Taiwania 65(4): 548‒558, 2020
DOI: 10.6165/tai.2020.65.548

NOTE

Contribution to the knowledge of the Lichen Mycota of Myanmar (I)twenty species newly recorded from Southern Myanmar
Yoshihito OHMURA1,*, Miyu SUGIMOTO2, Mu Mu AUNG3, Nobuyuki TANAKA1,2
1. Department of Botany, National Museum of Nature and Science, 4-1-1 Amakubo, Tsukuba, Ibaraki, 305-0005, Japan.
2. Graduate School of Agriculture, Ibaraki University, 3-21-1 Chuo, Ami, Inashiki, 300-0393, Japan.
3. Forest Research Institute, Forest Department, Ministry of Natural Resources and Environmental Conservation, Yezin, Nay Pyi
Taw, Myanmar.
*Corresponding author’s email: ohmura-y@kahaku.go.jp(Manuscript received 12 June 2020; Accepted 30 August 2020; Online published 9 November 2020)
ABSTRACT: Specimens of lichens collected from southern Myanmar including Tanintharyi and Yangon Regions were examined.
As the result of taxonomic examinations, the following 20 species were identified: Astrothelium macrocarpum, Bulbothrix
subscortea, Coccocarpia erythroxyli, C. palmicola, Cruentotrema thailandicum, Dirinaria aegialita, D. consimilis, Dyplolabia
afzelii, Flakea papillata, Glyphis cicatricosa, Graphis cf. caesiella, G. desquamescens, G. supracola, Malmidea bakeri, Physcia
undulata, Pyrenula mamillana, Pyxine dactyloschmidtii, Sarcographa labyrinthica, Trypethelium eluteriae, and Zwackhia prosodea.
All of them, except the two species of Coccocarpia, are new records for Myanmar. The ITS rDNA sequences were successfully
obtained from 17 samples for 14 species. The BLAST identities for Myanmar collections with the same species in GenBank range
from 89 to 100%.
KEY WORDS: BLAST, distribution, inventory, ITS rDNA, lichenized fungi, Southeast Asia, taxonomy.

INTRODUCTION
Myanmar is considered to hold important andirreplaceable biodiversity that is part of the Indo-Burma
Biodiversity Hotspot. Habitats range from the mangrove
forests and coral reefs of the Andaman Islands in the south
to the snow-capped peaks of Mt. Kakaboradzi (5,881 m)in the north, the highest mountain in Southeast Asia
(Mittermeier et al., 2004; Tanaka et al., 2018). Despitemuch attention paid to the investigation and conservationof plant and animal biodiversity (Tordoff et al., 2011), thelichen mycota has been poorly studied so far. Only 10
species and one variety of lichens were reported before thepresent study: i.e., Bulbothrix cf. tabacina (Mont. &Bosch) Hale, Coccocarpia erythroxyli (Spreng.)
Swinscow & Krog, C. palmicola (Spreng.) Arv. & D.J.
Galloway, Dirinaria picta (Sw.) Clem. & Shear,Gyronactis asiatica Ertz & Tehler, Leptogium
trichophorum Müll. Arg., L. trichophorum var.
fuliginosum Müll. Arg., Melaspilea heterocarpa (Fée)
Müll. Arg., Parmotrema sulphuratum (Nees & Flot.) Hale,
Pyrenula laevigata (Pers.) Arnold, and Rhizocarpon
subalboatrum (Nyl.) Müll. Arg. (Nylander and Crombie,
1883; Müller, 1889; Arvidsson, 1982; Ertz et al., 2015).
As part of the international cooperative project"Biological Inventory with special attention to
Myanmar" organized by the National Museum of Natureand Science, Japan and the Forest Department, Ministryof Natural Resources and Environmental Conservation,
Myanmar, the authors have since 2017 studied the lichen
548

mycota of Myanmar.
This paper contributes to the knowledge of the lichen
mycota of Myanmar. ITS rDNA sequences weregenerated from DNA extracted from identified specimens.

MATERIALS AND METHODS
Field surveys in Tanintharyi and Yangon Regions in
Myanmar were carried out from 16 to 25 January 2017.
A total of 159 specimens was collected. Twenty-fivespecimens are identified to species level and are reported
in this study, while the remaining specimens are still
under examination. All specimens are housed in theherbarium of the National Museum of Nature andScience, Japan (TNS), with some duplicates in theherbaria of the Forest Research Institute, Myanmar(RAF) and National Taiwan University, Taiwan (TAI).
Morphological observations were made using a
dissecting microscope (Olympus SZX16) and adifferential interference contrast microscope (Olympus
BX51). Anatomical examinations were made on hand cut
sections mounted in water. Ascospore measurements are
given as (minimum–) range including mean ± standard
deviation (–maximum) (n = number of measurements).
Chemical substances of lichens were examined using
thin layer chromatography (TLC) (Culberson and Johnson,1982). Solvent system B′ (hexane: methyl tert-butyl ether:
formic acid, 140: 72: 18) was used for all TLC analyses.
DNA extraction followed a modified CTAB protocol(Hosaka, 2009).

2020

Ohmura et al.: Lichens from southern Myanmar

Table 1. Identified taxa from the southern part of Myanmar and the BLAST results of the ITS rDNA sequence
Taxa

Voucher

Astrothelium
macrocarpum

Y. Ohmura 12285

Y. Ohmura 12392

Bulbothrix
subscortea

Y. Ohmura 12397

Y. Ohmura 12398

Coccocarpia
erythroxyli

Y. Ohmura 12336

Coccocarpia
palmicola

Y. Ohmura 12408

Cruentotrema
thailandicum
Dirinaria aegialita

Y. Ohmura 12327
Y. Ohmura 12281

Dirinaria consimilis Y. Ohmura 12396

Dyplolabia afzelii

Y. Ohmura 12324
Y. Ohmura 12411

GenBank
BLAST results with GenBank sequences. Identity; score; e-value
accession number (accession number, species) of the top 3 sequences (maximum) offor ITS rDNA
related taxa are shown.
LC573990
527/533 (99%), 948, 0.0 (LC127382, Astrothelium macrocarpum)
527/533 (99%), 948, 0.0 (LC127383, Astrothelium macrocarpum)
520/525 (99%), 941, 0.0 (LC127384, Astrothelium macrocarpum)
LC573991
783/793 (99%), 1406, 0.0 (LC127382, Astrothelium macrocarpum)
783/793 (99%), 1406, 0.0 (LC127383, Astrothelium macrocarpum)
786/800 (98%), 1397, 0.0 (AB759880, Astrothelium macrocarpum)
LC573992
527/527 (100%), 974, 0.0 (KM249896, Bulbothrix subscortea)
526/526 (100%), 972, 0.0 (KM249904, Bulbothrix subscortea)
527/528 (99%), 968, 0.0 (KM249907, Bulbothrix subscortea)
LC573993
526/527 (99%), 968, 0.0 (KM249896, Bulbothrix subscortea)
525/526 (99%), 966, 0.0 (KM249904, Bulbothrix subscortea)
526/528 (99%), 963, 0.0 (KM249907, Bulbothrix subscortea)
LC573994
726/744 (98%), 1267, 0.0 (KT947056, Coccocarpia erythroxyli)
724/744 (97%), 1256, 0.0 (KT947055, Coccocarpia erythroxyli)
718/743 (97%), 1218, 0.0 (KT947060, Coccocarpia palmicola)
LC573995
724/725 (99%), 1334, 0.0 (KT947060, Coccocarpia palmicola)
709/743 (95%), 1162, 0.0 (KT947056, Coccocarpia erythroxyli)
707/743 (95%), 1151, 0.0 (KT947055, Coccocarpia erythroxyli)
LC573996
483/545 (89%), 652, 0.0 (KP012902, Dyplolabia afzelii)
366/428 (86%), 425, 2e-114 (KR017123, Fissurina insidiosa)LC573997
532/533 (99%), 979, 0.0 (MK028196, Dirinaria aegialita)
531/533 (99%), 972, 0.0 (MK028169, Dirinaria aegialita)
531/533 (99%), 972, 0.0 (MK028168, Dirinaria aegialita)
LC573998
621/628 (99%), 1118, 0.0 (AB764068, Dirinaria aegialita)
552/560 (99%), 987, 0.0 (LC516424, Dirinaria consimilis)
532/538 (99%), 959, 0.0 (MK028174, Dirinaria picta)
N/A
N/A
LC573999
530/546 (97%), 911, 0.0 (KP012902, Dyplolabia afzelii)
334/375 (89%), 484, 2e-132 (MK503256, Dyplolabia afzelii)
LC574000
726/815 (89%), 994, 0.0 (KT232210, Flakea papillata)
N/A
N/A
N/A
N/A

Flakea papillata
Y. Ohmura 12329
Glyphis cicatricosa Y. Ohmura 12294
Graphis cf.
Y. Ohmura 12268
caesiellaGraphisY. Ohmura 12337
N/A
desquamescens
Graphis supracola Y. Ohmura 12268 pr.p. N/A
Malmidea bakeri Y. Ohmura 12366
LC574001
Y. Ohmura 12404
LC574002
Physcia undulata Y. Ohmura 12300
LC574003

N/A

Y. Ohmura 12413

LC574004

PyxineY. Ohmura 12276
dactyloschmidtii

LC574005

Sarcographalabyrinthica

Y. Ohmura 12399

LC574006

Y. Ohmura 12400
Y. Ohmura 12270

N/A
N/A

N/A
606/624 (97%), 1051, 0.0 (MK499341, Malmidea subaurigera)
605/623 (97%), 1050, 0.0 (MK499341, Malmidea subaurigera)
507/507 (100%), 937, 0.0 (MK028207, Physcia undulata)
507/507 (100%), 937, 0.0 (MK028206, Physcia undulata)
898/916 (98%), 1589, 0.0 (KT820154, Pyrenula massariospora)
845/925 (91%), 1245, 0.0 (KT820150, Pyrenula mamillana)825/908 (91%), 1201, 0.0 (KT820149, Pyrenula mamillana)
488/489 (99%), 898, 0.0 (MK028211, Pyxine retirugella)
487/489 (99%), 891, 0.0 (MK028217, Pyxine retirugella)
487/489 (99%), 891, 0.0 (MK028216, Pyxine retirugella)
410/488 (84%), 444, 3e-120 (KC592266, Leiorreuma sericeum)
338/389 (87%), 420, 5e-113 (KC592265, Leiorreuma sericeum)
329/381 (86%), 394, 3e-105 (NR_120278, Halegrapha chimaera)
N/A
N/A

Y. Ohmura 12388

N/A

N/A

Pyrenula
mamillana

Trypethelium
eluteriae
Zwackhia
prosodea549

Taiwania

For DNA amplification, 10 μL of PCR mix contained
1 μL genomic DNA extraction, 0.25 μL of each primer(10 pmol/μL) and 5 μL Emerald-Amp PCR Master Mix(TaKaRa Bio Inc.). PCR amplification of the ITS rDNA
region (including ITS1, 5.8S rDNA and ITS2) was
performed using the primer set of ITS1F (Gardes andBruns, 1993) as the 5′ primer and LR1 (Vilgalys and
Hester, 1990) as the 3′ primer. PCR cycling conditions
were 94°C (3 min), followed by 11 cycles of 95°C (30sec), 62°C to 52°C (30 sec) with annealing temperatures
lowered by 1°C between cycles, and 72°C (1 min),followed by 30 cycles at 52°C annealing temperature anda final extension at 72°C (7 min). Sequencing was doneon an ABI Prism 3130x genetic analyzer (Applied
Biosystems) using the BigDye Terminator ver. 3.1 Cycle
Sequencing Kit according to the manufacturer’s
instructions.
The obtained sequences were analyzed using the
GenBank BLAST search (blastn). Default settings forthe blastn option were used.
Voucher information and GenBank accession
numbers for the obtained ITS rDNA sequences areshown in Table 1.

RESULTS AND DISCUSSION
Astrothelium macrocarpum (Fée) Aptroot & Lücking,Lichenologist 48: 867. 2016.
Figs. 1A & 1B
For a detailed description and synonyms, see Aptrootand Lücking (2016). Ascospore size for the Myanmar
collections is (15.7–)19.1–21.9(–22.9) × (5.4–)7.0–8.2(–9.1) µm (n =53). Perispore is 2.1–6.5 μm thick.
Chemistry: Parietin.
Remarks: Ascospore size of the Myanmar collection
is slightly smaller than the reported size (21–28 × 7–11μm) in Aptroot and Lücking (2016). Because theirranges partially overlap, this study treats the difference
as intraspecific variation.
Astrothelium macrocarpum resembles Trypethelium
eluteriae Spreng. in having an olive-green to yellow
crustose thallus and yellow pseudostromata. However, it
is readily distinguished from the latter species by
erumpent pseudostromata with flattened top and 3septate ascospores.
In addition to the morphological and chemical
conformity, the BLAST results show high identities (98–99%) with A. macrocarpum sequences in GenBank(Table 1).
Distribution: Pantropical. Asia (Borneo, India,
Singapore, Sri Lanka, and Thailand), Oceania (Australia
and Papua New Guinea), North and South Americas(Brazil, Colombia, Costa Rica, Cuba, French Guiana,
Guyana, Panama, USA, and Venezuela), and Africa(Gabon and South Africa) (Aptroot and Lücking, 2016).
New to Myanmar.
550Vol. 65, No. 4

Specimens examined: MYANMAR. Tanintharyi Region:
Pha Chaung Inn Beach, Kanbauk Village, Yephyu Township,
N14°30′37″, E97°56′44″, on bark of Fabaceae, elev. 0 m, 18 January
2017, Y. Ohmura 12285 (TNS, RAF, TAI); Migyanughlaung,
Tanintharyi Nature Reserve, Yephyu Township, N14°40′18″,
E98°08′16″, on bark of broad-leaf tree, elev. 35 m, 23 January 2017, Y.
Ohmura 12392 (TNS, RAF, TAI).
Bulbothrix subscortea (Asahina) Marcelli & Benatti,
Mycosphere 3: 48. 2012.
Fig. 1C
For detailed descriptions, see Benatti (2012) andZhang et al. (2014).
Chemistry: Atranorin and salazinic acid.
Remarks: In addition to the morphological andchemical conformity, the BLAST results show highidentities (99–100%) with B. subscortea sequences in
GenBank (Table 1).
Distribution: Tropical to subtropical regions of Asia
(Cambodia, China, India, Japan, Nepal, and Taiwan)(Zhang et al., 2014). New to Myanmar.
Specimens examined: MYANMAR. Tanintharyi Region:
Migyanughlaung, Tanintharyi Nature Reserve, Yephyu Township,
N14°40′18″, E98°08′16″, on bark of Areca catechu, elev. 35 m, 23
January 2017, Y. Ohmura 12397 (TNS, RAF, TAI), Y. Ohmura 12398(TNS, RAF).

Coccocarpia erythroxyli (Spreng.) Swinscow & Krog,
Norweg. J. Bot. 23: 256. 1976.
Fig. 1DFor a detailed description and synonyms, see
Arvidsson (1982).
Chemistry: No lichen substance was detected by TLC.
Remarks: In addition to the morphological andchemical conformity, the BLAST results show highidentity (98%) with C. erythroxyli (the first hit) sequencein GenBank (Table 1).
Distribution: Tropical to temperate regions of Asia,
Oceania, North and South Americas, Europe, and Africa(Arvidsson, 1982).
Specimen examined: MYANMAR. Tanintharyi Region:
Kanbauk Village, Yephyu Township, N14°35′14″, E98°03′06″, onbark of broad-leaf tree, elev. 25 m, 20 January 2017, Y. Ohmura 12336(TNS, RAF).

Coccocarpia palmicola (Spreng.) Arv. & D.J. Galloway,
Bot. Not. 132: 242. 1979.
Fig. 1E
For a detailed description and synonyms, see
Arvidsson (1982).
Chemistry: No lichen substance was detected by TLC.
Remarks: In addition to the morphological andchemical conformity, the BLAST results show highidentity (99%) with C. palmicola sequence in GenBank(Table 1).
Distribution: Tropical to warm-temperate regions ofAsia, Oceania, North and South Americas, and Africa(Arvidsson, 1982).

2020

Ohmura et al.: Lichens from southern Myanmar

Specimen examined: MYANMAR. Tanintharyi Region:
Migyanughlaung, Tanintharyi Nature Reserve, Yephyu Township,
N14°40′18″, E98°08′16″, on bark of Areca catechu, elev. 35 m, 23
January 2017, Y. Ohmura 12408 (TNS, RAF).
Cruentotrema thailandicum Rivas Plata, Papong &Lumbsch, Fungal Divers. 52: 119. 2012.
Figs. 1F & 1G
For a detailed description, see Rivas Plata et al.
(2012). Ascospore size for the Myanmar collections is(15.1–)17.0–20.6(–23.6) × (5.9–)6.5–7.5(–8.1) μm(n=39). Perispore is 1.4–2.6 μm thick.
Chemistry: Red pigments (Rf classes 1, 2–3).
Remarks: This species resembles C. lirelliforme J.
Kalb, Polyiam & K. Kalb that has lirelliform ascomata,while C. thailandicum has angular-rounded ascomata(Kalb et al., 2016). The specimen from Myanmar has
both lirelliform and angular-rounded ascomata.
However, the ascospore size of the specimen from
Myanmar agrees with C. thailandicum (15–25 × 7–10μm), while C. lirelliforme has longer ascospores (20–30
× 7–10 μm) (Rivas Plata et al., 2012; Kalb et al., 2016).
Since both species were described on few specimens
collected at the respective type localities, further
taxonomic research including additional collections isneeded to examine their relationship.
The BLAST results show less than 89% identity with
Graphidaceae sequences in GenBank although an evalue with Dyplolabia afzelii shows 0.0 (Table 1). The
low homology results were due to the fact that ITS rDNA
sequences of Cruentotrema species are not registered in
GenBank.
Distribution: Asia (Thailand) (Rivas Plata et al.,2012). New to Myanmar.
Specimen examined: MYANMAR. Tanintharyi Region:
Padauk-kon Chaung, Thet Kae Kawt Compartment, Tanintharyi
Nature Reserve, Yephyu Township, N14°23′19″, E98°11′05″, on barkof broad-leaf tree, elev. 40 m, 19 January 2017, Y. Ohmura 12327(TNS, RAF).

Dirinaria aegialita (Afzel. ex Ach.) B.J. Moore,
Bryologist 71: 248. 1968.
Fig. 1H
For a detailed description and synonymy, see Elix(2009).
Chemistry: Atranorin, divaricatic acid and terpenes.
Remarks: In addition to the morphological andchemical conformity, the BLAST results show highidentity (99%) with D. aegialita sequences in GenBank(Table 1).
Distribution: Tropical to warm-temperate regions ofAsia, Oceania, North and South Americas, and Africa(Elix, 2009). New to Myanmar.
Specimen examined: MYANMAR. Tanintharyi Region:
Pha Chaung Inn Beach, Kanbauk Village, Yephyu Township,
N14°30′37″, E97°56′44″, on bark of Fabaceae, elev. 0 m, 18 January
2017, Y. Ohmura 12281 (TNS, RAF, TAI).

Dirinaria consimilis (Stirt.) D.D. Awasthi, J. Indian Bot.
Soc. 49: 135. 1970.
Fig. 1I
For a detailed description, see Elix (2009). For
synonymy, see Kashiwadani (1979).
Chemistry: Atranorin, sekikaic acid and terpenes.
Remarks: The BLAST results show 99% identitywith D. aegialita (the first hit), 99% with D. consimilis(the second hit) and 99% with D. picta (the third hit)sequences in GenBank (Table 1). The presence of
sekikaic acid in the Myanmar collection confirmed theidentification as D. consimilis. The other two species
contain divaricatic acid.
Distribution: Tropical to warm-temperate regions ofAsia, Oceania and Africa (Elix, 2009). New to Myanmar.
Specimen examined: MYANMAR. Tanintharyi Region:
Migyanughlaung, Tanintharyi Nature Reserve, Yephyu Township,
N14°40′18″, E98°08′16″, on bark of Areca catechu, elev. 35 m, 23
January 2017, Y. Ohmura 12396 (TNS, RAF).

Dyplolabia afzelii (Ach.) A. Massal., Neagen. Lich.: 6.
1854.
Figs. 1J & 1K
For a detailed description, see Archer (2006).
Ascospore size for the Myanmar collections is (14.9–)15.6–
18.0(–20.2) × (6.1–)6.9–8.3(–9.2) μm (n=63).
Chemistry: Lecanoric acid.
Remarks: In addition to the morphological andchemical conformity, the BLAST result shows highidentity (97%, the first hit) with D. afzelii sequence in
GenBank (Table 1). The low identity (89%) with thesecond sequence of this species is due to the many
ambiguous sites and short read of the sequence(MK503256).
Distribution: Pantropical. Asia (Cambodia, China,
India, Indonesia, Malaysia, the Philippines, Singapore,
Sri Lanka, Thailand, and Vietnam), Oceania (Australia
and Papua New Guinea), North and South Americas(Bermuda, Bolivia, Brazil, Colombia, Cuba, Dominica,
French Guiana, Guyana, Mexico, Peru, Suriname,
U.S.A., and Venezuela), and Africa (Cameroon and
Kenya) (Staiger, 2002; Archer, 2006; Singh and Singh,
2010; Moon et al., 2015; Buaruang et al., 2017). New to
Myanmar.
Specimens examined: MYANMAR. Tanintharyi Region:
Padauk-kon Chaung, Thet Kae Kawt Compartment, Tanintharyi Nature
Reserve, Yephyu Township, N14°23′19″, E98°11′05″, on bark of Ficus
elastica, elev. 40 m, 19 January 2017, Y. Ohmura 12324 (TNS, RAF,
TAI); Migyanughlaung, Tanintharyi Nature Reserve, Yephyu Township,
N14°41′09″, E98°08′59″, on bark of Anacardium occidentale, elev. 60 m,
23 January 2017, Y. Ohmura 12411 (TNS, RAF).

Flakea papillata O.E. Erikss., Syst. Ascom. 11: 14. 1992.
Fig. 1L
For a detailed description, see Perlmutter (2006).
Chemistry: No lichen substance was detected by
TLC. Thor and Kashiwadani (1996) reported zeorin andtwo other triterpenoids from Japanese collections.
551

Taiwania

Vol. 65, No. 4

Fig. 1. Lichens collected from southern Myanmar. A–B. Astrothelium macrocarpum (Y. Ohmura 12285, TNS) (A: ascomata; B:
ascospores). C. Bulbothrix subscortea (Y. Ohmura 12397, TNS). D. Coccocarpia erythroxyli (Y. Ohmura 12336, TNS). E.
Coccocarpia palmicola (Y. Ohmura 12408, TNS). F–G. Cruentotrema thailandicum (Y. Ohmura 12327, TNS) (F: ascomata; G:
ascospores). H. Dirinaia aegialita (Y. Ohmura 12281, TNS). I. D. consimilis (Y. Ohmura 12396, TNS). J–K. Dyplolabia afzelii (Y.
Ohmura 12411, TNS) (J: ascomata; K: ascospores). L. Flakea papillata (Y. Ohmura 12329, TNS). M. Glyphis cicatricosa (Y. Ohmura
12294, TNS). N–P. Graphis cf. caesiella (Y. Ohmura 12268, TNS) (N: ascomata; O: vertical section of ascoma; P: ascospores).
Scales: A, L, M, N = 0.5 mm; B, G, K, P =10 μm; C, E, F, J = 1 mm; D, H, I = 2 mm; O = 0.1 mm.

Remarks: This species is known as a strictly steriletaxon characterized by flake-like thalli of narrowly
arranged, irregularly wrinkled, small bluish-green lobes
that are not or only little incised. Molecular phylogenetic
analysis confirmed it to belong to Verrucariaceae
552(Muggia et al., 2009). The BLAST result shows only
89% identity with F. papillata (KT232210, U.S.A.
sample) sequence in GenBank (Table 1). The low
identity of the BLAST result indicates that F. papillata
is a heterogeneous sterile taxon with low morphological2020

Ohmura et al.: Lichens from southern Myanmar

differentiation. The differences observed in thallus
chemistry (Thor and Kashiwadani, 1996) might be apossible taxonomic feature to distinguish taxa within F.
papillata s. lat. Further study of this taxon is needed withrespect to morphological, chemical, and molecularphylogenetic data.
Distribution: Pantropical. Asia (Japan and Thailand),
Oceania (Australia and Fiji), North and South Americas(Argentina, Brazil, Cuba, Paraguay, Peru, and U.S.A.),
and Africa (Kenya) (Thor and Kashiwadani, 1996;
Perlmutter, 2006; Cáceres, 2007; Buaruang et al., 2017).
New to Myanmar.
Specimen examined: MYANMAR. Tanintharyi Region:
Padauk-kon Chaung, Thet Kae Kawt Compartment, Tanintharyi
Nature Reserve, Yephyu Township, N14°23′19″, E98°11′05″, on rock,elev. 40 m, 19 January 2017, Y. Ohmura 12329 (TNS, RAF, TAI).

Glyphis cicatricosa Ach., Syn. Meth. Lich.: 107. 1814.
Fig. 1M
For a detailed description and synonyms, see Archer(2009). Because the collected material was very poor,
anatomical measurements could not be performed.
Distribution: Tropical to warm-temperate regions ofAsia, Oceania, and North and South Americas (Archer,2009). New to Myanmar.
Specimen examined: MYANMAR. Tanintharyi Region:
Forest Department Office of Tanintharyi Region, Dawei (Tavoy),
N14°04′28″, E98°11′44″, on bark of tree branch, elev. 15 m, 17January 2017, Y. Ohmura 12294 (TNS).

Graphis cf. caesiella Vain., Acta Soc. Fauna Fl. Fenn.
7(2): 122. 1890.
Figs. 1N–P
For a detailed description and synonymy, see Archer(2009). Ascospore size for the Myanmar collection is(19.9–)22.8–30.8(–35.0) × (6.0–)6.4–7.2(–7.5) μm (n=19).
Perispore is 1.3–3.8 μm thick. The number of septa is 6–9.
Chemistry: Norstictic acid (major), salazinic acid(minor), and protocetraric acid (trace).
Remarks: According to Lücking (2009), the Graphis
caesiella aggregate consists of at least four species withsimilar morphology but different chemistry: G. bakeri(salazinic), G. caesiella (norstictic), G. dendrogramma(stictic), and G. supracola (protocetraric). The
morphology of the Myanmar collection agrees with G.
caesiella, but the chemistry does not fit to the current
circumscription of the species (it should contain only
norstictic acid). The Myanmar collection could be a new
chemotype for G. caesiella or a new species but further
study is needed to clarify the taxonomic position in
relation with these four taxa based on morphological,
chemical and molecular phylogenetical analyses.
Because of this taxonomic problem, this study treats the
Myanmar collection as G. cf. caesiella. See also remarksof G. supracola below.
Distribution: For G. caesiella, tropical to subtropicalregions of Asia (Japan and Thailand), Oceania (Australia,

Norfolk Island, Lord Howe Island, and Vanuatu), Northand South Americas (Brazil and Costa Rica), and Africa(the Seychelles) (Archer, 2009; Buaruang et al., 2017;
Ohmura and Kashiwadani, 2018). New to Myanmar.
Specimen examined: MYANMAR. Yangon Region:
Bogyoke Park, Kandawgyi Lake, N16°47′47″, E96°09′56″, on bark of
Azadirachta indica, elev. 20 m, 16 January 2017, Y. Ohmura 12268(TNS, RAF).

Graphis desquamescens (Fée) Zahlbr., Denkschr.
Kaiserl. Akad. Wiss. Wien, Math.-Naturwiss. Kl. 83:
108. 1909.
Figs. 2A–D
For a detailed description, see Archer (2009).
Ascospore size for the Myanmar collection is(19.3–)24.2–36.0(–45.3) × (4.6–)6.1–8.3(–9.6) μm(n=31). The number of septa is 7–9.
Chemistry: Norstictic acid.
Distribution: Tropical to warm-temperate regions ofAsia (China, Japan and Taiwan), Oceania (Australia and
New Zealand), and North and South Americas (Brazil,
Mexico and U.S.A.) (Nakanishi, 1966; Archer, 2009).
New to Myanmar.
Specimen examined: MYANMAR. Tanintharyi Region:
Kanbauk Village, Yephyu Township, N14°35′14″, E98°03′06″, onbark of broad-leaf tree, elev. 25 m, 20 January 2017, Y. Ohmura 12337(TNS, RAF).

Graphis supracola A.W. Archer, Aust. Syst. Bot. 14:267. 2001.
Fig. 2E–G
For a detailed description, see Archer (2009).
Ascospore size for the Myanmar collection is(23.1–)24.6–35.8(–42.8) × (6.5–)6.8–7.8(–8.5) μm(n=20). Perispore is 1.5–3.1 μm thick. The number of
septa is 7–9(–10).
Chemistry: Protocetraric acid.
Remarks: This collection was found after TLC
examinations of several spots on the specimen of G. cf.
caesiella. These species grew side by side and theboarder of colonies is not distinctly recognized. See also
remarks of G. cf. caesiella above.
Distribution: Pantropical (Lücking et al., 2009).
Asia (Cambodia and Thailand), Oceania (Australia),
North and South Americas (Brazil, Columbia and USA),
and Africa (Cameroon) (Archer, 2012; Joshi et al., 2016).
New to Myanmar.
Specimen examined: MYANMAR. Yangon Region: Bogyoke
Park, Kandawgyi Lake, N16°47′47″, E96°09′56″, on bark of Azadirachta
indica, elev. 20 m, 16 January 2017, Y. Ohmura 12268 pr.p. (TNS).

Malmidea bakeri (Vain.) Kalb, Rivas Plata & Lumbsch,
Biblioth. Lichenol. 106: 154. 2011.
Figs. 2H & 2I
For a detailed description, see Kalb et al. (2011).
Ascospore size for the Myanmar collections is (10.2–)11.1–
14.1(–16.0) × (4.3–)5.3–7.1(–8.4) μm (n=30).
Chemistry: Atranorin.
553

Taiwania

Vol. 65, No. 4

Fig. 2. Lichens collected from southern Myanmar. A–D. Graphis desquamescens (Y. Ohmura 12337, TNS) (A: ascomata; B: verticalsection of ascoma; C: inspersed hymenium; D: ascospore). E–G. Graphis supracola (Y. Ohmura 12268 pr.p., TNS). H–I. Malmidea
bakeri [H: ascomata and thallus (Y. Ohmura 12404, TNS); I. ascospores (Y. Ohmura 12366, TNS)]. J. Physcia undulata (Y. Ohmura
12300, TNS). K–L. Pyrenula mamillana (Y. Ohmura 12413, TNS) (K: ascomata; L: ascospores in ascus). M. Pyxine
dactyloschmidtii (Y. Ohmura 12276, TNS). Scales: A, E, H = 0.5 mm; B, F = 0.1 mm; C, D, G, I, L =10 μm; J = 2 mm; K, M = 1 mm.

554

2020

Ohmura et al.: Lichens from southern Myanmar

Remarks: This species is known to be close to M.
subaurigera (Vain.) Kalb, Rivas Plata & Lumbsch, but
the latter species has larger ascospores (17–21 × 9–13μm) and an orange-yellow medulla (Kalb et al., 2011).
The BLAST results show 97% identity with M.
subaurigera sequence in GenBank (Table 1). ITS rDNA
sequences of M. bakeri were not registered in GenBank
before this study.
Distribution: Tropical to subtropical regions of Asia
(India, the Philippines and Thailand) (Kalb et al., 2011;
Gupta and Sinha, 2016).
Specimens examined: MYANMAR. Tanintharyi Region:
Tanintharyi Nature Reserve, Yephyu Township, N14°44′18″,
E98°13′42″, on bark of broad-leaf tree, elev. 160 m, 21 January 2017, Y.
Ohmura 12366 (TNS, RAF); Migyanughlaung, Tanintharyi Nature
Reserve, Yephyu Township, N14°40′18″, E98°08′16″, on bark of Areca
catechu, elev. 35 m, 23 January 2017, Y. Ohmura 12404 (TNS, RAF).

Physcia undulata Moberg, Nordic J. Bot. 6: 861. 1986.
Fig. 2J
For a detailed description, see Moberg (2001).
Chemistry: Atranorin, zeorin and other terpenes.
Remarks: In addition to the morphological andchemical conformity, the BLAST results show highidentity (100%) with P. undulata sequences in GenBank(Table 1).
Distribution: Pantropical. Asia (India and Thailand),
Oceania (Australia and New Zealand), North and SouthAmericas (Argentina, Brazil, Chile, Colombia, Costa
Rica, and Ecuador), and Africa (Kenya and South Africa)(Moberg, 1990, 2001; Singh and Sinha, 2010; Buaruang
et al., 2017). New to Myanmar.
Specimen examined: MYANMAR. Tanintharyi Region:
Kanbauk Village, Yephyu Township, N14°35′32″, E98°03′28″, onbark of Moringa oleifera, elev. 35 m, 19 January 2017, Y. Ohmura
12300 (TNS, RAF, TAI).

Pyrenula mamillana (Ach.) Trevis., Consp. Verruc.: 13.
1860.
Figs. 2K & 2L
For detailed descriptions and synonyms, see Aptroot(2009, 2012). Ascospore size for the Myanmar
collection is (14.9–)16.0–18.8(–19.6) × (5.4–)6.1–7.1(–
7.5) μm (n=40). Perispore is 0.8–1.0 μm thick.
Chemistry: No lichen substance was detected by TLC.
Remarks: This species resembles Pyrenula
massariospora (Starbäck) R.C. Harris, but the ascospore
size is smaller (mostly <21 μm) than in the latter (mostly>20 μm) (Aptroot, 2012).
The BLAST results show 98% identity with P.
massariospora (the first hits) and 91% identity with P.
mamillana (the second and third hits) sequences in
GenBank but the e-values are 0.0 in all cases (Table 1).
Morphological features especially ascospore size of the
Myanmar collection are consistent with the descriptionof P. mamillana.
Distribution: Pantropical. Asia (India, Japan, thePhilippines, Sri Lanka, and Thailand), Oceania(Australia and Papua New Guinea), and South America(Brazil, Colombia, Costa Rica, and Guyanas) (Aptroot,
2009; Kashiwadani et al., 2009; Singh and Sinha, 2010;
Flakus et al., 2013; Buaruang et al., 2017). New to
Myanmar.
Specimen examined: MYANMAR. Tanintharyi Region:
Migyanughlaung, Tanintharyi Nature Reserve, Yephyu Township,
N14°41′09″, E98°08′59″, on bark of Mangifera indica, elev. 60 m, 23
January 2017, Y. Ohmura 12413 (TNS, RAF).

Pyxine dactyloschmidtii
Phytotaxa 59: 44. 2012.

Kalb

&

Mongkolsuk,

Fig. 2M
For a detailed description, see Mongkolsuk et al.
(2012).
Chemistry: Atranorin, testacein and terpenes (Rfclasses 4–5, 5, 5–6, 6).
Remarks: This species is morphologically similar to
P. retirugella Nyl. and P. boonpragobiana Kalb &Mongkolsuk. However, P. dactyloschmidtii is easily
distinguished from these species by the absence ofnorstictic acid (Mongkolsuk et al., 2012).
The BLAST results show high identity (99%) with P.
retirugella sequences in GenBank (Table 1). However,the ITS rDNA sequences for P. dactyloschmidtii were
not registered in GenBank before this study. Taxonomic
revision with regard to the presence or absence ofnorstictic acid is needed for P. retirugella and P.
dactyloschmidtii because of the close identity of theirITS DNA sequences.
Distribution:
Asia(IndiaandThailand)(Mongkolsuk et al., 2012; Zachariah et al., 2019). New
to Myanmar.
Specimen examined: MYANMAR. Tanintharyi Region:
Pha Chaung Inn Beach, Kanbauk Village, Yephyu Township,
N14°30′37″, E97°56′44″, on bark of broad-leaf tree, elev. 0 m, 18
January 2017, Y. Ohmura 12276 (TNS, RAF, TAI).

Sarcographa labyrinthica (Ach.) Müll. Arg., Mém. Soc.
Phys. Genève 29(8): 62. 1887.
Figs. 3A & 3B
For detailed descriptions and synonyms, see Staiger(2002) and Archer (2009). Ascospore size for the
Myanmar collections is (13.8–)16.8–23.4(–26.6) ×(5.3–)6.3–8.1(–9.3) μm (n=51).
Chemistry: Norstictic, stictic and constictic acids.
Remarks: The BLAST results show less than 87%
identity with Graphidaceae sequences in GenBank(Table 1). The low homology results were due to the factthat ITS rDNA sequences of Sarcographa species were
not registered in the GenBank.
Distribution: Pantropical. Asia (India, Indonesia,
Malaysia, the Philippines, Sri Lanka, and Thailand),
Oceania (Australia, New Zealand and Norfolk Island),North and South Americas (Brazil, Mexico and U.S.A.),
Africa, and Europe (Archer, 2009; Singh and Sinha,2010). New to Myanmar.
555

Taiwania

Vol. 65, No. 4

Fig. 3. Lichens collected from southern Myanmar. A–B. Sarcographa labyrinthica (Y. Ohmura 12399, TNS) (A: ascomata on
pseudostroma; B: ascospores in ascus). C–D. Trypethelium eluteriae (Y. Ohmura 12270, TNS) (C: ascomata; D: ascospores inascus). E–G. Zwackhia prosodea (Y. Ohmura 12388, TNS) (E: ascomata; F: vertical section of ascoma; G: ascospores in ascus).
Scales: A, C = 0.5 mm; B, G = 10 μm; D = 20 μm; E = 2 mm; F = 0.1 mm.

Specimens examined: MYANMAR. Tanintharyi Region:
Migyanughlaung, Tanintharyi Nature Reserve, Yephyu Township,
N14°40′18″, E98°08′16″, on bark of broad-leaf tree; elev. 35 m, 23
January 2017, Y. Ohmura 12399 (TNS, RAF, TAI); Migyanughlaung,
Tanintharyi Nature Reserve, Yephyu Township, N14°40′18″,
E98°08′16″, on bark of broad-leaf tree, elev. 35 m, 23 January 2017, Y.
Ohmura 12400 (TNS, RAF).
Trypethelium eluteriae Spreng., Anleit. Kenntn. Gew. 3:
351. 1804.
Figs. 3C & 3D
For a detailed description and synonyms, see Aptrootand Lücking (2016). Ascospore size for the Myanmar
collection is (36.6–)41.1–54.1(–60.9) × (6.8–)7.8–9.6(–
10.5) μm (n=61). Perispore is 1.4–3.9 μm thick. The
number of septa is 9–14.
Chemistry: Parietin.
Remarks: This species somewhat resembles
Astrothelium macrocarpum, but T. eluteriae is readily
distinguished by the morphology of pseudostromata andascospores (see the paragraph on A. macrocarpum).
Distribution: Pantropical. Asia (Cambodia, China,
Hong Kong, India, Indonesia, Japan, Java, thePhilippines, Sri Lanka, Taiwan, Thailand, and Vietnam),
Oceania (Australia, New Caledonia and Papua New
Guinea), North and South Americas (Brazil, Columbia,
556

Costa Rica, Cuba, El Salvador, French Guiana,
Galapagos, Guadeloupe, Guatemala, Mexico, Paraguay,
Surinam, U.S.A., and Venezuela), and Africa (Gambia,
Tanzania and Seychelles) (Aptroot and Lücking, 2016).
New to Myanmar.
Specimen examined: MYANMAR. Yangon Region:
Bogyoke Park, Kandawgyi Lake, N16°47′47″, E96°09′56″, on bark of
Mimusops sp., elev. 20 m, 16 January 2017, Y. Ohmura 12270 (TNS,
RAF).

Zwackhia prosodea (Afzel.) Ertz, Bull. Soc. Naturalistes
Luxemb. 113: 106. 2012.
Figs. 3E–G
For a detailed description, see Pentecost and James(2009, as ‘Opegrapha prosodea’). For synonymy, seeHafellner (2008). Ascospore size for the Myanmar
collection is (60.8–)66.9–85.1(–95.4) × (4.1–)4.9–6.9(–
7.7) μm (n=26). Perispore is 0.9–2.3 μm thick. The
number of septa is 12–15. Excipulum of ascoma is 109–
169 µm wide laterally and 98–221 µm at the base.
Chemistry: No lichen substance was detected by TLC.
Remarks: This species resembles Z. robusta (Vain.)Ertz and Z. viridis (Ach.) Poetsch & Schied. However, Z.
prosodea is distinguished from Z. robusta by the absenceof confluentic acid and from Z. viridis by the larger
ascomata (1–3 mm against 0.4–1 mm for Z. viridis) and2020

Ohmura et al.: Lichens from southern Myanmar

ascospores [50–80 μm against 25–60(–75) μm for Z.
viridis] (Ertz, 2009). But not only these ascospore sizes
but also those of the Myanmar collection widely
overlapped. Whereas, the size of ascoma exipulum for
the Myanmar collection is much thicker (109–169 µm
wide laterally and 98–221 µm at the base) than those of
Z. viridis [25–50 µm wide laterally and 15–80 µm at thebase according to Ertz (2009)], and the size fits to Z.
prosodea (105 – 200 µm at the base; Joseph et al., 2018).
Distribution: Tropical to subtropical regions of Asia
(China, India, Taiwan, and Thailand), Oceania(Australia), North America, Europe (Azores, Madeiraand U.K.), and Africa (Macaronesia) (Tavares, 1952;
Wei, 1991; Aptroot and Rodrigues, 2005; Pentecost andJames, 2009; Aptroot et al., 2010; Singh and Sinha, 2010;
Buaruang et al., 2017). New to Myanmar.
Specimen examined: MYANMAR. Tanintharyi Region:
Tanintharyi Nature Reserve, Yephyu Township, N14°44′12″,
E98°11′30″, on bark of broad-leaf tree, elev. 270 m, 22 January 2017,
Y. Ohmura 12388 (TNS, RAF, TAI).

ACKNOWLEDGMENTS
This study was carried out by the international cooperative
project “Biological Inventory with special attention to
Myanmar” as the integrated research initiated by the National
Museum of Nature and Science, Japan based on amemorandum of understanding (MoU) with the Forest
Department, Ministry of Natural Resources and Environmental
Conservation, Myanmar. We thank to Forest Department of
Myanmar for permitting and supporting our botanical surveys
in the protected areas; to P. Clerc for showing type specimens
collected from Myanmar and housed in G; to S. Joseph forproviding the information of Zwackhia spp.; to K.-O. Nam forassisting DNA experiments; and to two reviewers for carefulreading our manuscript and constructive comments. This study
is partly funded by the JSPS KAKENHI no. 18KK0210 for N.
Tanaka.

LITERATURE CITED
Aptroot, A. 2009. Pyrenulaceae. In: McCarthy, P. M. (ed.),Flora of Australia, Vol. 57, Lichens 5. ABRS, Canberra &CSIRO Publishing, Melobourne. pp. 84–194
Aptroot, A. 2012. A world key to the species of
Anthracothecium and Pyrenula. Lichenologist 44(1): 5–53.
Aptroot, A. and R. Lücking. 2016. A revisionary synopsis of
the Trypetheliaceae (Ascomycota: Trypetheliales).
Lichenologist 48(6): 763–982.
Aptroot, A. and A.F. Rodrigues. 2005. Additions to the
Azorean lichen flora. Arquipélago. Life Marine Sci. 22A:
71–75.
Aptroot, A., A.F. Rodrigues, F. Schumm, S. Câmara and R.
Gabriel. 2010. List of lichens and lichenicolous fungi(Fungi). In: Botges, P. A. V. et al. (eds.), Listagem dos
Organismos Terrestres e Marinhos dos Açores. A list of the
Terrestrial and Marine Biota from the Azores. Princípia,Cascais. pp. 59–79.
Archer, A.W. 2006. The lichen family Graphidaceae inAustralia. Bibl. Lichenol. 94. J. Cramer, Berlin, Stuttgart.
191 pp.

Archer, A.W. 2009. Graphidaceae. In: McCarthy, P. M. (ed.).
Flora of Australia, vol. 57, Lichens 5. ABRS, Canberra &CSIRO Publishing, Melobourne. pp. 84–194.
Archer, A.W. 2012. Graphis supracola A.W. Archer is notendemic to Australia. Australas. Lichenol. 71: 18–19.
Arvidsson, L. 1982. A monograph of the lichen genus
Coccocarpia. Op. Bot. 67: 1–96.
Benatti, M.N. 2012. Three resurrected species of the genus
Bulbothrix Hale (Parmeliaceae, lichenized fungi).
Mycosphere 3(1): 46–55.
Buaruang, K., K. Boonpragob, P. Mongkolsuk, E.
Sangvichien, K. Vongshewarat, W. Polyiam, A.
Rangsiruji, W. Saipunkaew, K. Naksuwankul, J. Kalb,
S. Parnmen, E. Kraichak, P. Phraphuchamnong, S.
Meesim, T. Luangsuphabool, P. Nirongbut, V.
Poengsungnoen, N. Duangphui, M. Sodamuk, S.
Phokaeo, M. Molsil, A. Aptroot, K. Kalb, R. Lücking
and H.T. Lumbsch. 2017. A new checklist of lichenized
fungi occurring in Thailand. MycoKeys 23: 1–91.
Cáceres, M.E.daS. 2007. Corticolous crustose andmicrofoliose lichens of northeastern Brazil. Libri Bot. 22:
1–168.
Culberson, C.F. and A. Johnson. 1982. Substitution of
methyl tert.-butyl ether for diethyl ether in the standardized
thin-layer chromatographic method for lichen products. J.
Chromatogr. 238(2): 483–487.
Elix, J.A. 2009. Physciaceae. In: McCarthy, P. M. (ed.), Floraof Australia, vol. 57, Lichens 5. ABRS, Canberra & CSIRO
Publishing, Melobourne. pp. 494–533.
Ertz, D. 2009. Revision of the corticolous Opegrapha species
from the Paleotropics. Bibl. Lichenol. 102. J. Cramer,
Berlin, Stuttgart. 176 pp.
Ertz, D., A. Tehler, M. Irestedt, A. Frisch, G. Thor and P.
van den Boom. 2015. A large-scale phylogenetic revisionof Roccellaceae (Arthoniales) reveals eight new genera.
Fungal Divers. 70(1): 31–53.
Gardes, M. and T.D. Bruns. 1993. ITS primers withenhanced specificity for Basidiomycetes - application to
the identification of mycorrhizae and rusts. Mol. Ecol. 2(2):113–118.
Gupta, P. and G.P. Sinha. 2016. New records of lichen taxa
from Uttar Pradesh, India. Crypt. Biodivers. Assess. 1(2):64–69.
Flakus, A., H.J. M. Sipman, K. Bach, P.R. Flakus, K.
Knudsen, T. Ahti, U. Schiefelbein, Z. Palice, A.
Jabłońska, M. Oset, R.I. Meneses and M. Kukwa. 2013.
Contribution to the knowledge of the lichen biota of Bolivia.
5. Pol. Bot. J. 58(2): 697–733.
Hafellner, J. 2008. Additions and corrections to the checklist
and bibliography of lichens and lichenicolous fungi of
insular Laurimacaronesia. IV. Fritschiana 64: 1–28.
Hosaka, K. 2009. Phylogeography of the genus Pisolithus
revisited with some additional taxa from New Caledonia
and Japan. Bull. Natl. Mus. Nat. Sci., Ser. B, Bot. 35: 151–
167.
Joseph, S., G.P. Sinha and V.S. Ramachandran. 2018
Taxonomic Revision of the Lichen Genus Opegraphasensu lato (Roccellaceae) in India. Indian Journal ofForestry Additional Series VI. Bishen Singh Mahendra Pal
Singh, Dehra Dun, India. pp. 1–172.
Joshi, S., D.K. Upreti, A.E. Egbe and J.-S. Hur. 2016. New
records of Graphis from Cameroon, with a key to African
species of Graphis. Mycotaxon 131(4): 925–937.

557

Taiwania
Kalb, K., E. Rivas Plata, R. Lücking and H.T. Lumbsch.
2011. The phylogenetic position of Malmidea, a new genus
for the Lecidea piperis- and Lecanora granifera-groups(Lecanorales, Malmideaceae) inferred from nuclear andmitochondrial ribosomal DNA sequences with specialreference to Thai species. Bibl. Lichenol. 106. J. Cramer,
Berlin, Stuttgart. pp. 143–168.
Kalb, K., W. Polyiam, E. Rivas Plata, P.A. Bawingan, K.
Kalb and R. Lücking. 2016. ‘Missing links’ alive? Noveltaxa represent morphological transitions between
distinctive phenotypes among extant Graphidaceae(lichenized Ascomycota: Ostropales). Phytotaxa 268(2):
110–122.
Kashiwadani, H. 1979. Lichens of Kohzu-shima Island, the
Izu Islands. Bull. Natl. Sci. Mus., Ser. B (Bot.), Tokyo 5:
97–106.
Kashiwadani, H., A. Aptroot and K.H. Moon. 2009. New
pyrenocarpous lichens in Japan, with the resurrection of thegenus Trypetheliopsis for Musaespora. In: Aptroot, A. et al.
(eds.): Biodiversity and Ecology of Lichens. Liber
Amicorum Harrie Sipman. Bibl. Lichenol. 99. J. Cramer inder Gebrüder Borntraeger Verlagsbuchhandlung, BerlinStuttgart. pp. 247–258.
Lücking, R. 2009. The taxonomy of the genus Graphis sensu
Staiger (Ascomycota: Ostropales: Graphidaceae).
Lichenologist 41(4): 319–362.
Lücking, R., A.W. Archer and A. Aptroot. 2009. A worldwide key to the genus Graphis (Ostroplaes: Graphidaceae).
Lichenologist 41(4): 363–452.
Mittermeier, R.A., P. Robles Gil, M. Hoffmann, J.D.
Pilgrim, T.M. Brooks, C. G. Mittermeier and G.A.B. da.
Fonseca. 2004. Hotspots Revisited: Earth’s Biologically
Richest and Most Endangered Ecoregions. Mexico City:
Cemex. 390 pp.
Moberg, R. 1990. The lichen genus Physcia in Central andSouth America. Nord. J. Bot. 10(3): 319–342.
Moberg, R. 2001. The lichen genus Physcia in Australia. In:
McCarthy, P. M. et al. (eds.), Lichenological Contributionsin Honour of Jack Elix. Bibl. Lichenol. 78. J. Cramer,
Berlin, Stuttgart. pp. 289–311.
Mongkolsuk, P., S. Meesim, V. Poengsungnoen and K.
Kalb. 2012. The lichen family Physciaceae in Thailand–I.
The genus Pyxine. Phytotaxa 59(1): 32–54.
Moon, K.H., M. Nakanishi, Y. Futagami and H.
Kashiwadani. 2015. Studies on Cambodian species ofGraphidaceae (Ostropales, Ascomycota) (II). J. Jpn. Bot.
90: 98–102.
Muggia, L., C. Gueidan, G. Perlmutter, O.E. Eriksson and
M. Grube. 2009. Molecular data confirm the position of
Flakea papillata in the Verrucariaceae. Bryologist 112(3):
538–543.
Müller, J. 1889. Lichenologische Beiträge 32. Flora(Regensburg) 72: 505–508.
Nakanishi, M. 1966. Taxonomical studies on the family
Graphidaceae of Japan. J. Sci. Hiroshima Univ., Ser. B, Div.
2 (Bot.) 11: 51–126.

558

Vol. 65, No. 4

Nylander, W. and J.M. Crombie. 1883. On a collection of
exotic lichens made in eastern Asia by the late Dr. A. C.
Maingay. J. Linn. Soc., Bot. 20(125): 48–69.
Ohmura, Y. and H. Kashiwadani. 2018. Checklist of Lichens
and Allied Fungi of Japan. National Museum of Nature andScience Monographs 49. Natl. Mus. Nat. Sci., Tokyo. 143
pp.
Pentecost, A. and P.W. James. 2009. Opegrapha Ach. (1809).
In: Smith, C. W. et al. (ed.) 2009. The Lichens of GreatBritain and Ireland. British Lichen Society, London. pp.
631–647.
Perlmutter, G.B. 2006. Flakea papillata in North America.
Bryologist 109(4): 566–569.
Rivas Plata, E., R. Lücking and H.T. Lumbsch. 2012. A newclassification for the family Graphidaceae (Ascomycota:
Lecanoromycetes: Ostropales). Fungal Divers. 52(1): 107–
121.
Singh, K.P. and G.P. Sinha. 2010. Indian Lichens: an
Annotated Checklist. Botanical Survey of India. Ministryof Environment and Forests, Salt Lake City, Kolkata. 571
pp.
Staiger, B. 2002. Die Flechtenfamilie Graphidaceae. Studienin Richtung einer natürlicheren Gliederung. Bibl. Lichenol.
85. J. Cramer, Berlin, Stuttgart. 526 pp.
Tanaka, N., S. Tagane, A. Naiki, M.M. Aung, N. Tanaka, S.
Dey, J. Mood and J. Murata. 2018. Contributions to the
Flora of Myanmar I: Nine taxa of monocots newly recorded
from Myanmar. Bull. Natl. Mus. Nat. Sci., Ser. B (Bot.) 44:
31–39.
Tavares, C.N. 1952. Contributions to the lichen flora ofMacaronesia. I. Lichens from Madeira. Port. Acta Biol. (B)
3: 308–391.
Thor, G. and H. Kashiwadani. 1996. Zeorin and two othertriterpenoids found in Flakea papillata. Systema
Ascomycetum 14: 87–90.
Tordoff, A.W., M.R. Bezuijen, J.W. Duckworth, J. R.
Fellowes, K. Koenig, E. H. B. Pollard and A. G. Royo.
2011. Ecosystem profile: Indo-Burma biodiversity hotspot,
2011 Update. Washington DC, Critical Ecosystem
Partnership Fund. 381 pp.
Vilgalys, R. and M. Hester. 1990. Rapid genetic identificationand mapping of enzymatically amplified ribosomal DNA
from several Cryptococcus species. J. Bacteriol. 172(8):4238–4246.
Wei, J. 1991. An enumeration of lichens in China.
International Academic Publishers, Beijing. 278 pp.
Zachariah, S.A., S. Nayaka, P. Gupta and S.K. Varghese.
2019. The lichen genus Pyxine (Caliciaceae) in Kerala statewith P. dactyloschmidtii as new to India. Hattoria 10: 109–
117.
Zhang, Y.Y., X.Y. Wang, J.W. Li, H.A. Shi, X. Ye and L.I.
Wang. 2014. Bulbothrix asiatica sp. nov., and other new
records of Parmeliaceae with bulbate cilia from Cambodia.
Bryologist 117(4): 379–385.