Bull. Natl. Mus. Nat. Sci., Ser. B, 47(2), pp. 59–69, May 21, 2021

Species Identiﬁcation Based on DNA of Selected Mushrooms from
Myanmar (1) Lactarius austrotorminosus and17 Other Taxa Newly Reported from Myanmar
Kentaro Hosaka1, *, Kyung-Ok Nam1, Wah Wah Linn2 and Mu Mu Aung2
1

Department of Botany, National Museum of Nature and Science,
4–1–1 Amakubo, Tsukuba, Ibaraki 305–0005, Japan
2
Forest Research Institute, Forest Department, Yezin, Nay Pyi Taw, Myanmar*E-mail: khosaka@kahaku.go.jp(Received 15 February 2021; accepted 24 March 2021)

Abstract A total of 33 specimens (22 species) of mushrooms collected from Myanmar werecharacterized and identiﬁed to species based on morphological observation and DNA barcoding
using the internal transcribed spacer (ITS) region of nuclear ribosomal DNA newly generated forthis study. The ITS sequences of many Myanmar materials showed signiﬁcant similarity (99% orhigher) to the existing data in GenBank, but some had much lower similarity, warranting further,more rigorous taxonomic studies. Results from the BLAST search indicated that the countries oforigin of the top hits varied. Although the majority of Myanmar materials showed close aﬃnities
with materials from the adjacent areas, such as Thailand, India or China, some materials had
signiﬁcant similarity in ITS sequences with those from geographically distant areas, such as
Argentina and French Guiana. A total of 18 species, including Lactarius austrotorminosus, arenewly reported from Myanmar.
Keywords : Agaricomycota, Agaricomycetes, BLAST, biogeography, Dacrymycetes, distribution,
fungi, internal transcribed spacer, inventory, Southeast Asia.

Introduction
To clarify fauna and ﬂora (including fungi) of
Myanmar and the surrounding areas, a joint
research team of National Museum of Nature andScience, Japan, and Forest Research Institute,
Myanmar has been conducting biological inventories in Myanmar since 2016. As a result, theﬁrst comprehensive ﬁeld guidebook of ﬂora andfauna (including fungi) has been published(Tanaka, 2021). The guidebook contains coloredphotographs and morphological descriptions of
selected species, and a total of 25 species ofmushrooms are illustrated.
Little is known for mushroom diversity in© 2021 National Museum of Nature and Science

Myanmar. The most comprehensive list of mushrooms in Myanmar was published by Thaung(2007), listing ca. 180 species. Since then, no
intensive surveys on mushrooms have been conducted until 2016. Since the beginning of thejoint research project described above, more than
400 additional specimens of mushroom were collected, including some new records from Myanmar (Hosaka et al., 2019, 2020). Mushroom
inventory in Myanmar is still at very preliminary
stage and many new records and species remain
to be described.
While the publication of the ﬁrst ﬁeld guidebook containing fungi is a signiﬁcant event, thebook does not cover the molecular characterization of mushrooms from Myanmar. Because
many mushroom species cannot easily be distin-

60

Kentaro Hosaka et al.

guished each other by morphological characters
alone, comparing them by DNA sequence data is
crucial. For example, Hosaka et al. (2020) demonstrated that materials of Sphaerobolus from
Myanmar are morphologically indistinguishable
from the existing species, but they clearly represent a new species in the genus by analyzing
DNA sequences of mitochondrial and nuclear
genes.
In this study, we characterized a total of 22
species of mushrooms covered in the ﬁeld guidebook (Hosaka and Linn, 2021) based on newly
generated DNA Hosaka and Linn of the internaltranscribed spacer (ITS) region of nuclear ribosomal DNA.
Materials and Methods
Collecting and Curation of Specimens
A total of 33 specimens used in this study
were collected during the years 2016–2017 fromvarious locations in Myanmar. The samples were
photographed under natural light in the ﬁeld andthen wrapped with aluminum foil for transport tothe laboratory. In the laboratory, the samples
were photographed again using artiﬁcial lighting
and more in-depth macroscopic observation was
conducted. The samples were dried with low heatand good air circulation using a food dehydrator
for 24 hours.
In addition to dried materials, small fragments(ca. 3 cubic millimeters each) of clean, sterile tissue from freshly collected materials were cut
using a clean razor blade. Contamination of visible soil particles and other materials was carefully avoided. The tissue fragments were soaked
in dimethyl sulfoxide (DMSO) buﬀer (Seutin etal., 1991) with an addition of 100 mM Tris-HCl(pH 8.0) and 0.1 M sodium sulﬁte (Na2SO3), andthey were stored at room temperature until further molecular experiment became possible, following the procedures of Hosaka (2009) andHosaka and Castellano (2008).
All specimens collected during the ﬁeldwork
were deposited at Forest Research Institute,
Myanmar, and the duplicate at the herbarium of

the National Museum of Nature and Science(TNS), Tsukuba, Japan. All tissue samples werestored in freezers (−80°C) at the Center for
Molecular Biodiversity Research, National
Museum of Nature and Science.
DNA Preparation, PCR, and Sequencing
DNA was extracted from the tissue fragments
stored in DMSO buﬀer. Tissues were groundunder liquid nitrogen using a mortar and pestle.
DNA extractions used a modiﬁed cetyltrimethylammonium bromide (CTAB) extraction followed
by glass milk puriﬁcation methods as summarized by Hosaka (2009) and Hosaka and Castellano (2008).
DNA sequence data were obtained from the
ITS region. The primer combination of ITS5 and
ITS4 (White et al., 1990) was used for PCR
ampliﬁcations. PCR reactions were carried out
using 20 μl reaction volumes each containing:
1 μl genomic DNA, 1 μl dNTPs (4 mM), 1 μl ofeach primer (8 μM), 0.5 units of Taq polymerase(TaKaRa, Tokyo, Japan), 2 μl MgCl2 (25 mM),
2 μl bovine serum albumin (BSA). PCR reactions
were performed using the following parameters:
95°C for 3 min; 35 cycles of 95°C for 1 min,
51°C for 45 sec, 72°C for 1 min; and 72°C for
15 min.
PCR products were electrophoresed in 1%
agarose gels stained with ethidium bromide andvisualized under UV light. When ampliﬁcation
bands were conﬁrmed, PCR products were thenpuriﬁed using the ExoSap-IT (Millipore, Molsheim, France) and directly sequenced using the
Big Dye Terminator Cycle Sequencing Kit on
ABI3500 (Applied Biosystems Inc., Foster City,
CA, USA), following the manufacturerʼs instructions.
Molecular Analyses
The obtained raw sequences were edited using
ATGC version 7.1.0 (GENETYX Corporation,
Tokyo, Japan). All edited ITS sequences wereﬁrst complied into a single FASTA ﬁle, and they
were then analyzed using the GenBank basic
local alignment search tool (BLAST) searchDNA barcoding of Myanmar mushrooms (1)(Zhang et al., 2000) to conﬁrm their phylogenetic aﬃnities with registered taxa. Default settings of blastn option, including “Standard databases (nr etc.)” with “Nucleotide collection (nr/nt)” optimized for “Highly similar sequences(megablast),” were used. All major scores
resulted from the BLAST search, e.g., “Total
Score”, “Query Cover” and “Per. Ident,” wererecorded along with taxon names and the countries of origin of the top hits. When the top hits
did not have information on speciﬁc epithet or
the country of origin, data from the next highest
hit were also recorded. BLAST search was conducted on February 2, 2021. After the BLAST
results were obtained, the distribution records ofeach species (including synonyms) weresearched using the Global Biodiversity Information Facility (GBIF) database (GBIF, 2021) andthe occurrence data were downloaded to comparewith literature.
Results and Discussion
Molecular Identiﬁcation of Specimens
Results from the BLAST search using theblastn option with default settings mostly corroborated the validity of our ﬁeld identiﬁcation of
the species (Tables 1 and 2). A total of 30 out of
33 sequences showed percent identities of 97%
or higher with the registered data. The twosequences identiﬁed as Craterellus atratus (KHMYA16-168) and Fistulina hepatica (KHMYA16-057) showed percent identities of
96.26% and 96.98%, respectively. The lowest
percent identity was 92.3% for the collection tentatively identiﬁed as Turbinellus ﬂoccosus (KHMYA16-017).
For this study, we have applied the speciesnames obtained from the top hits of the BLAST
search (Table 1). However, many taxa are likely
to contain multiple biological species. Furthermore, we have not conﬁrmed taxonomic identityof the registered data based on voucher specimens. It is therefore advisable that taxon names
used in this study are treated as tentative ones,
and more rigorous taxonomic studies should be

61

conducted in the future.
The countries of origin of the top BLAST hits
indicated that many have close aﬃnities withmaterials from Southeast Asia, South Asia andEast Asia (Table 2). Thailand was represented bya total of 7 sequences, followed by China (5) andIndia (3). Some sequences from distantly locatedregions were also represented as the top hit,including Geastrum schweinitzii (Argentina),
Phallus merulinus (French Guiana) and Psathyrella candolleana (Nigeria). No signiﬁcant hits
were observed from materials from Australia andNorth America. These results indicate a closeaﬃnity of the Myanmar mycoﬂora with that ofAsia, especially Southeast Asia. However, thedegree of taxon sampling is signiﬁcantly diﬀerent across species and countries, and therefore
the results should be interpreted with caution.
Specimens Molecularly Identiﬁed in this Study
A total of 33 specimens were identiﬁed to species based on morphological observation and ITS
sequences. Because morphological characters
were described in detail by Hosaka and Linn
(2021), we hereby provide the information oncollection and molecular data along with the distribution records of each species.
Amanita ceciliae (Berk. & Broome) Bas
Material examined: Myanmar: Mandalay, Than
Daung Village near Pyin Oo Lwin (21.851267,
96.374177), alt. 985 m, under Quercus spp., 31
Aug 2016, K. Hosaka (KH-MYA16-100).
Type locality: United Kingdom.
Remarks: The species is generally consideredwidely distributed in the Northern Hemisphere(Imazeki and Hongo, 1987; Hosaka and Linn,2021), which is corroborated by the GBIF
records (GBIF, 2021), showing specimen records
from North America, Europe, Japan and Taiwan,
but not from Southeast Asia. The ITS sequenceof material from Myanmar (KH-MYA16-100)
showed 99.62% identity with that of Japanese
specimen (Table 2), and they can be considered
conspeciﬁc. This is the ﬁrst record of the species
from Myanmar.

62

Kentaro Hosaka et al.
Table 1.

Mushrooms of Myanmar molecularly identiﬁed to species

ID*1

Taxon names

Voucher nos.*2

Accession nos.*3

1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33

Amanita ceciliae (Berk. & Broome) Bas
Craterellus atratus (Corner) Yomyart, Watling, Phosri, Piap. & Sihan.
Dacryopinax spathularia (Schwein.) G.W. Martin
Favolaschia calocera R. Heim
Fistulina hepatica (Schaeﬀ.) With.
Geastrum courtecuissei P.-A. Moreau & C. Lécuru
Laccaria vinaceoavellanea Hongo
Laccaria vinaceoavellanea Hongo
Laccaria vinaceoavellanea Hongo
Lactarius austrotorminosus H.T. Le & Verbeken
Lactiﬂuus piperatus (L.) Roussel
Lactiﬂuus volemus (Fr.) Kuntze
Lentinus sajor-caju (Fr.) Fr.
Lentinus sajor-caju (Fr.) Fr.
Lentinus squarrosulus Mont.
Lentinus squarrosulus Mont.
Lentinus squarrosulus Mont.
Leucocoprinus birnbaumii (Corda) Singer
Leucocoprinus birnbaumii (Corda) Singer
Neonothopanus nambi (Speg.) R.H. Petersen & Krisai
Phallus merulinus (Berk.) Cooke
Pisolithus albus (Cooke & Massee) Priest
Psathyrella candolleana (Fr.) Maire
Psathyrella candolleana (Fr.) Maire
Psathyrella candolleana (Fr.) Maire
Pycnoporus sanguineus (L.) Murrill
Russula virescens (Schaeﬀ.) Fr.
Schizophyllum commune Fr.
Schizophyllum commune Fr.
Schizophyllum commune Fr.
Schizophyllum commune Fr.
Trogia infundibuliformis Berk. & Broome
Turbinellus ﬂoccosus (Schwein.) Earle ex Giachini & Castellano

KH-MYA16-100
KH-MYA16-168
KH-MYA16-051
KH-MYA16-026
KH-MYA16-057
KH-MYA16-040
KH-MYA16-063
KH-MYA16-101
KH-MYA16-106
KH-MYA16-023
KH-MYA16-048
KH-MYA16-117
KH-MYA16-004
KH-MYA17-046
KH-MYA16-003
KH-MYA16-077
KH-MYA16-140
KH-MYA17-043
KH-MYA17-077
KH-MYA17-031
KH-MYA17-064
KH-MYA17-119
KH-MYA16-012
KH-MYA16-085
KH-MYA16-121
KH-MYA16-053
KH-MYA16-045
KH-MYA16-013
KH-MYA16-094
KH-MYA17-038
KH-MYA17-112
KH-MYA17-033
KH-MYA16-017

MW577302
MW577303
MW577304
MW577305
MW577306
MW577307
MW577308
MW577309
MW577310
MW577311
MW577312
MW577313
MW577314
MW577315
MW577316
MW577317
MW577318
MW577319
MW577320
MW577321
MW577322
MW577323
MW577324
MW577325
MW577326
MW577327
MW577328
MW577329
MW577330
MW577331
MW577332
MW577333
MW577334*1 The numbers are corresponding to those of Table 2.
*2 All specimens have been deposited under the same voucher numbers both in Japan (TNS) and Myanmar (Forest
Research Institute).
*3 The GenBank accession numbers of the ITS sequences newly generated for this study are shown.

Craterellus atratus (Corner) Yomyart, Watling,
Phosri, Piap. & Sihan.
Material examined: Myanmar: Mandalay, near
Pyin Oo Lwin (21.954638, 96.384396), alt.
929 m, under Quercus spp., 1 Sept 2016, K.
Hosaka (KH-MYA16-168).
Type locality: Brazil.
Remarks: The species has been reported from
Brazil (Corner, 1966), Guyana (Wilson et al.,2012; GBIF, 2021), and Thailand (Yomyart et al.,2012). The ITS sequence of material from Myanmar (KH-MYA16-168) showed 96.26% identitywith that of Thai specimen (Table 2), and they
can be considered conspeciﬁc. However, due totheir disjunct distribution pattern, the conspeciﬁcity of South American and Southeast Asianmaterials warrants further studies. This is the ﬁrst
record of the species from Myanmar. In addition,
Craterellus was not reported by Thaung (2007),and therefore, it could be the ﬁrst record of thegenus from Myanmar.
Dacryopinax spathularia (Schwein.) G.W.Martin
Material examined: Myanmar: Mandalay, Baw
Reserve Forest No. 85 near Pyin Oo Lwin
(21.908162, 96.541085), alt. 969 m, on hardwood,
29 Aug 2016, K. Hosaka (KH-MYA16-051).
Type locality: United States of America.

Uncultured fungus/Amanita ceciliae
Craterellus atratus
Dacryopinax spathularia
Favolaschia calocera
Fistulina hepatica
Geastrum courtecuissei
Uncultured Laccaria/L. vinaceoavellanea
Laccaria sp./L. vinaceoavellanea
Laccaria sp./L. vinaceoavellanea
Lactarius austrotorminosus
Lactarius piperatus
Lactarius volemus
Lentinus sajor-caju
Lentinus sajor-caju
Lentinus squarrosulus
Lentinus sp./L. squarrosulus
Lentinus squarrosulus
Leucocoprinus birnbaumii
Leucocoprinus birnbaumii
Neonothopanus nambi
Phallus merulinus
Pisolithus albus
Psathyrella candolleana
Psathyrella candolleana
Psathyrella candolleana
Pycnoporus sanguineus
Russula cf. virescens
Schizophyllum commune
Schizophyllum commune
Schizophyllum commune
Schizophyllum commune
Trogia infundibuliformis
Turbinellus ﬂoccosus

1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33

959/959
1122
883
1349
1332
1009
1109/1109
1245/1138
1125/1050
1330
1166
1197
1223
1234
1254
1096/1092
1212
1279
1286
1327
935
1155
1103
1131
1199
1158
1142
1164
1170
1166
1151
1242
1083

Total Score*3
99/99
100
97
98
98
94
100/100
98/91
100/96
100
100
99
100
100
100
100/100
99
100
99
100
100
99
100
100
99
98
98
100
99
100
100
99
97

QueryCover (%)*3
0/0
0
0
0
0
0
0/0
0/0
0/0
0
0
0
0
0
0
0/0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0

E Value*3
99.62/99.62
96.26
98.6
99.59
96.98
99.46
99.83/99.83
99.14/98.9
100/98.98
99.86
97.12
99.85
99.55
99.7
100
100/99.83
99.55
99.15
97.74
99.19
99.61
100
99.18
99.52
99.85
99.84
97.06
100
100
100
99.84
98.72
92.3

Per.Ident (%)*3
AB015694
AB445115
AJ537397
MW462898
KY474052
KF988437
AB777480/AB453023
MN585661/KJ411955
MN585660/KJ411955
EF141533
KF220042
HQ318260
KP283492
KP283492
KT273373
KJ654561/MG719283
JQ868749
MW191767
MH861267
JN571729
MF372141
JQ365190
KT273355
KY940508
KH940508
FJ372672
MT333231
MK647986
MH857808
MN341837
MH857808
MW504969
KJ411951

Accession nos. *4

BLAST results of the ITS sequences generated from materials of Myanmar
Japan
ThailandIndonesia
Indian.s./China
Argentina
ThailandChina/India
China/India
ThailandBelgium
Thailand
Malaysia
Malaysia
Nigeria
Indonesia/India
India
New ZealandJapan
Vietnam
French Guiana
ThailandNigeria
China
China
Thailand
China
South Africa
n.s./India
South Korea
n.s./India
ThailandIndia

Country of origin*5*1 The numbers are corresponding to those of Table 1.
*2 Taxon names of the BLAST top hits, followed by those of the next highest hits with speciﬁc epithet, if applicable, are shown.
*3 Values of the BLAST top hits, followed by those of the next highest hits with species level identiﬁcation, if applicable, are shown.
*4 GenBank nucleotide accession numbers of the BLAST top hits, followed by those of the next highest hits with species level identiﬁcation, if applicable, are shown.
*5 The countries of origin of the BLAST top hits, followed by those of the next highest hits, if applicable.

BLAST top hit*2

ID*1

Table 2.

DNA barcoding of Myanmar mushrooms (1)6364Kentaro Hosaka et al.

Remarks: The species is generally consideredwidespread (Hosaka and Linn, 2021), and the
GBIF records (GBIF, 2021) showed its distribution in all major continents except for Antarctica.
The ITS sequence of material from Myanmar(KH-MYA16-051) showed 98.6% identity withthat of Indonesian specimen (Table 2). Higher
diversity of Dacryopinax and related genera wererecently discovered (Shirouzu et al., 2009) andthe taxonomic status of Myanmar material warrants further studies. The species has previously
been recorded from Thailand (Chandrasrikul etal., 2008) and Myanmar (Thaung, 2007).
Favolaschia calocera R.Heim
Material examined: Myanmar: Mandalay, Pyin
Oo Lwin, National Kandawgyi Garden (21.992546,
96.471205), alt. 1114 m, on hardwood, 28 Aug
2016, K. Hosaka (KH-MYA16-026).
Type locality: Madagascar.
Remarks: Although it has not been proven yet,the species is thought to have originated in Madagascar (McMullan-Fisher et al., 2014), andmaterials from Australia and New Zealand are
generally considered exotic. The GBIF records(GBIF, 2021) showed its distribution in Europe,
Taiwan, Southeast Asia (Laos) and widely in theSouthern Hemisphere. The ITS sequence ofmaterial from Myanmar (KH-MYA16-026)
showed 99.59% identity with that of Indian specimen (Table 2). This is the ﬁrst record of the species from Myanmar.
Fistulina hepatica (Schaeﬀ.) With.
Material examined: Myanmar: Mandalay, Baw
Reserve Forest No. 85 near Pyin Oo Lwin
(21.908162, 96.541085), alt. 969 m, on Quercus
sp., 29 Aug 2016, K. Hosaka (KH-MYA16-057).
Type locality: Germany.
Remarks: The species is generally consideredwidespread (Imazeki and Hongo, 1989; Hosaka
and Linn, 2021), and the GBIF records (GBIF,
2021) showed its worldwide distribution in allcontinents except for Antarctica. Although norecords of the species from Southeast Asia
except for the Philippines are available from the

GBIF (GBIF, 2021), Chandrasrikul et al. (2008)
reported it from Thailand. The ITS sequence ofmaterial from Myanmar (KH-MYA16-057)
showed 96.98% identity with that of Chinese
specimen (Table 2). This is the ﬁrst record of thespecies from Myanmar.
Geastrum courtecuissei P.-A.Moreau & C.Lécuru
Material examined: Myanmar: Mandalay, Pyin
Oo Lwin, National Kandawgyi Garden(21.992546, 96.471205), alt. 1114 m, under bamboo, 28 Aug 2016, K. Hosaka (KH-MYA16-040).
Type locality: Guadeloupe.
Remarks: The species has originally been
described as G. schweinitzii (Hosaka and Linn,2021), but its sequence corresponds to the paratype of G. courtecuissei (Accioly et al., 2019).
The ITS sequence of material from Myanmar(KH-MYA16-040) showed 99.46% identity withthat of Argentinian specimen (Table 2), and they
can be considered conspeciﬁc despite their longgeographical distance from each other. This is
the ﬁrst record of the species from Myanmar andoutside of Central and South America, i.e.,
Argentina and Guadeloupe (Accioly et al., 2019).
Laccaria vinaceoavellanea Hongo
Materials examined: Myanmar: Mandalay, Baw
Reserve Forest No. 85 near Pyin Oo Lwin
(21.908162, 96.541085), alt. 969 m, under Quercus
sp., 29 Aug 2016, K. Hosaka (KH-MYA16-063);
Mandalay, Than Daung Village near Pyin Oo
Lwin (21.851267, 96.374177), alt. 985 m, under
Quercus sp., 31 Aug 2016, K. Hosaka (KHMYA16-101); Mandalay, Than Daung Village
near Pyin Oo Lwin (21.842197, 96.402772), alt.
963 m, under Quercus sp., 31 Aug 2016, K.
Hosaka (KH-MYA16-106).
Type locality: Japan.
Remarks: The species has been reported fromJapan, Korea, New Guinea (Imazeki and Hongo,
1987), China (Wang et al., 2004), and Southeast
Asia (Laos and Malaysia) (GBIF, 2021). The ITS
sequences of materials from Myanmar showed atleast 98.9% identity with those of Chinese,
Indian or Thai materials (Table 2), and they can

DNA barcoding of Myanmar mushrooms (1)be considered conspeciﬁc. This is the ﬁrst recordof the species from Myanmar. In addition, Laccaria was not reported by Thaung (2007), andtherefore, it could be the ﬁrst record of the genus
and the family Hydnangiaceae from Myanmar.
Lactarius austrotorminosus H.T.Le & Verbeken
Material examined: Myanmar: Mandalay, Pyin
Oo Lwin, National Kandawgyi Garden (21.
992546, 96.471205), alt. 1114 m, under broadleaf
trees, 28 Aug 2016, K. Hosaka (KHMYA16-023).
Type locality: Thailand.
Remarks: The species was originally reported
from Thailand (Le et al., 2007; GBIF, 2021), but
it has more recently been discovered from South
Korea (Lee et al., 2018). The ITS sequence ofmaterial from Myanmar (KH-MYA16-023)
showed 99.86% identity with that of Thai specimen (Table 2), and they can be considered conspeciﬁc. The type materials were collected under
Castanopsis (Le et al., 2007), but the collection
locality in Myanmar was dominated by a variety
of broadleaf trees and Pinus spp., and the presence of Castanopsis was not conﬁrmed (Hosaka
et al., 2019).
Lactiﬂuus piperatus (L.) Roussel
Material examined: Myanmar: Mandalay, Baw
Reserve Forest No. 85 near Pyin Oo Lwin
(21.908162, 96.541085), alt. 969 m, under Quercus sp., 29 Aug 2016, K. Hosaka (KHMYA16-048).
Type locality: Sweden [neotypiﬁed by De
Crop et al. (2014)].
Remarks: The species is generally consideredwidespread in temperate regions of the Northern
Hemisphere (Hosaka and Linn, 2021 as “Lactarius piperatus (L.) Pers.”), but there are also somerecords from Australia (Imazeki and Hongo,
1989; GBIF, 2021) and Southeast Asia (Chandrasrikul et al., 2008; GBIF, 2021). The ITS
sequence of material from Myanmar (KHMYA16-048) showed 97.12% identity with thatof Belgium specimen (Table 2). This is the ﬁrst
record of the species from Myanmar.

65

Lactiﬂuus volemus (Fr.) Kuntze
Material examined: Myanmar: Mandalay,
Shwe Kyaung Village near Pyin Oo Lwin
(21.801094, 96.419242), alt. 911 m, under Quercus
sp., 31 Aug 2016, K. Hosaka (KH-MYA16-117).
Type locality: Sweden [epitypiﬁed by Van dePutte et al. (2016)].
Remarks: The species is generally consideredwidespread in temperate regions of the Northern
Hemisphere [Imazeki and Hongo, 1989; Hosaka
and Linn, 2021 as “Lactarius volemus (Fr.) Fr.”],but there are also some scattered records from
Australia (GBIF, 2021). The species has beenreported from Thailand (Chandrasrikul et al.,2008) and China (Wang et al., 2004), and the ITS
sequence of material from Myanmar (KHMYA16-117) showed 99.85% identity with thatof Thai specimen (Table 2). It is the ﬁrst recordof the species from Myanmar.
Lentinus sajor-caju (Fr.) Fr.
Materials examined: Myanmar: Yangon, Kan
Daw Gyi Lake (16.798038, 96.164525), alt.
35 m, on hardwood, 27 Aug 2016, K. Hosaka(KH-MYA16-004); Bo Cho Island (10.673655,
98.246158), alt. 70 m, on hardwood, 19 May
2017, K. Hosaka (KH-MYA17-046).
Type locality: Old tropics but could not be
identiﬁed with certainty.
Remarks: The species has been reported
widely distributed in tropical and subtropicalregions of the world, including Africa, Australia,
and Southeast Asia, but it appears uncommon inSouth America (GBIF, 2021). The ITS sequencesof materials from Myanmar showed at least
99.55% identity with those of Malaysian material(Table 2), and they can be considered conspeciﬁc. The species has previously been recorded
from Thailand (Chandrasrikul et al., 2008) andMyanmar (Thaung, 2007).
Lentinus squarrosulus Mont.
Materials examined: Myanmar: Yangon, Kan
Daw Gyi Lake (16.798038, 96.164525), alt.
35 m, on hardwood, 27 Aug 2016, K. Hosaka(KH-MYA16-003); Mandalay, Baw Reserve For-

66

Kentaro Hosaka et al.

est No. 85 near Pyin Oo Lwin (21.905359,
96.533851), alt. 968 m, on hardwood, 29 Aug
2016, K. Hosaka (KH-MYA16-077); Myanmar:
Mandalay, near Pyin Oo Lwin (21.767522,
96.415126), alt. 884 m, on sclerotia, 31 Aug
2016, K. Hosaka (KH-MYA16-140).
Type locality: Old tropics but could not be
identiﬁed with certainty.
Remarks: The species has been reported
widely mostly from tropical regions of the world(GBIF, 2021). The ITS sequences of materials
from Myanmar showed at least 99.55% identitywith those of Indonesian, Indian and Nigerianmaterials (Table 2), and they can be considered
conspeciﬁc. The species has previously beenrecorded from Thailand (Chandrasrikul et al.,2008) and Myanmar (Thaung, 2007).
Leucocoprinus birnbaumii (Corda) Singer
Materials examined: Myanmar: Bo Cho Island(10.676936, 98.248632), alt. 19 m, in broadleaf
forest, 18 May 2017, K. Hosaka (KH-MYA17043); Lampi Island (10.841444, 98.231806), alt.
22 m, in broadleaf forest, 21 May 2017, N.
Tanaka (KH-MYA17-077).
Type locality: Czech Republic.
Remarks: The species has been recorded fromall continents except for Antarctica (GBIF,
2021), but its main distribution areas are generally considered tropical to subtropical regions(Imazeki and Hongo, 1987; Hosaka and Linn,2021). The ITS sequences of materials from
Myanmar showed 97.74% to 99.15% identitywith those of Japanese and New Zealand materials, respectively (Table 2), The species has beenrecorded from Thailand (Chandrasrikul et al.,2008), but this is the ﬁrst record of the species
from Myanmar.
Neonothopanus nambi (Speg.) R.H.Petersen &KrisaiMaterial examined: Myanmar: Bo Cho Island(10.676936, 98.248632), alt. 19 m, on hardwood,
18 May 2017, K. Hosaka (KH-MYA17-031).
Type locality: Paraguay.
Remarks: The species has been reported

mostly from tropical regions of Africa (Madagascar), Australia, Micronesia, South America,
China and Southeast Asia (Malaysia) (Capelari etal., 2011; GBIF, 2021). No records from Thailand, which is located next to Myanmar andtherefore expected to have similar mycoﬂora,
could be obtained. The ITS sequence of material
from Myanmar (KH-MYA17-031) showed
99.19% identity with that of Vietnamese specimen (Table 2), and they can be considered conspeciﬁc. This species is known as bioluminescent, but the bioluminescent property of
Myanmar materials has not been conﬁrmed yet(Hosaka and Linn, 2021).
Phallus merulinus (Berk.) Cooke
Material examined: Myanmar: Bo Cho Island(10.668087, 98.243671), alt. 43 m, in broadleaf
forest, 19 May 2017, K. Hosaka (KHMYA17-064).
Type locality: Indonesia.
Remarks: The species has been recorded fromSouth America (Cabral et al., 2014), India andSoutheast Asia (Indonesia, Philippines and Singapore) (GBIF, 2021). No records from Thailand,
which is located next to Myanmar and therefore
expected to have similar mycoﬂora, could beobtained. The ITS sequence of material from
Myanmar (KH-MYA17-064) showed 99.61%
identity with that of French Guianan specimen(Table 2), and they can be considered conspeciﬁc
despite their long geographical distance fromeach other. This is the ﬁrst record of the species
from Myanmar.
Pisolithus albus (Cooke & Massee) Priest
Material examined: Myanmar: Moeyungyi Wetland Wildlife Sanctuary (17.589505, 96.575639),
alt. 39 m, under Acacia auriculiformis and A.
mangium, 24 May 2017, K. Hosaka (KHMYA17-119).
Type locality: Australia.
Remarks: The species has been recorded from
Australia, New Zealand, India and Southeast
Asia (GBIF, 2021), but the records outside Australia are the results of the anthropogenic intro-

DNA barcoding of Myanmar mushrooms (1)duction with ectomycorrhizal host trees (Eucalyptus and Acacia) (Martin et al., 2002).
Accordingly, the material of Myanmar, which
was collected under Acacia, is also considered
exotic, but it is the ﬁrst record of the species
from Myanmar. The ITS sequence of material
from Myanmar (KH-MYA17-119) showed 100%
identity with that of Thai specimen (Table 2).
Psathyrella candolleana (Fr.) Maire
Materials examined: Myanmar: Yangon, Kan
Daw Gyi Lake (16.798038, 96.164525), alt.
35 m, in broadleaf forest, 27 Aug 2016, K.
Hosaka (KH-MYA16-012); Mandalay, Wet win
Protected Public Forest near Pyin Oo Lwin
(22.08978, 96.632556), alt. 713 m, in broadleaf
forest, 30 Aug 2016, K. Hosaka (KHMYA16-085); Mandalay, Shwe Kyaung Village
near Pyin Oo Lwin (21.801094, 96.419242), alt.
911 m, in broadleaf forest, 31 Aug 2016, K.
Hosaka (KH-MYA16-121).
Type locality: Sweden [neotypiﬁed by Örstadius et al. (2015)].
Remarks: This species is generally consideredwidespread (Imazeki and Hongo, 1987), but the
GBIF data showed only one record from Southeast Asia (Vietnam) (GBIF, 2021). The ITS
sequences of materials from Myanmar showed atleast 99.18% identity with those of Chinese or
Nigerian materials (Table 2), and they can beconsidered conspeciﬁc. The species has beenreported from Thailand (Chandrasrikul et al.,2008), but this is the ﬁrst record of the species
from Myanmar.
Pycnoporus sanguineus (L.) Murrill
Material examined: Myanmar: Mandalay, Baw
Reserve Forest No. 85 near Pyin Oo Lwin
(21.908162, 96.541085), alt. 969 m, on hardwood,
29 Aug 2016, K. Hosaka (KH-MYA16-053).
Type locality: Suriname.
Remarks: The species has been recorded
widely in tropical to temperate regions of theworld (GBIF, 2021), including Thailand (Chandrasrikul et al., 2008). The ITS sequence ofmaterial from Myanmar (KH-MYA16-053)

67

showed 99.84% identity with that of Thai specimen (Table 2). This is the ﬁrst record of the species from Myanmar.
Russula virescens (Schaeﬀ.) Fr.
Material examined: Myanmar: Mandalay, Baw
Reserve Forest No. 85 near Pyin Oo Lwin
(21.908162, 96.541085), alt. 969 m, under Quercus
sp., 29 Aug 2016, K. Hosaka (KH-MYA16-045).
Type locality: Germany.
Remarks: The species has been recorded
widely in the Northern Hemisphere, includingSoutheast Asia (Imazeki and Hongo, 1989;
Chandrasrikul et al., 2008; GBIF, 2021). The ITS
sequence of material from Myanmar (KHMYA16-045) showed 97.06% identity with thatof Chinese specimen (Table 2). This is the ﬁrst
record of the species from Myanmar.
Schizophyllum commune Fr.
Materials examined: Myanmar: Yangon, Kan
Daw Gyi Lake (16.798038, 96.164525), alt.
35 m, on hardwood, 27 Aug 2016, K. Hosaka(KH-MYA16-013); Mandalay, Wet win Protected
Public Forest near Pyin Oo Lwin (22.08978,
96.632556), alt. 713 m, on hardwood, 30 Aug
2016, K. Hosaka (KH-MYA16-094); Bo Cho
Island (10.676936, 98.248632), alt. 19 m, on
hardwood, 18 May 2017, K. Hosaka (KHMYA17-038); Lampi Island (10.690818,
98.24412), alt. 33 m, on hardwood, 23 May 2017,
K. Hosaka (KH-MYA17-112).
Type locality: Sweden.
Remarks: The species is considered cosmopolitan (Hosaka and Linn, 2021) and the GBIF
records showed its distribution in all continents
including Antarctica (GBIF, 2021). The ITS
sequences of materials from Myanmar showed atleast 99.84% identity with materials from various
countries (Table 2), and they can be considered
conspeciﬁc. The species has previously beenreported from Myanmar (Thaung, 2007).
Trogia infundibuliformis Berk. & Broome
Material examined: Myanmar: Bo Cho Island(10.676936, 98.248632), alt. 19 m, on hardwood,

68

Kentaro Hosaka et al.

18 May 2017, K. Hosaka (KH-MYA17-033).
Type locality: Sri Lanka.
Remarks: The species is probably widely distributed throughout tropical regions of the world,
but the GBIF data only showed scattered records
from Southeast Asia (Laos) and Africa (Democratic Republic of the Congo) (GBIF, 2021). Further literature search indicated that the species isalso distributed in India, Sri Lanka (Kumar andManimohan, 2009) and Thailand (Chandrasrikul
et al., 2008). The ITS sequence of material from
Myanmar showed 98.72% identity with that ofThai specimen (Table 2), and they can be considered conspeciﬁc. This is the ﬁrst record of thespecies from Myanmar.
Turbinellus ﬂoccosus (Schwein.) Earle ex Giachini & Castellano
Material examined: Myanmar: Mandalay, Pyin
Oo Lwin, National Kandawgyi Garden(21.992546, 96.471205), alt. 1114 m, under Pinus
sp., 28 Aug 2016, K. Hosaka (KH-MYA16-017).
Type locality: United States of America.
Remarks: The species has been recorded from
North America, Japan, China (Wang et al., 2004;
GBIF, 2021), India (Khaund and Joshi, 2014),
and Thailand (Chandrasrikul et al., 2008). The
ITS sequence of material from Myanmar showed
only 92.3% identity with that of Indian specimen(Table 2). It is, therefore, likely that the Myanmar material represents a new species to science.
This is the ﬁrst record of the species from Myanmar. In addition, Turbinellus (or Gomphus) was
not reported by Thaung (2007), and therefore, it
could be the ﬁrst record of the genus and thefamily Gomphaceae from Myanmar.
Acknowledgments
The authors thank Kazuo Nishibori, Megumi
Johnston, and Jun Yamazaki for assisting ourcuratorial work. We also thank the park rangers
and oﬃcers in Myanmar for assisting our ﬁeldwork. Thanks also goes to Dr. Tiara Cabral,
INPA, Brazil, for thoroughly reviewing the manuscript. This study was ﬁnanced in part by aresearch project of the National Museum of
Nature and Science entitled as “Biological inventory with special attention to Myanmar: investigations of the origin of southern elements of Japanese ﬂora and fauna” and JSPS KAKENHI
Grant Number 18KK0210.
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