High diversity of Ganoderma and Amauroderma (Ganodermataceae, Polyporales) in Hainan Island, China

Abstract

Species of Amauroderma and Ganoderma (Ganodermataceae) have been widely used as traditional medicines in Asia over many centuries. The genera are widely researched, owing to their beneficial medicinal properties and chemical constituents with potential nutritional and therapeutic uses. There are, however, taxonomic confusions surrounding the species in these genera, whose circumscription is often unclear. We surveyed species of Amauroderma and Ganoderma in Hainan Island in Southern China. In this paper, we provide data on 15 species found in the Wuzhishan and Jiangfengling mountains on the island. One species named Ganoderma ellipsoideum Hapuar., T.C. Wen & K.D. Hyde, sp. nov. is new to science and introduced with evidence from morphology and molecular analysis. The other species collected are described with photographs and compared with similar taxa. We provide a phylogeny for the two genera based on ITS sequence data and the taxonomic status of the species is briefly discussed.

Full text

Submitted 6 August 2018, Accepted 15 September 2018, Published 18 September 2018
Corresponding Author: Ting-Chi Wen – e-mail – tingchiwen@yahoo.com 931
High diversity of Ganoderma and Amauroderma (Ganodermataceae,
Polyporales) in Hainan Island, China
Hapuarachchi KK1,2,3, Karunarathna SC4, Raspé O5,6, De Silva KHWL7,8,
Thawthong A2,3, Wu XL9, Kakumyan P3, Hyde KD2,3,4 and Wen TC1*
1The Engineering Research Center of Southwest Bio–Pharmaceutical Resource Ministry of Education, Guizhou
University, Guiyang 550025, Guizhou Province, China
2Center of Excellence in Fungal Research, Mae Fah Luang University, Chiang Rai 57100, Thailand
3School of Science, Mae Fah Luang University, Chiang Rai 57100, Thailand
4Key Laboratory for Plant Diversity and Biogeography of East Asia, Kunming Institute of Botany, Chinese Academy of
Sciences, 132 Lanhei Road, Kunming 650201, China
5Botanic Garden Meise, Nieuwelaan 38, 1860 Meise, Belgium
6Fédération Wallonie-Bruxelles, Service général de l’Enseignement universitaire et de la Recherche scientifique, Rue
A. Lavallée 1, 1080 Bruxelles, Belgium
7Research Group of Aquatic Plant Ecology, Donghu Experimental Station of Lake Ecosystem, Institute of
Hydrobiology, Chinese Academy of Sciences, Wuhan, 430072, China
8University of Chinese Academy of Sciences, Beijing 100049, China
9Guizhou Academy of Sciences, Guiyang, 550009, Guizhou Province, China
Hapuarachchi KK, Karunarathna SC, Raspé O, De Silva KHWL, Thawthong A, Wu XL,
Kakumyan P, Hyde KD, Wen TC 2018 – High diversity of Ganoderma and Amauroderma
(Ganodermataceae, Polyporales) in Hainan Island, China. Mycosphere 9(5), 931–982, Doi
10.5943/mycosphere/9/5/1
Abstract
Species of Amauroderma and Ganoderma (Ganodermataceae) have been widely used as
traditional medicines in Asia over many centuries. The genera are widely researched, owing to their
beneficial medicinal properties and chemical constituents with potential nutritional and therapeutic
uses. There are, however, taxonomic confusions surrounding the species in these genera, whose
circumscription is often unclear. We surveyed species of Amauroderma and Ganoderma in Hainan
Island in Southern China. In this paper, we provide data on 15 species found in the Wuzhishan and
Jiangfengling mountains on the island. One species named Ganoderma ellipsoideum Hapuar., T.C.
Wen & K.D. Hyde, sp. nov. is new to science and introduced with evidence from morphology and
molecular analysis. The other species collected are described with photographs and compared with
similar taxa. We provide a phylogeny for the two genera based on ITS sequence data and the
taxonomic status of the species is briefly discussed.
Key words – 1 new species – ITS – Lingzhi – Morphology – Phylogeny
Introduction
Ganodermataceae (Basidiomycota) is a large, cosmopolitan family of polypores with five
accepted genera: Amauroderma Murril, Ganoderma P. Karst, Haddowia Steyaert, Humphreya
Steyaert and Polyporopsis Audet (Richter et al. 2015) including 592 epithets, of which about three
fourths are classified under the genus Ganoderma (www.indexfungorum.org, accessed 21 August
Mycosphere 9(5): 931–982 (2018) www.mycosphere.org ISSN 2077 7019
Article
Doi 10.5943/mycosphere/9/5/1

932
2018). Recently, Foraminispora Robledo, Costa-Rezende & Drechsler-Santos was introduced to
accommodate Porothelium rugosum (= Amauroderma sprucei) and Furtadoa Costa-Rezende,
Robledo & Drechsler-Santos was introduced to accommodate, Amauroderma brasiliensis (Singer)
Ryvarden and A. corneri Gulaid & Ryvarden, as well as a new species Furtadoa biseptata Costa-
Rezende, Drechsler-Santos & Reck (Costa-Rezende et al. 2017). Furthermore, some researchers
support the claim of legitimacy of Magoderna Steyaert to accommodate Ganoderma subresinosum
(Murrill) C.J. Humphrey with strong morphological and molecular evidence (Gomes-Silva et al.
2015, Costa-Rezende et al. 2016, 2017). Ganodermataceae is distinct from other families of
polypores in having peculiar double-walled basidiospores (with interwall pillars for Ganoderma)
and the inner wall of the Ganodermataceae spores is quite often coloured, and usually the surface is
ornamented (Donk 1964, Adaskaveg & Gilbertson 1988)). Furthermore, the basidiospores also bear
an apical, often shrunk umbo, and the apex appears as truncate (Smith & Sivasithamparam 2000).
Seo & Kirk (2000) described basidia of members of Ganodermataceae as relatively large and
ranging from typically clavate to pyriform. The hyphal system is usually trimitic, but occasionally
dimitic or monomitic, and is comprised of hyaline, thin-walled, branched, septate or aseptate and
clamped generative hyphae, as well as pigmented, thick-walled, arboriform or aciculiform and
branched skeletal hyphae, and usually colourless, terminal branched binding hyphae, if present.
Ganoderma and Amauroderma are the two most important genera in Ganodermataceae.
Ganoderma was established by Karsten (1881) with G. lucidum (Curtis) P. Karst as the type
species. This fungus is distributed all over the world in green ecosystems both in temperate and
tropical regions and grow as facultative parasites that can live as saprobes on rotting stumps and
roots (Turner 1981). Ganoderma species are not classified as edible mushrooms, as the basidiomes
are always corky and tough and do not have the fleshy texture characteristic of true edible
mushrooms (Hapuarachchi et al. 2015). Basidiomes grow on living, or more commonly, dead
trunks or branches of trees. One of two types of basidiomes is produced, depending on the species:
laccate basidiomes with a shiny upper surface, or non-laccate basidiomes with a dull upper surface
(Smith & Sivasithamparam 2003, Pilotti et al. 2004). The members of family Ganodermataceae
have been widely used as traditional medicines over many centuries in Asia (Zhou et al. 2015).
Species in Ganodermataceae are widely researched, because they contain many chemical
constituents with potential nutritional and therapeutic values and, therefore, have highly prized
medicinal value (De Silva et al. 2012a, b, 2013, Hapuarachchi et al. 2016a, b, 2017). Ganoderma
species are economically important not only because of their medicinal properties, but also because
of their phytopathogenicity (Dai et al. 2007, 2009). Ganoderma species cause white rot of hard
woods by decomposing lignin, cellulose and related polysaccharides (Hepting 1971, Adaskaveg et
al. 1991). The root and stem rots caused by Ganoderma species, result in forestry yield loss
worldwide, in important trees such as Elaeis guineensis (oil palm; Glen et al. 2009) and Hevea
brasiliensis (rubber; Monkai et al. 2016). Yu & Shen (2003) reported that Ganoderma (Lingzhi)
was successfully cultivated for the first time in 1969 in China. Ganoderma products, from different
parts of the basidiomes, mycelia or spores, are sold in the form of powder, coffee, tea or other
drinks, dietary supplements, syrups, tooth pastes, soaps and lotions (Chang & Buswell 1999, Lai et
al. 2004, Singh et al. 2013).
After the genus Ganoderma was established, Patouillard (1887) transferred other species to
Ganoderma, and Karsten (1889) introduced the genus Elfvingia for non-laccate Ganoderma
species. Furthermore, Patouillard (1889) introduced section Amauroderma for species with
spherical and subspherical basidiospores with uniformly thickened walls. Ganoderma P. Karst.
comprises sect. Amauroderma, Ganoderma, subgenera, Ganoderma and Trachyderma to date
(www.indexfungorum.org, accessed 21 August 2018). The genus Ganoderma was first divided into
two sub-genera, Ganoderma and Elfvingia by Karsten (1889). Various authors have used different
taxonomic characters for the identification of the species (Murrill 1902, 1903, Atkinson 1908,
Coleman 1927, Corner 1947). The pileipellis has been differently characterized by different
authors: Furtado (1965) referred to a derm of the palisadoderm type in Elfvingia and layer of cells
in a hymeniderm in Ganoderma, while Steyaert (1980) referred to a trichoderm in Elfvingia and

933
“hymenioderm” in subg. Ganoderma. Clemençon (2004) used “crustohymeniderm” for subg.
Ganoderma. However, Steyaert (1972, 1980) worked extensively on the genus and he introduced
many new species, transferred many names to the genus and removed several synonyms. Ryvarden
(1995), Gottlieb & Wright (1999a, b) studied the macro and micromorphology of this genus
further. Later, isoenzyme analysis techniques have been used to separate the species of Ganoderma
(Park et al. 1994, Gottlieb et al. 1995, 1998, Gottlieb & Wright 1999a, Smith & Sivasithamparam
2000). Furthermore, ribosomal DNA sequencing tools have been used to analyse the phylogenetic
relationship in Ganoderma lucidum complex (Moncalvo et al. 1995a, b, c). Hseu et al. (1996) used
RAPD–Polymerase chain reaction (PCR) and internal transcribed spacer (ITS) sequences to
differentiate the isolates of G. lucidum complex. Although phylogenetic analyses support the
monophyly of Ganoderma and its classification into two subgenera, there are many problems in the
resolution below the sub generic level (Moncalvo 2000, Hong & Jung 2004).
Despite their huge economical importance, the taxonomy of Ganoderma remains unclear,
because it has been plagued by confusions and misconceptions. Ganoderma is called “Ling Zhi”,
Chi–zhi” or “Rui–zhi” in China; “Reishi”, “Munnertake” or “Sachitake” in Japan and “Youngzhi”
in Korea (Wagner et al. 2003). Traditional Chinese books classified Ganoderma into six species
with reference to the colour of the basidiomes: Sekishi (red), Shishi (violaceous), Kokushi (black),
Oushi (yellow), Hakushi (white) and Seishi (blue), and were assigned to different species based on
different triterpenoid patterns (Szedlay 2002). Ganoderma was first reported from China by Teng
(1934), with four species including G. lucidum and one variety. Later, many researchers
extensively studied Chinese Ganodermataceae and introduced more than 100 species in four
genera: Ganoderma, Amauroderma, Haddowia and Humphreya (Zhao & Zhang 2000, Wu & Dai
2005, Cao et al. 2012). Ganoderma lucidum was accepted as the scientific binomial of “Lingzhi” in
many reports on Chinese edible and medicinal mushrooms, but it was recently described as G.
lingzhi based on strong molecular and morphological evidence (Cao et al. 2012, Dai et al. 2017).
Mycochemical and other studies reported different species numbers in the genus Ganoderma (Peng
et al. 2014, Yan et al. 2013). However, the majority of Ganoderma species reported from China
have not been subjected to systematic studies (Baby et al. 2015). There are 445 epithets listed in
Index Fungorum (accessed 21 August 2018) for Ganoderma, while Kirk et al. (2008) estimated
there were 80 species worldwide although part of them are synonyms.
Ganoderma species identification has been difficult and recently debated by many
researchers (Moncalvo et al. 1995a, Wang et al. 2009, Cao et al. 2012, Yao et al. 2013, Richter et
al. 2015, Zhou et al. 2015). A number of Ganoderma collections and species have been misnamed
because of the presence of heterogenic forms, taxonomic obstacles and inconsistencies in the way
the genus has been subdivided (Mueller et al. 2007). Ganoderma species have been reported to be
genetically heterogeneous because of geographical differentiation, and high genetic diversity is
maintained by out-crossing between generations (Miller et al. 1999, Pilotti et al. 2003). This high
genetic diversity is accompanied by substantial morphological variation, even within species (Hong
et al. 2001). Hence, naming a species within this genus often is problematic and traditional
taxonomic methods based only on morphology are insufficient for establishing a stable
classification system for Ganoderma (Hseu et al. 1996, Hong et al. 2002, Thawthong et al. 2017),
which has resulted in an uncertain nomenclature. This confusing situation is mainly the result of
various criteria used in identification by different authors. Some authors strictly focus on host–
specificity, geographical distribution and macro morphology of basidiomes, while other authors
only focus on spore characteristics as the primary taxonomic characteristics (Sun et al. 2006,
Ekandjo & Chimwamurombe 2012). Recently, some researchers have suggested using a
combination of morphological, chemotaxonomic and molecular methods to develop a more stable
taxonomy for this genus (Richter et al. 2015, Welti et al. 2015).
Amauroderma Murrill (1905) was introduced and typified by Amauroderma regulicolor
(Berk. ex Cooke) Murrill and this is a widespread tropical genus with 135 epithets listed in Index
Fungorum (accessed 21 August 2018). Species of Amauroderma usually occur on roots of living or
dead trees or, more rarely, are wood inhabiting (Furtado 1981, Ryvarden 2004) and cause white rot.

934
Members of this genus vary from stipitate to sessile with non-truncate, globose to subglobose,
verrucose to asperulate (smooth in only one species) basidiospores (Ryvarden 2004). Taxonomic
studies of Amauroderma in China have been carried out over many years by various researchers
(Teng 1936, Zhao et al. 1979, Zhao & Zhang 1987, Li & Yuan 2015, Song et al. 2016). Twenty
species have been recorded in China (Zhao & Zhang 2000), but among these, only six have been
confirmed as Amauroderma based on both morphological characters and phylogenetic analyses: A.
austrosinense J.D. Zhao & L.W. Hsu, A. concentricum Song, Xiao L. He & B.K. Cui, A. perplexum
Corner, A. rugosum (Blume & T. Nees) Torrend, A. subresinosum (Murrill) Corner and A.
yunnanense J.D. Zhao & X.Q. Zhang (Li & Yuan 2015). The other Amauroderma species recorded
from China, have not been fully studied yet.
Hainan Province is an island in southern China located between 18° 10´–20° 10´ N and 108°
37´–111° 05´ E (Fig. 1). It has both lowland areas in the northeast and mountainous terrain in the
central and southwestern regions. The average annual temperature and precipitations are 19.7 °C
and 2650 mm, respectively. This island has subtropical to tropical vegetations (Dai et al. 2011).
Several studies have been carried out over the past three decades to investigate the Ganoderma
diversity of Hainan Island (Wu et al. 1999, Li et al. 2010, Dai et al. 2011, Wang et al. 2012).
The objective of the present study is to document the members of Ganoderma and
Amauroderma found in Wuzhishan and Jiangfengling Mountains on Hainan Island with
descriptions, colour photographs, illustrations and a phylogeny. All the species of Amauroderma
and Ganoderma are compared with similar taxa. A checklist of species of Amauroderma and
Ganoderma reported from Hainan Island is also given in Table 1.
Check-list of the species of Amauroderma and Ganoderma known for Hainan Island
The authors of the scientific names in the following list are given according to Index
Fungorum, 2018 (http://www.indexfungorum.org/names/Names.asp) (Table 1).
Materials & Methods
Samples of Ganoderma were collected in August 2014 from Wuzhishan and Jiangfengling
Mountains, Hainan Province, China and dealt with as in Cao et al. (2012). The materials were
deposited at Guizhou University (GACP) and Mae Fah Luang University (MFLU) herbaria.
Morphological characteristics examination
Macro-morphological characters were described based on fresh material, and on the
photographs provided here. Colour codes (e.g. 3A3) are from Kornerup & Wanscher (1978).
Specimens were dried and placed separately in plastic bags. For micro-morphological observations,
basidiomes were examined under a stereo dissecting microscope (Motic SMZ 168 series) and
sections were cut with a razor blade, mounted in 5% KOH, and then observed, measured and
illustrated under a compound microscope (Nikon ECLIPSE 80i) equipped with a camera (Canon
600D). Measurements were made using Tarosoft (R) Image Frame Work v. 0.9.7. At least 20
basidiospores were measured from each mature specimen except for very scanty materials. The
basidiospore size was measured both with and without the myxosporium, but only spore sizes with
myxosporium were used for comparisons. Basidiospore dimensions are given as (a–) b–c–d (–e),
where a represents the minimum, b (mean average-standard deviation), c the average, d (mean
average+standard deviation) and e the maximum. Q, the length/width ratio (L/W) of a spore in side
view and Qm is the average, smallest and largest Q values given as Q. Pellis sections were taken
from the mature pileus portion and mounted in Melzer’s reagent for observation. The Facesoffungi
number is provided as explained in Jayasiri et al. (2015).
DNA extraction, PCR, and sequencing
Samples taken from dried basidiome were used to extract genomic DNA. Genomic DNA was
extracted using an EZgene TM Fungal gDNA Kit (Biomiga, CA, USA) according to the
manufacturer instructions. DNA concentrations were estimated visually on agarose gel by

935
comparing band intensity with 1 Kb DNA ladder (Invitrogen Biotech). The nuclear ribosomal
internal transcribed spacers region (ITS) was amplified using primers ITS5 and ITS4 (White et al.
1990). Reaction mixtures (20 µl) contained 1 µl template DNA (ca. 10 ng), 10 µl distilled water,
and 1 µl (10 µM) of each primer and 7 µl 2× BenchTopTM Taq Master Mix (Biomigas).
Amplification conditions were 2 min at 94 °C followed by 35 cycles of 95 C for 30 s, 59 °C for 30
s and 72 °C for 1 min, followed by a final extension at 72 °C for 10 min. Amplified PCR products
were verified by 1% agarose gel electrophoresis stained with ethidium bromide in 1x TBE. The
PCR products were sequenced by Invitrogen Biotechnology (Beijing).
Figure 1 – Location of Wuzshishan and Jiangfengling Mountains and other areas of collection in
Hainan province, China (This map was prepared using the freely available online map data (Global
Administrative Areas http://www.gadm.org/download, accessed March 6, 2017).

936
Table 1 List of Amauroderma and Ganoderma species known for Hainan Island
Taxa
Status
(verified/unverified)
Current
name
Reference
Amauroderma rugosum
(Blume & T. Nees) Torrend
(1920)
verified
Wu et al. 1999,
Li et al. 2010
Li & Yuan 2015,
This study
A. austrosinense J.D. Zhao &
L.W. Hsu (1984)
verified
Wu et al. 1999,
Li & Yuan 2015,
This study
Ganoderma ahmdii Steyaert
(1972)
unverified
Wu et al. 1999
G. amboinense (Lam.) Pat.
(1887)
unverified
Wu et al. 1999
G. annulare (Jungh.) Gilb.
(1962)
verified
G. australe
(Fr.) Pat.
(1889)
Wu et al. 1999,
Dai et al. 2011,
This study
G. applanatum (Pers.) Pat.
(1887)
verified
Wu et al. 1999,
This study
G. atrum J.D. Zhao, L.W. Hsu
& X.Q. Zhang (1979)
unverified
G. flexipes
Pat. (1907)
Wu et al. 1999,
Wang & Wu 2007
This study
G. australe (Fr.) Pat. (1889)
verified
Wu et al. 1999,
Li et al., 2010,
Dai et al. 2011,
This study
G. bawanglingense J.D.
Zhao & X.Q. Zhang (1987)
Unverified
Li et al. 2010
G. boninense Pat. (1889)
unverified
G.
orbiforme
(Fr.)
Ryvarden
2000
Wu et al. 1999,
Dai et al. 2011
G. brownii (Murrill) Gilb.
(1962)
unverified
Wu et al. 1999
G. calidophilum J.D. Zhao,
L.W. Hsu & X.Q. Zhang
(1979)
verified
Li et al. 2010,
Dai et al. 2011,
This study
G. capense (Lloyd) Teng
(1963)
unverified
Wu et al. 1999
G. chalceum var. chalceum
(Cooke) Steyaert (1967)
unverified
Wu et al. 1999
G. cochlear (Blume & T.
Nees) Merr. (1917)
unverified
Wu et al. 1999
G. cupreum (Cooke) Bres.
(1911)
verified
G.
orbiforme
Wu et al. 1999,
Wang et al. 2014,
This study
G. curtisii (Berk.) Murr.
(1908)
unverified
Wu et al. 1999
G. dahlii (Henn.) Aoshima
(1971)
unverified
Wu et al. 1999
G. densizonatum J.D. Zhao
& X.Q. Zhang (1986)
Verified
G.
orbiforme
Wu et al. 1999,
Wang et al. 2014

937
Table 1 Continued.
Taxa
Status
(verified/unverified)
Current
name
Reference
G. flexipes Pat. (1907)
verified
Wu et al. 1999,
Zhou et al. 2015,
This study
G. fornicatum (Fr.) Pat. (1889)
verified
G. orbiforme
Wu et al. 1999,
Wang et al. 2014,
This study
G. gibbosum ((Blume & T.
Nees) Pat. (1897)
verified
Dai et al. 2011,
This study
G. hainanense J.D. Zhao, L.W.
Hsu & X.Q. Zhang (1979)
unverified
G. flexipes
Wu et al. 1999,
Wang & Wu 2007,
This study
G. hoehnelianum Bres. (1912)
verified
Wu et al. 1999,
Wang & Wu 2010,
This study
G. jiangfenglingense X.L Wu.
(1996)
unverified
Wu et al. 1999
G. 1eytense Steyaert (1972)
unverified
Wu et al. 1999
G. limushanense J.D. Zhao &
X.Q. Zhang (1986)
verified
G. orbiforme
Wu et al. 1999,
Wang et al. 2014,
This study
G. lobatum (Schwein.) G.F.
Atk. (1908)
unverified
Wu et al. 1999
G. 1ucidum (W Curt. Fr.)
Karst. (1881)
unverified
Wu et al. 1999
G. luteomarginatum J.D. Zhao,
L.W. Hsu & X.Q. Zhang
(1979)
unverified
Wu et al. 1999
G. mastoporum (Lev.) Pat.
(1889)
verified
G. orbiforme
Wu et al. 1999,
Wang et al. 2014,
This study
G. multipileum Ding Hou.
(1950)
verified
Dai et al. 2011,
Wang et al. 2005,
Zhou et al. 2015
G. multiplicatum (Moat.)Pat.
(1889)
verified
Dai et al. 2004,
This study
G. nigrolucidum (Lloyd) D.A.
Reid (1975)
unverified
Dai et al. 2004,
Li et al. 2010
G. parviungulatum .D. Zhao &
X.Q. Zhang (1986)
unverified
G. flexipes
Wu et al. 1999,
Cao et al. 2012
This study
G. ramosissimum J.D. Zhao
(1989)
unverified
Wu et al. 1999
G. resinaceum Boud (1890)
verified
Wu et al. 1999
This study
G. shangsiense J.D. Zhao.
(1988)
Verified
G.
hoehnelianu
m
Li et al. 2010,
Wang & Wu 2010,
This study
G. sinense J.D. Zhao. L.W.
Hsu & X.Q. Zhang (1979)
verified
Dai et al. 2004,
Wang et al. 2005,
This study
G. subresinosum (Murrill) C.J.
Humphrey (1938)
verified
Wu et al. 1999,
Li et al. 2010
This study

938
Table 1 Continued.
Taxa
Status
(verified/unverified)
Current
name
Reference
G. tornatum (Pers.) Bres.
(1912)
verified
G. australe
Li et al. 2010,
This study
G. tropicum (Jungh.) Bres.
(1910)
verified
Li et al. 2010,
Zhou et al. 2015,
This study
G. valesiacum Boud. (1895)
unverified
Wu et al. 1999
G. weberianum (Bres. & Henn.
ex Sacc.) Steyaert (1972)
verified
Wu et al. 1999,
Wang et al. 2012
Sequence alignment and phylogenetic analysis
The taxon information and GenBank accession numbers used in the molecular analyses are
listed in Table 2. Quality of the newly obtained sequences from the Ganoderma specimens from
Hainan Province, China was checked by observing the chromatogram with BioEdit (Hall 1999) and
by examining BLAST search results according to Nilsson et al. (2012). The BLAST search was
also used to retrieve sequences from the closest matching taxa in Ganodermataceae. Seventy
nucleotide sequences representing 35 species of Ganodermataceae from Asia, America and Europe
were retrieved from GenBank. Those sequences and 19 newly generated sequences were aligned
using MAFFT v. 7.309 (Katoh & Standley 2013) online at
http://mafft.cbrc.jp/alignment/server/index.html, and the alignment was improved manually where
necessary using Bioedit. Maximum Likelihood (ML) analysis was performed using RAxML-HPC2
(Stamatakis 2014) on the CIPRES Science Gateway V. 3.3 (Miller & Blair 2009), with default
settings except the number of bootstrap replicates was set to 1,000. For Bayesian (BY) analysis,
GTR+I+G model of evolution was selected with MrModeltest 2.2 (Nylander 2004) as the best-fit
model. BY analyses were conducted with 2 runs of six simultaneous Markov chains and trees were
sampled every 100th generation. The analyses were stopped after 5,000,000 generations when the
average standard deviation of split frequencies was below 0.01. The convergence of the runs was
checked using TRACER v1.6 (Rambaut et al. 2013). The first 25% of the resulting trees were
discarded as burn-in, and PP were calculated from the remaining sampled trees. In both ML and
BY analyses, Coriolopsis trogii was selected as the outgroup. ML bootstrap values and Bayesian
posterior probabilities greater than or equal to 70% and 0.95, respectively, were considered as
significant support. The phylogenetic tree was visualized with FigTree version 1.4.0 (Rambaut
2012) available at http://tree.bio.ed.ac.uk/software/figtree/.
Table 2 ITS sequences used in the phylogenetic analysis
Species
Voucher/
strain
Origin
GenBank
accession
number
Reference
Amauroderma
rugosum (Blume &
T. Nees) Torrend
GACP1408
0910
Hainan, China
MH106869
This study
A. rugosum
GACP1408
0952
Hainan, China
MH106870
This study
A. rugosum
Zhou 523
Guangxi, China
KJ531674
Li & Yuan 2015
A. rugosum
Dai9553
Hainan, China
KJ531668
Li & Yuan 2015
A. rugosum
Cui8882
Guangdong, China
KJ531667
Li & Yuan 2015
Ganoderma sp.
JM95/5
Thailand
AF255122
Moncalvo &
Buchanan 2008

939
Table 2 Continued.
Species
Voucher/
strain
Origin
GenBank
accession
number
Reference
Ganoderma sp.
JM98/233
Yunnan, China
AF255116
Moncalvo &
Buchanan 2008
G. adspersum
(Schulzer) Donk
ITA 39
Unknown
EF060011
Terho et al. 2007
G. adspersum
PF263
Italy
JN176908
GenBank
G. applanatum
(Pers.) Pat.
K(M)12082
9
UK
AY884179
GenBank
G. applanatum
Dai8924
China
KU219987
Song et al. 2016
G. australe (Fr.) Pat.
GACP1408
1671
Hainan, China
MH106871
This study
G. australe
GACP1408
1134
Hainan, China
MH106872
This study
G. australe
GDGM258
31
China
JX195200
GenBank
G. cupreum
HMAS130
804
Australia
JX840345
GenBank
G. curtisii
CBS
100131
NC, USA
JQ781848
Zhou et al. 2015
G. curtisii
CBS
100132
NC, USA
KJ143967
Zhou et al. 2015
G. destructans
M.P.A. Coetzee,
Marinc. & M.J.
Wingf.
CMW4367
0
South Africa
KR183856
Coetzee et al.
2015
G. destructans
CMW4367
1
South Africa
KR183857
Coetzee et al.
2015
G. ellipsoideum
Hapuar., T.C. Wen &
K.D. Hyde, sp. nov.
(holotype)
GACP1408
0966
Hainan, China
MH106867
This study
G. ellipsoideum
GACP1408
0968
Hainan, China
MH106868
This study
G. ellipsoideum
GACP1408
1215
Hainan, China
MH106886
This study
G. flexipes Pat.
GACP1404
5450
Hainan, China
MH106873
This study
G. flexipes
Wei5200
(IFP)
Hainan, China
JN383978
Cao & Yuan
2013
G. flexipes
Wei5494
(IFP)
Hainan, China
JN383979
Cao & Yuan
2013
G. fornicatum (Fr.)
Pat.
BCRC3537
4
Taiwan
JX840349
Wang et al. 2014
G. fornicatum
TNM-
F0009926
China
JX840348
Wang et al. 2014
G. gibbosum (Cooke)
Pat.
XSD-34
Unknown
EU273513
GenBank
G. gibbosum
GACP1407
0442
Hainan, China
MH106880
This study
G. gibbosum
SFC201509
18-08
Korea
KY364271
Jargalmaa et al.
2017

940
Table 2 Continued.
Species
Voucher/
strain
Origin
GenBank
accession
number
Reference
G. hoehnelianum
Bres.
Dai12096
China
KU219989
Song et al. 2016
G. hoehnelianum
Dai11995
China
KU219988
Song et al. 2016
G. hoehnelianum
GACP1408
0913
Hainan, China
MH106881
This study
G. leucocontextum
T.H. Li, W.Q. Deng,
Dong M. Wang &
H.P. Hu
GDGM443
03
China
KJ027607
Li et al. 2015
G. leucocontextum
TL-2013
China
KF011548
Li et al. 2015
G. lipsiense sensu
auct.
FIN
131R610
Unknown
EF060004
Terho et al. 2007
G. lipsiense
NOR74/67/
5
Unknown
EF060002
Terho et al. 2007
G. lobatum (Cooke)
G.F. Atk.
JV 0402/24
Unknown
KF605677
GenBank
G. lobatum
JV
1212/10J
Unknown
KF605676
GenBank
G. lucidum (Curtis)
P. Karst.
Dai11593
(IFP)
Finland
JQ781852
Cao et al. 2012
G. lucidum
K175217
UK
KJ143911
Zhou et al. 2015
G. lucidum
MT2610
(BRNM)
Czech Republic
KJ143912
Zhou et al. 2015
G. lucidum
Dai2272
(IFP)
Sweden
JQ781851
Cao et al. 2012
G. lucidum
HKAS7108
8
Yunnan, China
KC222321
Yang & Feng
2013
G. lucidum
GICN04
Italy
AM906058
Guglielmo et al.
2008
G. lucidum
GIT099
Italy
AM269773
Guglielmo et al.
2008
G. lucidum
Rivoire
4195
France
KJ143909
Zhou et al. 2015
G. mastoporum Lév.)
Pat.
TNM-
F0018835
China
JX840351
Wang et al. 2012
G. mastoporum
TNM-
F0018838
China
JX840350
Wang et al. 2012
G. mastoporum
Gma-1
Unknown
GU213486
GenBank
G. mizoramense
Zothanz., Blanchette,
Held & C.W. Barnes
UMN-MZ4
India
KY643750
Crous et al. 2017
G. mizoramense
UMN-MZ5
India
KY643751
Crous et al. 2017
G. multipileum Ding
Hou
CWN0467
0
Taiwan, China
KJ143913
Wang et al. 2012
G. multipileum
Dai9447
Hainan, China
KJ143914
Wang et al. 2012

941
Table 2 Continued.
Species
Voucher/
strain
Origin
GenBank
accession
number
Reference
G. multiplicatum
(Mont.) Pat.
GACP1408
1328
Hainan, China
MH106879
This study
G. multiplicatum
Dai 13710
China
KU572489
Xing et al. 2016
G. orbiforme (Fr.)
Ryvarden
GACP1408
0918
Hainan, China
MH106878
This study
G. orbiforme
GACP1408
1239
Hainan, China
MH106876
This study
G. orbiforme
GACP1408
1108
Hainan, China
MH106874
This study
G. orbiforme
GACP1408
1329
Hainan, China
MH106877
This study
G. orbiforme
GACP1408
1235
Hainan, China
MH106875
This study
G. oregonense
Murrill 1908
CBS
265.88
OR, USA
JQ781875
Zhou et al. 2015
G. oregonense
CBS
266.88
OR, USA
JQ781876
Zhou et al. 2015
G. parvulum Murrill
URM83343
Brazil
JQ618246
Correia de Lima
et al. 2014
G. parvulum
URM80765
Brazil
JX310822
Correia de Lima
et al. 2014
G. pfeifferi Bres.
K(M)12081
8
UK
AY884185
Park et al.
2012
G. pfeifferi
874 (CAS–
IM)
Czech Republic
AM906059
Guglielmo et al.
2008
G. philippii (Bres. &
Henn. ex Sacc.) Bres.
E7098
Indonesia, Sumatra
islands
AJ536662
GenBank
G. philippii
E7425
Malaysia, Selangor
AJ608713
GenBank
G. resinaceum Boud.
DP107
Italy
AM906064
Guglielmo et al.
2008
G. resinaceum
GR102
India
GU451247
Mohanty et al.
2011
G. resinaceum
GACP
HNU02
Hainan, China
MH106883
This study
G. ryvardenii
HKAS5805
3
Cameroon
HM138671
Kinge & Mih
2011
G. sinense J.D. Zhao,
L.W. Hsu & X.Q.
Zhang
GACP1408
1236
Hainan, China
MH106882
This study
G. sinense
GS175
Unknown
DQ425014
Su et al. 2007
G. sinense
GS92
Unknown
DQ424982
Su et al. 2007
G. steyaertanum B.J.
Sm. & Sivasith.
II-121-1
Indonesia
KJ654427
Glen et al. 2014
G. steyaertanum
6-WN-15-
M-A
Indonesia
KJ654459
Glen et al. 2014

942
Table 2 Continued.
Species
Voucher/
strain
Origin
GenBank
accession
number
Reference
G. subresinosum
(Murrill) C.J.
Humphrey
GACP1408
1663
Hainan, China
MH106885
This study
G. subresinosum
3C-29
Indonesia
KJ654406
Glen et al. 2014
G. subresinosum
5-D-3-D-26
Indonesia
KJ654467
Glen et al. 2014
G. tropicum (Jungh.)
Bres.
BCRC3712
2 (TNM)
Taiwan, China
EU021457
Wang et al. 2009
G. tropicum
Dai9724
China
JQ781879
Cao et al. 2012
G. tropicum
GACP1408
1518
Hainan, China
MH106884
This study
G. tsugae Murrill
Dai3937
(IFP)
China
JQ781853
Cao et al. 2012
G. tsugae
12751b
(BJFC)
USA(CT)
KJ143919
Zhou et al. 2015
G. tsugae
AFTOL-
ID771
Unknown
DQ206985
Matheny et al.
2007
G. tsugae
Dai12760
USA
KJ143920
Zhou et al. 2015
G. wiiroense E.C.
Otto, Blanchette,
C.W. Barnes & Held
UMN-21-
GHA
Ghana
KT952363
Crous et al. 2015
G. wiiroense
UMN-20-
GHA
Ghana
KT952361
Crous et al. 2015
Coriolopsis trogii
(Berk.) Domański
TC-02
(TNM)
USA
JN164993
Jargalmaa et al.
2017
* BJFC, Beijing Forestry University; BRNM, Moravian Museum in Brno; CBS, Centraalbureau voor
Schimmelcultures, Utrecht, the Netherlands; CAS–IM, Academy of Sciences of Czech Republic Institute of
Microbiology Department of Experimental Mycology, Videnska, Czech Republic; CGMC China General
Microbiological Culture Collection Center, Chinese Academy of Sciences; Di. Va.P.R.A, Department of
Exploitation and Protection of the Agricultural and Forestry resources, University of Torino, Grugliasco
(Italy); FFPRI ,the Forest Products Research Institute; FWP, Fungi of West Pakistan, Jardin Botanique de
Belgique, Belgium; GACP, Herbarium of Guizhou Agricultural College (Guizhou University), Guiyang
City, Guizhou, China; HMAS, Mycological Herbarium of the Institute of Microbiology, Chinese Academy
of Sciences, HKAS; Kunming Institute of Botany, Chinese Academy of hidden flowers herbarium; IFP,
Institute of Applied Ecology, Chinese Academy of Sciences; JV, The private herbarium of Josef Vlasak;
KEW, the Royal Botanical Garden, Kew, Surrey, UK; KTCC, Korean Type Culture Collection; NY, the
New York Botanical Garden; Rivoire, the private collection of Bernard Rivoire; RYV, Herbarium of Leif
Ryvarden, Oslo, Norway; TNM, the Herbarium of the National Museum of Natural Science.
* Type specimens are in bold.
Results
Phylogeny
The tree topologies obtained from ML and BY were identical. Therefore, only the ML tree is
shown (Fig. 2). The 89 collections of Amauroderma and Ganoderma clustered in ten clades

943
(Amauroderma rugosum, G. applanatum, G. hoehnelianum, G. lucidum species complex, G.
orbiforme, G. resinaceum, G. sinense, G. subresinosum, laccate Ganoderma and non-laccate
Ganoderma) (Fig. 2). Our collections from Hainan Province, China clustered with all other
Amauroderma and Ganoderma species, including the holotypes (G. destructans, G. wiiroense, G.
ryvardenii, and G. mizoramense) and the paratype (G. wiiroense) in well-supported clades (Fig. 2).
Furthermore, the new species G. ellipsoideum clustered in the non-laccate Ganoderma species
clade with high support (BS = 91%, BPP = 1.0).
Sequences, A. rugosum (GACP14080910 and GACP14080952), G. australe
(GACP14081134 and GACP14081671), G. flexipes (GACP HNU5450), G. gibbosum
(GACP14070442), G. hoehnelianum (GACP14080913), G. multiplicatum (GACP14081328), G.
orbiforme (GACP14080918, GACP14081108, GACP14081235, GACP14081239 and
GACP14081329), G. resinaceum (GACP HNU02), G. sinense (GACP14081236), G. subresinosum
(GACP14081663) and G. tropicum (GACP14081518) obtained from collections, clustered in well-
supported clades forming monophyletic groups with, A. rugosum (BS = 99%, BPP = 1.0), G.
australe (BS = 100%, BPP = 1.0), G. flexipes (BS = 97%, BPP = 0.98), G. gibbosum (BS = 73%,
BPP = 0.97), G. hoehnelianum (BS = 100%, BPP = 1.0), G. multiplicatum (BS = 100%, BPP = 1),
G. orbiforme (BS = 99%, BPP = 0.99), G. resinaceum (BS = 95%, BPP = 1), G. sinense (BS =
100%, BPP = 0.99), G. subresinosum (BS = 100%, BPP = 1.0) and G. tropicum (BS = 100%,
BPP=1.0), respectively (Fig. 2).
Ganoderma lucidum sequences from Europe (Dai2272, GIT099, Dai11593, MT2610,
BR4195 and K175217) and China (HKAS71088), G. tsugae sequences from China (Dai3937) and
USA (12751b and Dai12760), G. leucocontextum sequences from China (GDGM44303 and TL-
2013) and G. oregonense sequences from USA (CBS265.88 and CBS266.88) clustered in the G.
lucidum species complex clade (BS = 96%, BPP = 1) (Fig. 2). Ganoderma lipsiense sequences
(NOR74/67/5, FIN131R610) (= G. applanatum) with two other G. applanatum sequences from
China (Dai8924) and Europe (K(M) 120829) clustered in a well-supported G. applanatum clade,
forming a monophyletic group (BS = 99%, BPP = 1) (Fig. 2). The Ganoderma resinaceum (GACP
HNU02) sequence obtained from collections from Hainan Province, together with G. resinaceum
sequences from India (GR102) and Italy (DP107) attained from GenBank, clustered in the well-
supported G. resinaceum clade (BS = 95%, BPP = 1) (Fig. 2). Therefore, we can conclude that
Asian and European G. resinaceum species are phylogenetically similar even they have different
morphological characteristics. Stalpers (1978) believed G. resinaceum is the correct name for the
fungus described as G. lucidum and few researchers considered the numerous names as synonyms
of G. resinaceum (Steyaert 1980, Ryvarden 1985). Moreover, it was believed that based on
molecular data, G. resinaceum is a species complex (Moncalvo et al. 1995a). Later, it was found
that G. resinaceum could not be distinguished phylogenetically from G. lucidum (Hong & Jung
2004). However, Mohanty et al. (2011) has mentioned Ganoderma resinaceum to be distinct from
G. lucidum and later, more researchers have suggested that G. resinaceum differed from G. lucidum
(Cao et al. 2012, Zhou et al. 2015). Ganoderma hoehnelianum (GACP14080913) sequence
obtained from our study, together with G. hoehnelianum sequences from China (Dai12096 and
Dai11995) retrieved from GenBank, clustered in the well-supported G. hoehnelianum clade (BS =
100%, BPP = 1.0). Furthermore, Ganoderma hoehnelianum groups in a sister clade to G.
resinaceum (Fig. 2).
Ganoderma curtisii (BS = 100%, BPP = 1.0), G. destructans (BS = 99%, BPP = 0.95), G.
flexipes (BS = 97%, BPP = 0.98), G. mizoramense (BS = 100%, BPP = 1.0), G. multipileum (BS =
74%, BPP = 0.95), G. multiplicatum (BS = 100%, BPP = 1.0), G. parvulum (BS = 100%, BPP =
1.0), G. philippii (BS = 100%, BPP = 1.0), G. steyaertianum (BS = 98%, BPP = 1.0), G. tropicum
(BS = 100%, BPP = 1.0) and G. wiiroense (BS = 100%, BPP = 1.0) clustered together in laccate
Ganoderma species clade (Fig. 2). Ganoderma tropicum sequence (GACP14081518) from
collections from Hainan Province, together with G. tropicum sequences (BCRC37122 and
Dai9724) from China obtained from GenBank, clustered in the well-supported G. tropicum clade
forming a monophyletic group (BS = 100%, BPP = 1.0) (Fig. 2). Previous phylogenetic evidence

944
indicates that G. tropicum is a distinct species (Cao et al. 2012, Wang et al. 2012, Yang & Feng
2013, Zhou et al. 2015) and this species can be widely found across subtropical and tropical Asia
(Moncalvo & Ryvarden 1997). Some researchers have shown in their phylogenies that G. tropicum
is phylogenically more similar to G. multipileum (Wang et al. 2012, Cao et al. 2012). In our studies,
G. tropicum resembles G. multipileum in morphology and habitat even though they are distinct
species as Wang et al. (2009) mentioned previously. Ganoderma flexipes sequence
(GACP14045450) obtained from collections from Hainan Province, together with G. flexipes
sequences from China (Wei5200 and Wei5494) attained from GenBank, clustered in the well-
supported G. flexipes clade (BS = 100%, BPP = 1.0) (Fig. 2). Ganoderma flexipes clustered with G.
wiiroense species originated from Ghana and further, they group in a sister clade to G. philippii
from Malaysia. With the aid of molecular evidence many researchers reported G. flexipes species in
China are more similar to G. multipileum and G. tropicum species found in China (Cao et al. 2012,
Yang & Feng 2013, Zhou et al. 2015). Our study here confirmed the same opinion. Ganoderma
multiplicatum sequence (GACP14081328) obtained from collections from Hainan Province,
together with G. multiplicatum sequences (Dai13710) from China retrieved from GenBank,
clustered in the well-supported G. multiplicatum clade forming a monophyletic group (BS = 100%,
BPP = 1.0) (Fig. 2).
Ganoderma adspersum (BS = 100%, BPP = 1.0)), G. australe (100%, BPP = 1.0), G.
ellipsoideum (BS = 91%, BPP = 1.0), G. gibbosum (BS = 73%, BPP = 0.97), G. lobatum (BS =
99%, BPP = 1.0) and G. pfeifferi (BS = 100%, BPP = 1.0) are clustered together in non-laccate
Ganoderma species clade (Fig. 2). Ganoderma australe sequences (GACP14081671 and
GACP14081134) obtained from our collections from Hainan Province, together with G. australe
sequence (GDGM25831) retrieved from GenBank, clustered in the well-supported G. australe
clade (100%, BPP = 1.0) (Fig. 2). Ganoderma australe forms a support clade (BS = 70%, BPP =
0.95) with Ganoderma adspersum, G. ellipsoideum, G. gibbosum and G. lobatum. Ganoderma
ellipsoideum is a new member of non laccate Ganoderma species and clustered in a separate clade
from G. gibbosum with strong molecular support (BS = 91%, BPP = 1) even they share some
similar morphologies (Fig. 2). Ganoderma gibbosum sequence (GACP14070442) obtained from
our study, together with G. gibbosum sequences (XSD-34 and SFC20150918-08) acquired from
GenBank, clustered in the well-supported G. gibbosum clade forming a monophyletic group (BS =
73%, BPP = 0.97) (Fig. 2).
Ganoderma orbiforme (GACP14080918, GACP14081108 GACP14081235, GACP14081239
and GACP14081329) sequences acquired from collections from Hainan Province, together with G.
mastoporum (TNM F0018835, TNM F0018838 and Gma 1) and G. fornicatum sequences (TNM
F0009926, BCRC35374) retrieved from GenBank clustered in the well-supported G. orbiforme
clade (BS = 99%, BPP = 0.99) (Fig. 2). Wang et al. (2012) proposed G. orbiforme is the earliest
valid name to use for the species G. mastoporum and G. fornicatum as previously described.
Ganoderma sinense sequence (GACP14081236) obtained from our collections from Hainan
Province, together with G. sinense sequences (GS175 and GS92) retrieved from GenBank,
clustered in the well-supported G. sinense clade forming a monophyletic group (BS = 100%, BPP =
1.0) (Fig. 2). Furthermore, this species form a support clade with G. orbiforme and, however
recognized as a species distinct from G. orbiforme.
Ganoderma subresinosum sequence (GACP14081663) acquired from our collections from
Hainan Province, together with G. subresinosum sequences (3C-29 and 5-D-3-D-26) from
Indonesia retrieved from GenBank, clustered in the well-supported G. subresinosum clade forming
a monophyletic group (BS = 100%, BPP = 0.99) (Fig. 2). Furthermore, G. subresinosum formed a
distinct lineage from Ganoderma which supports the claim of Costa-Rezende et al. (2017)
suggested, as Magoderna might be accepted at generic level. In the present phylogeny,
Amauroderma formed a distinct lineage from Ganoderma. Amauroderma rugosum sequences
(GACP14080910 and GACP14080952) obtained from our collections are clustered with A.
rugosum sequences (Cui8882, Dai9553 and Zhou523) which retrieved from the GenBank in a well-
supported clade forming a monophyletic group (BS = 99%. BPP = 1) (Fig. 2).

945
Figure 2 – Phylogram generated by maximum likelihood analysis of ITS sequences data. Bootstrap
support values for maximum likelihood, greater than 70% and Posterior Probabilities from
Bayesian Inference ≥ 0.95 are given above branches. The tree was rooted with Coriolopsis trogii.
The strain numbers and the countries of origin are mentioned after the species. Type species are
indicated in black bold.
Taxonomy
Key to the Genera
1. Basidiospores conspicuously truncated at maturity, ellipsoid; basidiomes sessile to stipitate: if
stipe present, never with a black zone between the stipe context and pileus context ......... Ganoderma

946
1. Basidiospores round; basidiomes often stipitate; with a black zone between the stipe context and
pileus context ................................................................................................................. Amauroderma
Key to the Species
1. Basidiomes laccate ……………………………………………………………………………….2
1. Basidiomes non laccate …………………………………………………………………………..9
2. Basidiomes distinctly stipitate ……………………………………………………………………3
2. Basidiomes sessile/or with a short stipe ………………………………………………………….5
3. Bovista type binding hyphae: present ………………………………………………......G. sinense
3. Bovista type binding hyphae: absent ……………………………………………………………..4
4. Pileus: suborbicular ………………………………………………………………G. calidophilum
4. Pileus: flabelliform ……………………………………………………………………..G. flexipes
5. Differentiated swollen zone at the point of attachment: present ………………………………….6
5. Differentiated swollen zone at the point of attachment: absent ……………………G. resinaceum
6. Pileus: suborbicular ……………………………………………………………………………….7
6. Pileus: spathulate ……………………………………………………………………...G. tropicum
7. Distinct concentric zones in upper surface of the pileus: present ………………………………...8
7. Distinct concentric zones in upper surface of the pileus: absent ………………..G. subresinosum
8. Pileus colour: dark brown ………………………………………………………G. hoehnelianum
8. Pileus colour: orange …………………………………………………………….G. multiplicatum
9. Spores shape: ellipsoid …………………………………………………………………………..10
9. Spores shape: elongate ………………………………………………………………G. orbiforme
10. Pileus: suborbicular …………………………………………………………………………….11
10. Pileus: spathulate ………………………………………………………………………………12
11. Spores length > 8 µm …………………………………………………………………G. australe
11. Spores length <8 µm ……………………………………………………………..G. applanatum
12. Margin in pileus: wavy ……………………………………………………………..G. gibbosum
12. Margin in Pileus: soft …………………………………………………………....G. ellipsoideum
Genus Ganoderma P. Karst., 1881, Rev. Mycol. (Toulouse) 3, p. 17.
= 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)
= Magoderna Steyaert (1972)
= Tomophagus Murrill, Torreya 5: 197 (1905)
= Trachyderma (Imazeki) Imazeki, Bull. Gov. Forest Exp. Stn Tokyo 57: 97 (1952)
= Whitfordia Murrill (1908)
Description (from Ryvarden 2004).
Basidiomes annual or perennial, stipitate to sessile; pileus surface with a thick, dull cuticle or
shiny and laccate with a thin cuticle or cuticle of clavate end cells; context cream coloured to dark
purplish brown, soft and spongy to firm-fibrous; pore surface cream coloured, bruising brown, the
pores regular, 4–7 per mm; tube layers single or stratified, pale to purplish brown; stipe when
present central or lateral; hyphal system dimitic; generative hyphae with clamps; skeletal hyphae
hyaline to brown, non-septate, often with long, tapering branches; basidia broadly ellipsoid,
tapering abruptly at the base; cystidia absent; basidiospores broadly to narrowly ellipsoid with a
truncate apex and apical germ pore, wall two-layered, endosporium brown and separated from the
hyaline exosporium by inter-wall pillars, negative in Melzer's reagent, 7–30 μm long.
Type species: Ganoderma lucidum (Leyss: Fr.) Karst.

947
Annotated list of Ganoderma species in the Hainan Island
Ganoderma cf. applanatum
Basidiome annual, sessile (usually with a distinctly contracted base), non-laccate, woody.
Pileus 10–15 × 4–7 cm, sub-dimidiate, sub-applanate; upper surface hard, several layers thick,
brown (6E4) to greyish brown (6E3), concentrically sulcate zones with turberculate bumps and
ridges and rivulose depressions, radially rugose, with irregularly ruptured crust overlying the pellis;
margin soft, 2–3 mm thick, rounded, concolourous with the rest of the pileus; lower surface greyish
yellow (4B3) to light brown (6D5). Hymenophore up to 15 mm long, indistinctly stratose; pores
initially whitish, light brown (6E4), 4–5 per mm; tubes circular or sub-circular. Context up to 3 cm
thick, dry, duplex; lower layer greyish orange (6B3), fibrous/pithy, composed of coarse loose
fibrils; upper layer greyish orange (6B3), woody. Basidiospores (n=50) (5.9–)6.3–7.4–8.1(–8.8) ×
(2.2–)4.2–5.8–6.3(–7) μm (Qm=1.8, Q=1.1–3.4, with myxosporium). (3.1–)4.0–5.7–6.4(–7.0) ×
(1.8–)2.5–2.8–3.5(–4.4) μm (Qm =2.1, Q=1.4–2.9, without myxosporium), ellipsoid, sometimes
truncate at one end, greyish orange (5B5) to light orange (5A5), eusporium bearing fine, short and
distinct echinulae, overlaid by a hyaline myxosporium. Pileipellis a hymeniderm, brown (6E4) to
greyish brown (6E3), composed of apically acanthus like branched cells, dextrinoid. Context
trimitic; generative hyphae (n=30) (1.7–)2.5–2.6–3.5(–3.6) μm in width, thin-walled, colorless,
with clamp connections; skeletal hyphae (n=30) (3.3–)3.7–4.3–5.9(–6.4) μm in width, brown (6E4)
to greyish brown (6E3), dextrinoid, thick-walled; binding hyphae (n=30) (2.9–)3.5–3.7–4.4(–5.4)
μm in width, brown (6E4) to greyish brown (6E3), thick-walled, frequently branched at apex,
septate, intertwined with the skeletal hyphae (Fig. 3).
Habitat – On a decaying wood log, accompanied in humus rich soil with over heavily rotted
litter on the ground.
Specimens examined – CHINA, Hainan Province, Jiangfengling Mountain, Coniferous
rainforest, 18° 44′ N, 108° 51′ E, elev. 550 m, collection date unknown, collector X.L Wu (GACP
HNU40), 9 August 2014, collector T.C Wen (GACP14080965, GACP14080967).
Notes – Ganoderma applanatum belongs in subgenus Elfvingia, which includes non-laccate
species. It has a worldwide distribution and, is a central species in G. applanatum-G. australe
complex. The type material originates from Europe and is preserved at the Rijksherbarium Leiden,
Netherlands. This species is characterized by having; a thin, acute margin in pileus, unbranched
terminal endings of skeletal hyphae, and (5.5–9.0) μm size, ellipsoid basidiospores (Pegler &
Young 1973, Ryvarden & Gilbertson 1993, Leonard 1998, Wu & Dai 2005). Our study
macroscopically and microscopically agrees with the description of Pegler & Young (1973).
Ganoderma australe can be distinguished from G. applanatum by having larger basidiospores
(Moncalvo & Ryvarden 1997). Richter et al. (2015) mentioned that G. lipsiense also belongs in this
complex, and has been treated by some researchers as the correct name for G. applanatum.
However, the Nomenclature Committee for Fungi has approved the G. applanatum is the correct
name for this taxon to conserve the basionym of G. applanatum, Boletus applanatus Pers. against
the earlier name, Boletus lipsiensis Batsch. Furthermore, in our phylogenetic analysis European
species of G. lipsiense and G. applanatum have been clustered in similar clade with 100% BS value
(Fig. 2). Ganoderma applanatum can be distinguished chemotaxonomically by the presence of
Benzopyranone derivatives such a Ganodermaaldehyde and Applanatins which have been isolated
from specimens originating from both North America and Asia (Ming et al. 2002, Wang et al.
2007). This species was first described from China by Teng (1963) and later Zhao (1988) re-
examined the specimen and synonymized with Ganoderma mirivelutinum J.D. Zhao. However,
Wang et al. (2009) proved that holotype of G. mirivelutinum is morphologically distinct from
Teng’s specimen, but identical to Ganoderma applanatum var. laevisporum C.J. Humphrey &
Leus-Palo which characterized by basidiospores with smooth inner and outer wall layers. Our
collections agree with the description provided by Ryvarden & Gilbertson (1993). Moreover, we
identify our specimens as Ganoderma cf. applanatum, since authentic type materials from the
Northern temperate hemisphere were not included in this study.

948
Figure 3 – Ganoderma cf. applanatum (GACP HNU40) a Upper surface. b Lower surface. c Pores
in the lower surface (4.5×). d Cut surface. e–g Spores (100×). h Generative hyphae (40×). i Binding
hyphae (40×). j Skeletal hyphe (40×). Scale bars: e–g = 10 μm, h–j = 5 μm.

949
Ganoderma australe (Fr.: Fr.) Pat., 1889, Bull. Soc. mycol. Fr. 5, p. 71
≡ Polyporus australis Fr.: Fr., 1828, Elench. Fung. 1, p. 108 (Type lost, initially described
from Pacific Is.).
(See Index Fungorum for other synonyms)
Basidiome annual, sessile, non-laccate, woody. Pileus; 6–13 × 5.5–6 cm, sub-orbicular,
plano-convex, applante, at center slightly swollen: upper surface brownish orange (6C4) to brown
(6D4), distinctly concentrically sulcate, with irregularly ruptured crust overlying the pellis: margin
soft, slightly lobate and concolorous with the pileus; lower surface yellowish white (4A2).
Hymenophore up to 10 mm long, indistinctly stratose; pores initially yellowish white (4A2), later
brown (6D6), 4–5 per mm; pores circular or sub-circular. Context up to 1.5 cm thick, dry, duplex;
lower layer, brown (6D6), fibrous/pithy, composed of coarse loose fibrils; upper layer dark brown
(7F8), woody. Basidiospores (n=40) (6.1–)7.6–9.2–10.8(–11.5) × (4.7–)5.3–7.6–7.9(–8.5) μm
(Qm=1.5, Q=0.9–2.6, with myxosporium). (4.9–)5.3–6.6–7.9(–8.5) × (3.1–)3.4–4.2–5.1(–5.8) μm
(Qm=1.5, Q=0.9–2.9, without myxosporium), ellipsoid, brownish orange (6C8) to brown (6D8),
with a brown eusporium bearing fine, short, and distinct echinulae, overlaid by a hyaline
myxosporium. Pileipellis a hymeniderm, (6–28) μm, brownish orange (6C8) to brown (6D8),
composed of apically acanthus like branched cells, dextrinoid. Context trimitic; generative hyphae
not observed; skeletal hyphae (n=30) (4.5–)3.1–3.9–4.1(–4.8) μm, thick-walled, sometimes
branched, brownish orange (6C8); binding hyphae (n=30) (3.5–)4.2–5.6–5.8(–6.1) μm, thick-
walled, branched, brownish orange (6C8) (Fig. 4).
Habitat – On a decaying wood log, accompanied in humus rich soil with over heavily rotted
litter on the ground.
Specimens examined – CHINA, Hainan Province, Jiangfengling Mountain, Coniferous
rainforest, 18° 44′ N, 108° 51′ E, elev. 550 m, 11 August 2014, collector T.C. Wen
(GACP14081175, GACP14081134, GACP14081161, GACP14081671), collection date unknown,
collector X.L Wu (GACP HNU42, GACP HNU5456).
Notes – Ganoderma australe (Fr.) Pat. 1889, belongs to G. applanatum-australe complex and
can be distinguished from G. applanatum by having larger basidiospores (Moncalvo & Ryvarden
1997). This species, common in the tropics, has never been recorded from Europe but was mistaken
for the domestic G. adspersum (Smith & Sivasithamparam 2000). The type specimen of G. australe
cannot be found or is destroyed and the only material deposited in the Royal Botanic Gardens Kew
is different from original collection (Ryvarden & Gilbertson 1993). Currently, the only neotype
available is from Europe. However, careful neotypification of G. australe is needed to resolve the
problem that persists in taxonomy and geographical distribution (Kaliyaperumal & Kalaichelvan
2008). Moncalvo & Ryvarden (1997) considered Ganoderma annulare as an illegitimate name,
since the basionym is illegitimate as nomenclaturly superfluous and earliest valid name to use is G.
australe. Ganoderma tornatum and G. australe have been synonymized many times, but some
researchers mistakenly prioritized G. tornatum over G. australe in publications. Furthermore, the
concept of G. tornatum as a variety of G. applanatum was not accepted (Humphrey & Leus 1931,
Steyaert 1975a, b, Ryvarden & Johnson 1980, Gilbertson & Ryvarden 1986, Ryvarden &
Gilbertson 1993). This species was first introduced by Teng (1963) from China and after that
several authors studies on this species (Tai 1979, Zhao et al. 1981, Zhao 1989, Bi et al. 1993, Teng
1996, Zhao & Zhang 2000, Dai et al. 2011). Our collections agree with the descriptions provided
by Moncalvo & Ryvarden (1997).
Ganoderma calidophilum J.D. Zhao, L.W. Hsu & X.Q. Zhang, Acta microbiol. Sin. 19(3): 270
(1979)
Basidiome annual, stipitate, laccate, corky. Pileus 4–7 × 3–4.5 cm, up to 1.5 cm thick at the
base, sub-orbicular, rotund, upper surface; reddish brown (8E8) to dark brown (8F8), radially
rugose with turberculate bumps and ridges and rivulose depressions, with irregularly ruptured crust
overlying the pellis, margin; blunt to truncate, greyish orange (5B3), lower surface; light brown
(7D5). Hymenophore up to 25 mm long, indistinctly stratose; pores initially greyish brown (7D3),

950
Figure 4 – Ganoderma australe (GACP14081671) a Upper surface. b Pores in the lower surface
(4.5×). c Pieces of the specimen. d Cut surface. e Cross section of the basidiome (4.5×). f–g Spores
(100×). h Generative hyphae (100×). i Skeletal hyphae (100×). Scale bars: f–g = 10 μm, h–i = 5
μm.

951
bruising brown (8E8), pores circular or sub-circular, 5–6 per mm. Context up to 4 mm thick,
duplex, dry; lower layer reddish brown (8E8), fibrous, composed of coarse loose fibrils; upper layer
dark brown (8F8), corky to woody. Stipe eccentric, dorsally lateral to nearly dorsal, sub-cylindrical,
dark brown (8F8), 6 × 13 cm, 1.4 cm thick at the base. Basidiospores (n=40) (7.3–)8.1–9.2–
13.8(14.6–) × (–4.8)5.3–7.6–7.9(–8.6) μm (Qm=1.5, Q=0.9–2.6 with myxosporium). (5.2–)6.1–6.6–
9.6(–9.9) × (3.5–)4.9–5.1–5.9(–6.3) μm (Qm=1.5, Q=0.9–2.9, without myxosporium), broadly
ellipsoid, light orange (5A5), brownish orange (6C8) to light brown (5A5), with a brown eusporium
bearing fine, short, and distinct echinulae, overlaid by a hyaline myxosporium. Pileipellis a
hymeniderm, brownish orange (6C8), clavate like cells, dextrinoid. Context trimitic; generative
hyphae (n=30) (1.7–)2.3–2.6–3.4(–3.9) μm, hyaline, thin-walled without clamp connections;
skeletal hyphae (n=30) (3.1–)4.5–5.4–6.5(–6.8) μm thick-walled, nearly solid, sometimes branched,
greyish brown (5B5) to brownish orange (6C8); binding hyphae (n=30) (2.5–)3.6–4.1–4.8(–5.4)
μm, thick-walled, branched, nearly solid, light orange (5A5) to brownish orange (6C8) (Fig. 5).
Habitat – On rotten wood, in dry dipterocarp forest and in upper mixed deciduous forest or
growing up from soil.
Specimens examined – CHINA, Hainan Province, Diaoluoshan National Nature Reserve, 18°
68′ N, 109° 95′ E, elv. 1058 m, collection date unknown, collector X.L. Wu (GACP HNU08,
GACP HNU86).
Notes – Ganoderma calidophilum was introduced by Zhao et al. (1979) from Hainan
Province, China as a stipitate wood-inhabiting polypore with a laccate pileus and broadly ellipsoid
basidiospores. Later, many Chinese authors described this species from China (Zhao et al. 1981,
Zhao 1989, Bi et al. 1993, Zhao & Zhang 2000). Wu et al. (1999), Li et al. (2010) and Dai et al.
(2011) included G. calidophilum in their polypore diversity checklists from Hainan Province. The
observed morphological characters of G. calidophilum in this study, are similar with previous
descriptions (Zhao et al. 1979, Bi et al. 1993) even we were unable to obtain molecular data from
our G. calidophilum specimen. Wang & Wu (2007) suggested that this species is a synonym of G.
flexipes and Cao et al. (2012) subsequently followed this conclusion. However, present study
confirms G. flexipes and G. calidophilum are morphologically different with pileus size, colour,
context and basidiospore size (Bi et al. 1993, Wang & Wu 2007). Hence, this species needs further
studies with authentic type materials, fresh collections and suitable molecular markers.
Ganoderma ellipsoideum Hapuar., T.C. Wen & K.D. Hyde, sp. nov.
Index Fungorum number: IF554384, Facesoffungi number: FoF04458
Etymology: refers to the basidiospore shape ‘ellipsoid’
Holotype: GACP14080966
Basidiome annual, sessile (usually with a distinctly contracted base), non laccate, woody. Pileus 3–
6.5 × 4–5 cm, up to 1.5 cm thick at the base, flabelliform, plano convex, applante, upper surface;
hard, several layers thick, brownish orange (5C4) to yellowish brown (5E8) crust overlies pellis,
not cracking, yellowish brown (5F8), containing fibrous pithy context, concentrically sulcate zones
with turberculate bumps and ridges and rivulose depressions, alternating brownish orange (5C4) to
yellowish brown (5E8) zones, differentiated zone at the point of attachment; margin 4 mm thick,
rounded and concolorous with the pileus; lower surface white (4A1) to yellowish white (4A2).
Hymenophore up to 10 mm long, indistinctly stratose; pores initially yellowish white (4A2), later
yellowish brown (5D8) to brownish yellow (5C8), pores circular or sub-circular. Context up to 1.5
cm thick, dry, duplex; lower layer yellowish brown (5F8), pithy, composed of coarse loose fibrils,
soft; upper layer dark brown (7F8), woody. Basidiospores (n=50) (5.3–)6.1–6.8–7.3(–7.8) × (2.9–
)3.7–4.2–4.6(–5.2) μm (Qm=1.6, Q =1.2–2.1, with myxosporium) (3.6–)4.2–4.7–5.3(–6.2) × (1.8–
)2.4–2.9–3.3(–3.8) μm (Qm=1.6, Q=0.9–2.6, without myxosporium), greyish orange (5B4),
elongate to ellipsoid, with a brown eusporium bearing fine, short and distinct echinulae, overlaid by
a hyaline myxosporium. Pileipellis a hymeniderm, yellowish brown (5D8) to brownish yellow
(5C8), composed of apically acanthus like branched cells, dextrinoid. Context trimitic; generative
hyphae (n=50) (0.6–)1.1–1.3–2.7(–3.1) μm in width, thin-walled, colorless; skeletal hyphae

952
(n=100) (0.7)1.2–1.6–3.3(3.8) μm in width, greyish orange (5B3), thick-walled, sometimes
branched; binding hyphae (n=75) (1.2–)1.5–2.3–4.1(–4.8) μm in width, greyish orange (5B3),
thick-walled, branched, intertwined with the skeletal hyphae (Fig. 6).
Figure 5 – Ganoderma calidophilum (GACP HNU86) a, b Upper surface. c Pores in the lower
surface (4.5×). d Lower surface. e Section of the context. f–i Spores (100×). j Skeletal hyphae
(100×). Scale bars: f–i = 10 μm, j–k = 5 μm.

953
Habitat – On a decaying wood log, accompanied in humus rich soil with over heavily rotted
litter in forest, mossy coniferous forests, producing basidiomata from late summer to late autumn.
Currently only known from Hainan Province, China.
Material examined – CHINA, Hainan Province, Jiangfengling Mountain, 18° 44′ N, 108° 51′
E, elev. 550 m, 9 August 2014, collector T.C. Wen (GACP14080966, holotype), (GACP14081215
and GACP14080968, paratypes).
Notes – Ganoderma ellipsoideum is a new member of genus Ganoderma (Fig. 6) and it is
distinguished by ellipsoid spores (6.1–7.3 × 3.7–4.6) μm, with a brown eusporium bearing fine,
short and distinct echinulae. It is clustered and morphologically similar with G. gibbosum (Blume
& T. Nees) Pat., which is characterized by the sub-flabellate to sub-dimidiate, non laccate pileus,
duplex, brown 4–5 mm thick context, trimitic hyphal system, brown, 7–14 mm long tubes, elongate
to ellipsoid spores, 6.6–8.3 × 5–6 μm, smooth hyaline outer wall, echinulate inner wall (Bi et al.
1993). Furthermore, it clustered with unidentified Ganoderma species recorded from Thailand
(JM95/5) and China (JM98/233) (description unavailable), which belong to the G australe complex
and the G. applanatum complex, respectively (Moncalvo & Buchanan 2008). The species is
currently only known from the type locality, Jiangfengling Mountain, Hainan, China.
Ganoderma flexipes Pat., Bull. Soc. mycol. Fr. 23(2): 75 (1907)
= Fomes flexipes (Pat.) Sacc. & Traverso, Syll. fung. (Abellini) 19: 710 (1910)
= Polyporus flexipes (Pat.) Lloyd, Mycol. Writ. 3 (Syn. Stip. Polyporoids): 104 (1912)
Basidiome annual, stipitate, laccate, corky. Pileus 1.5–4 × 0.5–2.5 cm, up to 0.2 cm thick at
the base, flabelliform, rotund, upper surface; reddish brown (8E8), concentrically sulcate zones
with turberculate bumps and ridges and rivulose depressions, with irregularly ruptured crust
overlying the pellis, margin; soft, yellowish brown (5E8), lower surface; light brown (7D5).
Hymenophore up to 10 mm long, indistinctly stratose; pores initially greyish brown (7D3), bruising
brown (8E8), pores circular or sub-circular or isodiametric, 3–4 per mm. Context up to 4 mm thick,
triplex, dry; lower layer reddish brown (8E8), fibrous, composed of coarse loose fibrils; upper layer
dark brown (8F8), corky to woody. Stipe eccentric, dorsally lateral to nearly dorsal, sub-cylindrical,
dark brown (8F8), 4 × 7 cm, 0.4 cm thick at the base. Basidiospores (n=25) (9.5–)9.2–10.2–11.5(–
12.2) × (6.8–)7.9–8.1–7.9(–8.4) μm (Qm =1.4, Q=0.9–2.1 with myxosporium). (8.2–)8.1–8.5–9.6(–
9.9) × (3.2–)4.9–5.5–5.9(–6.4) μm (Qm=1.4, Q=0.9–2.9, without myxosporium), ellipsoid, light
orange (5A5), brownish orange (6C8) to light brown (5A5), with a brown eusporium bearing fine,
short and distinct echinulae, overlaid by a hyaline myxosporium. Pileipellis a hymeniderm,
brownish orange (6C8), clavate like cells, dextrinoid. Context trimitic; generative hyphae (n=30)
(1.7–)2.3–2.6–3.4(–3.9) μm, hyaline, thin-walled without clamp connections; skeletal hyphae
(n=30) (3.1–)4.5–5.4–6.5(–6.8) μm thick-walled, nearly solid, sometimes branched, greyish brown
(5B5) to brownish orange (6C8); binding hyphae (n=30) (2.5–)3.6–4.1–4.8(–5.4) μm, thick-walled,
branched, nearly solid, light orange (5A5) to brownish orange (6C8)(Fig. 7).
Habitat – On rotten wood, in dry dipterocarp forest and in upper mixed deciduous forest or
growing up from soil.
Specimens examined – CHINA, Hainan Province, Jiangfengling Mountain, 18° 44′ N, 108°
51′ E, elev. 550 m, collection date unknown, collector X.L. Wu (GACP HNU42, GACP
HNU5450), 13 August 2014, collector T.C. Wen (GACP14081339).
Notes – Ganoderma flexipes was introduced by Patouillard (1907) from Vietnam and has
been recorded from China, India, Nepal and Pakistan (Steyaert 1972). Many Chinese authors
reported this species from China (Zhao et al. 1981, Zhao 1989, Zhao & Zhang 2000, Wang & Wu
2007). Ganoderma flexipes can easily be recognized by its small reddish brown pileus, long and
thin stipe, reddish brown to dark brown context, and ellipsoid or ovoid basidiospores. Our
collections fall within the range (8–9.9–13 × 5.5–6.5–8) μm, measured by Steyaert (1972) and fits
that of the lectotype (9–11 × 6–7.5) μm, measured by Ryvarden (1983) for G. flexipes.
Furthermore, the pore diameter and corky context composed of mostly fine and thick-walled

954
hyphae, in the collections from Hainan Province resemble those of authenticated specimens and of
the holotype (Steyaert 1972).
Figure 6 – Ganoderma ellipsoideum (GACP14080966, holotype) a Upper surface. b Lower
surface. c Cut surface. d Pores in the lower surface (4.5×). e Pieces of the specimen. f–i Spores
(100×). j Skeletal hyphae (100×). k Binding hyphae (100×). Scale bars: f–i = 10 μm, j–k = 5 μm.

955
Figure 7 – Ganoderma flexipes (GACP HNU5450) a, b Upper surface. c Lower surface. d Cut
surface. e Pores in the lower surface (5×). f–h Spores (100×). i Skeletal hyphae (100×). j Generative
hyphae (40×). k binding hyphae (100×). Scale bars: f–h = 10 μm, i–k =5 μm.

956
Ganoderma gibbosum (Blume & T. Nees) Pat., Ann. Jard. Bot. Buitenzorg, suppl. 1: 114 (1897)
= Polyporus gibbosus Blume & T. Nees, Nov. Act. Academiae Caesareae Leopoldino-
Carolinae Germanicae Naturae Curiosorum 13: 19, t. 4:1-4 (1826)
= Fomes amboinensis var. gibbosus (Blume & T. Nees) Cooke, Grevillea 13 (68): 118 (1885)
= Fomes gibbosus (Blume & T. Nees) Sacc. Syll. Fung. 6: 156 (1888)
= Scindalma gibbosum (Blume & T. Nees) Kuntze, Revisio generum plantarum 3 (2): 518
(1898)
Basidiome annual, sessile (usually with a distinctly contracted base), non laccate, woody.
Pileus 6–12 × 3–6 cm, up to 3 cm thick at the base, spathulate, upper surface; hard, several layers
thick, light brown (6D5) to light brown (6D6), crust overlies the pellis, concentric zones with
turberculate bumps and ridges and rivulose depressions; margin with numerous undulations and
irregularities, wavy, 2 mm thick, concolorous with the pileus; lower surface light brown (5D4).
Hymenophore up to 6 mm long, indistinctly stratose, light brown (5D4), pores circular or sub
circular, 4–5 pores per mm. Context up to 3 cm thick, dry, duplex, lower layer; brown (6E8), pithy,
composed of coarse loose fibrils, upper layer; dark brown (6F8), woody. Basidiospores (n=50)
(4.3–)6.9–7.6–9.2(–10.5) × (3.8–)4.6–5.6–5.7(–6.1) μm (Qm =1.7, Q=0.8–2.5, with myxosporium).
(3.6–)4.8–6.0–7.6(–8.7) × (2.0–)2.7–3.5–4.5(–6.7) μm (Qm=1.7, Q=0.8–3.2, without
myxosporium), ellipsoid to elongate, brown (6D8) to light brown (6D6), eusporium bearing fine,
short, and distinct echinulae, overlaid by a hyaline myxosporium, bitunicate. Pileipellis a
hymeniderm, brown (6D8) to light brown (6D6), composed of apically acanthus like branched
cells, dextrinoid. Context trimitic; generative hyphae (n=30) (1.3–)1.6–2.1–2.7(–3.6) μm in width,
thin-walled, hyaline; skeletal hyphae (n=30) (4.0–)4.8–4.7–5.2(–5.8) μm in width, brown (6F8),
dextrinoid, thick-walled; binding hyphae (n=30) (2.6–)3.6–4.7–5.8(–6.1) μm in width, brown
(6F8), thick-walled, branched, intertwined the skeletal hyphae (Fig. 8).
Habitat – On rotten wood, in dry dipterocarp forest and in upper mixed deciduous forest or
growing up from soil.
Specimens examined – CHINA, Hainan Province, Jiangfengling Mountain, 18° 44′ N, 108°
51′ E, elev. 550 m, collection date unknown, collector X.L. Wu (GACP HNU32), 15 August 2014,
collector T.C. Wen (GACP14070442, GACP14081517).
Notes – Ganoderma gibbosum is known from Australia (Saccardo 1888), China (Teng
1963, Zhao et al. 1981, Zhao 1989, Bi et al. 1993, Zhao & Zhang 2000), Indonesia (Java) and
Vietnam (Parmasto 1986), but the location of the type is not known (Moncalvo & Ryvarden 1997).
Ganoderma gibbosum was considered to be subspecies of G. applanatum (Zhao 1989). However,
G. gibbosum was renamed as G. incrassatum: a species which known only from Australia, based on
the monophyletic origin and low level of sequence variation (Smith & Sivasithamparam 2003). Wu
et al. (1999), Li et al. (2010) and Dai et al. (2011) included G. gibbosum in their Polypore diversity
checklists from Hainan Province. Our collections from Hainan Province agree well with the
descriptions provided by Bi et al. (1993) and Zhao & Zhang (2000).
Ganoderma hoehnelianum Bres. Annls mycol. 10(5): 502 (1912)
= Ganoderma shangsiense J.D. Zhao, Acta Mycol. Sin. 7(1): 17 (1988).
Basidiome annual, sessile (usually with a distinctly contracted base), laccate, woody. Pileus
3.1–5.4 × 1.9–3.5 cm, up to 1.5 cm thick at the base, clustered, suborbicular, plano concave,
triquetrous to sub applanate; upper surface hard, several layers thick, with brown (6D6) crust
overlies the pellis, containing fibrous pithy context, concentrically sulcate zones, alternating light
brown (6D6) and dark brown (6F8) zones, differentiated zone at the point of attachment, margin;
soft, 3 mm thick, rounded, yellowish white (4A2), lower surface; white (5A1). Hymenophore up to
5 mm long, indistinctly stratose, greyish orange (5B3), pores circular or sub circular. Context up to
1.5 cm thick, dry, triplex; lower layer brown (6F8) fibrous, composed of coarse loose fibrils, soft;
middle layer dark brown (6E8); upper layer light brown (6D6), woody. Basidiospores (n=50) (9.4–
)10.5–11.7–12.1(–12.5) × (9.4–) 9.7–10.8–11.1(–11.8) μm (Qm =1.1, Q=0.9–1.2, with
myxosporium), (8.9–)9.7–10.6–11.3(–11.6) × (7.2–)8.8–9.3–10.1(–10.5) μm (Qm=1.1, Q=0.8–1.1,

957
without myxosporium), broadly ovoid to subglobose, brown (6E8), with a brown eusporium
bearing fine, short and distinct echinulae, overlaid by a hyaline myxosporium, bitunicate. Pileipellis
a hymeniderm, light brown (6D6) to dark brown (6E8), composed of clavate cells, dextrinoid.
Context trimitic; generative hyphae (n=100) (0.3–)0.9–1.1–1.9(–2.8) μm in width, thin-walled,
hyaline; skeletal hyphae (n=40) (0.9–)1.5–1.9–(–4.0) μm in width, thick-walled, greyish orange
(6B3); binding hyphae (n=50) (1.4–)1.8–2.9–3.5(–5.6) μm in width, thick-walled, branched,
greyish orange (6B3), intertwined with skeletal hyphae (Fig. 9).
Habitat – On a decaying wood log, accompanied in humus rich soil with over heavily rotted
litter on the ground, mossy temperate mixed coniferous forests. Producing basidiomata from
summer to late autumn.
Specimens examined – CHINA, Hainan Province, Jiangfengling Mountain, Coniferous
rainforest, 18°44′N, 108°51′E, elev. 550 m, 9 August 2014, collector T.C. Wen (GACP14080913,
GACP14081635).
Notes – Ganoderma hoehnelianum was introduced by Bresadola (1912) from Java,
Indonesia. The context color, basidiospore characteristics, and cuticular composition are key
diagnostic features for the identification of G. hoehnelianum (Wang & Wu 2010). The latter
authors revealed, by examining the isotype, that G. hoehnelianum is an earlier name for
Ganoderma shangsiense (Wang et al. 2005), a species originally described from China. This
species was recorded as G. shangsiense in the Hainan Province polypore diversity checklists
provided by Li et al. (2010) and Dai et al. (2011). Our collections from Hainan Province agree well
with the descriptions provided by Bresadola (1912) and Wang & Wu (2010).
Ganoderma multiplicatum (Mont.) Pat., Bull. Soc. mycol. Fr. 5(2, 3): 74 (1889)
≡ Polyporus multiplicatus Mont. (1854)
(See Index Fungorum for synonyms)
Basidiome annual, sessile (usually with a distinctly contracted base), strongly laccate, woody.
Pileus 6–9 × 3.5–5 cm, up to 3 cm thick at the base, sub-orbicular, plano convex, sub applante;
upper surface hard, several layers thick, orange (6D8) to brown (7E8), crust overlies the pellis,
containing fibrous pithy context, swollen differentiated zone at the point of attachment; margin soft
or having irregularities, 1 cm thick, rounded, white (5A1); lower surface white (5A1) to orange
(5B2). Hymenophore up to 4 mm long, indistinctly stratose, white (5A1) to orange (5B2), pores
circular or sub circular or isodiametric. Context up to 3 cm thick, dry, duplex; lower layer brown
(7E8), fibrous, composed of coarse loose fibrils, soft; upper layer pale orange (5A3), woody.
Basidiospores (n=30) (4.8–)5.9–7.1–8.3(–9.9) × (3.3–)6.6–4.3–8.3(–9.6) μm (Qm=1.6, Q=1.2–2.0,
with myxosporium). (3.8–)4.5–5.3–6.2(–7.2) × (1.8–)2.3–2.6–3.5(–3.8) μm (Qm=1.3, Q=1.3–3.8,
without myxosporium), broadly ellipsoid to elongate, greyish orange (5B5) to brown (6E8),
eusporium bearing fine, short, and distinct echinulae, overlaid by a hyaline myxosporium,
bitunicate. Pileipellis a hymeniderm, greyish orange (5B5) to brown (6E8), composed of apically
acanthus like branched cells, dextrinoid. Context dimitic; skeletal hyphae (n=30) (2.6–)2.9–3.4–
3.8(–4.2) μm in width, light brown (6D4), thick walled, sometimes branched; binding hyphae
(n=30) (1.2–)2.1–2.6–3.3(–3.7) μm in width, light brown (6D4), thick walled, branched,
intertwined the skeletal hyphae (Fig. 10).
Habitat – On a decaying wood log, accompanied in humus rich soil with over heavily rotted
litter on the ground, mossy temperate mixed coniferous forests. Producing basidiomata from
summer to late autumn.
Specimens examined – CHINA, Hainan Province, Jiangfengling Mountain, Coniferous
rainforest, 18° 44 ′N, 108° 51′ E, elev. 550 m, 13 August 2014, collector T.C. Wen
(GACP14081328, GACP14081330).
Notes – Ganoderma multiplicatum (Mont.) Pat. was originally collected in French Guyana
(Moncalvo & Ryvarden 1997) and is characterized by subglobose to broadly ellipsoid
basidiospores (7–8 × 5–6 μm) with a dimitic or trimitic hyphal system. This species has described
from China by many authors (Zhao et al. 1979, Zhao et al. 1981, Zhao 1989, Zhao & Zhang 2000,

958
Wang & Wu 2007). The morphology of G. multiplicatum collections from Hainan Province agrees
well with that of the holotype as described by Gottlieb & Wright (1999a), Ryvarden (2000).
Figure 8 – Ganoderma gibbosum (GACP14070442) a Upper surface. b Lower surface. c Pores in
the lower surface. d Pores in the lower surface (4.5×). e Cut surface. f–j Spores (100×). K Binding
hyphae (100×). i Generative hyphae. m Skeletal hyphae (100×). Scale bars: f–i = 10 μm, j–k = 5
μm.

959
Figure 9 – Ganoderma hoehnelianum (GACP14080913). a Upper surface. b Lower surface. c Cut
surface. d Pores in the lower surface (4.5×). e Transverse section of the specimen showing spores
(40×). f–h Spores (100×). i Generative hyphae (100×). j Binding hyphae (100×). Scale bars: f–h =
10 μm, i–j = 5 μm.

960
Figure 10 – Ganoderma multiplicatum (GACP14081328) a Upper surface. b Lower surface.
c Pores in the lower surface (4.5×). d Cut surface. e Pieces of the specimen. f–i Spores (100×).
j Binding hyphae (100×). k Skeletal hyphae (100×). l Generative hyphae (100×). Scale bars: f–i
=10 μm, j–l =5 μm.
This species has been considered as similar to G. chalceum (Corner 1983) and was
synonymized as G. subamboinense (Ryvarden 2000). However, Correia de Lima et al. (2014)

961
suggested that G. chalceum and G. subamboinense are not synonyms of G. multiplicatum on the
basis of molecular data. Ganoderma multiplicatum was subsequently found in Africa (Steyaert
1980), Asia (Zhao 1989, Bhosle et al. 2010) and South America (Bolaños-Rojas et al. 2016).
Ganoderma orbiforme (Fr.) Ryvarden (as ‘orbiformum’) (2000)
≡ Polyporus orbiformis Fr. (1838)
= Ganoderma boninense Pat., Bull. Soc. mycol. Fr. 5(2, 3): 72 (1889)
= Ganoderma mastoporum (Lév.) Pat. [as ‘malosporum’], Bull. Soc. mycol. Fr. 5(2, 3): 75
(1889)
= Ganoderma fornicatum (Fr.) Pat. sensu Imazeki, Bull. Tokyo Sci. Mus. 1: 47 (1939)
= Ganoderma cupreum (Cooke) Bres, Annls mycol. 9(3): 268 (1911)
= Ganoderma subtornatum Murrill, Bull. Torrey bot. Club 34: 477 (1907)
= Ganoderma densizonatum J.D. Zhao & X.Q. Zhang, in Zhao, Zhang & Xu, Acta Mycol.
Sin. 5(2): 86 (1986)
= Ganoderma limushanense J.D. Zhao & X.Q. Zhang, Acta Mycol. Sin. 5(4): 219 (1986)
(See Index Fungorum for other synonyms)
Basidiome annual, sessile (usually with a distinctly contracted base), non laccate, woody.
Pileus 3–8 × 2.5–6 cm, up to 2 cm thick at the base, suborbicular, plano convex, applanate, upper
surface; hard, several layers thick, light brown (6D5), crust overlies the pellis, concentrically
sulcate zones with turberculate bumps and rivulose depressions, differentiated zone at the point of
attachment, radially rugose; margin with numerous undulations and irregularities, 3 mm thick,
rounded and concolourous with the pileus; lower surface light brown (6D4). Hymenophore up to
3mm long, indistinctly stratose, orange grey (6B2), pores circular or sub-circular. Context up to 2
cm thick, dry, duplex; lower layer dark brown (7F8), pithy, composed of coarse loose fibrils, soft;
upper layer light brown (5D4), woody. Basidiospores (n=47) (6.9–)7.6–8.3–10.3(–10.3) × (3.6–
)4.8–5.3– 5.8(–5.7) μm (Qm=1.8, Q=1.5–2.6, with myxosporium). (4.7–)5.8–6.5–7.8(–8.5) × (2.8–
)3.2–3.4–4.3(–5.6) μm (Qm=1.8, Q=1.5–2.6, without myxosporium), yellowish brown (5D8),
elongate, eusporium bearing fine, short, and distinct echinulae, overlaid by a hyaline myxosporium,
bitunicate. Pileipellis a hymeniderm, dark brown (7F8), composed of apically acanthus like
branched cells, dextrinoid. Context trimitic; generative hyphae (n=30) (0.4–)0.5–1.3–1.8(–2.6) μm
in width, thin-walled, hyaline; skeletal hyphae (n=30) (3.2–)3.8–4.2–4.8(–5.2) μm in width, light
brown (5D6), thick-walled; binding hyphae (n=30) (1.3–)1.1–1.5–2.6(–2.9) μm in width, light
brown (5D6), thick-walled, branched, intertwined the skeletal hyphae (Fig. 11).
Habitat – On a decaying wood log, accompanied in humus rich soil with over heavily rotted
litter on the ground in forest, mossy temperate mixed coniferous forests, producing basidiomata
from summer to late autumn.
Specimens examined – CHINA, Hainan Province, Jiangfengling Mountain, Coniferous
rainforest, 18° 44′ N, 108° 51′ E, elev. 550 m, 9 August 2014, collector T.C. Wen;
(GACP14080918, GACP14080953) 10 August 2014; (GACP14081069), 11 August 2014;
(GACP14081108, GACP14081182, GACP14081184, GACP14081185), 12 August 2014;
(GACP14081202, GACP14081232, GACP14081235, GACP14081239), 13 August 2014;
(GACP4081310, GACP4081329, GACP14081333, GACP14081340), Wuzshishan Mountain,
Coniferous rainforest, 18° 53′ N, 109° 38′ E, elev. 1240 m, 12 August 2014, collector T.C Wen;
(GACP14081666).
Notes – Ganoderma orbiforme (Fr.) Ryvarden was originally described as ‘Polyporus
orbiformis’ and identified by the strongly amyloid, irregular, tuberculate to blunted cuticle cells and
moderately large basidiospores (Ryvarden 2000). This is a. tropical species and was originally
described from Guinea in Africa, and also known from Bonin Island in the Pacific and in the
Neotropics (Ryvarden 2000). Ganoderma orbiforme possess greater variability in morphology than
other taxa in Ganoderma; rigid basidiocarp with a weakly to strongly laccate, partly laccate or dull
pileus, variably brown context, ellipsoid to ovoid basidiospores with fine and short echinulae, and
purplish brown pore surface at maturity (Wang et al. 2014). Ganoderma cupreum, G. fornicatum,

962
G. mastoporum, G. orbiforme, G. subtornatum and the species which originally described from
China; G. densizonatum and G. limushanense are morphologically very similar to one another in
basidiome texture, pilear cuticle structure, context and pore color and basidiospore characteristics.
Hence, based on morphological and molecular data, it was concluded, that the above mentioned
taxa are conspecific and G. orbiforme is the earliest valid name for use (Wang et al. 2014). Our
collections from Hainan Province agree with the descriptions provided by Ryvarden (2000).
Figure 11 – Ganoderma orbiforme (GACP14081235) a Upper surface. b Lower surface. c section
of the specimen. d Pores in the lower surface (4.5×). e–g Spores (100×). h Binding hyphae (100×).
i Skeletal hyphae (100×). Scale bars: e–g = 10 μm, h–i = 5 μm.

963
Ganoderma cf. resinaceum
Basidiome perennial, sessile, usually with a distinctly contracted base, laccate, corky. Pileus
7–12 × 4–10 cm, up to 1.5 cm thick at the base, sub-orbicular, rotund, upper surface; reddish brown
(8E8) to dark brown (8F8), concentrically sulcate zones with turberculate bumps and ridges and
rivulose depressions, radially rugose, with irregularly ruptured crust overlying the pellis, margin;
blunt to truncate, greyish orange (5B3), lower surface; light brown (7D5). Hymenophore up to 25
mm long, indistinctly stratose; pores initially greyish brown (7D3), bruising brown (8E8), pores
circular or sub-circular, 5–6 per mm. Context up to 4 mm thick, duplex, dry; lower layer reddish
brown (8E8), fibrous, composed of coarse loose fibrils; upper layer reddish brown (8E8), corky to
woody. Basidiospores (n=25) (9.3–)10.9–11.2–12.5(–12.8) × (8.5–)8.8–9.1–7.9(–8.3) μm (Qm=1.5,
Q=1.1–2.6 with myxosporium). (6.7–)6.9–7.1–7.9(–8.1) × (3.5–)4.9–5.1–5.9(–6.3) μm (Qm=1.5,
Q=0.9–2.9, without myxosporium), ellipsoid, light orange (5A5), brownish orange (6C8) to light
brown (5A5), with a brown eusporium bearing fine, short, and distinct echinulae, overlaid by a
hyaline myxosporium. Pileipellis a hymeniderm, brownish orange (6C8), clavate like cells,
dextrinoid. Context dimitic; skeletal hyphae (n=30) (3.1–)4.5–5.4–6.5(–6.8) μm, thick-walled,
nearly solid, sometimes branched, greyish brown (5B5); binding hyphae (n=30) (2.5–)3.6–4.1–
4.8(–5.4) μm, thick-walled, branched, nearly solid, light brown (7D5) (Fig. 12).
Habitat – On a decaying tree trunk, accompanied in humus rich soil with over heavily rotted
litter on the ground.
Specimens examined – CHINA, Hainan Province, Diaoluoshan National Nature Reserve, 18°
68′ N, 109° 95′ E, elv. 1058 m, collection date unknown, collector X.L. Wu (GACP HNU02,
GACP HNU58).
Notes – Ganoderma resinaceum was introduced by Boudier in 1889 from France (Patouillard
1889). This species is characterized by a pileus with varying colours, a fibrous spongy
homogeneous context, larger basidiospores and an amyloid pileipellis (Ryvarden 2004).
Ganoderma resinaceum has been described by Steyaert (1972), Ryvarden & Gilbertson (1993) by
emphasizing varying pileus color changes from shiny varnish laccate orange-yellow, brown,
ochraceous-brown to reddish-brown pileus on young basidiocarps, that turn black or blackish when
mature or ageing. This is because of different carotenoid pigments or/and secondary metabolites
produced by G. resinaceum. However, this colour evolution and shiny varnish appearance of the
pileus rarely applies to specimens of G. resinaceum growing in tropical climates (Ayissi &
Mossebo 2014). Steyaert (1980) grouped this species with Ganoderma parvulum as a complex due
to the variations and inconsistencies of taxonomy in Ganoderma. Ganoderma resinaceum may be
confused with G. boninense (= G. orbiforme) (Torres-Torres et al. 2015). However, the particular
combination of characters such as basidiomata color, fibrous spongy context without resinous
bands, almost cylindrical to clavate cuticle cells, and ellipsoid basidiospores, with free and
relatively thin pillars make it possible to identify this species as a species complex. Ganoderma
resinaceum may be confused with those of G. pfeifferi which has a similar resinous layer on the
upper crust in pileus, but this species has a dark brown to umber context and wider spores
Ganoderma lucidum has a lighter context without a darker zone above the tubes and no resinous
layer on the crust, thus making it much more glossy and shiny even in older specimens. The spores
also appear more coarsely warted than those of G. resinaceum (Ryvarden & Gilbertson 1993).
Moncalvo (2000) differentiated G. resinaceum and G. weberianum from other related species. This
specis may be confused with G. chalceum, which however has a black band in the context
(Ryvarden 2004). Ganoderma resinaceum has been widely used in organisms very dissimilar to the
species originally described by Boud and numerous new names have been designated for this
common species based on macro morphological variations. This species was described for the first
time in China by Zhao (1989) and later few Chinese authors recorded and described this species
(Wu et al. 1999, Zhao & Zhang 2000, Wu & Dai 2005). Our study agrees both macroscopically and
microscopically with the description from the holotype made by Ryvarden (2004) and Torres-
Torres et al. (2012). We identify our specimens as Ganoderma cf. resinaceum, since authentic type
materials were not included in this study. This species is distributed Southern and Central Europe

964
with a few isolated localities in southern Denmark, North Africa. Asia, North America and
widespread in the Paleotropics (Ryvarden & Gilbertson 1993).
Figure 12 – Ganoderma cf. resinaceum (GACP HNU02) a Upper surface. b Lower surface. c Cut
surface. d Pores in the lower surface (5×). e-h Spores (100×). i Cuticle cells (100×). j Binding
hyphae (100×). K Skeletal hyphae (100×). Scale bars: e–h = 10 μm, j–l = 5 μm.

965
Ganoderma tropicum (Jungh.) Bres., Annls mycol. 8(6): 586 (1910)
≡ Polyporus tropicus Jungh. (1838)
= Fomes tropicus (Jungh.) Cooke, Grevillea 14(no. 69): 19 (1885)
= Scindalma tropicum (Jungh.) Kuntze, Revis. gen. pl. (Leipzig) 3(2): 519 (1898)
Basidiome annual, sessile (usually with a distinctly contracted base), strongly laccate, woody.
Pileus 4–8 × 2.5–6 cm, up to 1 cm thick at the base, spathulate, plano convex; upper surface hard,
several layers thick, yellowish brown (5F8), orange (6D8) to brown (7E8), crust overlies the pellis,
containing fibrous pithy context, swollen differentiated zone at the point of attachment; margin
soft, 1 cm thick, rounded, white (5A1); lower surface white (5A1) to orange (5B2). Hymenophore
up to 4 mm long, indistinctly stratose, white (5A1) to orange (5B2), pores sub circular or
isodiametric. Context up to 3 cm thick, dry, duplex; lower layer brown (7E8), fibrous, composed of
coarse loose fibrils, soft; upper layer light brown (5A5), woody. Basidiospores (n=25) (10.8–)11.2–
12.1–12.8(–13.1) × (8.3–)9.6–10.1–10.8(11.1) μm (Qm=1.3, Q=0.9–1.5, with myxosporium). (7.9–
)8.8–9.1–10.2(–10.8) × (5.8–)6.4–7.3–7.8(–9.8) μm (Qm=1.3, Q=1.1–1.7, without myxosporium),
broadly ellipsoid, brownish orange (6C8) to light brown (5A5) eusporium bearing fine, short and
distinct echinulae, overlaid by a hyaline myxosporium, bitunicate. Pileipellis a hymeniderm,
greyish orange (5B5) to brown (6E8), composed of apically acanthus like branched cells,
dextrinoid. Context trimitic; generative hyphae (n=30) (0.4–)0.6–1.4–1.8(–2.6) μm in width, thin-
walled, hyaline skeletal hyphae (n=30) (2.8–)3.1–3.4–3.8(–4.3) μm in width, light brown (6D4),
thick walled, sometimes branched; binding hyphae (n=30) (1.1–)2.2–2.5–3.4(–3.8) μm in width,
light brown (6D4), thick walled, branched, intertwined the skeletal hyphae (Fig. 13).
Habitat – On a decaying wood log, accompanied in humus rich soil with over heavily rotted
litter on the ground in forest, mossy temperate mixed coniferous forests, producing basidiomata
from summer to late autumn.
Specimens examined – CHINA, Hainan Province, Wuzshishan Mountain, Coniferous
rainforest, 18° 53′ N, 109° 38′ E, elev. 1240 m, 15 August 2014, collector T.C. Wen
(GACP14081511, GACP14081518).
Notes – Ganoderma tropicum was introduced as Polyporus tropicus by Junghuhn (1838)
from Java, Indonesia and later, Bresadola (1910) transferred this species to Ganoderma.
Ganoderma tropicum is distributed from lowland tropical Asia and the subtropics (Steyaert 1972)
and its taxonomy has been well-resolved with a strong bootstrap support (Wang et al. 2012, Yang
& Feng 2013). This species is characterized by broadly ellipsoid to ellipsoid basidiospores with
thick echinulae and is considered as a member of the G. lucidum species complex (Zhou et al.
2015). Among the Chinese Ganoderma species, G. flexipes, G. multipileum, G. sichuanense, and G.
tsugae are the most similar species to G. tropicum, since they share a reddish-brown pileal surface,
similar basidiospores and cuticle cells, however G. tropicum differs from G. multipileum by having
strongly echinulate basidiospores (Cao et al. 2012). Ganoderma tropicum was first reported from
China by Tai (1979) and then few Chinese researchers have reported on this species (Zhao 1989, Bi
et al. 1993, Zhao & Zhang 2000). Our collections from Hainan Province agree well with the
descriptions provided by the latter researchers. Wu et al. (1999), Li et al. (2010) and Dai et al.
(2011) recorded this species from Hainan Province in their Polypore diversity checklists.
Ganoderma tropicum causes white root and butt rot on several species of Acacia in forest
plantations in China (Dai et al. 2007) and further it is one of the most aggressive pathogens that
causes basal stem rot in oil palm (Turner 1981, Wong et al. 2012).
Ganoderma sinense J.D. Zhao, L.W. Hsu & X.Q. Zhang, Acta Microbiol. Sin. 19(3): 272 (1979)
= Ganoderma formosanum T.T. Chang & T. Chen in Trans. Or. Mycol. Soc. 82: 731
(1984).
Misapplications:
Ganoderma japonicum (Fr. Lloyd in Teng, Fungi of China; 447 (1963): Tai, Syll. Fung.
Sin.: 469 (1979), Teng, Fungi of China: 326 (1996), non Polyporus japonicus Fr.,Epicrisis: 442
(1838) (Ganoderma japonicum (Fr.) Lloyd, Mycol. Writ. 3: Syn. Stip. Polyp.: 102(1912)).

966
Ganoderma lucidum (Leyss.) P. Karst. var. japonicum (Fr.) Bres. in Teng in Sinensia 5:199
(1934). non Polyporus japonicus Fr., Epicrisis: 442 (1838).
Figure 13 – Ganoderma tropicum (GACP14081518) a Upper surface. b Lower surface. c Cut
surface. d Pores in the lower surface (5×). e-g Spores (100×). h Binding hyphae (100×).
i Generative hyphae (100×). j Skeletal hyphae (100×). Scale bars: e–g = 10 μm, h–j = 5 μm.

967
Basidiome annual, stipitate, strongly laccate, corky. Pileus 7–8 × 5.0–5.5 cm, up to 0.5 cm
thick at the base, reniform, sub-dimidiate; upper surface dark brown (8F8), radially rugose,
concentrically sulcate, with irregularly ruptured crust overlying the pellis; margin blunt,
concolorous with the pileus; lower surface brown (7E8). Hymenophore up to 15 mm long,
indistinctly stratose; pores initially dark brown (8F8), bruising brown (7E8), pores circular or sub-
circular, 4–5 per mm. Context up to 1 cm thick, duplex, dry; lower layer brown (7D8), fibrous,
composed of coarse loose fibrils; upper layer dark brown (8F8), corky. Stipe eccentric, sub
cylindrical, concolorous with the pileus, 8 × 13 cm. Basidiospores (n=25) (10.6–)11.5–12.5–13.5(–
14.3) × (7.1–)7.7–8.3–8.9(–11.7) μm (Qm=1.5, Q=1.1–1.8 with myxosporium). (6.8–)9.2–10.2–
11.1(–11.7) × (5.4–)6.1–6.8–7.7(–8.4) μm (Qm=1.5, Q=1.1–1.8, without myxosporium), ellipsoid,
brownish orange (7C8) to reddish orange (7B8) with a brown eusporium bearing fine, short, and
distinct echinulae, overlaid by a hyaline myxosporium. Pileipellis a hymeniderm, brownish orange
(7C8), clavate like cells, dextrinoid. Context dimitic; generative hyphae (n=25) (0.5–1.6–3.3) μm
diam, colorless, thin-walled; skeletal hyphae (n=20) (3.3–4.5–5.2) μm thick-walled, nearly solid,
sometimes branched, orange (5A6); binding hyphae (n=20) (3.4–4.6–5.5) μm, thick-walled,
branched, nearly solid, light brown (5A5) (Fig. 14).
Habitat – On rotten wood, in dry dipterocarp forest and in upper mixed deciduous forest or
growing up from soil. Producing basidiomata from summer to late autumn.
Specimens examined – CHINA, Hainan Province, Jiangfengling Mountain, Coniferous
rainforest, 18° 44′ N, 108° 51′ E, elev. 550 m, collection date 12 August 2014, collector T.C. Wen
(GACP14081236, GACP14081260).
Notes – Ganoderma sinense originated from China and is characterized by slightly
longitudinally crested basidiospores and a uniformly brown to dark brown context. Our G. sinense
collections from Hainan Province agree with that of the holotype as described by Wang & Wu
(2007), moreover with descirptons given by Bi et al. (1993), Zhao & Zhang (2000), and Wu & Dai
(2005). This species was considered as G. lucidum or G. japonicum by Chinese authors (Teng
1934, 1939, 1963). Ganoderma sinense differs from G. lucidum in thin-fleshed basidiomata,
with long, slender stipes, rarely branched skeletal hyphae with bovista type binding hyphae
(Pegler & Yao 1996). Ganoderma japonicum (= G. dimidiatum) has a yellow pileus when
young, red brown or rusty brown at maturity and off-white context (Pegler & Yao 1996), and is
distinguished from the purplish black to black pileus and brown context in G. sinense.
Ganoderma formosanum introduced by Chang & Chen (1984) from Taiwan, is very similar with
G. sinense in morphology. However, Zhao & Zhang (2000) considered this species as a distinct
species based on the duplex context and ovoid basidiospores and this conclusion was
subsequently followed by Wu & Dai (2005). However, Wang & Wu (2007) studied both
holotypes of G. formosanum and G. sinense carefully and concluded that both species similar in
morphology. Furthermore, it was concluded that the latter species are synonyms and the earliest
valid name to be used is G. sinense based on molecular and morphological data (Moncalvo et al.
1995a, Wang & Wu 2007).
Ganoderma sinense is morphologically similar to G. orbiforme in having a purplish black to
black laccate pileus, uniformly brown context or with whitish streaks or patches near the cuticle, a
dorsally lateral or lateral stipe and a subtropical-tropical distribution. Ganoderma sinense
however, can easily be distinguished from G. orbiforme by its erect stipe, cuticle composed of
clavate cells, and ovoid basidiospores with few, long and thick echinulae (Wang et al. 2014).
Pharmacopoeia of People’s Republic of China 2000 edition (part one) has mentioned that the
general term “Lingzhi” covers both “Chizhi (G. lucidum)” and “Zizhi (G. sinense)”. However,
with the aid of molecular data, G. lucidum and G. sinense were clearly distinguished as two
different species (Zhou et al. 2008, Liao et al. 2015). Furthermore, based on chemotaxonomical
data, Oxygenated lanostane-type triterpenes have been proven as suitable markers to differentiate
G. lucidum and G. sinense (Fu et al. 2008). Among all Ganoderma species, only G. lucidum and
G. sinense have been approved to produce health products (Zhou et al. 2016).

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Figure 14 – Ganoderma sinense (GACP14081236) a, b Upper surface. c Lower surface. d Cut
surface. e Pores in the lower surface (5×). f–h Spores (100×). i Binding hyphae (40×). j generative
hyphae (40×). k Skeletal hyphae (40×). Scale bars: e–h = 10 μm, i–j = 5 μm.
Ganoderma subresinosum (Murrill) C.J. Humphrey, Mycologia 30(3): 332 (1938)
≡ Fomes subresinosus Murrill 1908
= Polyporus mamelliporus Beeli, Bull. Soc. R. Bot. Belg. 62: 62 (1929)
= Trachyderma subresinosum (Murrill) Imazeki, Bull. Gov. Forest Exp. Stn Tokyo 57: 119
(1952)
= Magoderna subresinosum (Murrill) Steyaert, Persoonia 7(1): 112 (1972)
= Amauroderma subresinosum (Murrill) Corner, Beih. Nova Hedwigia 75: 93 (1983)

969
Basidiome annual, sessile (with distinctly contracted base), weakly laccate, woody. Pileus
16–20 × 11–13 cm, up to 4 cm thick at the base, sub-orbicular; upper surface dark brown (8F4),
radially rugose, concentrically sulcate with irregularly ruptured crust overlying the context; margin
blunt or wavy, yellow brown; lower surface brownish orange (6C4). Hymenophore up to 20 mm
long, indistinctly stratose; pores initially greyish orange (5B3), bruising brownish orange (6C4),
pores circular, 3–5 per mm. Context up to 2 cm thick, duplex, dry; lower layer, light orange (5A4),
fibrous, composed of coarse loose fibrils; upper layer brownish orange (6C5), corky. Basidiospores
(n=25) (13.3–)13.9–15.3–16.5(–17.8) × (8.0–)9.2–10.2–11.1(–12.1) μm (Qm=1.5, Q=1.3–1.9 with
myxosporium). (11.6–)12.2–13.5–14.8(–16.3) × (6.3–)7.3–8.4–9.5(–10.4) μm (Qm=1.6, Q=1.3–2.2,
without myxosporium), ellipsoid to ellongate, orange (6A6), pale orange (6A3) to greyish orange
(5B4), with a brown eusporium bearing fine, short, and distinct echinulae, overlaid by a hyaline
myxosporium. Pileipellis a hymeniderm, light orange (5A4), clavate like cells, dextrinoid. Context
dimitic; generative hyphae not observed; skeletal hyphae (n=25) (2.2–3.1–4.0) μm, thick-walled,
nearly solid, sometimes branched, orange white (5A2); binding hyphae (n=20) (1.4–2.3–3.3) μm,
thick-walled, branched, nearly solid, brownish orange (6C4) (Fig. 15).
Habitat – On a decaying wood log, accompanied in humus rich soil with over heavily rotted
litter on the ground. Producing basidiomata from summer to late autumn.
Specimens examined – CHINA, Hainan Province, Wuzhishan Mountain, Coniferous
rainforest, 18° 53′ N, 109° 38′ E, elev. 1240 m, collection date 16 August 2014, collector T.C. Wen
(GACP14081663, GACP14081690).
Notes – Murrill (1908) introduced this species from the Philippines, as Fomes subresinosus,
which characterized by smooth and hyaline basidiospores. Humphrey (1938) transferred this
species to Ganoderma. Imazeki (1952) included this species in the genus Trachyderma as T.
tsunodae Imazeki. Steyaert (1972) introduced the genus Magoderna, typified by M. subresinosus
to accommodate species with dimidiate to pleuropodal basidiomata, anticlinal hyphae in the pilear
surface and ovoid-ellipsoid to spherical basidiospores without a truncate apex. Furthermore, the
genus has been considered as synonym of Amauroderma (A. subresinosum) (Corner 1983).
However, this species is now recorded as G. subresinosum in Index Fungorum
(www.indexfungorum/org/names/Names.asp). Recently some researchers suggested Magoderna
(M. subresinosum) might be accepted at generic level based on strong morphological and molecular
data (Gomes-Silva et al. 2015, Costa-Rezende et al. 2016, 2017). Ganoderma subresinosum was
recorded from China by Teng (1963), Tai (1979), Teng (1996) as Fomes subresinosum and later
reported by, Zhao (1989) and Zhao & Zhang (2000) as A. subresinosum. The known distribution of
this species extends from the Philippines to West Africa through Malaysia, Myanmar, India, Sri
Lanka, Borneo Island and Eastern and Central Africa (Steyaert 1972).
Genus Amauroderma P. Karst., 1881, Rev. Mycol. (Toulouse) 3, p. 17.
= Amauroderma (Pat.) Torrend, Brotéria, sér. Bot. 18: 121 (1920)
= Ganoderma sect. Amauroderma Pat., Bull. Soc. mycol. Fr. 5(2, 3): 75 (1889)
= Lazulinospora Burds. & M.J. Larsen, Mycologia 66(1): 97 (1974)
= Magoderna Steyaert, Persoonia 7(1): 111 (1972)
= Whitfordia Murrill, Bull. Torrey bot. Club 35: 407 (1908)
After Amauroderma was introduced, Torrend (1920) worked extensively on this genus in
South America. He published an important work based mainly on spore shape (globose or oblong,
never truncate) and the presence of a stipe (usually dull, like the pilear surface) and recorded 28
species of Amauroderma placed within three sections. Later, Amauroderma was carefully revised
by Furtado (1981), who recognized 27 species. He defined Amauroderma by the globose to
subglobose basidiospores, with double walls, stipitate basidiomes and a tropical distribution
pattern. Ryvarden (2004) described 21 species of Amauroderma from the Neotropics, using the
same genus circumscription. However, according to Index Fungorum (www.indexfungorum.org,
accessed 21 August 2018), there are 135 binomials of this genus. Amauroderma is a widespread
tropical genus that usually occurs on roots of living or dead trees or, is more rarely, wood

970
inhabiting (Furtado 1981, Ryvarden 2004) and cause white rots. Macroscopically, Amauroderma
shares similarities with Ganoderma, a similar basidiome shape of central or lateral stipe and laccate
or dull surface. Ganoderma can be separated from Amauroderma by its distinctly truncate
basidiospores, and most Ganoderma species grow on dead wood, while most Amauroderma species
grow in the ground from buried roots/woods (Ryvarden 2004). Amauroderma species are regarded
as economically valuable because of their important medicinal properties and pathogenicity (Dai et
al. 2007, 2009, Jiao et al. 2013, Chan et al. 2013). In China, taxonomic research on Amauroderma
began with Teng, who recorded ten species (Teng 1936, 1939, 1963). Later Zhao et al. (1979, 1983,
1984) and Zhao & Zhang (1986, 1987, 2000) reported 20 species, nine of which were new. Here,
we record two Amauroderma species found in Jiangfengling Mountain in Hainan Island, based on
micro- and macro morphological characteristics together with molecular data.
Figure 15 – Ganoderma subresinosum (GACP14081663) a Upper surface. b Lower surface. c Cut
surface. d Pores in the lower surface (5×). e–h Spores (100×). i Binding hyphae (100×). j Skeletal
hyphae (100×). Scale bars: e–h = 10 μm, i–j = 5 μm.

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Amauroderma austrosinense J.D. Zhao & L.W. Hsu, Acta Mycol. Sinica 3: 20. 1984.
Basidiome annual, stipitate, weakly laccate, corky. Pileus 3.5–5 × 3.5–4.5 cm, up to 0.5 cm
thick at the base, orbicular, convex; upper surface soft, only few layers, containing fibrous context,
alternating brownish orange (6C8) to light brown clay brown (6D4) concentrically zones, distinctly
differentiated zone at the point of attachment; margin rounded, brownish orange (6C8), 1–2 mm
thick; lower surface white (6A1). Hymenophore up to 2 mm long, orange white (6C2), pores
circular or sub-circular or isodiametric. Context up to 2 mm thick at the base, dry, duplex; lower
layer greyish brown (6D3), fibrous, soft, upper layer; orange white (6C2), composed of tightly
interwoven, finer fibrils. Stipe stout cylindric, concolorous with the pileus, dorsally lateral, 6.5 × 2
cm. Basidiospores (n=50) (6.8–)7.1–7.6–8.3(–9.6) × (5.2–)6.3–7.0–7.8(–9.5) μm (Qm=1.4, Q=0.9–
1.5, with myxosporium). (3.9–)4.8–5.5–6.2(–6.3) × (3.6–)4.3–4.9–5.6(–6.7) μm (Qm=1.2, Q=0.9–
2.5, without myxosporium), subglobose, brown (6E8) to dark brown 6F8), with a brown eusporium
bearing fine, short, and distinct echinulae, overlaid by a hyaline myxosporium. Pileipellis a
hymeniderm, brownish orange (6C8), composed of apically acanthus like branched cells. Context
trimitic; generative hyphae (n=20) (2.6–)1.9–2.5–3.1(–3.5) μm, hyaline, thin-walled with clamp
connections, rarely seen; skeletal hyphae (n=40) (1.9–)2.2–2.8–3.8(–4.0) μm, light brown (6D4),
thick-walled, sometimes branched; binding hyphae (n=40) (1.1–)1.7–3.1–3.9(–4.3) μm, thick-
walled, branched, light brown (6D4), intertwined the skeletal hyphae (Fig. 16).
Habitat – On rotten wood, in dry dipterocarp forest and in upper mixed deciduous forest or
growing up from soil. Producing basidiomata from summer to late autumn.
Specimens examined – CHINA, Hainan Province, Jiangfengling Mountain, coniferous
rainforest, 18° 44’ N, 108° 51’ E, elev. 550 m, collection date 10 August 2014, collector T.C. Wen
(GACP14081030, GACP14081045).
Notes – Amauroderma austrosinense described by Zhao et al. (1984) from Hainan Province
and is identified by its umbelliform basidiomes with distinct concentric rings, small pores and
basidiospores. Even we were unable to obtain DNA, our collections agree well with the description
of the holotype provided by Li & Yuan (2015). Later, this species was described by many Chinese
authors (Zhao 1989, Zhao & Zhang 2000, Wu & Dai 2005, Li & Yuan 2015). Amauroderma
camerarium (Berk.) J.S. Furtado is similar to A. austrosinense in its clay-colored, densely
concentrically zoned, and radially furrowed pileus (Li & Yuan 2015) and differs in its larger
basidiospores (12–15 × 10–13 µm) (Ryvarden 2004). Amauroderma leucosporum is similar to A.
austrosinense in having similar-sized subglobose to globose basidiospores, but it can be
distinguished by it’s finely to distinctly villous pileus and stipe (Corner 1983).
Amauroderma rugosum (Blume & T. Nees) Torrend, Brotéria, sér. bot. 18(no. 2): 127 (1920)
= Amauroderma amoiense J.D. Zhao & L.W. Hsu, Acta Mycol. Sinica 2: 164. (1983)
= Amauroderma wuzhishanense J.D. Zhao & X.Q. Zhang, Acta Mycol. Sinica 6: 208. (1987)
(See Index Fungorum for other synonyms)
Basidiome annual, stipitate, weakly laccate, corky. Pileus 2.5–3.5 × 2.0–2.5 cm, up to 0.5 cm
thick at the base, subreniform; upper surface brownish orange (6C4) to brown (6E8), radially
rugose, concentrically sulcate with irregularly ruptured crust; margin blunt or wavy, concolorous
with the pileus; lower surface greyish orange (5B5). Hymenophore up to 10 mm long, indistinctly
stratose; pores initially brownish orange (5C4), bruising brown (6E8), pores circular, 3–5 per mm.
Context up to 8 mm thick, duplex, dry; lower layer light brown (5D6), fibrous, composed of coarse
loose fibrils, brown (6E8), corky. Stipe eccentric, sub cylindrical, concolorous with the pileus, 5 × 7
cm, (0.5 along stipe) cm. Basidiospores (n=20) (11.8–)12.9–13.4–13.9(–14.1) × (10.2–)10.8–11.2–
11.8(–12.1) μm (Qm=1.2, Q=1.1–1.4, with myxosporium). (11.9–)12.1–12.9–13.1(–13.5) × (9.6–
)10.2–10.9–11.2(–11.8) μm (Qm=1.2, Q=0.9–1.3, without myxosporium), subglobose, greyish
orange (5B3), with a brown eusporium, overlaid by a hyaline myxosporium. Pileipellis a
hymeniderm, brownish orange (5C4), clavate like cells, dextrinoid. Context dimitic; generative
hyphae not observed; skeletal hyphae (n=25) (1.2–2.5–3.5) μm, thick-walled, nearly solid,

972
sometimes branched, orange white (5A2); binding hyphae (n=20) (0.6–1.7–2.6) μm, thick-walled,
branched, nearly solid, orange white (5A2) (Fig. 17).
Figure 16 – Amauroderma austrosinense (GACP14081030) a In the wild. b Upper surface.
c Lower surface. d Pores in the lower surface (4.5×). e Cut surface. f–h Spores (100×). i Binding
hyphae (40×). j Generative hyphae (40×). k Skeletal hyphae (40×). Scale bars: f–h = 10 μm, i–k = 5
μm.

973
Figure 17 – Amauroderma rugosum (GACP14081210) a Upper surface. b Lower surface. c Cut
surface. d Pores in the lower surface (5×). e–h Spores (100×). i Skeletal hyphae (100×). j Binding
hyphae (100×). Scale bars: e–h = 10 μm, i–j = 5 μm.
Habitat – On rotten wood, in dry dipterocarp forest and in upper mixed deciduous forest or
growing up from soil. Producing basidiomata from summer to late autumn.
Specimens examined – CHINA, Hainan Province, Jiangfengling Mountain, coniferous
rainforest, 18° 44′ N, 108° 51′ E, elev. 550 m, elev. 1350 m, collection date 9 August 2014,
collector T.C. Wen (GACP14080910, GACP14080952, GACP14080929, GACP14080956).
Notes – Amauroderma rugosum is a soil-inhabiting saprobe (Baran De 1991) and Blume & T.
Nees described this species as Polyporus rugosus on the basis of specimens from Java in 1826
(Torrend 1920). Berkeley (1856) introduced a new species Porothelium rugosum from Brazil.
Steyaert (1972) suggested Polyporus rugosus and Porothelium rugosum were similar species when
comparing their morphological descriptions. Patouillard (1889) transferred Polyporus rugosus to

974
Ganoderma (as G. rugosum). In 1894, Patouillard took up the name Porothelium rugosum Berk.
and changed it as Ganoderma sprucei Pat. since he considered that there were already an epithet
called “rugosum” in Ganoderma. Torrend (1920) transferred G. sprucei Pat. to Amauroderma (A.
rugosum). Furtado (1968) synonymized P. rugosum Berk. as Amauroderma sprucei (Pat.) Torrend.
Hence, it was a problem whether P. rugosum Berk. (= G. sprucei Pat.) and G. rugosum (BI. &
Nees) Pat. were similar species. Torrend therefore continued to use ‘sprucei’, the earliest epithet
available in Amauroderma. Considering the combination of Porothelium rugosum in
Foraminispora the epithet is available. However, Costa-Rezende et al. (2017) proposed
Foraminispora, a new genus to accommodate Porothelium rugosum (= Amauroderma sprucei)
with the aid of strong morphological and molecular data. According to Index Fungorum and
MycoBank A. rugosum (Blume & T. Nees) Torrend is now a legitimized species characterized by
mesopodal and often excentric, or often pleuropodal pileus with subglobose basidiospores. This
species was reported by Teng (1963) from China and later many Chinese researchers reported this
species (Tai 1979, Zhao et al. 1981, Zhao 1989, Bi et al. 1993, Teng 1996, Zhao & Zhang 2000).
Ryvarden & Johansen (1980) provided detailed description of this species and our collections agree
well with that description and moreover, the descriptions provided by latter Chinese authors.
Conclusions
Species concepts within Ganoderma and Amauroderma have evolved from morphological
species to phylogenetic and biological species. Macroscopic, microscopic, as well as molecular
data confirmed one novel species together with records of 12 known species of Ganoderma and
two known species of Amauroderma. This is the first comprehensive study of these two genera
from Hainan Province, China. However, we need epitypes, reference collections, fresh collections
and vouchered multigene nucleotide sequence data of more informative DNA markers to determine
the taxonomy of Ganoderma and Amauroderma. The present study provides a snapshot useful for
future taxonomic and phylogenetic studies, as well as for cultivation studies because of the high
potential to use in biotechnology.
Acknowledgements
This work was financed by the Science and Technology Foundation of Guizhou Province
(No. [2017]2511-1), and the Science Research Foundation of Guizhou University (No. 201309).
Dr. Shaun Pennycook is thanked for nomenclatural advices. Kalani K. Hapuarachchi is grateful to
Hansika Perera, Ishani Goonasekara and Monika Dayarathna for their valuable comments and
suggestions.
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