3.36 μm, Q =

2.18 (n = 30/1, with the turgid vesicular appendix included).

Materials examined: China, Hainan, Qiongzhong County, Limushan,

on stump of Hevea, 9 Sep. 2016, Cui 14443 (BJFC029311), Cui 14444

(BJFC029311). Singapore, Bukit Timah Nature Reserve, on fallen trunk of

angiosperm tree, 19 Jul. 2017, Dai 17828 (BJFC025360).

Notes: Ganoderma philippii was rstly described from Myanmar, and

it has a wide distribution in South-East Asia including South China,

Indonesia, Malaysia and Singapore (Steyaert 1972, Moncalvo &

Ryvarden 1997). Ganoderma philippii can be distinguished by the

sessile basidiomata with variable pilei, pale brown to purplish black

pileal surface with dense concentric zones, small and obovoid

basidiospores.

Ganoderma sanduense Hapuar. et al., Mycosphere 8: 274. 2019.

MycoBank MB 634622.

For a detailed description of Ganoderma sanduense, see

Hapuarachchi et al. (2019b).

Notes: The type locality of Ganoderma sanduense is Sandu County

of Guizhou Province in southwestern China. Ganoderma stratoideum

was also described from Sandu County, and it shares layered, reddish

black and laccate pilei, moderately sized pores (3–5 per mm), and

heterogenous context with G. sanduense (He & Yu 1989). However,

the small pilei (2–4 × 1–2.5 cm), greyish brown pore surface and

larger basidiospores (12.1–13.8 × 9.2–10.5 μm) differentiate G.

sanduense (Hapuarachchi et al. 2019b). The type specimen of G.

stratoideum has been lost, and its taxonomic status is doubtful.

Ganoderma shanxiense L. Fan & H. Liu, Phytotaxa 406: 132.

2019. MycoBank MB 830632.

For a detailed description of Ganoderma shanxiense, see Liu et

al. (2019).

Notes: Ganoderma shanxiense was described from Shanxi Province

in China and is characterised by its basidiospores with a tapering

and obtuse end at maturity (Liu et al. 2019). In the phylogenetic

tree, G. shanxiense clustered with G. chuxiongense (Fig. 1) which

Fig. 51. Basidiomata of Ganoderma philippii.

372

Sun et al.

Fig. 52. Microscopic structures of Ganoderma philippii (drawn from Cui 14443). A. Basidiospores. B. Hyphae from trama. C. Hyphae from context. Scale

bars = 10 μm.

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Ganodermataceae from China

was described from Yunnan Province, but the latter species differs

by a purplish black and laccate pileal surface, greyish white pore

surface with small pores (7–8 per mm), and broadly ellipsoid to

ovoid basidiospores (10–12 × 7–8.5 μm).

Ganoderma sichuanense J.D. Zhao & X.Q. Zhang, Acta. Mycol.

Sin. 2: 159. 1983. MycoBank MB 107984.

For a detailed description of Ganoderma sichuanense, see Zhao et

al. (1983) and Wang et al. (2012).

Notes: Ganoderma sichuanense was rst described from Sichuan

Province by Zhao et al. (1983). Since then, more specimens have

been collected from Guangdong, Guangxi of China and Sri Lanka.

Phylogenetically, G. weberianum grouped with G. sichuanense

which is consistent with Hapuarachchi et al. (2019b). Further studies

are needed to clarify the relationship between G. weberianum and

G. sichuanense.

Ganoderma sinense J.D. Zhao et al., Acta Microbiol. Sin. 19: 272.

1979. MycoBank MB 314325. Figs 53, 54.

Synonyms: Ganoderma austrofujianense J.D. Zhao et al., Acta

Microbiol. Sin. 19: 274. 1979.

Ganoderma luteomarginatum J.D. Zhao et al., Acta Microbiol. Sin.

19: 274. 1979.

Ganoderma formosanum T.T. Chang & T. Chen, Trans. Br. Mycol.

Soc. 82: 731. 1984.

Ganoderma guinanense J.D. Zhao & X.Q. Zhang, Acta Mycol. Sin.

6: 4. 1987.

Ganoderma mediosinense J.D. Zhao, Acta Mycol. Sin. 7: 205.

1988.

Description: Basidiomata annual, laterally stipitate, hard corky

to woody hard. Pilei solitary, variable, abelliform to reniform or

circular, up to 14 cm diam and 1.6 cm thick. Pileal surface reddish

brown to purplish black, strongly laccate, glabrous, with concentric

furrows and radial wrinkles; margin acute to obtuse, entire, slightly

incurved. Pore surface white when fresh, turning darker when

bruised, light buff when dry; pores circular to angular, 3–5 per mm;

dissepiments thin, mostly entire. Context heterogeneous, the upper

layer white to buff, the lower layer pale brown to yellowish brown,

with black melanoid lines, hard corky, up to 3 cm thick. Tubes

yellowish brown, non-stratied, up to 1.5 cm long. Stipe reddish

brown to purplish black, attened to cylindrical, up to 6 cm long

and 2 cm diam. Hyphal system trimitic; generative hyphae with

clamp connections; all hyphae IKI –, CB +; tissues darkening in

KOH. Generative hyphae in context colourless, thin-walled, 2–4

μm diam; skeletal hyphae in context pale yellowish brown, thick-

walled with a narrow lumen or sub-solid, arboriform and exuous,

3–6 µm diam; binding hyphae in context colourless, thick-walled,

branched and exuous, 1.5–2.5 μm diam. Generative hyphae in

tubes colourless, thin-walled, 1–3 μm diam; skeletal hyphae in

tubes pale brown to dark brown, thick-walled with a narrow lumen

or sub-solid, frequently arboriform and exuous, 2–4 μm diam;

binding hyphae in tubes colourless, thick-walled, branched and

exuous, 1–2 μm diam. Pileipellis composed of clamped generative

hyphae, thick-walled to sub-solid, apical cells clavate, inated, pale

yellow to yellowish brown, about 40–55 × 6–12 μm, forming a

Fig. 53. Basidiomata of Ganoderma sinense.

374

Sun et al.

regular palisade. Cystidia and cystidioles absent. Basidia absent;

basidioles barrel-shaped to clavate, colourless, thin-walled, 16–24

× 9–12 μm. Basidiospores ellipsoid, not obviously truncated, pale

yellowish brown, IKI –, CB +, double-walled with moderately thick

walls, exospore wall smooth, endospore wall with dense spinules,

(11–)11.6–13.2(–13.7) × (7–)7.3–8.5(–8.8) μm, L = 12.39 μm, W

= 7.99 μm, Q = 1.74 (n = 30/1, with the turgid vesicular appendix

included).

Materials examined: China, Hainan, Lingshui County, Diaoluoshan

Forest Park, on ground of angiosperm forest, 15 Jun. 2016, Cui 13825

(BJFC028691), Cui 13835 (BJFC028701); Guangxi, Jinxiu County,

Fig. 54. Microscopic structures of Ganoderma sinense (drawn from Cui 14574). A. Basidiospores. B. Apical cells from cuticle. C. Basidioles. D. Hyphae from

trama. E. Hyphae from context. Scale bars = 10 μm.

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Ganodermataceae from China

Dayaoshan Nature Reserve, on stump of angiosperm tree, 15 Jul. 2017,

Cui 14524 (BJFC029393), Cui 14526 (BJFC029395).

Notes: Ganoderma sinense was described from Hainan Province

by Zhao et al. (1979), and it can be easily distinguished in the

wild by its laterally stipitate basidiomata with dark reddish brown

to purplish black and strongly laccate pileal surface. In this

study, G. sinense clustered with G. japonicum with high support

(Fig. 1) which is consistent with previous study by Hapuarachchi

et al. (2019a). Ganoderma japonicum should be treated as

a synonym of G. sinense temporarily based on the previous

description and comments (Moncalvo & Ryvarden 1997, Liao

et al. 2015, Hapuarachchi et al. 2019a). Wang (2005) and Cao

(2013) also mentioned that G. austrofujianense, G. formosanum,

G. guinanense, G. luteomarginatum and G. mediosinense are

synonyms of G. sinense after studying the type specimens.

Ganoderma tropicum (Jungh.) Bres., Annls Mycol. 8: 586. 1910.

MycoBank MB 149294. Figs 55, 56.

Basionym: Polyporus tropicus Jungh., Verh. Batav. Genootsch.

Kunst. Wet. 17: 63. 1838.

Description: Basidiomata annual to perennial, usually sessile,

sometimes laterally stipitate, hard corky to woody hard. Pilei

solitary, variable, abelliform to shell-shaped or circular, up to 17 cm

diam and 3 cm thick. Pileal surface reddish brown to dark brown,

strongly laccate, glabrous, with obvious concentric zones; margin

acute to obtuse, entire or sometimes lacerated. Pore surface white

when fresh, turning darker when bruised, straw yellow when dry;

pores circular to angular, 4–6 per mm; dissepiments slightly thick to

moderately thick, mostly entire. Context dark brown, homogeneous,

with black melanoid lines, corky, up to 2.2 cm thick. Tubes brown,

non-stratied, up to 8 mm long. Stipe reddish brown to purplish

black, attened to cylindrical, up to 6 cm long and 1.2 cm diam.

Hyphal system trimitic; generative hyphae with clamp connections;

all hyphae IKI –, CB +; tissues darkening in KOH. Generative

hyphae in context colourless, thin-walled, 2.5–3.5 µm diam; skeletal

hyphae in context pale yellowish brown, thick-walled with a wide

to narrow lumen or sub-solid, frequently arboriform and exuous,

2.5–6 µm diam; binding hyphae in context colourless, thick-walled,

branched and exuous, 1–2 µm diam. Generative hyphae in tubes

colourless, thin-walled, 2–3 μm diam; skeletal hyphae in tubes pale

brown to dark brown, thick-walled with a narrow lumen or sub-solid,

frequently arboriform and exuous, 2–4 μm diam; binding hyphae

in tubes colourless, thick-walled, branched and exuous, 0.5–1 μm

diam. Pileipellis composed of clamped generative hyphae, thick-

walled to sub-solid, apical cells clavate, sometimes branched or

protuberant, inated and exuous, yellowish brown, about 19–32

× 4–9 μm, forming a regular palisade. Cystidia, cystidioles, basidia

and basidioles absent. Basidiospores ellipsoid, truncated, pale

yellowish brown, IKI –, CB +, double-walled with slightly thick

walls, exospore walls smooth, endospore walls with dense

spinules, (6.5–)7–8.5(–9) × (4.5–)5–6(–6.5) µm, L = 7.73 μm, W

= 5.66 μm, Q = 1.37 (n = 30/1, with the turgid vesicular appendix

excluded); (6.8–)7–9.6(–10) × (4–)4.5–6.2(–6.4) µm, L = 8.90

μm, W = 5.56 μm, Q = 1.60 (n = 30/1, with the turgid vesicular

appendix included).

Materials examined: China, Hainan, Haikou, Jinniuling Park, on stump of

Acacia, 7 Jun. 2016, Dai 16434 (BJFC022551); Guangdong, Maoming,

Dianbai, on dead tree of Casuarina, 3 Jun. 2019, Dai 19679 (BJFC031355);

Guangxi, Baise, Baise Uprising Memorial Park, on living tree of Acacia, 1

Jul. 2019, Cui 20029 (BJFC031703). Sri Lanka, Colombo, Dombagaskarda

Forest Reserve, on living tree of angiosperm, 27 Feb. 2019, Dai 19491

(BJFC031171). Vietnam, Ho Chi Minh City, Botanical Garden, on living

tree of Diospyros bejandii, 11 Oct. 2017, Cui 16369 (BJFC029668).

Fig. 55. Basidiomata of Ganoderma tropicum.

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Sun et al.

Notes: Ganoderma tropicum was described from Indonesia, and

it is characterised by the sessile to woody hard basidiomata, a

strongly laccate pileal surface with obvious concentric zones, dark

brown context, sometimes branched or protuberant apical cells of

pileipellis, and ellipsoid basidiospores with dense spinules on the

endospore walls (Steyaert 1972, Ryvarden 1981). These features

are consistent with the observation in this study, and all specimens

used in the phylogenetic analysis clustered together with high

support (Fig. 1).

Fig. 56. Microscopic structures of Ganoderma tropicum (drawn from Cui 16341). A. Basidiospores. B. Apical cells from cuticle. C. Hyphae from trama. D.

Hyphae from context. Scale bars = 10 μm.

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Ganodermataceae from China

Ganoderma tsugae Murrill, Bull. Torrey Bot. Club 29: 601. 1902.

MycoBank MB 239416. Figs 57, 58.

Description: Basidiomata annual, laterally stipitate, hard corky

to woody hard. Pilei solitary, abelliform, up to 20 cm diam and

4 cm thick. Pileal surface pale yellowish brown to reddish brown,

strongly laccate, glabrous, with distinct concentric furrows and

slightly radial wrinkles; margin acute to obtuse, lacerated like

petals, slightly incurved when dry. Pore surface white when fresh,

turning darker when bruised, pale yellowish brown when dry; pores

circular to angular, 4–6 per mm; dissepiments moderately thick,

mostly entire. Context heterogeneous, the upper layer white, the

lower layer pale brown to yellowish brown, without black melanoid

lines, hard corky, up to 2.2 cm thick. Tubes yellowish brown, non-

stratied, up to 1.5 cm long. Stipe reddish brown to purplish black,

attened to cylindrical, up to 6 cm long and 2 cm diam. Hyphal

system trimitic; generative hyphae with clamp connections; all

hyphae IK I–, CB +; tissues darkening in KOH. Generative hyphae

in context colourless, thin-walled, 2–3 µm diam; skeletal hyphae

in context pale yellowish brown, thick-walled with a narrow lumen

or sub-solid, arboriform and exuous, 2.5–6.5 µm diam; binding

hyphae in context colourless, thick-walled, branched and exuous,

0.5–2 µm diam. Generative hyphae in tubes colourless, thin-

walled, 1.5–3 μm diam; skeletal hyphae in tubes pale brown to dark

brown, thick-walled with a narrow lumen or sub-solid, frequently

arboriform and exuous, 1.5–4 μm diam; binding hyphae in tubes

colourless, thick-walled, branched and exuous, 0.5–2 μm diam.

Pileipellis composed of clamped generative hyphae, thick-walled,

apical cells clavate, inated, yellowish brown, about 30–45 × 5–10

μm, forming a regular palisade. Cystidia and cystidioles absent.

Basidia barrel-shaped, colourless, thin-walled, 22–28 × 12–18 µm;

basidioles broadly clavate, colourless, thin-walled, 18–23 × 11–14

µm. Basidiospores ellipsoid, truncated, pale yellowish brown, IKI

–, CB +, double-walled with moderately thick walls, exospore walls

smooth, endospore walls with dense spinules, 8–10(–11) × 5–6.2

µm, L = 9.12 μm, W = 5.49 μm, Q = 1.66 (n = 30/1, with the turgid

vesicular appendix excluded); 9–11(–12) × 5–6(–7) µm, L = 10.21

μm, W = 5.88 μm, Q = 1.74 (n = 30/1, with the turgid vesicular

appendix included).

Materials examined: China, Jilin, Fusong County, Songjianghe Forest

Park, on stump of Larix, 2 Aug. 2016, Cui 14110 (BJFC028978), Cui

14112 (BJFC028980); Xinjiang, Habahe County, Baihabahe Forest Park,

on stump of Larix, 10 Sep. 2015, Dai 15851 (BJFC019952), Dai 15856

(BJFC019957). USA, Connecticut, on living tree of Tusga, 19 Jul. 2012,

Dai 12751b (BJFC013059).

Notes: Ganoderma tsugae was described from the USA growing

on Tsuga canadensis by Murrill (1902); it has been reported

occurring on several genera of coniferous trees such as Abies,

Larix, Picea and Tsuga (Steyaert 1980, Adaskaveg & Gilbertson

1986). Ganoderma tsugae grouped in the G. lucidum complex in

the phylogenetic analyses (Fig. 1).

Ganoderma weberianum (Bres. & Henn. ex Sacc.) Steyaert,

Persoonia 7: 79. 1972. MycoBank MB 314330. Figs 59, 60.

Fig. 57. Basidiomata of Ganoderma tsugae.

378

Sun et al.

Basionym: Fomes weberianus Bres. & Henn. ex Sacc., Syll. Fung.

(Abellini) 9: 174. 1891.

Synonyms: Ganoderma microsporum R.S. Hseu, Mycotaxon 35:

36. 1989.

Ganoderma tenue J.D. Zhao et al., Acta Microbiol. Sin. 19: 271.

1979.

Description: Basidiomata annual, sessile or with short stipe, usually

growing together, imbricate, hard corky to woody hard. Pilei solitary,

abelliform, applanate, up to 14 cm diam and 8 mm thick. Pileal

surface reddish brown to purplish black or near black, strongly

laccate, glabrous, with concentrical bands and slightly radial

wrinkles; margin acute, lacerated like petal, slightly incurved when

Fig. 58. Microscopic structures of Ganoderma tsugae (drawn from Cui 14554). A. Basidiospores. B. Apical cells from cuticle. C. Basidia and basidioles. D.

Hyphae from trama. E. Hyphae from context. Scale bars = 10 μm.

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Ganodermataceae from China

dry. Pore surface white when fresh, turning darker when bruised,

pale yellow when dry; pores circular to angular, 4–6 per mm;

dissepiments thin to slightly thick, mostly entire. Context greyish

brown, homogeneous, without black melanoid lines, hard corky,

up to 2 mm thick. Tubes pale brown to pale grey, non-stratied,

up to 6 mm long. Hyphal system trimitic; generative hyphae with

clamp connections; all hyphae IKI –, CB +; tissues darkening in

KOH. Generative hyphae in context colourless, thin-walled, 2–4 μm

diam; skeletal hyphae in context pale yellowish brown, thick-walled

with a wide to narrow lumen or sub-solid, frequently arboriform and

exuous, 2.5–7 μm diam; binding hyphae in context colourless,

thick-walled, branched and exuous, 2–3 μm diam. Generative

hyphae in tubes colourless, thin-walled, slightly swollen at the distal

end, 2–3.5 μm diam; skeletal hyphae in tubes pale brown to dark

brown, thick-walled with a narrow lumen or sub-solid, frequently

arboriform and exuous, 2–4.5 μm diam; binding hyphae in tubes

colourless, thick-walled, branched and exuous, 1.5–3 μm diam.

Pileipellis composed of clamped generative hyphae, thick-walled to

sub-solid, apical cells clavate, inated, golden yellow, about 60–90

× 6–12 μm, forming a regular palisade. Cystidia and cystidioles

absent. Basidia barrel-shaped, colourless, thin-walled, 15–35 ×

13–18 μm; basidioles in shape like the basidia, colourless, thin-

walled, 14–25 × 9–15 μm. Basidiospores subglobose to broadly

ellipsoid, not obviously truncated, pale yellowish brown, IKI –, CB

+, double-walled with slightly thick walls, exospore wall smooth,

endospore wall with dense spinules, 6–7 × 4–6 μm, L = 6.53 μm,

W = 5.13 μm, Q = 1.27 (n = 30/1, with the turgid vesicular appendix

excluded).

Materials examined: China, Guangdong, Yangjiang, Jinshan Botanical

Garden, on rotten stump of angiosperm tree, 3 Jun. 2019, Dai 19673

(BJFC031349); Maoming, Dianbai Region, on living tree of Casuarina,

3 Jun. 2019, Dai 19682 (BJFC031358); Zhanjiang, Guangdong Ocean

University (Huguang Campus), on living tree of Cinnamomum, 4 Jun.

2019, Dai 19689 (BJFC031365). Vietnam, Ho Chi Minh City, United

Palace Park, on living tree of Cynometra dongnaiensis, 10 Oct. 2017, Cui

16359 (BJFC029658), Cui 16360 (BJFC029659).

Notes: Ganoderma weberianum was rst described from

the Samoa Islands, and it probably has a worldwide tropical

distribution (Steyaert 1972). Moncalvo et al. (1995) and Smith

& Sivasithamparam (2003) mentioned that the ITS sequences

of G. weberianum is almost the same to G. microsporum and

suggested that G. microsporum should be treated as a synonym

of G. weberianum. Cao (2013) regarded Ganoderma tenue as a

synonym of G. weberianum based on the similar morphological

characters such as reddish brown to purplish black pileal surface,

apical cells of pileipellis regularly palisaded, and subglobose to

broadly ellipsoid basidiospores.

Ganoderma weixiense Karun. & J.C. Xu, Phytotaxa 423: 78.

2019. MycoBank MB 646645.

Fig. 59. Basidiomata of Ganoderma weberianum.

380

Sun et al.

For a detailed description of Ganoderma weixiense, see Ye et al.

(2019).

Notes: Ganoderma weixiense was described from high altitude areas

(altitude > 2 000 m) of Weixi County and Jinning in Yunnan Province.

Ganoderma leucocontextum has similar morphological characters

and close phylogenetic relationship with G. weixiense except the size

of pores and basidiospores (details in notes of G. leucocontextum). In

addition, G. leucocontextum shares similar climate and altitude habits

with G. weixiense. In the current study, these two species grouped

together, and it is quite difcult to separate them in morphology,

ecology and phylogeny. The taxonomic status of G. weixiense needs

to be claried in further studies.

Fig. 60. Microscopic structures of Ganoderma weberianum (drawn from Cui 16360). A. Basidiospores. B. Apical cells from cuticle. C. Basidia and basidioles.

D. Hyphae from trama. E. Hyphae from context. Scale bars = 10 μm.

381www.studiesinmycology.org

Ganodermataceae from China

Ganoderma williamsianum Murrill, Bull. Torrey Bot. Club 34: 478.

1907. MycoBank MB 141987.

Synonym: Ganoderma meijiangense J.D. Zhao, Acta Mycol. Sin.

7: 16. 1988.

For a detailed description of Ganoderma williamsianum, see Wang

&Wu (2010).

Notes: Ganoderma williamsianum was described from the

Philippines and is easily confused with G. acontextum, G. australe

and other non-laccate Ganoderma species in the wild as they all

shared a hard and dull pileal surface. Ganoderma williamsianum

is widely distributed in tropical areas of the Philippines, China,

Malaysia, Singapore, Sri Lanka, Thailand and Vietnam. Wang &

Wu (2010) studied specimens of G. williamsianum collected from

Hainan and Yunnan provinces in China and the type specimen of

G. meijiangense, and the latter was treated as a synonym of G.

williamsianum.

Key to accepted species of Ganoderma in China

1a. Endospore wall smooth .......................................................................................................................................................................... 2

1b. Endospore wall ornamented .................................................................................................................................................................. 3

2a. Basidiomata annual, pileal surface chestnut-colour, tubes non-stratied ......................................................................... G. castaneum

2b. Basidiomata perennial, pileal surface greyish brown, tubes stratied .............................................................................. G. yunlingense

3a. Pileal surface dull ................................................................................................................................................................................... 4

3b. Pileal surface laccate ........................................................................................................................................................................... 13

4a. Pilei imbricate, margin lacerated like petals ......................................................................................................................... G. puerense

4b. Pilei solitary, margin entire ..................................................................................................................................................................... 5

5a. Basidiospores subglobose ........................................................................................................................................... G. hoehnelianum

5b. Basidiospores broadly ellipsoid to ellipsoid or ovoid .............................................................................................................................. 6

6a. Tubes stratied ....................................................................................................................................................................................... 7

6b. Tubes non-stratied ................................................................................................................................................................................ 8

7a. Context homogeneous; basidiospores 5.5–7 × 4.1–5.2 μm ............................................................................................. G. applanatum

7b. Context heterogeneous; basidiospores 7–12 × 5–8 μm ........................................................................................................ G. australe

8a. Pores < 4 per mm ........................................................................................................................................................ G. tongshanense

8b. Pores > 4 per mm .................................................................................................................................................................................. 9

9a. Context without black melanoid lines; apical cells in cuticle branched ........................................................................... G. ellipsoideum

9b. Context with black melanoid lines; apical cells in cuticle unbranched ................................................................................................. 10

10a. Distributed in higher altitudes ............................................................................................................................................... G. alpinum

10b. Distributed in lower altitudes .............................................................................................................................................................. 11

11a. Apical cells in cuticle irregularly branched or with protuberances ............................................................................... G. williamsianum

11b. Apical cells in cuticle unbranched or without protuberances .............................................................................................................. 12

12a. Pileal surface reddish brown to greyish brown, pores angular .......................................................................................... G. gibbosum

12b. Pileal surface greyish brown to nearly black, pores circular ......................................................................................... G. guangxiense

13a. Basidiomata sessile ........................................................................................................................................................................... 14

13b. Basidiomata stipitate or with constricted short stipe .......................................................................................................................... 17

14a. Basidiospores almond-shaped .................................................................................................................................. G. angustisporum

14b. Basidiospores ellipsoid to ovoid ......................................................................................................................................................... 15

15a. Apical cells in cuticle irregularly branched or with protuberances ....................................................................................... G. mutabile

15b. Apical cells in cuticle unbranched or without protuberances .............................................................................................................. 16

16a. Basidiomata small, pileal surface yellowish brown to reddish brown; basidiospores > 4 μm in width ............. G. bubalinomarginatum

16b. Basidiomata large, pileal surface dark brown to near black; basidiospores < 4 μm in width ................................................ G. philippii

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Sun et al.

17a. Pores < 3 per mm ......................................................................................................................................................... G. magniporum

17b. Pores > 3 per mm .............................................................................................................................................................................. 18

18a. Pileal surface nearly black ................................................................................................................................................................. 19

18b. Pileal surface pale brown to yellowish brown or reddish brown ......................................................................................................... 23

19a. Stipe short or constricted at base, < 4 cm in length ........................................................................................................................... 20

19b. Stipe obviously long, > 4 cm in length ................................................................................................................................................ 21

20a. Basidiospores subglobose to broadly ellipsoid, < 6 μm in width ................................................................................... G. weberianum

20b. Basidiospores broadly ellipsoid to ellipsoid or ovoid, > 6 μm in width ................................................................................ G. orbiforme

21a. Basidiospores truncated .................................................................................................................................................. G. sanduense

21b. Basidiospores not obviously truncated ............................................................................................................................................... 22

22a. Pore surface grey to pale brown, pores 5–6 per mm; basidiospores 8–11 × 5.5–7 μm ....................................................... G. ahmadii

22b. Pore surface white to buff, pores 3–5 per mm; basidiospores 11.6–13.2 × 7.3–8.5 μm ....................................................... G. sinense

23a. Pore surface yellowish when fresh ..................................................................................................................................................... 24

23b. Pore surface white to cream or greyish white when fresh .................................................................................................................. 25

24a. Pilei lobate, with dark red pileal surface ...................................................................................................................... G. chuxiongense

24b. Pilei semicircle, shell-like, reniform to circular, with yellowish brown to reddish brown pileal surface .................................... G. lingzhi

25a. Distributed in temperate or subtropical areas .................................................................................................................................... 26

25b. Distributed in subtropical or tropical areas ......................................................................................................................................... 31

26a. Context white to cream ............................................................................................................................................ G. leucocontextum

26b. Context buff to brown ......................................................................................................................................................................... 27

27a. Growing on coniferous trees .................................................................................................................................................. G. tsugae

27b. Growing on broad-leaf trees ............................................................................................................................................................... 28

28a. Basidiospores < 4 μm in width .......................................................................................................................................... G. weixiense

28b. Basidiospores > 4 μm in width ........................................................................................................................................................... 29

29a. Context with black melanoid lines ................................................................................................................................. G. sichuanense

29b. Context without black melanoid lines ................................................................................................................................................. 30

30a. Context heterogeneous; basidiospores 9–11 × 5.5–7 μm ..................................................................................................... G. lucidum

30b. Context homogeneous; basidiospores 11–13 × 8–9.5 μm .............................................................................................. G. shanxiense

31a. Stipe short or constricted at base, < 6 cm in length ........................................................................................................................... 32

31b. Stipe obviously long, > 6 cm in length ................................................................................................................................................ 34

32a. Growth on palm trees ....................................................................................................................................................... G. boninense

32b. Growth on other trees ........................................................................................................................................................................ 33

33a. Basidiomata small; apical cells in cuticle unbranched, basidiospores 10.7–12.8 × 5.7–9 μm ............................. G. subangustisporum

33b. Basidiomata large; apical cells in cuticle branched or with protuberances, basidiospores 7–9.6 × 4.5–6.2 μm ................. G. tropicum

34a. Basidiomata laterally stipitate ............................................................................................................................................................. 35

34b. Basidiomata dorso-laterally stipitate .................................................................................................................................................. 36

35a. Pilei solitary .................................................................................................................................................................. G. casuarinicola

35b. Pilei imbricate .................................................................................................................................................................. G. multipileum

36a. Context homogeneous, without black melanoid lines ..................................................................................................... G. subexipes

36b. Context heterogeneous, with black melanoid lines ............................................................................................................................ 37

37a. Basidiospores larger, 8.5–12.6 × 7.2–9.1 μm ............................................................................................................... G. calidophilum

37b. Basidiospores smaller, 7–9 × 4.8–5.5 μm ............................................................................................................................. G. exipes

383www.studiesinmycology.org

Ganodermataceae from China

Haddowia Steyaert, Persoonia 7: 108. 1972. MycoBank MB 17717.

Type species: Haddowia longipes (Lév.) Steyaert

Description: Basidiomata annual, laterally stipitate, corky. Pilei

solitary, sub-orbicular to abelliform. Pileal surface yellowish

brown to reddish brown or blackish brown, laccate, tomentose,

concentrically zonate and furrowed, radial rugose. Pore surface

white when fresh becoming straw yellow when dry; pores circular

to angular; dissepiments thick, entire. Context white, corky. Hyphal

system trimitic; generative hyphae colourless, thin-walled, with

clamp connections; skeletal hyphae colourless to pale yellow, thick-

walled, arboriform and exuous; binding hyphae colourless, sub-

solid, branched and exuous. Basidiospores globose to ellipsoid,

non-truncated, yellow to pale yellowish brown, double-walled

with thick walls, exospore wall smooth and covering longitudinal

crests, endospore wall with two longitudinal crests and transverse

membranes.

Notes: Haddowia has similar basidiomata in shape and colour to

Ganoderma, but differs by its non-truncated basidiospores which

are double and thick-walled, a smooth exospore wall covering

longitudinal crests, and an endospore wall with two longitudinal

crests and transverse membranes. Although no outer wall on the

basidiospores of Haddowia was observed by Steyaert (1972), and

a smooth exospore wall exists according to the scanning electron

micrographs of Haddowia species taken by Costa-Rezende et

al. (2020b) and the current study (Fig. 8 E). In the phylogenetic

analyses, Ha. longipes and Ha. macropora formed a distinct well-

supported clade of Ganodermataceae (Fig. 1).

Haddowia macropora B.K. Cui, Vlasák & Y.F. Sun, sp. nov.

MycoBank MB 839663. Figs 61, 62.

Diagnosis: Differs from other species in the genus by its yellowish

brown to reddish brown pileal surface and large pores.

Etymology: macropora (Lat.), refers to the large pores.

Typus: French Guiana (holotype JV 1908/46).

Description: Basidiomata annual, laterally stipitate, corky. Pilei

solitary, sub-orbicular to abelliform, up to 4 cm diam and 1 mm

thick. Pileal surface yellowish brown to reddish brown, strongly

laccate, glabrous, with obvious concentric furrows and irregularly

radial wrinkles; margin obtuse, entire, wavy when dry. Pore surface

cream when fresh becoming dark when bruised; pores angular to

irregular, 1–2 per mm; dissepiments slightly thick, entire. Context

cream, without dark melanoid lines, soft corky, up to 1.5 mm thick.

Tubes concolorous with context, up to 8.5 mm long. Stipe reddish-

brown to purple-black, cylindrical and solid, up to 16.5 cm long

and 6 mm diam. Hyphal system trimitic; generative hyphae with

clamp connections, all the hyphae IKI + (dextrinoid), CB +; tissues

darkening in KOH. Generative hyphae in context colourless, thin-

walled, 2–3 μm diam; skeletal hyphae in context colourless to

pale yellow, thick-walled with a wide to narrow lumen or sub-solid,

frequently arboriform and exuous, 2–5 μm diam; binding hyphae

in context colourless, thick-walled, rarely branched and exuous,

1–1.5 μm diam. Generative hyphae in tubes colourless, thin-walled,

2–3 μm diam; skeletal hyphae in tubes pale yellow, thick-walled

with a wide to narrow lumen or sub-solid, frequently arboriform and

exuous, 2–4 μm diam; binding hyphae in tubes colourless, thick-

Fig. 61. Basidiomata of Haddowia macropora.

384

Sun et al.

walled, rarely branched and exuous, up to 1.5 μm diam. Pileipellis

composed of clamped generative hyphae, thick-walled to sub-

solid, apical cells clavate, inated and exuous, yellowish brown,

about 32–48 × 6–12 μm, forming a regular palisade. Cystidia and

cystidioles absent. Basidia barrel-shaped, colourless, thin-walled,

23–32 × 14–17 μm; basidioles clavate, colourless, thin-walled,

21–30 × 7–15 μm. Basidiospores subglobose to broadly ellipsoid,

pale yellowish brown, IKI + (dextrinoid), CB +, double-walled

with distinctly thick walls, exospore wall smooth and covering

longitudinal crests, endospore wall with intermittently longitudinal

crests and transverse membranes, (13.6–)14–15.5(–16) × (12–)

12.3–13.8(–14) μm, L = 14.64 μm, W = 13.08 μm, Q = 1.12 (n =

60/1).

Notes: Haddowia macropora has the typical basidiospores of

Haddowia which differ from other genera in Ganodermataceae

having intermittently longitudinal crests and transverse membranes

on the endospore walls (Steyaert 1972). It can further be

Fig. 62. Microscopic structures of Haddowia macropora (drawn from JV 1908/46). A. Basidiospores. B. Apical cells from cuticle. C. Basidia and basidioles.

D. Hyphae from trama. E. Hyphae from context. Scale bars = 10 μm.

385www.studiesinmycology.org

Ganodermataceae from China

distinguished by the yellowish brown to reddish brown pileal

surface, and large pores. Haddowia longipes as the type species in

this genus is quite different to Ha. macropora by the blackish brown

pileal surface, thick context (4–5 mm), and ellipsoid basidiospores

(12–19 × 10–14.5 μm, Steyaert 1972).

Key to accepted species of Haddowia

1a. Pileal surface yellowish brown to reddish brown, context thin (< 1.5 mm) ....................................................................... Ha. macropora

1b. Pileal surface blackish brown, context thick (4–5 mm) ........................................................................................................ Ha. longipes

Humphreya Steyaert, Persoonia 7: 98. 1972. MycoBank MB

17778.

Type species: Humphreya lloydii (Pat. & Har.) Steyaert.

For a detailed description of Humphreya, see Steyaert (1972).

Notes: Humphreya basidiospores have a typical ornamentation

of reticulate or disjointed crests on the endospore walls which are

different from other genera of Ganodermataceae (Steyaert 1972).

Until now, Humphreya contained three species, viz., Hu. eminii,

Hu. endertii and Hu. lloydii. No sequence data are available for

Humphreya; thus, it is not included in the phylogenetic analyses of

Ganodermataceae.

Key to accepted species of Humphreya

1a. Basidiospores more than 20 μm in length ................................................................................................................................ Hu. eminii

1b. Basidiospores less than 20 μm in length ................................................................................................................................................ 2

2a. Basidiomata up to 12 cm diam; basidiospores with reticulate ornamentation on the endospore wall ..................................... Hu. lloydii

2b. Basidiomata about 3 cm diam; basidiospores with disjointed crests on the endospore wall ................................................ Hu. endertii

Magoderna Steyaert, Persoonia 7: 111. 1972. MycoBank MB

18011.

Type species: Magoderna subresinosum (Murrill) Steyaert

Description: Basidiomata annual, sessile or stipitate, woody hard.

Pilei solitary, sub-orbicular to infundibuliform. Pileal surface hair

brown to coal black, dull to slightly shiny, glabrous, with concentric

furrows and radial wrinkles. Pore surface buffy brown to pale grey

brown when dry; pores circular to angular; dissepiments thick,

entire. Context cream to buff, sometimes with dark melanoid

lines, woody hard. Hyphal system trimitic; generative hyphae

colourless, thin-walled, with clamp connections; skeletal hyphae

near colourless, sub-solid, arboriform and exuous; binding hyphae

colourless, thick-walled, branched and exuous. Basidiospores

ellipsoid to ovoid, non- truncated, pale yellow, double and thick

walled, exospore wall faintly verrucose, endospore wall with dense

spinules.

Notes: Magoderna has a brous and pale white context, anticlinal

hyphae in pileipellis, and ellipsoid to ovoid basidiospores

without a truncated apex. Until now, M. infundibuliforme and M.

subresinosum were accepted in Magoderna. In the phylogenetic

analyses, the Magoderna clade was well supported and grouped

with the Neoganoderma clade (Fig. 1).

Magoderna subresinosum (Murrill) Steyaert, Persoonia 7: 112.

1972. MycoBank MB 317117. Figs 63, 64.

Basionym: Fomes subresinosus Murrill, Bull. Torrey Bot. Club 35:

410. 1908.

Description: Basidiomata annual, sessile, woody hard. Pilei solitary,

abelliform, up to 12 cm diam and 3 cm thick. Pileal surface coal

black, slightly shiny, glabrous, sticky, with obvious concentric

furrows and strong radial wrinkles; margin subacute to obtuse,

entire, incurved when dry. Pore surface pale greyish brown when

dry; pores circular to angular, 4–5 per mm; dissepiments distinctly

thick, entire. Context cream to pale wood brown, without dark

melanoid lines, woody hard and brous, up to 1.3 cm thick. Tubes

pale yellowish brown, up to 1.6 cm long. Hyphal system trimitic;

generative hyphae with clamp connections, all hyphae IKI –, CB

+; tissues slightly darkening in KOH. Generative hyphae in context

colourless, thin-walled, 2–4 μm diam; skeletal hyphae in context

colourless, sub-solid, arboriform and exuous, 2–6 μm diam;

binding hyphae in context colourless, thick-walled, branched and

exuous, up to 1 μm diam. Generative hyphae in tubes colourless,

thin-walled, 2–3 μm diam; skeletal hyphae in tubes colourless,

sub-solid, arboriform and exuous, 2–4 μm diam; binding hyphae

in tubes colourless, thick-walled, branched and exuous, up to

1 μm diam. Pileipellis composed of clamped generative hyphae,

thick-walled, apical cells clavate, inated, dark brown, about 20–

35 × 5–9 μm, anticlinal, forming a regular palisade. Cystidia and

cystidioles absent. Basidia barrel-shaped, colourless, thin-walled,

20–25 × 18–20 μm; basidioles in shape like the basidia, colourless,

thin-walled, 18–20 × 11–15 μm. Basidiospores ellipsoid to ovoid,

non-truncated, pale yellow, IKI –, CB +, double-walled with slightly

thick walls, exospore wall faintly verrucose, endospore wall with

dense spinules, (14.7–)14.9–17.3(–18) × (9.6–)9.8–11.3(–11.8)

μm, L = 15.88 μm, W = 10.52 μm, Q = 1.51 (n = 60/2).

Materials examined: China, Guangdong, Maoming, on living tree of

Casuarina, Jun. 2017, Cui 14579 (BJFC029448), Cui 14580 (BJFC029449),

Cui 14581 (BJFC029450). Malaysia, Selangor, Kota Damansara, National

Forest Reserve, on dead angiosperm tree, 17 Apr. 2018, Dai 18626

(BJFC026914); on stump of angiosperm tree, 6 Dec. 2019, Cui 18262

(BJFC035121), Cui 18280 (BJFC035139).

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Sun et al.

Notes: Magoderna subresinosum is widely distributed in tropical

and subtropical areas of Asia and Africa (Steyaert 1972). The

taxonomic status of M. subresinosum was controversial for a long

time. Humphrey (1938) regarded it as Ganoderma subresinosum

due to the ganodermoid basidiomata. Corner (1983) suggested

it should be placed in Amauroderma based on a similar hyphal

system and similar basidiospores. However, it can be distinguished

by a slightly shiny and sticky coal black pileal surface, pale and

brous context, ellipsoid to ovoid and non-truncated basidiospores

with faintly verrucose exospore wall and dense spinules on the

endospore wall.

Fig. 63. Basidiomata of Magoderna subresinosum.

Key to accepted species of Magoderna

1a. Pileal surface snuff brown and dull; basidiospores smaller (9–10.5 × 8–9 μm ......................................................... M. infundibuliforme

1b. Pileal surface coal black and slightly shiny; basidiospores larger (14.9–17.3 × 9.9–11.3 μm) ..................................... M. subresinosum

Neoganoderma B.K. Cui & Y.F. Sun, gen. nov. MycoBank MB

840978.

Diagnosis: Differs from other genera by its at to convex pilei

with brown pileal surface, cream context, slightly truncated

basidiospores with longitudinal ridges on the endospore wall which

are equal in length to the basidiospores.

Etymology: neoganoderma (Lat.), refers to the genus producing

Ganoderma-like basidiomata and the distribution in the Neotropics.

Type species: Neoganoderma neurosporum (J.S. Furtado) B.K.

Cui & Y.F. Sun

Description: Basidiomata annual, laterally stipitate or sessile, corky.

Pilei solitary, at to convex. Pileal surface reddish brown to dark

brown, dull, glabrous, concentrically zonate and furrowed. Pore

surface cream to pale cinnamon brown; pores circular. Context

pallid white or cream, soft corky. Hyphal system dimitic; generative

hyphae colourless, thin-walled, branched, with clamp connections;

skeletal hyphae colourless to pale yellow, terminal arboriform or

unbranched. Basidiospores ellipsoid, slightly truncated, pale

yellow, double and thick-walled, endospore wall with longitudinal

ridges which equal in length to the basidiospores.

Notes: So far, Neoganoderma includes N. neurosporum which

has only been collected from Neotropics. The ganoderma-like

basidiomata and haddowia-like ornamentation of endospore

wall make Neoganoderma easily confused with Ganoderma and

Haddowia, but Neoganoderma has unique basidiospores with

longitudinal ridges on the endospore wall without obvious traverse

ridges which are equal in length to the basidiospores. In the

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Ganodermataceae from China

phylogenetic analyses, Neoganoderma formed an independent

clade distinct from other genera within Ganodermataceae and, so

far, it is monotypic (Fig. 1).

Neoganoderma neurosporum (J.S. Furtado) B.K. Cui & Y.F. Sun,

comb. nov. MycoBank MB 840979.

Basionym: Ganoderma neurosporum Furtado, Persoonia 4: 386.

1967.

Description: Basidiomata annual, laterally stipitate or sessile,

corky. Pilei solitary, at to convex, up to 15 cm diam and 3 cm thick.

Pileal surface reddish brown to dark brown, dull, glabrous, with

concentric furrows; margin obtuse, entire. Pore surface cream to

pale cinnamon brown; pores circular, 4–5 per mm. Context pallid

white or cream, soft corky, up to 2.5 cm thick. Tubes pale greyish

brown, up to 1.5 cm long. Stipe dark brown, slightly swollen at

base, up to 10 cm long and 1.5 cm diam. Hyphal system dimitic;

Fig. 64. Microscopic structures of Magoderna subresinosum (drawn from Cui 18280). A. Basidiospores. B. Apical cells from cuticle. C. Basidia and basidioles.

D. Hyphae from trama. E. Hyphae from context. Scale bars = 10 μm.

388

Sun et al.

generative hyphae with clamp connections, colourless, thin-walled,

branched, 2–5 μm diam; skeletal hyphae terminal arboriform or

unbranched, colourless to pale yellow, 3–7 μm diam. Pileipellis

composed of irregularly to slight anticlinal skeletal hyphae and thin

generative hyphae. Basidiospores ellipsoid, slightly truncated, pale

yellow, IKI–, double-walled with thick walls, endospore wall with

longitudinal ridges which equal in length to basidiospores, 16–20

× 11–15 μm.

Notes: The brief description of Neoganoderma neurosporum

was taken from Ryvarden (2004b) and Costa-Rezende et al.

(2020b). According to the records, N. neurosporum is known from

dead wood of deciduous trees in Neotropics. Neoganoderma

neurosporum was placed in Haddowia by its similar endospore wall

ornamentation, but no obvious traverse ridges were observed in

N. neurosporum under SEM. Besides, N. neurosporum formed an

independent lineage in the phylogenetic analyses (Fig. 1). More

detailed description of N. neurosporum need to be made from

future collections.

Sanguinoderma Y.F. Sun et al., Persoonia 44: 224. 2020.

MycoBank MB 828433.

Type species: Sanguinoderma rude (Berk.) Y.F. Sun et al.

For a detailed description of Sanguinoderma, see Sun et al. (2020).

Notes: Sanguinoderma was established by Sun et al. (2020) and

10 species were included in this genus. In this study, six new

species are described based on the main distinguishing character

of Sanguinoderma i.e., the fresh pore surface changes rapidly to

blood red when bruised; there are other morphological features that

differentiate it too.

Sanguinoderma guangdongense B.K. Cui & Y.F. Sun, sp. nov.

MycoBank MB 839664. Figs 65, 66.

Diagnosis: Differs from other species in the genus by its dark pileal

surface with shades of brown concentric zones and dense radial

lines, brous context.

Etymology: guangdongense (Lat.), refers to the holotype of this

species located at Guangdong.

Typus: China, Guangdong, Huizhou, on ground, 19 May 2019, Cui 17259

(holotype BJFC034117).

Additional materials examined: China, Guangdong, Shaoguan, Danxiashan

Nature Reserve, on ground, 4 Jun. 2019, Cui 17240 (BJFC034098);

Yunnan, Yuxi, Longquan Park, on ground of forest, 16 Aug. 2019, Dai

20419 (BJFC032087). Thailand, Chiang Mai, Doi Saket, on ground, 24

Jul. 2016, Dai 16724 (BJFC022831).

Description: Basidiomata annual, centrally to laterally stipitate, hard

corky to woody hard. Pilei solitary, sub-orbicular to umbelliform, up

to 8 cm diam and 7 mm thick. Pileal surface dark yellowish brown to

near black, dull, tomentose, with shades of brown concentric zones

and dense radial lines; margin obtuse, entire, wavy and obviously

incurved when dry. Pore surface pale straw yellow when fresh

Fig. 65. Basidiomata of Sanguinoderma guangdongense.

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Ganodermataceae from China

becoming blood red when bruised and then quickly darkening;

pores circular to angular or irregular, 5–7 per mm; dissepiments

slightly thick, entire. Context wood brown to dark straw yellow, with

dark melanoid lines, hard corky, brous, up to 4 mm thick. Tubes

pale straw yellow to dark yellowish brown, up to 3 mm long. Stipe

slightly darker than pileal surface, cylindrical and hollow, slightly

swollen at base, up to 9.5 cm long and 6 mm diam. Hyphal system

trimitic; generative hyphae with clamp connections, all hyphae IKI

–, CB +; tissues darkening in KOH. Generative hyphae in context

colourless, thin-walled, 4–5 μm diam; skeletal hyphae in context

pale yellow, thick-walled with a wide lumen, arboriform and exuous,

3–7 μm diam; binding hyphae in context colourless, thick-walled,

rarely branched and exuous, 1–2 μm diam. Generative hyphae

in tubes colourless, thin-walled, 3–4 μm diam; skeletal hyphae in

Fig. 66. Microscopic structures of Sanguinoderma guangdongense (drawn from Cui 17259). A. Basidiospores. B. Apical cells from cuticle. C. Basidia and

basidioles. D. Cystidioles. E. Hyphae from trama. F. Hyphae from context. Scale bars = 10 μm.

390

Sun et al.

tubes pale yellow, thick-walled with a wide lumen, arboriform and

exuous, 3–6 μm diam; binding hyphae in tubes colourless, thick-

walled, rarely branched and exuous, up to 2 μm diam. Pileipellis

composed of clamped generative hyphae, slightly thick-walled,

apical cells clavate, faintly inated, dark yellowish brown, about

28–37 × 6–10 μm, forming a regular palisade. Cystidia absent;

cystidioles fusiform, colourless, thin-walled, 14–20 × 4–7 μm.

Basidia barrel-shaped to clavate, colourless, thin-walled, 18–25

× 10–15 μm; basidioles clavate, colourless, thin-walled, 14–20

× 8–13 μm. Basidiospores subglobose to broadly ellipsoid, pale

yellow, IKI –, CB +, double-walled with slightly thick walls, exospore

wall smooth, endospore wall with dense spinules, (9.4–)9.7–10.8(–

11.2) × (8.8–)9–9.8(–10.2) μm, L = 10.26 μm, W = 9.28 μm, Q =

1.1 (n = 60/2).

Notes: Sanguinoderma guangdongense can be distinguished

by a dark brown to almost black pileal surface with shades of

brown concentric zones and dense radial lines. Sanguinoderma

microporum shares the woody hard basidiomata and similar

ornamentation of pilei with Sa. guangdongense, but Sa. microporum

has a pale pileal surface, extremely thick dissepiments of micro

pores and larger basidiospores (11–12 × 8.7–9.8 μm, Sun et al.

2020).

Sanguinoderma infundibulare B.K. Cui & Y.F. Sun, sp. nov.

MycoBank MB 839665. Figs 67, 68.

Diagnosis: Differs from other species in the genus by its funnel-

shaped pilei, yellowish brown to greyish brown pileal surface with

dense and radial ne wrinkles.

Etymology: infundibulare (Lat.), refers to the funnel-shaped pilei.

Typus: China, Guangdong, Shaoguan, Danxiashan Nature Reserve, on

ground, 4 Jun. 2019, Cui 17248 (holotype BJFC034106).

Additional materials examined: China, Guangdong, Shaoguan, Danxiashan

Nature Reserve, on ground of angiosperm forest, 17 Dec. 2017, Dai 18148

(BJFC025677), Dai 18149 (BJFC025678), Dai 18151 (BJFC025680); 4

Jun. 2019, Cui 17238 (BJFC034096), Cui 17256 (BJFC034114).

Description: Basidiomata annual, centrally to laterally stipitate, hard

corky. Pilei solitary, funnel-shape, up to 7.5 cm diam and 6 mm thick.

Pileal surface yellowish brown to greyish brown, dull, tomentose, with

obvious concentric zones, dense and radial ne wrinkles; margin

slightly acute to obtuse, entire and slightly wavy when dry. Pore

surface greyish white when fresh becoming to blood red when bruised

and then quickly darkening; pores circular to angular, 4–6 per mm;

dissepiments slightly thick, entire. Context pale wood brown to greyish

brown, sometimes with dark melanoid lines, corky, up to 4 mm thick.

Tubes pale grey to greyish brown, up to 2 mm long. Stipe concolorous

with pileal surface, cylindrical and hollow, slightly swollen at base, up to

10 cm long and 7 mm diam. Hyphal system trimitic; generative hyphae

with clamp connections, all hyphae IKI –, CB +; tissues darkening in

Fig. 67. Basidiomata of Sanguinoderma infundibulare.

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Ganodermataceae from China

KOH. Generative hyphae in context colourless, thin-walled, 3–4 μm

diam; skeletal hyphae in context pale yellow, thick-walled with a

wide to narrow lumen or sub-solid, arboriform and exuous, 3–7 μm

diam; binding hyphae in context colourless, sub-solid, branched and

exuous, 1–2 μm diam. Generative hyphae in tubes colourless, thin-

walled, 3–4 μm diam; skeletal hyphae in tubes pale yellowish brown,

thick-walled with a wide to narrow lumen or sub-solid, arboriform and

exuous, 3–6 μm diam; binding hyphae in tubes colourless, sub-solid,

branched and exuous, up to 2 μm diam. Pileipellis composed of

clamped generative hyphae, thick-walled, apical cells clavate, faintly

inated and with obvious septa, pale yellowish brown, about 23–30

× 6–11 μm, forming a regular palisade. Cystidia absent; cystidioles

clavate and apices constricted, colourless, thin-walled, 20–35 × 2–6

μm. Basidia barrel-shaped to clavate, colourless, thin-walled, 22–30 ×

14–18 μm; basidioles in shape like the basidia, colourless, thin-walled,

15–25 × 9–19 μm. Basidiospores subglobose to broadly ellipsoid, pale

yellow, IKI –, CB +, double-walled with slightly thick walls, exospore

wall smooth, endospore wall with slightly dense spinules, (10–)10.2–

12(–12.2) × (8.5–)9–10.2(–10.6) μm, L = 11.04 μm, W = 9.53 μm, Q

= 1.16 (n = 60/1).

Fig. 68. Microscopic structures of Sanguinoderma infundibulare (drawn from Cui 17248). A. Basidiospores. B. Apical cells from cuticle. C. Basidia and

basidioles. D. Cystidioles. E. Hyphae from trama. F. Hyphae from context. Scale bars = 10 μm.

392

Sun et al.

Notes: Sanguinoderma infundibulare was collected from the

subtropical areas of China. It is like Amauroderma preussii in the

thin and funnel-shaped pilei with obvious radial wrinkles, but A.

preussii was described from Cameroon with darker and incurved

pilei when dry, larger pores (2–4 per mm) and smaller basidiospores

(7–9.1–11.5 × 6.5–8.5–10 μm, Steyaert 1972, Hapuarachchi et al.

2018a).

Sanguinoderma longistipitum B.K. Cui & Y.F. Sun, sp. nov.

MycoBank MB 839666. Figs 69, 70.

Diagnosis: Differs from other species in the genus by basidiomata

with small pilei and long stipe.

Etymology: longistipitum (Lat.), refers to the basidiomata with a

long stipe.

Typus: China, Yunnan, Honghe, Huanglianshan Forest Park, on ground of

forest, 11 Aug. 2019, Dai 20696 (holotype BJFC032363).

Additional materials examined: China, Yunnan, Jinghong, Xishuangbanna

Botanical Garden, on ground, 23 Jul. 2014, Dai 13891 (BJFC017621);

Hainan, Ledong County, Jianfengling Nature Reserve, on ground of

angiosperm forest, 19 Jun. 2016, Cui 13903 (BJFC028769). Thailand,

Chiang Rai, Doi Mae Salong, on ground of angiosperm forest, 22 Jul.

2016, Dai 16635 (BJFC022745).

Description: Basidiomata annual, laterally stipitate, hard corky.

Pilei solitary, sub-orbicular to abelliform, auricular or spathulate,

up to 4 cm diam and 5 mm thick. Pileal surface greyish brown

to almost black, dull, glabrous, with concentric zones and radial

wrinkles; margin obtuse, entire, slightly wavy and incurved when

dry. Pore surface greyish white when fresh becoming to blood

red when bruised and then quickly darkening; pores circular to

angular, 6–8 per mm; dissepiments moderately thick, entire.

Context wood brown to greyish brown, sometimes with dark

melanoid lines, corky, up to 2 mm thick. Tubes dark grey, up to

4 mm long. Stipe concolorous with pileal surface, cylindrical and

solid, swollen at base, up to 14.5 cm long and 5 mm diam. Hyphal

system trimitic; generative hyphae with clamp connections, all

hyphae IKI –, CB +; tissues darkening in KOH. Generative hyphae

in context colourless, thin-walled, 3–4 μm diam; skeletal hyphae

in context pale yellow, thick-walled with a wide to narrow lumen

or sub-solid, frequently arboriform and exuous, 3–6 μm diam;

binding hyphae in context colourless, sub-solid, branched and

exuous, 1–2 μm diam. Generative hyphae in tubes colourless,

thin-walled, 3–4 μm diam; skeletal hyphae in tubes pale yellow,

thick-walled with a wide to narrow lumen or sub-solid, arboriform

and exuous, 3–5 μm diam; binding hyphae in tubes colourless,

sub-solid, frequently branched and exuous, up to 2 μm diam.

Pileipellis composed of clamped generative hyphae, thick-

walled, apical cells nger-shape, with multiple obvious septa,

pale greyish brown, about 22–40 × 5–10 μm, forming a regular

palisade. Cystidia absent; cystidioles fusiform, colourless, thin-

walled, 16–22 × 5–10 μm. Basidia barrel-shaped, colourless, thin-

walled, 20–27 × 12–17 μm; basidioles in shape like the basidia,

colourless, thin-walled, 19–25 × 5–16 μm. Basidiospores broadly

ellipsoid, pale yellow, IKI –, CB +, double-walled with slightly thick

walls, exospore wall smooth, endospore wall with dense spinules,

10–11(–11.3) × (8–)8.4–9.6(–9.8) μm, L = 10.58 μm, W = 8.94 μm,

Q = 1.18–1.19 (n = 60/2).

Notes: Sanguinoderma longistipitum is a distinct species on account

of its auricular or spathulate pilei with a long stipe. Sanguinoderma

guangdongense has a similar distribution with Sa. longistipitum

which can be collected from Yunnan Province, but the former

can be distinguished by larger basidiomata with sub-orbicular to

umbelliform pilei, shorter stipe and subglobose basidiospores.

Sanguinoderma longistipitum is similar with the small specimens

of Sa. rugosum, which also have broadly ellipsoid basidiospores

in similar size (10.2–11.3 × 8.3–9.2 μm, Sun et al. 2020). However,

the longer stipe and fusiform cystidioles of Sa. longistipitum

distinguish it from Sa. rugosum. Amauroderma auriscalpium which

was described from the Neotropics has similar-shaped pilei, but the

basidiospores in A. auriscalpium are subglobose and smaller (6–8

μm, Torrend 1920).

Fig. 69. Basidiomata of Sanguinoderma longistipitum.

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Ganodermataceae from China

Sanguinoderma melanocarpum B.K. Cui & Y.F. Sun, sp. nov.

MycoBank MB 839667. Figs 71, 72.

Diagnosis: Differs from other species in the genus by its small and

sub-orbicular pilei, coal black pileal surface with alternating dark to

light concentric furrows and strong radial wrinkles.

Etymology: melanocarpum (Lat.), refers to the coal black pileal

surface.

Typus: Malaysia, Selangor, Kota Damansara, Community Forest Reserve,

on ground, 16 Apr. 2018, Dai 18603 (holotype BJFC026891).

Additional material examined: Malaysia, Selangor, Taman Botani Negara

Shah Alam, on stump of angiosperm tree, 12 Apr. 2018, Dai 18512

(BJFC026801).

Description: Basidiomata annual, laterally stipitate, hard corky to

woody hard. Pilei solitary, sub-orbicular to abelliform, up to 4.5

Fig. 70. Microscopic structures of Sanguinoderma longistipitum (drawn from Dai 20696). A. Basidiospores. B. Apical cells from cuticle. C. Basidia and

basidioles. D. Cystidioles. E. Hyphae from trama. F. Hyphae from context. Scale bars = 10 μm.

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Sun et al.

cm diam and 5 mm thick. Pileal surface coal black when fresh,

dull, glabrous, with alternating dark to light concentric furrows

and strong radial wrinkles, centre navel-shaped; margin obtuse,

entire, wavy and incurved when dry. Pore surface cream to

greyish white when fresh becoming to blood red when bruised

and then quickly darkening; pores circular to angular, 6–8 per

mm; dissepiments distinctly thick, entire. Context straw yellow to

yellowish brown, with dark melanoid lines, hard corky, up to 3

mm thick. Tubes greyish brown to dark grey, up to 3 mm long.

Stipe concolorous with pileal surface, cylindrical and hollow,

slightly swollen at base, up to 12 cm long and 5 mm diam. Hyphal

system trimitic; generative hyphae with clamp connections, all

the hyphae IKI + (slightly dextrinoid), CB +; tissues darkening in

KOH. Generative hyphae in context colourless, thin-walled, 3–5

μm diam; skeletal hyphae in context pale yellow, thick-walled with

a wide to narrow lumen or sub-solid, arboriform and exuous,

3–7 μm diam; binding hyphae in context colourless, sub-solid,

branched and exuous, 1–1.5 μm diam. Generative hyphae in

tubes colourless, thin-walled, 3–4 μm diam; skeletal hyphae in

tubes pale yellow, thick-walled with a wide to narrow lumen or

sub-solid, slightly arboriform and exuous, 3–6 μm diam; binding

hyphae in tubes colourless, sub-solid, branched and exuous,

up to 1.5 μm diam. Pileipellis composed of clamped generative

hyphae, thick-walled to sub-solid, apical cells clavate, faintly

constricted and exuous, yellowish brown, about 28–32 × 5–7

μm, forming a regular palisade. Cystidia and cystidioles absent.

Basidia barrel-shaped, colourless, thin-walled, 20–25 × 12–17

μm; basidioles clavate, colourless, thin-walled, 16–22 × 10–16

μm. Basidiospores subglobose to broadly ellipsoid, pale yellow,

IKI + (slightly dextrinoid), CB +, double-walled with slightly thick

walls, exospore wall smooth, endospore wall with dense spinules,

(10–)10.4–11.8(–12) × (8.8–)9–10.5(–10.8) μm, L = 10.98 μm, W

= 9.77 μm, Q = 1.10–1.15 (n = 60/2).

Notes: Sanguinoderma melanocarpum was collected from

Malaysia and it has sub-orbicular pilei, a dark pileal surface with

strongly concentric furrows and radial wrinkles which are similar

to Sa. rugosum, but Sa. rugosum differs from Sa. melanocarpum

by dark brown pilei, slightly thick dissepiments of pores, clavate

cystidioles and elliptical basidiospores (10.2–11.3 × 8.3–9.2 μm)

without amyloid or dextrinoid reaction (Sun et al. 2020).

Fig. 71. Basidiomata of Sanguinoderma melanocarpum.

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Ganodermataceae from China

Sanguinoderma microsporum B.K. Cui & Y.F. Sun, sp. nov.

MycoBank MB 839668. Figs 73, 74.

Diagnosis: Differs from other species in the genus by having the

smallest basidiospores.

Etymology: microsporum (Lat.), refers to the small basidiospores.

Typus: Thailand, Chiang Mai, Doi Saket, on ground, 24 Jul. 2016, Dai

16726 (holotype BJFC022833).

Additional materials examined: China, Hainan, Ledong County,

Jianfengling Nature Reserve, on ground of angiosperm forest, 19 Jun.

2016, Cui 13897 (BJFC028763), Cui 13901 (BJFC028767).

Description: Basidiomata annual, centrally to laterally stipitate,

hard corky. Pilei solitary, near orbicular, up to 4.5 cm diam and 3

mm thick. Pileal surface dark yellowish brown to almost black, dull,

glabrous, with concentric zones and radial wrinkles; margin acute

to obtuse, entire, incurved when dry. Pore surface pale yellowish

brown to pale grey when dry, becoming to blood red when bruised

Fig. 72. Microscopic structures of Sanguinoderma melanocarpum (drawn from Dai 18603). A. Basidiospores. B. Apical cells from cuticle. C. Basidia and

basidioles. D. Hyphae from trama. E. Hyphae from context. Scale bars = 10 μm.

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Sun et al.

and then quickly darkening; pores circular to angular, 5–7 per mm;

dissepiments slightly thick, entire. Context straw yellow, without

dark melanoid lines, corky, up to 2 mm thick. Tubes pale straw

yellow, up to 1 mm long. Stipe concolorous with pileal surface,

cylindrical and hollow, up to 11 cm long and 7 mm diam. Hyphal

system trimitic; generative hyphae with clamp connections, all

hyphae IKI –, CB +; tissues darkening in KOH. Generative hyphae

in context colourless, thin-walled, 3–5 μm diam; skeletal hyphae

in context yellowish brown, thick-walled with a wide to narrow

lumen or sub-solid, arboriform and exuous, 4–7 μm diam; binding

hyphae in context colourless, sub-solid, branched and exuous,

1–2 μm diam. Generative hyphae in tubes colourless, thin-walled,

3–4 μm diam; skeletal hyphae in tubes pale yellow, thick-walled

with a wide to narrow lumen or sub-solid, arboriform and exuous,

3–6 μm diam; binding hyphae in tubes colourless, sub-solid,

branched and exuous, up to 1.5 μm diam. Pileipellis composed

of clamped generative hyphae, slightly thick-walled, apical cells

clavate, slightly inated, reddish brown, about 30–40 × 5–8 μm,

forming a regular palisade. Cystidia and cystidioles absent. Basidia

clavate, colourless, thin-walled, 14–23 × 9–11 μm; basidioles in

shape like the basidia, colourless, thin-walled, 12–17 × 5–10 μm.

Basidiospores subglobose to broadly ellipsoid, pale brown, IKI –,

CB +, double-walled with slightly thick walls, exospore wall smooth,

endospore wall with dense spinules, (4.3–)4.7–5.6(–5.9) × (4–)4.3–

5.2(–5.4) μm, L = 5.22 μm, W = 4.79 μm, Q = 1.06–1.12 (n = 60/2).

Notes: Sanguinoderma microsporum is unique in the genus due

to its small basidiospores. It is like Sa. melanocarpum in the

almost black orbicular pilei with long stipe, but the latter can be

distinguished by smaller pores (6–8 per mm) with distinctly thick

dissepiments, harder context with dark melanoid lines, and larger

basidiospores (10.4–11.8 × 9.0–10.5 μm) with slightly dextrinoid

reaction.

Sanguinoderma tricolor B.K. Cui & Y.F. Sun, sp. nov. MycoBank

MB 839669. Figs 75, 76.

Diagnosis: Differs from other species in the genus by its hard

basidiomata with concentric zonate pileal surface in three different

colours when fresh.

Etymology: tricolor (Lat.), refers to the pileal surface with obvious

concentric zones in three different colours.

Typus: Malaysia, Selangor, Kota Damansara, National Forest Reserve, on

ground, 7 Dec. 2019, Cui 18292 (holotype BJFC035151).

Additional materials examined: Malaysia, Selangor, Kota Damansara,

National Forest Reserve, on ground, 6 Dec. 2019, Cui 18242

(BJFC035101); Forest Research Institute of Malaysia, on stump of Hopea,

15 Apr. 2018, Dai 18574 (BJFC026862).

Description: Basidiomata annual, laterally stipitate, hard corky to

woody hard. Pilei solitary, abelliform to reniform, up to 12 cm diam

and 1 cm thick. Pileal surface rust colour, dark brown to almost

black when fresh, dull, glabrous, with obvious concentric zones in

different colours and radial wrinkles; margin obtuse, entire, very

Fig. 73. Basidiomata of Sanguinoderma microsporum.

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wavy and incurved when dry. Pore surface cream to yellowish

brown when fresh becoming to blood red when bruised and then

quickly darkening, or unchanging in old specimens; pores circular,

5–7 per mm; dissepiments extremely thick (about 0.11–0.14 mm

thick), entire. Context pale straw yellow to wood brown, without

dark melanoid lines, hard corky, up to 3 mm thick. Tubes dark straw

yellow to pale brown, up to 8 mm long. Stipe concolourous with

pileal surface, cylindrical and solid, swollen at base, up to 5.5 cm

long and 1.5 cm diam. Hyphal system trimitic; generative hyphae

with clamp connections, all hyphae IKI –, CB +; tissues darkening

in KOH. Generative hyphae in context colourless, thin-walled, 3–4

μm diam; skeletal hyphae in context pale golden yellow, thick-

walled with a wide to narrow lumen or sub-solid, arboriform and

exuous, 3–7 μm diam; binding hyphae in context colourless, sub-

solid, branched and exuous, 1–2 μm diam. Generative hyphae

in tubes colourless, thin-walled, 3–5 μm diam; skeletal hyphae

Fig. 74. Microscopic structures of Sanguinoderma microsporum (drawn from Dai 16726). A. Basidiospores. B. Apical cells from cuticle. C. Basidia and

basidioles. D. Hyphae from trama. E. Hyphae from context. Scale bars = 10 μm.

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Sun et al.

in tubes pale yellowish brown, thick-walled with a wide to narrow

lumen or sub-solid, arboriform and exuous, 3–6 μm diam; binding

hyphae in tubes colourless, sub-solid, branched and exuous, up

to 2 μm diam. Pileipellis composed of clamped generative hyphae,

thick-walled to sub-solid, apical cells clavate, inated and exuous,

yellowish brown, about 18–27 × 5–7 μm, forming a regular

palisade. Cystidia and cystidioles absent. Basidia barrel-shaped to

clavate, colourless, thin-walled, 15–25 × 10–15 μm; basidioles in

shape like the basidia, colourless, thin-walled, 14–17 × 9–13 μm.

Basidiospores subglobose to broadly ellipsoid, pale yellow, IKI –,

CB +, double-walled with slightly thick walls, exospore wall smooth,

endospore wall with dense spinules, (10–)10.2–11.5(–12) × (8.3–)

8.8–10.2(–10.5) μm, L = 10.91 μm, W = 9.53 μm, Q = 1.13–1.16

(n = 60/2).

Notes: Sanguinoderma tricolor is a distinct species on account of

its pileal surface with concentric zones in three different colours

when fresh. It can be confused with Sa. microporum by having hard

basidiomata, and small pores (5–7 per mm) with extremely thick

dissepiments (about 0.12–0.16 mm thick), but Sa. microporum

differs from Sa. tricolor by the monochromatic pileal surface, context

with dark melanoid lines, larger and more elliptical basidiospores

(11–12 × 8.7–9.8 μm, Q = 1.23–1.28, Sun et al. 2020).

Fig. 75. Basidiomata of Sanguinoderma tricolor.

Key to accepted species of Sanguinoderma

1a. Pore dissepiments extremely thick ......................................................................................................................................................... 2

1b. Pore dissepiments thin to distinctly thick ................................................................................................................................................ 3

2a. Pileal surface pale yellowish brown, pore surface yellowish brown, context with dark melanoid lines .......................... Sa. microporum

2b. Pileal surface rust brown to almost black, pore surface white to pale yellow, context without dark melanoid lines ................ Sa. tricolor

3a. Pore dissepiments lacerate, tubes fascicular when dry ..................................................................................................... Sa. laceratum

3b. Pore dissepiments entire, tubes unchanged when dry .......................................................................................................................... 4

4a. Pores ≤ 4 per mm .................................................................................................................................................................................. 5

4b. Pores > 4 per mm .................................................................................................................................................................................. 7

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Ganodermataceae from China

5a. Pores sinuate; basidiospores > 13.5 μm in length ............................................................................................................. Sa. sinuosum

5b. Pores circular to irregular; basidiospores < 13.5 μm in length ............................................................................................................... 6

6a. Pore dissepiments thin; basidiospores globose to subglobose ....................................................................................... Sa. bataanense

6b. Pore dissepiments slightly thick; basidiospores subglobose to broadly ellipsoid ....................................................................... Sa. rude

7a. Basidiospores < 6 μm in length ..................................................................................................................................... Sa. microsporum

7b. Basidiospores > 6 μm in length .............................................................................................................................................................. 8

8a. Pileal surface coal black; basidiospores slightly dextrinoid in Melzer’s reagent ........................................................ Sa. melanocarpum

8b. Pileal surface brown to almost black; basidiospores IKI– in Melzer’s reagent ....................................................................................... 9

9a. Pileipellis composed of apical cells with obvious septa ....................................................................................................................... 10

9b. Pileipellis composed of apical cells without obvious septa .................................................................................................................. 12

10a. Basidiomata with long stipe; apical cells of pileipellis digitate, with multiple septa ..................................................... Sa. longistipitum

10b. Basidiomata with short stipe; apical cells of pileipellis clavate, with simple septa ............................................................................. 11

11a. Pilei applanate, reniform, pileal surface dark brown to almost black ............................................................................. Sa. elmerianum

11b. Pilei funnel-shape, pileal surface yellowish brown to greyish brown ........................................................................... Sa. infundibulare

12a. Cystidioles absent .............................................................................................................................................................................. 13

12b. Cystidioles present ............................................................................................................................................................................. 14

13a. Pore surface yellowish green when fresh; basidiospores subglobose to broadly ellipsoid ............................................ Sa. avovirens

13b. Pore surface pale grey when fresh; basidiospores reniform ............................................................................................ Sa. reniforme

14a. Basidiomata sessile to subsessile; basidiospores ≥ 14 μm in length ............................................................................. Sa. perplexum

14b. Basidiomata stipitate; basidiospores < 14 μm in length ..................................................................................................................... 15

15a. Pileal surface with shades of brown concentric zones and slender radial lines, context brous ........................... Sa. guangdongense

15b. Pileal surface with concentric furrows and radial wrinkles, context corky ......................................................................... Sa. rugosum

Sinoganoderma B.K. Cui, J.H. Xing & Y.F. Sun, gen. nov.

MycoBank MB 839661.

Diagnosis: Differs from other genera by its ganodermoid

basidiomata, applanate pilei with pale yellow pileal surface, cream

context, thin dissepiments of pores, truncated basidiospores with

an uneven or foveolate exospore wall and solid spinules on the

endospore wall.

Etymology: sinoganoderma (Lat.), refers to the genus producing

ganoderma-like basidiomata and distributed in China.

Type species: Sinoganoderma shandongense (J.D. Zhao & L.W.

Xu) B.K. Cui et al.

Description: Basidiomata annual, stipitate, corky. Pilei solitary,

abelliform to shell-shaped, applanate. Pileal surface pale yellow

to reddish brown, slightly laccate, glabrous, with concentric furrows

and radial wrinkles. Pore surface near white when fresh; pores

circular; dissepiments thin, entire. Context cream to pale wood

brown, without dark melanoid lines, soft corky. Hyphal system

trimitic; generative hyphae colourless, thin-walled, with clamp

connections; skeletal hyphae near colourless to pale yellow, with

narrow lumen or sub-solid, arboriform and exuous; binding hyphae

colourless, thick-walled, branched and exuous. Basidiospores

ellipsoid to ovoid, truncated, pale yellowish brown, double-walled

and distinctly thick-walled, exospore wall uneven or foveolate,

endospore wall with solid spinules.

Notes: Sinoganoderma is established due to its pale yellow pileal

surface, cream context, thin dissepiments of pores, and truncated

basidiospores with an uneven or foveolate exospore wall and solid

spinules on the endospore wall. It is composed of one species

which has been only collected from China. The ornamentation

of the exospore wall of basidiospores observed under SEM is

similar to Foraminispora, but the hollow and columnar spinules

which persist until the exospore wall forming visible holes make

Foraminispora different from other genera in Ganodermataceae. In

the phylogenetic analyses, Sinoganoderma formed an independent

clade distinct from other genera within Ganodermataceae and, so

far, is monotypic (Fig. 1).

Sinoganoderma shandongense (J.D. Zhao & L.W. Xu) B.K. Cui,

J.H. Xing & Y.F. Sun, comb. nov. MycoBank MB 839662. Figs 77,

78.

Basionym: Ganoderma shandongense J.D. Zhao & L.W. Xu, Acta

Mycol. Sin. 5: 90. 1986.

Description: Basidiomata annual, laterally stipitate, corky. Pilei

solitary, abelliform to shell-shaped, applanate, up to 6.5 cm

diam and 2.5 mm thick. Pileal surface pale yellow to reddish

brown, slightly laccate, glabrous, with concentric furrows and

radial wrinkles; margin obtuse, entire. Pore surface near white

when fresh; pores circular, 3–5 per mm; dissepiments thin, entire.

Context cream to pale wood brown, without dark melanoid lines,

soft corky, up to 6 mm thick. Tubes cream, up to 4.3 mm long.

Stipe purplish-red, slightly laccate, cylindrical and solid, slightly

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Sun et al.

swollen at base, up to 5.6 cm long and 1.2 cm diam. Hyphal system

trimitic; generative hyphae with clamp connections, all hyphae IKI

–, CB +; tissues slightly darkening in KOH. Generative hyphae

in context colourless, thin-walled, 2–4 μm diam; skeletal hyphae

in context near colourless to pale yellow, with narrow lumen or

sub-solid, arboriform and exuous, 2–5 μm diam; binding hyphae

in context colourless, thick-walled, branched and exuous, 1–2

diam. Generative hyphae in tubes colourless, thin-walled, 3–4 μm

diam; skeletal hyphae in tubes near colourless, almost sub-solid,

arboriform and exuous, 2–4 μm diam; binding hyphae in tubes

colourless, thick-walled, branched and exuous, up to 1.5 μm diam.

Pileipellis composed of clamped generative hyphae, thick-walled,

apical cells clavate, inated and exuous, pale yellowish brown,

about 37–45 × 5–9 μm, forming a regular palisade. Cystidia and

cystidioles absent. Basidia barrel-shaped to clavate, colourless,

thin-walled, 13–19 × 10–18 μm; basidioles in shape like the basidia,

Fig. 76. Microscopic structures of Sanguinoderma tricolor (drawn from Cui 18292). A. Basidiospores. B. Apical cells from cuticle. C. Basidia and basidioles.

D. Hyphae from trama. E. Hyphae from context. Scale bars = 10 μm.

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Ganodermataceae from China

colourless, thin-walled, 12–16 × 8–10 μm. Basidiospores ellipsoid

to ovoid, truncated, pale yellowish brown, IKI –, CB +, double-

walled with distinctly thick walls, exospore wall uneven or foveolate,

endospore wall with solid spinules, (11.9–)12–13.2(–13.3) × (7.8–)

8–9(–9.2) μm, L = 12.66 μm, W = 8.42 μm, Q = 1.49–1.52 (n = 60/2,

with the turgid vesicular appendix included).

Materials examined: China, Shandong, Changqing County, Liantaishan

Forest Park, on stump of Albizia, 24 Aug. 2015, Dai 15785 (BJFC019889),

Dai 15786 (BJFC019890); on living tree of Albizia, 24 Aug. 2015, Dai 15787

(BJFC019891), Dai 15788 (BJFC019892), Dai 15790 (BJFC019894), Dai

15791 (BJFC019895); on living tree of Albizia, 6 Aug. 2019, Dai 20243

(BJFC031911), Dai 20244 (BJFC031912).

Notes: Sinoganoderma shandongense was rstly described by

Zhao & Zhang (1986a) as Ganoderma shandongense. It has

typical ganodermoid morphology, but Si. shandongense formed

an independent lineage in the phylogenetic analyses (Fig. 1). It is

worth mentioning that the sequences of two specimens (xsd08032

and xsd08085) uploaded into GenBank as G. ramosissimum are

identical with Si. shandongense based on phylogenetic results in

this study, and therefore the name of the two specimens (xsd08032

and xsd08085) needs to change to Si. shandongense.

Tomophagus Murrill, Torreya 5: 197. 1905. MycoBank MB 18657.

Type species: Tomophagus colossus (Fr.) Murrill

Description: Basidiomata annual, sessile, soft corky. Pilei solitary,

abelliform. Pileal surface pale straw yellow to reddish brown,

slightly laccate, glabrous, without ornamentation or not obvious.

Pore surface white to straw yellow; pores circular; dissepiments

thick, entire. Context white to wood brown, soft corky. Hyphal system

dimitic; generative hyphae colourless, thin-walled, branched, with

clamp connections; skeletal hyphae colourless to pale yellow, thick-

walled, thick-walled with narrow lumen or sub-solid, arboriform,

strongly collapsed and exuous. Basidiospores ellipsoid to ovoid,

truncated, yellow to pale yellowish brown, double-walled with

distinctly thick walls, exospore wall slightly foveolate to verrucose

or reticulate, endospore wall with short and irregular ridges.

Notes: Tomophagus has typical ganodermoid basidiospores, but

its pale white and soft context make it different from other genera

(Murrill 1905, Le et al. 2012). Two species, To. colossus and To.

cattienensis are accepted in Tomophagus. In the phylogenetic

analyses, Tomophagus formed an independent clade within

Ganodermataceae (Fig. 1).

Tomophagus cattienensis X.T. Le & Moncalvo, Mycol. Prog. 11:

777. 2012. MycoBank MB 561806. Figs 79, 80.

Description: Basidiomata annual, sessile, soft corky. Pilei solitary,

abelliform, up to 8 cm diam and 3 cm thick. Pileal surface pale straw

yellow when fresh, slightly laccate, glabrous, without ornamentation;

Fig. 77. Basidiomata of Sinoganoderma shandongense.

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Sun et al.

margin obtuse, entire, and incurved when dry. Pore surface pale

wood brown to straw yellow becoming dark when bruised; pores

circular to oval, 3–5 per mm; dissepiments moderately thick, entire.

Context wood brown, with dark resinous lines, soft corky, slightly

brous and powdery, up to 3 cm thick. Tubes pale greyish brown,

corky, up to 5 mm long. Hyphal system dimitic; generative hyphae

with clamp connections, all hyphae IKI –, CB +; tissues darkening

in KOH. Generative hyphae in context colourless, thin-walled,

branched, 3–4 μm diam; skeletal hyphae in context pale yellow,

thick-walled with narrow lumen or sub-solid, arboriform, strongly

collapsed and exuous, 3–5 μm diam. Generative hyphae in tubes

colourless, thin-walled, branched, 3–4 μm diam; skeletal hyphae

in tubes colourless to pale yellow, thick-walled with narrow lumen

or sub-solid, arboriform, strongly collapsed and exuous, 3–4

μm diam. Pileipellis composed of clamped generative hyphae,

thin- to slightly thick-walled, apical cells clavate, faintly inated,

Fig. 78. Microscopic structures of Sinoganoderma shandongense (drawn from Dai 20244). A. Basidiospores. B. Apical cells from cuticle. C. Basidia and

basidioles. D. Hyphae from trama. E. Hyphae from context. Scale bars = 10 μm.

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Ganodermataceae from China

colourless to pale yellow, about 32–45 × 4–7 μm, forming a regular

palisade. Cystidia and cystidioles absent. Basidia barrel-shaped to

clavate, colourless, thin-walled, 18–27 × 10–13 μm; basidioles in

shape like the basidia, colourless, thin-walled, 15–20 × 9–12 μm.

Basidiospores ellipsoid to ovoid, truncated, pale yellowish brown,

IKI –, CB +, double-walled with distinctly thick walls, exospore wall

slightly foveolate, endospore wall with short and irregular ridges,

(10–)10.2–11.8(–11.9) × (6.3–)6.5–7.8(–8) µm, L = 11.09 μm, W

= 7.22 μm, Q = 1.54 (n = 60/1, with the turgid vesicular appendix

excluded); (11.2–)11.3–12.7(–13) × (6.8–)7–8(–8.2) µm, L = 12.04

μm, W = 7.39 μm, Q = 1.63 (n = 60/1, with the turgid vesicular

appendix included).

Material examined: Malaysia, Selangor, Jeram, on dead tree of Elaeis, 10

Apr. 2018, Dai 18487 (BJFC026776).

Notes: Tomophagus cattienensis was described from South

Vietnam, and it can be distinguished from To. colossus by its pale

red-brown and laccate pileal surface, pale brown context when

dry, and slightly larger basidiospores (17.5–21.5 × 11.5–14.5

μm, Le et al. 2012). The specimen of To. cattienensis used in this

study was collected from Malaysia, and it grouped with another

To. cattienensis specimen in the phylogenetic analyses (Fig. 1).

However, the specimens used in Le et al. (2012) showed different

morphological characters to our specimen, such as a reddish brown

pileal surface, larger pores (2–3 per mm), and larger basidiospores

(17.5–21.5 × 11.5–14.5 μm).

Fig. 79. Basidiomata of Tomophagus cattienensis.

Key to accepted species of Tomophagus

1a. Pileal surface reddish brown; basidiospores smaller (10.2–11.8 × 6.5–7.8 µm) ............................................................ To. cattienensis

1b. Pileal surface yellow; basidiospores larger (14–20 × 9–14 µm) .......................................................................................... To. colossus

Trachydermella B.K. Cui & Y.F. Sun, gen. nov. MycoBank MB

840976.

Diagnosis: Differs from other genera by its sessile basidiomata with

atly abelliform pilei, trachytic and ochraceous to yellowish brown

pileal surface, watery context.

Etymology: trachydermella (Lat.), refers to the genus having

trachytic pileal surface.

Type species: Trachydermella tsunodae (Yasuda ex Lloyd) B.K.

Cui & Y.F. Sun

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Sun et al.

Description: Basidiomata annual, sessile, soft when fresh. Pilei

solitary, atly abelliform. Pileal surface ochraceous to yellowish

brown when fresh, dull, glabrous, trachytic, with concentric zones

and radial wrinkles. Pore surface wood brown when dry; pores

circular; dissepiments thick, entire. Context cream, watery when

fresh and turning hard corky when dry. Hyphal system trimitic;

generative hyphae colourless, thin-walled, with clamp connections;

skeletal hyphae pale yellow, thick-walled with narrow lumen or

sub-solid, arboriform and exuous; binding hyphae colourless,

thick-walled, rarely branched and exuous. Basidiospores ellipsoid

to ovoid, truncated, pale yellow, double-walled with distinctly thick

walls, exospore wall verrucose to vermicular, endospore wall with

conspicuous spinules.

Notes: Trachyderma is an illegitimate name as homonym of a

lichen genus in Pannariaceae and was renamed as Trachydermella

Fig. 80. Microscopic structures of Tomophagus cattienensis (drawn from Dai 18487). A. Basidiospores. B. Apical cells from cuticle. C. Basidia and basidioles.

D. Hyphae from trama. E. Hyphae from context. Scale bars = 10 μm.

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Ganodermataceae from China

in this study. Trachydermella is similar to Tomophagus in the pale

white and soft context when fresh, but pale straw yellow pilei with

non-obvious ornamentation, dimitic hyphal system with branched

generative hyphae, and smaller basidiospores of Tomophagus

can distinguish them easily (Murrill 1905, Le et al. 2012). In the

phylogenetic analyses, Trachydermella formed an independent

clade within Ganodermataceae (Fig. 1).

Trachydermella tsunodae (Yasuda ex Lloyd) B.K. Cui & Y.F. Sun,

comb. nov. MycoBank MB 306952. Figs 81, 82.

Basionym: Polyporus tsunodae Yasuda ex Lloyd, Mycol. Writ.

(Cincinnati) 5: 792. 1918.

Description: Basidiomata annual, sessile, soft corky when fresh.

Pilei solitary, atly abelliform, up to 8 cm diam and 3 mm thick.

Pileal surface ochraceous to yellowish brown when fresh, dull,

glabrous, with obvious dark concentric zones and radial wrinkles;

margin obtuse, entire, incurved when dry. Pore surface wood

brown when dry; pores circular, 3–5 per mm; dissepiments

moderately thick, entire. Context cream to pale wood brown,

with dark melanoid lines, watery when fresh and turning hard

corky when dry, up to 1.5 mm thick. Tubes straw yellow, up to 1.2

mm long. Hyphal system trimitic; generative hyphae with clamp

connections, all the hyphae IKI + (dextrinoid), CB +; tissues

darkening in KOH. Generative hyphae in context colourless,

thin-walled, 2–3 μm diam; skeletal hyphae in context pale yellow,

thick-walled with narrow lumen or sub-solid, arboriform and

exuous, 2–5 μm diam; binding hyphae in context colourless,

thick-walled, rarely branched and exuous, 1–1.5 μm diam.

Generative hyphae in tubes colourless, thin-walled, 2–3 μm

diam; skeletal hyphae in tubes pale yellow, thick-walled with

narrow lumen or sub-solid, arboriform and exuous, 2–4 μm

diam; binding hyphae in tubes colourless, thick-walled, rarely

branched and exuous, up to 1.5 μm diam. Pileipellis composed

of clamped generative hyphae, thin- to slightly thick-walled, apical

cells clavate, exuous, yellowish brown, about 25–43 × 3–7

μm, forming a patchy palisade. Cystidia and cystidioles absent.

Basidia barrel-shaped, colourless, thin-walled, 25–35 × 22–24

μm; basidioles in shape like the basidia, colourless, thin-walled,

20–27 × 12–20 μm. Basidiospores ellipsoid to ovoid, truncated,

pale yellow, IKI + (dextrinoid), CB +, double-walled with distinctly

thick walls, exospore wall verrucose to vermicular, endospore

wall with conspicuous spinules, (19.4–)19.8–21.5(–22) × (12.5–)

12.7–15(–15.2) μm, L = 20.48 μm, W = 13.96 μm, Q = 1.47 (n =

60/1, with the turgid vesicular appendix included).

Material examined: China, Guizhou, on dead tree of Litsea cubeba, 17

Jun. 2000, Dai 3221c (BJFC018543).

Notes: The holotype of Trachydermella tsunodae was collected from

Japan, and is distinguished by its pale white and watery context

when fresh, large basidia (25–35 × 22–24 μm) and basidioles

(20–27 × 12–20 μm), and truncated basidiospores with verrucose

to vermicular exospore walls. The specimens of Tr. tsunodae used

in this study were collected from Guizhou Province in southwest

China. Our observations of Tr. tsunodae are generally consistent

with the original description, but the obvious binding hyphae

observed in this specimen are contrary to the dimitic hyphal system

recorded by Imazeki (1952).

Fig. 81. Basidiomata of Trachydermella tsunodae.

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Sun et al.

DISCUSSION

Ganodermataceae is one of the most important families of

macrofungi with many species having important ecological and

economic values (Pilotti 2005, Dai et al. 2009, Glen et al. 2009,

Cao et al. 2012, Chan et al. 2013, Zhou et al. 2015, Jong et al.

2017, Rodríguez-Couto 2017, Zhang et al. 2019, Wang et al.

2021). As species of the family have medicinal, agricultural

and biotechnological application, accurate classication of

Ganodermataceae has been pursued for a long time (Murrill 1905,

Donk 1948, Imazeki 1952, Steyaert 1972, Moncalvo & Ryvarden

1997, Ryvarden 2004b, Costa-Rezende et al. 2017, 2020b, Sun

et al. 2020). Ganodermataceae is a large and complex family and

although many studies have focused on Ganodermataceae, the

species diversity, geographic distribution, species classication,

taxonomy and phylogeny of Ganodermataceae remained uncertain.

Donk (1948) proposed Ganodermataceae as a family, but it

was not widely accepted until the sixth edition of the “Dictionary

of Fungi” published by Ainsworth & Bisby (1971). Since then,

Ganodermataceae has been treated as a family with unique

Fig. 82. Microscopic structures of Trachyderma tsunodae (drawn from Dai 3221c). A. Basidiospores. B. Apical cells from cuticle. C. Basidia and basidioles.

D. Hyphae from trama. E. Hyphae from context. Scale bars = 10 μm.

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Ganodermataceae from China

double-walled basidiospores with particular ornamentation

on the endospore walls in the Polyporales (Ryvarden 2004b,

Costa-Rezende et al. 2017, 2020b, Hapuarachchi et al. 2019b).

Based on morphological observations and the six-gene

combined phylogenetic analyses, 14 genera were conrmed

in Ganodermataceae: Amauroderma, Amaurodermellus,

Cristataspora, Foraminispora, Furtadoella gen. nov., Ganoderma,

Haddowia, Humphreya, Magoderna, Neoganoderma gen. nov.,

Sanguinoderma, Sinoganoderma gen. nov., Tomophagus and

Trachydermella gen. nov. Besides the four new genera, detailed

descriptions for Ganoderma, Haddowia, Humphreya, Magoderna,

Sanguinoderma and Tomophagus are also provided in this study.

The details of Amauroderma, Amaurodermellus, Cristataspora,

Foraminispora and Furtadoella gen. nov. are well-presented in

Costa-Rezende et al. (2017, 2020b) and Sun et al. (2020). In

our work, SEM micrographs of basidiospores of 10 genera in

Ganodermataceae were presented (Fig. 8). The keys for six genera

of Ganodermataceae are provided, and 56 species including 23

new species are described and illustrated.

Neoganoderma was presented as a new genus with one

species, N. neurosporum. The study of N. neurosporum is limited

due to the scarcity of specimens from the Neotropics, and detailed

description was provided by Ryvarden (2004b) and Costa-Rezende

et al. (2020b). Neoganoderma can be distinguished by its unique

basidiospores with longitudinal ridges which equal in length to

basidiospores on the endospore wall.

Sinoganoderma has similar morphological characters with

Ganoderma, such as abelliform pilei with pale reddish brown and

laccate pileal surface, truncated basidiospores with ornamentation

on the endospore wall. However, Sinoganoderma differs by its

applanate pilei with paler pileal surface, cream context, large pores

with thin dissepiments, truncated basidiospores with an uneven

or foveolate exospore wall and solid spinules on the endospore

wall (Fig. 8 H). Sinoganoderma shandongense is the only species

recognised in Sinoganoderma. It was originally described as G.

shandongense from temperate areas of Shandong Province, East

China (Zhao & Zhang 1986a).

Ganoderma is the largest genus in Ganodermataceae including

459 taxa recorded in Index Fungorum (http://www.indexfungorum.

org/) as of 17 April 2021. Considering previous studies and the

current morphological and phylogenetic analyses, 181 species of

Ganoderma are conrmed, including 16 new species; 40 species

was conrmed for China (Moncalvo & Ryvarden 1997, Ryvarden

2004b, Cao et al. 2012, Dai 2012, Hapuarachchi et al. 2018b, Xing

et al. 2018, Hapuarachchi et al. 2019b, Decock & Ryvarden 2020,

Ryvarden 2020). In this study, 95 taxa of Ganoderma with available

molecular data were involved in the phylogenetic analyses and

divided into 10 clades based on the laccate or dull pileal surface

(Fig. 1). The species in subclade I (84 % ML, 1.00 BPP) have

laccate pileal surface except G. mirabile. Subclade II (100 % ML,

1.00 BPP) and subclade III (100 % ML, 1.00 BPP) constituted

monophyletic laccate species. Subclade IV (100 % ML, 1.00 BPP)

and subclade VI (98 % ML, 1.00 BPP) were only composed of the

species with a dull pileal surface. Subclade V (93 % ML, 1.00 BPP)

included the species with dull pileal surface except G. destructans,

G. dunense, G. mutabile and G. pfeifferi which have visibly

obviously laccate pileal surface. Except for G. hoehnelianum and

G. puerense sp. nov., subclade VII (75 % ML, 0.99 BPP), subclade

VIII, subclade IX (100 % ML, 1.00 BPP) and subclade X (99 % ML,

1.00 BPP) were formed by the laccate species. According to the

evolutionary progress in the phylogenetic tree, it might be assumed

that Ganoderma species have evolved from laccate to dull. Whether

the feature of laccate or dull pileal surface can be the key evidence

for reconstructing classication system of Ganoderma remains to

be explored in the future.

Fryssouli et al. (2020) performed a single-gene phylogenetic

analysis for 80 Ganoderma species based on ITS sequences,

and the Ganoderma species were divided into ve clades: Clade

A (including clusters A1, A2, A3), Clade B, Clade C (including

clusters C1, C2), Clade D (including clusters D1, D2, D3, D4),

Clade E (including clusters E1, E2, E3, E4, E5). Clade A consisted

of both laccate species and dull species, and these species form

four different subclades (subclade VII, subclade VIII, subclade

IX, subclade X) in our study. Clade B consisted of dull species,

which is corresponding to subclade VI composed of the species

with dull pileal surface in our study. Clade C includes six species

from Paleotropics with laccate pileal surface in corresponding to

subclade II in our study. Clade D included four clusters, cluster D1

only include G. mbrekobenum, which is corresponding to subclade

III in our study, while clusters D2, D3 and D4 all including laccate

species which formed subclade I in our study. Clade E (including

clusters E1, E2, E3, E4, E5) is consisted of species with both laccate

and dull pileal surface, cluster E1 only includes dull species which

formed subclade IV in our study, while clusters E2, E3, E4 and E5

including both laccate and dull species which formed subclade V

in our study. This indicated that the division of Ganoderma species

by ITS based analysis by Fryssouli et al. (2020) is different from

the division of Ganoderma species by our multiple gene-based

analysis (Fig. 1). Moreover, the ITS based phylogenetic analysis

(Fig. 2) for species of Ganoderma and related genera in our current

study shown that species of Ganoderma were mixed together with

other genera, which indicated that the single ITS based analysis

is not sufcient to investigate the relationship of Ganoderma and

related genera.

Amauroderma s. str., Foraminispora, Furtadoella and

Sanguinoderma were separated from Amauroderma s. lat. based

on morphological and phylogenetic studies (Costa-Rezende et al.

2017, Sun et al. 2020). As of 17 April 2021, 137 taxa of Amauroderma

had been recorded in Index Fungorum (http://www.indexfungorum.

org/), among them 58 species were conrmed as independent

species (Table 2). According to Costa-Rezende et al. (2020a), 24

Amauroderma species were reported from the Neotropics, among

those 16 species were phylogenetically supported. The previous

studies have recorded 24 species of Amauroderma in China (Zhao

& Zhang 2000, Li & Yuan 2015, Song et al. 2016), however, 16

of them have been demonstrated as synonyms or with confused

nomenclatures (Steyaert 1972, Moncalvo & Ryvarden 1997, Li &

Yuan 2015, Sun et al. 2020), and the others need to be abandoned

because of incorrect descriptions or misidentications; for the

time being, there are no species of Amauroderma s. str. Known

from China. Foraminispora was established by Costa-Rezende et

al. (2017) and typied by Fo. rugosa from the Neotropics. Sun et

al. (2020) proposed that, Fo. austrosinensis, Fo. concentrica, Fo.

yinggelingensis and Fo. yunnanensis which were described from

China should be included in Foraminispora based on similar spore

ultrastructure characters and phylogenetic analysis. Furtadoella

consisted of three species from the Neotropics based on soft

basidiomata with dull pileal surface and pale context, dimitic hyphal

system in trama but a monomitic hyphal system in context, with

both clamped and simple-septate generative hyphae (Costa-

Rezende et al. 2017). Sanguinoderma is composed of species

from tropical Asia, Africa and Oceania, with the colour of fresh pore

surface changing to blood red when bruised. In this study, six new

species were described with effective morphological differences

408

Sun et al.

and phylogenetic support, and a total of 16 species were conrmed

in Sanguinoderma.

Costa-Rezende et al. (2020b) established two genera in

Ganodermataceae with adequate analyses, which was further

conrmed in this study. Amaurodermellus is proposed to contain

the Neotropical species: Amauroderma ovisporum based on

amaurodermoid basidiomata with ovoid and non-truncated

basidiospores. Cristataspora is composed of Ganoderma coffeatum

and G. aviporum, which are distinguished by stipitate basidiomata

with white context, and truncated basidiospores with vertical or

transverse ridges on the endospore walls. Ganoderma coffeatum

was recorded by Zhao & Zhang (2000) as Humphreya coffeata in

China, but the specimens stored in HMAS has been determined as

G. lucidum through morphological observation.

Haddowia was established by Steyaert (1972) including Ha.

aëtii and Ha. longipes based on non-truncated basidiospores with

longitudinal ridges partly connected with short transverse walls

on exospore walls (Fig. 8 E). Zmitrovich (2018) combined Ha.

aëtii to Ganoderma as G. aetii. In this study, Ha. macropora was

described from French Guiana as a new species with yellowish

brown pileal surface, large pores, intermittently longitudinal crests

and transverse membranes on the exospore walls.

Humphreya has unique basidiospores with truncated apex

and reticular or erratic irregularly ridged double walls. Humphreya

coffeata has been combined to Cristataspora due to the

longitudinally orientated crests as an independent phylogenetic

branch (Costa-Rezende et al. 2020b). The taxonomic status of Hu.

eminii, Hu. endertii and Hu. lloydii need to be further claried based

on more specimens.

In comparison to other genera, Magoderna has anticlinal

hyphae in the pileipellis, ellipsoid to ovoid basidiospores with faintly

verrucose exospore wall and tiny spinules on the endospore wall

(Fig. 8 F). Magoderna infundibuliforme, M. subresinosum and

M. vansteenisii were rst included in Magonderna when it was

established as a genus (Steyaert 1972). Magoderna vansteenisii

has since been combined as Sanguinoderma rugosum by Corner

(1983) without type specimen examination. And no available

specimen of M. infundibuliforme was examined in this study, so, it

should be kept as separate species in Magoderna before studying

type specimens.

Tomophagus can be distinguished by the pale and soft context,

and dimitic hyphal system with branched generative hyphae (Le

et al. 2012). Tomophagus colossus and To. cattienensis were

included into Tomophagus and formed an independent clade with

good support in the phylogenetic tree (Fig. 1). One specimen

of To. cattienensis collected from Malaysia was examined by

macro-morphology, and microscopic examinations together with

ultrastructural observations. The exospore wall of the basidiospores

in Tomophagus was slightly foveolate to verrucose to reticulate

which resembles Ganoderma, but the ornamentation of the latter is

deeper (Fig. 8D, I). The detailed descriptions of Tomophagus given

in this study make it more credible and recognisable as genus in

Ganodermataceae.

Trachyderma was renamed as Trachydermella due to its

illegitimacy, and has one species so far, Tr. tsunodae. In this

study, Tr. tsunodae showed to be an independent clade with high

support in the phylogenetic tree (Fig. 1). The specimen collected in

southwestern China was conrmed by the watery context when fresh

and large hyphae, spores and basidia, since no sequence could be

generated. The ornamentation of basidiospores in Trachydermella

has been observed under SEM, and shows a similar verrucose to

reticulate exospore wall to that in Amauroderma and Furtadoella.

However, the distinctly truncated apex of the basidiospores

distinguishes Trachydermella from Amauroderma and Furtadoella

(Fig. 8A, C, J).

After several studies, 642 taxa of Ganodermataceae were

recorded in Index Fungorum (http://www.indexfungorum.org/) as

of 10 March 2022, among which Amauroderma which has 141

records, Amaurodermellus has one species, Cristataspora has

two species, Foraminispora has ve species, Furtadoella has

three species, Ganoderma has 467 records, Haddowia has three

records, Humphreya has four records, Magoderna has three

records, Tomophagus has two species, Trachydermella has one

species and Sanguinoderma has 10 species. According to the

nomenclatural study of Ganodermataceae by Moncalvo & Ryvarden

(1997) and other studies, there are four major reasons to abandon

some species: i) they are synonyms of already named species, ii)

the type specimens are lost or immature, iii) they represent invalid

names, and iv) there are errors in sequencing and nomenclature.

Based on these reasons, we have now conrmed 278 species in

the world (Table 2), and 145 of them have molecular data.

In the past few decades, many scholars have focused on

exploring the diversity of Ganodermataceae with 130 taxa of

Ganodermataceae in China (Zhao & Zhang 2000, Cao & Yuan

2012, Dai 2012, Wang & Wu 2014, Li et al. 2015, Zhou et al. 2015,

Hapuarachchi et al. 2018b, Xing et al. 2018, Ye et al. 2019, Sun et

al. 2020, He et al. 2021). Previously, the mostly recorded species

were recognised only based on morphological characters. However

in recent years, phylogenetic analyses have applied to the studies

of Ganodermataceae. As a result of these studies, more than

30 species have been regarded as synonyms, and several taxa

should be abandoned due to being immature specimens without

basidiospores or confusing original descriptions (Wang 2005, Cao

2013, Li & Yuan 2015, Xing 2019). Sun et al. (2020) classied seven

species originally reported as Amauroderma in China as other

species, and Costa-Rezende et al. (2020b) combined Humphreya

coffeatum to Cristataspora. In this study, the remnant of the species

was checked based on morphological comparisons and geographic

distribution together with phylogenetic analyses. For the time

being, 59 species of Ganodermataceae are recognised in China,

of which only one without available sequences. The recorded taxa

of Ganodermataceae reported in China and its current taxonomic

status are presented in Table 3.

Just recently, two additional new species of Ganoderma, G.

dianzhongense and G. esculentum were described by He et al.

(2021), the morphological characters and molecular evidence were

sufcient to recognize their legitimacy even if they were not included

in our current phylogenetic analyses. Both of them have been listed

in Tables 2 and 3 along with other species of Ganoderma in China.

Since ITS and nLSU sequences were rst used to identify

Ganoderma species in Moncalvo et al. (1995), many DNA

sequences have been uploaded to GenBank (https://www.ncbi.

nlm.nih.gov/). These sequences are mostly generated from eight

genes: ITS, nLSU, rpb1, rpb2, tef1, tub, mtSSU and nSSU; rpb1

and tub are not widely for Ganodermataceae due to insufcient

quantity. To evaluate the practicability and reliability of six genes,

the phylogenetic analyses of Ganodermataceae based on ITS,

nLSU, rpb2, tef1, mtSSU and nSSU sequences were carried out

respectively (Figs 2–7). The internal transcribed spacer region

(ITS) was considered as the universal barcode of fungi (Schoch

et al. 2012), but its limitation in identifying complex groups or

potential species cannot be ignored (Badotti et al. 2017) even if

ITS is the most abundant gene region in Ganodermataceae. Loci

such as rpb2 and tef1 are very useful for identifying the species

409www.studiesinmycology.org

Ganodermataceae from China

Table 3. Taxonomic status of Ganodermataceae reported from China.

Genus Taxa Current status References

Amauroderma (24) A. amoiense = Sanguinoderma rugosum Li & Yuan (2015)

A. auriscalpium Nomenclature unclear and holotype sterile Moncalvo & Ryvarden (1997)

A. austrosinense = Foraminispora austraosinensis Sun et al. (2020)

A. bataanense = Sanguinoderma bataaense Sun et al. (2020)

A. concentricum = Foraminispora concentrica Sun et al. (2020)

A. conjunctum Inconsistent with original description This study

A. dayaoshanense = Pyrrhoderma sendaiense Li & Yuan (2015)

A. elmerianum = Sanguinoderma elmerianum Sun et al. (2020)

A. exile Specimens lost and only distributed in Neotropics This study

A. fujianense = Ganoderma fornicatum Li & Yuan (2015)

A. guangxiense Type specimen lost This study

A. hongkongense Holotype polluted and sterile This study

A. jiangxiense Ganodermoid basidiocarps and sterile This study

A. longgangense Ganodermoid basidiocarps and sterile This study

A. nigrum Nomenclature unclear Moncalvo & Ryvarden (1997)

A. perplexum = Sanguinoderma perplexum Sun et al. (2020)

A. preussii = Sanguinoderma infundibulare sp. nov. This study

A. rude = Sanguinoderma rude Sun et al. (2020)

A. rugosum = Sanguinoderma rugosum Sun et al. (2020)

A. schomburgkii = Sanguinoderma elmerianum This study

A. sikorae = Amauroderma preussii Steyaert (1972)

A. subresinosum = Magoderna subresinosum Sun et al. (2020)

A. wuzhishanense = Amauroderma rugosum Li & Yuan (2015)

A. yunnanense = Foraminispora yunnanensis Sun et al. (2020)

Ganoderma (104) G. ahmadii √This study

G. albomarginatum Nomenclature repeated with same specimen Xing (2019)

G. amboinense Nomenclature unclear Moncalvo & Ryvarden (1997)

G. angustisporum √Xing et al. (2018)

G. annulare = Ganoderma australe Ryvarden (1989)

G. applanatum √Dai (2012)

G. atrum = Ganoderma exipes Cao (2013)

G. australe √Dai (2012)

G. austrofujianense = Ganoderma sinense Cao (2013)

G. bawanglingense = Ganoderma australe This study

G. bicharacteristicum Holotype sterile Xing (2019)

G. boninense √Wang (2005)

G. brownii Inconsistent with original description Wang (2005)

G. calidophilum √This study

G. cantharelloideum = Ganoderma lucidum Cao (2013)

G. capense = Ganoderma weberianum Wang (2005)

G. casuarinicola √Xing et al. (2018)

G. chalceum Inconsistent with original description This study

G. chenghaiense = Ganoderma multipileum Cao (2013)

G. chiungchungense Description and type specimen unclear Xing (2019)

G. cochlear Nomenclature unclear and type specimen lost Moncalvo & Ryvarden (1997)

G. colossus = Tomophagus colossus Cao (2013)

G. crebrostriatum = Ganoderma mastoporum Cao (2013)

G. cupulatiprocerum = Ganoderma duropora Zhao & Zhang 2000

G. curtisii Inconsistent with original description Xing (2019)

410

Sun et al.

Table 3. (Continued).

Genus Taxa Current status References

G. daiqingshanense = Ganoderma multiplicatum Cao (2013)

G. densizonatum = Ganoderma orbiforme Wang et al. (2014)

G. dianzhongense √He et al. (2021)

G. diaoluoshanense = Ganoderma mastoporum Cao (2013)

G. dimidiatum Description and type specimen unclear This study

G. donkii Inconsistent with original description Moncalvo & Ryvarden (1997)

G. duropora Inconsistent with original description This study

G. ellipsoideum √Hapuarachchi et al. (2018b)

G. esculentum √He et al. (2021)

G. exipes √Dai (2012)

G. formosanum = Ganoderma sinense Cao (2013)

G. fornicatum = Ganoderma orbiforme Wang et al. (2014)

G. fulvellum = Fomes fulvellus This study

G. gibbosum √Luangharn et al. (2020)

G. guinanense = Ganoderma sinense Cao (2013)

G. guizhouense Description and type specimen unclear Xing (2019)

G. hainanense = Ganoderma exipes Cao (2013)

G. hoehnelianum √This study

G. jianfenglingense Description and type specimen unclear Xing (2019)

G. kunmingense Holotype sterile Cao (2013)

G. leucocontextum √Li et al. (2015)

G. limushanense = Ganoderma orbiforme Wang et al. (2014)

G. lingzhi √Cao et al. (2012)

G. lobatum Inconsistent with original description This study

G. lucidum √This study

G. luteomarginatum = Ganoderma sinense Cao (2013)

G. magniporum √This study

G. mastoporum = Ganoderma orbiforme Wang et al. (2014)

G. mediosinense = Ganoderma sinense Cao (2013)

G. meijiangense = Ganoderma williamsianum Wang & Wu (2010)

G. microsporum = Ganoderma weberianum Cao (2013)

G. mirabile Climate different with type locality This study

G. mirivelutinum = Ganoderma australe This study

G. mongolicum Unlike Ganoderma Wang (2005)

G. multipileum √Dai (2012)

G. multiplicatum Inconsistent with original description This study

G. mutabile √Cao & Yuan (2012)

G. neojaponicum Inconsistent with original description Wang (2005)

G. nigrolucidum Inconsistent with original description Wang (2005)

G. nitidum Climate different with type locality This study

G. ochrolaccatum Inconsistent with original description This study

G. orbiforme √Wang et al. (2014)

G. ostracodes Inconsistent with original description Wang (2005)

G. parviungulatum = Ganoderma exipes Cao (2013)

G. petchii Inconsistent with original description This study

G. pfeifferi Inconsistent with original description Wang (2005)

G. philippii √Dai (2012)

G. ramosissimum Holotype sterile Cao (2013)

G. renii Description unclear and type specimen lost Cao (2013)

411www.studiesinmycology.org

Ganodermataceae from China

Table 3. (Continued).

Genus Taxa Current status References

G. resinaceum Inconsistent with original description Xing (2019)

G. rotundatum = Ganoderma multiplicatum Cao (2013)

G. sanduense √Hapuarachchi et al. (2019b)

G. sanmingense Holotype sterile Cao (2013)

G. shandongense = Sinoganoderma shandongense comb. nov. This study

G. shangsiense = Ganoderma hoehnelianum Wang & Wu (2010)

G. shanxiense √Liu et al. (2019)

G. sichuanense √Wang et al. (2012)

G. simaoense Holotype sterile Cao (2013)

G. sinense √Dai (2012)

G. stipitatum = Ganoderma tropicum Wang (2005)

G. stratoideum Description unclear and type specimen lost Xing (2019)

G. subumbraculum = Ganoderma weberianum Wang (2005)

G. tenue = Ganoderma weberianum Cao (2013)

G. theaecola = Ganoderma multiplicatum Cao (2013)

G. tibetanum Description and type specimen unclear Xing (2019)

G. triangulum = Ganoderma australe This study

G. tropicum √Dai (2012)

G. trulla Inconsistent with original description This study

G. tsugae √Dai (2012)

G. tsunodae = Trachydermella tsunodae comb. nov. This study

G. ungulatum = Ganoderma australe This study

G. valesiacum Inconsistent with original description Wang (2005)

G. weberianum √Dai (2012)

G. weixiense √Ye et al. (2019)

G. williamsianum √Dai (2012)

G. wuhuense Description unclear and type specimen lost Xing (2019)

G. wuzhishanense Molecular sequence error Xing (2019)

G. xingyiense Description unclear and type specimen lost Xing (2019)

G. zhenningense Description and type specimen unclear Xing (2019)

Haddowia (1) Ha. longipes √Dai (2012)

Humphreya (1) Hu. coffeatum Inconsistent with original description This study

in Ganodermataceae, but the instability of them usually produces

uncontrollable mutations. Compared to other genes, nLSU,

mtSSU and nSSU are so conservative that it is hard to delimit

the species in Ganodermataceae using just these genes. Thus,

phylogenetic analyses based on a gene locus alone is insufcient

and a combined multi-gene dataset with ITS, nLSU, rpb2, tef1,

rpb1 and tub, is better recommended for phylogenetic analyses

of Ganodermataceae. In this study, 1 382 sequences from 391

specimens were used in the phylogenetic analyses which included

63 type specimens. There were 817 sequences newly generated

and uploaded to GenBank, including 132 sequences of ITS, 139

sequences of nLSU, 83 sequences of rpb2, 124 sequences of

tef1, 150 sequences of mtSSU and 189 sequences of nSSU. The

reliability of these sequences was referenced by literature citations,

released information on NCBI and practical application, which

suggests that the sequences used in this study should be the basis

of future phylogenetic analyses of Ganodermataceae.

ACKNOWLEDGEMENTS

We express our gratitude to Prof Yu-Cheng Dai (China) for providing

specimens and pictures for our study. The curators of herbaria of HMAS,

HKAS, GDGM, IFP and Drs Tom May (Australia), Hai-Jiao Li (China), Jie

Song (China), Lu-Lu Shen (China), Mei-Ling Han (China), Jun-Liang Zhou

(China) and Ms. Xing Ji (China), Yan Wang (China), Yu-Li Han (China)

are thanked for help during eld collections and molecular studies. Drs

Shuang-Hui He (China), Fang Wu (China), Hai-Sheng Yuan (China), Yu-

Lian Wei (China), Li-Wei Zhou (China), Jun-Zhi Qiu (China), Hai-Xia Ma

(China), Bo Zhang (China), Shi-Liang Liu (China), Ming Zhang (China),

De-Wei Li (USA) are thanked for companionship during eld collections.

Dr Sheng-Hua Wu (China) is thanked for providing valuable suggestions

on this study. The research was supported by the National Natural Science

Foundation of China (Nos. U2003211, 31870008, 31670016), the Scientic

and Technological Tackling Plan for the Key Fields of Xinjiang Production

and Construction Corps (No. 2021AB004) and Beijing Forestry University

Outstanding Young Talent Cultivation Project (No. 2019JQ03016).

412

Sun et al.

DECLARATION ON CONFLICT OF INTEREST

The authors declare that there is no conict of interest.

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FOREST PATHOL

In September 2022, hazelnut trees in six orchards in the Çarşamba district of Samsun province, a major hazelnut production area in the Black Sea region of Türkiye, showed symptoms of leaf yellowing, shoot dieback, decline, basal stem rot and the presence of brown bracket‐like sessile basidiocarps. Approximately 5%–10% of hazelnut trees were affected, highlighting the significant impact of the disease. A comprehensive morphological analysis was conducted to characterize fungal isolates obtained from symptomatic tissues as G . adspersum . This analysis incorporated features observed in basidiomata from declining trees and those reproduced after isolates were cultivated on wheat grain for spawn production and grown on an oak sawdust‐based substrate. Confirmation of identification was achieved through the utilization of the internal transcribed spacer (ITS), large subunit (LSU) and small subunit (SSU) loci, along with sequencing and subsequent phylogenetic analysis. The G . adspersum isolates generally exhibited no significant growth differences at similar temperatures but demonstrated enhanced growth at 30°C, while growth was completely inhibited at 40°C. Incompatible reactions between isolates confirmed that distinct genotypes were present, highlighting genetic diversity within the species. In the pathogenicity trials, employing a single infested wheat grain per inoculation at the wounded site, hazelnut suckers exhibited distinct brown discolouration surrounding the inoculation site following a 2.5‐month incubation period. Significantly, lesions exceeding 3 cm in length were observed, providing clear evidence of vigorous pathogenic activity by G . adspersum . This study is the first report of G . adspersum causing butt rot in hazelnuts, underscoring the significance of this finding for hazelnut cultivation and providing a foundation for future research and disease management strategies.

Two new species of Ganoderma (Ganodermataceae, Basidiomycota) from Southwest China

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Full-text available

Jun 2024

Ganoderma is a large and diverse genus containing fungi that cause white rot to infect a number of plant families. This study describes G. phyllanthicola and G. suae as new species from Southwest China, based on morphological and molecular evidence. Ganoderma phyllanthicola is characterized by dark brown to purplish black pileus surface with dense concentric furrows, pale yellow margin, irregular pileipellis cells, small pores (5–7 per mm) and ellipsoid to sub-globose basidiospores (8.5–10.0 × 6.0–7.5 µm). Ganoderma suae is characterized by reddish brown to oxblood red pileus surface and lead gray to greyish-white pore surface, heterogeneous context, wavy margin and almond-shaped to narrow ellipsoid basidiospores (8.0–10.5 × 5.0–7.0 μm). The phylogeny of Ganoderma is reconstructed with multi-gene sequences: the internal transcribed spacer region (ITS), the large subunit (nrLSU), translation elongation factor 1-α gene (TEF-1α) and the second subunit of RNA polymerase II (RPB2). The results show that G. suae and G. phyllanthicola formed two distinct line-ages within Ganoderma. Descriptions, illustrations and phylogenetic analyses results of the two new species are presented.

A New Species of Ganoderma (Ganodermataceae, Polyporales) from Southern China and Optimum Condition for Mycelia Production

Article

Full-text available

Feb 2024

The present study sought to propose Ganoderma guixiense sp. nov. as a new species based on phenotypic and genotypic evidence. Phylogenetic analyses were carried out based on the internal transcribed spacer (ITS), the large subunit of nuclear ribosomal RNA gene (nLSU), and the second subunit of RNA polymerase II (RPB2) sequence data. G. guixiense has been characterized by pileate basidiomata, long stipe, in addition to reddish-black zonate pileal surface. Basidiospores are broadly ellipsoid with one end tapering at maturity, and measuring 9–12.8 × 6.5–9.3 μm. Basidia are oval to subglobose. This study marks the first exploration of the biological characteristics of G. guixiense. The result indicated that the optimal medium of mycelial growth was observed on malt extract agar (MEA) and yeast extract peptone dextrose agar (YPD) while the optimal temperature was found to be 25–30 °C with pH range of 6–7.

Ganoderma pfeifferi Bres. 1889 and Ganoderma resinaceum Boud. 1889 as Potential Therapeutic Agents: A Comparative Study on Antiproliferative and Lipid-Lowering Properties

Preprint

Full-text available

Jun 2024

Medicinal mushrooms, especially Ganoderma species hold immense promise for production of a wide range of bioactive compounds with various effects. The biochemical potential of indigenous fungal strains, specific to a region, could play a critical role in the continuous search for novel strains with superior activities on a global scale. This research focused on the ethanolic (EtOH) and hot-water (H2O) extracts of fruiting bodies of two wild-growing Ganoderma species: G. pfeifferi and G. resinaceum with an aim to assess its nutritional (total carbohydrate content-TCC), and mineral composition in relation to bioactive properties: antioxidant, antiproliferative and lipid-lowering. Atomic absorption spectrophotometry (AAS) revealed that G. pfeifferi is a promising source of minerals that are essential for numerous physiological functions in the human body like bone health and muscle and nerve function, with calcium (Ca 4.55 ± 0.41 mg/g d.w.) and magnesium (Mg 1.33 ± 0.09 mg/g d.w.), being the most abundant macro element present. Zinc (Zn), manganese (Mn), and chromium (Cr) were particularly notable, with concentrations ranging from 21.49 to 41.70 mg/kg d.w. The EtOH extract of G. pfeifferi demonstrated significantly elevated levels of TCC, essential macromolecules for energy and structural functions in the body, with higher quantities of all three standard carbohydrates detected in this type of extract. Similar to revealed composition the same species, G. pfeifferi stood out as the most prominent antioxidant agent, with H2O extract being stronger than EtOH in ABTS assay (86.85 ± 0.67 mg TE/g d.w.), while EtOH extract displayed the highest anti OH• radical scavenging ability (IC50 = 0.18 ± 0.05 μg/mL) as well as the most notable reducing potential among all. The highest antiproliferative effect against the breast cancer cell line (MCF-7), demonstrated the H2O extracts from G. resinaceum with the most pronounced activity after 24 hours (IC50 = 4.88 ± 0.50 μg/mL) which surpasses that of the standard compound, ellagic acid (IC50 = 33.94 ± 3.69 μg/mL). Administration of both Ganoderma extracts mitigated diabetic lipid disturbances and exhibited potential renal and hepatic protection in vivo on white Wistar rats by preservation of kidney function parameters in G. resinaceum H2O pre-treatment (urea: 6.27 ± 0.64 mmol/L, creatinine: 50.00 ± 6.45 mmol/L) and the reduction of ALT levels (17.83 ± 3.25 U/L) com-pared to diabetic control groups treated with saline (urea: 46.98 ± 6.01 mmol/L, creatinine: 289.25 ± 73.87 mmol/L, and ALT: 60.17 ± 9.64 U/L). These results suggest that pre-treatment with G. resinaceum H2O extracts may have potential antidiabetic properties. In summary, detected microelements are vital for maintaining overall health, supporting metabolic processes, and protecting against various chronic diseases. Further research and dietary assessments could help determine the full potential and applications of two underexplored Ganoderma species native to Serbia in nutrition and health supplements.

Perspectivas do potencial imunomodulador e antineoplásico de polissacarídeos e outras biomoléculas de Ganoderma spp.

Article

Full-text available

Apr 2024

Os polissacarídeos derivados de espécies fúngicas pertencentes ao gênero Ganoderma têm sido reconhecidos como agentes importantes tanto na modulação do sistema imunológico quanto na inibição do crescimento tumoral. Este estudo objetivou destacar a capacidade antineoplásica e imunomoduladora in vitro dos polissacarídeos de Ganoderma. Foi demonstrado que esses polissacarídeos são capazes de estimular a atividade de células imunes, fortalecendo assim resposta imunológica contra células tumorais. Adicionalmente, esses compostos naturais mostram-se capazes de prevenir, bloquear ou mesmo reverter o desenvolvimento do câncer, com potencial para mitigar os efeitos colaterais associados aos tratamentos convencionais. É importante ressaltar que esses compostos também exibem atividades antineoplásicas, retardando o crescimento e a disseminação de tumores. O potencial imunomodulador e antineoplásico dos polissacarídeos de Ganoderma pode representar uma fonte promissora de tratamentos adjuvantes e alternativos para o câncer e outras doenças relacionadas ao sistema imunológico. Diante dos desafios e limitações da quimioterapia, os medicamentos fitoterápicos emergem como alternativas promissoras no tratamento do câncer. Nesse contexto, os polissacarídeos extraídos de Ganoderma, um cogumelo medicinal, despertam crescente interesse devido às suas propriedades comprovadas de quimioprevenção e antineoplásicas.

Four New Fungal Species in Forest Ecological System from Southwestern China

Article

Full-text available

Mar 2024
J. Fungi

Four new wood-inhabiting fungi were found in Southwestern China within the genera Phanerochaete, Phlebiopsis, Asterostroma, and Vararia of the families Phanerochaetaceae and Peniophoraceae, belonging to the orders Polyporales and Russulales individually. Combined with their morphological characteristics and molecular biological evidence, the present study describes them as new fungal taxa. Asterostroma yunnanense is characterized by the resupinate, membranaceous to pellicular basidiomata with a cream to salmon-buff hymenial surface, hyphal system dimitic bearing simple-septa, thin- to thick-walled, yellowish brown asterosetae with acute tips, and thin-walled, echinulate, amyloid, globose basidiospores. Phanerochaete tongbiguanensis is characterized by the resupinate basidiomata with a white to cream hymenial surface, a monomitic hyphal system with simple-septa generative hyphae, the presence of subclavate cystidia covered with a lot of crystals, and oblong ellipsoid basidiospores (6–9 × 3–4.5 µm). Phlebiopsis fissurata is characterized by the membranaceous, tuberculate basidiomata with a buff to slightly brown hymenial surface, a monomitic hyphal system with simple-septa, conical cystidia, and broadly ellipsoid. Vararia yingjiangensis is characterized by a corky basidiomata with a pinkish buff to cinnamon-buff hymenial surface, cracking, yellowish dichohyphae with slightly curved tips, subulate gloeocystidia, and thick-walled, ellipsoid basidiospores (6.5–11.5 × 5–7 µm). The phylogenetic analyses of ITS + nLSU revealed that the two new species were nested into the genera Phanerochaete and Phlebiopsis within the family Phanerochaetaceae (Polyporales), in which Phanerochaete tongbiguanensis was sister to P. daliensis; Phlebiopsis fissurata was grouped with P. lamprocystidiata. Two new species were clustered into the genera Asterostroma and Vararia within the family Peniophoraceae (Russulales), in which Asterostroma yunnanense was sister to A. cervicolor; Vararia yingjiangensis formed a single branch.

Fresh-eating Lingzhi becomes possible: Comparative evaluation of nutritional and taste profile of Ganoderma tsugae at different fruiting body morphogenesis stages

Article

May 2024
LWT-FOOD SCI TECHNOL

Strategy on rapid discrimination of different varieties based on the combination of HS-GC-IMS and DNA Mini-barcode, spore powder of Ganoderma as a case study

Article

Feb 2024
MICROCHEM J

Whole-genome sequencing and comparative genomic analyses of the medicinal fungus Sanguinoderma infundibulare in Ganodermataceae

Article

Full-text available

Feb 2024

Sanguinoderma infundibulare is a newly discovered species of Ganodermataceae known to have high medicinal and ecological values. In this study, the whole-genome sequencing and comparative genomic analyses were conducted to further understand Ganodermataceae's genomic structural and functional characteristics. Using the Illumina NovaSeq and PacBio Sequel platforms, 88 scaffolds were assembled to obtain a 48.99-Mb high-quality genome of S. infundibulare. A total of 14,146 protein-coding genes were annotated in the whole genome, with 98.6% of complete benchmarking universal single-copy orthologs (BUSCO) scores. Comparative genomic analyses were conducted among S. infundibulare, Sanguinoderma rugosum, Ganoderma lucidum, and Ganoderma sinense to determine their intergeneric differences. The 4 species were found to share 4,011 orthogroups, and 24 specific gene families were detected in the genus Sanguinoderma. The gene families associated with carbohydrate esterase in S. infundibulare were significantly abundant, which was reported to be involved in hemicellulose degradation. One specific gene family in Sanguinoderma was annotated with siroheme synthase, which may be related to the typical characteristics of fresh pore surface changing to blood red when bruised. This study enriched the available genome data for the genus Sanguinoderma, elucidated the differences between Ganoderma and Sanguinoderma, and provided insights into the characteristics of the genome structure and function of S. infundibulare.

Polyhydroxy Sterols Isolated From Ganoderma sinense Spores and Their Cytotoxic Activities

Article

Full-text available

Jan 2024
NAT PROD COMMUN

Background: The fruiting body of Ganoderma sinense Zhao, Xu et Zhang iscertified as an authoritative medical material in the Chinese Pharmacopoeia 2020 edition and has been used as a crude drug for a long time. G sinense spores are crucial part of the fruiting body of G sinense and exhibited antitumor and immune-enhancing activities according to our previous reports. However, there were few studies about the chemical composition of G sinense spores. This study was aimed at the structure determination and antitumor effect of the sterols isolated from G sinense spores. Methods: Modern chromatographic methods were applied to isolate compounds from the ruptured spores of G sinense. Their chemical structures were elucidated by analyses of one-dimensional and two-dimensional ¹H nuclear magnetic resonance, and high-resolution electrospray ionization mass spectrometry data. The cytotoxicities of the isolated compounds against 3 tumor cell lines, human non-small cell lung cancer A549 and 95D cells, and human hepatocellular carcinoma HepG2 cells were measured by Cell Counting Kit-8. Results: A novel ergosterol derivative, (22E,24R)-3β,5β,6α,7α,14β-pentahydroxyergosta-8,22-dien-15-one (1), and 5 known sterols, (22E,24R)-3β,5α,9α,14β-tetrahydroxyergosta-7,22-dien-6-one (2), (22E,24R)-ergosta-7,9(11),22-triene-3β,5α,6β-triol (3), (22E,24R)-ergosta-7,9(11),22-triene-3β,5α,6β,14α-tetrol (4), β-daucosterine (5), and (22E,24R)-ergosta-7,22-diene-3β,5α,6β,9α-tetrol (6), were obtained from the ruptured spores of G sinense. Compound 1 exhibited cytotoxicity against A549 cells and HepG-2 cells with IC50 values of 65.12 ± 4.76 and 97.34 ± 6.36 μM, respectively. Compounds 1, 2, and 4 were obtained from G sinense for the first time. Conclusion: This paper is a continuation of an investigation of the chemical ingredients from G sinense spores and their antitumor effect. Only compound 1 showed weak cytotoxicities against A549 cells and HepG-2 cells.

Taxonomic study on a threatened polypore, Polyporus pseudobetulinus, and a morphologically similar species, P. subvarius

Article

Full-text available

Jan 2011

A taxonomic study based on morphological and phylogenetic characters revealed that the specimens referred to as ‘Polyporus pseudobetulinus’ in Japan actually represent two distinct species, namely, P. pseudobetulinus and P. subvarius. These cam be distinguished by characteristics of the pileus surface, the presence/absence of clamp-connections on generative hyphae, and the shape of the basidiospores. P. subvarius is newly reported from Japan following the original description from China. The phylogenetic positions, detailed descriptions, and illustrations are provided for both species.

Phylogenetic analyses and morphological characters reveal two new species of Ganoderma from Yunnan province, China

Article

Full-text available

Nov 2021

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.

Ganoderma (Ganodermataceae, Basidiomycota) Species from the Greater Mekong Subregion

Article

Full-text available

Sep 2021
J. Fungi

The cosmopolitan fungal genus Ganoderma is an important pathogen on arboreal plant hosts, particularly in tropical and temperate regions. It has long been used as a traditional medicine because of its medicinal properties and chemical constituents. In this study, Ganoderma collections were made in the Greater Mekong Subregion (GMS), encompassing tropical parts of Laos, Myanmar, Thailand, Vietnam, and temperate areas in Yunnan Province, China. The specimens used in this study are described based on micro-macro-characteristics and phylogenetic analysis of combined ITS, LSU, TEF1�, and RPB2 sequence data. In this comprehensive study, we report 22 Ganoderma species from the GMS, namely, G. adspersum, G. applanatum, G. australe, G. calidophilum, G. ellipsoideum, G. flexipes, G. gibbosum, G. heohnelianum, G. hochiminhense, G. leucocontextum, G. lucidum, G. multiplicatum, G. multipileum, G. myanmarense, G. orbiforme, G. philippii, G. resinaceum, G. sichuanense, G. sinense, G. subresinosum, G. williamsianum, and G. tsugae. Some of these species were reported in more than one country within the GMS. Of these 22 species, 12 were collected from Yunnan Province, China; three were collected from Laos; three species, two new records, and one new species were collected from Myanmar; 15 species and four new records were collected from Thailand, and one new species was collected from Vietnam. Comprehensive descriptions, color photographs of macro and micro-characteristics, the distribution of Ganoderma within the GMS, as well as a phylogenetic tree showing the placement of all reported Ganoderma from the GMS are provided.

High diversity of Ganoderma and Amauroderma (Ganodermataceae, Polyporales) in Hainan Island, China

Article

Full-text available

Sep 2018

The Ganoderma weberianum-resinaceum lineage: multilocus phylogenetic analysis and ... 95 The Ganoderma weberianum-resinaceum lineage: multilocus phylogenetic analysis and morphology confirm G. mexicanum and G. parvulum in the Neotropics

Article

Full-text available

Oct 2019

Many species of Ganoderma exhibit a high phenotypic plasticity. Hence, particularly among them, the morphological species concept remains difficult to apply, resulting in a currently confused taxonomy; as a consequence, the geographical distribution range of many species also remains very uncertain. One of the areas with a strong uncertainty, as far as morphological species concept is concerned, is the Neotropics. It is common that names of species described from other regions, mainly from northern temperate areas, have been applied to Neotropical species. The aim of the present study was to determine which species might lay behind the G. weberianum complex in the Neotropics, using morphological studies and phylogenetic inferences based on both single (ITS) and multilocus (ITS, rpb2, and tef1-α) sequences. The results indicated that G. weberianum sensu Steyaert, which is the usually accepted concept for this taxon, was absent from the Neotropics. In this area, G. weberianum sensu Steyaert encompassed at least two phylogenetic species, which are tentatively, for the time being, identified as belonging to G. mexicanum and G. parvulum. These two species could be distinguished morphologically, notably by the ornamentation or its absence on their chlamydospores. The results also showed that additional species from the Neotropics might still exist, including, e.g., G. perzonatum, but their circumscription remains uncertain until now because of the paucity of material available. Furthermore, it was found that the current concept of G. resinaceum embraced a complex of species. A peer-reviewed open-access journal MycoKeys Launched to accelerate biodiversity research RESEARCH ARTICLE Milay Cabarroi-Hernández et al. / MycoKeys 59: 95-131 (2019) 96

A global meta-analysis of ITS rDNA sequences from material belonging to the genus Ganoderma (Basidiomycota, Polyporales) including new data from selected taxa

Article

Full-text available

Dec 2020

Ganoderma P. Karst. is a cosmopolitan genus of white-rot fungi which comprises species with highly-prized pharmaceutical properties, valuable biotechnological applications and of significant phytopatho-logical interest. However, the status of the taxonomy within the genus is still highly controversial and ambiguous despite the progress made through molecular approaches. A metadata analysis of 3908 nuclear ribosomal internal transcribed spacer (ITS) rDNA sequences obtained from GenBank/ENA/DDBJ and UNITE was performed by targeting sequences annotated as Ganoderma, but also sequences from environmental samples and from material examined for the first time. Ganoderma taxa segregated into five main lineages (Clades A to E). Clade A corresponds to the core of laccate species and includes G. shanxiense and three major well-supported clusters: Cluster A.1 ('G. lucidum sensu lato') consists of taxa from Eurasia and North America, Cluster A.2 of material with worldwide occurrence including G. resinaceum and Cluster A.3 is composed of species originating from all continents except Europe and comprises G. lingzhi. Clade B includes G. applanatum and allied species with a Holarctic distribution. Clade C comprises taxa from Asia and Africa only. Clade D consists of laccate taxa with tropical/subtropical occurrence, while clade E harbours the highest number of non-laccate species with a cosmopolitan distribution. The 92 Ganoderma-associated names, initially used for sequences labelling, correspond to at least 80 taxa. Amongst them, 21 constitute putatively new phylospecies after our application of criteria relevant to the robustness/support of the terminal clades, intra-and interspecific genetic divergence and available biogeographic data. Moreover , several other groups or individual sequences seem to represent distinct taxonomic entities and merit further investigation. A particularly large number of the public sequences was revealed to be insufficiently and/or incorrectly identified, for example, 87% and 78% of entries labelled as G. australe and G. lucidum, respectively. In general, ITS demonstrated high efficacy in resolving relationships amongst most of the Ganoderma taxa; however, it was not equally useful at elucidating species barriers across the entire genus and such cases are outlined. Furthermore, we draw conclusions on biogeography by evaluating species occurrence on a global scale in conjunction with phylogenetic structure/patterns. The sequence variability assessed in ITS spacers could be further exploited for diagnostic purposes.

Notes on Some Basidiomycetes from the Orient

Article

Sep 2018

C. J. Humphrey

Type studies in the Polyporaceae 14. Species described by N. Patouillard, either alone or with other mycologists

Article

Jan 1983

Leif Ryvarden

A laccase Gl-LAC-4 purified from white-rot fungus Ganoderma lucidum had a strong ability to degrade and detoxify the alkylphenol pollutants 4-n-octylphenol and 2-phenylphenol

Article

Dec 2020
J HAZARD MATER

In this study, the ability of laccase Gl-LAC-4, purified from Ganoderma lucidum, to degrade and detoxify two representative alkylphenol pollutants, 4-n-octylphenol and 2-phenylphenol, was systematically studied. Gl-LAC-4 laccase had a very strong ability to degrade high concentrations of 4-n-octylphenol, 2-phenylphenol, and alkylphenol mixtures. The degradation speed of Gl-LAC-4 toward 2-phenylphenol was very fast. Gl-LAC-4 displayed strong tolerance for a variety of metal salts and organic solvents in the degradation of alkylphenols. Gl-LAC-4 showed strong tolerance for high concentrations of various metal salts, such as MgSO4, MnSO4, Na2SO4, CuSO4, ZnSO4, CdSO4, and K2SO4, in the degradation of 4-n-octylphenol and 2-phenylphenol. In the case of the same metal cation, the inhibitory effect of the metal salt with Cl⁻ as the anion on the degradation of 4-n-octylphenol and 2-phenylphenol by laccase was stronger than that of the metal salt with SO⁴²⁻ as the anion. An increase in the number of chloride ions caused a greater inhibitory effect on alkylphenol degradation by laccase. Gl-LAC-4 exhibited strong tolerance for glycerol, ethylene glycol, butanediol, propylene glycol, and organic solvent mixtures in the degradation of alkylphenols. Gl-LAC-4 treatment significantly reduced or eliminated the phytotoxicity of 4-n-octylphenol and 2-phenylphenol.

A Beauveria phylogeny inferred from nuclear ITS and EF1-α sequences: evidence for cryptic diversification and links to Cordyceps teleomorphs

Article

Mar 2005

Beauveria is a globally distributed genus of soil-borne entomopathogenic hyphomycetes of interest as a model system for the study of entomo-pathogenesis and the biological control of pest insects. Species recognition in Beauveria is difficult due to a lack of taxonomically informative morphology. This has impeded assessment of species diversity in this genus and investigation of their natural history. A gene-genealogical approach was used to investigate molecular phylogenetic diversity of Beauveria and several presumptively related Cordyceps species. Analyses were based on nuclear ribosomal internal transcribed spacer (ITS) and elongation factor 1-alpha (EF1-α) sequences for 86 exemplar isolates from diverse geographic origins, habitats and insect hosts. Phylogenetic trees were inferred using maximum parsimony and Bayesian likelihood methods. Six well supported clades within Beauveria, provisionally designated A–F, were resolved in the EF1-α and combined gene phylogenies. Beauveria bassiana, a ubiquitous species that is characterized morphologically by globose to subglobose conidia, was determined to be non-monophyletic and consists of two unrelated lineages, clades A and C. Clade A is globally distributed and includes the Asian teleomorph Cordyceps staphylinidaecola and its probable synonym C. bassiana. All isolates contained in Clade C are anamorphic and originate from Europe and North America. Clade B includes isolates of B. brongniartii, a Eurasian species complex characterized by ellipsoidal conidia. Clade D includes B. caledonica and B. vermiconia, which produce cylindrical and comma-shaped conidia, respectively. Clade E, from Asia, includes Beauveria anamorphs and a Cordyceps teleomorph that both produce ellipsoidal conidia. Clade F, the basal branch in the Beauveria phylogeny includes the South American species B. amorpha, which produces cylindrical conidia. Lineage diversity detected within clades A, B and C suggests that prevailing morphological species concepts underestimate species diversity within these groups. Continental endemism of lineages in B. bassiana s.l. (clades A and C) indicates that isolation by distance has been an important factor in the evolutionary diversification of these clades. Permutation tests indicate that host association is essentially random in both B. bassiana s.l. clades A and C, supporting past assumptions that this species is not host specific. In contrast, isolates in clades B and D occurred primarily on coleopteran hosts, although sampling in these clades was insufficient to assess host affliation at lower taxonomic ranks. The phylogenetic placement of Cordyceps staphylinidaecola/bassiana, and C. scarabaeicola within Beauveria corroborates prior reports of these anamorph-teleomorph connections. These results establish a phylogenetic framework for further taxonomic, phylogenetic and comparative biological investigations of Beauveria and their corresponding Cordyceps teleomorphs.

Species diversity, systematic revision and molecular phylogeny of Ganodermataceae (Polyporales, Basidiomycota) with an emphasis on Chinese collections

Abstract

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