Journal of

Fungi
Article
Ganoderma (Ganodermataceae, Basidiomycota) Species from
the Greater Mekong Subregion
Thatsanee Luangharn 1,2,3 , Samantha C. Karunarathna 1,2 , Arun Kumar Dutta 4 , Soumitra Paloi 5 ,
Itthayakorn Promputtha 6,7 , Kevin D. Hyde 1,3,8 , Jianchu Xu 1,2 and Peter E. Mortimer 1, *

1 Centre for Mountain Futures (CMF), Kunming Institute of Botany, Kunming 650201, China;
[email protected] (T.L.); [email protected] (S.C.K.); [email protected] (K.D.H.);
[email protected] (J.X.)
2 CIFOR-ICRAF, World Agroforestry Centre (ICRAF), Kunming 650201, China
3 Center of Excellence in Fungal Research, Mae Fah Luang University, Chiang Rai 57100, Thailand
4 Department of Botany, West Bengal State University, Barasat 700126, India; [email protected]
5 National Center for Genetic Engineering and Biotechnology (BIOTEC), National Science and Technology
Development Agency (NSTDA), 113 Thailand Science Park, Phahonuyothin Rd., Khlong Nueng,
Khlong Luang, Pathum Thani 12120, Thailand; [email protected]
6 Department of Biology, Faculty of Science, Chiang Mai University, Chiang Mai 50200, Thailand;
[email protected]
7 Research Center in Bioresources for Agriculture, Industry and Medicine, Chiang Mai University,
Chiang Mai 50200, Thailand
8 Institute of Plant Health, Zhongkai University of Agriculture and Engineering, Haizhu District,
Guangzhou 510225, China
* Correspondence: [email protected]






Abstract: The cosmopolitan fungal genus Ganoderma is an important pathogen on arboreal plant
Citation: Luangharn, T.; hosts, particularly in tropical and temperate regions. It has long been used as a traditional medicine
Karunarathna, S.C.; Dutta, A.K.; Paloi, because of its medicinal properties and chemical constituents. In this study, Ganoderma collections
S.; Promputtha, I.; Hyde, K.D.; Xu, J.; were made in the Greater Mekong Subregion (GMS), encompassing tropical parts of Laos, Myanmar,
Mortimer, P.E. Ganoderma Thailand, Vietnam, and temperate areas in Yunnan Province, China. The specimens used in this
(Ganodermataceae, Basidiomycota)
study are described based on micro-macro-characteristics and phylogenetic analysis of combined
Species from the Greater Mekong
ITS, LSU, TEF1α, and RPB2 sequence data. In this comprehensive study, we report 22 Ganoderma
Subregion. J. Fungi 2021, 7, 819.
species from the GMS, namely, G. adspersum, G. applanatum, G. australe, G. calidophilum, G. ellip-
https://doi.org/10.3390/jof7100819
soideum, G. flexipes, G. gibbosum, G. heohnelianum, G. hochiminhense, G. leucocontextum, G. lucidum,
Academic Editor: Kuang R. Chung
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
Received: 29 July 2021 more than one country within the GMS. Of these 22 species, 12 were collected from Yunnan Province,
Accepted: 22 September 2021 China; three were collected from Laos; three species, two new records, and one new species were
Published: 30 September 2021 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-
Publisher’s Note: MDPI stays neutral and micro-characteristics, the distribution of Ganoderma within the GMS, as well as a phylogenetic
with regard to jurisdictional claims in tree showing the placement of all reported Ganoderma from the GMS are provided.
published maps and institutional affil-
iations. Keywords: two new species; biogeography; ecological aspects; Lingzhi; medicinal mushroom;
morphology

Copyright: © 2021 by the authors.

Licensee MDPI, Basel, Switzerland. 1. Introduction
This article is an open access article
Ganoderma was established by Karsten [1] with Polyporus lucidus (Curtis) Fr. (=Gan-
distributed under the terms and
oderma lucidum (Curtis) P. Karst.) as the type species [2]. The genus has a worldwide
conditions of the Creative Commons
distribution but is predominantly found in tropical and temperate regions, including
Attribution (CC BY) license (https://
creativecommons.org/licenses/by/
Africa, America, Europe, and Asia [3–6]. Most members of Ganoderma are pathogenic
4.0/).
in nature, inflicting various diseases to plants such as white rot and stem rot as well as

J. Fungi 2021, 7, 819. https://doi.org/10.3390/jof7100819 https://www.mdpi.com/journal/jof

J. Fungi 2021, 7, 819 2 of 83

wood decay [3,6–9]. The genus is circumscribed by sessile to stipitate basidiomata with
double-walled basidiospores and interwall pillars [1,2]. Kirk et al. [10] mentioned that
globally there are 80 species of Ganoderma, while two global fungal databases viz. Index
Fungorum [11] and MycoBank [12] hold 459 and 503 records, respectively.
In light of an earlier taxonomic framework, Ganoderma included three subgenera
viz. Ganoderma P. Karst., Elfvingia P. Karst., and Trachyderma Imazeki [13]. The subgenus
Ganoderma includes two sections. Ganoderma and Phaenema contain laccate species (a cutis
surface consisting of a palisade of inflated hyphal ends) with a shiny upper surface, whereas
all non-laccate species (palisade is absent) with a dull upper surface are under the subgenus
Elfvingia [14,15]. The subgenus Trachyderma has been deemed illegitimate and is treated as
a synonym of Ganoderma due to the presence of the lichenized genus Trachyderma [15].
Ganoderma mushrooms have long been used as traditional medicine in many Asian
countries [4]. In the Pharmacopoeia of the People’s Republic of China, Ganoderma species
are recorded as being used over two millennia [16]. Traditional Chinese books proposed
and classified Ganoderma species based on basidiocarp coloration [17]. Ganoderma species
are suitable sources of natural bioactive compounds of high and low molecular weight,
especially polysaccharides, protein, sterols, and triterpenoids [18–20]. These compounds
are known to possess extensive therapeutic properties, such as antibacterial, antifun-
gal, antiviral, anticancer, antitumor, anti-inflammatory, anti-hypotensive, and antiox-
idative agents [15,21–25]. Moreover, these compounds also treat many immunological
diseases [15,22], making Ganoderma species a popular functional food [26].
The Greater Mekong Subregion (GMS) consists of Cambodia, Lao People’s Demo-
cratic Republic, Myanmar, Yunnan Province of the People’s Republic of China, Thailand,
and Vietnam. The GMS is a biodiversity hotspot with rich fungal diversity [27], as well
as featuring a high diversity of endemic and endangered organisms. A variable climate,
vegetation type, and habitat primarily characterize the GMS, supporting the existence of
thousands of macrofungal species, many of which remain unknown [28–31]. Wild edible
and medicinal macrofungi comprise an important source of income for people residing in
the GMS. Past work has shown that the GMS is a hotpot of Ganoderma diversity, and thus
of interest to the broader community of scientists working on this genus, yet no studies
have attempted to bring all this information into one space and update the knowledge
the known distribution of species, as well as introduce newly discovered species. Thus,
our study seeks to clarify the status of Ganoderma in the GMS, where it is both an important
medicinal and plant pathogenic mushroom.
There is much debate among taxonomists regarding how to best resolve nomenclatural
confusion within Ganoderma. This stems from the high degree of phenotypic plasticity
found in Ganoderma species [4,32–36]. Ganoderma lucidum was broadly defined from the
original concept [34], with variations occurring in macro-characteristics of the basidiomes,
resulting in many synonyms and general redundancy in the taxonomy of this species and
genus [34,37].

1.1. Biogeography of Ganoderma

Ganoderma lucidum (Curtis) P. Karst., the type species of Ganoderma, was described
based on a specimen collected from the U.K. Moncalvo et al. [38] concluded that G. lucidum
was distributed across northern and southern Europe and likely extended into China.
The name Ganoderma lingzhi (Lingzhi) has been adopted in place of G. lucidum for specimens
native to East Asia [4]. Among Ganoderma species, several species similar to G. lucidum have
also been described worldwide: G. multipileum [39], G. Sichuanense [4,16,40] from China;
G. resinaceum from Europe [41]; and G. oregonense, G. sessile, G. tsugae, and G. zonatum from
the U.S. [42,43]. These species are accepted as members of the G. lucidum complex.
Taxonomic treatments of Ganoderma species remain unclarified due to the high de-
gree of phenotypic plasticity [44,45]. Some Ganoderma species are distributed worldwide,
such as G. applanatum and G. australe, while others are known only from specific locali-
ties [2,46]. Limited holotypes and unsettled geographical distributions have resulted in
 China; G. resinaceum from Europe [41]; and G. oregonense, G. sessile, G. tsugae, and G. zona‐
tum from the U.S. [42,43]. These species are accepted as members of the G. lucidum com‐
plex.
Taxonomic treatments of Ganoderma species remain unclarified due to the high de‐
J. Fungi 2021, 7, 819 gree of phenotypic plasticity [44,45]. Some Ganoderma species are distributed worldwide, 3 of 83
such as G. applanatum and G. australe, while others are known only from specific localities
[2,46]. Limited holotypes and unsettled geographical distributions have resulted in much
ambiguity among Ganoderma taxa. Ganoderma has been reported from tropical Laos, My‐
much ambiguity
anmar, Thailand,among Ganoderma
and Vietnam taxa.
[30,31] andGanoderma
temperate has been[6,31]
China reported from
in the tropical
GMS. Laos,
Members
Myanmar, Thailand, and Vietnam [30,31] and temperate China [6,31] in the
of Ganoderma can be of significant importance in horticulture since they infect landscape GMS. Members
of Ganoderma
plants, can perennial
fruit trees, be of significant importance
crops, and in horticulture
economically importantsince
treesthey infect
[27,29], landscape
which can be
plants, fruit trees, perennial crops, and economically important trees [27,29],
resulted in the death of affected trees. Ganoderma has been recorded mostly in a wide range which can
be resulted in the death of affected trees. Ganoderma has been recorded mostly
of tree species, especially on hardwood trees [8]. Ganoderma has not been reported as host‐ in a wide
range of tree
specific. Theyspecies, especially
are often found on hardwood
in natural forests [8]. Ganoderma
trees(deciduous has not
forests) andbeen reportedon
dominated as
host-specific. They are often found in natural forests (deciduous forests)
plantations trees [30,31]. Although Ganoderma has long been regarded as one of the genera and dominated
on plantations
with a worldwide treesdistribution [30], theGanoderma
[30,31]. Although diversity andhas long been regarded
composition as one in
of Ganoderma of the
the
genera with a worldwide distribution [30], the diversity and composition
GMS remains poorly understood. Here, we summarize and update the Ganoderma species of Ganoderma
in the GMS remains
distribution in the GMSpoorly understood.
based on locationHere,
datawefrom
summarize
known and the Ganoderma
update(Figure
collections 1). The
species distribution in the GMS based on location data from known collections (Figure 1).
extant Ganoderma species in the GMS are compiled in Table S1, and species diversity (SD)
The extant Ganoderma species in the GMS are compiled in Table S1, and species diversity
hotspots of Ganoderma in GMS are shown in Figure 2.
(SD) hotspots of Ganoderma in GMS are shown in Figure 2.

Figure 1. Distribution of Ganoderma species in the GMS.

gi 2021, 7, x FOR PEER REVIEW
J. Fungi 2021, 7, 819 4 of 83

2. Species diversity (SD) hotspots of Ganoderma species in the GMS.

Figure Figure
2. Species diversity (SD) hotspots of Ganoderma species in the GMS.
1.2. Ecological Aspects of Ganoderma
1.2. Ecological Aspects
Wu and of suggested
Dai [47] Ganoderma that the morphology of Ganoderma species varies based
on factors such as climate, nutrition, vegetation, and geography; moreover, morphology
Wu and Dai [47] suggested that the morphology of Ganoderma species varies b
is not associated with the genetic material of a particular species. Basidiome features are
on factors suchbyasthe
influenced climate, nutrition,
interaction vegetation,
of both intrinsic (geneticand
andgeography;
physiological)moreover,
and extrinsicmorph
is not (environmental)
associated with the[48–51].
factors genetic material
Therefore, of a particular
comprehensive species.ofBasidiome
documentation species ecol- feature
ogy is necessary when studying Ganoderma. Criteria for identification
influenced by the interaction of both intrinsic (genetic and physiological) vary: some authorsand ext
strictly focus on geographical distribution, host-specificity, and macro-characteristics of
(environmental)
basidiomes, whilefactors
others[48–51].
focus onTherefore, comprehensive
spore morphology as the primarydocumentation of species
taxonomic characteris-
ogy isticnecessary when
[52,53]. In the studying
present Ganoderma.
study, ecological factorsCriteria for identification
such as collection period, season,vary: some au
climate,
host plant, and substrate of Ganoderma species were recorded.
strictly focus on geographical distribution, host‐specificity, and macro‐characterist
Ganoderma species are known as both important wood-decaying fungi and pathogens
basidiomes, while others focus on spore morphology as the primary taxonomic char
that can survive on a wide range of hosts [54]. These fungi decay lignin, cellulose,
istic [52,53]. In the present
and hemicellulose, resultingstudy, ecological
in a severe factors
loss of woody such
plant as collection
strength [9,55]. Theyperiod,
possess seaso
mate, host plant, and substrate of Ganoderma species were recorded.
lignocellulose-decomposing enzymes useful for bioenergy production and bioremedia-
tion [56]. Diseases
Ganoderma species caused
are by Ganoderma
known species
as both result in lower
important yields in economically
wood-decaying fungi and p
important trees [54].
gens that In can survive on a wide range of hosts [54]. These fungi decay lignin, cellu
this paper, we update biogeography records of Ganoderma by reporting species
and hemicellulose,
distributed acrossresulting
the Greaterin a severe
Mekong loss of
Subregion woody
(GMS) plant
as well strengthmorpholog-
as conducting [9,55]. They po
lignocellulose‐decomposing enzymes useful for bioenergy productionYunnan
ical and phylogenetic studies on Ganoderma collected from temperate regions in and biorem
Province, China, and tropical regions of Laos, Myanmar, Thailand, and Vietnam (Table 1).
tion [56]. Diseases caused by Ganoderma species result in lower yields in economicall
portant trees [54].
In this paper, we update biogeography records of Ganoderma by reporting sp
distributed across the Greater Mekong Subregion (GMS) as well as conducting mo
logical and phylogenetic studies on Ganoderma collected from temperate regions in
J. Fungi 2021, 7, 819 5 of 83

Table 1. Species of Ganoderma, host plants, diseases, and main regions of distribution within the Greater Mekong Subregion.

Ganoderma Species Host Plant Diseases Region Reference

Pterocarpus sp. Wood decay Thailand This study
G. adspersum
Mangifera indica Wood decay Laos This study
Acer sp. Vietnam [57]
G. applanatum Rhizophora apiculata, Machilus Wood decay, Thailand,
[30,58], this study
yunnanensis, and hardwoods Butt rot China, Myanmar, Laos
Machilus yunnanensis Wood decay Yunnan, China This study
G. applanatum Artocarpus sp., and Dipterocarpus sp. Wood decay Thailand This study
Laos, Thailand,
Anisoptera costata and Shorea robtusa Wood decay, rotten wood [30,59], this study
and Myanmar
G. australe
Camellia sinensis var. assamica Root rot Thailand [60]
Castanopsis sp. and Machilus
G. calidophilum Wood decay Yunnan, China This study
yunnanensis
G. casuarinicola Pinus kesiya White rot Thailand [31], this study
G. ellipsoideum Unknown Wood decay Thailand This study
China, Myanmar,
G. flexipes Pinus sp. and unknown [6,30,61], this study
Vietnam
Albizia mollis, Machilus yunnanensis,
Yunnan, China This study
and Pinus sp.
Albizia lebbeck, Dendrocalamus strictus,
Wood decay, rotten
G. gibbosum Dipterocarpus sp., Mangifera indica, Thailand This study
wood
P. pterocarpum, and Pinus sp.
Hardwoods Wood decay China, Thailand [30], this study
G. hochimin-hensis Areca sp. Root rot Vietnam This study
G. hoehnelianum Unknown Wood decay Myanmar This study
G. leucocontextum Cyclobalanopsis glauca and C. glauca Wood decay, root rot Yunnan, China [62], this study
Quercus sp. Wood decay Yunnan, China This study
G. lucidum Acacia sp., Dendrocalamus strictus,
Wood decay Thailand This study
and Pterocarpus sp.
Unknown decayed hardwood Wood decay Yunnan, China [30], this study
G. multipileum
Pinus merkusii Wood decay Thailand This study
G. multiplicatum Quercus sp. Wood decay Yunnan, China This study
G. myanmarense Unknown tree Wood decay Myanmar This study
G. nasalaense Not mention Wood decay Laos [36]
G. neojaponicum Near hardwood roots Wood decay Myanmar [30]
G. orbiforme Albizia mollis and Indochinese sp. Wood decay Thailand This study
Unknown decayed hardwood Wood decay China, Laos This study
Elaeis guineensis Basal stem rot Thailand [63]
G. philippii Unknown Root rot Thailand This study
G. resinaceum Albizia mollis Basal stem rot Yunnan, China This study
G. sichuanense Quercus sp. China [4,40]
Pterocarpus sp., Mangifera indica,
Sesbania grandiflora, and Peltophorum Wood decay Thailand This study
pterocarpum
Castanopsis sp., Castanea sp.,
Wood decay,
Cyclobalanopsis sp, and Graucoides Yunnan, China [4], this study
root rot
schotky
Albizia mollis and Quercus sp. Rotten wood Yunnan, China [30], this study
G. sinense Dendrocalamus strictus and Wood decay, Rotten
Thailand This study
Dipterocarpus sp. wood
J. Fungi 2021, 7, 819 6 of 83

Table 1. Cont.

Ganoderma Species Host Plant Diseases Region Reference

Peltophorum pterocarpum and
G. subresinosum Wood decay Thailand This study
Castanopsis sp.
G. thailandicum Pinus merkusii Wood decay Thailand [31], this study
G. tropicum Dipterocarpus sp. Wood decay Thailand [31], this study
G. tsugae Larix sp., Picea sp. and Tsuga sp. Butt rot, Wood decay China [57,64], this study
G. williamsianum Unknown Wood decay Myanmar This study

2. Materials and Methods

2.1. Study Sites
Ganoderma specimens were collected from the temperate parts of Yunnan Province,
China, and tropical parts of Laos, Myanmar, Thailand, and Vietnam (Figure S1). Detailed
collection site information, such as location, climate, the monthly temperature during the
rainy season, host trees species, and native forest type, are provided in Table 2.

Table 2. Regional information for collection sites in the Greater Mekong Subregion, where collections occurred.

Study Sites China Myanmar Laos Thailand Vietnam

Climate Type Temperate Tropical Tropical Tropical Tropical
December 2016;
September– October–December 2017;
Collecting date December 2017; July 2019 June–July 2018 June June 2019
September– and November 2018
November 2018
Monthly
16–22 ◦ C 26–30 ◦ C 25–32 ◦ C 25–38 ◦ C 28–32 ◦ C
temperature
Coniferous forest, Coniferous forest,
Mixed deciduous Deciduous forest
dry evergreen forest, deciduous forest,
Forest type forest and tropical and tropical rain Deciduous forest
and evergreen coniferous dry evergreen forest,
rain forest. forest
forest and tropical rain forest.
Acacia sp.,
Albizia lebbeck, Artocarpus
Albizia mollis, spp., Castanopsis spp.,
Castanopsis spp., Dendrocalamus strictus,
Fagus spp., Machilus Dipterocarpus spp., Garuga
Unidentified tree
Host tree species yunnanensis, Pinus spp., Mangifera indica pinnata, Indochinese spp., Areca spp.
species
Pterocarpus macrocarpus, Maerus siamensis,
Quercus spp., and Hevea Mangifera indica,
brasiliensis Pterocarpus macrocarpus,
Peltophorum pterocarpum,
and Pinus spp.

2.2. Sample Collection and Isolation

Fresh basidiomes of Ganoderma species were collected from China, Laos, Myanmar,
Thailand, and Vietnam. The samples were photographed and transported back to the
laboratory, where fresh macroscopic details were described. The cultures were aseptically
isolated by using heat sterilized forceps, transferring sections of internal tissue from fruiting
bodies onto potato dextrose agar (PDA) medium and incubated at 25–30 ◦ C, for 1–3 weeks,
under dark conditions [59]. After incubation, the agar surface was fully covered with white
mycelium. The pure stock culture was then covered with mineral oil and deposited in the
voucher culture collection of the Mae Fah Luang University culture collection (MFLUCC)
Chiang Rai, Thailand. The cultures are being maintained at 4 ◦ C for further studies.
The voucher samples were then air-dried at 40 ◦ C for 48 h until they were completely
dehydrated. Finally, the material was deposited in the herbarium of Mae Fah Luang
University (MFLU Herb.) with duplicates in the herbarium of Kunming Institute of Botany,
Academia Sinica (HKAS), Yunnan Province, China.
J. Fungi 2021, 7, 819 7 of 83

2.3. Morphological Study

Macro-characteristics were described following the method of Lodge et al. [65], while
color notations were recorded following Ridgeway [66]. Macroscopic characteristics were
determined according to the methodology described by Largent [67]. To observe mi-
croscopic characteristics, free-hand sections were made under a dissecting microscope
(OLYMPUS SZ61) and mounted on a glass slide containing 3–5% KOH, 1–3% Congo red,
and Melzer’s reagent for highlighting all tissues [68]. Microphotography was performed
with a Nikon ECLIPSE Ni (Nikon, Tokyo, Japan) compound microscope, with a Canon
EOS 600D (Tokyo, Japan) digital camera fitted on the top of the microscope. Basidiospores
and hyphal system sizes, colors, and shapes were recorded and photographed. Basid-
iospore measurements were taken using the Tarosoft® Image Framework program v. 0.9.0.7.
For measuring basidiospore statistics, the Tulloss’ standard format [69] was followed [Q
= L/W], where Q, the quotient of basidiospore length to width (L/W) in side view and
Qm, the mean of Q-values ± SD. The calculation was performed by measuring at least
50 basidiospores from each basidiomata [70]. Photographs were edited in Adobe Illustrator
CS v. 3.0.

2.4. DNA Extraction, PCR Amplification, and Sequencing

Dried internal tissues of the basidiomes were grounded, and total DNA was extracted
using the Biospin Fungus Genomic DNA Extraction Kit (BioFlux® ). The ITS, LSU, RPB2,
and TEF1α genes were amplified by polymerase chain reaction (PCR). The PCR ampli-
fications were performed in a total volume of 25 µL of PCR mixtures containing 9.5 µL
ddH2 O, 12.5 µL of PCR master mix, 1 µL of DNA template, and 1 µL of each primer
(10 µM). PCR amplification was carried out using primer pairs ITS5/ITS4 for internal
transcribed spacer rDNA region (ITS1, 5.8S rDNA and ITS2), LROR/LR5 for the nuclear
ribosomal large subunit 28S rDNA (LSU), fRPB2-5F/fRPB2-7cR for the partial RNA poly-
merase second largest subunit region (RPB2) [71–73], and EF1-983F/EF1-2218R for the
partial translation elongation factor 1-alpha (TEF1α) [74]. The PCR cycling amplification
conditions differed following amplified markers. For ITS and LSU, the following condition
was used: 3 min at 94 ◦ C, followed by 35 cycles of 95 ◦ C for 30 s, 55 ◦ C for 1 min, 72 ◦ C
for 1 min, followed by a final extension at 72 ◦ C for 10 min. The amplification condition
for TEF1α consisted of initial denaturation at 5.30 min at 95 ◦ C, followed by 35 cycles of
94 ◦ C for 1 min, 57 ◦ C for 30 s and 72 ◦ C for 1.30 min, followed by a final extension at 72 ◦ C
for 10 min. The cycling profile of 3 min at 94 ◦ C followed by 35 cycles of 95 ◦ C for 1 min,
52 ◦ C for 2 min and 72 ◦ C for 1 min, followed by a final extension at 72 ◦ C for 10 min,
was used for RPB2. The sequencing of PCR products was carried out by Sangon Biotech
(Shanghai) Co., Ltd., Shanghai, China. The nuclear ribosomal internal transcribed spacer
region (nrITS) of the fungi was amplified, and the sequence was deposited in GenBank to
obtain the accession number.

2.5. Sequence Alignment and Phylogenetic Analyses

Ganoderma specimens’ sequences were subjected to standard BLAST searches of Gen-
Bank to determine the primary identity of the Ganoderma species. All the other sequences
of taxa closely related to our Ganoderma species were retrieved from GenBank. Sequences
with high similarity indices were determined from BLAST searches to find the closest
matches with taxa and from recently published literature [31,37,75]. All sequences used to
construct the phylogenetic tree are listed in Table 3. Sanguinoderma rugosum (Blume and
T. Nees) Torrend (Cui 9011), and Tomophagus colossus (TC-02) were used as the outgroup
taxa [62,76]. Sequences were aligned with MAFFT online server [77] and manually adjusted
using Bioedit v. 7.2.5 [78] and Clustal X softwares [79]. Alignments were checked manually
and optimized to allow maximum sequence similarity. Gaps were treated as missing data.
Maximum parsimony (MP) analysis was performed using the PAUP beta 10 software
version 4.0 [80].
J. Fungi 2021, 7, 819 8 of 83

Clades inferred from the MP analyses were further assessed by maximum likeli-
hood (ML) bootstraps with 1000 replicates using random step-wise sequence additions,
performed using RAxML-HPC2 on XSEDE v. 8.2.8 [81] on the CIPRES webportal [82],
and carried out using raxmlGUI v. 1.3.1 [83]. The best-fitting substitution model for
each single gene partition and the concatenated data set was determined in MrModeltest
2.3 [84]. Bayesian posterior probabilities (PP) with the GTR+I+G model were used for
each partition. Bayesian Markov chain Monte Carlo (MCMC) analyses were conducted
in MrBayes v. 3.2.2 [85]. The number of generations was set at 3,500,000, with trees being
sampled every 100th generations (a total of 35,000 trees), resulting in an average standard
deviation of split frequencies below 0.01. Based on the tracer analysis, the first sampled
topologies of 8,750 trees representing 25% of total trees were discarded in the burn-in phase.
The remaining 26,250 trees were used for calculating posterior probability (PP) values in
the majority rule consensus tree.
Phylogenetic trees were visualized with FigTree v. 1.4.0 [86] and edited using Microsoft
Office PowerPoint 2010 before being exported to Adobe Illustrator CS v. 3.0. Maximum
likelihood (ML) and maximum parsimony (MP) bootstrap values equal to or greater than
70% and Bayesian posterior probabilities (BP) equal to or greater than 0.95 are presented
above the branches.

Table 3. GenBank accession numbers for ITS, LSU, TEF1α, and RPB2 sequence data of the taxa used in this study and
procured from GenBank based on the earlier studies for conducting phylogenetic analyses. Details of newly amplified
sequences are represented in bold.

Fungal GenBank Accession no.

Voucher Locality References
Species ITS LSU RPB2 TEF1α
G. adspersum GACP15061220 Thailand MK345425 – MK371437 MK371431 [30]
G. adspersum MFLU 19-2177 Laos MN396652 – MN423113 – This study
G. adspersum MFLU 19-2178 Thailand MN396653 – MN423114 MN423149 This study
G. adspersum MFLU 19-2220 Thailand MN396655 MN428663 MN423116 MN423151 This study
Cui 13817
G. angustisporum Fujian, China MG279170 – MG367507 MG367563 [37]
(holotype)
Guangdong,
G. angustisporum Cui 14578 MG279171 – – MG367564 [37]
China
G. applanatum FIN131R610 – EF060004 – – – GenBank
G. applanatum MFLU 19-2175 Thailand MN396333 – – – This study
G. applanatum MFLU 19-2188 China MN396332 – – – This study
Dai 12588 Durban, South
G. aridicola KU572491 – – KU572502 [87]
(holotype) Africa
Hainan Island,
G. australe GACP14081671 MH106871 – – – [88]
China
G. australe GACP14061914 China MK345428 – – MK371432 [30]
G. australe MFLU 13-0534 Thailand KP142173 – – MN423152 [59]
G. australe HKAS 97397 China MN396656 MN428664 – – This study
G. australe MFLU 19-2171 Lao MN396657 – – – This study
G. austroafricanum CBS138724 South Africa KM507324 KM507325 – – [9]
G. boninense WD 2028 Japan KJ143905 KU220015 KJ143964 KJ143924 [76]
G. boninense WD 2085 Japan KJ143906 – KJ143965 KJ143925 [76]
G. boninense GbHap1 – MK713555 – – – GenBank
G. boninense GB001 – KX092000 – – – GenBank
Yunnan,
G. calidophilum MFLU 19-2174 MN398337 – – – This study
China
DMC 322
G. carocalcareum Cameroon EU089969 – – – [46]
(holotype)
G. carocalcareum DMC 513 Cameroon EU089970 – – – [46]
Dai 16336 Guangdong,
G. casuarinicola MG279173 – MG367508 MG367565 [37]
(holotype) China
G. casuarinicola HKAS104639 Thailand MK817650 MK817654 MK840868 MK871328 [31]
G. chocoense QCAM3123 Ecuador MH890527 – – – [89]
G. curtisii CBS 100132 USA JQ781849 – KJ143967 KJ143927 [76]
G. destructans CMW43670 South Africa KR183856 KR183860 – – [9]
G. destructans CMW42146 South Africa MG020245 – – MG020200 [9]
G. ecuadorense ASL799 Ecuador KU128524 KX228350 – – [90]
G. ecuadorense PMC126 Ecuador KU128525 KU128529 – – [90]
J. Fungi 2021, 7, 819 9 of 83

Table 3. Cont.

Fungal GenBank Accession no.

Voucher Locality References
Species ITS LSU RPB2 TEF1α
GACP14080966
G. ellipsoideum Hainan, China MH106867 – – – [88]
(holotype)
G. ellipsoideum GACP14080968 Hainan, China MH106868 – – – [88]
G. ellipsoideum MFLU 19-2221 Thailand MN398339 – – MN423157 This study
G. enigmaticum Dai 15970 Africa KU572486 – MG367513 KU572496 [91]
G. enigmaticum Dai 15971 Africa KU572487 – MG367514 KU572497 [91]
G. flexipes Wei 5494 China JN383979 – – – [6]
Yunnan,
G. flexipes MFLU 19-2198 MN398340 MN428665 – – This study
China
SFC20150630-
G. gibbosum Korea KY364264 – – – [92]
23
Yunnan,
G. gibbosum HKAS 97411 MN398341 – – This study
China
G. gibbosum MFLU 19-2176 Thailand MN396311 – MN423118 – This study
G. gibbosum MFLU 19-2190 Laos MN396310 – MN423117 – This study
MFLU 19-2224
G. hochiminhense Vietnam MN398324 MN396390 – MN423176 This study
(holotype)
G. hochiminhense MFLU 19-2225 Vietnam MN396662 MN396391 – MN423177 This study
G. hoehnelianum Dai11995 Yunnan, China KU219988 – MG367497 MG367550 [91]
G. hoehnelianum MFLU 19-2168 Myanmar MN396316 – MN423123 MN423158 This study
G. leucocontextum Dai 15601 China KU572485 – MG367516 KU572495 [37]
Yunnan,
G. leucocontextum HKAS 97401 MN396317 MN428670 MN423124 – This study
China
Wu 1006-38
G. lingzhi Hubei, China JQ781858 – JX029980 JX029976 [4]
(holotype)
G. lobatum JV1212/10J USA KF605676 – – KU572501 GenBank
G. lucidum Rivoire 4195 France KJ143909 – KJ143969 – [76]
G. lucidum Cui 14404 Sichuan, China MG279181 – MG367519 MG367573 [37]
Yunnan,
G. lucidum MFLU 19-2161 MN396338 – MN423135 MN423168 This study
China
G. lucidum MFLU 19-2162 Thailand MN396341 – MN423138 – This study
LIP
Martinica,
G. martinicense SWMart08-55 KF963256 – – – [93]
France
(holotype)
UMN7-3GHA
G. mbrekobenum Ghana KX000896 KX000897 – – [90]
(holotype)
UMN-MZ4
G. mizoramense India KY643750 – – – [94]
(holotype)
G. multiplicatum Dai 13122 China KU572488 – – KU572498 [95]
Yunnan,
G. multiplicatum MFLU 19-2152 MN401405 – – MN423171 This study
China
Taiwan PRC,
G. multipileum CWN 04670 KJ143913 – KJ143972 KJ143931 [87]
China
G. multipileum MFLU 19-2166 Thailand MN401406 – MN423142 MN423172 This study
G. mutabile Yuan 2289 Yunnan, China JN383977 – – – [76]
MFLU 19-2167
G. myanmarense Myanmar MN396330 MN428672 – – This study
(holotype)
G. myanmarense MFLU 19-2169 Myanmar – MN398325 – – This study
GACP17060211
G. nasalaense Laos MK345441 MK346831 – – [30]
(holotype)
G. neojaponicum ASI 7032 Korea JQ520193 – – – [96]
G. orbiforme Cui 13918 Hainan, China MG279186 – MG367522 MG367576 [37]
G. orbiforme GACP14061420 Laos MK345447 MK346833 – – [30]
G. orbiforme MFLU 17-1933 Thailand MN401408 – MN423144 – This study
G. oregonense CBS 265.88 USA JQ781875 – KJ143974 KJ143933 [76]
G. philippii E7098 Malaysia AJ536662 – – – [97]
G. philippii E7425 Malaysia AJ608713 – – – [97]
G. philippii MFLU 19-2222 Thailand MN401410 MN398326 – MN423174 This study
G. philippii MFLU 19-2223 Thailand MN401411 MN398327 – MN423175 This study
QCAM6422
G. podocarpense Ecuador MF796661 MF796660 – – [94]
(holotype)
G. resinaceum HMAS86599 England AY884177 – JF915435 – GenBank
G. resinaceum CBS 194.76 Netherlands KJ143916 – – KJ143934 [76]
Yunnan,
G. resinaceum MFLU 19-2153 MN398315 MN398328 – – This study
China
J. Fungi 2021, 7, 819 10 of 83

Table 3. Cont.

Fungal GenBank Accession no.

Voucher Locality References
Species ITS LSU RPB2 TEF1α
HKAS 58053 Cameroon,
G. ryvardenii HM138671 – – – [98]
(holotype) Africa
Cameroon,
G. ryvardenii HKAS 58054 HM138672 – – – [98]
Africa
GACP18012501
G. sanduense China MK345450 – – – [30]
(holotype)
G. sanduense GACP18012502 China MK345451 – – – [30]
G. sessile JV 1209/9 USA KF605629 – – KJ143936 [76]
G. sessile JV 1209/27 USA KF605630 – KJ143976 KJ143937 [76]
Shandong,
G. shandongense Dai 15785 MG279190 – MG367526 MG367580 [37]
China
Shandong,
G. shandongense Dai 15791 MG279192 – MG367528 MG367582 [37]
China
HMAS 42798
G. sichuanense Sichuan, China JQ781877 – – – [4]
(holotype)
G. sichuanense MFLU 19-2164 Thailand MN396324 – MN423130 MN423163 This study
Yunnan,
G. sichuanense HKAS 97398 MN396319 – MN423126 MN423159 This study
China
G. sichuanense CGMCC5.2175 Sichuan, China KC662402 – – – [99]
G. sinense Wei 5327 Hainan, China KF494998 KF495008 MG367529 KF494976 [37]
G. sinense MFLU 19-2172 Thailand MN398319 MN398332 MN423146 – This study
Yunnan,
G. sinense MLFU 19-2173 MN398316 MN398329 – – This study
China
G. steyaertianum MEL:2382783 Australia KP012964 – – – GenBank
G. steyaertianum 6 WN 20B Indonesia KJ654462 – – – [100]
7-SU-3-C-70
G. subresinosum Indonesia KJ654472 – – – GenBank
(M)-B
G. subresinosum MFLU 17-1912 Thailand MN398321 – – – This study
HKAS104640
G. thailandicum Thailand MK848681 MK849879 MK875831 MK875829 [31]
(holotype)
G. thailandicum HKAS104641 Thailand MK848682 MK849880 MK875832 MK875830 [31]
G. tropicum Yuan 3490 Yunnan, China JQ781880 – – KJ143938 [4]
G. tropicum Dai 16434 Hainan, China MG279194 – MG367532 MG367585 [37]
G. tropicum HKAS 97486 Thailand MH823539 MH823540 MH883621 – [75]
G. tsugae Dai12751b USA KJ143919 – KJ143977 KJ143939 [76]
Yunnan,
G. tsugae HKAS 97406 MG279195 – MG367533 MG367586 This study
China
G. valesiacum CBS428.84 USA JQ520218 – – – [96]
G. weberianum CBS219.36 Philippines JQ520219 – – – [96]
G. weberianum GanoTK17 Cameroon JN105705 – – – GenBank
UMN-20-GHA
G. wiiroense Ghana KT952361 – – – [101]
(para type)
G. williamsianum Dai 16809 Thailand MG279183 – MG367535 MG367588 [101]
G. williamsianum MFLU 19-2170 Myanmar MN398323 MN398334 – – This study
G. wuzhishanense GACP14081689 Hainan, China KU994772 – – – [76,102]
G. zonatum FL-02 USA KJ143921 – KJ143979 KJ143941 [76]
Sanguinoderma Guangdong,
Cui 9011 KJ531664 – MG367506 KU572504 [79]
rugosum China
Tomophagus colossus TC-02 Vietnam KJ143923 – – KJ143943 [76]

3. Results
In this study, we morphologically and phylogenetically analyze and report 22 Gan-
oderma species from the GMS, namely, G. adspersum, G. applanatum, G. australe, G. cali-
dophilum, G. ellipsoideum, G. flexipes, G. gibbosum, G. heohnelianum, G. hochiminhense, G. leu-
cocontextum, 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. Of these 22 species, 12 were from Yunnan Province, China; three were
from Laos; three species, two new records, and one new species were from Myanmar;
15 species and four new records were from Thailand; and one new species was from Viet-
nam. The phylogenetic and morphological analyses results of the 22 Ganoderma species
are detailed below.
J. Fungi 2021, 7, 819 11 of 83

3.1. Phylogenetic Analyses

Phylogenetic analyses were inferred from the combined data set of ITS, LSU, RPB2,
and TEF1α sequences of 114 taxa, of which 112 taxa belong to the genus Ganoderma, and the
remaining 2, Sanguinoderma rugosum Blume and T. Nees (Cui 9011) and Tomophagus colossus
(Fr.) Murrill, Torreya (TC-02) are the outgroup taxa. The data set comprised 3524 charac-
ters with gaps (637 characters for ITS, 866 characters for LSU, 1016 characters for RPB2,
and 1005 characters for TEF1α), of which 2459 characters were constant, 855 characters were
variable and parsimony-informative, and 210 characters were parsimony-uninformative.
Tree topologies resulted from the ML analysis were similar to that of the MP
J. Fungi 2021, 7, x FOR PEER REVIEW 12 and
of 91 Bayesian
analysis. Hence, the best-scoring ML tree is shown in Figure 3.

Figure 3. Cont.
J. Fungi 2021, 7, 819 12 of 83
J. Fungi 2021, 7, x FOR PEER REVIEW 13 of 91

J. Fungi 2021, 7, x FOR PEER REVIEW 14 of 91

Figure 3. Maximum likelihood phylogenetic tree ML obtained from the DNA sequence data of ITS, LSU, RPB2, and TEF1α
Figure 3. Maximum likelihood phylogenetic tree ML obtained from the DNA sequence data of ITS, LSU, RPB2, and TEF1α
data sets.
dataBootstrap values
sets. Bootstrap BS from
values maximum
BS from maximumlikelihood
likelihood ML, left,maximum
ML, left, maximum parsimony
parsimony MP,MP, middle,
middle, equalequal to or greater
to or greater
than 70%
thanand
70%Bayesian posterior
and Bayesian probabilities
posterior PP, right,
probabilities equal
PP, right, to or
equal to greater than
or greater 0.95
than are
0.95 indicated
are indicatedabove
above or
or below
below the nodes
nodes as MLBS/MPBS/PP.
as MLBS/MPBS/PP. The tree isThe tree is
rooted rooted
with with Sanguinoderma
Sanguinoderma rugosumrugosum Cui and
Cui 9011 9011 Tomophagus
and Tomophagus colossus
colossus TC‐02.
TC-02. Newspecies,
New
species, new record species, and known species obtained in this study are indicated in bold black.
new record species, and known species obtained in this study are indicated in bold black.
Ganoderma specimens used for this study, based on the collections made from China,
Laos, Myanmar, Thailand, and Vietnam, were all placed within the Ganoderma clade. The
phylogenetic tree includes 14 laccate clades, one non‐laccate clade, and an outgroup clade.
In this study, of a total of 22 Ganoderma species, 13 species (viz. G. australe, G. calidophilum,
J. Fungi 2021, 7, 819 13 of 83

Ganoderma specimens used for this study, based on the collections made from China,
Laos, Myanmar, Thailand, and Vietnam, were all placed within the Ganoderma clade.
The phylogenetic tree includes 14 laccate clades, one non-laccate clade, and an outgroup
clade. In this study, of a total of 22 Ganoderma species, 13 species (viz. G. australe, G. cali-
dophilum, G. flexipes, G. gibbosum, G. leucocontextum, G. applanatum, G. lucidum, G. multi-
plicatum, G. resinaceum, G. sanduense, G. sichuanense, G. sinense, G. tsugae) from Yunnan
Province, China, 3 Ganoderma species (G. adspersum, G. australe, G. gibbosum) from Laos, 2
Ganoderma species (G. hoehnelianum, G. williamsianum) from Myanmar, and 11 Ganoderma
species (G. adspersum, G. applanatum, G. ellipsoideum, G. gibbosum, G. lucidum, G. multipileum,
G. orbiforme, G. philippii, G. sichuanense, G. sinense, G. subresinosum) are reported from Thai-
land. Furthermore, two taxa, viz. G. myanmarense from Myanmar and G. hochiminhense from
Vietnam, are described here as new species. The tree topologies provided considerably
high support in the terminal nodes but failed to recover deeper nodes with high statistical
support. The details of 13 Ganoderma species clades are provided in the following:
Clade 1 was statistically unsupported and comprised four species, viz. G. orb-
iforme, G. ecuadorense, G. sinense, and G. nasalaense. Three sequences of the laccate G. orb-
iforme from China, Laos, and Thailand clustered together with significant support values
(MLBS = 87%/MPBS = 91%/PP = 1.00), suggesting all of them to be the morphotype of
the same taxon. However, G. orbiforme is closely related to G. ecuadorense but differed
by strong statistical support (MLBS = 95%/MPBS = 91%/PP = 1.00). Two isolates of
G. sinense from China and Thailand clustered together (MLBS = 100%/MPBS = 86%/PP
= 0.99) and show genotypic closeness with the taxon G. nasalaense (MLBS = 100%/MPBS
= 91%/PP = 1.00). Clade 2 consists of the laccate G. angustisporum, G. ryvardenii, G. zona-
tum, and G. hochiminhense (MFLU 19-2224 and MFLU 19-2225). Two sequences of the
newly described species, G. hochiminhense, from Vietnam clustered together with strong
support values (MLBS = 100%/MPBS = 98%/PP = 0.99) and comes sister to G. ryvardenii
and G. zonatum although, this position is statistically unsupported.
Clade 3 was strongly supported (MLBS = 97%/MPBS = 99%/PP = 1.00) and contains
the laccate G. casuarinicola from China (Dai 16336) and Thailand (HKAS 104639), two se-
quences of G. thailandicum (HKAS 104640 and HKAS 104641) from Thailand, G. enigmaticum
(Dai 15970 and Dai 15971) from Africa, G. aridicola (Dai 12588) from South Africa and G. cali-
dophilum (MFLU 19-2174) from China. Basal to Clade 3, two species viz. G. williamsianum
and G. mbrekobenum formed Clade 4. The newly generated sequence of G. williamsianum
from Myanmar (MFLU 19-2170) clustered with the same taxon sequence, previously de-
posited in the GenBank nucleotide database from Thailand (Dai 16809), with full support
values (MLBS = 100%/MPBS = 100%/PP = 1.00). The molecular data confirmed the pres-
ence of G. williamsianum in Myanmar, and this holds the first distributional record of the
taxon from Myanmar.
Clade 5 consists of the non-laccate G. adspersum from Thailand (MFLU 19-2178 and
MFLU 19-2220) and Laos (MFLU 19-2177) (MLBS = 93%/MPBS = 75%/PP = 0.95); holotype
sequence of G. ellipsoideum from China (GACP14080966 and GACP14080968) and Thailand
(MFLU 19-2221) (MLBS = 97%/MPBS = 96%/PP = 0.95); G. gibbosum from Laos (MFLU
19-2190), Thailand (MFLU 19-2176), China (HKAS 97411), and Korea (SFC20150630-23)
(MLBS = 100%/MPBS = 93%/PP = 0.95); G. lobatum (JV 1212/10J) from USA; G. mutabile
(Yuan 2289) from China; and five sequences of G. australe from China (HKAS 97397,
GACP14061914, GACP14081671), Thailand (MFLU 13-0534), and Laos (MFLU 19-2171)
(MLBS = 100%/MPBS = 100%/PP = 1.00). However, this clade is statistically unsupported.
Clade 6 is formed by the cluster of Ecuadorean Ganoderma species viz. G. chocoense
(QCAM 3123) and G. podocarpense (QCAM6422). This clade was unsupported by the
maximum parsimony analysis but weakly supported by the maximum likelihood and
Bayesian analyses (MLBS = 71%/PP = 0.95).
Clade 7 is statistically unsupported. This clade comprises of the isolate of the laccate
G. multipileum from Thailand (MFLU 19-2216) and Taiwan, PRC (CWN 04670) (MLBS
= 100%/MPBS = 76/PP = 0.95) and shows its genetic closeness with that of G. steyaer-
J. Fungi 2021, 7, 819 14 of 83

tianum, G. mizoramense and G. destructans. Two newly amplified sequences of G. philippii

from Thailand (MFLU 19-2222 and MFLU 19-2223) clustered with strong support val-
ues (MLBS = 100%/MPBS = 99/PP = 1.00) with the same species sequence, previously
deposited from Malaysia (E7098 and E7425). The Chinese (Yuan 3490 and Dai 16434) and
Thailand (HKAS 97486) strains of G. tropicum comes basal to this clade but, this position is
not supported statistically. In Clade 8, two newly amplified sequences of G. sichuanense from
China (HKAS 97398) and Thailand (MFLU 19-2164) clustered with strong support values
(MLBS = 96%/MPBS = 87%/PP = 0.99) with the sequence of the same species G. sichua-
nense (CGMCC52175). Moreover, one of the Chinese sequences, G. lingzhi (Wu1006-38),
clustered with all the three sequences of G. sichuanense, we suggesting wrong identification.
earlier deposited in GenBank from China, G. curtisii stands sister to G. sichuanense, and this
position is strongly supported (MLBS = 100%/MPBS =76%/PP = 0.99).
Clade 9 comprised five small subclades. The laccate G. sessile along with G. neojapon-
icum and G. valesiacum formed a subclade with moderate support (MLBS = 84%/MPBS
= 81%/PP = 0.99). Sister to this subclade, there remains another subclade containing
the Chinese G. resinaceum along with sequences of the same taxon from the Netherlands
(CBS 194.76) and England (HMAS86599) but, this position is statistically unsupported.
The newly generated sequence of G. hoehnelianum from Myanmar (MFLU 19-2168) formed
the next subclade with that of the Chinese collection (Dai 11995) and showed its ge-
netic closeness with the laccate G. carocalcareum (DMC 322 and DMC 513) and G. aus-
troafricanum (CBS138724) with full support values (MLBS = 100%/MPBS = 100%/PP =
1.00). This subclade was followed by the cluster of two species viz. G. weberianum (CBS
219.36, GanoTK 17) and G. sichuanense (HMAS42798), but the position of these two taxa
was statistically unsupported. Two Chinese sequences of G. sanduense (GACP18012501 and
GACP18012502) form the next subclade. The basal subclade contained two newly gener-
ated sequences of G. applanatum from Thailand (MFLU 19-2175) and China (MFLU 19-2188)
along with the previously deposited sequence of the same taxon with strong support val-
ues (MLBS = 100%/MPBS = 100%/PP = 1.00). The newly described taxon, G. myanmarense
from Myanmar (MFLU 19-2167 and MFLU 19-2169), falls in Clade 10 where the new species
shows its genetic similarity with that of the laccate G. wiiroense from Ghana (UMN-20-GHA)
and G. destructans from South Africa (CMW43670). G. flexipes remains basal to this cluster
(Wei5494 and MFLU 19-2198). The newly generated sequence of G. flexipes (MFLU 19-2198)
shows its full identity with one of the previously deposited Chinese G. flexipes sequences
(Wei5494) and clustered with full support values (MLBS = 100%/MPBS = 100%/PP = 1.00).
Clade 11 was enriched with the G. lucidum species complex where four sequences of
G. lucidum from three countries viz. Thailand (MFLU 19-2162), China (MFLU 19-2161 and Cui
14404), and France clustered together (MLBS = 100%/MPBS = 100%/PP = 1.00) and revealed
G. leucocontextum (Dai 15601 and HKAS 97401), G. tsugae (Dai 12751b, HKAS 97406), G. orego-
nense as sister taxa with strong support values (MLBS = 100%/MPBS = 99%/PP = 1.00).
Two sequences of the taxon G. shandongense (Dai 15785 and Dai 15791) formed Clade
12. This small clade was followed by another clade, Clade 13, which comprised of four
sequences of G. boninense (Clade 13).
Clade 14 was the extreme basal clade where the sequence of G. subresinosum (MFLU
17-1912), collected from Thailand (MFLU 17-1912), clustered with full support (MLBS = 100%/
MPBS = 100%/PP = 1.00) with one of the earlier sequences of the same taxon, deposited in
the nucleotide sequence database.

3.2. Taxonomy
Ganoderma P. Karst., Revue Mycologique Toulouse. 3(9): 17 (1881)
= Dendrophagus Murrill, Bull. Torrey bot. Club. 32(9): 473 (1905)
= Elfvingia P. Karst., Bidr. Känn. Finl. Nat. Folk. 48: 333 (1889)
= Friesia Lázaro Ibiza, Revista Real Acad. Ci. Madrid. 14: 587 (1916)
= Ganoderma subgen. Trachyderma Imazeki, Bull. Tokyo Sci. Mus. 1: 49 (1939)
= Tomophagus Murrill, Torreya. 5: 197 (1905)
J. Fungi 2021, 7, 819 15 of 83

= Trachyderma (Imazeki) Imazeki, Bull. Gov. Forest Exp. Stn Tokyo. 57: 97 (1952)
Type species: Ganoderma lucidum (Leyss: Fr.) Karst.
Notes: (≡) is homotypic, or nomenclatural, synonyms, (=) is heterotypic, or taxonomic,
synonyms.
Description: Basidiomes annual, dimidiate, sessile or sub-stipitate to stipitate. Pileus
subdimidiate to dimidiate, flabelliform, perennial, stipitate or sessile. Pileus surface non-
laccate (dull) or weakly to strongly laccate, glossy, shiny, smooth, spathulate, shallow,
furrows, sulcate, several layers thick, with thin- to thick-cuticle cells or cuticle of clavate
end cells, thicker at the base than the margin, thin- to thick-crust overlaying the pileus,
consistency hard, consistency hard, light weight when dried. Pileus color variable, light
yellow to yellow, light brown, slightly brown to dark brown, sometimes homogeneous
reddish gray to reddish-yellow. Context brown to dark brown, grayish orange to orange,
sometimes grayish-yellow, mostly soft, sometimes spongy to firm-fibrous. Hymenophore
di-trimitic, heterogeneous, non-septate or septate, usually yellow, slightly light orange,
or light brown to brown, sometimes with melanoid bands. Tubes are hard, woody when
dried. Tube layers single or stratified, pale to purplish-brown, almost hyaline with clamp
connections, occasionally branched at apex, thin- to thick-walled. Stipe central or lateral,
glossy with a distinct cuticle. Margin actively growing, entirely white when fresh, round,
soft and smooth when young, slippery when touched from youth to maturity, and tough
when broken. Pores 4–7 in number per mm, angular, entire, subcircular to circular, regular,
cream or white when young, light yellow, light orange to brown when mature. Pore surface
usually white to cream when fresh, turning yellowish-white to pale yellow on drying,
some sections reddish gray to brown, and brownish gray when wet.
Hyphal structure: Hyphal system di-trimitic, including generative, skeletal, and binding
hyphae; mostly generative hyphae with clamp connections, hyaline, brown, non-septate,
or septate, often with long and tapering branches. Basidia broadly ellipsoid, tapering
abruptly at the base. Cystidia absent. Basidiospores broadly to narrowly ellipsoid or
oblong, sometimes globose to subglobose, with double walls, truncate apex, apical germ
pore present, usually with light brown to brown endosporium, hyaline exosporium with
thin inter-walled pillars, hyaline endosporium with thick outer walls, and some very thin
exosporium.
Ecology: mostly on hardwoods, trunks, and stumps, occurring on several different
living tree host species.
Notes: Justo et al. [103] treated Ganodermataceae as a synonym of Polyporaceae
and included the genus Ganoderma under Polyporaceae. Later, Cui et al. [104] excluded
Ganoderma from Polyporaceae due to the presence of double-walled basidiospores, unlike
Polyporaceae. So, the distinctiveness of the genus Ganoderma lies in the presence of double-
walled and truncate basidiospores. Species with a laccate, glossy surface are present in
both Ganodermataceae and Polyporaceae as centrally and laterally stipitate species.

3.2.1. Taxonomy of Ganoderma from China

Ganoderma angustisporum J.H. Xing, B.K. Cui and Y.C. Dai, Mycokeys 34: 98 (2018)
Taxonomy and phylogenic analysis were described in Xing et al. [37]
Notes: Ganoderma angustisporum is characterized by annual, sessile, broadly basidiomes,
strongly laccate on the upper surface of basidiomes, white pore surfaces, and almond-
shaped, slightly truncate, narrow 9.0–11.3 × 4.0–5.2 µm basidiospores. It is a group of
white-rot fungi that predominantly grow on living Casuarina equisetifolia in Fujian Province,
China.
Ganoderma applanatum (Pers.) Pat., Hymenomyc. Eur. (Paris): 143 (1887) (Figure 4)
≡ Boletus lipsiensis Batsch, Elenchus fungorum. Continuatio prima.: 183, t. 25:130
(1786)
≡ Scindalma lipsiense (Batsch) Kuntze, Revisio generum plantarum. 3(2): 518 (1898)
≡ Polyporus lipsiensis (Batsch) E.H.L. Krause, Basidiomycetes Rostochienses.: 54 (1928)
J. Fungi 2021, 7, 819 16 of 83

≡ Agaricus lipsiensis (Batsch) E.H.L. Krause, Basidiomycetum Rostochiensium, Suppl.

4: 142 (1932)
= Boletus applanatus Pers., Observationes mycologicae. 2: 2 (1799)
= Polyporus merismoides Corda, Deutschlands Flora, Abt. III. Die Pilze Deutschlands. 3:
139 (1837)
= Polyporus stevenii Lév., Annls Sci. nat., Bot.: 91 (1844)
= Polyporus leucophaeus Mont., Sylloge generum specierumque plantarum cryptoga-
marum.: 157 (1856)
= Polyporus leucophaeum Mont. (1856)
= Polyporus incrassatus Berk., Journal of the Linnean Society. Botany. 16: 41 (1877)
= Polyporus concentricus Cooke, Grevillea. 9(49): 13 (1880)
= Fomes gelsicola Berl., Malpighia. 3: 373 (1889)
= Fomes nigriporus Lázaro Ibiza, Revista de la Real Academia de Ciencias Exactas
Fisicas y Naturales Madri. 14: 662 (1916)
= Ungularia subganodermica Lázaro Ibiza, Revista de la Real Academia de Ciencias
Exactas Fisicas y Naturales Madri. 14: 674 (1916)
= Fomes longoporus Lloyd, Mycological Writings. 6(62): 940 (1920)
Facesoffungi number: FoF 06249
Description: Basidiomes annual, perennial, sessile. Pileus 1.5–5.8 cm in length, 0.5–4.5 cm
in width, and up to 1.5 cm thick at the base, sessile (without stipe), perennial, subdimidiate,
sub-flabelliform to flabelliform, usually flat, convex, imbricate, umbonate or uneven, rarely
ungulate, glabrous when present, broadly attached when mature, often with undefined
concentric zones at the center that extend to the margin, thick at the base, slightly soft at the
margin when mature. Pileus surface shiny, silky, smooth, and soft when young, hard when old,
frequently furrowed and shallow sulcate, undulating, somewhat spathulate to uneven on the
upper surface when mature, covered by irregularly ruptured thick crust, slightly non-laccate
(dull) and faded from when mature to old, compact and hard when mature, woody to corky
when old. Pileus color is usually homogenous with grayish-orange (6B3–6B5) at the center,
slight brownish-orange (6C4), orange white (6A2), to pale orange (6A3), with yellowish-gray
(4B2) at the margin when mature. Context up to 0.3–1 cm thick at the base, mostly light brown
(7D5), brown (6E8) to dark brown (7F6–7F8) of cuticle cells, with walls varying in thickness to
subsolid hyphae, some fibrous pithy context, usually separated by layers of context tissue
at the base, and some occurred woody lines. Tube woody, hard, often dark brown (7F7–7F8)
when dried, with sulcate at different levels. Stipe almost sessile and broadly attached when
present, with a differentiated zone at the point of attachment. Margin up to 1 cm thick, white
(5A1), yellowish-gray (4B2) when mature, turns light brown (6D4) to brown (6E8) when
scratched or bruised, often slippery when wet, soft when young, thinner than the center.
Pore 4–6 in number per mm, subcircular to circular, sometimes angular. Pore surface initially
white (7A1), grayish-orange (7C3–7C4) when mature, turning to light brown (7D6) to brown
(7D8) when scratched or bruised.
Hyphal structure: Hyphal system trimitic; generative hyphae 0.8–2.6 µm (x = 2.1,
n = 30) in diam, almost hyaline, with clamp connections, abundant, thin-walled and
occasionally thick-walled; skeletal hyphae 2.1–4.6 µm width (n = 30), usually thick-walled,
hyaline, sometimes branched; binding hyphal 1.6–3.3 µm width (n = 30), thick-walled and
occasionally thin-walled, branched, and intertwined with the skeletal hyphae. Pileipellis a
hymeniderm, grayish brown (6E4), which is composed of apically acanthus-like branched
cells. Basidiospores mostly ellipsoid with double walls, with a size range of (9.8–)10.4–11.1–
11.9(–12.1) × (7.3–)8.0–8.6–9.2(–9.9) µm, (x = 11.3 × 8.7 µm, n = 50) µm, with Q = 1.79–1.86,
L = 11.23 µm, W = 6.12 µm (including myxosporium), (6.2–)7.6–8.6–9.7(–10.4) × (5.0–)
5.8–7.1–8.2(–8.9) µm (x = 8.6 × 7.1 µm, n = 50) µm, with Q = 1.19–1.24, L = 8.59 µm,
W = 7.12 µm (excluding outer myxosporium), brownish-orange (7D4) to brown (7D7–
7D8) in KOH, and reddish-brown (8E6) to dark brown (8F4) in Melzer’s reagent. Basidia
14–20 × 8–10 µm, with four sterigmata.
J. Fungi 2021, 7, 819 17 of 83
J. Fungi 2021, 7, x FOR PEER REVIEW 19 of 91

Figure
Figure 4.4. Morphology
Morphologyof ofGanoderma
Ganodermaapplanatum
applanatum(HKAS
(HKAS107254,
107254,MFLU
MFLU19‐2188): (a,b)
19-2188): mature
(a,b) basidiomes;
mature (c)(c)
basidiomes; pore charac‐
pore char-
teristics; (d–f) hyphae of trama in KOH; (g,h) generative hyphae of context in KOH; (i–k) basidiospores. Scale bars: (a,b)
acteristics; (d–f) hyphae of trama in KOH; (g,h) generative hyphae of context in KOH; (i–k) basidiospores. Scale bars:
= 2 cm, (c) = 1000 μm, (d–h) = 20 μm, (i–k) = 5 μm.
(a,b) = 2 cm, (c) = 1000 µm, (d–h) = 20 µm, (i–k) = 5 µm.
Ganoderma australe
Ecology: Solitary (Fr.) Pat.,
on stump Bull. Soc.yunnanensis.
of Machilus mycol. Fr. 5(2–3): 65 (1889) (Figure 5)
≡ Polyporus australis Fr., Elenchus
Specimens examined: CHINA, Yunnan Province,Fungorum. 1: 108 (1828)25◦ 090 3500 N, 99◦ 090 4900 E,
Baoshan,
≡ Fomes australis (Fr.) Cooke, Grevillea. 14(69): 18 (1885)
1973 m elev., 11 November 2017, T. Luangharn, HKAS 107254, MFLU 19-2188.
≡ Placodes
Notes: australis
Ganoderma (Fr.) Quél.,
lipsiense Enchiridion
has been treated byFungorum in Europa
some researchers as media et praesertim
the correct name for
in Gallia Vigentium. 171 (1886)
G. applanatum [15]. G. applanatum (=G. lipsiense) belongs in the subgenus Elfvingia, which is
≡ Fomes by
characterized applanatus
distinctivevar. australeisspecies,
non-laccate (Fr.) Cleland andacute
a thin and Cheel, Journal
margin of of
theProceedings of
pileus, and un-
the Royal Society of New South Wales. 51: 518 (1918)
branched terminal endings of skeletal hyphae, with ellipsoid basidiospores [47,105–107]. G. ap-
≡ Ganoderma
planatum applanatum
causes white butt rot on subsp. australe (Fr.)
angiosperm treesBourdot and Galzin,
and is widely Bulletin
distributed de la [64].
in China So‐
ciété Mycologique de France. 41: 184 (1925)
Hence, our specimen of G. applanatum, collected from a temperate region of China, is described
based≡onGanoderma applanatum
morphological f. australe
characteristics and(Fr.) Bourdotphylogenetic
molecular and Galzin,data.
Bulletin
Ourde la Société
results agree
Mycologique
well with thosedeofFrance.
Ryvarden41: 184
and (1925)
Gilbertson [105].
≡ Elfvingia australis
Ganoderma australe(Fr.)
(Fr.) G. Cunn.,
Pat., Bull. Bulletin of the
Soc. mycol. Fr. New Zealand
5(2–3): Department
65 (1889) (Figure 5) of In‐
dustrial Research. 164: 256 (1965)
J. Fungi 2021, 7, 819 18 of 83

≡ Polyporus australis Fr., Elenchus Fungorum. 1: 108 (1828)

≡ Fomes australis (Fr.) Cooke, Grevillea. 14(69): 18 (1885)
≡ Placodes australis (Fr.) Quél., Enchiridion Fungorum in Europa media et praesertim
in Gallia Vigentium. 171 (1886)
≡ Fomes applanatus var. australeis (Fr.) Cleland and Cheel, Journal of Proceedings of
the Royal Society of New South Wales. 51: 518 (1918)
≡ Ganoderma applanatum subsp. australe (Fr.) Bourdot and Galzin, Bulletin de la Société
Mycologique de France. 41: 184 (1925)
≡ Ganoderma applanatum f. australe (Fr.) Bourdot and Galzin, Bulletin de la Société
Mycologique de France. 41: 184 (1925)
≡ Elfvingia australis (Fr.) G. Cunn., Bulletin of the New Zealand Department of
Industrial Research. 164: 256 (1965)
= Polyporus tornatus Pers., Botanique (Nagpur). 5: 173 (1827)
= Polyporus scansilis Berk., Journal of the Linnean Society. Botany. 16: 53 (1877)
= Fomes annularis Lloyd, Mycol. Writ. 4(40): 6 (1912)
= Ganoderma tornatum var. tornatum (Pers.) Bres., Hedwigia. 53(1–2): 55 (1912)
= Fomes konigsbergii Lloyd (1915)
= Fomes polyzonus Lloyd, Synopsis of the genus Fomes. (7): 269 (1915)
= Fomes pseudoaustraleis Lloyd, Synopsis of the genus Fomes. (7): 269 (1915)
= Fomes undatus Lázaro Ibiza, Revista de la Real Academia de Ciencias Exactas Fisicasy
Naturales Madri. 14: 661 (1916)
Facesoffungi number: FoF 06242
Description: Basidiomes annual, perennial, subdimidiate, sessile. Pileus 14–28 cm in
length, 12–32 cm in width, and 1.4–3.2 cm thick. Pileus flabelliform, spathulate, subdimidi-
ate, umbonate, single, sulcate, large, obtuse from host, radial from the center extending the
margin, broadly attached, often thick at the center, slightly soft at the margin, consistency
hard, and tough to break when dried. Pileus surface convex, corky, furrowed, spathulate,
mostly umbonate or uneven, non-laccate (dull) on maturity or in old, usually slippery
where the new hyphae are in active development (margin), slightly concentrically sulcate at
the center toward margin, smooth, covered with thick and hard crust, irregularly ruptured
crust overlying the surface, woody, and corky when dried, with cracked crust when mature,
and tough to break when dried. Pileus color often brown (6E7–6E8) at the base, reddish-
orange (7B7–7B8), brownish-orange (7C6–7C8), almost covered with grayish-red (8C4–8C5)
on the upper surface when old, slight reddish-brown (8F8, 9E7–9E8) close to the margin.
Context up to 0.5–2 cm thick near stipe, fibrous, composed of coarse loose fibrils, brown
(6D7–6D8) to dark brown (6F7), with reddish-brown (8D8–8D9) coarse loose fibrils, covered
with thick crust. Tube 0.4–1.5 cm long, brown (7D8) to dark brown (6F8). Stipe sessile,
broadly attached. Margin soft when young, slippery when fresh, blunt when mature, white
(4A1). Pore 4–6 in number per mm, subcircular to circular, sometimes angular. Pore surface
initially white (4A1), slight to pale yellow (4A3) when mature, turned brownish-orange
(6C7–6C8) when scratched or bruised or discolored when touched.
Hyphal structure: Hyphal system trimitic, dense and hard, thick-walled, typically
with narrow lumen, flexuous, and many branches, usually brownish-orange (6C5–6C7)
in KOH; generative hyphae 2.2–3.8 µm broad (n = 30), thin-walled, hyaline, tapering at
branch, with clamp connections; skeletal hyphae 2.9–4.2 µm broad (n = 30), sometimes
branched, nearly solid, thick-walled; binding hyphae 2.6–4.0 µm broad (n = 30), thick-
walled, branched, more or less solid; hymenial with sword-like apices in the context.
Pileipellis a hymeniderm, usually brownish-orange (6C5) to brown (6E8), composed of
apically acanthus-like branched cells. Basidiospores mostly ellipsoid to broadly ellip-
soid, double walls, (6.2-)7.1–9.4–10.4(-11.8) × (5.2-)6.0–7.4–8.9(-9.7) µm (x = 9.4 × 7.4 µm,
n = 50) µm, with Q = 1.24–1.30, L = 9.42 µm, W = 7.43 µm (including myxosporium),
(5.3-) 6.7–7.8–9.6(-10.5) × (4.5-)5.1–5.7–6.3(-7.2) µm (x = 7.8 × 5.7 µm, n = 50) µm, with
Q = 1.31–1.38, L = 7.85 µm, W = 5.83 µm (excluding outer myxosporium), overlaid by
hyaline, apically brown, bearing a fine, distinct, short, echinulae truncate, turgid vesicular
 Ganoderma australe is distinguished from G. applanatum by the larger dimensions of
its basidiospores, different stipe features, thickness of the cuticle, and color of the context
layer, all of which were considered in delimiting G. applanatum and G. australe [2,105,107].
G. applanatum is confined to northern temperate regions, while G. australe can be found in
J. Fungi 2021, 7, 819 tropical and subtropical regions [112,113]. There are reports on the occurrence in Australia
19 of 83
[14], China [30], New Zealand [114], southern India [51,95], Taiwan, PRC [115], and Thai‐
land [30,59]. Ganoderma australe is a cosmopolitan species, which is known to cause white
rot on woody material. It shows parasitic or pathogenic behavior on a wide range of both
appendix,
dead inner broadleaved
and living wall orange (6B8) to brownish-orange
deciduous trees [116–118].(6C8) or lightaustrale
Ganoderma brown is(7D5–7D6)
distributed to
brown (7E7–7E8), outer wall mostly reddish-brown
worldwide, especially in tropical regions [14]. (8E7–8E8, 8F7) in 5% KOH.

Figure
Figure 5.
5. Morphology
Morphology of of Ganoderma
Ganoderma australe
australe (HKAS
(HKAS 97397):
97397): (a,b)
(a,b) mature
mature basidiomes;
basidiomes; (c)
(c) margin;
margin; (d)
(d) pore
pore characteristics;
characteristics;
(e) mycelia of tube layers; (f–h) context hyphae as seen in Melzer’s reagent; (i–l) basidiospores. Scale
(e) mycelia of tube layers; (f–h) context hyphae as seen in Melzer’s reagent; (i–l) basidiospores. Scale bars: (a) =(a)5 cm,
bars: (b)
= 5 cm,
= 3 cm, c = 1 cm, d = 500 μm, e = 20 μm, f = 20 μm, g,h = 10 μm, i–l = 5 μm.
(b) = 3 cm, (c) = 1 cm, (d) = 500 µm, (e) = 20 µm, (f) = 20 µm, (g,h) = 10 µm, (i–l) = 5 µm.

Ganoderma calidophilum
Habitat: Solitary, J.D.living
growing on Zhao,Neocinnamomum
L.W. Hsu and delavayi
X.Q. Zhang,
(Lec.)Acta Mycologica
H. Liou. tree or,
Sinica 19: 270 (1979) (Figure 6)
decaying stump, and living Fagus spp.
Facesoffungi number:
Specimens examined: FoF 06244
CHINA, Yunnan Province, Kunming Botanical Garden, 25◦ 080 3900 N,
◦ 0 0
102 44 30 E, 1956 m, 27 September 2016, T. Luangharn, HKAS 97397.
Notes: Ganoderma australe belongs under the subgenus Elfvingia [38]. This fungus was
initially described from the Pacific Islands [108]. G. australe belongs to the G. applanatum-
australe complex [2]. Ganoderma australe was established as a non-laccate (dull) pilei.
The type specimen of this fungus is missing, while the neotype specimen is available in
Europe [105]. Ganoderma applanatum and G. australe from Europe have been confused
based on the macro-characteristic features [107]. The typification of G. australe remains
unresolved and was exemplified by several authors [61,109–111], and its similar cultural
characteristics also showed the phenotypic plasticity in morphological level and a higher
level of nucleotide divergence in the ITS rDNA region that made G. australe a complex
species [45].
J. Fungi 2021, 7, 819 20 of 83

Ganoderma australe is distinguished from G. applanatum by the larger dimensions of

its basidiospores, different stipe features, thickness of the cuticle, and color of the context
layer, all of which were considered in delimiting G. applanatum and G. australe [2,105,107].
G. applanatum is confined to northern temperate regions, while G. australe can be found
in tropical and subtropical regions [112,113]. There are reports on the occurrence in Aus-
tralia [14], China [30], New Zealand [114], southern India [51,95], Taiwan, PRC [115],
and Thailand [30,59]. Ganoderma australe is a cosmopolitan species, which is known to
cause white rot on woody material. It shows parasitic or pathogenic behavior on a wide
range of both dead and living broadleaved deciduous trees [116–118]. Ganoderma australe
is distributed worldwide, especially in tropical regions [14].
Ganoderma calidophilum J.D. Zhao, L.W. Hsu and X.Q. Zhang, Acta Mycologica
Sinica 19: 270 (1979) (Figure 6)
Facesoffungi number: FoF 06244
Description: Basidiomes annual, stipitate, subdimidiate. Pileus 3–7 cm in length, 2–4 cm
in width, and 0.2–1 cm thick. Pileus subdimidiate to dimidiate, spathulate, stipitate, sulcate,
umbonate, radial from the center extending to the margin, tough to break when dried, often
thick at the center, slightly soft at the margin, and light in weight when dried. Pileus surface
corky, convex, furrowed, glabrous, glossy, incised, shiny, spathulate, shallow, sulcate when
fresh, umbonate or uneven, laccate and glossy when mature, weakly laccate when old and
in regions of developing hyphae (margin), slightly concentrically sulcate, layers smooth
at the center when young, irregularly ruptured crust overlying the context, and tough
to break when dried. Pileus color usually homogenous with brownish-red (8C7–8C8),
brownish-red (9C7–9C8), reddish-brown (9D6–9D7) center, extending brownish-orange
(6C7–6C8) toward the stipe, brownish-red (9C8) from the center to light brownish-orange
(6C8), and usually light brown (6D8) at the margin when old. Context up to 0.2–0.6 cm thick
near stipe, dry, fibrous, composed of coarse loose fibrils, brownish-orange (6C5–6C8) upper
layers when fresh, brown (6D7) at lower layers, dark brown (8F7) when dried, covered
with thin crust, trimitic hyphal system. Tube 0.3–0.9 cm in length, brown (7D8). Stipe 5–
14 cm long, cylindrical, almost stipitate with broadly, irregularly ruptured crust overlying,
strongly laccate with brown (7D8) when mature, dark brown (8F8) when old, and woody or
corky when dried. Margin soft when young, laccate when mature, weakly laccate to laccate
when old, blunt when old, usually light brown (6D8) when mature to old. Pore 4–5 in
number per mm, subcircular to circular, sometimes angular. Pore surface initially pale
orange (5A3) to brownish-orange (6D8) when mature, discolored when touched, brown
(6E8) when scratched or bruised.
Hyphal structure: Hyphal system trimitic hyphal, usually brownish-orange (6C5–6C7)
in KOH; generative hyphae 1.2–3.2 µm broad (n = 30), thin-walled, hyaline, without
clamp connections; skeletal hyphae 3.2–6.4 µm broad (n = 30), sometimes branched, nearly
solid, thick-walled, without clamp connections; binding hyphae 2.4–5.2 µm broad (n = 30),
usually thin to thick-walled, many branches, nearly solid, hymenial with sword-like apices
in the context. Basidiospores mostly ellipsoid to broadly ellipsoid, with double wall, with a
size range of (7.4-)8.5–11.9–12.6(-13.7) × (6.3-)7.2–8.3–9.1(-9.6) µm (x = 11.9 × 8.3 µm,
n = 50) µm, with Q = 1.39–1.45, L = 11.92 µm, W = 8.35 µm (including myxosporium),
(6.8-)7.6–10.4–11.3(-12.8) × (5.4-)6.3–7.0–7.6(-8.1) µm (x = 10.4 × 7.1 µm, n = 50) µm, with
Q = 1.43–1.49, L = 10.39 µm, W = 7.12 µm (excluding outer myxosporium), overlaid by
hyaline, apically and echinulae, truncate, turgid vesicular appendix, inner wall orange
(6B8) to deep orange (6A8), reddish-orange (7A8, 7B7–7B8) or yellowish-red (8B8), outer
wall usually reddish-brown (8D7–8D8; 8E8) in 5% KOH.
Habitat: Solitary, near the hardwood root of Castanopsis spp., living tree of Machilus
yunnanensis.
Specimens examined: CHINA, Yunnan Province, Baoshan, 25◦ 060 2900 N, 99◦ 080 2900 E,
1973 m elev., 11 November 2017, T. Luangharn, MFLU 19-2174.
Notes: Ganoderma calidophilum is a species originally described from Hainan Province,
China, by Zhao [119]. Several reports have confirmed that this fungus is mentioned in
J. Fungi 2021, 7, 819 21 of 83

Hainan Province polypore diversity checklists [13,61,120,121]. This fungus is distinctive in

these forms, featuring a laccate pileus with broadly ellipsoid basidiospores with double
walls, and it is widely found across subtropical and tropical Asia [108,119,122]. Wang and
Wu [112] suggested that G. calidophilum is a synonym of G. flexipes. However, the evaluated
G. calidophilum and G. flexipes are different in terms of pileus color, pileus shape and size,
context, and basidiospore size [112]. In this study, we present our G. calidophilum collection
J. Fungi 2021, 7, x FOR PEER REVIEW 23 of 91
from Yunnan Province, China, based on taxonomic and phylogenetic analyses. Our strain
is similar to the described strain of Wang and Wu [119], Zhao et al. [119], and Bi et al. [123].

Figure6.
Figure 6. Morphology
Morphology of of Ganoderma
Ganodermacalidophilum
calidophilum(MFLU
(MFLU19-2174):
19‐2174):(a,b)
(a,b)mature
maturebasidiomes;
basidiomes;(c)
(c)pore
pore
characteristics; (d–f) context hyphae in Melzer’s reagent; (g) tube layer hyphae in Melzer’s reagent;
characteristics; (d–f) context hyphae in Melzer’s reagent; (g) tube layer hyphae in Melzer’s reagent;
(h–k) basidiospores. Scale bars: (a) = 2 cm, (b) = 5 cm, (c) = 1000 μm, (d–g) = 20 μm, (h–k) = 5 μm.
(h–k) basidiospores. Scale bars: (a) = 2 cm, (b) = 5 cm, (c) = 1000 µm, (d–g) = 20 µm, (h–k) = 5 µm.

Ganodermaflexipes
Ganoderma flexipesPat.,
Pat., Bulletin
Bulletin de de la Société
la Société Mycologique
Mycologique de France.
de France. 23(1):
23(1): 75 75
(1907)
(1907) (Figure
(Figure 7) 7)
≡≡Fomes
Fomesflexipes
flexipes(Pat.)
(Pat.)Sacc.
Sacc.and
andTraverso,
Traverso,Sylloge
SyllogeFungorum.
Fungorum.19: 19:710
710(1910)
(1910)
≡≡Polyporus
Polyporusflexipes
flexipes(Pat.)
(Pat.)Lloyd,
Lloyd,Synopsis
Synopsisofofthe
thestipitate
stipitatePolyporoids.
Polyporoids.(7):
(7):104
104 (1912)
(1912)
Facesoffunginumber:
Facesoffungi number: FoF
FoF 06245
06245
Description: Basidiomes annual or perennial, stipitate. Pileus 0.5–3.2 cm in length, 0.5–
3 cm broad, up to 0.5 cm thick at the base. Pileus stipitate, sub‐reniform to reniform, or
subflabellate to flabellate, concentrically sulcate zones with tuberculate, glabrous when
young to maturity, bumps when mature, often tough to break when dried. Pileus surface
J. Fungi 2021, 7, 819 22 of 83

Description: Basidiomes annual or perennial, stipitate. Pileus 0.5–3.2 cm in length,

0.5–3 cm broad, up to 0.5 cm thick at the base. Pileus stipitate, sub-reniform to reniform,
or subflabellate to flabellate, concentrically sulcate zones with tuberculate, glabrous when
young to maturity, bumps when mature, often tough to break when dried. Pileus surface
shiny, smooth, and soft when young, frequently furrowed and shallow sulcate on upper
surface, undulating, somewhat spathulate to uneven when mature, covered by an irregu-
larly ruptured thin crust, faded or weakly laccate when young, and laccate when mature.,
and woody when old. Pileus color usually homogenous with reddish-brown (8E8) to dark
brown (9F7–9F8) at the center, slight to the margin from mature to old. Context up to
0.1–0.6 cm thick at the base, very dry, brown (7D7–7D8) to reddish-brown (8E7), containing
fibrous pithy context and corky when old. Tube hard, often dark brown (7F8). Stipe almost
3–12 cm in length, 0.3–1.5 in width, sub-cylindrical to cylindrical, often dark brown (7F8),
and strongly laccate from mature to old. Margin soft when young, laccate when mature,
some wavy, often light brown (6D5–6D6) on the upper surface. Pore 4–5 in number per mm,
subcircular to circular, sometimes angular. Pore surface initially grayish-orange (6B4–6B6),
turns brown (7D7) to reddish-brown (8D5–8D7) when scratched or bruised, discolored
when touched.
Hyphal structure: Hyphal system trimitic, bearing clamp connections, hyaline, thick-
walled, tapering branch, some swollen differentiated zone at the point of attachment;
generative hyphae (1.8-)2.2–2.9–3.4(-3.8) µm broad (n = 30), thin-walled, hyaline, un-
branched, with clamp connections; skeletal hyphae (3.0-)3.8–4.8–5.4(-6.2) µm broad (n = 30),
with walls varying in thickness, with subsolid, binding hyphae (2.2-)2.8–3.8–4.5(-5.1) µm
broad (n = 30), usually thick-walled, appearing alongside Bovista hyphae, and many
branches, usually light yellow (4A4–4A5) to yellowish-orange (4B8) of thin-walled and
orange (6A7) to deep orange (6A8) of thick-walled in Melzer’s reagent. Pileipellis a hymeni-
derm, dark brown (6F8), composed of apically acanthus-like branched cells. Basidiospores
mostly ellipsoid to broadly ellipsoid with double wall at maturity, (8.1-)8.8–9.7–10.6(-11.2)
× (6.1-)6.6–7.7–9.7(-10.4) µm (x = 9.7 × 7.7 µm, n = 50) µm, with Q = 1.08–1.15, L =
9.68 µm, W = 7.72 µm (including myxosporium), (7.3-)7.8–8.3–8.7(-9.2) × (4.0-)4.6–5.4–
5.8(-6.2) µm (x = 10.2 × 6.4 µm, n = 50) µm, with Q = 1.51–1.57, L = 8.34 µm, W = 5.39 µm
(excluding outer myxosporium), overlaid by hyaline, dextrinoid, echinulae, inner wall ech-
inulate brown (5D8, 7E6–7E8), and outer wall usually dark brown (7E8) to reddish-brown
(8E6–8E8) in Melzer’s reagent.
Habitat: Solitary on the decaying hardwood of Pinus spp.
Specimens examined: CHINA, Yunnan Province, Baoshan, 25◦ 060 2900 N, 99◦ 080 2900 E,
1973 m elev., November 2017, T. Luangharn, MFLU 19-2189.
Notes: Ganoderma flexipes is originally described from Vietnam by Patouillard [124].
It has been recorded from China, India, Laos, Nepal, and Pakistan [4,13,30,110,122]. G. flex-
ipes is characterized by its small reddish-brown pileus, long and thin stipe, usually reddish-
brown to dark brown context, and ellipsoid or ovoid basidiospores [125]. Among the
Chinese Ganoderma species, G. flexipes is one of the most similar species to G. sichuanense
as they share a reddish-brown pileal surface, similar basidiospores, and cuticle cells [4].
Our G. flexipes from China is very similar to the description of Ryvarden [125] and Ha-
puarachchi et al. [30], and basidiospores are within the range of 9.7–10.2 × 6.4–7.7 µm.
J. Fungi 2021, 7, 819 23 of 83
J. Fungi 2021, 7, x FOR PEER REVIEW 25 of 91

Figure7.
Figure 7. Morphology
Morphologyof ofGanoderma
Ganodermaflexipes (MFLU
flexipes (MFLU19‐2189): (a,b)
19-2189): mature
(a,b) basidiomes;
mature (c) pore
basidiomes; char‐
(c) pore
acteristics; (d) hyphae of tube layers in Melzer’s reagent; (e,f) context hyphae in Melzer’s reagent;
characteristics; (d) hyphae of tube layers in Melzer’s reagent; (e,f) context hyphae in Melzer’s reagent;
(g–i) basidiospores. Scale bars: (a) = 3 cm, (b) = 2 cm, (c) = 500 μm, (d) = 30 μm, (E,F) = 20 μm, (G–I)
(g–i) basidiospores. Scale bars: (a) = 3 cm, (b) = 2 cm, (c) = 500 µm, (d) = 30 µm, (e,f) = 20 µm,
= 5 μm.
(g–i) = 5 µm.

Ganodermagibbosum
Ganoderma gibbosum(Blume
(Blumeand andT.T.Nees)
Nees)Pat.,
Pat.,Ann.
Ann.Jard.
Jard.Bot.
Bot.Buitenzorg,
Buitenzorg,suppl.
suppl.1:1:
114 (1897) (Figure
114 (1897) (Figure 8) 8)
≡≡Polyporus
Polyporus gibbosus
gibbosus (Blume
(Blume andand T. Nees).,
T. Nees)., Nov. Nov. Act. Academiae
Act. Academiae CaesareaeCaesareae Leo‐
Leopoldino
poldino Carolinae
Carolinae Germanicae Germanicae Naturae Curiosorum.
Naturae Curiosorum. 13: 19, (1826)
13: 19, t. 4(1–4) t. 4(1–4) (1826)
≡ Fomes amboinensis var. gibbosus (Blume and T.
≡ Fomes amboinensis var. gibbosus (Blume and T. Nees) Cooke, Nees) Cooke,Grevillea.
Grevillea.13(68):
13(68):118
118
(1885)
(1885)
≡≡Fomes
Fomesgibbosus
gibbosus(Blume
(Blumeand andT.T.Nees)
Nees)Sacc.
Sacc.Syll.
Syll.Fung.
Fung.6:6:156 156(1888)
(1888)
≡ Scindalma gibbosum (Blume and T. Nees) Kuntze., Revisio
≡ Scindalma gibbosum (Blume and T. Nees) Kuntze., Revisio generum plantarum generum plantarum3(2):
3(2):
518(1898)
518 (1898)
Facesoffunginumber:
Facesoffungi number:FoF FoF06246
06246
Description:Basidiomes
Description: Basidiomesannual
annualoror perennial,
perennial, sessile.
sessile. Pileus
Pileus 8–218–21
cm cm in length,
in length, 6–136–13
cm
cm in width, and 1–3.5 cm thick, convex, imbricate, umbonate, uneven,
in width, and 1–3.5 cm thick, convex, imbricate, umbonate, uneven, ungulate, subflabellate, ungulate, subfla‐
bellate, subdimidiate,
subdimidiate, usually
usually round whenround when
present, present, somewhat
primordial, primordial,roundsomewhat round
and plump and
when
plump somewhat
young, when young, somewhat
imbricate, when imbricate, when flabelliform
seen from above seen from (fan-shaped),
above flabelliform
broadly(fan‐
at-
shaped),
tached, broadly
thick at theattached, thicksoft
base, slightly at at
thethe
base, slightly
margin when soft at thePileus
mature. margin whennon-laccate
surface mature. Pi‐
leus surface
(dull), non‐laccate
furrowed, (dull), furrowed,
incised, sulcate, smooth when incised,
young, sulcate,
usuallysmooth when
silky, soft, young,
and usually
slippery sur-
J. Fungi 2021, 7, 819 24 of 83

face when fresh, undulating on the upper surface, somewhat spathulate to uneven, with a
crust (0.2–0.4 mm), woody from mature to older, and lined or cracked crust occurs when
old. Pileus color usually homogenous with grayish-orange (6B3–6B6), brownish-orange
(6C5–6C6), and brown (6D7–6D8) at the base extending to the margin of mature fruiting
bodies. Context up to 0.5–1.8 cm thick, compact and hard, trimitic hyphal, with clamp con-
nections, hyaline, with walls varying in thickness with simple septa, composed of narrow
and sparingly branched; generative hyphae 1.3–3.2 µm broad (n = 30) with hyaline; skeletal
hyphae 2.8–4.7 µm broad (n = 30), usually thick-walled; binding hyphal 2.1–4.2 µm width
(n = 30) with walls varying in thickness. Hymenophore reddish-brown (8D7). Tube layers
0.4–1.2 cm in length. Stipe almost sessile and broadly attached when present. Margin
blunt-edged, wavy, slippery from young, softer, and often white (8A1) when youth to
maturity, and light brown (6D5) when old, the present yellow line between the edge of the
margin and close to the underside of basidiomes. Pore 4–7 in number per mm, subcircular
to circular. Pore surface white (11A1) when present, pale yellow (4A3) to grayish-yellow
(4B3–4B4) when scratched or bruised, discolored when touched.
Hyphal structure: Hyphal system trimitic hyphal, with clamp connections, usually
reddish-brown (8D7–8D8); generative hyphae (1.2-)1.5–2.4–3.0(-3.6) µm broad (n = 30),
thin-walled and hyaline; skeletal hyphae (2.7-)3.2–3.6–4.2(-4.8) µm broad (n = 30), dex-
trinoid, abundant thick wall; binding hyphae (2.5-)3.0–3.5–4.0(-4.4) µm broad (n = 30),
thick wall, branched, usually intertwined the generative and skeletal hyphae, mostly dark
brown near the tube layers, appearing alongside Bovista-type ligative hyphae, hymenial,
sword-like apices at the context. Pileipellis a hymeniderm, dark brown (6D8), composed
of apically acanthus-like branched cells, dextrinoid. Basidiospores mostly ellipsoid to
broadly ellipsoid or oblong with double walls, (5.8-)6.2–7.2–8.4(-9.2) × (5.4-)5.7–5.4–6.8(-7.7)
µm (x = 7.3 × 5.6 µm, n = 50) µm, with Q = 1.48–1.52, L = 7.32 µm, W = 5.68 µm (includ-
ing myxosporium), (4.8-)5.2–6.0–6.7(-7.2) × (4.6-)4.9–5.5–5.7(-6.2) µm (x = 6.2 × 5.6 µm,
n = 50) µm, with Q = 1.08–1.14, L = 6.24 µm, W = 5.67 µm (including myxosporium), over-
laid by hyaline, dextrinoid, echinulae, echinulate brown inner wall, light yellow (4A4–4A5)
to grayish-yellow (4B5–4B6) in 5% KOH. Basidia not seen.
Habitat: Solitary on decaying hardwood of Machilus yunnanensis, living tree of Albizia
mollis and Pinus spp.
Specimens examined: CHINA, Yunnan Province, Kunming Institute of Botany garden,
25◦ 080 3900 N, 102◦ 440 3000 E, 1956 m elev., 31 December 2016, T. Luangharn, HKAS 97411.
Notes: Ganoderma gibbosum belongs to the family Ganodermataceae, which was first
described in Australia [126]. G. gibbosum has been recorded from China [40], India [95],
Korea [92], Laos [30], and Thailand [30]. This species is distinctive in having non-laccate
basidiomes and ellipsoids with double-walled basidiospores [40]. Ganoderma gibbosum has
been reported to cause white rot and several other diseases in hard woods [7] and is widely
distributed in both tropical and temperate areas [4]. It was considered to be a subspecies of
G. applanatum [4], while G. applanatum was the earlier name of G. australe [127]. Ganoderma
australe and G. gibbosum were renamed as G. incrassatum based on their monophyletic
origin [128] since it had been well recognized that G. applanatum was synonymized with
G. applanatum.
J. Fungi 2021, 7, 819 25 of 83
J. Fungi 2021, 7, x FOR PEER REVIEW 27 of 91

Figure 8.
Figure 8. Morphology
Morphology of of Ganoderma
Ganoderma gibbosum
gibbosum (HKAS
(HKAS 97411):
97411): (a,b)
(a,b) upper
upper surface
surface ofof mature
mature basidi‐
basid-
omes; (c) margin; (d) under surface of mature basidiomes; (e) pore characteristics; (f) hyphae of
iomes; (c) margin; (d) under surface of mature basidiomes; (e) pore characteristics; (f) hyphae of tube tube
layers in Melzer’s; (g–i) basidiospores in Melzer’s; (j–l) context hyphae in Congo red. Scale bars:
layers in Melzer’s; (g–i) basidiospores in Melzer’s; (j–l) context hyphae in Congo red. Scale bars:
(b,d) = 3 cm, (c) = 1 cm, (e) = 500 μm, (g–i) = 5 μm, (j–l) = 20 μm.
(b,d) = 3 cm, (c) = 1 cm, (e) = 500 µm, (g–i) = 5 µm, (j–l) = 20 µm.

Ganoderma leucocontextum
Ganoderma leucocontextum T.H.T.H. Li,
Li, W.Q.
W.Q. Deng,
Deng, Sheng
Sheng H.
H. Wu,
Wu, D.M.
D.M. Wang
Wang andand H.P.
H.P.
Hu, Mycotaxon 56: 82 (2015) (Figure
Hu, Mycotaxon 56: 82 (2015) (Figure 9) 9)
Facesoffungi number:
Facesoffungi number: FoF
FoF 06247
06247
Description: Basidiomes flabelliform, subdimidiate,stipitate.
Description: Basidiomes flabelliform, subdimidiate, stipitate. Pileus
Pileus 6–14
6–14 cm
cm in
in length,
length,
4–12 cm in width, and 1–3.2 cm thick. Pileus flabelliform, spathulate, stipitate, subdimid‐
4–12 cm in width, and 1–3.2 cm thick. Pileus flabelliform, spathulate, stipitate, subdimidiate
iate to dimidiate, umbonate, somewhat semicircular, plump, concentrically sulcate zone,
J. Fungi 2021, 7, 819 26 of 83

to dimidiate, umbonate, somewhat semicircular, plump, concentrically sulcate zone, broad

and thick at the base, mostly radial from the center extending to the margin, tough to
break when dried, often thick at the center, slightly soft at the margin, light in weight when
dried, and not woody. Pileus surface convex, furrowed, imbricate, incised, glossy, shiny,
spathulate, shallow sulcate when fresh, umbonate or uneven, usually smooth layers at
center when young to age, non-laccate to weakly laccate when present, strongly laccate
and glossy when mature, weakly laccate where the new hyphae are in active development
(margin), irregularly ruptured crust overlying the context, and tough to break when dried.
Pileus color usually homogenous with orange (6A7) and deep orange (6A8) at the center
toward stipe, extending deep orange (5A8) from the center, slight deep yellow (4A8) where
the new hyphae are in active development when mature, usually red (11B7–11B8) at
the center, and orange (6A7) to deep orange (6A8)–(6B8) extending to the upper margin
surface from mature to old. Context up to 0.3–2.4 cm thick near stipe, white context when
fresh, yellowish-white (1A2) when dried, soft and fibrous, trimitic hyphal, with clamp
connections, hyaline, with walls varying in thickness with simple septa, and unbranched.
Tubes 0.3–1.2 cm in length. Stipe 3–10 cm in length, 4–7 cm in width, sub-cylindrical to
cylindrical, almost stipitate, broad at the base, some presented short stipitate, strongly
laccate with dark brown (8F7–8F8) to grayish ruby (12E6–12E7) when mature, and grayish
brown (8E4) when old. Margin wavy, softer, slippery when young, laccate when mature,
strongly laccate when old, orange yellow (4A7) to deep yellow (4D8) where the new hyphae
are in active development, deep orange (6A7–6A8) to brown (6D8) from mature to old.
Pore 4–6 in number per mm, subcircular, some circular, or angular. Pore surface white
(11A1) when present, yellowish-white (2A2) when mature, brownish-orange (6C7–6C8)
when scratched or bruised, discolored when touched.
Hyphal structure: Hyphal system trimitic, usually golden brown (5D7), yellowish-
brown (5D8) to reddish-brown (8D7–8D8) in KOH; generative hyphae 2.3–5.2 µm broad
(n = 30), thin-walled, hyaline, with clamp connections; skeletal hyphae 2.6–5.5 µm broad
(n = 30), thick-walled, unbranched or rearly branched; binding hyphae 1.8–4.2 µm width
(n = 30), usually thin to thick-walled, branched, hymenial with sword-like apices in the
context. Basidiospores mostly ellipsoid to broadly ellipsoid, double walls, (8.8-)9.3–10.7–
11.4(-12.6) × (6.8-)7.4–8.3–8.7(-9.2) µm (x = 10.5 × 8.4 µm, n = 50) µm, with Q = 1.22–
1.28, L = 10.52 µm, W = 8.41 µm (including myxosporium), (7.8-)8.1–8.5–8.8(-9.1) × (5.2-)
5.7–6.0–6.5(-6.9) µm (x = 8.3 × 6.2 µm, n = 50) µm, with Q = 1.32–1.38, L = 8.34 µm,
W = 6.18 µm (excluding outer myxosporium), overlaid by hyaline, apically echinulae,
truncate, some turgid, vesicular appendix, inner walled echinulate, golden yellow (5B7),
grayish-orange (5D6) to yellowish-brown (5D7), outer walled reddish-brown in 5% KOH.
Cystidia absent. Cultures characteristics white mycelial after incubation at 30 ◦ C for
10 days.
Habitat: Solitary, on the decaying hardwood of unknown tree.
Specimens examined: CHINA, Yunnan Province, Baoshan, 25◦ 090 3500 N, 99◦ 090 4900 E,
1973 m elev., 26 October 2016, J. Xu, HKAS 97401.
Notes: Ganoderma leucocontextum was introduced by Li et al. [62] from the Tibet Au-
tonomous Region of China. This species can be easily recognized by its stipitate, white
context, thick stipe, broadly ellipsoid basidiospores (9–12.5 × 7–9 µm), coarse echinulae,
mostly regular cuticle hyphae, and its deciduous wood habitat [62]. The holotype is similar
to G. lucidum from Europe [62]; however, the illustrated differences in macro-characteristics
of the European G. lucidum are smaller basidiospores (7–12 × 6–8 µm), a deeper-colored
context that is usually rust-colored, becoming dark purple to brown in older portions [105].
Additionally, G. leucocontextum also resembles the widely cultivated G. lucidum (G. lingzhi)
in East Asia [62], but the Chinese G. lucidum has a deeper-colored context and is even darker
near the tube layer, with shorter cutis elements (20–40 × 7–15 µm) and smaller spores
(8–11.5 × 5.5–8.5 µm) (including myxosporium) than G. leucocontextum leucocontextum [5].
Our G. leucocontextum collection from Hainan Province agrees well with the descriptions
provided by Li et al. [62].
J. Fungi 2021, 7, 819 27 of 83
J. Fungi 2021, 7, x FOR PEER REVIEW 29 of 91

Figure 9.
Figure 9. Morphology
MorphologyofofGanoderma
Ganodermaleucocontextum:
leucocontextum:(a)(a)
mature basidiomes
mature of the
basidiomes strain
of the HKAS
strain HKAS
97401; (b,c) mature basidiomes of the strain MFLU 19‐2160; (d) pore characteristics; (e) culture
97401; (b,c) mature basidiomes of the strain MFLU 19-2160; (d) pore characteristics; (e) culture after
after incubation at 25 °C for 14 days; (f) culture after incubation at 25 °C for 21 days; (g,h) context
incubation at 25 ◦ C for 14 days; (f) culture after incubation at 25 ◦ C for 21 days; (g,h) context hyphae
hyphae in KOH; (i–l) basidiospores. Scale bars: (a,e,f) = 2 cm, (b,c) = 4 cm, (d) = 500 μm, (g,h) = 30
in KOH;
μm, (i–l) (i–l) basidiospores. Scale bars: (a,e,f) = 2 cm, (b,c) = 4 cm, (d) = 500 µm, (g,h) = 30 µm,
= 5 μm.
(i–l) = 5 µm.
Ganoderma lucidum (Curtis) P. Karst., Revue Mycologique Toulouse. 3(9): 17 (1881)
Ganoderma lucidum (Curtis) P. Karst., Revue Mycologique Toulouse. 3(9): 17 (1881)
(Figure 10)
(Figure 10)
≡ Boletus rugosus Jacq., Flora Austriaca. 2: 44, f. 169 (1774)
≡ Boletus rugosus Jacq., Flora Austriaca. 2: 44, f. 169 (1774)
≡ Boletus lucidus Curtis, Fl. Londinensis. 4: 72, t. 224 (1781)
≡ Boletus lucidus Curtis, Fl. Londinensis. 4: 72, t. 224 (1781)
≡ Polyporus lucidus (Curtis) Fr., Systema Mycologicum. 1: 353 (1821)
≡ Polyporus lucidus (Curtis) Fr., Systema Mycologicum. 1: 353 (1821)
≡≡Grifola
Grifolalucida
lucida(Curtis)
(Curtis)Gray,
Gray,AAnatural
naturalarrangement
arrangementofofBritish Britishplants.
plants.1:1:644
644(1821)
(1821)
≡ Fomes lucidus (Curtis) Cooke, Grevillea. 13(68):
≡ Fomes lucidus (Curtis) Cooke, Grevillea. 13(68): 118 (1885) 118 (1885)
≡≡Placodes
Placodeslucidus
lucidus(Curtis)
(Curtis)Quél.,
Quél., Enchiridion
Enchiridion Fungorum
Fungorum in Europa
in Europa media
media et praeser‐
et praesertim
tim in Gallia Vigentium.:
in Gallia Vigentium.: 170 (1886)170 (1886)
≡≡Phaeoporus
Phaeoporuslucidus
lucidus(Curtis)
(Curtis)J.J.Schröt.,
Schröt.,Kryptogamen-Flora
Kryptogamen‐Floravon vonSchlesien.
Schlesien.3-1(4):
3‐1(4):491
491
(1888)
(1888)
== Boletus
Boletus flabelliformis
flabelliformis Leyss.,
Leyss., Flora
Flora halensis.:
halensis.: 219
219 (1761)
(1761)
= Agaricus pseudoboletus Jacq., Miscellanea austriaca
= Agaricus pseudoboletus Jacq., Miscellanea austriaca ad ad botanicum,
botanicum, chemiam
chemiam et et histo-
histo‐
riam naturalem spectantia. 1: 26, t. 41
riam naturalem spectantia. 1: 26, t. 41 (1773) (1773)
== Boletus
Boletus obliquatus
obliquatus Bull.,
Bull., Herbier
Herbier dede la
la France.
France. 1:
1: t.t. 77 (1781)
(1781)
== Boletus
Boletus vernicosus
vernicosus Bergeret,
Bergeret, Phytonomatotechnie
Phytonomatotechnieuniverselle.
universelle. 1: 1: 99
99 (1783)
(1783)
== Agaricus
Agaricus lignosus
lignosus Lam.,
Lam., Encyclopédie
Encyclopédie Méthodique,
Méthodique, Botanique.
Botanique. 1-1:
1‐1: 51
51 (1783)
(1783)
= Boletus dimidiatus Thunb., Fl. Japonica.: 348, f. 39 (1784)
J. Fungi 2021, 7, 819 28 of 83

= Boletus dimidiatus Thunb., Fl. Japonica.: 348, f. 39 (1784)

= Boletus castaneus Weber, Suppl. Fl. hols.: 13 (1787)
= Boletus laccatus Timm, Flora megapolitanae Prodomus exhibeus plantas ductatus
Megapolitano.: 269 (1788)
= Boletus crustatus J.J. Planer, Index Plantarum quas in Agro Erfurtensi sponte prove-
nientes.: 280 (1788)
= Agarico igniarium trulla Paulet, Traité des champignons. 2: 95, pl. 10:1-2 (1793)
= Boletus verniceus Brot., Flora Lusitanica. 2: 468 (1804)
= Ganoderma ostreatum Lázaro Ibiza, Revta R. Acad. Cienc. exact. fis. nat. Madr.:
110(1916)
= Ganoderma nitens Lázaro Ibiza, Revta R. Acad. Cienc. exact. fis. nat. Madr.: 104
(1916)
Facesoffungi number: FoF 06250
Description: Basidiomes imbricate, reniform, stipitate. Pileus up to 2–5 cm in length,
2–4 cm in width, and 0.8–2.2 cm thick. Pileus stipitate, reniform, imbricate, irregular,
some laterally, and flabelliform with a contracted, concentrically sulcate zone, irregularly
ruptured crust overlying the context, radial or branched from the center extending to
the margin, tough to break when dried, often thick at the center, slightly soft at margin,
and leathery when aged, tough to break when dried. Pileus surface weakly laccate when
present, strongly laccate and glossy when mature, weakly laccate where the new hyphae
are in active development (margin), smooth layer at the center from young to age, usually
furrowed, incised, undulate to sulcate, somewhat spathulate to uneven, some woody or
corky when old. Pileus color usually yellowish-red (8B7–8B8) at the center, slight to reddish-
orange (7B7–7B8), and orange (6A7–6A8) on upper pileus surface. Context up to 0.4–1.4 cm
thick at the base, abundant thick-walled, subsolid hyphae, concentric lines of various
shade, bearing clamp connections, light brown (6D6) to brown (6D8, 6E8), presenting dark
brown (6F8) melanoid bands. Tube hard, often brown (7D7) to dark brown (7F7). Stipe
up to 8–16 cm in length, up to 0.6–1.8 cm in width, central stipe, cylindrical, thick with
uneven at the base (up to 1.8 cm), usually dark brown (7F7–7F8), laccate, and cracked
when old. Margin often 0.4–1 cm, orange (6A7–6A8) on upper surface, and reddish-yellow
(4A8) under surface, thin and soft than the center. Pore (75–)110–145(–165) µm, circular,
some angular, 4–6 in number per mm. Pore surface white (11A1) to light brown (7D6),
turning brown (7D7–7D8) to dark brown (6F6) when scratched or bruised.
Hyphal structure: Hyphal system trimitic, with clamp connections, hyaline, thin-walled
with abundant thick-walled with simple septa, sparingly branched, swollen by melanoid
bands, usually pale orange (5A3), light orange (5A5), to reddish-orange (8A7–8A8) in KOH;
generative hyphae up to 1.7–3.2 µm broad (n = 30), almost hyaline, usually thin to thick-
walled, with clamp connections, and sparingly branched and flexuous; skeletal hyphae
3.0–6.4 µm broad (n = 30), usually thick-walled with clamp, and abundantly branched and
flexuous; binding hyphae 2.0–5.6 µm broad (n = 30), usually thick-walled with abundant
branches, and occurring melanoid bands. Basidiospores ellipsoid to broadly ellipsoid,
some globose with double walls, with a truncate apex, with double wall, mostly overlaid
by hyaline myxosporium, eusporium bearing fine, short, and distinct, coarse, echinulae,
hyaline, turgid, vesicular appendix, (7.7-)8.4–9.4–10.6(-11.5) × (5.2-)5.9–6.3–7.1(-8.4) µm,
(x = 9.5 × 6.4 µm, n = 50) µm, with Q = 1.47–1.52, L = 9.52 µm, W = 6.34 µm (including myx-
osporium), (6.0-)6.9–7.3–8.1(-8.5) × (4.6-)4.9–5.3–5.8(-6.2) µm (x = 7.5 × 5.2 µm, n = 50) µm,
with Q = 1.41–1.47, L = 7.52 µm, W = 5.24 µm (excluding outer myxosporium), brownish-
orange (6C8), (6D8) to brown (6E5) of endosporium (inner wall) with brown (7E7–7E8)
exosporium (outer wall) in Congo red, brownish-orange (6C8) in 5% KOH, and yellowish-
brown in Melzer’s reagent.
Habitat: Solitary, on decaying hardwood of Quercus sp. in the native forest.
Specimens examined: CHINA, Yunnan Province, Honghe, 23◦ 210 5000 N, 103◦ 220 2400 E,
874 m elev., 15 August 2017, T. Luangharn, MFLU 19-2161.
J. Fungi 2021, 7, 819 29 of 83

Notes: Ganoderma lucidum (Curtis) P. Karst. was originally reported from temperate
England [2]. Previously, it was characterized as Boletus lucidus Curtis and then Polyporus
lucidus (Curtis) Fr. (1821) [1]. The species P. lucidus was characterized by having a laccate
pileus and stipe. The molecular phylogenetic analyses indicated that the G. lucidum from
Europe is not conspecific to the Chinese G. lucidum; thus, the European G. lucidum remained
the true G. lucidum, and the Chinese G. lucidum was proposed as G. lingzhi [4], and most of
the collections named G. lucidum in East Asia were not conspecific with the G. lucidum found
in Europe [129]. Ganoderma lucidum is relatively common in Europe, while its geographic
J. Fungi 2021, 7, x FOR PEER REVIEW
distribution in East Asia, East Africa, Europe, North America, Asia, and other parts of 32 of
the91

world is largely unknown [38].

Figure 10.
Figure 10. Morphology
Morphology of of Ganoderma
Ganoderma lucidum
lucidum (MFLU
(MFLU 19-2161):
19‐2161): (a,b)
(a,b) mature
mature basidiomes;
basidiomes; (c)
(c)pore
pore
characteristics; (d) hyphae of pore characteristics in KOH; (e) hyphae of trama in KOH; (f,g) gener‐
characteristics; (d) hyphae of pore characteristics in KOH; (e) hyphae of trama in KOH; (f,g) generative
ative and skeletal hyphae of context in KOH; (h) hyphae of tube layers in KOH; (i–k) basidiospores.
and skeletal hyphae of context in KOH; (h) hyphae of tube layers in KOH; (i–k) basidiospores. Scale
Scale bars: (a,b) = 2 cm, (c) = 1000 μm, (d) = 150 μm, (e–g) = 20 μm, (h) = 15 μm, (i–k) = 5 μm.
bars: (a,b) = 2 cm, (c) = 1000 µm, (d) = 150 µm, (e–g) = 20 µm, (h) = 15 µm, (i–k) = 5 µm.

Ganoderma
Several multiplicatum
studies of Ganoderma(Mont.) Pat.,
have used theBulletin
name G. delucidum
la Société
forMycologique de France
any laccate Ganoderma
5: 74 (1889) (Figure 11)
species, as Ganoderma are ly highly variable, often resulting in taxonomic and phyloge-
netic ≡ Polyporusespecially
confusion, multiplicatus
withMont., Annales
regards des Sciences
to G. lucidum Naturelles
[38]. The taxonomyBotanique. 1: 128
of Ganoderma
(1854)
has been a constant topic of debate due to the high levels of phenotypic plasticity in its
≡ Fomes multiplicatus (Mont.) Cooke, Grevillea. 14 (69): 18 (1885)
≡ Scindalma multiplicatum (Mont.) Kuntze, Revisio generum plantarum. 3 (2): 519
(1898)
Facesoffungi number: FoF 06251
J. Fungi 2021, 7, 819 30 of 83

species. Several characteristics of Ganoderma are similar to G. lucidum, such as G. mul-

tipileum [39], G. oregonense [42], G. resinaceum [41], G. tsugae, G. lucidum, G. sichuanense,
and G. sinense [4,32,33,35,40,130] from China. Cao et al. [4] have clarified a different new
species, Chinese G. lucidum as G. lingzhi, which has an East Asian distribution. The most
striking characteristics that differentiate G. lucidum from G. lingzhi are the presence of
melanoid bands in the context, a yellow pore surface, and thick dissepiments (80–120 µm)
at maturity [4]. The molecular evidence reveals G. lucidum and G. sinense as two clear
different species [130].
In China, the first-reported G. lucidum was illustrated based on collections from
Guizhou Province [124]. Then, Teng [131] reported more collections from different regions
of China, and many subsequent collections have been reported [4,13,47,121,132]. Recently,
this fungus has been reported to be distributed worldwide based on gross similarity of
features, e.g., in Europe [105], Asia [13,133], America [134,135], and Africa [112]. Our col-
lection from Yunnan Province, China, also agrees well with the descriptions provided from
Asia.
Ganoderma multiplicatum (Mont.) Pat., Bulletin de la Société Mycologique de France
5: 74 (1889) (Figure 11)
≡ Polyporus multiplicatus Mont., Annales des Sciences Naturelles Botanique. 1: 128
(1854)
≡ Fomes multiplicatus (Mont.) Cooke, Grevillea. 14 (69): 18 (1885)
≡ Scindalma multiplicatum (Mont.) Kuntze, Revisio generum plantarum. 3 (2): 519
(1898)
Facesoffungi number: FoF 06251
Description: Basidiomes annual or perennial, stipitate with short base. Pileus 1.5–
7.5 cm in length, 0.5–4 cm in width, and up to 1.5 cm thick at the base. Pileus dimidiate,
flabelliform, reniform, usually flat, convex, imbricate, umbonate or uneven, rarely ungulate,
glabrous when present, often with undefined concentric zones at the center that extend to
the margin, and thick at the base, slightly soft at margin when mature. Pileus surface shiny,
silky, smooth, and soft when young, non-laccate (dull) when mature, hard and woody
when old, frequently furrowed and shallow sulcate, undulating, somewhat spathulate
to uneven on upper surface when mature, covered by irregularly ruptured thick crust,
slightly dull and faded when mature to old, compact and hard when mature, woody to
corky from mature to old. Pileus color usually homogenous with grayish-orange (6B3) at
the center slight to brownish-orange (6C4) and pale orange (6A3), usually yellowish-gray
(4B2) at the margin when mature, and brown (6E8) when dried. Context up to 0.4–1 cm
thick at the base, mostly brown (6E8) to dark brown (7F6–7F8) of cuticle cells, and dark
brown (6F6) melanoid bands, thick-walled, some fibrous pithy context, usually separated
by layers of context tissue at the base. Tube woody hard, often with dark brown (7F7–7F8)
when dried, with sulcate at different levels. Stipe short stipitate, dark brown (7F7), and a
differentiated zone at the point of attachment. Margin up to 1 cm thick, initially white
(5A1), yellowish-gray (4B2) when mature, turns light brown (6D4) to brown (6E8) when
scratched or bruised, often slippery when wet, softer when young, thinner than the center.
Pore 4–7 in number per mm, subcircular to circular, some angular. Pore surface initially
white (7A1) to yellowish-white (1A2), becoming pale orange (5A3) when mature, light
brown (7D6) to brown (7D8) when handled, scratched, and bruised.
Hyphal structure: Hyphal system trimitic; generative hyphae 2.1–4.8 µm (x = 2.2,
n = 30) in diam, clamp, almost hyaline, thin to thick-walled, composed of narrow and
spare branches; skeletal hyphae 3.2–6.5 µm width (n = 30), usually thick-walled, hya-
line, some branched and intertwined hyphae; binding hyphal 2.4–5.7 µm width (n = 30),
thick-walled, many branches, and comprised Bovista-type ligative hyphae. Pileipellis a
hymeniderm, brown (6E8), composed of apically acanthus-like branched cells, dextrinoid.
Basidiospores mostly ellipsoid with double walls, (7.8-)8.7–10.8–12.2(-13.3) × (6.9-)7.4–
9.1–10.0(-10.7) µm, (x = 10.7 × 9.1 µm, n = 50) µm, with Q = 1.15–1.22, L = 10.79 µm,
W = 9.13 µm (including myxosporium), (5.4-)5.9–6.6–7.1(-7.7) × (4.9-)5.4–5.8–6.2(-6.7) µm
J. Fungi 2021, 7, 819 31 of 83

(x = 6.6 × 5.8 µm, n = 50) µm, with Q = 1.11–1.17, L = 6.64 µm, W = 5.82 µm (excluding
outer myxosporium), inner walled deep orange (6A8), brownish-orange (7C7–7C8) in KOH,
outer walled dark brown (7E6), dark brown (7E8) to reddish-brown (8E8) in KOH. Cutis
usually composed of clavate cells.
Habitat: Solitary on stump of Quercus spp.
Specimens examined: CHINA, Yunnan Province, Jinning District, 24◦ 410 1700 N, 102◦ 130 1500 E,
1912 m elev., 8 October 2017, T. Luangharn, MFLU 19-2152.
Notes: Ganoderma multiplicatum was originally collected from French Guyana [2].
This species has a distinctive form with its reddish-black pileus, a not fully homoge-
nous context, tuberculate hyphal ends in cuticle cells, with small subglobose to broadly
ellipsoid basidiospores (7–8 × 5–6 µm) [7,136–138]. Ganoderma multiplicatum has been con-
sidered most similar to G. chalceum [113], also considered a synonym of G. subamboinense
Henn. [136], but Correia de Lima et al. [139] illustrated that G. chalceum and G. subam-
boinense are in different clades, suggesting they are not synonymous. Our G. multiplicatum
specimen was collected from Yunnan Province, China. It is similar to the original de-
scription, showing ellipsoid basidiospores, while sub-globous basidiospores could not be
observed. This species has been reported from Africa [140], Asia [61], China [13,61,119,122],
J. Fungi 2021, 7, x FOR PEER REVIEW 34 of 91
India [141], Myanmar [30], Taiwan, PRC [122], and neotropical regions of Brazil, Colombia,
and Venezuela [138].

Figure11.
Figure 11. Morphology
Morphology of of Ganoderma
Ganodermamultiplicatum
multiplicatum(MFLU
(MFLU19-2152):
19‐2152):(a)
(a)mature
maturebasidiomes;
basidiomes;(b)(b)pore
pore
characteristics;(c)
characteristics; (c)hyphae
hyphaeofoftube
tube layers
layers in
in KOH;
KOH; (d–f)
(d–f) context
context hyphae
hyphae inin KOH;
KOH; (g–i)
(g–i) basidiospores.
basidiospores.
Scale bars: (a) = 2 cm, (b) = 1000 μm, (d–f) = 20 μm, (g–i) = 5 μm.
Scale bars: (a) = 2 cm, (b) = 1000 µm, (d–f) = 20 µm, (g–i) = 5 µm.

Ganoderma resinaceum
Ganoderma resinaceumBoud.,Boud.,Bulletin
Bulletinde
delalaSociété
SociétéMycologique
Mycologiquede deFrance.
France. 5;5;72
72
(1889)(Figure
(1889) (Figure12) 12)
≡Fomes
≡ Fomesresinaceus
resinaceus(Boud.)
(Boud.)Sacc.,
Sacc.,Sylloge
SyllogeFungorum.
Fungorum.9:9:179
179(1891)
(1891)
≡ Scindalma resinaceum (Boud.) Kuntze, Revisio generum plantarum.
≡ Scindalma resinaceum (Boud.) Kuntze, Revisio generum plantarum. 3(2); 3(2);519
519(1898)
(1898)
≡ Friesia resinacea (Boud.) Lázaro Ibiza, Revta R. Acad. Cienc. exact. fis. nat.
≡ Friesia resinacea (Boud.) Lázaro Ibiza, Revta R. Acad. Cienc. exact. fis. nat. Madr.: Madr.:
591 (1916)
591 (1916)
≡Ganoderma
≡ Ganodermalucidum
lucidumsubsp.
subsp.resinaceum
resinaceum(Boud.)
(Boud.) Bourdot
Bourdot and
and Galzin,
Galzin, Bulletin
Bulletin de
delala
SociètèMycologique
Sociètè Mycologiquede deFrance.
France. 41;
41; 177
177 (1925)
(1925)
≡Ganoderma
≡ Ganodermalucidum
lucidum var.
var. resinaceum
resinaceum (Boud.)
(Boud.) Maire,
Maire, Fungi
Fungi Catalaunici:
Catalaunici: Contribu‐
Contributions
átions á lètude
lètude de la Mycologique
de la Flore Flore Mycologique de la Catalogne:
de la Catalogne: 38 (1933)
38 (1933)
= Ganoderma chaffangeonii Pat., Bulletin de la Société Mycologique de France. 5: 74
(1889)
= Polyporus polychromus Copel., Annales Mycologici. 2 (6): 507 (1904)
= Ganoderma praelongum Murrill, North American Flora. 9 (2): 121 (1908)
= Ganoderma argillaceum Murrill, North American Flora. 9 (2): 122 (1908)
J. Fungi 2021, 7, 819 32 of 83

= Ganoderma chaffangeonii Pat., Bulletin de la Société Mycologique de France. 5: 74

(1889)
= Polyporus polychromus Copel., Annales Mycologici. 2 (6): 507 (1904)
= Ganoderma praelongum Murrill, North American Flora. 9 (2): 121 (1908)
= Ganoderma argillaceum Murrill, North American Flora. 9 (2): 122 (1908)
= Ganoderma pulverulentum Murrill, North American Flora. 9 (2): 121 (1908)
= Ganoderma subperforatum G.F. Atk., Botanical Gazette Crawfordsville. 46 (5): 337
(1908)
= Ganoderma areolatum Murrill, Bulletin of the New York Botanical Garden. 8: 149
(1912)
= Mensularia vernicosa Lázaro Ibiza, Revista de la Real Academia de Ciencias Exactas
Fisicas y Naturales Madri. 14: 740 (1916)
= Ganoderma subtuberculosum Murrill, Lloydia. 7 (4): 326 (1945)
Facesoffungi number: FoF 06252
Description: Basidiomes annual, perennial, short stipitate. Pileus 1.5–12.5 cm in length,
1–7 cm in width, and up to 2 cm thick at the base. Pileus dimidiate, flabelliform, reniform,
convex, imbricate, umbonate or uneven, some ungulate, concentric zones at the center
that extend to the margin, broadly attached, thick at the base, slightly soft at the margin
when mature. Pileus surface glossy, shiny, silky, smooth, and soft when young, laccate
when mature, furrowed and shallow sulcate, undulating, somewhat spathulate to uneven
on upper surface when mature, covered by irregularly ruptured thin crust, slightly dull
and faded when mature to old, compact and hard when mature, woody to corky when
mature to old. Pileus color reddish-brown (10E7–10E8) at the center, slight to yellowish-red
(8B7–8B8), reddish-orange (7A7–7A8), and light orange (5A5–5A6) closed to the margin,
and white (4A1) at the margin. Context up to 0.4–1 cm thick at the base, mostly grayish-
yellow (4C6) to dark brown (7F6–7F8) cuticle cells, and dark brown (6F6) melanoid bands,
thick-walled, some fibrous pithy context, usually separated by layers of context tissue at
the base. Tube woody hard, often dark brown (7F7–7F8) when dried, concolorous with
pore surface, and sulcate at different levels. Stipe short stipitate, usually reddish-brown
(10E7–10E8), and a differentiated zone at the point of attachment. Margin up to 1.5 cm
thick, initially white (5A1), yellowish-gray (4B2) when mature, turning light brown (6D4)
to brown (6E8) when scratched or bruised, often slippery when wet, softer when young,
thinner than the center. Pore 4–7 in number per mm, angular to circular. Pore surface
initially white (7A1) to yellowish-white (1A2), becoming light orange (5A5) when mature,
light brown (7D6) to brown (7D8) when handled, scratched, or bruised.
Hyphal structure: Hyphal system trimitic; generative hyphae 2.1–4.7 µm (x = 3.6, n = 30)
in diam, clamp, almost hyaline, thin-walled, composed of sparse branches; skeletal hyphae
3.2–6.2 µm width (n = 30), usually thick-walled, hyaline, some branched and intertwined
hyphae; binding hyphal 2.8–5.1 µm width (n = 30), thick-walled and occasionally thick-
walled, without septate hyphae, many branches, and composed of Bovista-type ligative
hyphae. Basidiospores mostly ellipsoid with double walls, (7.6-)8.4–9.4–10.5(-11.3) × (6.5-)
7.1–8.4–9.0(-9.8) µm, (x = 9.3 × 8.2 µm, n = 50) µm, with Q = 1.10–1.16, L = 9.31 µm,
W = 8.24 µm (including myxosporium), (6.5-)7.1–8.2–9.1(-9.8) × (4.8-)5.3–5.7–6.8(-7.3) µm
(x = 8.1 × 5.6 µm, n = 50) µm, with Q = 1.42–1.48, L = 8.13 µm, W = 5.62 µm (excluding
outer myxosporium), inner walled orange (5A6) to deep orange (5A7–5A8, 6A8) in KOH
and grayish brown (5C5–5C6) in Melzer’s reagent, outer walled dark brown (7E6–7E8) to
reddish-brown (8E8) in KOH and light brown (6D5–6D6) to brown (6D7–6D8) in Melzer’s
reagent.
Habitat: Solitary, on living tree of Albizia mollis (Wall.) Boiv.
Specimens examined: CHINA, Yunnan Province, Kunming Institute of Botany, 25◦ 080 3900 N,
102 440 3000 E, 1962 m, 12 July 2017, T. Luangharn, MFLU 19-2153.
◦

Notes: Ganoderma resinaceum was introduced by Boudier in 1889 from France [41].
This species has also been described by Steyaert [110] and Ryvarden and Gilbertson [105].
Ganoderma resinaceum is distinctively characterized by variable pileus coloration, a fi-
J. Fungi 2021, 7, 819 33 of 83

brous spongy homogeneous context, larger basidiospores, and an amyloid pileipellis [7].
This species is considered to have characteristics similar to G. pfeifferi in its upper crust
resinous layers. However, this species has a dark brown to umber context and wider
spores. In addition, G. resinaceum also shares similarities with G. lucidum, while G. lucidum
possesses a varying light context without a dark zone above the tubes and no resinous layer
on the crust [105,140]. Ganoderma resinaceum was evaluated to the species complex base on
molecular evidence [38], but in the phylogenetic analysis, it cannot be distinguished from
G. lucidum [142]. However, several researchers suggested that G. resinaceum differs from
G. lucidum [4,76,143].
Ganoderma sanduense Hapuar., T.C. Wen and K.D. Hyde, Mycosphere 10, 274 (2019)
Taxonomy and phylogenetic analyses are shown in Hapuarachchi et al. [30].
Notes: Ganoderma sanduense is characterized by its ferruginous laccate pileus, orbicular,
strongly laccate, several layers thick, basidiospores 12.1–13.8 × 9.2–10.5 µm, relatively
large broadly ellipsoid to ellipsoid basidiospores, with a light brown eusporium bearing
J. Fungi 2021, 7, x FOR PEER REVIEW
fine, hyaline, short, and distinct echinulae. This fungus is solitary on rotten wood in 36 dry
of 91
dipterocarp forests and in upper-mixed deciduous forests from Guizhou Province, China.

Figure12.
Figure 12. Morphology
Morphology ofof Ganoderma
Ganodermaresinaceum
resinaceum(MFLU
(MFLU19-2153):
19‐2153):(a–c)
(a–c)mature
maturebasidiomes;
basidiomes;(d)
(d)pore
pore
characteristics; (e) hyphae of pore in KOH; (f) hyphae of tube layers in KOH; (g,h) context hyphae
characteristics; (e) hyphae of pore in KOH; (f) hyphae of tube layers in KOH; (g,h) context hyphae
in KOH; (i) basidiospores in Melzer’s reagent; (j,k) basidiospores in KOH. Scale bars: (a–c) = 4 cm,
in KOH; (i) basidiospores in Melzer’s reagent; (j,k) basidiospores in KOH. Scale bars: (a–c) = 4 cm,
(d) = 1000 μm, (e) = 150 μm, (f) = 30 μm, (g,h) = 50 μm, (i–k) = 5 μm.
(d) = 1000 µm, (e) = 150 µm, (f) = 30 µm, (g,h) = 50 µm, (i–k) = 5 µm.
Ganoderma sanduense Hapuar., T.C. Wen and K.D. Hyde, Mycosphere 10, 274 (2019)
Taxonomy and phylogenetic analyses are shown in Hapuarachchi et al. [30].
Notes: Ganoderma sanduense is characterized by its ferruginous laccate pileus, orbicu‐
lar, strongly laccate, several layers thick, basidiospores 12.1–13.8 × 9.2–10.5 μm, relatively
large broadly ellipsoid to ellipsoid basidiospores, with a light brown eusporium bearing
J. Fungi 2021, 7, 819 34 of 83

Ganoderma sichuanense J.D. Zhao and X.Q. Zhang, Acta mycol. sin.: 159 (1983)
(Figure 13)
Facesoffungi number: FoF 06248
Description: Basidiomes annual or perennial, stipitate. Pileus 0.5–3.2 cm in length,
0.5–3 cm in width, up to 1 cm thick at the base. Pileus reniform to circular, or subflabellate
when seen from above, concentrically sulcate zones with turberculate, glabrous when youth
to maturity, bumps when mature, often tough to break when dried, often with undefined
concentric zones at the center that extend to the margin, thick at the center, slightly soft
at the margin. Pileus surface shiny, silky, smooth, and soft when young, hard and woody
old, frequently furrowed and shallow sulcate on upper surface, undulating, somewhat
spathulate to uneven when mature, covered by irregularly ruptured thin crust, and strongly
laccate from mature to old. Pileus color usually homogenous with yellowish-red (8A7–8A8)
at the center, slight reddish-orange (7A7–7A8), and reddish-brown (8E8) at the deep-sulcate
margin from mature to old. Context up to 0.2–0.8 cm thick at the base, some thin-walled,
with abundant thick-walled to subsolid hyphae, containing fibrous pithy context, bearing
clamp connections, with dark brown (7F7) melanoid bands occurring. Tube hard and
woody, thin-walled, frequently branched, with clamped connection, and often dark brown
(7F7–7F8) when dried. Stipe up to 3–812 cm in length, up to 0.3–1 cm in width, centrally
stipitate, almost sub-cylindrical to cylindrical, concolorous with the pileus, often reddish-
brown (8E7–8E8), and strongly laccate from mature to old. Margin soft when young,
strongly laccate when mature, some wavy, slippery when wet, smooth, softer, thinner than
the base, and soft than the center, often deep orange (5A8) to golden yellow (5B7–5B8) from
mature to old. Pore 4–6 in number per mm, subcircular to circular, sometimes angular.
Pore surface initially white (11A1), pale yellow (3A3) to yellow (3A7) when mature, turns
light brown (7D5), brown (7D7–7D8) to dark brown (7F6–7F8) when scratched or bruised,
becoming discolored when touched.
Hyphal structure: Hyphal system trimitic, with clamp connections, hyaline, thin to
thick-walled, tapering at branch, sometimes swollen at the attachment point, composed
of some narrow hyphae; generative hyphae (1.3–)1.8–2.3–2.6(–2.8) µm broad (n = 30),
hyaline, and thin-walled; skeletal hyphae (2.1–)2.5–3.9–4.8(–5.2) µm broad (n = 30) abun-
dant with walls varying in thickness, unbranched, sometimes subsolid; binding hyphae
(1.7–)2.1–2.8–3.6(–4.3) µm broad (n = 30), usually with walls varying in thickness, narrow
to subsolid, usually presenting as orange white (6A2), pale orange (6A3) to light orange
(5A5) of thin-walled, and pale red (6A3) of thick-walled, with subsolid in KOH. Pileipellis
a hymeniderm, light brown (6D6), clavate-like cells, dextrinoid. Basidiospores mostly
ellipsoid, some oblong with double walls, (8.0-)8.6–9.6–10.5(-11.0) × (6.2-)6.7–8.4–9.6(-10.1)
µm (x = 9.5 × 8.3 µm, n = 50) µm, Q = 1.11–1.17, L = 9.49 µm, W = 8.31 µm (including myx-
osporium), (7.0-)7.4–7.9–8.5(-9.0) × (4.2-)4.5–5.6–5.9(-6.4) µm (x = 7.8 × 5.6 µm, n = 50) µm,
with Q = 1.36–1.41, L = 7.80 µm, W = 5.62 µm (excluding outer myxosporium), overlaid
by hyaline, dextrinoid, echinulae, inner wall echinulate with grayish-orange (6B5–6B6) to
brownish-orange (7D4–7D5), and outer walled usually dark brown (7E8–7E8) to reddish-
brown (8E6–8E8) in KOH.
Habitat: Solitary on the living tree of Graucoides schotky.
Specimens examined: CHINA, Yunnan Province, Xishan Forest Park, 24◦ 570 5300 N,
102 530 1000 E, 2013 m elev., 29 October 2016, T. Luangharn, HKAS 97398.
◦

Notes: Ganoderma sichuanense was originally described from the Sichuan Province,
China, in 1983 [5]. However, G. sichuanense was published in 1983 [40] but has not been
widely used. Ganoderma sichuanense was verified as “G. lucidum” (Lingzhi) based on both
morphological and molecular data. This fungus was distinguished from other Ganoderma
species. Ganoderma sichuanense was characterized by its distinctive substipitate to stipitate,
flabellate to reniform, radially rugose pileus, laccate with a verrucose or tuberculose upper
surface, pore surface yellowish when young, becoming brown or black when bruised,
and small spores [13]. Originally the basidiospores were described as (7.4–9.5 × 5–7)
µm [40], then updated to (7.8–10.4 × 5.2–6.4) µm [13,61], and (9–11.5 × 6.5–8) µm [5].
J. Fungi 2021, 7, 819 35 of 83

The study basidiospores were 7.8–9.5 × 5.6–8.3 µm, which is in the range of the original
report, which is not distinct from those of basidiospores found in other reports. Cao et al. [4]
stated that G. sichuanense differs from G. lingzhi as its sessile basidiocarps and smaller basid-
iospores (7.4–9.2 × 5–6.6) µm, with distinctive yellow context, thick dissepiments, absence
of concentric growth zones in the context, basidiospore size, yellow pore surface, and pres-
ence of melanoid bands upon maturity [4,37]. Ganoderma sichuanense was yellowish-brown,
with a dark brown eusporium bearing thick echinulae, overlaid by a hyaline myxosporium.
However, among the Chinese Ganoderma species, G. flexipes, G. multipileum, G. sichuanense,
J. Fungi 2021, 7, x FOR PEER REVIEW
G. tropicum, and G. tsugae are the most similar species to G. lingzhi because they share 38 of 91a

reddish-brown pileal surface, similar basidiospores, and cuticle cells [4].

Figure13.
Figure 13.Morphology
MorphologyofofGanoderma
Ganodermasichuanense
sichuanense(HKAS
(HKAS 97398):
97398): (a–c)
(a–c) mature
mature basidiomes;
basidiomes; (d)(d) hy‐
hyphae
phae of trama in KOH; (e,f) context hyphae in KOH; (g) pore characteristics; (h) hyphae of pore in
of trama in KOH; (e,f) context hyphae in KOH; (g) pore characteristics; (h) hyphae of pore in KOH;
KOH; (i,j) basidiospores. Scale bars: (a,b) = 1 cm, (c) = 3 cm, (d) = 20 μm, (e,f) = 30 cm, (g) = 1000 μm,
(i,j) basidiospores. Scale bars: (a,b) = 1 cm, (c) = 3 cm, (d) = 20 µm, (e,f) = 30 cm, (g) = 1000 µm,
(i,j) = 5 μm.
(i,j) = 5 µm.
Ganoderma sinense J.D. Zhao, L.W. Hsu and X.Q. Zhang, Acta Mycologica Sinica. 19:
272 (1979) (Figure 14)
= Ganoderma formosanum T.T. Chang and T. Chen. Transactions of the British Myco‐
logical Society. 82(4): 731 (1984)
J. Fungi 2021, 7, 819 36 of 83

Ganoderma sinense J.D. Zhao, L.W. Hsu and X.Q. Zhang, Acta Mycologica Sinica. 19:
272 (1979) (Figure 14)
= Ganoderma formosanum T.T. Chang and T. Chen. Transactions of the British Mycolog-
ical Society. 82(4): 731 (1984)
Facesoffungi number: FoF 06253
Description: Basidiomes annual, stipitate, subdimidiate. Pileus 2–6 cm in length,
2–4 cm in width, and 0.3–1 cm thick. Pileus stipitate, subdimidiate to dimidiate, flabelli-
form, spathulate, umbonate, radial from the center extending to the margin, tough to break
when dried, often thick at the center, slightly soft at the margin, light in weight when dried,
and without woody. Pileus surface laccate, convex, some radial furrowed to furrowed, im-
bricate, incised, glossy, shiny, spathulate, shallow sulcate when fresh, umbonate or uneven,
strongly laccate and glossy when mature, and weakly laccate where the new hyphae are
in active development (margin), usually smooth layers at the center when young to age,
irregularly ruptured crust overlying the context, and leathery when age when break. Pileus
color usually homogenous with brownish-red (8C7–8C8) to reddish-brown (8D7–8D8)
at the center toward stipe, extending brownish-red (9C8) from the center, slight to the
margin when mature, usually reddish-brown (8E5–8E8) upper margin surface when old.
Context up to 0.3–1 cm thick near stipe, dry, upper layer brownish-orange (6C8) when
fresh, grayish-orange (5B5) at lower layers, with dark brown (8F7) when dried, soft and
fibrous, covered with thin crust, some present woody, trimitic hyphal, hyaline, thin to
thick-walled with simple septa, with branched. Tube 0.3–0.6 cm in length, brown (7D8).
Stipe 4–16 cm in length, sub-cylindrical to cylindrical, almost stipitate with broadly and
thick at the base, irregularly ruptured crust overlying, usually strongly laccate with brown
(7D8) to dark brown (8F8) when mature, and dark brown (8F8) when old. Margin soft,
some wavy, laccate when mature, weakly laccate when old, brownish-orange (6D8) when
mature to old. Pore 4–6 in number per mm, subcircular to circular. Pore surface white
(11A1) to yellowish-white (2A2) when mature, discolored when touched, brownish (6E7)
to dark brown (6F7) when scratched or bruised.
Hyphal structure: Hyphal system trimitic, with clamp connections, usually light or-
ange (5A5), orange (5A7), golden yellow (5B7–5B8), sometimes brownish-red (8C7) in
KOH; generative hyphae 1.3–2.4 µm broad (n = 30), hyaline, thin-walled, with clamp
connections; skeletal hyphae 3.1–5.2 µm broad (n = 30), usually hyaline, thick-walled,
unbranched, and solid; binding hyphae 2.9–5.2 µm width (n = 30), thin to thick-walled,
with branched, hymenial with sword-like apices in the context. Basidiospores mostly ellip-
soid to broadly ellipsoid, with double walls, with size range of (9.5-)10.2–11.4–12.3(-13.1)
× (7.0-)7.6–8.4–9.3(-10.2) µm (x = 11.2 × 8.5 µm, n = 50) µm, with Q = 1.28–1.36, L = 11.24
µm, W = 8.50 µm (including myxosporium), (8.3-)9.4–10.3–11.5(-12.3) × (6.1-)6.7–7.1–7.5
(-8.0) µm (x = 10.2 × 7.2 µm, n = 50) µm, with Q = 1.41–1.47, L = 10.32 µm, W = 7.13 µm
(excluding outer myxosporium), overlaid by hyaline, apically, short echinulae, truncate,
some turgid vesicular appendix, inner wall echinulate, orange (5A7), deep orange (5A8,
5B8), orange (6B8), with brownish-orange (6B8), outer wall usually brownish-red (8C7–8C8)
in 5% KOH.
Habitat: Solitary on decaying and living tree of Albizia mollis (Wall.) Boiv., living tree
Quercus sp.
Specimens examined: CHINA, Yunnan Province, Baoshan, 25◦ 090 3500 N, 99◦ 090 4900 E,
1973 m elev., 11 November 2017, T. Luangharn, MLFU 19-2173.
Notes: Ganoderma sinense was described from China, characterized by a uniformly
brown to dark brown context and slightly longitudinally crested basidiospores [122].
This species was considered as a species with high phenotypic [13,36]. Our G. sinense
collection was obtained from Yunnan Province, China, and agrees well with the description
of the holotype as described by Wang and Wu [122]. Several reports have also illustrated
G. sinense from China [13,47,123]. Ganoderma sinense is considered to have characteristics
similar to G. lucidum, while G. sinense illustrates differences in macro- characteristics
in its thin pileus, long stipes, and rarely branched skeletal hyphae with Bovista-type
J. Fungi 2021, 7, 819 37 of 83

binding hyphae [106], and these two different species are distinguished in reports [130].
Ganoderma sinense is also reported to have similar characteristics with G. formosanum; hence,
J. Fungi 2021, 7, x FOR PEER REVIEW
G. formosanum was treated as synonymous, and consequently, the earliest used valid 40 of 91
name
was G. sinense [122].

Figure14.
Figure 14. Morphology
Morphology of of Ganoderma
Ganodermasinense
sinense(MFLU
(MFLU19-2173):
19‐2173):(a–c)
(a–c)mature
maturebasidiomes;
basidiomes;(d–f)
(d–f)context
context
hyphae in
hyphae in KOH;
KOH; (g)
(g) pore
porecharacteristics;
characteristics;(h–j)
(h–j)basidiospores in in
basidiospores KOH. Scale
KOH. bars:
Scale (a–c)(a–c)
bars: = 2 cm,
= 2(d–f)
cm,
= 20 μm, (g) = 1000 μm, (h–j) = 5 μm.
(d–f) = 20 µm, (g) = 1000 µm, (h–j) = 5 µm.

Ganoderma tsugae
Ganoderma tsugae Murrill,
Murrill, Bulletin
Bulletinofofthe
theTorrey
TorreyBotanical
BotanicalClub. 29; 601
Club. 29; (1902) (Fig‐
601 (1902)
ure 15) 15)
(Figure
≡Fomes
≡ Fomestsugae
tsugae(Murrill)
(Murrill)Sacc.
Sacc.and
andD.D.Sacc.,
Sacc.,Sylloge
SyllogeFungorum.
Fungorum.17: 17:123
123(1905)
(1905)
≡ Polyporus tsugae (Murrill) Overh.: 714 (1915)
≡ Polyporus tsugae (Murrill) Overh.: 714 (1915)
== Polyporus
Polyporus metallicus
metallicus Lloyd,
Lloyd, Mycological
Mycological Writings.
Writings. 66 65):
65): 1099
1099 (1920)
(1920)
Facesoffunginumber:
Facesoffungi number: FoF
FoF 06254
06254
Description: Basidiomes annual, subdimidiate, stipitate. Pileus 2–16 cm in length, 2–9
cm in width, and 0.5–3 cm thick at the base. Pileus stipitate, subdimidiate to dimidiate,
flabelliform, spathulate, umbonate, concentrically sulcate zone, radial from the center ex‐
tending to the margin, tough to break when dried, often thick at the center, slightly soft at
the margin, light in weight when dried, with woody or corky when dried. Pileus surface
J. Fungi 2021, 7, 819 38 of 83

Description: Basidiomes annual, subdimidiate, stipitate. Pileus 2–16 cm in length,

2–9 cm in width, and 0.5–3 cm thick at the base. Pileus stipitate, subdimidiate to dimidiate,
flabelliform, spathulate, umbonate, concentrically sulcate zone, radial from the center
extending to the margin, tough to break when dried, often thick at the center, slightly soft
at the margin, light in weight when dried, with woody or corky when dried. Pileus surface
laccate, convex, radial furrowed, imbricate, incised, glossy, shiny, spathulate, shallow
sulcate, umbonate or uneven, strongly laccate and glossy when mature, and weakly laccate
where the new hyphae are in active development (margin), usually smooth layers at the
center when young to age, irregularly rugose, irregularly ruptured, thin crust overlying
the context, tough to break when dried. Pileus color usually homogenous with brownish-
red (8C7–8C8) to reddish-brown (8D7–8D8) at the center toward the stipe and margin
surface when mature to old. Context up to 0.4–2.2 cm thick near the stipe, brownish-orange
(7C7–7C8) to brown (7D8) on the upper layers, brownish-red (8C6) when dried, soft and
fibrous, covered with thin crust, some present woody, dimitic hyphal, hyaline, thin-walled
with simple septa, branched. Tube 0.3–1.6 cm in length, with dark brown (7F5). Stipe
4–10 cm in length, 2 cm thick, sub-cylindrical to cylindrical, almost stipitate and broad and
thick at the base, irregularly ruptured crust overlying, usually strongly laccate with brown
(7D8) to dark brown (8F8) when mature, usually dark brown (8F8) when old. Margin soft,
some wavy, laccate when mature, and strong laccate when old, brownish-red (8C7–8C8)
to reddish-brown (8D7–8D8) from mature to old. Pore 4–6 in number per mm, circular or
angular. Pore surface yellowish-white (4A2) when present to yellowish-white (2A2) when
mature, discolored when touched, brownish (6E7) when scratched or bruised.
Hyphal structure: Hyphal system trimitic, with clamp connections; generative hy-
phae 3.1–4.8 µm broad (n = 30), hyaline, thin-walled, with clamp connections; skeletal
hyphae 3.1–6.8 µm broad (n = 30), usually hyaline, thick-walled, non-septate, unbranched,
and solid; binding hyphae 3.9–5.0 µm width (n = 30), with walls varying in thickness,
with many branches, some hymenial with sword-like apices in the context. Basidiospores
mostly ellipsoid to broadly ellipsoid, with double walls, with size range of (9.7-)10.6–
12.7–14.3(-15.8) × (7.3-)8.4–10.7–11.5(-12.4) µm (x = 12.7 × 10.5 µm, n = 50) µm, with
Q = 1.18–1.24, L = 12.68 µm, W = 10.48 µm (including myxosporium), (8.3-)9.4–10.8–12.6
(-13.1) × (6.1-)6.9–7.6–8.3(-9.2) µm (x = 10.7 × 7.6 µm, n = 50) µm, with Q = 1.36–1.45,
L = 10.68 µm, W = 7.59 µm (excluding outer myxosporium), overlaid by hyaline, apically
and short echinulae, truncate and turgid vesicular appendix, inner walled echinulate,
brownish-orange (6C7–6C8), outer walled usually dark-brownish (6F7–6F8) in 5% KOH.
Habitat: Solitary, on decaying Quercus spp. tree.
Specimens examined: CHINA, Yunnan Province, Jinning District, 24◦ 410 1700 N, 102◦ 130 1500 E,
1973 m elev., 11 November 2017, JC. Xu, HKAS 97406.
Notes: Ganoderma tsugae has been treated as a synonym of G. lucidum [144–146].
This fungus is characterized by a laccate and concentric yellowish-red pileus, stipitate,
fan-shaped, sulcated with a yellow margin, ovoid, verrucose, and truncated basidiospores.
G. tsugae is widely distributed across the USA [35,109]. The phylogenic analysis supported
G. tsugae as an independent species distinct from G. lucidum, as it grows exclusively on
conifers, especially on Tsuga and Abies species, while G. lucidum inhabits mostly angiosper-
mous trees [76]. According to Loyd et al. [35], G. tsugae is similar to G. oregonense as
they share a distinctly white context tissue, rough basidiospores, and are predominately
associated with conifers decay.
J. Fungi 2021, 7, 819 39 of 83
J. Fungi 2021, 7, x FOR PEER REVIEW 43 of 93

Figure15.
Figure 15.Morphology
Morphologyof ofGanoderma
Ganoderma tsugae (HKAS 97406):
tsugae (HKAS 97406):(a,b)
(a,b) mature
maturebasidiomes;
basidiomes;(c) (c)
hyphae
hyphaeof tube
of
tube layers; (d–g) context hyphae in KOH; (h) pore characteristics; (i,j) basidiospores. Scale =bars:
layers; (d–g) context hyphae in KOH; (h) pore characteristics; (i,j) basidiospores. Scale bars: (a,b) 4 cm,
(c) = =
(a,b) (c) == 30
cm,(d–g)
304cm, 30 cm, (h) = 500
µm,(d–g) µm,
= 30 µm, = 5=µm.
(i,j)(h) 500 µm, (i,j) = 5 µm.

3.2.2.
3.2.2.Taxonomy
Taxonomyof ofGanoderma
GanodermafromfromLaos
Laos
Ganoderma adspersum
Ganoderma adspersum (Schulzer)Donk
(Schulzer) DonkProc.
Proc. K. Ned.Akad.
K. Ned. Akad.Wet.,
Wet.,Ser.
Ser.C,C, Biol.
Biol. Med.
Med. Sci.
Sci. 72(3):
72(3): 273 273 (1969)
(1969) (Figure
(Figure 16) 16)
≡≡ Polyporus adspersus Schulzer, Flora.: 11 (1878)
Polyporus adspersus Schulzer, Flora.: 11 (1878)
= Polyporus linhartii Kalchbr., Fungi Hong. 252 (1884)
= Polyporus linhartii Kalchbr., Fungi Hong. 252 (1884)
= Ganoderma europaeum Steyaert, Bulletin du Jardin Botanique de l'État à Bruxelles. 31:
= Ganoderma europaeum Steyaert, Bulletin du Jardin Botanique de l'État à Bruxelles. 31:
70 (1961)
70 (1961)
Facesoffungi number: FoF 06241
Facesoffungi
Description: number: FoF
Basidiomes 06241 subdimidiate, sessile. Pileus 3–22 cm in length,
annual,
2–14 cmDescription:
broad, and Basidiomes annual,
1–4 cm thick subdimidiate,
at the base. Pileussessile.
sessile,Pileus 3–22 cm
perennial, in length, 2–14
subdimidiate to
dimidiate,
cm broad,flabelliform,
and 1–4 cm spathulate,
thick at the umbonate,
base. Pileusconcentrically sulcate
sessile, perennial, zone, somewhat
subdimidiate round
to dimidiate,
flabelliform, spathulate, umbonate, concentrically sulcate zone, somewhat round and
J. Fungi 2021, 7, 819 40 of 83

and plump when young, somewhat imbricate with flabelliform (fan-shaped) when seen
from above, broadly attached, radial from the center extending to the margin, tough to
break when dried, thick at the base, slightly soft at the margin when mature, light in
weight with woody or corky when dried. Pileus surface non-laccate (dull), convex, radially
furrowed, incised, spathulate, shallow sulcate, usually silky, soft, and smooth when young,
and slippery surface when fresh, thick crust overlaying the context, a differentiated zone at
the point of attachment, and tough to break when dried. Pileus color usually homogenous
with reddish-orange (7A8) to brown (7D7–7D8) at the center when mature, golden yellow
(5B7), brownish-orange (5C5–5C6, 6D5) when old toward the stipe and margin surface.
Context up to 0.5–2.5 cm thick near stipe, brown (7D8) to brownish-red (8F8) when mature
or dried, soft and fibrous, covered with hard and thick crust, woody when old, trimitic
hyphal system present, hyaline, thin to thick-walled, branched. Tube 0.5–1.5 cm in length,
usually homogenous with orange (5A7) to dark orange (5A8), reddish-orange (7A7–7A8),
and grayish-red (8C7). Stipe 1–5 cm in length, 6 cm thick at the base, almost sessile or some
short, stipitate, broad and thick at the base, usually non-laccate, and brown (7D8) to dark
brown (8F8) when mature. Margin 0.5–4 cm thick, round, soft, brown (7D8) when mature
to old, and usually concolourous with the pileus. Pore 4–6 in number per mm, subcircular
to circular. Pore surface yellowish-white (2A2) when mature, discolored when touched,
brown (7D8) when scratched or bruised.
Hyphal structure: Hyphal system di-trimitic, with clamp connections, orange (6A7) to
deep orange (6A8), brownish-yellow (6C8) to brownish-orange (7C8); generative hyphae
1.4–2.8 µm broad (n = 30), hyaline, thin-walled, with clamp connections; skeletal hyphae 2.1–
4.4 µm broad (n = 30), usually hyaline, thick-walled, and solid; binding hyphae 1.5–3.6 µm
broad (n = 30), with walls varying in thickness, with many branches, some hymenial with
sword-like apices in the context. Pileipellis a hymeniderm, brown (6E8) to dark brown
(7F6), which is composed of apically clavate-like branched cells. Basidiospores mostly
ellipsoid to broadly ellipsoid, sometimes ovoid, with double walls, with a size range of
(6.9-)7.5–9.1–9.8(-10.6) × (4.7-)5.4–6.4–7.0(-7.7) µm (x = 9.1 × 6.4 µm, n = 50) µm, with Q
= 1.38–1.45, L = 9.09 µm, W = 6.41 µm (including myxosporium), (5.6-)6.3–7.6–8.4(-9.2)
× (4.2-)4.7–5.6–6.1(-6.6) µm (x = 7.6 × 5.7 µm, n = 50) µm, with Q = 1.35–1.40, L = 7.6
µm, W = 5.52 µm (excluding outer myxosporium), overlaid by hyaline, apically, and short,
echinulae, a truncate and turgid vesicular appendix, light yellow (4A4–4A5), grayish-
yellow (4B3–4B4) to brownish-orange (5C5–5C6), (5B8) of inner wall, outer wall usually
yellowish-brown (5D8, 5E7–5E8) to brown (6D7–6D8) in 5% KOH.
Habitat: Solitary, near the roots of a living Mangifera indica tree.
Specimens examined: LAOS, Luang Namtha Province, 20◦ 350 4700 N, 101◦ 040 0700 E, 935 m
elev., 20 June 2018, T. Luangharn, MFLU 19-2177.
Notes: Ganoderma adspersum was first reported by Donk [147], who described it as Poly-
porus adspersus Schulzer. Ganoderma adspersum is characterized by a distinctive non-laccate,
sessile, and applanate pileus. Ganoderma adspersum is often confused with G. applanatum,
G. australe, and Polyporus [148]. Ryvarden [149] and Ryvarden and Gilbertson [105] consid-
ered the correct name of G. adspersum as a synonym of G. australe, with G. adspersum can be
differentiated from G. applanatum by its thicker at the base, and larger basidiospores, while
G. applanatum tends to emerge sharply at right angles [105,110], with molecular analysis
also supporting the differentiation [45,76,150–153]. Our collections agree well with the
description provided by Ryvarden and Gilbertson [105].
J. Fungi 2021, 7, 819 41 of 83
J. Fungi 2021, 7, x FOR PEER REVIEW 44 of 91

Figure16.
Figure 16. Morphology
Morphology of ofGanoderma
Ganoderma adspersum
adspersum (MFLU
(MFLU19-2177):
19‐2177): (a)
(a) mature
maturebasidiomes;
basidiomes; (b)
(b)pore
pore
characteristics; (c) hyphae of pore; (d) hyphae of tube layers in KOH; (e,f) context hyphae
characteristics; (c) hyphae of pore; (d) hyphae of tube layers in KOH; (e,f) context hyphae in KOH;in KOH;
(g–j) basidiospores. Scale bars: (a) = 3 cm, (b) = 1000 μm, (d,e) 20 μm, (f) = 10 μm, (g–j) = 5 μm.
(g–j) basidiospores. Scale bars: (a) = 3 cm, (b) = 1000 µm, (d,e) 20 µm, (f) = 10 µm, (g–j) = 5 µm.

Ganodermaaustrale
Ganoderma australe(Fr.)
(Fr.)Pat.,
Pat.,Bull.
Bull.Soc.
Soc. mycol.
mycol. Fr.
Fr. 5(2,
5(2, 3):
3): 65
65 (1889)
(1889) (Figure
(Figure 17)
17)
Facesoffungi number: FoF
Facesoffungi number: FoF 06242 06242
Description: Basidiomes
Description: Basidiomes annual,
annual,perennial,
perennial,sessile.
sessile.Pileus
Pileus6–11
6–11cm cmin inlength,
length,2–6.5
2–6.5cmcm
broad,and
broad, and0.8–2
0.8–2cm
cm thick.
thick. Pileus
Pileus single,
single, flabelliform,
flabelliform, subdimidiate,
subdimidiate, spathulate,
spathulate, umbonate,
umbonate,
sulcate,obtuse
sulcate, obtusefrom
fromthethe host,
host, broadly
broadly attached,
attached, consistency
consistency hardhard and
and tough
tough when
when mature,
mature,
tough to break when dried, often thick at the center, slightly soft at the margin,
tough to break when dried, often thick at the center, slightly soft at the margin, and usually and usu‐
ally woody and corky when dried. Pileus surface corky, convex, furrowed,
woody and corky when dried. Pileus surface corky, convex, furrowed, spathulate, mostly spathulate,
mostly umbonate
umbonate or uneven,or uneven,
usually usually non‐laccate
non-laccate (dull)mature
(dull) when when mature to old, smooth
to old, smooth layers
layers when
when present,
present, deep at
deep sulcate sulcate at thethick
the center, center,
andthick
hardand hard
crust, crust, irregularly
irregularly ruptured
ruptured crust crust
overlying
overlying
the surface,the surface,dark
presented presented
browndark (7F8)brown
cracked (7F8) cracked
crust crust
when old, when
and toughold,toand tough
break when to
break when
dried. Pileusdried. Pileushomogeneous
color often color often homogeneous
with pale redwith palereddish-orange
(7A5), red (7A5), reddish‐orange
(7A6–7A7),
(7A6–7A7),
brown (7D8),brown (7D8),
to orange redto(8B7–8B8)
orange red on (8B7–8B8) on the upper
the upper surface surface
of the base of the
closed basemargin
to the closed
when
to themature
margintowhen
old. mature
Contextto upold.
to 0.5–1.2
Context cmupthick near stipe,
to 0.5–1.2 fibrous,
cm thick nearcomposed of coarse
stipe, fibrous, com‐
posed of coarse loose fibrils, brown (6D7–6D8), dark brown (6F7) to reddish‐brown (8D8,
J. Fungi 2021, 7, 819 42 of 83

loose fibrils, brown (6D7–6D8), dark brown (6F7) to reddish-brown (8D8, 8D9), covered
with thick crust, trimitic hyphal, thick-walled, dense with simple septa, typically with
narrow lumen, flexuous, and many branches. Tube 0.4–1 cm in length, brown (7D8) to
dark brown (6F8). Stipe sessile with broad attached. Margin white (4A1) when present
to mature, soft and slippery when growing fresh, shallow sulcate at the margin, covered
and blunt when old. Pore 4–6 in number per mm, subcircular to circular, sometimes
J. Fungi 2021, 7, x FOR PEER REVIEW 46 of 91
angular. Pore surface initially white (4A1), slightly yellowish-white (3A2) when mature,
brownish-red (8C4–8C5) when scratched, bruised, or discolored when touched.

Figure17.
Figure 17.Morphology
MorphologyofofGanoderma
Ganodermaaustrale
australe (MFLU
(MFLU 19‐2171):
19-2171): (a,b)
(a,b) mature
mature basidiomes;
basidiomes; (c–e)
(c–e) con‐
context
text hyphae in KOH; (f–i) basidiospores. Scale bars: (a,b) = 2 cm, (c–e) = 20 μm, (f–i)
hyphae in KOH; (f–i) basidiospores. Scale bars: (a,b) = 2 cm, (c–e) = 20 µm, (f–i) = 5 µm. = 5 μm.

Ganoderma
Hyphal gibbosum
structure: Hyphal(Blume
systemand T. Nees)
trimitic, withPat., Ann.connections,
clamp Jard. Bot. Buitenzorg, suppl. 1:
usually brownish-
114 (1897)
orange (Figurein18)
(6C5–6C7) KOH; generative hyphae 2.0–3.4 µm broad (n = 30), thin-walled, hyaline,
Facesoffungi
tapering number:
branches, with clampFoF 06243
connections; skeletal hyphae 3.1–4.5 µm broad (n = 30), usually
Description: Basidiomes annual or perennial,
thick-walled, sometimes branches, nearly solid; binding sessile, subflabellate,
hyphae 2.5–3.9 µmorwidth
subdimidiate.
(n = 30),
Pileus 2–16
usually cm in length,
thick-walled, many2–9 cm broad,
branches, and
nearly 0.5–2.3
solid, and cm thick. Pileus
hymenial conks, convex,
with sword-like apices imbri‐
in the
cate, umbonate, uneven, ungulate, usually round when occurring, primordial, somewhat
round and plump when young, flabelliform (fan‐shaped) when seen from above, broadly
attached when mature, thick at the base when mature. Pileus surface non‐laccate, smooth
when young, silky, soft, and slippery surface when fresh, furrowed on the surface with
J. Fungi 2021, 7, 819 43 of 83

context. Pileipellis a hymeniderm, brown (7D8), composed of apically acanthus-like branched

cells, dextrinoid. Basidiospores mostly ellipsoid to broadly ellipsoid, with double walls, with a
size range of (6.5-)7.6–10.1–11.4(-12.5) × (5.9-)6.7–8.5–9.2(-10.3) µm (x = 7.2 × 5.9 µm, n =
50) µm, with Q = 1.19–1.26, L = 7.24 µm, W = 5.92 µm (including myxosporium), (5.1-)6.2–
8.3–9.7(-10.9) × (4.4-)5.6–6.8–7.7(-8.8) µm (x = 8.2 × 6.8 µm, n = 50) µm, with Q = 1.17–1.26,
L = 8.23 µm, W = 6.79 µm (excluding outer myxosporium), overlaid by hyaline, brown
apically, bearing fine, distinct, short, echinulae, truncate, turgid vesicular appendix, inner
wall light brown (6D4–6D5) to brown (7E7–7E8), and outer wall usually reddish-brown
(8E5–8E6, 8F7) in 5% KOH.
Habitat: Solitary, on the decaying hardwood of Canarium spp. tree species.
Specimens examined: LAOS, Luang Namtha Province, 20◦ 350 4700 N, 101◦ 040 0700 E, 935 m
elev., 20 June 2018, T. Luangharn, MFLU 19-2171.
Ganoderma gibbosum (Blume and T. Nees) Pat., Ann. Jard. Bot. Buitenzorg, suppl. 1:
114 (1897) (Figure 18)
Facesoffungi number: FoF 06243
Description: Basidiomes annual or perennial, sessile, subflabellate, or subdimidiate.
Pileus 2–16 cm in length, 2–9 cm broad, and 0.5–2.3 cm thick. Pileus conks, convex, imbri-
cate, umbonate, uneven, ungulate, usually round when occurring, primordial, somewhat
round and plump when young, flabelliform (fan-shaped) when seen from above, broadly
attached when mature, thick at the base when mature. Pileus surface non-laccate, smooth
when young, silky, soft, and slippery surface when fresh, furrowed on the surface with
sulcate to undulating, somewhat spathulate to uneven, incised, compact, hard, and woody
when older, covered with a tough crust (0.1–0.2 mm), usually dull and faded when mature
to old, and some occurred the lined or cracked crust when older. Pileus color brownish-
orange (5C5), reddish white (7A2) at the base, and homogenous with grayish-orange (6B3),
brownish-orange (7C5), and light brown (6D4) toward the center of maturity fruiting
bodies, white (6A1) at the margin, and usually the color changes to dark brown (8F8) upon
touch, becoming grayish-red (8C4–8F6), reddish-brown (8E6) to dull red (10C3) when old.
Context up to 0.3–1.3 cm thick, trimitic hyphal with clamp connections, hyaline, with walls
varying in thickness, simple septate, composed of narrow, and sparse branches; generative
hyphae 1.0–3.4 µm broad (n = 30), with walls varying in thickness, and hyaline; skeletal
hyphae 4.0–6.4 µm broad (n = 30) with thick walls; binding hyphal 2.0–6.5 µm broad
(n = 30). Hymenophore up to 3 mm in length, with reddish-brown (8D7). Tube layers
0.2–0.8 cm in length, up to 80–163 µm in width, and non-presented when young. Stipe
almost sessile, broadly attached when present. Margin wavy, blunt, slippery when wet,
thinner at the base and soft than the center, often white (8A1) from youth to maturity,
and light brown (6D5) when old. Pore 4–7 in number per mm, when fresh, angular, subcir-
cular to circular. Pore surface white (11A1) to orange white (6A2) when fresh, scratched or
bruised, and discolored when touched.
Hyphal structure: Hyphal system trimitic, with clamp connections, usually reddish-
brown (8D7–8D8); generative hyphae 1.0–3.8 µm broad (n = 30), with walls varying in
thickness, hyaline, and unbranched; skeletal hyphae 4.2–6.4 µm broad (n = 30), light brown
(7D6) to brown (7D8) in Melzer’s reagent with dextrinoid, usually thick-walled; binding
hyphae 1.8–6.4 µm broad (n = 30), brown (6F8) to reddish-brown (8D8) in Melzer’s reagent,
thick-walled, many branches, the generative and skeletal hyphae usually intertwined,
mostly dark brown (6F7) near the tube layers; Bovista-type ligative hyphae, hymenial
with sword-like apices in the context. Pileipellis a hymeniderm, brown (6D8) to light
brown (6D6), composed of apically acanthus-like branched cells, dextrinoid. Basidiospores
mostly ellipsoid to oblong ellipsoid or broadly ellipsoid, with double walls, with a size
range of (4.4-)6.8–8.2–9.5(-10.2) × (3.6-)4.2–5.2–5.8(-6.5) µm (x = 8.3 × 5.4 µm, n = 50) µm,
with Q = 1.49–1.56, L = 8.34 µm, W = 5.44 µm (including myxosporium), (3.6-)5.3–6.0–
7.2 (-8.3) × (2.8-)3.9–4.5–5.4(-6.2) µm (x = 6.2 × 4.5 µm, n = 50) µm, with Q = 1.34–1.40,
L = 6.24 µm, W = 4.51 µm (excluding outer myxosporium), overlaid by hyaline, dextrinoid,
J. Fungi 2021, 7, 819 44 of 83

echinulae, inner wall echinulate brown, light brown (6D6–6D8) to brown (6E8) in 5% KOH,
and reddish-brown (8F6) to dark brown (8F8) in Melzer’s reagent. Basidia not seen.
Habitat: Occasionally on decaying wood of Pinus spp.
Specimens examined: LAOS, Luang Prabang, 19◦ 510 5100 N, 102◦ 110 3900 E, 589 m elev.,
12 July 2018, T. Luangharn, MFLU 19-2190.
Ganoderma nasalaense Hapuar., Pheng. and K.D. Hyde, Mycosphere 10(1): 272 (2019)
Taxonomy and phylogeny analysis are shown in Hapuarachchi et al. [30].
Notes: Ganoderma nasalaense is characterized by its duplex context, rigid basidiomes,
purplish-black laccate crust, dark brown to gray on the upper pileus surface, brown tube
layer, purplish-brown pore surface, trimitic hyphal system, and relatively large, broadly el-
lipsoid to ellipsoid light brown basidiospores (12.1–13.8 × 9.2–10.5) µm, truncate, with fine
and short echinulae (10–12 × 6.5–7.5) µm, and a cuticle composed of strongly amyloid,
clavate cells, usually with several irregular lobes or protuberances (30–80 × 3–10.5)
J. Fungi 2021, 7, x FOR PEER REVIEW 48 of 91 µm.
This fungus is solitary on decaying hardwood tree trunks in Huaphanh Province, Laos.

Figure 18.
Figure 18. Morphology
MorphologyofofGanoderma
Ganodermagibbosum
gibbosum (MFLU
(MFLU 19‐2190): (a,b)
19-2190): mature
(a,b) basidiomes;
mature (c–e)(c–e)
basidiomes; con‐ con-
text hyphae as seen in Melzer’s reagent; (f–h) basidiospores. Scale bars: (a,b) = 2 cm, (c–e) = 20 μm,
text hyphae as seen in Melzer’s reagent; (f–h) basidiospores. Scale bars: (a,b) = 2 cm, (c–e) = 20 µm,
(f–h) = 5 μm.
(f–h) = 5 µm.
Ganoderma nasalaense Hapuar., Pheng. and K.D. Hyde, Mycosphere 10(1): 272
(2019)
Taxonomy and phylogeny analysis are shown in Hapuarachchi et al. [30].
Notes: Ganoderma nasalaense is characterized by its duplex context, rigid basidiomes,
purplish‐black laccate crust, dark brown to gray on the upper pileus surface, brown tube
J. Fungi 2021, 7, 819 45 of 83

3.2.3. Taxonomy of Ganoderma from Myanmar

Ganoderma hoehnelianum Bres., Annales Mycologici. 10 (5): 502 (1912) (Figure 19)
Facesoffungi number: FoF 06260
Description: Basidiomes annual or perennial, sessile. Pileus is 0.5–4 cm in length,
0.5–3 cm broad, and up to 0.5 cm thick at the base. Pileus is applanate, umbonate, sub-
reniform to reniform, or subflabellate to glabrous from youth to maturity, small in size,
obtuse from host, often with undefined concentric zones at the center that extend to
the margin, thick at the center, slightly thin and soft at the margin, and tough to break
when dried. Pileus surface shiny, smooth, and soft when young, frequently furrowed and
shallowly sulcate on the upper surface, which is undulating and somewhat spathulate to
uneven in maturity, covered by a thin crust, faded or weakly laccate when young, laccate
when mature, and woody when old. Pileus color homogenous, reddish-brown (8E4–8E8) to
dark brown orange (9F5–9F8) at the center when mature. Context up to 0.1–0.3 cm thick at
the base, brown (7D7–7D8), reddish-brown (8E7–8E8), and dark brown (7F7), abundantly
thick-walled, with clamp connections, subsolid hyphae, containing a fibrous pithy context.
Tube layers hard and corky, branched, with clamped connections, often brown (7D7) to
dark brown (7F8). Stipe almost sessile, broadly attached when present. Margin obtuse
from the center, soft and smooth when young, laccate when mature, slightly wavy and
slippery when wet, often yellowish-white (3A2) on the upper surface, and pale yellow
(4A3) under the margin. Pores 4–6 in number per mm. Pore surface usually white (11A1)
to light orange (5A4) when young, grayish-orange (6B4–6B6) when mature, turning light
brown (7D5–7D6), brown (7D7), and reddish-brown (8D5–8D7) when scratched or bruised,
becoming discolored when touched.
Hyphal structure: Hyphal system is dimitic, bearing clamp connections, hyaline,
with walls varying in thickness with simple septa and some swollen differentiated zones
at the point of attachment, composed of several narrow hyphae, and sparingly branched;
generative hyphae (1.8-)2.4–3.2–3.7(-4.2) µm broad (n = 30) are thin-walled and hyaline;
skeletal hyphae (3.3-)3.9–5.2–5.8(-6.2) µm broad (n = 30) have walls of varying thickness,
sometimes subsolid; binding hyphae (2.3-)2.9–4.2–4.9(-5.6) µm broad (n = 30) are usually
thick-walled with many branches, and appear alongside Bovista hyphae, which are usually
present from orange (5A6–5A7) to deep orange (6A8) and thin-walled, as well as also
reddish-brown (8D7–8D8, 9D8) to brownish-red (9C8) in Melzer’s reagent. Basidiospores
are mostly ellipsoid and featuring several ovoid with double walls, (6.7-)7.5–9.8–11.8(-13.2)
× (5.7-)6.4–7.8–9.6(-10.8) µm (x = 9.7 × 7.8 µm, n = 50) µm, with Q = 1.22–1.28, L = 9.73 µm,
W = 7.78 µm (including myxosporium), (5.8-)6.4–8.5–9.3(-10.2) × (4.8-)5.3–6.1–6.8(-7.4) µm
(x = 8.4 × 6.4 µm, n = 50) µm, with Q = 1.29–1.35, L = 8.43 µm, W = 6.41 µm (excluding
outer myxosporium), overlaid by a hyaline, dextrinoid, and echinulate whose inner wall
presents as brownish-orange (5C5–5C6) to brownish-yellow (5C7–5C8) and dark brown
(6F8) to reddish-brown (8E7–8E8) in Melzer’s reagent.
Specimens examined: MYANMAR, Chin State, Tedim Township, 13 July 2019, P. E.
Mortimer, MFLU 19-2168.
Notes: Ganoderma hoehnelianum was introduced by Bresadola in 1912 from Indonesia.
Of distinctive note are its context color, basidiospore characteristics, and cuticular cells [154].
Wang and Wu [154] reported that the original Chinese G. hoehnelianum was an earlier name
for G. shangsiense, and this fungus was also recorded as G. shangsiense in China’s Hainan
Province [30,91,120,121]. Our G. hoehnelianum is first recorded from Myanmar, and its
description is consistent with the descriptions provided by Wang and Wu [154].
J. Fungi 2021, 7, 819 46 of 83
J. Fungi 2021, 7, x FOR PEER REVIEW 50 of 91

Figure19.
Figure 19. Morphology
Morphology characteristics
characteristicsofofGanoderma
Ganodermahoehnelianum
hoehnelianum(MFLU
(MFLU19-2168):
19‐2168):(a)
(a)young
youngbasid-
ba‐
sidiomes; (b) pore characteristics; (c–e) context hyphae in Melzer’s reagent; (f–i) basidiospores.
iomes; (b) pore characteristics; (c–e) context hyphae in Melzer’s reagent; (f–i) basidiospores. Scale
Scale bars: (a) = 2 cm, (b) = 1000 μm, (c–e) = 20 μm, (f–i) = 5 μm.
bars: (a) = 2 cm, (b) = 1000 µm, (c–e) = 20 µm, (f–i) = 5 µm.

Ganodermamyanmarense
Ganoderma myanmarenseKarunarathna,
Karunarathna, Mortimer,
Mortimer, & Luangharn,
& Luangharn, sp. nov.
sp. nov. (Figure
(Figure 20)
20) FacesofFungi number: FoF 06262
FacesofFungi
Index Fungorum number: FoFIF06262
number: 556794
Index Fungorum
Diagnosis: Ganodermanumber: IF 556794
myanmarense is characterized by its shell-like pileus with strongly
laccateDiagnosis:
appearanceGanoderma myanmarense
colored orange, goldenisyellow
characterized by its shell‐like
at center, extending pileus with
reddish-yellow and
strongly
yellow atlaccate
margin, appearance colored orange,
usually homogenous golden yellow
reddish-brown at center,
when mature, extending
a white reddish‐
to light
yellowthat
yellow andindicates
yellow atactive
margin, usually homogenous
development reddish‐brown
on the margin of the pilealwhen mature,
surface, a white
a white pore
to light when
surface yellowfresh,
that an
indicates
orange,active
deepdevelopment on the margin
orange to reddish-orange of the pileal
context, surface,ofa
and absence
white pore
melanoid surface when fresh, an orange, deep orange to reddish‐orange context, and
bands.
absence of melanoid
Holotype: MYANMAR, bands.Chin State, Tedim Township, on dead wood of Casternopsis sp.,
13 JulyHolotype: MYANMAR,
2019, LT2019 Chin State, Tedim Township, on dead wood of Casternopsis
(MFLU 19-2167).
sp., 13 July 2019,The
Etymology: LT2019 (MFLU
species epithet19‐2167).
“myanmarense” refers to the country Myanmar, where
the holotype specimen was collected.
Etymology: The species epithet “myanmarense” refers to the country Myanmar, where
the holotype specimen was collected.
J. Fungi 2021, 7, 819 47 of 83
J. Fungi 2021, 7, x FOR PEER REVIEW 52 of 91

Figure20.
Figure 20.Characteristics
Characteristicsof
ofGanoderma
Ganodermamyanmarense:
myanmarense:(a)
(a)the
theupper
uppersurface
surfaceofofmature
maturebasidiomes
basidiomesofof
the strain MFLU 19‐2167; (b) the lower surface of mature basidiomes of strain MFLU 19‐2167; (c) the
the strain MFLU 19-2167; (b) the lower surface of mature basidiomes of strain MFLU 19-2167; (c) the
upper surface of mature basidiomes of the strain MFLU 19‐2169; (d,e) pore characteristics; (f) hy‐
upper surface of mature basidiomes of the strain MFLU 19-2169; (d,e) pore characteristics; (f) hyphae
phae of tube layers in KOH; (g) context hyphae in KOH; (h) context hyphae with clamp connections
of
intube
KOH; layers in KOH;
(i) hyphae (g) context
of trama hyphae
in KOH; in KOH; (h) context
(j–l) basidiospores. hyphae
Scale bars: (a,c) =with
2 cm,clamp
(d) = connections
500 μm, (e) =
in KOH; (i) hyphae of trama in
150 μm, (f–i) = 20 μm, (j–l) = 5 μm.KOH; (j–l) basidiospores. Scale bars: (a,c) = 2 cm, (d) = 500 µm,
(e) = 150 µm, (f–i) = 20 µm, (j–l) = 5 µm.
Ganoderma williamsianum Murrill, Bulletin of the Torrey Botanical Club 34: 478
Description: Basidiomes annual or perennial, stipitate. Pileus up to 5–16 cm in length,
(1907) (Figure 21)
4–15 cm in width, and 1–2.5 cm thick at the base. Pileus shell-like (involute from the margin
≡ Elfvingia williamsiana (Murrill) Imazeki, Bulletin of the Government Forest Exper‐
toward the center), sub-reniform to reniform or subflabellate to circular when viewed from
imental Station Meguro. 57: 106 (1952)
J. Fungi 2021, 7, 819 48 of 83

above, often with undefined zones at the center that extend to the margin, with a thick
center that is slightly soft at the margin, and tough to break when dried. Pileus surface
shiny, silky, smooth, and soft when fresh, hard when old, furrowed, sulcate to undulating,
somewhat spathulate to uneven, incised, faded or weakly laccate when young, strongly
laccate on maturity or when old. Pileus color homogenous, orange (5A6–5A7, 6B7–6B8),
golden yellow (5B6), and yellowish-red (8A7–8A8) at the center, slightly reddish-orange
(7A7–7A8) and reddish-brown (8E8) where deeply sulcate, reddish-yellow (4A7) and yellow
(2A6) at the margin. Context up to 0.6–1.5 cm thick at the base, with some areas thin-walled,
thick-walled, along with subsolid hyphae, bearing clamp connections, and absence of
melanoid bands. Tube layers woody and hard, usually thin-walled, frequently branched,
with clamped connections, and often dark brown (7F7) when dried. Stipe up to 3–8 cm in
length, up to 1.3–4 cm in width, centrally stipitate, nearly sub-cylindrical to cylindrical,
concolorous with the pileus, often dark brown (9F7–9F8) to violet-brown (11F7–11F8),
strongly laccate from maturity to old. Margin obtuse from the center, strongly laccate at
the edge, occasionally wavy, slippery when wet, smoother, softer, and thinner, at the base,
often yellow (2A6), and deep orange (5A8) to golden yellow (5B7–5B8) on maturity to old.
Pores 4–6 in number in number per mm, (60–)80–125(–165) µm, angular, subcircular to
circular. Pore surface usually white (11A1), turnins light brown (7D5), brown (7D7–7D8) to
dark brown (7F6–7F8) when scratched or bruised, becomes discolored when touched.
Hyphal structure: Hyphal system trimitic, with walls of varying thickness, clamp
connections present, simple septate hyphae, hyaline, narrow and sparingly branched;
generative hyphae (1.2-)1.8–2.2–2.4(-2.7) µm broad (n = 30), hyaline, thin-walled; skeletal
hyphae (1.8-)2.5–3.8–4.5(-5.0) µm broad (n = 30), abundant and thick-walled, sometimes
subsolid; binding hyphae (1.5-)1.9–2.7–3.3(-4.3) µm broad (n = 30), usually thick-walled.
Basidiospores (8.2-)9.3–11.6–12.5(-13.6) × (5.3-)6.0–7.1–7.8(-8.6) µm, (Qm = 1.7, Q = 1.2–2.4,
including myxosporium) (n = 50), (x = 11.5 × 7.1 µm, n = 50) µm, with Q = 1.57–1.65,
L = 11.52 µm, W = 7.13 µm (including myxosporium), (7.5-)8.3–9.1–10.4(-11.9) × (4.9-)5.4–
6.2–6.8(-7.5) µm (x = 9.0 × 6.3 µm, n = 50) µm, with Q = 1.40–1.48, L = 9.04 µm, W = 6.27 µm
(excluding outer myxosporium), mostly ellipsoid to broadly ellipsoid, or globose, double
walls, overlaid by hyaline; exosporium (outer wall) smooth, hyaline, endosporium (inner
wall) coarse and echinulate, with turgid vesicular appendix, and pale yellow inner wall that
can also present in KOH, and yellowish-brown (5E6–5E7) to brown (6E6–6E7) in Melzer’s
reagent, outer wall pale orange (6A3–6A4) to orange (6A6–6A7) in KOH, reddish-orange
(7A8), grayish-red (7B6–7B8), and dark brown (7F7–7F8) in Melzer’s reagent.
Habitat: Solitary near the hardwood root of unknown tree species.
Additional specimens: MYANMAR, Chin State, Tedim Township, 13 July 2019, P. E.
Mortimer, MFLU 19-2167 (holotype) and MFLU 19-2169 (paratype).
Ganoderma williamsianum Murrill, Bulletin of the Torrey Botanical Club 34: 478
(1907) (Figure 21)
≡ Elfvingia williamsiana (Murrill) Imazeki, Bulletin of the Government Forest Experi-
mental Station Meguro. 57: 106 (1952)
Facesoffungi number: FoF 06261
Description: Basidiomes annual or perennial, stipitate. Pileus 0.5–1.5 cm in length,
0.5–1 cm in width, and up to 0.5 cm thick at the base. Pileus sub-reniform to reniform,
or subflabellate to circular when young, often with concentric zones at the center that
extend to the margin, thick at the center, slightly soft at the margin, tough to break when
dried. Pileus surface shiny, silky, and soft when young, generally furrowed, smooth, sulcate
to undulating, somewhat spathulate to uneven when mature, hard and woody when old,
incised on the surface, faded or weakly laccate when young, and usually laccate when
mature. Pileus color usually homogenous, orange (5A7–5A8) to deep orange (5B7–5B8),
slightly white (5A1), and yellowish-white (3A2, 4A2) at the margin when young. Context
is up to 0.3–1.5 cm thick at the base, which is usually thick-walled, with abundant walls
varying in thickness, subsolid hyphae containing a fibrous pithy context with clamp con-
nections, and brown (7D8) melanoid bands when mature. The tube layers hard, frequently
J. Fungi 2021, 7, 819 49 of 83

branched with clamp connections, and often dark brown (7F8). Stipe 1.5 cm in length, 1 cm
in width when young, lateral to nearly dorsal, entrally stipitate, almost sub-cylindrical to
cylindrical, concolorous with the pileus, thick when young, and often brownish-orange
(7C6) to dark brown (9F7–9F8). Margin obtuse from the center, blunted when young, occa-
sionally wavy, slippery when wet, smooth and soft when young (1.5 cm), and often white
(3A1) to yellowish-white (3A2) when young. Pore surface white (11A1) to yellowish-white
J. Fungi 2021, 7, x FOR PEER REVIEW 54 of 91
(3A2), turns light brown (7D5) to brown (7D7–7D8) when scratched or bruised, becomes
discolored when touched.

Figure 21.
Figure 21. Morphology
Morphology characteristics
characteristicsofofGanoderma
Ganodermawilliamsianum
williamsianum(MFLU(MFLU 19‐2170): (a) (a)
19-2170): young ba‐
young
sidiomes; (b–d)
basidiomes; context
(b–d) hyphae
context inin
hyphae Melzer’s
Melzer’s reagent. Scale
reagent. bars:
Scale (a)(a)
bars: = 2=cm, (b–d)
2 cm, = 20
(b–d) μm.
= 20 µm.

3.2.4.Hyphal
Taxonomy of Ganoderma
structure: Hyphal system from Thailand
trimitic, bearing clamp connections, hyaline, with walls
of varying thickness, simple septate, swollen differentiated
Ganoderma adspersum (Schulzer) Donk Proc. K. Ned. zones at the
Akad. point
Wet., Ser.ofC,attachment,
Biol. Med.
composed of
Sci. (Figure 22)several narrow hyphae, and sparingly branched; generative hyphae (1.6-)1.9–
2.3–2.5(-2.6) µm broad
Facesoffungi (n = 30),
number: FoF thin-walled
06241 and hyaline; skeletal hyphae (3.1-)3.4–3.9–4.3
(-4.8) Description:
µm broad (n = 30), with
Basidiomes wallsto
annual ofperennial,
varying thickness,
applanate,with some subsolid;
subdimidiate. Pileusbinding
2–14 cm
hyphae (1.6-)1.9–2.6–2.9(-3.4)
in length, 2–7 cm in width, and broad (n
µm0.5–1.8 cm=thick
30), usually
at base.thick-walled, appearing
Pileus subdimidiate alongside
to dimidiate,
Bovista hyphae
flabelliform, with many
spathulate, branches,
umbonate, whose thick sulcate
concentrically walls usually presentorasshort
zone, sessile lightstipitate,
orange
(5A5), orange (5A6–5A7, 6B7–6B8) to deep orange (5A8, 6A8), and reddish-brown
distinctly contracted base, somewhat round and plump when young, somewhat imbricate (8D7–
8D8) to brownish-red (9C8) in Melzer’s reagent. Basidiospores not observed.
when viewed from above, flabelliform (fan‐shaped), usually broadly attached with radial
fromSpecimens examined:
center extending to MYANMAR,
the margin, toughChinwhen
State,break,
Tedimthick
Township,
at base,13 July 2019,
slightly soft atP.the
E.
Mortimer, MFLU 19-2170.
margin when mature, light weight when dried, and woody and corky texture when dried.
Pileus surface non‐laccate (dull), convex, radial furrowed, incised, spathulate, shallow sul‐
cate, usually silky, soft, smooth when young, and slippery surface when fresh, thick crust
overlying the context, differentiated zone at the point of attachment, several layers thick,
and leathery when broken. Pileus color usually homogenous with brown (7D7–7D8, 7E7–
7E8, 8D8) at center toward stipe to margin surface when mature. Context up to 0.3–1.3 cm
J. Fungi 2021, 7, 819 50 of 83

Notes: Ganoderma williamsianum belongs to the group of laccate Ganoderma. This fungus
was originally reported in the Philippines and is easily recognized, with its small, dense,
ungulate of pileus with pale, yellow pores, large spores, and a short skeletal [2,113]. Among
the Ganoderma species, there are some similarities between G. williamsianum and G. brownii,
such as both having yellow pores [109]; however, G. brownii can be differentiated from
G. williamsianum by its dull pileus with a hard crust [155], skeletal hyphae with occasional
branching, and smaller basidiospores [109,110,137,155]. Ganodermawilliamsianum is the ear-
liest valid name for G. meijiangense [154], containing similarities to G. meijiangense, such as
both having sessile, annual crust basidiomes, and white margin, but G. williamsianum can
be differentiated from G. meijiangense by its distinguishing dark brown context, without
any layer of black crust and a distinct cuticular composition [61]. Our G. williamsianum
collection from Myanmar marks a new record, as it shares traits similar to Moncalvo and
Ryvarden [2], with sessile, annual crust-like basidiomes, and white margin, trimitic hyphal
system bearing clamp connections, hyaline, and walls of varying thickness.

3.2.4. Taxonomy of Ganoderma from Thailand

Ganoderma adspersum (Schulzer) Donk Proc. K. Ned. Akad. Wet., Ser. C, Biol. Med.
Sci. (Figure 22)
Facesoffungi number: FoF 06241
Description: Basidiomes annual to perennial, applanate, subdimidiate. Pileus 2–14 cm
in length, 2–7 cm in width, and 0.5–1.8 cm thick at base. Pileus subdimidiate to dimidiate,
flabelliform, spathulate, umbonate, concentrically sulcate zone, sessile or short stipitate,
distinctly contracted base, somewhat round and plump when young, somewhat imbricate
when viewed from above, flabelliform (fan-shaped), usually broadly attached with radial
from center extending to the margin, tough when break, thick at base, slightly soft at the
margin when mature, light weight when dried, and woody and corky texture when dried.
Pileus surface non-laccate (dull), convex, radial furrowed, incised, spathulate, shallow
sulcate, usually silky, soft, smooth when young, and slippery surface when fresh, thick
crust overlying the context, differentiated zone at the point of attachment, several layers
thick, and leathery when broken. Pileus color usually homogenous with brown (7D7–
7D8, 7E7–7E8, 8D8) at center toward stipe to margin surface when mature. Context up
to 0.3–1.3 cm thick near the stipe, brown (7D8) to brownish-red (8F8) when mature and
dried, soft and fibrous, covered with hard and thick crust, woody when old, trimitic
hyphal, hyaline, with walls varying in thickness, with branches. Tube 0.2–1 cm in length,
usually homogenous with orange (5A7) to dark orange (5A8), reddish-orange (7A7–7A8),
and grayish-red (8C7). Stipe 1–3.8 cm in length, 3.5 cm thick at base, almost sessile,
some shortly stipitate, broadly thick at base, usually non-laccate, brown (7D8) to dark
brown (8F8) when mature. Margin round, soft, occurring brown (7D8) from mature to old,
presented numerous undulations, and usually concolorous with the pileus. Pores 4–6 in
number in number per mm, subcircular to circular. Pores surface yellowish-white (2A2)
when mature, turns brown (7D8) when scratched or bruised.
Hyphal structure: Hyphal system di-trimitic, with clamp connections; generative hy-
phae 1.2–2.8 µm broad (n = 30), hyaline, thin-walled, with clamp connections; skeletal
hyphae 2.0–4.4 µm broad (n = 30), usually hyaline, thick-walled, solid; binding hyphae
1.6–3.7 µm width (n = 30), with walls varying in thickness, many branches, without clamp
connections, some hymenial with sword-like apices in the context. Basidiospores mostly el-
lipsoid, sometimes ovoid with double walls, (7.0-)7.7–9.0–9.9(-10.7) × (4.8-)5.3–6.5–7.1 (-7.8)
µm (x = 8.9 × 6.7 µm, n = 50) µm, with Q = 1.30–1.37, L = 8.91 µm, W = 6.69 µm (including
myxosporium), (5.8-)6.4–7.7–8.6(-9.4) × (4.1-)4.5–5.6–6.0(-6.5) µm (x = 7.7 × 5.5 µm, n = 50)
µm, with Q = 1.35–1.42, L = 7.69 µm, W = 5.52 µm (excluding outer myxosporium), over-
laid by hyaline, apically and shortly echinulate, truncate and turgid vesicular appendix,
inner wall brownish-yellow (5C7–5C8) and light brown (6D7–6D8), outer wall usually
brownish-orange (7C8), brown (7D7–7D8) to dark brown (7E7) in 5% KOH.
Habitat: Solitary on decaying Pterocarpus sp.
J. Fungi 2021, 7, 819 51 of 83

J. Fungi 2021, 7, x FOR PEER REVIEW 56 of T.

Specimens examined: THAILAND, Kanchanaburi Province, 10 November 2018, 91
Luangharn, MFLU 19-2178.

Figure 22.
Figure 22. Morphology
Morphology of of Ganoderma
Ganoderma adspersum
adspersum (MFLU
(MFLU 19-2178):
19‐2178): (a)
(a) mature
mature basidiomes;
basidiomes; (b)
(b) pores
pores
characteristics; (c) hyphae of tube layers in KOH; (d,e) hyphae of thama in KOH; (f–h) basidiospores.
characteristics; (c) hyphae of tube layers in KOH; (d–f) hyphae of thama in KOH; (g–j) basidiospores.
Scale bars: (a) = 2 cm, (b) = 1000 μm, (c,e) = 20 μm, (f–h) = 5 μm.
Scale bars: (a) = 2 cm, (b) = 1000 µm, (c–f) = 20 µm, (g–j) = 5 µm.

Ganodermaapplanatum
Ganoderma applanatum(Batsch)
(Batsch)G.F.
G.F. Atk.,
Atk., Annales
Annales Mycologici
Mycologici 6: 189
6: 189 (1908)
(1908) (Figure
(Figure 23)
23) Facesoffungi number: FoF 06249
Facesoffungi
Description: number: FoF
Basidiomes 06249
annual or perennial, subdimidiate to dimidiate, sessile. Pileus
Description: Basidiomes annual
2–12 cm in length, 2.5–6 cm in width, or and
perennial,
0.8–3.1subdimidiate
cm thick at the to dimidiate,
base. Pileussessile. Pi‐
sessile,
leus 2–12 cm in length, 2.5–6 cm in width, and 0.8–3.1 cm thick at the base.
perennial, subdimidiate to dimidiate, subflabelliform to flabelliform, convex, imbricate, Pileus sessile,
perennial, or
umbonate subdimidiate to ungulate,
uneven, rarely dimidiate,withsubflabelliform
broadly attachedto flabelliform,
when mature, convex, imbricate,
thicker at base,
umbonate
slightly orat
soft uneven,
marginrarely
whenungulate,
mature. with
Pileusbroadly
surface attached when
furrowed, mature, thicker
tuberculate at base,
to undulate,
slightly and
uneven, soft incised
at margin
whenwhen
old, mature. Pileus
non-laccate surface
(dull), furrowed,
compact and hardtuberculate to undulate,
when mature, woody
uneven,
to and incised
corky texture whenwhen
matureold,tonon‐laccate (dull),
old, covered withcompact and (0.1–0.25
a thin crust hard when mm)mature, woody
overlies the
to corky texture when mature to old, covered with a thin crust (0.1–0.25
pileus, and some cracked crust when old. Pileus color differentiated zone with peach redmm) overlies the
pileus,toand
(7A4) some cracked(7A6,
reddish-orange crust7B6)
whenatold.
base,Pileus
towardcolor
to differentiated
brownish-orange zone(7C6–7C8)
with peach red
with
(7A4) to reddish‐orange (7A6, 7B6) at base, toward to brownish‐orange (7C6–7C8) with
radius light brown (7D5) zone at center, and extend to dark brown (6F8) closed to active
mycelium (margin) of maturity fruiting bodies. Context up to 0.5–2.3 cm thick, mostly
brown (6E8) to dark brown (7F6–7F8) of cuticle cells, upper layers light orange (5A5),
J. Fungi 2021, 7, 819 52 of 83

radius light brown (7D5) zone at center, and extend to dark brown (6F8) closed to active
mycelium (margin) of maturity fruiting bodies. Context up to 0.5–2.3 cm thick, mostly
brown (6E8) to dark brown (7F6–7F8) of cuticle cells, upper layers light orange (5A5), lower
reddish-brown (8D8) with fibrous, some fibrous pithy context, usually separated layers
of context tissue at base, and some occurred woody line. Tubes 0.8–2 cm in length, up to
70–160 µm in width, with sulcate at different levels. Stipe almost sessile (without stipe)
with broadly attached when present, with differentiated zone at the point of attachment.
Margin up to 1 cm thick, round, soft, often white (5A1), turning to light brown (6D4) and
brown (6E8) when scratched or bruised, slippery when wet, soft when young, and thin
than the center. Pore angular, subcircular, 4–6 in number per mm. Pore surface white (7A1)
when fresh, quickly turning to light brown (7D6) to brown (7D8) when handled, scratched,
and bruised.
Hyphal structure: Hyphal system trimitic; generative hyphae 0.5–2.8 µm (x = 2.4, n
= 30) in diam, clamp connections, almost hyaline, abundant thin-walled and occasion-
ally thick-walled, composed of narrow and sparingly branched; skeletal hyphae 2.7–4.9
µm broad (n = 30), usually thick-walled, hyaline, sometimes branched; binding hyphae
1.2–3.6 µm broad (n = 30), thick-walled, some branched, and intertwined the skeletal hy-
phae. Pileipellis a hymeniderm, brown (6E4), which composed of apically acanthus-like
branched cells. Basidiospores mostly ellipsoid to broadly ellipsoid, sometimes subcircular
with double walls, overlaid by hyaline, exosporium (outer wall) hyaline, endosporium
(inner wall) coarse echinulate, with turgid vesicular appendix, truncate at the distal end,
(10.0-)10.5–11.3–11.9(-12.4) × (7.2-)8.0–8.8–9.4(-10.2) µm, (x = 11.2 × 8.6 µm, n = 50) µm,
with Q = 1.28–1.33, L = 11.23 µm, W = 8.62 µm (including myxosporium), (6.3-)7.4–8.5–9.3
(-10.1) × (4.9-)6.3–7.4–8.7(-9.3) µm (x = 8.4 × 7.5 µm, n = 50) µm, with Q = 1.09–1.13,
L = 8.42 µm, W = 7.58 µm (excluding outer myxosporium), brown (7D7–7D8) in KOH,
and reddish-brown (8E6) to dark brown (8F4) in Melzer’s reagent. Basidia 14–20 × 8–10 µm,
with four sterigmata.
Habitat: Solitary on rotten wood, dead trunks, and decaying stumps of Artocarpus spp.,
and Dipterocarpus spp.
Specimens examined: THAILAND, Kanchanaburi Province, 10 November 2018, T.
Luangharn, MFLU 19-2175.
Ganoderma australe (Fr.) Pat., Bull. Soc. mycol. Fr. 5(2, 3): 65 (1889)
Facesoffungi number: FoF 02906
Characteristics follow Luangharn et al. [59].
Description: Basidiomes applanate, spathulate. Pileus 2–7 cm in length, 1–15 cm in
width, and 0.5–3 cm thick near the base. Pileus circular, applanate, spathulate, sometimes
flabelliform clusters when young, dimidiate, semicircular at maturity, smooth when present.
Pileus surface convex, furrow, glabrous, glossy, laccate, and consistently hard when fresh,
and tough and light in weight when dry. Pileus color distinct concentric zones with light
brown (6D4) to brown (6E8) at the center, slightly pale orange (5A3) to white (5A1) at the
margin when young, becoming reddish-brown (8E6) to dark brown (9F8) at the center,
dull red (8C4), and pale red (7A3) to white (8A1) at the margin at old age on the upper
surface, brown to dark brown when dried, separated by a layer of context, usually brown
to gray in winter or may fade as weathering destroys pigments on the pileus surface.
Tube layers 0.2–1.2 cm in length, 50–180 µm in width, thick-walled. Stipe 3–5 cm in length,
1.5–3.5 cm in width, 1.3–3.3 cm thick, applanate, with umbo that slightly extended at the
base. Margin up to 0.5–3 cm thick, thinner and lighter than the base, soft, round, pale yellow
(4A3) to grayish brown (5D3) and reddish gray (8B2), changing to grayish-orange (6E3)
when touched, thick toward the margin and downward toward the poreless marginal part
of hymenophore. Pore angular, 4–6 in number in number per mm. Pore surface white (2A1)
to pale yellow (2A3) in growing specimen, immediately discolored when bruised, cream to
grayish brown (5D3) when fresh, with brownish gray (5C2) to brown (4D7) when dried.
J. Fungi 2021, 7, 819 53 of 83
J. Fungi 2021, 7, x FOR PEER REVIEW 58 of 91

Figure 23.
Figure 23. Morphology
MorphologyofofGanoderma
Ganodermaapplantum
applantum (MFLU
(MFLU 19‐2175):
19-2175): (a,b)
(a,b) mature
mature basidiomes;
basidiomes; (c) sec‐
(c) section
tion
of theofbasidiomes;
the basidiomes; (d)characteristics;
(d) pore pore characteristics; (e) hyphae
(e) hyphae of poreof pore (f)
layers; layers; (f) generative
generative hyphae
hyphae of context of
context in Melzer’s reagent; (g,h) thick‐walled with sparing branched of generative and skeletal hy‐
in Melzer’s reagent; (g,h) thick-walled with sparing branched of generative and skeletal hyphae
phae of context in Melzer’s reagent; (i–k) basidiospores in Melzer’s reagent. Scale bars: (a,b) = 2 cm,
of context in Melzer’s reagent; (i–k) basidiospores in Melzer’s reagent. Scale bars: (a,b) = 2 cm,
(d) = 50 μm, (e) = 150 μm, (f) = 10 μm, (g) = 30 μm, (h) = 20 μm, (i–k) = 5 μm.
(d) = 50 µm, (e) = 150 µm, (f) = 10 µm, (g) = 30 µm, (h) = 20 µm, (i–k) = 5 µm.
Ganoderma
Hyphal australe
structure: Hyphal(Fr.)system
Pat., Bull. Soc. mycol.
di-trimitic, Fr. 5(2, generative
contextual 3): 65 (1889)hyphae, binding,
Facesoffungi number: FoF 02906
and skeletal hyphae, generative hyphae, 1–5 µm broad, with clamp connections, hyaline,
Characteristics
thin-walled; bindingfollow
hyphae Luangharn et al. [59].
2–6 µm broad, thin to thick-walled, branched, with clamp
Description: Basidiomes applanate,
connections; skeletal hyphae, hyaline, pale spathulate.
brown toPileus
brown,2–7 cm in length,
thick-walled, 2–71–15 cm in
µm broad.
width, and 0.5–3 cm thick near the base. Pileus circular, applanate, spathulate,
Basidiospores (6.4)6.9–9.3–10.4(11.1) × (5.8)6.4–7.9–8.8(9.7) µm, (x = 9.4 × 7.7 µm, n = 50), sometimes
flabelliform
with clusters
Q = 1.20–1.21, L =when young,
9.63 µm, W =dimidiate, semicircular
7.96 µm (including at maturity,(5.4)6.3–7.0–8.1(8.9)
myxosporium), smooth when pre‐
sent.
× Pileus surface convex,
(3.4)3.8–5.9–6.5(7.3) µm, (xfurrow,
= 7.1 ×glabrous,
5.8 µm, nglossy,
= 50) µm,laccate,
withandQ =consistently
1.19–1.25, L hard
= 7.29when
µm,
fresh,
W and
= 5.93 µmtough and light
(excluding in weight
outer when dry.reddish-brown,
myxosporium), Pileus color distinct
mostly concentric zones with
broadly ellipsoid at
light brown (6D4) to brown (6E8) at the center, slightly pale orange (5A3)
maturity, some distinct tapering at the distal end, truncate, double wall, thick-walled inner to white (5A1)
at the margin Basidia
endosporium. when young, becoming
not observed. reddish‐brown
Cultures (8E6)
characteristics to dark
turned brown
white after(9F8) at the
incubation
center,
◦ dull red (8C4), and pale red (7A3)
at 30 C for 10 days. Odor distinctive when dried. to white (8A1) at the margin at old age on the
upper surface,
Habitat: brownon
Solitary to hardwood
dark brown whenrobtusa,
Shorea dried, separated by a layer
or rotten wood, deadoftrunks,
context, usually
decaying
brown to gray
hardwood, in winter
decaying or may
stumps, andfade as weathering
occasionally destroys
occurring on pigments on the
standing trees orpileus
trunkssur‐
of
face. Tube
many layerstrees.
broad-leaf 0.2–1.2 cm in length, 50–180 μm in width, thick‐walled. Stipe 3–5 cm in
length, 1.5–3.5 examined:
Specimens cm in width, 1.3–3.3 cm Chiang
THAILAND, thick, applanate, with August
Mai Province, umbo that slightly
2012; extended
T. Luangharn,
at the base. Margin up
MFLU 13-0534, MFLUCC 12-0527. to 0.5–3 cm thick, thinner and lighter than the base, soft, round,
pale yellow (4A3) to grayish brown (5D3) and reddish gray (8B2), changing to grayish‐
J. Fungi 2021, 7, 819 54 of 83

Ganoderma casuarinicola J.H. Xing, B.K. Cui and Y.C. Dai., MycoKeys 34: 93–108
(2018)
Facesoffungi number: FoF 06130
Taxonomy and phylogeny analysis are shown in Luangharn et al. [31]
Notes: Ganoderma casuarinicola was collected on a Pinus kesiya stump in a pine forest.
This fungus is distinctive by its strongly laccate, shallow sulcate, reddish-brown pileus
surface, lateral stipe, white pore surface, and brown context. Thai G. casuarinicola shows
its annual, applanate to dimidiate, 3–16 cm long and 1.5–3 cm wide pileus, larger than
Guangdong collection. Our G. casuarinicola collections show longer tubes of 6–14 mm,
while the tubes of the Guangdong collection are 9 mm long; however, our collections reveal
a thinner margin (0.8–1.2 cm thick) than the Guangdong collection (2 cm thick). However,
the type of G. casuarinicola from the Guangdong collection does not have the melanoid
band [37], while our collection features a dark brown melanoid band. Micro- characteristics
are dense light brown to brown context layers; walls of varying thickness in generative
hyphae; thin-walled binding hyphae; and a thick-walled skeletal. Our G. casuarinicola
collection has mostly distinctive yellowish-brown basidiospores, with a smaller size range
of (8.7)10.8–13.5(14.4) × (6.6)7.6–8.9(9.8) µm than the type of G. casuarinicola (8.3-)9.0–10.2
(-11.5) × (4.5-)5.0–6.0(-7.0) µm (including myxosporium).
Ganoderma ellipsoideum Hapuar., T.C. Wen and K.D. Hyde, Mycosphere. 9(5): 951
(2018) (Figure 24)
Facesoffungi number: FoF 06255
Description: Basidiomes annual, sessile. Pileus 3–9 cm in length, 2.2–5 cm in width,
and 1.5–3.5 cm thick at the base. Pileus annual, convex, imbricate, sessile, umbonate,
uneven, ungulate, subflabellate or subdimidiate, somewhat imbricate, when seen from
above flabelliform (fan-shaped), usually round, when present primordial, somewhat round
and plump when present, broadly attached, thick at base, slightly soft at margin when
mature. Pileus surface non-laccate (dull), furrowed, incised, sulcate, smooth when young,
undulating on the upper surface, somewhat spathulate to uneven, covered by a thin and
hard crust (0.1–0.4 mm), and woody when older. Pileus color usually homogenous with
reddish-brown (9E7–9E8) to dark brown (9F7–9F8) at the base at the center, extending
white (4A1) to brown (7E7) on the upper margin surface of mature fruiting bodies. Context
up to 0.5–2.5 cm thick, compact and hard, trimitic hyphal, with clamp connections, hyaline,
thin-walled with simple septa, sparingly branched; generative hyphae 1.2–3.7 µm broad
(n = 30), hyaline, simple septate, with clamp connections; skeletal hyphae 1.8–4.2 µm broad
(n = 30), usually thick-walled, unbranched; binding hyphal 2.0–4.8 µm width (n = 30)
with sparingly branched, thick-walled, without clamp connection. Hymenophore usually
brown (7D8) to reddish-brown (8E8). Tube layers 0.5–2.2 cm in length. Margin blunt-edged,
wavy, slippery when young, and often white (8A1), where the new hyphae are in active
development when young to mature. Pore 4–6 in number per mm, angular, subcircular to
circular. Pore surface white (11A1) to pale yellow (2A3), turns brown (7E7) to dark brown
(7F7–7F8) when scratched or bruised.
Hyphal structure: Hyphal system trimitic, with clamp connections, with brown (7E7);
generative hyphae (1.2-)1.6–2.4–3.0(-3.8) µm broad (n = 30), branched, thin-walled, hya-
line, with grayish-yellow (4B5) in KOH; skeletal hyphae (1.7-)3.1–3.8–4.3(-4.8) µm broad
(n = 30), dextrinoid, abundant thick-walled, with unbranched; binding hyphae (2.1-)3.1–
3.7–4.2(-4.6) µm broad (n = 30), thick-walled, frequently branched, usually intertwined the
generative and skeletal hyphae, mostly brown (7E7) to dark brown (7F5) near the tube
layers; Bovista-type ligative hyphae, hymenial with sword-like apices in the context. Basid-
iospores mostly ellipsoid with double walls, (4.8-)5.3–6.6–7.2(-7.7) × (3.1-)3.5–4.3–5.0(-5.4)
µm (x = 6.8 × 4.5 µm, n = 50) µm, with Q = 1.34–1.43, L = 6.28 µm, W = 4.52 µm (includ-
ing myxosporium), (3.6-)4.1–5.5–6.0(-6.4) × (1.7-)2.1–2.8–3.3(-3.7) µm (x = 5.49 × 2.83 µm,
n = 50) µm, with Q = 1.83–1.92, L = 5.50 µm, W = 2.94 µm (including myxosporium), over-
laid by hyaline, dextrinoid, echinulae, inner wall echinulate brownish-yellow (5C7–5C8) to
brown (7D7–7D8) in 5% KOH. Basidia not seen.
J. Fungi 2021, 7, 819 55 of 83

Habitat: Solitary on rotten wood of Acacia sp.

Specimens examined: THAILAND, Chiang Mai Province, Mae Taeng, Mushroom
J. Fungi 2021, 7, x FOR PEER REVIEW 61 of 91
Research Centre, 19◦ 070 20000 N, 98◦ 410 4400 E, 652 elev., 14 June 2019, P. E. Mortimer,
MFLU 19-2221.

Figure24.
Figure 24. Morphology
Morphology characteristics
characteristicsof
ofGanoderma
Ganodermaellipsoideum
ellipsoideum(MFLU
(MFLU19-2221):
19‐2221): (a)
(a)the
theupper
uppersur-
face of mature basidiomes when fresh; (b) the lower surface of mature basidiomes when fresh. fresh.
surface of mature basidiomes when fresh; (b) the lower surface of mature basidiomes when (c) the
(c) the upper surface of mature basidiomes when dried; (d) margin; (e,f) pore characteristics; (g)
upper surface of mature basidiomes when dried; (d) margin; (e,f) pore characteristics; (g) hyphae of
hyphae of tube layers; (h,i) hyphae from trama in KOH; (j–l) hyphae from trama in KOH; (m–o)
tube layers; (h,i) hyphae from trama in KOH; (j–l) hyphae from trama in KOH; (m–o) basidiospores
basidiospores in KOH reagent. Scale bars: (a–d) = 2 cm, (e,f) = 500 μm, (g) = 200 μm, (g–l) = 20 μm,
in KOH
(m,n) = 5reagent.
μm. Scale bars: (a–d) = 2 cm, (e,f) = 500 µm, (g) = 200 µm, (g–l) = 20 µm, (m,n) = 5 µm.

Ganoderma gibbosum (Blume and T. Nees) Pat., Ann. Jard. Bot. Buitenzorg, suppl. 1:
Ganoderma gibbosum (Blume and T. Nees) Pat., Ann. Jard. Bot. Buitenzorg, suppl. 1:
114 (1897) (Figure 25)
114 (1897) (Figure 25)
Facesoffungi number: FoF 06246
Description: Basidiomes annual or perennial, sessile. Pileus 2–12 cm in length, 2–8 cm
in width, and 0.5–2.8 cm thick. Pileus sessile (without stipe), subflabellate or subdimidiate,
convex, imbricate, umbonate, uneven, round when occurring, primordial, round and
plump when youth, with broadly attached when mature, and thicker at base slightly soft
at margin when mature. Pileus surface non‐laccate (dull), smooth, soft, slightly dull and
J. Fungi 2021, 7, 819 56 of 83
J. Fungi 2021, 7, x FOR PEER REVIEW 63 of 91

Figure25.
Figure 25. Ganoderma
Ganoderma gibbosum
gibbosum(MFLU
(MFLU19-2176):
19‐2176): (a)
(a)the
theupper
uppersurface
surfaceof
ofmature
maturebasidiomes;
basidiomes;(b)
(b)the
the
lower surface of mature basidiomes; (c) margin; (d) context morphology; (e) pore characteristics;
lower surface of mature basidiomes; (c) margin; (d) context morphology; (e) pore characteristics; (f)
tube layers; (g) generative hyphae of context in Melzer’s reagent; (h,i) skeletal and binding hyphae
(f) tube layers; (g) generative hyphae of context in Melzer’s reagent; (h,i) skeletal and binding hyphae
of context in Melzer’s reagent; (j) hyphae and clamp connections of context in Melzer’s reagent; (k,l)
of context in Melzer’s reagent; (j) hyphae and clamp connections of context in Melzer’s reagent;
basidiospores in Melzer’s reagent. Scale bars: (a–c) = 2 cm, (d) = 1 cm, (e) = 500 μm, (f) = 200 μm, (g–
(k,l) basidiospores
i,k,l,) in μm.
= 5 μm, (j) = 20 Melzer’s reagent. Scale bars: (a–c) = 2 cm, (d) = 1 cm, (e) = 500 µm, (f) = 200
µm, (g–i,k,l,) = 5 µm, (j) = 20 µm.
Ganoderma lucidum (Curtis) P. Karst., Revue Mycologique Toulouse. 3(9): 17 (1881)
Facesoffungi number: FoF 06246
(Figure 26)
Description: Basidiomes annual or perennial, sessile. Pileus 2–12 cm in length, 2–8 cm
Facesoffungi number: FoF 06250
in width, and 0.5–2.8 cm thick. Pileus sessile (without stipe), subflabellate or subdimidiate,
Description: Basidiomes annual or perennial, sub‐reniform to reniform, stipitate. Pi‐
convex, imbricate, umbonate, uneven, round when occurring, primordial, round and
leus up to 2–6 cm in length, 1.5–3.0 cm in width, and 0.8–2.0 cm thick at the base. Pileus
plump when youth, with broadly attached when mature, and thicker at base slightly soft
stipitate, sub‐reniform to reniform, undefined imbricate, irregular, some laterally, and fla‐
at margin when mature. Pileus surface non-laccate (dull), smooth, soft, slightly dull and
belliform with a contracted, concentrically sulcate zone, irregularly ruptured crust over‐
faded when mature to old, usually silky when young, furrowed, tuberculate to undulate,
lying the context, radial from center extending to the margin, tough when broken, often
uneven, and incised when mature, compact and hard when mature, woody when mature to
thickcovered
old, at center
withslightly
a thicksoft at margin,
crust, and some andcracked
leathery when
crust whenageold.
when broken.
Pileus colorPileus surface
light orange
smooth layer at center from young to old, usually furrowed, incised, undulate
(6A4–6A5) to grayish orange (6B3–6B5) at base toward the center of mature fruiting bodies to sulcate,
somewhat
become spathulate (30E2–30E7)
grayish-green to uneven, someon thewoody
upper or corkywhen
surface texture whentoold,
mature old.weakly
Contextlaccate
up to
when present,
0.4–2.3 cm thick, strongly laccate
tri-dimitic and glossy
hyphal, when mature,
with clamps and usually
connections, weakly laccate
brownish-orange where
(7C7–7C8)
thereddish-brown
to new hyphae are in activemostly
(8D7–8D8), development (margin).
dark brown (7F7) Pileus
near thecolor
tubeusually
layers, brownish‐red
Bovista-type
ligative hyphae, hymenial with sword-like apices in the context. Tube (6A7–6A8)
(8C7–8C8) at the center, slight to reddish‐orange (7B7–7B8), and orange 0.1–0.6 cmon the
long,
upper pileus surface. Context up to 0.4–1.4 cm thick at base, abundant thick‐walled,
up to 80–160 µm in width, and sulcate at different levels. Stipe almost sessile with broadly sub‐
solid hyphae,
attached whenconcentric lines ofwavy,
present. Margin various shades, bearing
blunt-edged, slipperyclamp
when connections, lightyoung,
wet, soft when brown
(6D6) tothan
thinner brownthe (6D8, 6E8),softer
base and with than
darkthebrown (7F7)
center, and melanoid bands.
often white Tube
(8A1) usually
when young hard,
to
brown (7D7) to dark brown (7F7). Stipe up to 8–16 cm in length, up to
mature. Pore angular, subcircular, 4–6 in number per mm in fresh. Pore surface initial 0.8–1.5 cm in width,
eccentric
white stipe,
(7A1), turns cylindrical
light brownto slightly
(7D6) toflattened,
brown (7D8) laccate,
when and reddish‐brown
scratched (8D7–8D8,
or bruised, with a
8E7–8E8) from mature
slippery surface when fresh. to old. Margin often 0.5–1.3 cm, orange (6A7–6A8) upper surface,
and reddish‐yellow (4A8) under surface, thinner than the base, and softer than the center.
J. Fungi 2021, 7, 819 57 of 83

Hyphal structure: Hyphal system trimitric hyaline, with walls varying in thickness
with simple septa, composed of narrow and sparingly branched; generative hyphae 1.3–4.6
µm broad (n = 30), thin-walled and hyaline hyphae; skeletal hyphae 4.0–7.3 µm broad (n =
30), thick-walled and hyaline hyphae; binding hyphae 2.8–6.3 µm broad (n = 30), usually
with walls varying in thickness. Pileipellis a hymeniderm, brownish-orange (6C8) to brown
(6D8), composed of apically acanthus-like branched cells, dextrinoid. Basidiospores (4.2-)
6.5–8.1–10.3(-11.2) × (3.9-)4.8–4.6–5.7(-6.4) µm (x = 8.2 × 4.7 µm, n = 50) µm, with Q =
1.70–1.76, L = 8.38 µm, W = 4.85 µm (including myxosporium), (4.5–)6.1–7.2–8.1(-9.2) ×
(3.9-)4.5–5.3–5.2(-6.9) µm (x = 7.4 × 5.4 µm, n = 50), with Q = 1.31–1.38, L = 7.53 µm,
W = 5.60 µm (excluding outer myxosporium), ellipsoid or some globose with double walls,
overlaid by hyaline, dextrinoid, distinct echinulate, inner wall echinulate brown, light
brown (6D4) to brown (6E8) in 5% KOH. Basidia not seen.
Habitat: Solitary on standing trees of Dendrocalamus strictus.
Specimens examined: THAILAND, Chiang Rai Province, 19◦ 480 2000 N, 100◦ 040 1900 E,
680 m elev., October 2017, T. Luangharn, MFLU 19-2176.
Ganoderma lucidum (Curtis) P. Karst., Revue Mycologique Toulouse. 3(9): 17 (1881)
(Figure 26)
Facesoffungi number: FoF 06250
Description: Basidiomes annual or perennial, sub-reniform to reniform, stipitate. Pileus
up to 2–6 cm in length, 1.5–3.0 cm in width, and 0.8–2.0 cm thick at the base. Pileus stipitate,
sub-reniform to reniform, undefined imbricate, irregular, some laterally, and flabelliform
with a contracted, concentrically sulcate zone, irregularly ruptured crust overlying the
context, radial from center extending to the margin, tough when broken, often thick
at center slightly soft at margin, and leathery when age when broken. Pileus surface
smooth layer at center from young to old, usually furrowed, incised, undulate to sulcate,
somewhat spathulate to uneven, some woody or corky texture when old, weakly laccate
when present, strongly laccate and glossy when mature, and usually weakly laccate where
the new hyphae are in active development (margin). Pileus color usually brownish-red
(8C7–8C8) at the center, slight to reddish-orange (7B7–7B8), and orange (6A7–6A8) on
the upper pileus surface. Context up to 0.4–1.4 cm thick at base, abundant thick-walled,
subsolid hyphae, concentric lines of various shades, bearing clamp connections, light brown
(6D6) to brown (6D8, 6E8), with dark brown (7F7) melanoid bands. Tube usually hard,
brown (7D7) to dark brown (7F7). Stipe up to 8–16 cm in length, up to 0.8–1.5 cm in width,
eccentric stipe, cylindrical to slightly flattened, laccate, and reddish-brown (8D7–8D8,
8E7–8E8) from mature to old. Margin often 0.5–1.3 cm, orange (6A7–6A8) upper surface,
and reddish-yellow (4A8) under surface, thinner than the base, and softer than the center.
Pore 4–6 in number per mm, (70–)110–145(–160) µm, subcircular to circular, sometimes
angular. Pore surface white (11A1) when present, yellowish-white (2A3) to light brown
(7D6) from young to mature, turning brown (7D7–7D8) to dark brown (6F6) when scratched
or bruised.
Hyphal structure: Hyphal system trimitic, with clamp connections, hyaline, walls vary-
ing in thickness with simple septa, sparingly branched, swollen by melanoid bands, usually
brownish-orange (6C7–6C8) to brown (6D7–6D8) in KOH; generative hyphae up to (1.8–)
2.0–2.3–2.7(–3.0) µm broad (n = 30), usually thin-walled, some thick-walled, with clamp con-
nections, branched, and almost hyaline; skeletal hyphae (3.2–)4.3–5.4–6.1(–6.8) µm broad
(n = 30), usually thick-walled with clamp connections, with unbranched; binding hyphae
(2.4-)2.9–4.4–5.0(-5.9) µm broad (n = 30), walls usually varying in thickness with abundant
branched and present the melanoid bands. Basidiospores ellipsoid, some subglobose to
globose with double walls, with a truncate apex, size range of (8.2-)8.8–9.8–10.5 (-11.4) ×
(5.8-)6.2–6.8–7.5(-8.2) µm, (x = 10.0 × 6.9 µm, n = 50) µm, with Q = 1.51–1.57, L = 10.65 µm,
W = 6.92 µm, (including myxosporium), (6.3-)7.1–7.6–8.2(-8.4) × (4.8-)5.4–5.7–6.1(-6.5) µm,
(x = 7.5 × 5.8 µm, n = 50) µm, with Q = 1.26–1.31, L = 7.57 µm, W = 5.89 µm (excluding
outer myxosporium) (n = 50), brownish-orange (6C8, 6D8) to brown (6E5) of endosporium
(inner wall) with brown (7E7–7E8) exosporium (outer wall) in Congo red, brownish-orange
J. Fungi 2021, 7, 819 58 of 83

(6C8) in 5% KOH, and yellowish-brown (5D8) in Melzer’s reagent, mostly overlaid by

hyaline, coarse echinulate, hyaline turgid vesicular appendix.
Habitat: Solitary on decaying hardwood of Dendrocalamus strictus in dry evergreen
forest.
J. Fungi 2021, 7, x FOR PEER REVIEW Specimens examined: THAILAND, Chiang Rai Province, 19◦ 480 2000 N, 100◦ 040 19
65 00ofE,
91
680 m elev., 15 October 2017, T. Luangharn, MFLU 19-2162.

Figure 26.
Figure 26. Morphology
Morphology ofof Ganoderma
Ganoderma lucidum
lucidum(MFLU
(MFLU19‐2162):
19-2162):(a,b)
(a,b)mature
maturebasidiomes; (c)(c)
basidiomes; pore
pore
characteristics; (d) generative hyphae of context in KOH; (e,f) generative. skeletal and binding
characteristics; (d) generative hyphae of context in KOH; (e,f) generative. skeletal and binding
hyphae of context in KOH; (g–i) basidiospores. Scale bars: (g,i) = 2 cm, (c) = 200 μm, (d–f) = 20 μm,
hyphae of context in KOH; (g–i) basidiospores. Scale bars: (g,i) = 2 cm, (c) = 200 µm, (d–f) = 20 µm,
(g–i) = 5 μm.
(g–i) = 5 µm.
Ganodermamultipileum
Ganoderma multipileumDing
DingHou.
Hou. (1950)
(1950) (Figure
(Figure27)
27)
Facesoffungi number: FoF
Facesoffungi number: FoF 0625606256
Description: Basidiomes annual, stipitate. Pileus up to 6–11 cm in length, 4–9 cm in
width, and 1–3 cm thick at base. Pileus sub‐reniform to reniform, subflabellate to flabel‐
late, concentrically sulcate zone, fleshed at center slightly to margin, radial or branched
from center extend to the margin when seen from above, tough when break, often thick
at center slightly soft at margin, and leathery when age when break. Pileus surface weakly
J. Fungi 2021, 7, 819 59 of 83

Description: Basidiomes annual, stipitate. Pileus up to 6–11 cm in length, 4–9 cm in

width, and 1–3 cm thick at base. Pileus sub-reniform to reniform, subflabellate to flabellate,
concentrically sulcate zone, fleshed at center slightly to margin, radial or branched from
center extend to the margin when seen from above, tough when break, often thick at
center slightly soft at margin, and leathery when age when break. Pileus surface weakly
laccate to strong laccate at center when mature to age, and faded or week laccate at active
mycelial (margin), smooth, irregularly ruptured crust overlying the context, some woody
or corky texture when old, usually furrowed, incised, sulcate to undulating, and somewhat
spathulate to uneven on the surface. Pileus color usually homogenous brownish-red
(8C7–8C8) at the base, slight yellowish-red (8A7–8A8, 8B7–8B8) at center, and light brown
(7D5) to brown (7D7–7D8) on the upper margin surface. Context up to 0.3–1.2 cm thick
at the base, with walls varying in thickness, subsolid hyphae, bearing clamp connections,
usually light brown (6D4–6D6) to brown (6D8, 6E8) of hyphae, and dark brown (7F7)
melanoid bands in KOH. Tube hard, usually thin-thick-walled, often with brown (7D7) to
dark brown (7F7). Stipe up to 4–9 cm in length, up to 1–2.5 cm in width, almost eccentric,
sub-cylindrical to cylindrical, plump, strong laccate, often homogeneous with red (9A6) to
brownish-red (9C6), and dark red (10C7) when mature to old. Margin often white (4A1) to
pale yellow (4A4) where the new hyphae are in active development, light brown (7D5) to
brown (7D7) when bruised, strong laccate, wavy, slippery when wet, softer, thin than the
base, and soft than the center. Pore angular, 4–6 in number per mm, (99-)120–154(-170) µm
in diam, subcircular to circular. Pore surface white (11A1) when present, pale yellow (4A3)
to orange white (5A2) when young to mature, light brown (6D4) with age, turning to light
brown (7D6), brown (7D7–7D8) when dried or scratched and bruised.
Hyphal structure: Hyphal system trimitic, with clamp connections, hyaline, walls vary-
ing in thickness with simple septa, sparingly branched, usually light orange (5A4–5A5)
to orange (5A6–5A7) in KOH; generative hyphae up to (1.6-)2.2–3.5–4.6(-5.2) µm broad
(n = 30), usually thick-walled, unbranched, flexuous, and almost hyaline; skeletal hyphae
(3.8-)4.2–5.1–5.9(-6.7) µm broad (n = 30), usually thick-walled, unbranched, sometimes
subsolid; binding hyphae (1.4-)2.2–3.6–4.4(-5) µm broad (n = 30), usually walls vary-
ing in thickness, with flexuous, abundant branched, with Bovista-type binding hyphae,
and occurred the melanoid bands. Basidiospores mostly ellipsoid, some ovoide, trun-
cate at maturity, with double walls, (7.6-)8.8–11.7–12.4(-13.1) × (4.9-)5.3–6.1–6.9(-7.4) µm,
(x = 11.8 × 6.3 µm, n = 50) µm, with Q = 1.84–1.89, L = 11.84 µm, W = 6.32 µm (includ-
ing myxosporium), (5.9-)6.9–8.2–9.3(-10.6) × (4.3-)4.9–5.6–6.0(-6.5) µm, (x = 8.3 × 5.8 µm,
n = 50) µm, with Q = 1.40–1.46, L = 8.36 µm, W = 5.85 µm (excluding outer myxospo-
rium), mostly overlaid by hyaline, brownish-orange (6C8), (6D8) of exosporium (outer
wall), endosporium (inner wall) coarse echinulae, with hyaline turgid vesicular appendix,
with orange (6A7), (6B7) in KOH.
Habitat: Solitary on decaying stump of Pinus merkusii.
Specimens examined: THAILAND, Prachuap Khiri Khan Province, 12◦ 080 5200 N, 99◦ 450 4100 E,
491 m elev., 26 June 2018, T. Luangharn, MFLU 19-2166.
Notes: Ganoderma multipileum was originally reported from Taiwan, PRC [44], which
was presented over half a century ago from Taiwan, PRC [39]. This fungus was the
earliest valid name with G. lucidum from tropical Asia [44]; however, Wang et al. [33]
verified that G. multipileum is the correct name for this tropical fungus. This fungus is a
distinctive form with its laccate to strong laccate pileus, stipitate, rarely sessile, irregularly
ruptured crust overlying the context, yellow-brown to dark brown context, cream pore
surface, flattened or sub-cylindrical, lateral, horizontally lateral, with ovoid to ellipsoid
basidiospores (7.6–13.5 × 5.5–7.5 µm) (with myxosporium), 6.5–10.5 × 4.5–6.5 µm (without
myxosporium)), mostly truncate, brown, with a dark brown [33]. Some researchers have
shown in their phylogenies that G. tropicum is similar to G. multipileum [4,5], which is
considered most resemble G. tropicum in morphology and habitat even though they are
distinct species [33] and G. flexipes [4,76,129]. Our G. multiplicatum was collected from
Thailand, are similar to the original description by Wang et al. [33] by its showed laccate to
J. Fungi 2021, 7, 819 60 of 83

J. Fungi 2021, 7, x FOR PEER REVIEW

strong 67 of 91
laccate pileus, radial or branched from center extends to the margin, with ellipsoid
basidiospores.

Figure 27.
Figure 27. Morphology
Morphologycharacteristics
characteristicsofofGanoderma
Ganodermamultipileum
multipileum(MFLU
(MFLU 19‐2166): (a,b)(a,b)
19-2166): mature
mature
basidiomes; (c) pore characteristics; (d) generative hyphae of context in KOH; (e–g) generative.
basidiomes; (c) pore characteristics; (d) generative hyphae of context in KOH; (e–g) generative.
skeletal and binding hyphae of context in KOH; (h) basidiospores with tube layers; (i–k) basidio‐
skeletal and binding hyphae of context in KOH; (h) basidiospores with tube layers; (i–k) basidiospores
spores in KOH. Scale bars: (a,b) = 3 cm, (c) = 1000 μm, (d–g) = 20 μm, (h) = 50 μm, (i–k) = 5 μm.
in KOH. Scale bars: (a,b) = 3 cm, (c) = 1000 µm, (d–g) = 20 µm, (h) = 50 µm, (i–k) = 5 µm.

Ganodermaorbiforme
Ganoderma orbiforme(Fr.)(Fr.)Ryvarden,
Ryvarden,Mycologia.
Mycologia.92(1):
92(1):187
187(2000)
(2000)Figure
Figure28) 28)
≡ Polyporus orbiformis Fr., Epicrisis Systematis Mycologici.:
≡ Polyporus orbiformis Fr., Epicrisis Systematis Mycologici.: 463 (1838) 463 (1838)
≡≡Fomes
Fomesorbiformis
orbiformis(Fr.)
(Fr.)Cooke,
Cooke,Grevillea.
Grevillea.1414(69):
(69):18
18(1885)
(1885)
≡ Scindalma orbiforme (Fr.) Kuntze, Revisio generum
≡ Scindalma orbiforme (Fr.) Kuntze, Revisio generum plantarum. plantarum.3(2):3(2):519
519(1898)
(1898)
≡≡Ganoderma
Ganodermalucidum
lucidumvar.
var.orbiformis
orbiformis(Fr.)
(Fr.)Rick,
Rick,Iheringia.
Iheringia.7:7:201
201(1960)
(1960)
≡≡Ganoderma
Ganodermaorbiformum
orbiformum(Fr.)
(Fr.)Ryvarden
Ryvarden(2000)
(2000)
== Ganoderma
Ganoderma mastoporum
mastoporum (Lév.)
(Lév.)Pat.,
Pat.,Bulletin
BulletindedelalaSociété
SociétéMycologique
Mycologiquede deFrance.
France.5:
5:
71 (1889)
71 (1889)
== Ganoderma
Ganoderma fornicatum
fornicatum (Fr.)
(Fr.)Pat.,
Pat.,Bulletin
Bulletinde delalaSociété
SociétéMycologique
Mycologiquede deFrance.
France.5:5:71
71
(1889)
(1889)
== Ganoderma
Ganoderma boninense
boninense Pat.,
Pat., Bulletin
Bulletindedela
laSociété
SociétéMycologique
Mycologiquede deFrance
France5:5:72 72(1889)
(1889)
== Ganoderma
Ganoderma subtornatum
subtornatumMurrill,
Murrill,Bulletin
Bulletinofofthe
theTorrey
TorreyBotanical
BotanicalClub.
Club.34:34:477
477(1907)
(1907)
== Ganoderma
Ganoderma cupreum
cupreum(Cooke)
(Cooke)Bres.,
Bres.,Annales
AnnalesMycologici.
Mycologici.9: 9: 268
268 (1911)
(1911)
== Ganoderma
Ganoderma densizonatum
densizonatum J.D.J.D. Zhao
Zhaoand andX.Q.
X.Q.Zhang,
Zhang,Acta
Actamycol.
mycol.sin.:
sin.: 86
86 (1986)
(1986)
== Ganoderma
Ganoderma limushanense
limushanenseJ.D. J.D.Zhao
ZhaoandandX.Q.
X.Q.Zhang,
Zhang,Acta
Actamycol.
mycol.sin.:
sin.: 219
219 (1986)
(1986)
Facesoffungi number: FoF 06257
Description: Basidiomes annual or perennial, sessile. Pileus 4–11 cm in length, 3–6 cm
in width, and 1–2.4 cm thick at base. Pileus sessile, flabelliform or spathulate, convex, im‐
bricate, umbonate, uneven, ungulate, sub‐reniform, sub‐orbicular, subdimidiate, obtuse
from host, broadly attached, somewhat imbricate, thicker at base, slightly soft at margin
J. Fungi 2021, 7, 819 61 of 83
J. Fungi 2021, 7, x FOR PEER REVIEW 69 of 91

Figure 28.
Figure 28. Morphology
Morphology of of Ganoderma
Ganoderma orbiforme
orbiforme (MFLU
(MFLU 17-1933):
17‐1933): (a,b)
(a,b) mature
mature basidiomes;
basidiomes; (c)(c)pore
pore
characteristics; (d,e) context hyphae in KOH; (f–h) basidiospores in KOH. Scale bars: (a) =
characteristics; (d,e) context hyphae in KOH; (f–h) basidiospores in KOH. Scale bars: (a) = 2 cm,2 cm, (c)
= 1000 μm, (d,e) = 20 μm, (f–h) = 5 μm.
(c) = 1000 µm, (d,e) = 20 µm, (f–h) = 5 µm.

Ganoderma philippii
Facesoffungi number: FoFBres.06257
and Henn. ex Sacc. Bres, Iconographia mycological. 21:
1014,Description:
t. 1014 (1932) (Figure 29)
Basidiomes annual or perennial, sessile. Pileus 4–11 cm in length, 3–6 cm
≡ Fomes
in width, and philippii
1–2.4 cm Bres. and
thick at Henn. ex Sacc.,
base. Pileus Sylloge
sessile, Fungorum.or9:spathulate,
flabelliform 180 (1891) convex,
≡ Scindalma philippii (Bres. and Henn. ex Sacc.) Kuntze, Revisio
imbricate, umbonate, uneven, ungulate, sub-reniform, sub-orbicular, subdimidiate, generum plantarum.
obtuse
3(2): 519
from host,(1898)
broadly attached, somewhat imbricate, thicker at base, slightly soft at margin
when=mature.
Fomes pseudoferreus
Pileus color Wakef., Bulletin of Miscellaneous
usually homogenous Informations
with grayish-orange of thebrownish-
(6B3–6B6), Royal Bo‐
tanical Gardens
orange (6C5–6C6),Kew. and1918: 208 (6D7–6D8)
brown (1918) at base extending to the margin of maturity
Facesoffungi
to old. number:
Pileus surface FoFnon-laccate
partly 06258 (dull) to weakly laccate, faded texture when
Description:incised,
old, furrowed, Basidiomes annual,
sulcate, sessile. Pileus
undulating, 2–9 cm inspathulate
and somewhat length, 2–5to cm in width,
uneven on and
the
0.4–3.8surface,
upper cm thicksilky
at theandbase.
softPileus
whenannual, convex,when
fresh, woody sessile, umbonate,
mature ungulate,
to older, usually
and covered
roundcompact
with when present,
and hard primordial, plump when
crust (0.1–0.25 mm). present,
Contextbroadly attachedcm
up to 0.4–1.0 and thicktrimitic
thick, at base,
hyphae
slightly with clamp
soft at margin.connections, hyaline,
Pileus surface walls varying
non‐laccate in thickness
to weakly laccate, with simpleincised,
furrowed, septa,
sparingly branched.
sulcate, smooth when Hymenophore brownon
young, undulating (6D7–6D8)
the uppertosurface,
reddish-brown (8D7).
spathulate Tube layers
to uneven, cov‐
0.3–1.2 cmaincrust
ered with length, lightmm),
(0.1–0.3 brown (6D4–6D5)
cracked to brown
crust when (7D7).
old, and Stipe
woody almost
when old.sessile
Pileus with
color
broadly attached when present, with brownish-orange (6C8). Margin soft, wavy, blunt-
J. Fungi 2021, 7, 819 62 of 83

edged, slippery when young, with grayish-orange (6B5–6B6) to brownish-orange (6C8)

when old. Pore 4–6 in number per mm, circular or subcircular, or angular. Pore surface
white (11A1), when present, turns light brown (7D5) to brown (7D7–7D8) when scratched
or bruised, becoming discolored when touched.
Hyphal structure: Hyphal system trimitic, with clamp connections, usually orange
(6A6–6A8) to brownish-orange (6C7–6C8) in KOH; generative hyphae 1.4–5.0 µm broad
(n = 30), hyaline, thin-walled with clamp connections; skeletal hyphae 2.6–5.6 µm broad
(n = 30), usually thick-walled, unbranched; binding hyphae 1.2–4.8 µm width (n = 30),
usually walls varying in thickness, many branches, hymenial with sword-like apices
in the context. Basidiospores mostly ellipsoid to oblong ellipsoid or broadly ellipsoid,
with double walls, (7.1-)7.9–9.4–11.2(-11.8) × (5.2-)5.97–6.7–7.1(-7.7) µm (x = 9.6 × 6.8 µm,
n = 50) µm, with Q = 1.38–1.44, L = 9.63 µm, W = 6.82 µm (including myxosporium),
(6.4-)5.2–6.0–6.7(-10.7) × (3.9-)4.5–5.3–6.1(-6.6) µm (x = 6.2 × 5.1 µm, n = 50) µm, with
Q = 1.20–1.27, L = 6.28 µm, W = 5.10 µm (excluding outer myxosporium), overlaid by
hyaline, echinulate, inner wall echinulate brown, some turgid vesicular appendix, light
yellow (4A5) to reddish-yellow (4B7–4B7) in 5% KOH. Basidia not seen.
Habitat: solitary, on the living tree of Albizia mollis in deciduous forest, and living tree
of Indochinese spp.
Specimens examined: THAILAND, Chiang Rai Province, 19◦ 480 2000 N, 100◦ 040 1900 E,
680 m elev., 21 October 2017, T. Luangharn, MFLU 17-1933.
Notes: Ganoderma orbiforme (Fr.) Ryvarden was first described as Polyporus orb-
iformis, with the original specimen from the tropical region of Guinea in Africa [136].
The fungus was characterized by its distinctive weakly laccate surface or some dull surface
areas, brown context, brown pore surface, and brown tube layer, and ellipsoid or ovoid
basidiospore. Ganoderma cupreum, G. densizonatum, G. fornicatum, G. limushanense,
G. multiplicatum, and G. subtornatum are similar to G. orbiforme; however, taxonomy and
molecular analysis treated those taxa as the earliest valid names for G. orbiforme [7,156].
This fungus has been recorded from China, Laos, Myanmar, and Thailand [30], with our
collection is also the collection of G. orbiforme from Thailand. Our collection from Thailand
agrees well with the description by Ryvarden [136] and Wang et al. [156] reported that this
fungus posed rigid basidiomes, purplish-black laccate crust, brown pore surface, and tube
layer, with ellipsoid or ovoid basidiospores, fine and short echinulate, clavate cells usually
with several irregular lobes.
Ganoderma philippii Bres. and Henn. ex Sacc. Bres, Iconographia mycological. 21:
1014, t. 1014 (1932) (Figure 29)
≡ Fomes philippii Bres. and Henn. ex Sacc., Sylloge Fungorum. 9: 180 (1891)
≡ Scindalma philippii (Bres. and Henn. ex Sacc.) Kuntze, Revisio generum plantarum.
3(2): 519 (1898)
= Fomes pseudoferreus Wakef., Bulletin of Miscellaneous Informations of the Royal
Botanical Gardens Kew. 1918: 208 (1918)
Facesoffungi number: FoF 06258
Description: Basidiomes annual, sessile. Pileus 2–9 cm in length, 2–5 cm in width,
and 0.4–3.8 cm thick at the base. Pileus annual, convex, sessile, umbonate, ungulate,
usually round when present, primordial, plump when present, broadly attached and thick
at base, slightly soft at margin. Pileus surface non-laccate to weakly laccate, furrowed,
incised, sulcate, smooth when young, undulating on the upper surface, spathulate to
uneven, covered with a crust (0.1–0.3 mm), cracked crust when old, and woody when
old. Pileus color usually homogenous, brown (7E7–7E8) at base at the center, extending to
brownish-orange (6C8), and white (4A1) when present, slightly yellowish-white (2A2) on
the upper margin surface of mature fruiting bodies. Context consists of trimitic hyphae,
up to 0.3–1.8 cm thick, sparingly branched, walls varying in thickness, compact and hard,
with clamp connections, hyaline; generative hyphae 1.8–3.3 µm in width (n = 30), brownish-
yellow (5C7–5C8), thin-walled, simple septa, hyaline, with clamp connections; skeletal
hyphae 2.8–4.9 µm broad (n = 30), brownish-yellow (5C7–5C8), usually thick-walled,
J. Fungi 2021, 7, 819 63 of 83

unbranched; binding hyphae 2.0–4.2 µm width (n = 30), brownish-yellow (5C7–5C8) to

brownish-orange (6C7–6C8), sparingly branched, walls varying in thickness, and without
clamp connections. Hymenophore heterogeneous, brown (7C7–7C8, 7D8), and melanoid
band present when mature. Tube layers 0.3–3.8 cm in length. Margin blunt-edged, wavy,
and often white (8A1) where the new hyphae are in active development to yellowish-
white (2A2) from young to mature. Pore 4–7 in number per mm, subcircular to circular.
J. Fungi 2021, 7, x FOR PEER REVIEW 71 of 91
Pore surface white (11A1) when present, turning brown (7E7) to dark brown (7F7–7F8)
when scratched or bruised.

Figure29.
Figure 29.Morphology
MorphologyofofGanoderma
Ganoderma philippii
philippii (MFLU
(MFLU 19‐2222):
19-2222): (a)(a) mature
mature basidiomes
basidiomes of collection
of the the collec‐
MFLU 19-2222; (b,c) mature basidiomes of the collection MFLU 19-2223; (d) morphology of the of
tion MFLU 19‐2222; (b,c) mature basidiomes of the collection MFLU 19‐2223; (d) morphology the
tube
tube layers; (e,f) pore characteristics; (g,h) context hyphae; (i) hyphae from tube layers; (j–l) basidi‐
layers; (e,f) pore characteristics; (g,h) context hyphae; (i) hyphae from tube layers; (j–l) basidiospores
ospores in KOH. Scale bars: (a–d) = 2 cm, (e) = 500 μm, (f) = 150 μm, (g–i) = 20 μm, (j–l) = 5 μm.fi.
in KOH. Scale bars: (a–d) = 2 cm, (e) = 500 µm, (f) = 150 µm, (g–i) = 20 µm, (j–l) = 5 µm.fi.
Ganoderma sichuanense S.H. Wu, Y. Cao, and Y.C. Dai. Fungal Diversity. 56(1): 54
(2012) (Figure 30)
Facesoffungi number: FoF 06248
Description: Basidiomes annual or perennial, stipitate. Pileus 15 cm in length, up to 10
cm in width, and 2 cm thick. Pileus shell‐like (involute from margin into the center), sub‐
flabellate or reniform to circular when seen from above, often with undefined concentric
J. Fungi 2021, 7, 819 64 of 83

Hyphal structure: Hyphal system trimitic, with clamp connections, usually brown (7E7);
generative hyphae (1.7-)2.1–2.5–2.9(-3.3) µm broad (n = 30), branched, thin-walled, hya-
line, unbranched, grayish-yellow (4B5) in KOH, with clamp connections; skeletal hyphae
(2.9-)3.4–4.0–4.4(-5.0) µm broad (n = 30), dextrinoid, abundant thick-walled, with un-
branched; binding hyphae (2.1-)2.8–3.3–3.8(-4.3) µm broad (n = 30), walls varying in
thickness, frequently branched, generative and skeletal hyphae usually intertwined, mostly
brown (7E7) to dark brown (7F5) near the tube layer; Bovista-type ligative hyphae, hy-
menial with sword-like apices in the context. Basidiospores mostly oblong, double
walls, (5.8-)6.4–7.3–7.8(-8.2) × (4.1-)4.6–6.1–6.7(-7.1) µm (x = 7.4 × 6.0 µm, n = 50) µm,
with Q = 1.01–1.12, L = 7.32 µm, W = 6.98 µm (including myxosporium), (5.2-)5.7–6.5–7.1
(-7.5) × (3.7-)4.3–5.5–6.1(-6.5) µm (x = 6.62 × 5.58 µm, n = 50) µm, with Q = 1.14–1.24,
L = 6.56 µm, W = 5.48 µm (including myxosporium), overlaid by hyaline, dextrinoid, ech-
inulate, inner wall echinulate, grayish-orange (5B4–5B5) in 5% KOH, and yellowish-brown
(5E8) to light brown (6D8). Basidia not seen.
Habitat: Solitary on rotten wood of unknown tree species.
Specimens examined: THAILAND, Chiang Mai Province, Mae Taeng, Mushroom Re-
search Centre, 19◦ 070 2000 N, 98◦ 410 4400 E, 770 m elev., 14 June 2019, P. E. Mortimer, MFLU 19-
2222 and MFLU 19-2223.
Notes: Ganoderma philippii was introduced as Fomes philippii by Bresadola and Hen-
nings in 1891 [126] and later transferred to Ganoderma [157]. Ganoderma philippii causes red
root-rot disease, one of the most economically important diseases across a wide range of
commercial perennial woody crops of tropical Acacia species [158]. Ganoderma philippii is dis-
tributed across Asia, from the Philippines in the north to southern Papua New Guinea [111].
This species is characterized by its sessile, non-laccate pileus surface, with melanoid bands
that form a layer distinct from the context, with a di-trimitic hyphal system with clamped
connections overlaid by hyaline, echinulate, inner wall echinulate, sometimes turgid vesic-
ular appendix basidiospores [26]. In this study, our new record of G. philippii collected from
Thailand is described based on characteristics and phylogenetic analyses. Our collection
agrees well with the description by Wang et al. [33].
Ganoderma sichuanense S.H. Wu, Y. Cao, and Y.C. Dai. Fungal Diversity. 56(1): 54
(2012) (Figure 30)
Facesoffungi number: FoF 06248
Description: Basidiomes annual or perennial, stipitate. Pileus 15 cm in length, up to
10 cm in width, and 2 cm thick. Pileus shell-like (involute from margin into the center),
subflabellate or reniform to circular when seen from above, often with undefined con-
centric zones at center and extend to the margin, thick at center slightly soft at margin,
and leathery when age when break. Pileus surface usually laccate, faded or week laccate
when young, and strongly laccate when mature to age, shiny, silky, smooth, and soft
when fresh, furrowed on the surface with sulcate to undulating, somewhat spathulate to
uneven, incised, hard, and woody when old, and some occurred the brown (7D5) lined
when older. Pileus color usually homogenous at base at the center with red (9A6–9A7,
9B7–9B8), brownish-red (9C6–9C8), and dark red (10C7–10C8), extended to the margin with
reddish-yellow (4A7), but do not change the color when touched. Context up to 0.3–0.5 cm
thick at base, abundant thick-walled, some thin-walled, with subsolid hyphae, bearing
clamp connections, and occurred the dark brown (7F7–7F8) melanoid bands. Tube layers
usually thin-walled, frequently branched with clamp connections, and hard and woody
when mature, often brown (7D7). Stipe up to 3–8 cm in lenge, up to 1–3 cm in width,
centrally stipitate, almost sub-cylindrical to cylindrical, often red (9A6) to brownish-red
(9C6), and dark red (10C7) when mature. Margin obtuse from the center, some wavy,
slippery when wet, softer, strong laccate edged, thin than the base and soft than the center,
often reddish-yellow (4A7, 4B7–4B8), deep yellow (4A8), and orange (5A6–5A7) to golden
yellow (5B7–5B8) when mature to old. Pore angular, subcircular to circular, 4–7 in number
per mm, (40-)80–140(-155) µm. Pore surface white (11A1) to yellowish-white (3A2) when
young to mature, turning yellow (3B8) to olive-yellow (3C7–3C8) when dried, as well as
J. Fungi 2021, 7, 819 65 of 83

J. Fungi 2021, 7, x FOR PEER REVIEW discolored when touched, turns light brown (7D5), brown (7D7–7D8) to73dark
becoming of 91
brown (7F6–7F8) when scratched or bruised.

Figure30.
Figure 30. Morphology
Morphology of of Ganoderma
Ganodermasichuanense
sichuanense(MFLU
(MFLU19-2164):
19‐2164):(a,b)
(a,b)mature
maturebasidiomes;
basidiomes;(c)(c)pore
pore
characteristics; (d–f) walls varying in thickness sparing branched of generative and skeletal
characteristics; (d–f) walls varying in thickness sparing branched of generative and skeletal hyphae ofhyphae
of context in upper part in Melzer’s reagent; (g) generative hyphae of context in lower part in
context in upper part in Melzer’s reagent; (g) generative hyphae of context in lower part in Melzer’s
Melzer’s reagent; (h,i) thick‐walled sparing branched of generative and skeletal hyphae of context
reagent; (h,i) thick-walled sparing branched of generative and skeletal hyphae of context in lower
in lower part in Melzer’s reagent; (j–l) basidiospores in Melzer’s reagent. Scale bars: (a,b) = 2 cm, (c)
part
= 500inμm,
Melzer’s
(d) = reagent; (j–l) basidiospores
50 μm, (e,f,h,i) = 20 μm, (g)in= Melzer’s reagent.
30 μm, (j–l) Scale bars: (a,b) = 2 cm, (c) = 500 µm,
= 5 μm.
(d) = 50 µm, (e,f,h,i) = 20 µm, (g) = 30 µm, (j–l) = 5 µm.
Ganoderma sinense J.D. Zhao, L.W. Hsu, and X.Q. Zhang, Acta Mycologica Sinica.
Hyphal structure: Hyphal system trimitic, with bearing clamp connections, hyaline,
19: 272 (1979) (Figure 31)
with walls varying in thickness with simple septa, composed of narrow and sparingly
Facesoffungi number: FoF 06253
branched; generative hyphae (1.3-)1.6–1.8–2.2(-2.4) µm broad (n = 30), thin-walled, hyaline,
Description: Basidiomes annual, stipitate, subdimidiate. Pileus 4–10 cm in length, 6–
branched, with clamp connections; skeletal hyphae (1.6-)2.2–3.5–4.2(-4.9) µm broad (n = 30),
12 cm in width, 0.3–1.3 cm thick at base. Pileus stipitate, subdimidiate to dimidiate, flabel‐
walls varying in thickness, sometimes subsolid; binding hyphae (1.2-)1.5–1.7–2.0(-2.3) µm
liform, spathulate, umbonate, concentrically sulcate zone, radial from center extending to
broad (n = 30), usually thick-walled with narrow to subsolid, with pale orange (5A3) to
the margin, tough when broken, often thick at center, slightly soft at margin, light in
light orange (5A4) of thin-walled, and orange (6B7) to brownish-orange (6C8) of thick-
weight when dried, and non‐woody. Pileus surface laccate, convex, some radial furrowed,
imbricate, incised, glossy, shiny, spathulate, shallow sulcate when fresh, umbonate or un‐
even, strongly laccate and glossy when mature, and weakly laccate where the new hyphae
J. Fungi 2021, 7, 819 66 of 83

walled in Melzer’s reagent, and occurred the melanoid bands. Pileipellis a hymeniderm,
brown (7D7) with clavate-like cells, dextrinoid. Basidiospores mostly ellipsoid or oblong
ellipsoid, truncate at maturity, with double walls, (8.0-)9.4–11.3–12.2(-13.3) × (4.9-)5.4–6.5–
7.1(-7.4) µm, (x = 11.2 × 6.6 µm, n = 50) µm, with Q = 1.64–1.51, L = 11.24 µm, W = 6.69 µm
(including myxosporium), (7.8-)8.1–8.6–9.9(-11.7) × (4.9-)5.2–5.9–6.7(-7.4) µm (Qm = 1.5,
Q = 1.0–2.0, excluding outer myxosporium) (n = 50), (x = 8.7 × 5.8 µm, n = 50) µm, with
Q = 1.46–1.53, L = 8.72 µm, W = 5.84 µm (excluding outer myxosporium), overlaid by
hyaline, exosporium (outer wall) smooth and hyaline, endosporium (inner wall) coarse
echinulate, with turgid vesicular appendix, light brown (7D7) to brown (7D8) in KOH,
and reddish-brown (8E6) to dark brown (8F4) in Melzer’s reagent. Basidia 12–17 × 6–9 µm,
with four sterigmata.
Habitat: Occasionally occurring on the stump of decaying Pterocarpus sp., living tree.
Specimens examined: THAILAND, Prachuap Khiri Khan Province, 12◦ 080 5200 N, 99◦ 450 4100 E,
491 m elev., 25 June 2018, T. Luangharn, MFLU 19-2164.
Ganoderma sinense J.D. Zhao, L.W. Hsu, and X.Q. Zhang, Acta Mycologica Sinica. 19:
272 (1979) (Figure 31)
Facesoffungi number: FoF 06253
Description: Basidiomes annual, stipitate, subdimidiate. Pileus 4–10 cm in length,
6–12 cm in width, 0.3–1.3 cm thick at base. Pileus stipitate, subdimidiate to dimidiate, fla-
belliform, spathulate, umbonate, concentrically sulcate zone, radial from center extending
to the margin, tough when broken, often thick at center, slightly soft at margin, light in
weight when dried, and non-woody. Pileus surface laccate, convex, some radial furrowed,
imbricate, incised, glossy, shiny, spathulate, shallow sulcate when fresh, umbonate or
uneven, strongly laccate and glossy when mature, and weakly laccate where the new
hyphae are in active development (margin), usually smooth layers at center from young
to old age, irregularly ruptured crust overlying the context, and leathery when broken in
old age. Pileus color usually homogenous and brownish-red (8C7–8C8) to reddish-brown
(8D7–8D8) at center toward stipe and extending brownish-red (9C8) from the center slight
to the margin when mature, usually reddish-brown (8E5–8E8) upper margin surface when
old. Context up to 0.3–0.8 cm thick near the stipe, dried, upper layers brownish-orange
(6C8) when fresh, lower layers grayish-orange (5B5), dark brown (8F7) when dried, soft and
fibrous, covered with thin crust, some present woody, trimitic hyphae, hyaline, walls vary-
ing in thickness with simple septa, with branches. Tube 0.3–0.9 cm in length, brown (7D8).
Stipe 3–12 cm in length, sub-cylindrical to cylindrical, almost stipitate, with broad and thick
at base, irregularly ruptured crust overlying, usually strongly laccate, brown (7D8) to dark
brown (8F8) when mature, usually dark brown (8F8) when old. Margin soft, some wavy,
laccate when mature, weakly laccate when old, brownish-orange (6D8) from mature to
old. Pore 4–6 in number per mm, subcircular to circular. Pore surface white (11A1) when
present, to yellowish-white (2A2) when mature, turns brownish (6E7) to dark brown (6F7)
when scratched or bruised.
Hyphal structure: Hyphal system trimitic, with clamp connections, usually light orange
(5A5) to orange (5A7), golden yellow (5B7–5B8), sometimes brownish-red (8C7) in KOH;
generative hyphae 1.2–2.1 µm broad (n = 30), hyaline, thin-walled, with clamp connections;
skeletal hyphae 3.2–5.4 µm broad (n = 30), usually hyaline, thick-walled, unbranched, and solid;
binding hyphae 3.3–5.7 µm width (n = 30), walls varying in thickness, with many branches,
hymenial with sword-like apices in the context. Pileipellis a hymeniderm, grayish-orange (5B5)
to brown (6E8), clavate-like cells, dextrinoid. Basidiospores mostly ellipsoid to broadly ellipsoid,
double walls, (9.8-)10.4–11.7–12.5(-13.4) × (7.3-)7.7–8.9–10.0(-10.6) µm (x = 11.8 × 9.0 µm, n = 50)
µm, with Q = 1.29–1.34, L = 11.82 µm, W = 9.02 µm (including myxosporium), (8.9-)9.8–10.4–
12.0 (-12.8) × (6.2-)6.8–7.2–7.9(-8.3) µm (x = 10.6 × 7.3 µm, n = 50) µm, with Q = 1.41–1.47,
L = 10.47 µm, W = 7.25 µm (excluding outer myxosporium), overlaid by hyaline, apically
and shortly echinulate, truncate, some turgid vesicular appendix, inner wall echinulate,
orange (5A7), deep (5A8, 5B8), orange (6B8), brownish-orange (6B8), outer wall usually
brownish-red (8C7–8C8) in 5% KOH.
J. Fungi 2021, 7, 819 67 of 83

Habitat: Solitary, on the living tree of Dendrocalamus strictus and Dipterocarpus sp.
J. Fungi 2021, 7, x FOR PEER REVIEW Specimens examined: THAILAND, Nakhon Phanom Province, 22 December 75 of 91
2018,
T. Luangharn, MFLU 19-2172.

Figure 31.
Figure 31. Morphology
Morphology of ofGanoderma
Ganodermasinense
sinense(MFLU
(MFLU19‐2172):
19-2172):(a,b) mature
(a,b) basidiomes;
mature basidiomes;(c) (c)
porepore
characteristics; (d–f) generative skeletal and binding hyphae of context in KOH; (g–j) basidio‐
characteristics; (d–f) generative skeletal and binding hyphae of context in KOH; (g–j) basidiospores
spores in KOH. Scale bars: (a,b) = 2 cm, (c) = 1000 μm, (d–f) = 20 μm, (g–j) = 5 μm.
in KOH. Scale bars: (a,b) = 2 cm, (c) = 1000 µm, (d–f) = 20 µm, (g–j) = 5 µm.

Ganoderma subresinosum
Ganoderma subresinosum(Murrill)
(Murrill)C.J.
C.J.Humphrey,
Humphrey,Mycologia.
Mycologia. 30 30 (3):
(3): 332
332 (1938)
(1938)
(Figure32)
(Figure 32)
≡≡Fomes
Fomessubresinosus
subresinosusMurrill,
Murrill,Bulletin
Bulletinofofthe
theTorrey
TorreyBotanical
BotanicalClub.
Club.35: 35:410
410(1908)
(1908)
≡≡Trachyderma
Trachyderma subresinosum
subresinosum (Murrill)
(Murrill)Imazeki,
Imazeki,Bulletin
Bulletinofofthethe
Government
Government Forest Ex‐
Forest
perimental Station
Experimental Meguro.
Station Meguro. 57:57:
119119(1952)
(1952)
≡≡Magoderna
Magodernasubresinosum
subresinosum(Murrill)
(Murrill)Steyaert,
Steyaert,Persoonia.
Persoonia.7(1):
7(1):112
112(1973)
(1973)
≡≡Amauroderma
Amaurodermasubresinosum
subresinosum(Murrill)
(Murrill)Corner,
Corner,Beihefte
BeiheftezurzurNova
NovaHedwigia.
Hedwigia.75: 75:9393
(1983)
(1983)
== Polyporus
Polyporus mamelliporus
mamelliporus Beeli,
Beeli, Bulletin
Bulletin de
de la
la Société
Société Royale
Royale dede Botanique
Botanique de de Belgique.
Belgique.
62(1):
62(1): 62
62 (1929)
(1929)
Facesoffungi
Facesoffunginumber:
number: FoF
FoF 06259
06259
Description: Basidiomes annual, subdimidiate,sessile.
Description: Basidiomes annual, subdimidiate, sessile. Pileus
Pileus 6–136–13
cm incmlength,
in length,
4–10
4–10
cm incm in width,
width, and 0.6–2
and 0.6–2 cm thick.
cm thick. PileusPileus stipitate,
stipitate, flabelliform,
flabelliform, spathulate,
spathulate, umbonate,
umbonate, sub‐
subdimidiate to dimidiate,
dimidiate to dimidiate, single,
single, concentrically
concentrically sulcate
sulcate zone,
zone, radial
radial from
from center
center extending
extending to
to
thethe margin,tough
margin, toughwhen
whenbroken,
broken,often
oftenthicker
thickeratatcenter,
center, slightly
slightly soft at margin,
margin, andandlight
light
in
in weight
weight when
when dried.
dried. Pileus
Pileus surface
surface convex, glossy,
glossy, shiny,
shiny,usually
usually frequently
frequently furrowed,
furrowed,
shallow
shallow sulcate, mostly rugulose, spathulate, umbonate to uneven, laccate when mature,
sulcate, mostly rugulose, spathulate, umbonate to uneven, laccate when mature,
strongly laccate when old, concentrically sulcate with irregularly ruptured crust overlying
the context, hard from mature to old, and woody or corky texture when old. Pileus color
usually dark brown (9F7–9F8) from center to the margin. Context up to 0.3–1 cm thick
near the base, dry, upper layers grayish‐orange (5B6) when fresh, lower layers grayish‐
J. Fungi 2021, 7, 819 68 of 83

strongly laccate when old, concentrically sulcate with irregularly ruptured crust overlying
the context, hard from mature to old, and woody or corky texture when old. Pileus color
usually dark brown (9F7–9F8) from center to the margin. Context up to 0.3–1 cm thick near
the base, dry, upper layers grayish-orange (5B6) when fresh, lower layers grayish-orange
(5B5), dark brown (8F7) when dried, covered with crust, woody when dried, trimitic
hyphae, hyaline, walls varying in thickness, with simple septa, with branches. Tube
0.4–1.2 cm in length, brown (7D8). Stipe almost sessile, blunt, broadly attached, and thick
at base, irregularly ruptured crust overlying, and usually strongly laccate with brown dark
brown (9F7) from mature to old. Margin strongly laccate and dark brown (9F7) when
J. Fungi 2021, 7, x FOR PEER REVIEW 77 of 91
mature to old. Pore 4–5 in number per mm, subcircular to circular. Pore surface pale yellow
(4A3) to pale orange (5A3), turns dark brown (6F7) when scratched or bruised.

Figure32.
Figure 32. Morphology
Morphologyof ofGanoderma
Ganodermasubresinosum
subresinosum(MFLU
(MFLU17-1912):
17‐1912):(a)
(a)mature
maturebasidiomes;
basidiomes;(b)
(b)pore
characteristics; (c–e) generative hyphae of context in KOH; (f–h) basidiospores in KOH. Scale Scale
pore characteristics; (c–e) generative hyphae of context in KOH; (f–h) basidiospores in KOH. bars:
bars: (a) = 2 cm, (b) = 1000 μm, (c–e) = 20 μm, (f–h) = 5 μm.
(a) = 2 cm, (b) = 1000 µm, (c–e) = 20 µm, (f–h) = 5 µm.

Ganoderma
Hyphal thailandicum
structure: T. Luangharn,
Hyphal system P.E.with
di-trimitic, Mortimer,
clamp S.C. Karunarathna,
connections, and
usually J.C.
light
Xu, MycoKeys 59: 55 (2019)
yellow (3A5) to grayish-yellow (3B5), pale yellow (4A5), light orange (5A5) to orange (5B8)
Facesoffungi
in KOH; generativenumber: FoF 06129
hyphae 1.0–2.1 µm broad (n = 30), thin-walled, hyaline, with clamp con-
Index
nections; Fungorum
skeletal hyphaenumber:
2.8–5.2IFµm
556535
broad (n = 30), usually unbranched or few branches,
For characteristics, see Luangharn et al. [31].
Notes: Ganoderma thailandicum is characterized by its laccate, deep magenta close to
stipe, brownish‐red at center, and light yellow around active development toward the
margin on pileal surface, white pore surface, brownish‐red context, and absence of mela‐
J. Fungi 2021, 7, 819 69 of 83

and thick-walled; binding hyphae 2.5–5.0 µm width (n = 30), walls varying in thickness,
many branches, some hymenial with sword-like apices in the context. Pileipellis a hymeni-
derm, light orange (5A5), clavate-like cells, dextrinoid. Basidiospores mostly ellipsoid to
broadly ellipsoid, double walls, (11.6-)12.1–13.5–14.5(-15.8) × (8.1-)8.9–11.3–11.9(-12.5) µm
(x = 13.2 × 11.4 µm, n = 50) µm, with Q = 1.12–1.19, L = 13.24 µm, W = 11.44 µm (including
myxosporium), (10.2-)10.9–12.8–13.7(-14.6) × (6.8-)7.9–8.9–9.6(-10.2) µm (x = 12.6 × 8.8 µm,
n = 50) µm, with Q = 1.41–1.46, L = 12.63 µm, W = 8.84 µm (excluding outer myxosporium),
overlaid by hyaline, apically and echinulate, truncate, inner wall usually orange (6A7–6A8)
to reddish-golden (6C7) in KOH, brownish-orange (5C5–5C6) in Melzer’s reagent, outer
wall usually reddish-orange (7A8, 7B8) in 5% KOH and brown (6D7–6D8) in Melzer’s
reagent.
Habitat: Solitary on the decaying hardwood of Peltophorum pterocarpum and Castanop-
sis sp.
Specimens examined: THAILAND, Chiang Rai Province, 20◦ 150 0300 N, 100◦ 140 1700 E,
732 m elev., 9 October 2017, T. Luangharn, MFLU 17-1912.
Notes: Ganoderma subresinosum was introduced as Fomes subresinosus with the spec-
imen from the Philippines [43]. Next, Humphrey [159] verified this fungus species to
the genus Ganoderma, and then Imazeki [160] included this species in the genus Tra-
chyderma. Ganoderma subresinosum is a species that is distributed worldwide, known
from the Philippines to other Asian countries, and distributed across Eastern and Central
Africa [30,47,100,110]. This fungus is distinctive in form with its laccate pileus, sessile,
dark brown pileus surface, and concentrically sulcate with irregularly ruptured crust over-
lying the context, with ellipsoid to elongate basidiospores. Ganoderma subresinosum was
regarded as a synonym of Trachyderma tsunodae Imazeki [160], Magoderna subresinosus [110],
and Amauroderma subresinosum [113]. In China, this fungus was reported by Chinese re-
searchers as Fomes subresinosum [132,161,162] and verified to A. subresinosum by Zhao [61]
and Zhao and Zhang [13]. Recently, several studies have suggested that those three synony-
mous fungal species are different from G. subresinosum based on emergent morphological
and molecular data [163,164]. However, this fungus name is in the Index Fungorum as
G. subresinosum. So, in this study, we present the first record of G. subresinosum from
Thailand.
Ganoderma thailandicum T. Luangharn, P.E. Mortimer, S.C. Karunarathna, and J.C.
Xu, MycoKeys 59: 55 (2019)
Facesoffungi number: FoF 06129
Index Fungorum number: IF 556535
For characteristics, see Luangharn et al. [31].
Notes: Ganoderma thailandicum is characterized by its laccate, deep magenta close
to stipe, brownish-red at center, and light yellow around active development toward
the margin on pileal surface, white pore surface, brownish-red context, and absence of
melanoid band.
Ganoderma tropicum (Jungh.) Bres., Annales Mycologici 8(6): 586 (1910)
≡ Polyporus tropicus Jungh., Praemissa in floram cryptogamicam Javae insulae: 63
(1838)
≡ Fomes tropicus (Jungh.) Cooke, Grevillea. 14 (69): 19 (1885)
≡ Scindalma tropicum (Jungh.) Kuntze, Revisio generum plantarum. 3 (2): 519 (1898)
Facesoffungi number: FoF 05068
For characteristics, see Luangharn et al. [75].
Notes: Ganoderma tropicum was introduced as Polyporus tropicus by Junghuhn [165] with
the specimen from Indonesia. Phylogenetic analysis has been well resolved with the aid of
molecular data [5,129] and transferred this species to Ganoderma [166], where it is considered
a member of the G. lucidum species complex [76,167]. The fungus is characterized by its
distinctive reddish-brown pileal surface, dark brown context near the tubes, dense context
layer, thick near the base, with strongly echinulate basidiospores. Its distribution is highly
variable worldwide, scattered across tropical Asian regions, mainland China [4,5,30], South
J. Fungi 2021, 7, 819 70 of 83

America [137], and Taiwan, PRC [33], and it causes white root and butt rot on several
tree species [8]. Furthermore, there are some similarities between G. tropicum and other
Ganoderma species. According to Cao et al. [4], among the Chinese Ganoderma species,
G. flexipes, G. multipileum, G. sichuanense, and G. tsugae are similar to G. tropicum, having
a reddish-brown pileus surface, dark brown context, ellipsoid basidiospores, strongly
echinulate basidiospores, and irregular cuticle cells. Our new record of G. tropicum from
Northern Thailand was described based on characteristics together with phylogenetic
analyses, the details of which are shown in Luangharn et al. [75].
Taxonomy of Ganoderma from Vietnam
Ganoderma hochiminhense Karunarathna, Mortimer, & Luangharn, sp. nov. (Figure 33)
Facesoffungi number: FoF 06334
Index Fungorum number: IF 556794
Diagnosis: Ganoderma hochiminhense is characterized by its strongly laccate appear-
ance, with a reddish-brown color near the stipe, a deep orange to brownish-yellow at
the center, a white that indicates active development on the margin of the pileal surface,
a yellowish-white pore surface when fresh, an orange, deep orange to reddish-orange
context, and absence of melanoid bands.
Holotype: Vietnam, Hochiminh City, on Areca sp. (as described by the seller), 12 June
2019, LT2019 Gano 305 (MFLU 19-2224) and LT2019 Gano 306 (MFLU 19-2225).
Etymology: The specific epithet “hochiminhense” refers to the place in Vietnam from
where the holotype specimen was collected.
Description: Basidiomes sessile, orbicular. Pileus up to 0.8–2.2 cm in length, 0.5–1.4 cm
width, and 1.3 cm thick at base. Pileus orbicular, undulated, tuberculate, sulcate, single
or fused at the base, broadly attached, hard when dried. Pileus surface distinctively
laccate when young, strongly laccate when mature or dried. Pileus color reddish-brown
from the base, light to deep orange, golden yellow, brownish- to reddish-yellow at the
center, with deep yellow to orange yellow at the margin of the upper surface, generally
white where new hyphae are in active development, glossy, shiny, smooth, spathulate,
sulcate when fresh, with a thin crust overlying the pileus, which is thicker at the base
than at the margin, light weight when dried. Hymenophore mostly orange to brownish-
orange in KOH, up to 0.8 cm thick, with a dense but not fully homogenous context
layer, bearing distinct layers of concentric growth zones at the center that extend to the
margins, thick near the base, non-corky or woody texture when dried, bearing a simple
septum at base, tough to break when dried, melanoid bands absent. Context reddish-
orange in KOH; generative hyphae up to 2.72–3.82 µm (x = 3.54, n = 50) in diam, almost
colorless, thin-walled, some expanded at the apex, unbranched, with clamp connections;
skeletal hyphae-dominant, up to 3.56–7.47 µm (x = 5.85, n = 50), orange to brown, thick-
walled, unbranched, without clamp connections; binding hyphae 3.23–5.96 µm (x = 4.32,
n = 50), almost colorless, walls varying in thickness, with some narrow lumen to subsolid,
frequently branched, tortuous, and interwoven at the distal end. Tubes hard, brown, up to
0.2–0.5 cm long. Tubes layers generative hyphae 2.21–3.03 µm in diam, pale brown to
brown, thin-walled, some thick-walled, with clamp connections, unbranched; skeletal
hyphae 3.23–6.15 µm in diam, distinctly brown, thick-walled, some narrow lumen to
subsolid, frequently branched; binding hyphae 2.28–4.86 µm in diam, brownish-yellow,
thick-walled to solid, and frequently branched. Stipe short stipe, laccate when developing
to maturity, and strongly laccate from maturity to old age. Margin white when present to
maturity, light brown to brown between the young to mature stages, turning light brown
when dry, and silky, soft, and slippery to the touch between youth and maturity, usually
bruising when touched and tough to break. Pore angular to round, 4–6 in number per mm,
with dissepiments slightly thick to thick. Pore surface white when fresh, yellowish-white
when dried, turns brown to dark brown when touched.
J. Fungi 2021, 7, 819 71 of 83
J. Fungi 2021, 7, x FOR PEER REVIEW 80 of 91

Figure 33.
Figure 33. Characteristics
Characteristics ofofGanoderma
Ganoderma hochiminhense: (a) basidiomes
hochiminhense: of the of
(a) basidiomes strain
the MFLU 19‐2224;19-2224;
strain MFLU (b)
mature basidiomes of the strain MFLU 19‐2224; (c) mature basidiomes of the strain MFLU 19‐2225;
(b) mature basidiomes of the strain MFLU 19-2224; (c) mature basidiomes of the strain MFLU 19-2225;
(d,e) pores characteristics; (f) tube layers; (g) hyphae from tube layers; (h,i) hyphae from context in
(d,e)
KOH; pores characteristics;
(j) clamp connection(f) tube
from layers;(k)
context; (g)hyphae
hyphaefromfromtrama
tube in
layers;
KOH;(h,i) hyphae
(l) pores from context in
characteristics;
KOH;
(m–p) (j) clamp connection
basidiospores in KOH.from Scalecontext; (k) =hyphae
bars: (b,c) from= trama
2 cm, (d,e) in KOH;
1000 μm, (g–k) =(l)
20pores
μm, Lcharacteristics;
= 150 μm,
(m–p) basidiospores
(m–p) = 5 μm. in KOH. Scale bars: (b,c) = 2 cm, (d,e) = 1000 µm, (g–k) = 20 µm, L = 150 µm,
(m–p) = 5 µm.
J. Fungi 2021, 7, 819 72 of 83

Hyphal structure: Hyphal system trimitic, with tissues yellow, light orange to or-
ange and brown in KOH; generative hyphae up to 2.70–3.25 µm (x = 3.09, n = 50) in
diam, almost colorless, thin-walled, unbranched, with clamp connections, some slightly
swollen at the distal end; skeletal hyphae-dominant, up to 3.49–7.67 µm broad (x = 5.98,
n = 50), pale brown to distinctly brown, thick-walled, unbranched, without clamp con-
nections, occasionally with narrow lumen, some subsolid, interwoven; binding hyphae
3.12–5.87 µm broad (x = 4.28, n = 50), grayish-orange, walls varying in thickness, frequently
branched, tortuous, and interwoven in the distal end. Basidia clavate, 4-sterigmatic,
10.2–14.3 × 8.6–12.5 µm, yellowish to pale brown in KOH. Basidiospores ellipsoid, some-
times broadly ellipsoid or almond-shaped at maturity, reddish-yellowish, light brown to
brownish-orange, slightly truncate, double walls, exospore smooth, endospore with coarse
echinulate, (6.8)8.5–10.4(11.8) × (5.8)6.9–9.3(10.2) µm (x = 9.4 × 8.8 µm, n = 50), with Q
= 1.02–1.14, L = 9.38 µm, W = 8.82 µm (including myxosporium), (6.6)7.9–9.4(10.9) ×
(5.4)6.3–8.5(9.3) µm (x = 9.0 × 7.7 µm, n = 50) µm, with Q = 1.08–1.21, L = 8.98 µm, W = 7.73
µm (excluding outer myxosporium), strongly echinulate, cuticle cells irregular. Culture
characteristics white active mycelium on PDA, reaching 8 cm diam after 14 days at 25 ◦ C.
Habitat: solitary, on Areca spp. stumps.
Specimens examined: VIETNAM, Hochiminh City local market, 8◦ 540 3200 N, 98◦ 310 0900 E,
427 m elev., 12 June 2019, S.C. Karunarathna, LT2019 Gano 305 (MFLU 19-2224, holotype)
and LT2019 Gano 306 (MFLU 19-2225, paratype).

4. Discussion and Conclusions

The present study demonstrates the nomenclatural status, characteristics, phyloge-
netic analysis, host preference, growing season, climate, and substrate details of Ganoderma
species from the GMS. Detailed characteristics and molecular analyses allow us to define
the Ganoderma species used in our study as a distinctive well-supported clade, within Gano-
derma, with new insights to resolve species delimitation. Altogether, this study describes a
total of six new records and two new species of Ganoderma comprising two new records
(G. hoehnelianum and G. williamsianum) as well as a new species (G. myanmarense) from
Myanmar, four new records (G. ellipsoideum, G. multipileum, G. philippii, G. subresinosum)
from Thailand, and one new species (G. hochiminhense) from Vietnam.
Ganoderma orbiforme specimens were collected from living Albizia mollis and Indochinese
spp. in a deciduous forest from Thailand. These G. orbiforme specimens from the Thailand
group in the non-laccate Ganoderma clade with G. orbiforme from Laos [30] and China [37].
The Thai G. orbiforme shares similar features as mentioned by Hapuarachchi et al. [30] and
Wang et al. [156], such as possessing an annual to perennial, flabelliform or applanate pileus,
with a weakly laccate surface, the presence of several thick layers, an undulate margin,
and subcircular or circular pores. The pileus size of the Thai collection of G. orbiforme
(4–11 × 3–6 cm) is within the range of Asian collections (3.5–21 × 2.5–12 cm) [63] but
exceeds the Chinese collection (3–8 × 2.5–6 cm) [88]. These findings are consistent with
Ryvarden [136], who demonstrated that G. orbiforme are widely distributed in tropical and
temperate regions. The present study also reports an additional record of G. orbiforme from
a tropical region in Thailand.
Three G. sinense were collected from decaying as well as living Albizia mollis (Wall.)
Boiv. and Quercus spp. in temperate Yunnan Province, China, and two collections were
gathered from living Dendrocalamus strictus (Roxb.) Nees. and Dipterocarpus spp. in tropical
Thailand. The present collections of G. sinense grouped together as a sister taxon to
the G. nasalaense from Laos. Ganoderma sinense is regarded as having a high degree of
phenotypic plasticity and genetic diversity and was associated with substantial intraspecific
morphological variation [88,167]. When considering the morphological differences between
these, the Chinese G. sinense has a smaller-sized pileus (2–6 × 2–4 cm) and brownish-
orange context, while the Thai G. sinense has a larger-sized pileus (4–10 × 6–12 cm) with
a brown to dark brown context. However, the Chinese and Thai G. sinense also share
some characteristics, such as having stipitate, subdimidiate to dimidiate basidiocarps
J. Fungi 2021, 7, 819 73 of 83

with a laccate radial pileus that extends from the center to the margin, a colored dark
brown, cylindrical stipe, and basidiospores measuring 8.5–11.8 × 7.2–10.6 µm. The current
findings of G. sinense are in agreement with Hapuarachchi et al. [88], who demonstrated
morphological variations across G. sinense collected from Hainan and Guizhou Provinces,
China. In this study, G. sinense from Yunnan Province, China, and Thailand were also
recorded.
Ganoderma tropicum was introduced by Luangharn et al. [75] based on the collection
made from northern Thailand. In this study, phylogenetic analysis demonstrated that this
fungus served as the sister clade to G. multipileum, G. parvulum, and G. destructans, and the
Thai G. tropicum also grouped together with the Chinese G. tropicum, forming a sister clade
with G. multiplicatum and G. philippii.
The present study describes a new species, G. hochiminhense (MFLU 19-2224 and
MFLU 19-2225), from Vietnam. Phylogenetic analyses revealed G. hochiminhense collections
grouped as sister taxa to the laccate Ganoderma clade, and morphological characteristics
and molecular analyses provided insights to resolve species delimitation. Ganoderma
hochiminhense grouped together as a distinct clade with 100% ML, 98% MP, and 0.99 PP
support. Ganoderma hochiminhense forms a sister clade with G. zonatum from the U.S. [76],
the holotype G. ryvardenii from Cameroon [98], and the holotype G. angustisporum from
Africa [37]. However, G. hochiminhense, G. zonatum, and G. ryvardenii share similarities and
are sessile, laccate to strongly laccate on the upper pileus surface, and ellipsoid to broadly
ellipsoid with truncated apices basidiospores at maturity [76,98].
Although in the present phylogenetic tree G. hochiminhense clustered with G. zonatum
and G. ryvardenii, their macro- and micro-characteristics differ. Ganoderma hochiminhense
can be easily distinguished from G. zonatum by the size range of its small fruiting bodies
(0.8–2.2 × 0.5–1.4 cm diam), single or fused fruiting bodies on hosts, orbicular when
present, undulated pileus, reddish-brown at base, deep orange, golden yellow, reddish-
yellow at its center from maturity to old age, deep yellow and orange yellow on the
margin, white pore, reddish-orange context, (6.6–11.8 × 5.4–10.2 µm) basidiospore size
range, while the fruiting bodies of G. zonatum are brown on the upper pileus surface,
have cream pores, are finely echinulated, (9.4–8.8 × 9.0–7.7) basidiospore size range.
Ganoderma ryvardenii differs from G. hochiminhense by its large pileus size range (13 × 8.5 cm),
annual, dimidiate, or circular pileus, reddish margin, dark brown context, and basidiospore
size range (9.0–14.0 × 4–8 µm). Ganoderma zonatum was regarded as a species distributed
in subtropical to tropical climates [57,136,168], and our G. hochiminhense was also collected
from a tropical region. Consequently, we propose that taxa in this clade comprise species
isolated from tropical regions. This study confirmed that the new G. hochiminhense was
collected on Areca spp. stumps in tropical Vietnam. Considering overall morphology,
G. hochiminhense appears to be similar to G. boninense Pat. However, G. boninense is readily
distinguished from G. hochiminhense by its thicker and broader basidiocarp, reddish bay
pileus with reddish-brown pores, and differently sized basidiospores 10–12 × 7–8 µm in
G. boninense vs. 6.8–11.8 × 5.8–10.2 µm (including myxosporium), 6.6–10.9 × 5.4–9.3 µm
(excluding outer myxosporium) in G. hochiminhense [41].
For detailed characteristics and phylogenetic analyses of G. casuarinicola, G. thai-
landicum, and G. enigmaticum, see Xing et al. [37] and Luangharn et al. [31]. In the present
study, G. calidophilum collected from China formed a sister clade with the above three
Ganoderma species, and the clade is distinguished with suitable statistical support values.
Although G. calidophilum is sister to the laccate clade, its macro-characteristics are very
distinctive, such as a stipitate, subdimidiate to dimidiate pileus, homogenous brownish-red,
reddish-brown at the center, brownish-orange toward the stipe, light brown at the margin
from maturity to old age, brownish-orange context, and cylindrical stipe.
Ganoderma williamsianum was collected from tropical Myanmar, and the phyloge-
netic analysis grouped the present collection with G. williamsianum collected from Thai-
land [37]. Both G. williamsianum strains grouped as sister taxa to the holotype G. mbrekobe-
num from Ghana [90], with low support. The description of our collected G. williamsianum
J. Fungi 2021, 7, 819 74 of 83

is distinctive, having characteristics such as small fruiting bodies with a size range of
0.5–1.5 × 0.5–1 cm, rarely with a stipe-like base, subflabellate to circular fruiting bodies,
smooth and soft, weakly laccate, and homogenous orange to deep orange context, though
the basidiospore could not be observed. The current study allows for comparison between
G. williamsianum and G. mbrekobenum. G. mbrekobenum is macro-micro morphologically very
different from G. williamsianum, with its distinctive dimidiate, lateral stipe, dimitic hyphal
system, with ovoid to broadly ellipsoid basidiospore [90]. Xing et al. [37] also reported
the collection of G. williamsianum from Thailand with providing a phylogenic analysis of
their specimens but failing to provide any morphological descriptions. Ganoderma william-
sianum has been reported as a species distributed across China [154], Indonesia [110,155],
Malaysia [110], and the Philippines [110,169]. Here, this study introduces a new record of
G. williamsianum from Myanmar.
Ganoderma adspersum was collected near the root of the living tree Mangifera indica
L. in Laos, while the Thai G. adspersum strain was collected from the decaying hardwood
of Pterocarpus spp. Macro-characteristics of our collected G. adspersum share features
with what has already been described by Moncalvo et al. [45], such as having a non-
laccate, sessile, perennial, subdimidiate to dimidiate pileus, a sulcate, brown pileus surface
and context, and a yellowish-white pore surface, with a similar basidiospore size range
(6.9–10.6 × 4.7–7.8). However, some characteristics are also quite different, as the G. adsper-
sum from Laos is mostly ellipsoid to broadly ellipsoid, with brownish-orange inner walls
of basidiospores, while the Thai G. adspersum is mostly ellipsoid, with brownish-yellow
inner walls and brown to dark brown outer walls. It was observed that our Thai G. adsper-
sum had a di-trimitric hyphal system, while Hapuarachchi et al. [30] reported that their
Thai G. adspersum was composed of a trimitic hyphal system. This fungus is frequently
reported in temperate and tropical climates [45,152], where it grows on many hardwood
tree species [152,170]. Ganoderma adspersum has been reported in Europe [171], India [150],
Korea [92], Italy [172], Myanmar, and Thailand [30]. In this study, additional specimens of
G. adspersum from Laos and Thailand are presented for use in updated descriptions of this
species using detailed morphological observations and phylogenetic studies.
This study introduces a new record of G. ellipsoideum, collected from Chiang Mai
Province, Thailand, growing on rotten wood. The newly collected G. ellipsoideum serves as
the sister group to G. adspersum (ML = 97%, MP = 96%, PP = 0.95). It is also correlative to
the results by Hapuarachchi et al. [88]. Our collection from Thailand allows us to conduct
comparisons with the holotypes of G. ellipsoideum from Hainan Province, China. The current
G. ellipsoideum specimen shares similarities with the holotype collection in its annual, sessile,
several-layers-thick circular or subcircular pores, and presence of ellipsoid basidiospores.
However, the Chinese holotype of G. ellipsoideum presents quite a different basidiospore size
range (6.1–7.3 × 3.7–4.6 µm), including myxosporium, while the Thai G. ellipsoideum was
(4.8–7.7 × 3.1–5.4 µm) including myxosporium. The Chinese G. ellipsoideum was initially
yellowish-white, later having a yellowish-brown to brownish-yellow pore surface, while
Thai G. ellipsoideum presented white (11A1) to pale yellow (2A3) coloration. G. ellipsoideum
also serves as the sister group to the G. gibbosum and G. austral clade. Our G. gibbosum
collections were collected from China, Laos, and Thailand. G. gibbosum macro- and micro-
characteristics are quite different. For variations in G. gibbosum, see Luangharn et al. [173],
while G. australe from Thailand has been described in Luangharn et al. [5].
Ganoderma multipileum was collected from Thailand after the rainy season. Phylo-
genetic analysis showed our G. multipileum specimen matched closely within the clade
to G. multipileum (CWN 04670) from Taiwan. Our collection was grouped as a sister to
G. steyaertianum B. J. Smith and Sivasith, G. mizoramense Zothanzama, Blanchette, Held,
C.W., Barnes, G. martinicense Welti and Courtec, and G. destructans M.P.A. Coetzee, Mar-
inc. and M.J. Wingf. These Ganoderma species, including G. steyaertianum, G. mizoramense,
G. martinicense, G. destructans, and G. multipileum, share similarities in that they have a
laccate to strongly laccate upper pileus surface at maturity. Although G. steyaertianum can
be easily distinguished from G. multipileum by its distinctive kidney-shaped basiodomata
J. Fungi 2021, 7, 819 75 of 83

with a dark brown pileus surface, this species is also similar to G. multipileum in that its
basiodomata rarely extend completely to the margin [128]. Ganoderma mizoramense can be
differentiated from G. multipileum by its reddish-brown when fresh to liver-brown when
dry upper pileus surface, dark-brownish to dark-reddish brown context, and lower surface
white when fresh [94]. Ganoderma martinicense can be differentiated from G. multipileum
by its generally sessile basidiomata and dark-cinnamon-brown context [35], while our
G. multipileum specimen rarely displayed sessile basidiomata and had a light brown to
brown context; however, phylogenetic analysis revealed G. martinicense as a close relative
to the Asian taxon G. multipileum [35,93]. G. destructans is significantly different, with its
pileus containing a creamy and soft context and presenting mostly ovoid basidiospores [9].
Ganoderma philippii was collected in tropical Chiang Mai Province, Thailand. Phy-
logenetic analysis reveals it to be close to Malaysian G. philippii strains [97]. Ganoderma
philippii is a unique species that grows especially actively in heavy soils with high water
content [111]. Macro- and micro-characteristics were also correlative to results by Singh
et al. [26], who reported this fungus as easily identifiable by its non-laccate pileus sur-
face and sessile basidiome mostly presenting oblong basidiospores. Ganoderma philippii
formed a sister clade to G. multiplicatum and G. tropicum. G. philippii are very different,
with their distinctive non-laccate, often convex, umbonate, and plump pileus and sessile,
with mostly oblong basidiospores, while G. multiplicatum is laccate to strongly laccate,
radial or branched from the center, extending to the margin, and presenting mostly as
ellipsoid basidiospores [33]. G. tropicum was collected from Thailand; although it closed
a sister clade with G. multiplicatum, its macro- and micro-characteristics are noticeably
different and were detailed by Luangharn et al. [31].
In this study, our G. sichuanense specimens were collected in the temperate climate
zone of Yunnan Province, China, and a tropical part of Thailand. Phylogenetic analysis
placed our collections as sister taxa to the laccate Ganoderma clade and close to the pre-
viously reported G. sichuanense from China [99]. Our collections also share macro- and
micro-characteristics similar to the holotype described of G. lingzhi [4], with their shell-
like, reniform to circular pileus, reddish-brown pileal surface, strongly laccate, stipitate,
and mostly present ellipsoid basidiospores that truncate at maturity; however, G. lingzhi
was verified to G. sichunense in the records of Index Fungorum. Among the Chinese Gano-
derma species, G. flexipes, G. multipileum, G. sichuanense, G. tropicum, and G. tsugae are the
species that most closely resemble G. sichuanense, as their macro- and micro-characteristics
share a reddish-brown pileal surface, similar basidiospores, and cuticle cells. Nevertheless,
differences between these Ganoderma species have been detailed in Dai et al. [34] and
Yao et al. [174].
Ganoderma resinaceum has long been reported in temperate climates [38,76], and our col-
lection is also from a temperate part of Yunnan Province, China. Our Chinese G. resinaceum
is close to the G. resinaceum collected in England and the Netherlands, with suitable clade
support. Our result is also similar to the phylogenetic analysis by Douanla-Meli and
Langer [46]. However, G. resinaceum from England and the Netherlands lacks macro-micro
descriptions. Moreover, G. resinaceum collections have also been reported from India [143],
although the macro- and micro-characteristics of these Indian specimens differ from our
Yunnan collections. The Yunnan strain of G. resinaceum is distinctive with its short stipe,
varying reddish-brown upper pileus surface, grayish-yellow to dark brown context covered
by an irregularly ruptured thin crust, and larger pileus size range (1.5–12.5 × 1–7 cm),
while the Indian strain is stipitate, with a brown upper pileus surface, round margin, light
brown context, and smaller pileus size range (8–9 × 6–7.5 cm) [143]. Therefore, we con-
clude that despite phylogenetic similarities placing the various specimens, from temperate
and tropical regions, of G. resinaceum in the same clade, there remains a high degree of
morphological variance within this species [38,46,76,143].
We introduce G. hoehnelianum as a new record from Myanmar. This mushroom
was collected on the decaying stump of an unidentified dicotyledonous tree species in
tropical Myanmar. The phylogeny of our G. hoehnelianum specimen is close to the Chinese
J. Fungi 2021, 7, 819 76 of 83

G. hoehnelianum collection [91] and is well supported. Our collection shares similar macro-
and micro-characteristics to Wang and Wu [154] and Hapuarachchi et al. [88], with its
distinctive annual, sessile, plano concave, dark brown upper pileus surface and context,
and concentrically sulcate zones with broadly ovoid basidiospores. Our G. hoehnelianum
grouped with the G. austroafricanum H. Xing, B.K. Cui, and Y.C. Dai and G. carocalcareum
Douanla-Meli clade. G. austroafricanum is distinctive from G. hoehnelianum, with its reddish-
brown pileus surface, smooth surface, round margin, with subglobose basidiospores.
Ganoderma carocalcareum collected from a tropical region in Cameroon is distinctive from
G. hoehnelianum, with its friable context, and thick toward the margin and downwards to
the pore margin (part of hymenophore), red-brown to brown-orange margin, and forming
a concentrical aporoid zone [46].
Ganoderma applanatum was collected from temperate China and tropical Thailand.
Phylogenetic analysis shows our G. applanatum clustering as a distinctive group with
suitable support. The name G. lipsiense has been treated and corrected to G. applanatum [15],
and it is mentioned that G. applanatum also belongs to this complex species. Macro-
and micro-characteristics of our collections are similar to those described in detail in
Ryvarden and Gilbertson [105]. Our Chinese and Thai G. applanatum collections have quite
different macro-characteristics. The Chinese G. applanatum shows a smaller pileus size range
(1.5–5.8 × 0.5–4.5 cm) and is up to 1.5 cm thick at the base, while the Thai G. applanatum
was 2–12 × 2.5–6 cm and 0.8–3.1 cm thick at the base. The Chinese G. applanatum is
distinctive and non-laccate, faded from maturity to old age, while Thai G. applanatum had
differentiated zones of peach red and reddish-orange at the base, brownish-orange with
the radius, light brown at the center, and extending to dark brown close to active mycelium
(margin). Our results are also correlative to Wang et al. [5], who has reported on the high
morphological variation within Chinese Ganoderma. Ganoderma species from different
geographic areas have also shown separate lineages in phylogenetic analyses [38,153].
In this study, we describe a new Ganoderma species, G. myanmarense, collected in
tropical Myanmar. The two G. myanmarense collections grouped as sister taxa to the laccate
Ganoderma clade, and their characteristics and molecular analyses provided insights to
resolve species delimitation. This new species, G. myanmarense, forms a sister clade with
G. wiiroense E.C. Otto, Blanchette, C.W. Barnes and Held. from Ghana, G. destructans M.P.A.
Coetzee, Marinc, M.J. Wingf. from South Africa, and our G. flexipes from China. Ganoderma
wiiroense has an annual, dimidiate pileus; sessile, yellowish-brown to dark-reddish-brown
basidiocarps [101], while our G. myanmarense has its distinctive sectorial to shell-shaped,
sub-reniform to reniform pileus; golden yellow, yellowish-red at center, slightly reddish-
orange and reddish-brown (8E8) basidiocarps. Ganoderma destructans has its distinctive
globular pileus, with creamy soft non-poroid tissues [9], and G. flexipes is very different
from G. myanmarense by having sub-reniform to reniform pileus, small pileus size range
(0.5–3.2 × 0.5–3 cm), homogenous reddish-brown to dark brown upper pileus surface,
and sub-cylindrical to the cylindrical stipe.
Our G. lucidum collections came from both China and Thailand. Phylogenetic anal-
ysis showed our strains clustered with G. lucidum from France, with suitable statistical
support. The results indicated that our G. lucidum grouped as sister taxa to the laccate
G. leucocontextum from China. G. lucidum has a high degree of morphological variability,
often resulting in taxonomic and phylogenetic confusion [38]. However, our Chinese and
Thai G. lucidum specimens displayed a degree of morphological variability. The Chinese
G. lucidum possessed a cylindrical stipe, and when viewed from above, one can observe
the presence of a central stipe at the center of the pileus, while the Thai G. lucidum pre-
sented a cylindrical and eccentric stipe. However, both strains also shared key features,
with their distinctive stipitate, furrowed, incised, undulate to sulcate, undefined imbricate,
yellowish-red, brownish-red to reddish-orange with laccate upper pileus surface, light
brown to dark brown context, and presence of dark brown melanoid bands. Our results are
also in agreement with those of Wang et al. [5], who reported that Chinese G. lucidum has a
deeper-colored context that is even darker near the tube layer. Although both G. lucidum
J. Fungi 2021, 7, 819 77 of 83

and G. leucocontextum form a sister clade, their morphological characteristics are different.
Ganoderma leucocontextum shows its distinctive white context, thickset stipe, and broadly
ellipsoid basidiospores (9.5–12.5 × 7–9 µm). For descriptive details, please see Li et al. [62].
Ganoderma tsugae is one of the Ganoderma species that share similar characteristics with
G. lucidum [4]. In this study, we collected G. tsugae from Yunnan Province, China. In the phy-
logenetic analysis, it clustered with G. tsugae from the USA [76] and grouped as a sister to
G. oregonense [76,175]. However, G. tsugae has smaller basidiospores (9–11 × 6–8 µm) [176]
than G. oregonense (13–17 × 8–10 µm) [109].
In this study, we introduce a new record of G. subresinosum collected in Thailand
during the rainy season. Phylogenetic analysis indicated our G. subresinosum to be a
distinct clade with suitable support. Macro- and micro-characteristics of G. subresinosum
are similar to Hapuarachchi et al. [30], who demonstrated that this fungal species has
a distinctively annual, sessile, laccate, dark brown upper pileus surface, concentrically
sulcate, irregularly grayish-orange when dried and woody when old, presenting ellipsoid
to elongate basidiospores.
Based on comprehensive characteristics and molecular analyses, we report 23 Gan-
oderma species from GMS, including G. adspersum, G. applanatum, G. australe, G. cali-
dophilum, 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, G. tropicum, and G. tsugae, of which 13 Ganoderma species were
collected from Yunnan Province, China, 3 species from Laos, three species from Myanmar,
and 12 species from Thailand, including G. hochiminhense from Vietnam and G. myan-
marense from Myanmar. We noted a high degree of intraspecies morphological variability
between specimens collected from different parts of this region, confirming why it is hard
to correctly identify Ganoderma to the species level using only morphological observations
and why phylogenetic analyses are crucial in maintaining correct taxonomic placement
within this group.

Supplementary Materials: The following are available online at https://www.mdpi.com/article/

10.3390/jof7100819/s1, Figure S1: Collection sites of specimens used in this study in China, Laos,
Myan-mar, Thailand, and Vietnam. Black triangles indicate collection locations for the specimens;
Table S1: The extant Ganoderma species in the GMS.
Author Contributions: Conceptualization, T.L., S.C.K., P.E.M., I.P. and J.X.; methodology, T.L., S.C.K.,
A.K.D. and P.E.M.; formal analysis, T.L., S.P. and S.C.K.; investigation, T.L. and S.C.K.; data curation,
T.L., S.C.K., A.K.D., T.L. and S.P.; writing—original draft preparation, T.L.; writing—review and
editing, J.X., K.D.H., S.C.K. and P.E.M.; visualization, P.E.M.; supervision, J.X. and P.E.M.; S.C.K.;
project administration, J.X. and P.E.M.; funding acquisition, J.X. and P.E.M. All authors have read and
agreed to the published version of the manuscript.
Funding: This work was supported by the National Science Foundation of China (NSFC) (grant
number Y71L481211), the CAS President’s International Fellowship Initiative (PIFI) young staff under
(grant number 2020FYC0002), the National Science Foundation of China (NSFC) (grant number
31851110759), and the Thailand Science Research and Innovation (TSRI) grant “Macro fungi diversity
research from the Lancang-Mekong Watershed and surrounding areas” (grant number DBG6280009).
Acknowledgments: We appreciate the kind support given by the University of Chinese Academy of
Sciences, Beijing 100049, China; Centre for Mountain Futures (CMF), Kunming Institute of Botany,
Kunming 650201, Yunnan, China and Center of Excellence in Fungal Research, Mae Fah Luang
University, Chiang Rai, Thailand. This research work was partially supported by Chiang Mai
University. Cin Khan Lian at Ar Yone Oo social development association, Yangon, Myanmar,
is thanked for his help on field work. We thank the Germplasm Bank of Wild Species, Kunming
Institute of Botany, Kunming 650201, Yunnan, China, for enabling our molecular laboratory work.
Kasiphat Limsakul, Wilawan Punyaboon, Thilina Sudarshana Nimalrathna and Deping Wei are
acknowledged for their invaluable assistance. We acknowledge the Kunming Institute of Botany,
the Chinese Academy of Sciences, for providing the laboratories and instruments for molecular work.
Shaun Pennycook is thanked for help in naming the new fungal species.
J. Fungi 2021, 7, 819 78 of 83

Conflicts of Interest: The authors declare no conflict of interest.

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