Bull. Natl. Mus. Nat. Sci., Ser. B, 37(2), pp. 55–61, May 22, 2011

Notes on Eight Threatened Species of Lichens in Japan
Yoshihito Ohmura
Department of Botany, National Museum of Nature and Science,
Amakubo 4–1–1, Tsukuba, 305–0005 Japan
E-mail: ohmura-y@kahaku.go.jp(Received 17 February 2011; accepted 23 March 2011)

Abstract New localities and/or molecular data are provided for the following eight threatened
species of lichens in Japan: Chiodecton congestulum, Flavopunctelia soredica, Icmadophila japonica, Lethariella togashii, Pseudocyphellaria argyracea, Thysanothecium scutellatum, Usnea filipendula, and U. nipparensis. ITS rDNA sequences were successfully obtained from all species
except for C. congestulum, and were compared with the known sequences retrieved from GenBank.
Key words : distribution, ITS rDNA, lichenized Ascomycota, Red List.

situation of each species as well as molecular
data.

Introduction
The current Red List of Japanese lichens iscomprised of 152 taxa: 152 species, 1 subspecies, and 3 varieties (Ministry of the Environment, 2007; Inoue et al., 2010). Molecular dataof these lichens are of potentially great value fortaxonomic, phylogenetic, phytogeographic, andecological purposes.
During recent field surveys in Japan, fresh materials of eight threatened species were collected
in 2009–2010 from both new and already known
localities, and the sequences of internal transcribed spacer of ribosomal DNA (ITS rDNA)
were successfully obtained from all extracted
DNA samples except for Chiodecton congestulum. The purpose of this study is to provide information of new localities, and/or the present
Table 1.

Materials and Methods
All collections examined are stored at the
National Museum of Nature and Science, Tokyo(TNS) unless otherwise indicated. Two specimens of Chiodecton congestulum are stored at
Uppsala University (UPS).
Lichen substances were examined by thin layer
chromatography (TLC) (Culberson and Johnson,1982). Only the solvent B system was used here.
Materials for molecular analysis and the associated GenBank accession numbers are shown in
Table 1. DNA was extracted from 5–10 mg oflichen thalli by using a FastDNA SPIN Kit (MPBiomedicals) following the method of Ohmura etSpecimen vouchers for DNA analysis and the GenBank accession numbers obtained in this study
SpeciesCollection number of Y. Ohmura (in TNS)

GenBank accession no.

Flavopunctelia soredica
Icmadophila japonica
Lethariella togashii
Pseudocyphellaria argyracea
Thysanothecium scutellatum
Usnea filipendula
U. nipparensis

7164
6762
6735
6418
6773
6771
6282

AB623069
AB623070
AB623071
AB623072
AB623073
AB623074
AB623075

56

Yoshihito Ohmuraal. (2006).
PCR amplification of the ITS rDNA region(including ITS1, 5.8S rDNA, and ITS2) was performed using ITS1F (Gardes and Bruns, 1993) as
the 5 primer and LR1 (Vilgalys and Hester,1990) as the 3 primer. PCR reactions were performed according to the method of Ohmura et al.
(2006) by using PuReTaq Ready-To-go PCRbeads (GE Healthcare) and a thermal cycler(Gene Amp PCR system 9700, Applied Biosystems). PCR products were purified using an
ExoSAP-IT (GE Healthcare). Sequencing was
carried out using a BigDye Terminator ver. 3.1
Cycle Sequencing Kit on an ABI Prism 3130x
genetic analyzer (Applied Biosystems) according
to the manufacturer’s instructions. DNA strands
were assembled and manually corrected using
ATGC ver. 6.03 (Genetyx). The obtained sequences in this study were compared with the
known sequences of related species registered in
GenBank by using BLAST, a homology search
program (Altschul et al., 1997).
Results and Discussion
Chiodecton congestulum Nyl., Bull. Soc. Linn.
Normandie, sér. 2, 2: 106 (1868).
Chiodecton congestulum was well revised taxonomically by Thor (2002). This species has
been reported from India, Fiji, Australia,
Malaysia, and Japan (Thor, 2007). In Japan, it
has been known from western Honshu, Kyushu,
Ogasawara Islands, and Yaeyama Islands (Thor,
2002). New localities were found at Mt. Horaiji
in Aichi Prefecture and Nagatani Shrine in Kochi
Prefecture.
Fresh materials of this species were also collected from Miyajima Island in Hiroshima Prefecture. Although DNA was extracted from thespecimens, the ITS rDNA sequence could not be
obtained.
As reported by Thor (2002), this species grows
on rocks and tree barks. The specimens collectedfrom Aichi and Hiroshima Prefectures were saxicolous, and those collected from Kochi Prefecture were corticolous.

Specimen examined. Japan. Honshu. Prov.
Mikawa (Pref. Aichi): Mt. Horaiji, Kadoya,
Horai-cho, Minamishitara-gun, on rock, elevation about 315 m, 5 December 1996, Y. Ohmura
2840. Prov. Aki (Pref. Hiroshima): Mt. Misen,
Miyajima Island, Hatsukaichi-city, on rock (granite), elevation 259 m, 12 January 2010, M. Kono
(41 and 42), Y. Ohmura & Y. Koda. Shikoku.
Prov. Tosa (Pref. Kochi): Nagatani Shrine, Nagasawa, Tsuno-mura Village, on coniferous tree,
elevation 590–610 m, 30 October 2006, G. Thor
21515 (UPS, det. G. Thor, not seen); ditto, on
stump, G. Thor 21516 (UPS, det. G. Thor, notseen).
Flavopunctelia soredica (Nyl.) Hale, Mycotaxon, 20: 682 (1984).
Flavopunctelia soredica is a well-known foliose species of Parmeliaceae and is easily distinguished from similar species on the basis of thepresence of soralia along the margin of lobes andusnic and lecanoric acids as major substances in
the cortex and medulla respectively. Although F.
soredica is widely distributed in the Northern
Hemisphere, its presence is restricted in central
Honshu, Japan, where it has been found on barksof Betula, Castanea, Chamaecyparis, Larix,
Pinus, Prunus, and Zelkova (Asahina, 1941;
Kurokawa and Kashiwadani, 1988; Saitama
Museum of Natural History, 1988; Kashiwadani,
1991; Harada et al., 2003; Morita and Harada,
2006). Additional specimens were collected from
Arakoda (Saku-city) at 730 m alt. and Sugadaira(Ueda-city) at 1577 m alt. in Nagano Prefecture.
They were found on barks of Larix and Quercus.
Although the known localities are restricted to anarrow region in Japan, F. soredica appears togrow well in these conditions.
ITS rDNA sequences were successfully obtained from two specimens that were both collected from the same locality of Sugadaira but
from different trees: Betula ermanii and Quercus
crispula. Each ITS rDNA sequence is 495 bp inlength, and there are no differences between
them. The known F. soredica sequence from New
York, U.S.A. [GenBank accession no. AY773128

Eight Threatened Lichens in Japan57

Icmadophila japonica (Zahlbr.) Rambold &Hertel, Biblioth. Lichenol., 53: 230 (1993).
 Glossodium japonicum Zahlbr., Bot. Mag.
Tokyo, 41: 336 (1927).
Icmadophila japonica is a well-known speciesof lichens under the name of Glossodium japonicum Zahlbr. by Japanese lichenologists (seeRambold et al., 1993). This species generally
grows on decayed wood of coniferous trees in an
old-growth forest, and its distribution is restricted
to Far East Russia and Japan. In Japan, it grows
widely from Hokkaido to Kyushu (Sato, 1967),
but the populations in Japan are decreasing
according to recent field surveys (Inoue et al.,2010).
ITS rDNA sequence was successfully obtained
from one specimen collected from Mt. O-akan,
Hokkaido. The sequence of ITS rDNA region is466 bp in length. No reference ITS rDNA
sequence of Icmadophila has been registeredin GenBank.
Further molecular phylogenetic analysis basedon ITS rDNA sequences and other appropriate
markers is needed to examine the independenceof Glossodium and Icmadophila (s. str.) sincemorphology of the ascomata is quite different
between them.
Specimens examined. Japan. Hokkaido. Prov.
Kushiro: Mt. O-akan, on decayed stump, elevation 494 m, 10 August 2009, Y. Ohmura 6762.

Bot., 23: 95 (1976).
Lethariella togashii was revised taxonomicallyby Asahina (1952), Krog (1976), and Obermayer(1997). The distribution of this species is restricted to Far East Russia and Japan (Skirina, 2006).
In Japan, it has been collected from Hokkaido
and central Honshu (e.g. around Lake Yamanaka,
Mt. Fuji), but the habitats in the latter locality
were heavily damaged and the populations arealmost disappeared (Kashiwadani and Inoue,
2000).
The ITS rDNA sequence was successfully
obtained from one specimen collected at Mt.
O-akan, Hokkadio. The sequence of the ITS
rDNA region is 501 bp in length. According to
the BLAST result, the top scores of known related species were obtained with Letharia
columbiana (Nutt.) J.W. Thomson [AF115762(Thell et al., unpublished); identity434/461 bp,94%], L. vulpina (L.) Hue [GQ398408 (Altermann et al., unpublished); identity428/456,
cashmeriana
Krog.
93%],
Lethariella[DQ980014 (Crespo et al., 2007); identity521/457, 92%], and L. sernanderi (Motyka)
Obermayer [AF297744 (Kroken, unpublished);
identity420/459, 91%]. Although large-scale
phylogenetic analysis based on three ribosomal
DNA regions and RPB1 gene supports a monophyletic clade consisting of Letharia columbiana
and Lethariella cashmeriana (Crespo et al.,2007), a narrower scale analysis has not yet been
performed to confirm generic independence between them. The genus Letharia is morphologically separated from Lethariella mainly by thelacerate axis. The yellow-colored thallus due tovulpinic acid is distinct in Letharia spp., but thissubstance is also present as a major substance inLethariella togashii. The delimitation and independence between these genera should be revisedin the further researches.
Specimen examined. Japan. Hokkaido. Prov.
Kushiro: Mt. O-akan, on bark of Picea glehnii,
elevation 728 m, 10 August 2009, Y. Ohmura
6735.

Lethariella togashii (Asahina) Krog, Norw. J.

Pseudocyphellaria argyracea (Delise) Vain.,(Thell et al., 2005)] is different in only two
residues from Japanese materials (identity
493/495 bp, 99%).
Specimens examined. Japan. Honshu. Prov.
Shinano (Pref. Nagano): along the route from
Sugadaira Pasture to Mt. Azumaya, Ueda-city, onbark of Quercus crispula, elevation 1577 m, 30
May 2010, Y. Ohmura (7119 and 7164) & A.
Okamura; ditto, on bark of Betula ermanii, Y.
Ohmura (7121 and 7167) & A. Okamura; Arakoda, Saku-city, on bark of Larix leptolepis, elevation about 730 m, 17 July 1972, H. Kashiwadani
10332.

58

Yoshihito Ohmura

Fig. 1. Pseudocyphellaria argyracea. A. Habitat in Kita-iwo Island. B. Isidia along the margins of lobes and on
pseudocyphellae (Y. Ohmura 6418). Scale1 mm.

Eight Threatened Lichens in Japan

Hedwigia, 37: 35 (1898).
[Fig. 1]
Pseudocyphellaria argyracea is characterized
by the rosette-forming to irregularly spreading
foliose thallus with laminal white soralia andpseudocyphellae on the upper surface, and thesimple to coralloid isidia on pseudocyphellae andthe margins of lobes. This species is widely distributed in the Palaeotropics from East Africa to
India, Japan, New Zealand, South America, andthe islands of the Pacific Ocean (Galloway et al.,2001). In Japan, it has been collected from
Kyushu and Bonin Islands ( Ogasawara Islands)(Kurokawa, 1957, 1969; Kurokawa and Kashiwadani, 1988). Rich populations of this species
are newly found in Kita-iwo Island of Sulphur
Islands.
The ITS rDNA sequences were obtained fromfive specimens of Kita-iwo Island. They are 455
bp in length and no differences are found among
the sequences. The identity with the known sequence of this species from Réunion in the Indian Ocean is 93% (427/456 bp) [EU558727(Högnabba et al., unpublished)].
Specimens examined. Japan. Sulphur Islands:
Kita-iwo Island, on bark of Eurya japonica, elevation 522 m, 18 June 2009, Y. Ohmura 6394;
ditto, elevation 448 m, Y. Ohmura 6418 and
6419; ditto, on bark of Elaeocarpus sylvestrisvar. pachycarpus, elevation 490 m, Y. Ohmura
6404; Sakakigamine, Kita-iwo Island, on tree
trunk, elevation about 750 m, 19 June 2009, T.
Yamaguchi s. n. (Herb. Y. Ohmura 6452).
Thysanothecium scutellatum (Fr.) D.J. Galloway, Nova Hedwigia, 36: 390 (1982).
Thysanothecium scutellatum is a well-known
fruticose species of Cladoniaceae. It has been reported from Oceania, Southeast Asia, and East
Asia (Galloway and Bartlett, 1982; Wei et al.,
1994; Kashiwadani, 2008). In Japan, it has been
known from central to western Honshu, Shikoku,
and Kyushu where it grows on barks of Cryptomeria, Pinus, Washingtonia, and decayedstump elevation at 5 to 700 m (Asahina, 1956;
Okamoto and Iwatsuki, 1992; Kurokawa andKashiwadani, 1988; Kashiwadani, 1998, 2008;

59

Okamoto, 2010).
The ITS rDNA sequence was successfully obtained from one specimen collected in Miyajima
Island, Hiroshima Prefecture, and it is 551 bp inlength. According to the BLAST result, the top
score of known related species is obtained with
Notocladonia cochleata (Müll. Arg.) S. Hammer( Ramalea cochleata Müll. Arg.) [AF453267(Stenroos et al., 2002); identity514/525 bp,97%]. As discussed by Hammer (2003), thegenus Notocladonia is related to Thysanothecium
as well as Cladia and Squamella in the morphology. The identity of ITS rDNA sequences between T. scutellatum and N. cochleata is veryhigh. The main morphological difference between the genera is the branching patterns ofpodetia. Further phylogenetic studies of thesegenera are needed based on the ITS rDNA
sequences and other appropriate markers.
Specimens examined. Japan. Honshu. Prov.
Mikawa (Pref. Aichi): Mt. Horaiji, Kadoya,
Horai-cho, Minamishitara-gun, on base of decayed stump, elevation about 450 m, 5 December
1996, Y. Ohmura 2843. Prov. Yamashiro (Kyotofu): Takenouchi-cho, Sakyo-ku, Kyoto-city, onbark of Cryptomeria japonica, elevation about
140 m, 6 December 2005, H. Kashiwadani 47808
and 47809. Prov. Ise (Pref. Mie): Kitayama, Isecity, on bark of Cryptomeria japonica, 15 June
1976, C. Chuma 3786. Prov. Aki (Pref. Hiroshima): Miyajima Island, on bark of Pinus densiflora, elevation about 5 m, 10 January 1993, Y.
Ohmura 337; ditto, 19 March 1995, Y. Ohmura
936; ditto, on bark of Pinus thunbergii, elevation
2 m, 26 August 2009, Y. Ohmura 6773.
Usnea filipendula Stirt., Scott. Natural. 6: 104(1881).
Usnea filipendula is characterized by the fruticose thallus with a jet blacked base and the elongated terminal branches with convex soralia, andthe presence of salazinic acid as a major substance. For a detailed description, see Ohmura(2001). This species is widely distributed in circumpolar boreal to northern temperate regions(Thomson, 1984). In Japan, however, it has been

60

Yoshihito Ohmura

known only from two localities on Hokkaido(Ohmura and Onimaru, 2010).
The ITS rDNA sequence of the Japanese specimen is 492 bp in length. It is 100% identical to
the sequence from Sweden (492/492 bp)[AJ457149 (Articus et al., 2002)].
Specimens examined. Japan. Hokkaido. Prov.
Kitami: Hinami, Bihoro-cho, Abashiri-gun, onbark of Abies sachalinensis, elevation 330 m, 11
August 2009, Y. Ohmura 6771.
Usnea nipparensis Asahina, Lich. Jap. 3: 91(1956).
Usnea nipparensis is easily distinguished fromother related Japanese Usnea species by therounded soralia which are distinctly stipitate andlarger than branch diameter, and the presence of
caperatic acid as a major substance. For a detailed description, see Ohmura (2001). This
species is distributed in Japan, Korea, Taiwan,
India, and Nepal. In Japan, it has been known
from central Honshu.
A new locality was found in Nishizawa Valley,
Yamanashi Prefecture. The ITS rDNA sequenceswere successfully obtained from two specimens,
and they are both 494 bp in length. The sequences are 100% identical with that from Yatsugatake Mts., Nagano Prefecture in Japan(494/494 bp) [AB051652 (Ohmura, 2002)].
Specimens examined. Japan. Honshu. Prov.
Kai (Pref. Yamanashi): Nishizawa Valley, Mitomi-Kawaura, Yamanashi-city, on bark of Larix
kaempferi, elevation 1160 m, 8 February 2009, Y.
Ohmura (6274 and 6282), A. Crespo & P.K.
Divakar.
Acknowledgments
I thank Y. Hirayama and T. Hosaka for assisting with the molecular experiments and G. Thor
for providing information on one new locality of
Chiodecton congestulum. This study is partly
funded by the Grant-in-Aid for Young Scientists(B) of the Japan Society for the Promotion of
Science (no. 21770097).

References
Altschul, S. F., Madden, T. L., Schäffer, A. A., Zhang, J.,
Zhang, Z., Miller, W. and Lipman, D. J. 1997. Gapped
BLAST and PSI-BLAST: a new generation of protein
database search programs. Nucleic Acids Research 25:
3389–3402.
Articus, K., Mattsson, J.-E., Tibell, L., Grube, M. and
Wedin, M. 2002. Ribosomal DNA and b -tubulin data
do not support the separation of the lichens Usnea florida and U. subfloridana as distinct species. MycologicalResearch 106: 412–418.
Asahina, Y. 1941. Lichenologische Notizen XV. Journalof Japanese Botany 17: 71–76.
Asahina, Y. 1952. Lichenologische Notizen §85–87. Journal of Japanese Botany 27: 293–296.
Asahina, Y. 1956. Lichenologische Notizen §117–119.
Journal of Japanese Botany 31: 65–70.
Crespo, A., Lumbsch, H. T., Mattsson, J.-E., Blanco, O.,
Divakar, P. K., Articus, K., Wiklund, E., Bawingan, P.
A. and Wedin, M. 2007. Testing morphology-based hypotheses of phylogenetic relationships in Parmeliaceae(Ascomycota) using three ribosomal markers and thenuclear RPB1 gene. Molecular Phylogenetics and Evolution 44: 812–824.
Culberson, C. F. and Johnson, A. 1982. Substitution of
methyl tert.-butyl ether for diethyl ether in the standardized thin-layer chromatographic method for lichen
products. Journal of Chromatography 238: 483–487.
Galloway, D. J. and Bartlett, J. K. 1982. The lichen genus
Thysanothecium Mont. & Berk., in New Zealand. Nova
Hedwigia 36: 381–398.
Galloway, D. J., Kantvilas, G. and Elix, J. A. 2001.
Pseudocyphellaria. In: McCarthy, P. M. (ed.), Flora ofAustralia. Volume 58A, Lichens 3, pp. 47–77.
ABRS/CSIRO Australia, Melbourne.
Gardes, M. and Bruns, T. D. 1993. ITS primers with enhanced specificity for Basidiomycetes—application to
the identification of mycorrhizae and rusts. Molecular
Ecology 2: 113–118.
Hammer, S. 2003. Notocladonia, a new genus in the
Cladoniaceae. Bryologist 106: 162–167.
Harada, H., Anzai, T. and Yamamoto, Y. 2003. Materials
for the study of distributions of lichenized fungi (4).
Flavopunctelia soredica. Lichenology 2: 23–24.
Inoue, M., Ohmura, Y., Kashiwadani, H., Nakanishi, M.,
Matsumoto, T., Miyawaki, H., Moon, K. H. and Yoshida, K. 2010. Revised Red List—supplement. Lichens,
pp. 1–26. Ministry of Environment, Tokyo (in Japanese).
Kashiwadani, H. 1991. Lichens. In: Committee of Shiojiri-city History (ed.), Plants of Shiojiri-city. History of
Shiojiri-city 1 (Nature), pp. 702–715, 986–995. Shiojiri-city (in Japanese).

Eight Threatened Lichens in Japan
Kashiwadani, H. 1998. Lichenes minus cogniti exsiccati.
Fasc. V (Nos. 101–125). National Science Museum,
Tokyo.
Kashiwadani, H. 2008. Lichenes minus cogniti exsiccati.
Fasc. XV (Nos. 351–375). National Science Museum,
Tokyo.
Kashiwadani, H. and Inoue, M. 2000. Lichens. In: Environment Agency. Threatened Wildlife of Japan –Red
Data Book 2nd ed.—Vol. 9, Bryophytes, Algae,
Lichens, Fungi, pp. 259–312. Japan Wildlife ResearchCenter, Tokyo.
Krog, H. 1976. Lethariella and Protousnea, two newlichen genera in Parmeliaceae. Norwegian Journal of
Botany 23: 83–106.
Kurokawa, S. 1957. Habitats of some lichens. Miscellanea
Bryologica et Lichenologica 11: 1 (in Japanese).
Kurokawa, S. 1969. Lichens of Chichijima Island of theBonin Islands collected by Dr. H. Inoue. Bulletin of the
National Science Museum 12: 685–692, pl. 1–3.
Kurokawa, S. and Kashiwadani, H. 1988. Indices to taxa
distributed under lichenes rariores et critici exsiccati,
114 pp. National Science Museum, Tokyo.
Ministry of the Environment. 2007. Revision of Red Listfor mammalia, fishes in brackish and fresh waters, insects, shells, plant I and II. http://www.env.go.jp/press/
press.php?serial8648 (in Japanese).
Morita, T. and Harada, H. 2006. Materials for the study of
distributions of lichenized fungi (17). Flavopuncteliasoredica occurs in Karuizawa-machi, Nagano-ken,Central Japan. Lichenology 5: 179.
Obermayer, W. 1997. Studies on Lethariella with special
emphasis on the chemistry of the subgenus Chlorea.
Bibliotheca Lichenologica 68: 45–66.
Okamoto, T. 2010. New locality and substrate of Thysanothecium scutellatum (Fr.) D. J. Galloway (Cladoniaceae, lichenized Ascomycetes). Hikobia 15:
361–362.
Okamoto, T. and Iwatsuki, Z. 1992. New localities of four
rare lichens in Japan. Hikobia 11: 213–215.
Ohmura, Y. 2001. Taxonomic study of the genus Usnea(lichenized Ascomycetes) in Japan and Taiwan. Journalof the Hattori Botanical Laboratory 90: 1–96.
Ohmura, Y. 2002. Phylogenetic evaluation of infragenericgroups of the genus Usnea based on ITS regions in
rDNA. Journal of the Hattori Botanical Laboratory 92:
231–243.
Ohmura, Y., Kawachi, M., Kasai, F., Watanabe, M. M. andTakeshita, S. 2006. Genetic combinations of symbionts

61

in a vegetatively reproducing lichen, Parmotrema tinctorum, based on ITS rDNA sequences. Bryologist 109:
43–59.
Ohmura, Y. and Onimaru, K. 2010. Materials for the distribution of lichens in Japan (16) Usnea filipendula
Stirt. Journal of Japanese Botany 85: 190–192.
Rambold, G., Triebel, D. and Hertel, H. 1993. Icmadophilaceae, a new family in the Leotiales. Bibliotheca
Lichenologica 53: 217–240.
Saitama Museum of Natural History (ed.) 1988. A catalogue of the materials in the Saitama Museum of Natural History, No. 3, Lichens. 73 pp. Saitama Museum of
Natural History, Nagatoro.
Sato, M. 1967. Distibution and substratum of the Japanese lichens, (1) Glossodium japonicum Zahlbr. Miscellanea Bryologica et Lichenologica 4: 75–77 (in Japanese with English summary).
Skirina, I. F. 2006. Lethariella togashii (Parmeliaceae), arare new species to the lichen flora of Russia from thesouthern Far East. Botanicheskii Zhurnal (St. Petersburg) 91: 1114–1116.
Stenroos, S., Hyvönen, J., Myllys, L., Thell, A. and Ahti,
T. 2002. Phylogeny of the genus Cladonia s. lat.
(Cladoniaceae, Ascomycetes) inferred from molecular,
morphological, and chemical data. Cladistics 18:
237–278.
Thell, A., Herber, B., Aptroot, A., Adler, M. T., Feuerer,
T. and Kärnefelt, E. I. 2005. Preliminary phylogeographic study of Flavopunctelia and Punctelia inferredfrom rDNA ITS-sequences. Folia Cryptogamica Estonica 41: 115–122.
Thomson, J. W. 1984. American Arctic Lichens. 1. The
Macrolichens. Columbia University Press. New York.
Thor, G. 2002. The genera Chiodecton, Dichosporidium,
Graphidastra and Pulvinodecton (Lichens) in Japanand Taiwan, with notes on certain species-rich oldgrowth forests. Journal of Japanese Botany 77: 47–58.
Thor, G. 2007. The genera Chiodecton, Dichosporidium
and Erythrodecton in Peninsular Malaysia. Bibliotheca
Lichenologica 95: 543–548.
Vilgalys, R. and Hester, M. 1990. Rapid genetic identification and mapping of enzymatically amplified ribosomal DNA from several Cryptococcus species. Journalof Bacteriology 172: 4238–4246.
Wei, J., Jiang, Y. and Guo, S. 1994. Studies on the lichenfamily Cladoniaceae in China III. A new genus toChina: Thysanothecium. Mycosystema 7: 23–27.