Bull. Natn. Sci . Mus ., Tokyo, Ser. B, 27(1), pp. 11 ~21 , March 22,2001

Taxonomic Notes on Rhizocarpon Growing in
Solfatara Fields in Japan
Masakane lnoue
Department of Natural and Environmental Sciences,
Faculty of Education and Human Studies,
Akita University, Akita, 010-8502 Japan

Abstract Five species of Rhizocarpon growing in a solfataric field are reported forJapan. R. vulcani Mas. Inoue is new to science. R. oederi (Weber) Korber is new tothe flora of Asia. Taxonomic and chemical data are provided.
Key words:

lichens, taxonomy, Rhizocarpon , active volcanic area, solfatara, JapanThe Japanese archipelago has numerous volcanoes. It is known that the vegetation near solfataras in the volcanic areas are severely affected by sulphurous gases including H 2 S, S0 2 and others, and that characteristic communities such as Cladonia
vulcani - Descampsia ftexuosa-community and Caricetum angustisquamae-community are developed. There exist some ecological studies on lichens in this harsh habitat of Japan (Glime and Iwatsuki , 1990; Fahselt, 1995; Inoue, 1996; etc.), but mostcrustose lichens growing there are still unknown. In this paper I will present somenotes on five species of Japanese Rhizocarpon growing near solfataras; R. atrobrunnescens, R. badioatrum , R. lavatum , R. oederi and R. vulcani.
Nylander (1890) added Rhizocarpon atrobrunnescens, R. badioatrum (asLecidea atroalba) and R. lavatum to the lichen flora of Japan based on the collections
from Mt. Fuji , and the mountain was the only locality of these lichens previously
known and their existence in Japan were not reconfirmed except for R. badioatrum by
Fahselt (1995). The type and authentic specimens of possibly related species fromextra-Japan have been restudied for purposes of comparison.
In the list of specimens examined, the locality is followed by the altitude, accession number and the date collected. The specimens listed here are preserved in theherbarium of Akita University and in the herbarium of the National Science Museum,
Tokyo (TNS).

1)

Rhizocarpon atrobrunnescens (Nyl.) Zahlbr.
(Plate I: a)Cat. Lich. Univ. 4: 347 (1926).- Lecidea atrobrunnescens Nyl., Lich. Jap. p.
82 ( 1890). Type: Japan , Mt. Fuji "in limite arborum", - holotype in H (H-Nyl.
9893), vidi.
Thallus dark brown, polished, peltate or subsquamulose-areolate; areolae sub-

12

Masakane Inouescattered, with epinecral layer, more or less round, concave with blackish margin . Hypothallus black, visible between the areolae.
Apothecia reaching a diameter of 0.5 mm , black, appressed-adnate or subimmersed in the thallus; disc epruinose, plane or convex, with somewhat thick margin;
margin almost level with the disc surface. Excipulum thin , 30- 50 J..lm thick, darkbrown, paraplectenchymatous; hyphae obliquely subradiating, with a thin wall. Epithecium blackish green. Hymenium 120--150 J..lm high. Subhymenium colorless,15- 20 J..lm high. Hypothecium dark brown, reaching to !50 J..lm high. Paraphyses
branched, anastomosed. Asci clavate, I 00- 130 X 35---45 J..lm, 8-spored. Spores(30- )32---40(---42) X 16- 18(- 20) J..lm , muriform , with 5- 7 transverse septa and 1- 2 longitudinal septa, colorless at first, soon becoming brown, with halo.
Reactions: thallus & medulla P-, K- , KC-, C-. Chemical substances: two unidentified minor constituents (Fig. I). Habitat: on rocks . Range: endemic to Japan.
Rhizocarpon atrobrunnescens is closely related to Rhizocarpon bolanderi(Tuck.) Herre, known from northern North America, Greenland and Scandinavia, because of having peltate or squamulose areolae which are dark brown and polished,
and of having numerous-celled muriform spores. However, R. bolanderi has asci producing I or 2 bigger spores (32- 68 X 18- 32 J..lm, Thomson 1997) per ascus .
Rhizocarpon atrobrunnescens is also known to occur in the areas where solfataras are absent.
(Solfataras are present in the locality) Specimens examined : HOKKAIDO. Prov.
Kamikawa: Anseikako "Ansei Crater" of Mt. Furano, 1400 m alt. , M. lnoue no.
7960b (23 Aug. , 1974). HONSHU. Pref. Nagano: Mt. Asama, 2400 m, M. lnoue no.
11342 (23 Aug., 1976), 2130 m, M. Inoue no. 11349 (23 Aug. , 1976). (Solfataras areabsent from the locality) Representative specimens examined: HOKKAIDO. Prov.
Kushiro: Mt. Oakan, 1350 m, M. lnoue no . 8781 (26 July, 1974). Prov. Kamikawa:
Mt. Midori ofMts. Daisetsu, 1840m, M. Inoue no. 11664 (8 Aug., 1974).

2) Rhizocarpon badioatrum (Florke ex Spreng.) Th. Fries(Plate I: b)
Lich. Scand. 613 (1874).- Lecidea badioatra Florke ex Spreng., Neue Entdeck. 2: 95 (1821). Type : Europe?, non vidi. - Lecidea atroalba Ach. ; Nylander,
Lich. Jap. 82 ( 1890). - Rhizocarpon badioatrum var. atroalbum ; Zahlbr. , Cat. Lich .
Univ. 4:326 (1927); Sato, Cat. Lich. Jap. (ed. 2) 47 (1965).
Thallus brown or pale brown, irregularly areolate; areolae plane or slightly bullate, dispersed on a black hypothallus, or in part, contiguous. Hypothallus prominent, black, visible between the areolae.
Apothecia black, reaching a diameter of 1.0 mm, between or partly on the areolae, appressed-adnate, somewhat thickly marginate; margin almost level with the discsurface; disc epruinose, plane or convex. Excipulum thin , 30- 50 J..lm thick, darkbrown with violet tinge, paraplectencymatous; hyphae obliquely subradiating, with athin wall. Epithecium brown or pale brown, K + violet. Hymenium I 00- 120 J..lm

Taxonomic notes on Rhizocarpon

13

Rf-value

1.0-

Solvent sytem A

0

0.6-

c::>

-

0
0

0

0.2 -

I

1

3

2

4

Rf-value

1.0

Solvent sytem B

0.6-

0

-

=
c::::>

0.2 1

.

2

0

C)

3

4

Fig. 1. Chromatograms of Rhizocarpon atrobrunnescens in so lvent systems A and B comparing
with norstictic ac id, etc . 1. atranorin; 2. Rhizocarpon atrobrunnescens ; 3. norstictic acid; 4.
stictic acid. (solvent system A: a mixture of 180 ml of benzene, 45 ml of dioxane, and 5 ml ofacetic ac id; so lvent system 8 : a mixture of 100 ml of n-hexane , 80 ml of ethyl ether, and 20 mlof formi c acid.)

high. Subhymenium 15- 20 pm high, pale brown or colorless. Hypothecium concolorous to the exciplum , reaching to 150 pm high. Paraphyses branched, anastomosed.
Asci clavate, 80- 100X20- 35pm, 8-spored. Spores colorless at first , soon becomingbrown, !-septate, with halo, 20- 23 X 11 - 13 pm.
Reaction : thallus & medulla P-, K- , KC- , C-. Chemical substances : two unidentified minor constituents (same as those of R. atrobrunn escens, Fig. I). Habitat: onrocks . Range: Japan; Italy (Nimis & Poelt, 1987); Germany (Feuerer, 1978); Sweden(Ma1me, 1914); Svalbard Isls. (Elvebakk & Hertel , 1997); Greenland (Lynge, 1934;
Thomson, 1997); American arctic (Thomson, 1997); Australia (Filson, 1996); India(Schubert & Klement, 1966)
Rhizocarpon badioatrum is most remini scent of Rhizocarpon hochstetteri(Korb.) Vainio, known from northern Hemisphere including Japan (Inoue, 1997),

14

Masakane Inoue

which resembles in having similar thallus and spore morphology. However, R.
hochstetteri has spores which are always colorless except for those of very old stage
and producing stictic acid.
Kiimmerling ( 1991) reported stictic acid from R. badioatrum , and Follmann &Huneck ( 1971) reported an unidentified phenolic substance. On the contrary Japanese
representatives show different chemistry. There might be exsisting some chemotypes
in R. badioatrum.
Nylander (1890) reported Lecidea atroalba "Ach." from Japan (Fuji-Ichigome,on lava), which Sato ( 1965) mentioned as a synonym of Rhizocarpon badioatrum var.
atroalbum "(L.) Malme" when he summarized the Japanese lichens known up to1958 with no attempt at taxonomic revision and was based chiefly on Zahlbruckner's
Catalogus, but unfortunately I could not find out any voucher specimen from Nylander Herbarium at Helsinki University both on 1977 and 1994.
Rhizocarpon badioatrum is also known to occur in the areas where solfataras areabsent.
(Solfataras are present in the locality) Specimens examined: HOKKAIDO. Prov.
Kamikawa: Ansei Kako (Ansei-Crater) of Mt. Furano, 1400 m, M. Inoue no. 7960 a(23 Aug., 1974). Prov. Nemuro: Kamuiwakka of Shiretoko Pen., 740 m, coil. D. Fahaselt (H43) , M. Inoue no. 23558 (17 Oct., 1989); Mt. Kawayu-iouzan , 150 m, M.
Inoue no. 9039 (15 Aug., 1974). Prov. Kushiro : Mt. Meakan, 1250m, coli. D.
Fahaselt (H38) , M. lnoue no. 23561 (14 Oct. , 1989); 1255m, coil. D. Fahaselt (H39) ,M. Inoue no. 23562 (14 Oct. , 1989); 1260m, coli. D. Fahaselt (H29, 37), M. lnoue
nos. 23559, 23560 (14 Oct., 1989). Prov. Shiribeshi : Niseko Heights, 900- 1000 m,
M. Inoue no. 24679 (16 July, 1984). HONSHU. Pref. Gunrna: Mt. Motoshirane, 20002140m, M. Inoue no. 24462 (3 June, 1984). (Solfataras are absent from the locality)
Representative specimens examined: HOKKAIDO. Prov. Abashiri: Mt. Rausu of
Shiretoko Pen. , 1650 m, M. Inoue no. 7750 (23 July, 1974). Prov. Kushiro: Mt.
Oakan, 1350 m, M. lnoue no . 8755 (26 July, 1974). Prov. Tokachi: Mt. Yuniishikari Mt. Otofuke, 1640 m, M. Inoue no . 8377 (4 Aug., 1974). HONSHU. Pref. Yamanashi: Mt. Kitadake , 3050m, M. Inoue no. 12193 (19 Aug., 1978). Additional specimens examined: SWEDEN, Vastmanland, Arboga, Sfielund, on a boulder, coli. G.
Kjellmert, s.n. (25 March, 1953), TNS. AUSTRIA, Osztaler Alpen, Nord tiro! ,Kaunergrat, 2220- 2230 m, coli. H. Doppelbaur & A. Schroppel (Aug., 1961 ), Lich.
Alpium no. 169, TNS.
(Plate I: c)3) Rhizocarpon lavatum (Fr.) Hazslin
Magy. Birod. Zuzumo-Fior. , 206 (1884). - Lecidea atroalba var. lavata Fries,
Nov. Sched. Critic. p. 18 ( 1827). Type: Sweden, Smaland, Femsjo, coli. E. Fries, nonvidi.-Lecidea lavata Ach. ; Nyl. Lich. Jap. p. 82 (1890).
Thallus thin to medium , ochraceous or in part brownish gray, irregularly
cracked-areolate; areolae minute, contiguous, or in part more or less dispersed, sur-Taxonomic notes on Rhizocarpon

15

face mat. Hypothallus indistinct or forming a black hypothalline line.
Apothecia up to 0.7 mm wide, black, adnate, well constricted at the base; discepruinose, flat with a prominent entire margin at the juvenile stage, then becomingslightly convex with + I - obliterated margin, with umbo. Excipulum up to 70 11m
thick, dark brown, paraplectenchymatous; hyphae perpendicularly radiating 5- 7 11m
thick, with a thin wall. Epithecium brown. Hymenium (110)120-140(150)/lm high.
Subhymenium 15-20 11m high, pale brown. Hypothecium reaching to 150(- 200) 11m
high , dark brown. Paraphyses slender, 2- 31-lm thick, well branched, anastomosed;
apices not swollen. Asci clavate, 105- 130X25- 30 11m, 8-spored. Spores 23- 33X
12- 20 11m, with halo, colorless, muriform, with 4- 6 transverse septa and l- 2 longitudinal septa.
Chemical reactions: thallus & medulla P-, K-, KC- , C- ; Chemical substances: nolichen substances demonstrated on TLC. Habitat: on rocks close to running water andare temporary inundated. Range: Japan; Europe (Feuerer, 1991; Purvis et al. , 1992;
Santesson, 1993); North America (Wong & Brodo, 1992); Svalbard Isls. (Elvebakk &Hertel, 1997)
Rhizocarpon lavatum is distinctive by the ochraceous or brownish gray thallus,
adnate apothecia with umbonate disc, the colorless muriform spores and by the peculiar habitat. This species is known in Japan only from three present gatherings, but
the materials are well developed, and actually this species is forming large communities at the site.
This species is most reminiscent of Rhizocarpon oederi, since these two species
have a similar thallus , apothecia and a chemistry. However, R. oederi has a lower hymenium (up to 100 11m high) and smaller parallel 3-septated spores (18-20 X8- l01-lm).
Rhizocarpon lavatum is known as a lake- and stream-side species (Purvis et al.,
1992; Wong & Brodo, 1992, etc.). The Japanese representatives are also occurring inthe similar habitat where are temporary inundated (Plate 11: b). But Sukawa, where is
the only known locality of this species at present, is situated on a volcanic area whereabundant active solfatars are formed among or on lavas, and the running water near
the habitat is acidic and contains extremely high level heavy metals on the contrary toa mention of Purvis et al. (1992) "on siliceous rocks subject to inundation by unpolluted streams and lakes". I measured the pH values and the electroconductivity,
which is one of the parameters of salinity, of the stream water close to the habitat at
Sukawa; pH value is 2. 72-2.78 and the electroconductivity is 11 08- 11141-LS/cm.
Nylander (1890) reported this species (as Lecidea lavata) from Japan (Mt. Fuji,
Oumagayeshi - ltchigome ), but unfortunately I could not find out the voucher specimen from the Nylander Herbarium at Helsinki University on both 1977 and 1994.
Specimens examined. HONSHU. Pref. Akita: Sukawa, on the mountain side of
Mt. Kurikoma, 1080-1100 m, M. Inoue nos. 25773 & 25774 (15 Sept., 1997), 27632( 18 June, 2000). An additional specimen examined. Sweden, Bohuslii: Skafto, Klu-

16

Ma sakane lnouebban, coli. A. H. Magnusson , s.n. ( 16 July, 1922), TNS.

4)

Rhizocarpon oederi (Weber) Korber(Plate 1: d)Parerga Lich., p. 232 ( 1861 ). - Lichen oederi Web., Spicil. Flor. Goettingens.,
p. 182 ( 1778). Type: Germany, Harz, non vidi .
Thallus contiguous, thin to medium , ochraceous, irregularly cracked-areolate,surrounded by brownish hypothalline lines; areolae angular, flat , smooth , more or lesspolished.
Apothecia up to 0.5 mm wide, black, subimmersed to appressed-adnate on thethallus or among the areolae, not constricted at the base; margin prominent; di sc
epruinose, with umbo . Excipulum up to 60 J.lm thick, paraplectenchymatous, darkbrown . Epithecium blackish green or paler. Hymenium (60)70-90 J.lm high. Subhymenium 15- 20 J.lm high , pale brown. Hypothecium reaching to I 00( 150) J.lm high ,dark brown. Paraphyses slender, 2- 2.5 J.lm thick, well branched, anastomosed. Asciclavate, 50- 60X 15- 20 J.lm , 8-spored. Spores colorless, oblong, with halo , 3-septated,
16- 21 X7- IO,um.
Reaction : thallus & medulla P-, K-, KC- , C-. Chemical substances: no lichen
substances demonstrated on TLC. Habitat: on rocks. Range: Japan; Svalbard (Elvebakk & Hertel , 1997); North & Central Europe (Feuerer, 1978 & 1991 ; Santesson,
1993 ; Schade, 1935); North America (Thomson, 1967; Wong & Brodo, 1992).
Diagnostic characteristics for this species are: the ochraceous thallus which issurrounded by brownish hypothalline lines, the subimmersed to appressed-adnate
apothecia with umbo , and the colorless 3-septated spores.
Yamada Pass, where is one of the known localities ofthis species, is not situatedon the site close to the so lfataras but on the site about 1.3 km northwest of the craterof Mt. Shirane, where numerous so lfataras are formed, being affected by sulphurous
gasses.
Thi s species was previously known only from Arctic region, Europe and North
America. However, the range has now exended to Asia.
(So lfataras are present near the sites) Specimens exam ined. HONSHU. Pref.
Nagano : Yamada Pass in Mt. Shirane, 2060 m, M. Inoue nos. 1764 b & 11890 (20July, 1972), 2100 m, M. Inoue nos. 24745b, 24746 & 24750 (2 June, 1984). (Solfataras are absent from the site) Specimens examined. HOKKAIDO. Prov. Tokachi: Mt.
lshikari , 1860 m, M. Inoue no. 8470 (5 Aug. , 1974). Additional specimens examined.
Slovakia, Carpates, 500 malt. co li. Ki sze ly & Vezda (30 Aug. , 1974), Vezda, Lich se l.
Exs. no . 1295, TNS. Finland, "Tavasti a meridionali, Ronni ", coli. C. Leopold (in1876), Norrlin & Nylander's Exs. no. 343 , TNS .
5)

Rhizocarpon vulcani Mas . lnoue, sp. nov.
(Pl ate II : a)Thallus crustaceus, epilithicus, modice incrassatus aut sat tenuis , atrofuscescens,irregulariter areolatus aut verrucoso-areolatus; areolae minutae, leviter convexae, di s-Taxonomic notes on Rhizocarpon

17

persae aut discretae. Hypothallus nigricantus. Apothecia ad 0.5 mm lata, tota adnata,ad basin leviter constricta, disco vulgo piano, saepius umbonata, nigra, nudo, margine prominente, persistente. Excipulum 30- 50 Jlm latis, fusco-graucescens. Epithecium aeruginoso-fuligineum. Hymenium (90-)100- 120 Jlm crassum. Subhymenium
15- 20 Jlm crassum, decoloratum aut pallidum. Hypothecium totum fusco-nigrum, ad
I 00 Jlm crassum. Paraphyses graciles, passim ramoso-connexae. Asci clavati. Sporae
8-nae, ellipsoideae, decolores , murales, 22- 26 Jlm longae, 11-15 Jlm crassae.
Thallus dark brown, thickness medium , irregularly areolate or verrucose-areolate; areolae minute, surface mat, plane or slightly convex, or at times granulated,
more or less dispersed on a black hypothallus, or in part, contiguous; medulla I-. Hypothallus black, visible between the areolae.
Apothecia up to 0.5 mm wide, adnate or appressed-adnate, black, between or
partly on the areolae , slightly constricted at the base; disc epruinose, plane, at times
with umbo ; margin prominent, thick. Excipulum thin, 30- 50 Jlm thick, dark brownwith green tinge, paraplectenchymatous; hyphae obliquely subradiating, with a thinwall. Epithecium blackish green. Hymenium (90-) I 00- 120 Jlm high. Subhymenium
15- 20 Jlm high, colorless or pale brown. Hypothecium dark brown, reaching to
I 00 Jlm high; hyphae irregularly arranged. Paraphyses slender, 1.5- 2.5 Jlm thick, wellbranched, anastomosed; apices not or slightly thickened. Asci clavate, 80-100 X 2535 Jlm , 8-spored. Spores 22- 26 X 11 - 15 Jlm, colorless, ellipsoid, muriform, with 3 (orrarely 4) transverse septa and I longitudinal septa, with halo.
Reaction : P-, K- , KC- , C-. Chemical substances : no lichen substances demonstrated on TLC. Type: Japan , Honshu, Pref. Akita , Senpoku-gun, Tazawako-machi,
Tamagawa spa, 760 malt. , on rock , 15 Oct. , 1983 . M. Inoue no. 16569, - holotype
in TNS . Habitat: on rocks in volcanic areas where abundant solfataras are spouting
acidic gases. Range: Endemic to Japan.
Rhizocarpon vulcani reminds Rhizocapon grande (Fiorke ex Flotow) Arnold,
which is one of the well known species in northern Hemisphere and New Zealandand is known from the Shirakami Mountains of Japan (Inoue & Moon, 1996), because of having dark brown thallus of which areolae are dispersed on a black hypothallus and of having muriform spores. However, R. grande can be easily distinguished by having dark brown bigger spores (30- 3 5 X 16- 18 Jlm) with 4 to 7 transverse septa and I to 2 longitudinal septa, and by containing gyrophoric acid as amajor substance.
Spores of most samples of R. vulcani including type specimen always colorless,however, a few ones have pale brown spores which are old and changing from normal
shape.
Rhizocarpon vulcani is distributed from Hokkaido to Kyshu and is occurring involcanic areas where solfataras are spouting hot smokes and gases; perhaps it might
be one of the common lichens growing in this kind of peculiar habitat like Cladonia
vulcani. At the Tamagawa spa area Inoue ( 1996) reported the pH value 3.5 to 4 .6

18

Masakane lnoue

from the water remaining just after a rainfall in small hollows on the rocks, on which
R. vulcani is growing (Plate ll: c).
Specimens examined. HOKKAIDO. Prov. Shiribeshi: Ooyu-numa, Noboribetsu spa, 230m, M. Inoue no. 24739 (20 July, 1984), Niseko Heights, 900- IOOOm,
M. lnoue nos. 24683 & 24686 (16 July, 1984). Prov. Iburi: Mt. Nisekoan-nupuri,
1150 m, coil. H. Kashiwadani no. 15588 (20 Sept., 1979), TNS. HONSHU. Pref. Aomori: Shimokita Pen., Mt. Osorezan, lakeshore of Usori-ko, 210 m, M. lnoue no.
25610 ( 18 Oct. , 1996). Pref. Akita: Yugama in Mt. Yakeyama of Hachimantai,
1280 m, M. Inoue no. 25708 (16 May, 1998); Tamagawa spa, 760 m, M. Inoue nos.
23746,23748,23750,23751,23755-76 (18 July, 1995); Mt. Kurikoma, near Showaike, 1300 m, M. Inoue no. 25770 & 25777 (15 Sept., 1997); Mt. Yamabushi,
Kawarage-jigoku, 800 m, M. Inoue no. 27628 (5 May, 2000). Pref. Nagano: Mt.
Asama, Yunodaira, 20 I 0 m, M. lnoue no. 11325 (23 Aug. , 1976); Mt. On take,
2630m, M.Inoue no. 11497 (17 Aug. , 1976). KYUSHU. Pref. Nagasaki: Unzen spa
in Mt. Unzen, 600 m, M. Inoue no. 2555b (28 March, 1996). Pref. Miyazaki: Ebino
Heights (Oojigoku on the foot of Mt. Karakuni), 1350--1400 m, M. lnoue nos.
25493-25495 , 25497,25498,25506, 25527, 25529,25539 (27 March, 1996).
Acknowledgements
I am particularly indebted to directors and curators of the following herbaria forsending type and authentic specimens on loan: Drs. T. Ahti & 0. Vitikainen of
Helsinki University (H) and Dr. H. Kashiwadani (TNS). This study was partly supported by a Gramt-in-Aid for Scientific Research (C), Japan Society for the Promotion of Science to M. Inoue, no. I 0640676.
References
Elvebakk, A. & H. Hertel. 1997. A catalogue of Svalbard lichens. In Elvebakk, A. & P. Prestrud (eds.):
A catalogue of Svalbard plants, fungi , algae, and cyanobacteria. Norsk Polarinstitutt Skrifter, 198:271 - 359.
Fahselt, D. 1995. Growth form and reproductive character of lichens near active fumaroles in Japan.
Symbiosis, 18:211 - 231.
Feuerer, T. 1978. Zur Kenntnis der Flechtengattung Rhizocarpon in Bayern . Bet: Baye1: Bot. Ges ., 49:

59- 135.
Feuerer, T. 1991 . Revision der europaischen Arten der Flechtengattung Rhizocarpon mit nichtgelbem
Lager und vielzelligen Sporen. Bib/. Licheno/., 39: 1- 218, 1991 .
Filson, R. B. 1996. Checklist of Australian lichens and allied fungi. Flora of Australia supplementary series Number 7, pp. 204.
Follmann, G. and S. Huneck. 1971. Zur Phytochemie und Chemotaxonomie der Krustenftechten-Gattung
Rhizocarpon. Nova Hedwigia , 22:621 - 639.
Glime, J. M. and Z. Iwatsuki. 1990. Niche characteristics ofCladonia lichens associated with geothermal
vents in Japan. Ecological Research, 5: 131 - 141.

Taxonomic notes on Rhizocarpon

19lnoue, M. 1996. Lichens in acidic conditions. Biological Science (Tokyo) , 48: 25- 34. (in Japanese).
lnoue, M. 1997. Little known crustose lichens bearing lecideine black apothecia from Japan. Bull. Natn.
Sci. Mus. , Tokyo , ser. B, 23(2): 43- 58.
Inoue, M. & K.-H. Moon . 1996. Lichens of Shirakami-sanchi, Northern Japan. Scientific Studies of Shirakami-sanchi, pp. 233- 247. Association for National Park, Tokyo. (in Japanese)
Kiimmerling, H. 1991. Zur kenntnis der Flechtenflora am Hohen Meissener und=in seinem Yorland(Hessen) unter besonderer Beriicksichtigung chemischer Merkmale. Bib/. Lich. , 41: 1- 315.
Lynge, B. 1932. A revision of the genus Rhizocarpon (Ram.) Th. Fr. in Greenland. Skrifter om Svalbard
og Jshavet, 47 : 1- 30.
Malme, G. 0 . 1914. Yastra Jamtlands Rhizocarpon-Arten. Svensk Bot. Tidsk. , 8: 273- 294.
Nimis, P. & J. Poelt. 1987. The lichens and lichenicolous fungi of Sardinia (Italy). Studia geobotanica, 7(Suppl .): 1- 269
Nylander, W. 1890. Lichenes Japoniae. 122 pp., Paris.
Purvis, 0 . W., B. J. Coppins, D. L. Hawksworth, P W. lames and D. M. Moore (ed.). 1992. The lichen
flora of Great Britain and Ireland. Natural History Museum Publications in association with The
British Lichen Society. 710 pp., London.
Santesson, 1. 1993. The lichens and lichenicolous fungi of Sweden & Norway. 240 pp., Lund.
Schade, A. 1935. Die sachsischen Arten der Flechtengattung Rhizocarpon (Ram.) Th. Fr. (Beihefl) Bot.
Centralbl., 54: 75- 107.
Schubert, R. & 0. Klement. 1966. Beitrag zur Flechtenflora von Nord- und Mittelindien. Nova Hedwigia, 11: 1- 73.
Thomson, 1. W. 1967. Notes on Rhizocarpon in the arctic. Nova Hedwigia , 14:421-481.
Thomson, 1. W. 1997. American Arctic Lichens. 2. The Microlichens. The University of Wisconsin
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Wong, P. & I. Brodo. 1992. The lichens of Southern Ontario, Canada. Syllogeus, 69: l- 79.

20

Masakane lnouea. Rhizocwpon alrobrunnescens (Nyl. ) Zahlbr. (M. ln oue no. 7960 b). b. Rhizocarpon
badioalrum (Fliirke ex Spreng.) Th. Fri es (M. lnoue no. 7960 a). c. Rhi::ocalponlava/um (Fr.)
Hazs lin (M. lnoue no. 27632). d. Rhizocarpon oederi (Weber) Kiirber (M. ln o ue no. 1764 b).
X IO

Pl ate l.

Taxonomic notes o n Rhizoccupon

Pl ate 11 . a. Rhizocarpon vu/cani Mas. lno ue (M. lno ue no. 16569, ho lotype). X 10 . b. Habitat o f
Rhi::ocmpon /avarwn (a rrow) where is c lose to running water and is temporary inundated.
(S ukawa of Mt. Kuri koma). c. The type loca lity o f Rhi::ocarpon vulcan i (Tamagawa-Onsen"s pa" where active so lfata ras are spoutin g hot smo kes and gases).

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