Bull. Natn. Sci. Mus., Tokyo, Ser. B, 32(4), pp. 161–166, December 22, 2006

Noteworthy Species of the Genus Ramalina (Ascomycotina:Ramalinaceae) in China
Hiroyuki Kashiwadani1, Kwang Hee Moon2, Shouyu Guo3,Yun-Cheng Dai4 and Xi-Ling Chen4
1
Department of Botany, National Science Museum,4–1–1 Amakubo, Tsukuba, Ibaraki 305–0005 Japan
E-mail: hkashiwa@kahaku.go.jp
2
Laboratory of Mycology, Biological Sciences-Systematics and Ecology,
College of Natural Sciences, Seoul National University,San 56–1, Sillim 9 dong, Gwanak-gu, Seoul 151–742 Korea
3Key Laboratory of Systematic Mycology & Lichenology,
Institute of Microbiology, Chinese Academy of Sciences, Beijing 100080, P.R. China
4
Institute of Applied Ecology, Chinese Academy of Sciences,Wenhua Road 72, Shenyang 10016, P.R. China

Abstract Ramalina almquistii, R. hokkaidensis, R. intermedia, R. obtusata, R. pentecostii, R.
pertusa and R. sekika are reported from China for the first time. R. pentecostii and R. sekika are thesecond report for the species. R. pseudosekia was reduced to a synonym of R. pollinaria.
Key words : Ramalina almquistii, R. hokkaidensis, R. intermedia, R. obtusata, R. pentecostii, R.
sekika, lichens.

In the course of our taxonomic study of thegenus Ramalina basing on herbarium specimens
kept in the Institute of Microbiology, Academia
Sinica (HMAS), the Institute of Applied Ecology,
Academia Sinica (IFP) and the National ScienceMuseum, Tokyo (TNS), we found following
seven species to be new to China; R. almquistii,
R. hokkaidensis, R. intermedia, R. obtusata andR. pertusa. In addition, R. pentecostii and R.
sekika were found and they are the second reportfor the species in China. The present paper presents characteristic features for each species, including chemical variations, based on Chinesespecimens.
Materials and Methods
The specimens used for the present study wereprimarily based on the herbarium specimens kept
in HMAS, IFP and TNS. Specimens collected by
Kashiwadani and Moon in China in 1995 and1999 were also studied. The secondary productsof the specimens examined were determined bythin layer chromatography (TLC) using theamended procedures of Culberson & Johnson(1982). To evaluate anatomical variation within
the thallus and apothecia, sections were cut byhand with a razor blade. These were mounted in
GAW solution and the anatomy and spores wereexamined with standard light microscopic procedures.
Notes on Species
Ramalina almquistii Vain., Ark. Bot. 8: 17(1909).
Ramalina almquistii is a fistulose species characterized by saxicolous habit, discontinuous
medullary hyphae, absence of soredia or isidia,
short fusiform ascospores and by the presence of
sekikaic or divaricatic acids.
Although this species is widely distributed in
East Asia, it has never been recorded from China.
Three specimens collected on rocks at Prov.

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Hiroyuki Kashiwadani et al.

Liaoning were now identified with the divaricaticacid race of this species. New to China.
Specimens examined. China. Prov. Lianing:
Laotieshan, on rock, elevation 250 m, October 7,1981, X. Chen et al. 5762 (IFP); the same locality, September 1, 1931, Y. Kobayashi 13 (TNS);
Kuandian, Baishilazi Reserve, on rock, 1245 m
alt, X. Chen 6844 (IFP) .
Ramalina hokkaidensis Kashiw., Bull. Natn.
Sci. Mus. Ser. B, 12: 89 (1986).
This species was well studied by Kashiwadani(1986) for Japanese specimens. It very much resembles Ramalina conduplicans Vain., a species
widely distributed in East Asia, in having similar
laciniae with punctiform pseudocyphellae. However, it can be distinguished from the latter by thecanaliculate laciniae tapering towards the apices
and by the presence of stenosporic acid. R.
conduplicans has dorsiventral laciniae even in
the distal branches and produces homosekikaic,
sekikaic or divaricatic acids as major chemicalsubstances. New to China.
Specimen examined. China. Prov. Heilongjiang: Liangshui forest centre, Dailing, 300m, October 6, 1975, X. Chen 3863 & 3907-1(IFP). Prov. Sichuan, Liangshan Co.: N slope ofMt. Luoji, Xichang City (27°45N, 102°20E),
on twigs of Rhododendron sp., elevation
2205–2245 m, September 21, 1999, H. Kashiwadani 42376 (TNS).
Ramalina intermedia (Del. ex Nyl.) Nyl., Flora
56: 66 (1873).
(Fig. 1a & 1b)
Thallus saxicolous, up to 3 cm long. Laciniae
discrete or forming swards, sparingly and irregularly branched, growing from a delimited or moreor less expanded holdfast. Laciniae greenish yellow, solid, 0.3–1.5(–3) mm wide, flattened towards the basal parts, terete and more or lesscylindrical in distal parts, often ending in nodular
or granular soredia. Pseudocyphellae sparse, elliptical, flat. Soralia present, laminal or subterminal to terminal, soredia granular, covered byouter layers. Laciniae 150–400 m m thick; uppercortex indistinct, 3–5 m m thick; chondroid tissue

weakly cracked, continuous; medulla loose.
Apothecia very rare, laminal or submarginal, up
to 0.7 mm in diam.; disc flat to more or less convex, without white margin; thalline exciple entire
or sorediate; hymenium 35–40 m m high; subhymenium 15–20 m m thick, proper exciple 30–40
m m thick; ascospores shortly fusiform, 11–12
4–4.5 m m. Pycnidia not seen.
Chemistry: (Race 1) usnic, homosekikaic,sekikaic and 4-O-methylnorhomosekikaic acids.
(Race 2) usnic, divaricatic and 4-O-demethyldivaricatic acids.
Ramalina intermedia includes a series offorms ranging from discrete and shrubby thalli
growing from a delimited holdfast up to 3 cm
high with irregularly or dichotomously branchedand dorsiventral laciniae near the base (Fig. 1a),to small tufted thalli growing from more or lessexpanded holdfast up to 1 cm high with more orless terete and weakly branched laciniae (Fig.
1b). Similar morphological variations were also
reported for R. polymorpha (Ach.) Ach. by
Acharius (1797) and Krog and James (1977).
When Kashiwadani (1987) revised Japanese
species of R. intermedia, he reported the presence of homosekikaic and sekikaic acids as
major chemical substance. In addition to the race
1 containing homosekikaic acid agg., divaricaticacid containing specimens (race 2) were also
found for this species. The two chemical races
were inseparable morphologically and this chemical difference seems to have no taxonomicvalue.
Ramalina intermedia resembles R. pollinaria(Westr. ex Ach.) Ach., a saxicolous species widely distributed in the world, in having granularsoredia covered by outer layers (Kashiwadani andMoon 2003). However, it can be readily distinguished from the latter by the narrower laciniae
and by the absence of evernic and obtusatic acids
which are constant components for R. pollinaria.
It might be confused with R. polymorpha (Ach.)
Ach., a species widely distributed in the Northern Hemisphere, which differs in having nochemical substances.
During the present study, we found this species

Noteworthy Species of the Genus Ramalina (Ascomycotina: Ramalinaceae) in China

163

Fig. 1. Ramalina intermedia (Del. Ex Nyl.) Nyl., a) a plant growing among mosses (Z. Congfu 3487, IFP,bar1 mm), b) plants growing on exposed rock (H. Kashiwadani 39240, TNS, bar5 mm).

from three localities in China. They coincide wellwith the exsiccate specimens of this species preserved in TNS both in morphology and chemistry. The specimen from Mt. Changbai, Jilin
Province, was collected among mosses on rocks
and determined as the race 1. The specimensfrom Ping Shui Shan, Jiansi Province were collected on exposed rocks along river and determined as the race 2. This is a new report for thisspecies from China.
Specimens examined. (Race 1) China. Prov.
Liaoning: Laotie-shan, on rock, X. Chen et al.
5772 (IFP). Prov. Jilin. Mt. Changbai, among
mosses on ground, around the lake near the summit, elevation about 2150 m, August 30, 1983 m,C. Zhao 3487 (IFP).
(Race 2) China. Liaoning Prov.: Snake Island,Dalian, on rock around the summit, May 13,
1973, Zhang Lizhu 016-1 (IFP). Jiang Ci Sang:
Ping Shui Shan, Jing Gang Shan Nature Reserve,
on rocks, elevation 900 m, July 12, 1995, H.
Kashiwadani 39240 & 39323 (TNS).
Exsiccata examined. M.E. Hale: Lich. Amer.
Exs. 171 (race 1, TNS). V. Räsänen: Lich. Fenn.
Exs. 28 (race 1, H). W.A. Weber: Lich. Exs.
Colol. 605 (race 1, TNS).
Ramalina obtusata (Arnold) Bitt., Pringsheim
Jahrb. Wiss. Bot. 36: 435 (1901).
(Fig. 2)
Ramalina obtusata is characterized by corticolous habit, shrubby and fistulose thallus growing from a narrow holdfast, inflated and more orless pellucid thalli without side branches, sparseand small fenestrations, continuous medullary
hyphae without isolated clumps of hyphae andthe presence of evernic and obtusatic acids as
major chemical substances.
Historical review for this species has been provided by Krog and James (1977). Although fertile specimens for this species has not been reported, one specimen (Fig. 2) from Mt. Chanbai,Prov. Jilin has characteristic features written
above, which coincide well with those of the lectotype of R. obtusata (Arnold: Lich. Exs. 577b—
isolectotype in BM!). New to China.
Specimen examined. China. Prov. Jilin: 14 km
from Baishan Station, Mt. Changbai, on bark of
Picea yezoensis, elevation 1270 m, June 19,1980, C. Zhao 0320 (IFP).
Ramalina pentecostii Krog & Swinscow, Norw.
J. Bot. 23: 167 (1976).
(Fig. 3a, 3b, 3c)
Ramalina pentecostii is characterized by its
corticolous or saxicolous habit (Krog & Swinscow 1976), weakly branched subpendulous
laciniae showing bilateral appearance (Fig. 3a),marginal soralia (Fig. 3b) and shortly linear
pseudocyphellae, smooth chondroid layer without cracks (Fig. 3c) and by the absence of
medullary chemical substance. Apothecial laminal, 0.8–1.5 mm in diameter; disc with whitemargin; ascospores shortly fusiform, 11–125–6m m in size.
Ramalina pentecostii is a poorly known lichen

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Fig. 2.

Hiroyuki Kashiwadani et al.

Ramalina obtusata (Arnold) Bitt., showing a habit (C. Zhao 0320, IFP). Bar1 mm.

Fig. 3. Ramalina pentecostii Krog & Swinscow (Z. Congfu 2501, IFP). a) habit (bar1 cm), b) laciniae showing
bilateral appearance by marginal soralia (bar0.5 mm), c) cross section of branches, showing smooth chondroid tissue (bar100 m m).

Noteworthy Species of the Genus Ramalina (Ascomycotina: Ramalinaceae) in China

and it has been so far known only from Uganda,
East Africa (Type collection: Uganda, Toro,Ruwenzori, R.E. Pentecost 115—holotype inBM!). In the course of the present study, we
found several specimens comparable to thespecies collected in Mt. Tianshan, Xinjiang Prov.,
northwest China. These specimens are quite
identical morphologically as well as chemicallywith the type specimen. One of them (Z. Congfu
25019 is fertile and the description of apothecia
is given above, though Krog and Swinscow(1976) reported only sterile specimens.
This species is unique in having dorsiventral
laciniae with marginal continuous soralia and in
producing no chemical substances. This characteristic lichen is not easily mistaken for any otherspecies of Ramalina found in Asia. Specimens ofsubpendulous thalli of R. farinacea (Pers.) Ach.,a corticolous widespread species in temperate regions in the world, might be confused with thisspecies, but it differs in having repeatedly
branched laciniae with discrete soralia and in
producing depsidones such as protocetraric,
norstictic or salazinic acids as major chemical
substances.
This is the second record for R. pentecostii in
China. All specimens from China were collectedon branches of Picea sp., at elevations between
1900 to 2260 m.
Specimens examined. China. Prov. Xinjiang,Mt. Tianshan, Bayi forest centre, Picea sp.,
2260 m alt, August 31, 1982, C. Zhao 2242(IFP); the same locality, on dead wood, 1900 malt, September 1, 1982 (IFP); Takesi forest centre, on Picea sp., 1950 m alt., September 24,1982, C. Zhao 2501 (IFP); Zhaosu forest centre,
on Picea sp., 2260 m alt, September 26, 1982, C.
Zhao 2480 (IFP); Gongliu forest center, September 29, 1982, on Picea sp., 1900 m alt., September 29, 1982, C. Zhao 2629 (IFP).
Ramalina pertusa Kashiw., Mem. Natn. Sci.
Mus. 18: 102. (1985)
Although Ramalina pertusa has been reported
widely distributed in Japan and Taiwan (Kashiwadani 1985, Kashiwadani et al. 2006), it has165

never been reported from mainland China. One
specimen collected at Huanre, Prov. Liaoning, is
now identified as this species. New to China.
Specimen examined. China. Prov. Liaoning:
Huanren County, on dead twigs, 800 m, September 11, 1986, X. Chen & S. Liu 6702 (IFP).
Ramalina pollinaria (Westr. ex Ach.) Ach.,
Lich. Univ.: 608 (1810).
Basionym. Lichen pollinarius Westr., Kgl.
Vitensk. Acad. Nya Handl. 16: 56 (1795). Typecollection: Suesia (H-Ach. 1831D!).
Ramalina pseudosekika Asahina, J. Jpn. Bot.
16: 139 (1940). Type collection: North Korea,
Prov. Pyongannam-do, Gija-rim, Pyongyang,1930, H. Den s.n. (herb. Asahina 334—holotypein TNS), syn. nov., tlc: usnic, evernic and obtusatic acids.
Ramalina pollinaria is a well-known speciesof the genus characterized by the saxicolous
habit, solid and sparingly branched laciniae with
granular soredia, narrowly fusiform ascospores(12.5–143.5–4.5 m m in size) and by the presence of evernic and obtusatic acids as major
medullary substances.
Asahina (1940) described Ramalina pseudosekika Asahina from North Korea. The holotype specimen, however, is quite identical morphologically and chemically with the holotype ofR. pollinaria and can be simply reduced to a synonym of the latter species.
R. pollinaria seems to be widely distributed inChina.
Representative specimens examined. China.
Prov. Hebei, Weixian Co: Mt. Xiaowutai, onrock, 1700 m alt., August 15, 1982 Liu Shun 79(IFP). Prov. Heilongjiang: Laotudingzi hill Dahailin, on rock, August 28, 1973, Gao Qian
9218-1 (IFP); Laoli hill, Wudalianchi, on rock,300 m alt., X. Chen 3518 (IFP). Prov. Jilin, Helong Co: Shiliping, on rock 700 m alt., September 12, 1983, C. Zhao 3958 (IFP); north hill,
Ji’an town, on rock 120 m alt., September 13,1963, X. Chen 2469 (IFP); Ice field, Mt. Changbai, on rock, 1750 m alt., August 25, 1977, X.
Chen 4996 (IFP). Prov. Liaoning: Fengcheng

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Hiroyuki Kashiwadani et al.

Co.: Pheonix Hill, July 22, 1984, Gao De’en &Liu Shun 0252 (IFP); Kuandian Co.: Fotanggou,Baishilazi Reserve, on rock, 1000 m alt., November 18, 1988, X. Chen 7058 (IFP). Prov. Shandong: Mt. Laoshan, October 16, 1985, on rock,
X. Chen & Liu Shun 226 (IFP).
Ramalina sekika Asahina, J. Jpn. Bot. 16: 138(1941)
Ramalina sekika is a saxicolous species similar to R. pollinaria, which differs in having evernic and obtusatic acids as chemical substances. Itis apparently one of a rare species of the genus,
having been so far known only from the type collection in China (Type collection: Laotie-shan,Port Arthur, Prov. Liaoning, China, July 19,1940, Y. Asahina 333—holotype in TNS!, tlc:
sekikaic, 4-O-demethyl sekikaic and salazinic() acids). In the present study, we found additional localities for this species in Provinces
Lianing and Jilin, China as cited below. This is
the second report for the species.
Representative specimens examined. China.
Prov. Liaoning: Xianrendong, Zhuanghe City, onrock in Pinus densiflora forest, X. Chen et al.
5801 (IFP); Laotieshan, Lushun, on rock, September 1, 1931, Y. Kobayasi 14 (TNS); Huangnichuan, Lushan, on rock, March 19, 1963, X.
Chen 1191 (IFP); Huishan Hill, Shenyang, April5, 1973, X. Chen s.n. (IFP). Prov. Jilin: Badaogou, Changbai Co., on rock, 450 m alt., Septem-ber 5, 1963, X. Chen 2340 (IFP).
Acknowledgements
We are most grateful to Dr. S. Kurokawa,
Botanic Gardens of Toyama for his critical reading of the manuscript. This study was supported
in part by JSPS KAKENHI to H. Kashiwadani(no. 17570089) and MEXT KAKENHI to H.
Koyama (no. 07041151).
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