Bull. Natn. Sci. Mus., Tokyo, Ser. B, 26(4), pp. 139- 153, December 22, 2000

Japanese Species of Lecidella (Lichenes, Lecanoraceae) (II)'l
Masakane Inoue
Department of Natural and Environmental Sciences,
Faculty of Education and Human Studies, Akita University,
Akita 0 I 0- 8502, Japan
E-mail: ebinoue@ipc.akita-u.ac.jp

Abstract Five species of Lecidella are reported for Japan. L. elaeochroma var.
caesitia (Va in .) Mas. lnoue is a new combination. L. elaeochromoides and L. euphorea are new to the flora of Japan. Taxonomic and chemical data are provided.
Key words:

lichens, taxonomy, Lecidella , Japan

4) Lecidella elaeochroma (Ach.) Hazsl. , Magy. Birod. Zuzmo-Fior, 197 (1884)
Lecidella elaeochroma has a world-wide distribution, but has never been adequately typified . Three sheets of Lecidea elaeochroma, which together consist of 13
fragments collected from Gallia, Svecia and Lapponia, are preserved in the Acharian
Herbarium at Helsinki (H-Ach.). In the original description , Acharius (1803) did notcite any locality except "Habitat in cortice Alni", so l could not select any relevant
type specimen from the Herbarium in 1977 and 1994.
Poelt & Yezda ( 1981) recognized two varieties under this species on the basis ofthallus color; one is var. elaeochroma which is "Lager weisslich bis grau odergelblichgrau, graugriin ohne mit gelblichen Ton", and the other is var. flavicans
which is " Lager schwefelgelb bis gelbgri.in". On the basis of such a distinction,
Japanese specimens were the latter variety. An examination of the holotype of var.
flavicans (as Lecidea anomala v L. flavicans (H-Ac h. 326D, in H, vidi), however, revealed another important character, that is , var. flavicans has rimulose thalli (Fig. II). In contrast, Japanese representatives have a verrucose-granulate or verrucose-leprose thallus (Fig. l-2a, 2b), and therefore, they are not assignable to var.flavicans.
Since careful typification of L. elaeochroma and much more understanding of
the many other infra specific taxa would be necessary before a final decision could bemade , l have considered it preferable to maintain the var. caesitia described from
Hokkaido, Japan by Yainio ( 1921 ).

11Continued from Bull. Natn. Sci. Mus. , Tokyo, Ser. B, 23(4): pp. 127- 13 6, 1997.

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Masakane lnoue

I. Ho lotype of Lee idea a noma/a v L. flavicans A eh. (H-Ach. 3260) , ea. X I 0. 2a & 2b.
Lecidella e/a eochroma var. caesitia (Vain.) M as. lnoue (M. lnoue no . 9061 ). b. X I 0. 3.
Lecidella e/a eochromoides (Nyl.) Knoph & Hertel (M. lnoue no.16870) , X20. 4a & 4b.
Lecidel/a enteroleucella (Ny l. ) Herte l. a. (M . lno ue no. 7469). b. (M. lno ue no. 3444), X 10.
5. Lecidel/a euphorea (F iiirke) Hertel (M. lnoue no. 82 11 ). 6a & 6b. Lecidella sendaiensis(Zahlbr.) Knoph & Le uckcrt (M. lno uc no. 14833). b. X I 0.

Fig. I.

Lecidella elaeochroma (Ac h.) Choisy var. caesitia (Va in. ) Mas. lnoue, comb. nov.
(Fig. 1-2)
Basionym. Lee idea olivacea var. caesitia Va in. , Bot. Mag. , Tokyo 35: 69 ( 1921 ).
Type: Japan, Hokka ido, Prov. Kushi ro, Akkeshi, leg. A. Yasuda 334,- holotype inTUR ( H- Va in . 23755), vidi.- Lecidea elaeochroma var. caesitia (Va in.) Zahlbr. , Cat.
Lich. 3 : 57 1 ( 192 5).- Lecidea parasema a uct. non (Ach.) Ac h.; Mull. Arg., Nuov.
G iorn . Bot. !tal. 24 : 197 ( 1892).- Lecidea olivacea var. euphoreoides auct. non

Japanese Species of Lecidella (11 )141(Vain.) Vain. ; Vain. , Bot. Mag., Tokyo 32: 160 (1918), 35: 68 ( 1921 ).-Lecidea
spumosula Zahlbr. , Ann . Mycol. 14: 51 ( 1916), nom. illegit. , Art . 64, non Lee idea
spumosula Yasuda 1915 ( = Bacidia spumosula (Yasuda) Yasuda 1925).
Thallus effuse, thin to medium, continuous, verrucose-granulate or verrucoseleprose, sometimes granulose-diffracted, greenish gray with yellow or sulfurous
tinge, bordered by a + I - distinct black hypothalline line; medulla l-.
Apothecia up to 1.5 mm in diameter, adnate, black, moderately or prominently
constricted at the base; disc epruinose, plane w ith a prominent entire margin at thejuvenile stage, then becoming convex with an obliterated flexuous margin; margin
sometimes polished and paler than the disc. Excipulum 50- 80(- 100) pm thick , dirty
violet externa lly, the interior part violet-brown; hyphae perpendicularly radiating,5- 6 pm thick , with a thick wall. Epithecium blue-green or pale blue green. Hymenium (60- )80-1 00(- 120) pm high , sometimes interpenetrated with oily granules. Subhymenium reaching to 250 pm high , with irregularly shaped cavities, violet-brown ,composed of perpendicular hyphae. Paraphyses simp le or rarely branched, lax , slender, 1.5- 2 pm thick ; apices not or slightly clavate. Asci 50- 80(- I 00) X 13-15 ,u.m,clavate; tholus well advanced, I+ intensive blue. Spores with obtuse ends,(8- ) I 0- 16 X 5- 9(- 10) ,urn.
Reaction: thallus P+ yellow (faintly), K+ yellow, KC -, C-; medulla P+
yellow (faint) , K + yellow, KC -, C+ orange. Chemical substances: atranorin,
arthothelin ( + / -),and thiophanic acid.
Habitat: On bark in the lowlands to mountain regions.
Range: Known only from Japan .
Lecidella elaeochroma var. caesitia is characterized by a verrucose-granulate orverrucose- leprose thallus, viol et- brown excipulum as in the case of subhymenium,
and atranorin and xanthones ( c + ).
This variety is certa inly very near to saxicolous L. asema, but is distinguishedfrom it by a violet-brown subhymenium, since that of L. asema is golden brown.
Vainio ( 1918, 1921) reported Lee idea olivacea var. euphoreoides from Mt.
Akagi (A. Yasuda no. 71 , in TNS , vidi) and Gunma Prefecture (A. Yasuda no . 196, in
TNS , vidi). However, the above cited collections are not var. euphoreoides but var.
caesitia ; the thalli ne morphology of var. euphoreoides is similar to that of var. caesitia but var. euphoreoides (holotype, Finland, Lapponia Kemensis , leg. Vainio, HVainio 23746, in TUR, vidi) lacks atranorin and has a colorless subhymenium.
Yasuda ( 1915) validly published Lee idea spumosula "Zahlbr." based on the private communication from Zahlbruckner that was mentioned in the description. Perhaps the Zahlbruckner's publication might have been delayed due to disorder caused
by World War l ( 1914- 1918), and unfortunately Zahlbruckner's proposal of a new
species appeared the next year. In the envelope of the syntype of Lecidea spumosula"Zahlbr." preserved in TNS (Yasuda no . I 00), the number of which was cited by
Zahlbruckner ( 1916), there are six fragments of which external morphology is quite

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Masakane lnoue

similar. However, three fragments (Yasuda no. I 00 A-C) bear acicular septatedspores and are assignable to Bacidia sensu la to and the others (Yasuda no . I 00 0 - F)bear ellipsoid simple spores assignable to Lecidella. Even from the original descriptions, it is obvious that their proposals were based on different fragments; Yasuda( 1915) proposed Bacidia and Zahlbruckner (1916) proposed Lecidella. Unfortunately
Lecidea spumosula Zahlbr. (Zahlbruckner 1916) should be regarded as a laterhomonym of Lecidea spumosula "Zahlbr." Yasuda (Yasuda 1915) 2 ).
Specimens examined. HOKKAIDO. Prov. Kamikawa: Mt. Yubari , M. Oshio no .
12277 (13 Aug., 1965); Mt. Ashibetsu, M. Inoue no. 8842 (31 July, 1974). Prov. Nemuro : Sashirui of Shiretoko Peninsula, I 0 m, M. Inoue no . 9176 (21 July, 1974);
Kunbetsu of Shibetsu-cho, 80 m, M. Inoue no . 9091 & 9096 (21 July, 1974). Prov.
Kushiro: Forests around the Kushiro Marsh , M. lnoue no. 22180, 22188, 22198,
22205, 22210, 22212-13, 22216-17, 22230, 22235 , 22241, 22249, 22264, 22273,
22275-76, 22279-22281, 23823, 23942 & 24048 (2- 6 Sept. , 1992); Beppo of Kushiro-mura, 80 m, M. Inoue no. 9057 (20 July, 1974); Miyama of Kushiro-mura, !I 0 m,
M. lnoue no. 9058 , 9061 , 9069 & 9070 (20 July, 1974); Tenpoku Pass, 580m, Y.
Takahashi no. 6109 (5 Aug., 1981 ). Prov. Shiribeshi : Kamoenai , 150 m, M. Inoue no.
9126 & 9128 (2 Sept., 1974); Tomaru Pass, 800m, M. lnoue no. 9142 (2 Sept.,1974). Prov. Tokachi: Taiki-cho, 80 m, M. Inoue no. 4009 (4 July, 1973). Prov. Hidaka: Mt. Hidakaporoshiri, 750 m, M. lnoue no . 7862 (29 July, 1974); Mt. Apoi , I 0 m,
M. lnoue no . 4127 (5 July, 1973). Prov. Iburi: Noboribetsu, 250 m, M. Inoue no .
24734 (July, 1984); Horonai of Tomakomai-shi, 50 m, H. Kashiwadani no. 9461 (28Oct. , 1971 ), TNS. Prov. Hiyama: Okushiri Island, 50 m, M. lnoue no. 26138 (28Sept., 1987). Prov. Oshima: Mt. Yokotsu , M. Oshio no . 11485 & 11540 (6 Aug. ,1965). HONSHU. Pref. Aomori: Asamushi Seashore, coli. Y. Asahina no. 9 (5 Aug.,
1924), TNS; Mt. Iwaki , M. Oshio no. 8661 (2 Aug., 1962); River side of Akaishi ,570 m, Mts. Shirakami , M. lnoue no. 22954 (2 July, 1983). Pref. lwate: Yakebashiri ,Mt. Iwate, 680m, M.lnoue no . 21862 (3 Sept., 1991), 960m , M. lnoue no. 21896-97(3 Sept., 1991 ). Pref. Akita: Mt. Futatumori, Mts Shirakami, 970 m, M. lnoue no .
22587 ( 17 Sept., 1993); Mt. Kodake, Mts. Shirakami , 770 m, M. lnoue no. 22464 (5Sept., 1993); Mt. Moriyoshi, 650 m, M. lnoue no. 19843 (8 Oct. , 1987); Mt. Akitakoma, 450 m, M. lnoue no. 15868 (30 Aug., 1982); near Tsurunoyu Spha, 700 m, M.
lnoue no . 16552, 16554 & 16550 (7 Aug. , 1983); Mt. Honzan of Oga Peninsula,
570m, M.lnoue no. 16855, 16859, 16861 & 16863 (5 Aug. , 1983 ), 640m, K. Sasaki
no. 258 (25 Oct., 1991), 650m, K. Sasaki no . 211 , 212 & 214 (10 Aug. , 1991); Mt.
Kenashi of Oga Peninsula, 640m, M. Inoue no. 24224 (I June, 1991), 650m, M.
lnoue no . 24379 (27 March, 1994); Nibetsu of Akita-shi, coli. T. Makino, s. n. ( I 0
21

Bacidia spumosu/a (Yasuda) Ya suda , Flechten Japans 28, 192 5.- Lecidea spumosula Yasuda , Bot.
Mag ., Tokyo 29: 319, 191 5. Type: Japan , Pref. Gunma , Mt. Akagi, Yasuda I OOA- Iectotype in TNS, selected here.

Japanese Species of Lecidella (!I)

143

Sept., 1927), TNS; Mt. Ootakiyama of Akita-shi, 70 m, M. Inoue no. 16583 (Apr.,
1983); Mt. Chokai, 610m, M. lnoue no. 16507, 16509 & 16523 (3 May, 1983),
910 m, M. lnoue no. 16511 (3 May, 1983 ); Mt. Yamabushi , I 060 m, M. Inoue no .
16563 (8 Oct. , 1983); Takamatsu ofYuzawa-shi , 820m, M. Inoue no. 16564 (8 Oct.,1983); Mt. Kurikoma, 1400 m, M. Inoue no. 16700 & 16709 (22 Aug. , 1983). Pref.
Yamagata : Mts. Asahi, 600 m, M. Inoue no. I 0612 (30 Aug. - 2 Sept., 1975), 940 m,
M. Inoue no. 10616 (30 Aug.- 2 Sept., 1975), 1150m, M. lnoue no. 10496 (12- 15
Aug., 1975), 1360m, M. Inoue no. 10605 (30 Aug.- 2 Sept. , 1975) & 1410m, M .
Inoue no. I 0592 (30 Aug.-2 Sept., 1975); Mt. Gassan, M. Oshio no. 3969 (26 July,
1960); Mt. Iide, 730 m, M. Inoue no . I 0874 (7- 10 Aug. , 1975), I 030 m, M. Inoue no.
10864 (7- 10 Aug. , 1975), 1430m & M. Inoue no . 10886 (7- 10 Aug., 1975). Pref.
Miyagi: Mt. Kurikoma, 850m, M. Inoue no . 11750 (27 Aug. , 1975) & 1350m, M.
Inoue no. 10414 (28 Aug. , 1975). Pref. Tochigi: Nikko, Yatabe no. 292 (13 Aug.,1890), as Lecidea parasema (Miill. Arg. 1892), TNS . Pref. Gunma: Mt. Akagi , coil.
A. Tsunoda s. n. (17 Dec., 1912, Yasuda no. 71 ), as Lecidea olivacea var. euphoreoides (Vainio 1918), TNS; Mt. Akagi , coil. A. Tsunoda no. 512 (19 Sept., 1912
& 20 Dec., 1917, Yasuda no. I 00 0 - F), as Lee idea spumosula "Kouzuke", coil. A.
Tsunoda s. n. (I 0 Dec., 1917, Yasuda no. 196), as Lee idea olivacea var. euphoreoides(Vainio 1918), TNS; Mt. M yogi, 720 m, M. Inoue no . 11252 (I 0 Aug., 1976). Pref.
Niigata: Mt. Myoko, M. Oshio no . 4109 & 4114 (28-31 July, 1960); Pref. Toyama:
Mt. Shirakimine, 1410 m, M . lnoue no. 25788 (13 Oct. , 1997) & 1430 m, M. Inoueno. 25782 (13 Oct. , 1997). Pref. Ishikawa: Mt. Hakusan, 740 m, S. Nakanishi no.
66209, 1280 m, S. Nakanishi no . 67042 (2 June, 1967); Fujidaira-dani-guchi of Noto
Peninsula, !Om, M. Inoue no. 12011 (8 Oct. , 1976). Pref. Nagano: Mt. Jii , 1210m,
M. Inoue no. 5505-06, 1330 m, M. Inoue no. 5558 (1 Aug., 1973); Mt. Harinoki,
1600 m, M. Inoue no . 6041 & 6067 (17 Aug., 1973); Mt. Togakushi, 1180 m, M .
Inoue no. 719 (28 July, 1972), 1200 m, M. lnoue no. 9201-03 (29 May, 1974),
1220 m, M. lnoue no. 708 & 739 (28 July, 1972); Mt. Arafune, 1220 m, M. Inoue no.
5226 & 5191 (24 July, 1973); Mt. Hiraofuji , 1150 m, M. Inoue no. 1717 (29 July,1972); Mt. Akarusan , I 000 m, M. Inoue no. 11273 (13 Aug., 1976); Utsukushigahara
Heights, M. lnoue no. 24652, 24657 & 24658 (20 May, 1976); Mt. Anpeiji, 1480 m,
M. lnoue no. 13125, 13144 & 1210m, M. Inoue no. 13479 (2- 10 Aug. , 1979); Mt.
Utsugi, 1700m, M. Inoue no . 6896 (11 Aug., 1973); Mt. !se ofNagiso-cho, 1280m,
M. Inoue no. 439 (19 Aug., 1972). Pref. Yamanashi: Mt. Kitadake, 1730 m, M. lnoue
no . 12418 ( 19 Aug. , 1978). Pref. Kanagawa: Hakone, coil. Y. Asahina no. 98 (31Aug. , 1926), TNS. Pref. Shizuoka: Kagosaka Pass, coli. Y. Asahina, s. n. (4 June,
1926); Subashiri-guchi , Mt. Fuji, coil. Y. Asahina, s. n. (21 Aug., 1929), TNS. Pref.
Shiga: Mt. Ryozen , 400 m, M. Oshio no . I 0770 ( 16 June, 1965), 660 m, M. Oshio no.
I 0844 (16 June , 1965), 750 m, M. Oshio no. I 0824 ( 16 June, 1965). Pref. Mie: Mt.
Fujiwara, M. Oshio no. 4921 (5 Aug. , 1960). Pref. Tottori: Mt. Hyonosen, 860 m, M.
Inoue no. 7649 (11 May, 1974), 1210m, M. Inoue no . 7620 (11 May, 1974); Mt.

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Masakane lnoue

Daisen, 1140m, M. Inoue no. 7683 (13 May, 1974), 1180m, M. lnoue no. 7677 &
7694 (13 May, 1974), 1210m, M. Inoue no . 7700 (13 May, 1974). Pref. Hiroshima:
Taishaku Gorge , 520 m, M. lnoue no . 7066 (18 Apr. , 1973); Nakayama Pass near
Taishaku Gorge, 600m, M. lnoue no. 7094 & 7106 (18 Apr. , 1973); Mt. Osorakan,
I 030 m, M . lnoue no. 11125 (27 June, 1976), 1180 m, M. Inoue no . 11120 (27 June,
1976), 1250m, M. Inoue no. 11110 (27 June , 1976), 1400m, M. 1noue no. 11134 (27June, 1976); Mt. Togo, M. Oshio no . 6476 ( 11 June , 1961); Mt. Kanmuri , 720m, M.
Inoue no . 11843, 11857 & 11860 (28 May, 1977), 790 m, M. lnoue no. 11861 (28May, 1977), 950 m, M. Inoue no. 11850 (28 May, 1977); Uchiguro Pass of Togauchicho, 980 m , M. Inoue no. 11094 (26 June , 1976); Mt. Hiba, 870 m, M. lnoue no .
7576 (12 Oct. , 1973), IOOOm , M. lnoue no . 7558 (12 Oct. , 1973), 1090m, M. lnoue
no. 7531 (12 Oct., 1973), 1250m, M. Inoue no. 7600 (12 Oct., 1973). SHIKOKU.
Pref. Ehime: Mt. Saragamine, M. Oshio no . 8006 (4 May, 1962); Himetsuru-daira,1280 m, M. lnoue no. Ill 05 ( 16 July, 1976). Pref. Kochi : Masakinomori of Niyodomura, 1400m, K. Une no. 1512 & 1521 (24 Aug. , 1978). KYUSHU. Pref. Fukuoka:
Mt. Hiko , M. Oshio no . 5101 (4 Sept., 1960). Pref. Nagasaki: Mt. Ariake ofTsushima Island, 330 m , K. Terada (6 May, I 975, M. Inoue no . I 0970); Mt. Nagaura of
Nishisonogi Peninsula, 560 m , M. Inoue no. 9681 , 9684, 9685 & 9707 (9 Dec.,
1974); Mt. Unzen , 1070m (Nodake), M. lnoue no. 9804 (11 Dec., 1975), I lOOm
(Nodake), M. lnoue no. 9802 (11 Dec. , 1975), 1290m (Kunimi), M. lnoue no . 9822
(11 Dec., 1975), 1330 m (Myoken), M. Inoue no . 9802 ( 11 Dec., 1975). Pref.
Miyazaki : Mt. Ishido, I 070 m, M. lnoue no. I 0206 (22 March , 1975), 1100 m, M.
lnoue no. 10213 (22 March , 1975), 1170m, M. lnoue no. I 0168 (22 March , 1975),
1250 m, M. Inoue no. I 0184 (22 March , 1975). Additional specimens examined: (asLecidella elaeochroma). Canada, British Columbia, Moresby Island, I. M. Brodo no.
17251 in 27 June, 1971 (Lich. Can. Exs. no . 126, TNS); U.S.A. , Colorado, Boulder
County, W. A. Weber & M. Klackenbrink in 29 June , 1976 (Lich. Exs. Univ. Colorado Mus. no . 520, TNS).
5) Lecidella elaeochromoides (Nyl.) Knoph & Hertel in Knoph(Fig. 1-3)
Bib I. Lich. 36: 103 ( 1990). -Lecidea paras ema (Ach.) Ach . var. elaeochromoides Nyl. , Bull. Soc. Linn. Normandie, ser. 2., 6: 310 (1873). Type : France, " Pyrenees orientales", east from Port Yendres, non vidi.- Lecidea elaeochoromoides(Nyl.) Flag., Lich. alg. 258 ( 1895).
Thallus dark greenish gray, thin to medium , effuse, continuous, irregularly
cracked-areolate or rimose-areolate , or at times granulose-ditfract. Hypothallus indistinct.
Apothecia minute , black, up to 0.4- 0.5 mm in diameter, adnate , moderately towell constricted at the base; disc epruinose, plane with a prominent entire margin.
Excipulum 30-40(- 50) J.lm thick , violet or reddish brown with green tinge externally,
the interior part golden-brown ; hyphae perpendicularly radiating, 5- 6 J.lm thick , with

Japanese Species of Lecidella (11 )145

a thick wall. Epithecium blue-green. Hymenium 50- 60(- 80) ,urn high , not inspersed.
Subhymenium golden brown, up to 100 ,urn high , with irregularly-shaped cavities,
composed of perpendicul ar hyphae. Paraphyses lax , simple with a few branches;
apices slightly clavate. Asci clavate, 45- 65 X 12- 18 ,urn; tholus rather thin, I + intensive blue. Spores with obtuse ends, I 1- 15 X 6- 8 ,urn.
Reaction: thalus & medull a P- , K- , C+ orange (faint), KC + orange. Chemical substances: atranorin, arthothelin , thuringione (+ I -).
Habitat: on non-calcareous rocks in lower mountains of Hon shu.
Range: Japan ; Europe, Canary lsl s., Algeria, Morocco , Nepal , China, North
America (Knoph 1990, Knoph et al. 1995b).
Lecidella enteroleucella, which is one of the common representatives ofLecidella in lowlands of Japan, is most reminiscent of L. elaeochromoides, in that it
has a similar thallus , apothecial size and chemistry. Lecidella enteroleucella , however, has subimmersed to appresed-adnate apothecia and colorless subhymenium.
Lecidella carpathica , which has a golden-brown subhymenium , is distinguishedfrom L. elaeochromoides by having areolae which are bullate or subgranulate-conglomerate, wider apothecia (up to I mm in diameter) , and a different chemistry (C+ ).
Lecidella elaeochromoides has previously been reported only from Nepal andChina in Asia; its range is now extended to Japan.
Specimens examined. HONSHU. Pref. Akita: Mt. Ganmori, Shirakami Mts. ,950 m, coli. K. Ogasawara (25 July, 1983), M. lnoue no. 22828; Lake side of
Towadako, 400 m , M. Inoue no . 16595 & 16600 (Oct., 1981 ); Mt. Honzan, Oga Pen. ,550 m, M. lnoue no . 16870 ( 13 June, 1982). Pref. Gunma: lwamoto, Tone-gun, coli.
A. Tsunoda (14 Feb. , 1921 ), Yasuda no . 528 , TNS. Pref. lbaraki: Mt. Tsukuba, coil.
Y. Asahina no . 34 (March , 1924), TNS. Pref. Shizuoka: Mishima, coil. Y. Asahina, s.
n. (7 Jan. , 1931 ), TNS. Additional specimens examined. France, Ollioulas, coil. G.
C lauzade in 19 May, 1966 (Vezda, Lich . sel. Exsic. no . 488, TNS); Romania, Dobrogea, distr. Constanta, coil. A. Vezda in 5th Aug. , 1970 (Vezda, Lich. se!. Exs. no.
906, TNS); U.S.A. , California, Santa Barbara County, Channel Isl s, coil. Weber &Bratt in 8th Jan ., 1986 (Weber, Lich. Exs. no. 655 , TNS).
6) Lecidella enteroleucella (Nyl.) Hertel(Fig. 1-4)
Khumbu Hi mal 6: 330 ( 1977). -Lecidea enteroleucella Nyl. in Nyl. & Cromb.,
Journ . Linn. Soc. Lond., Bot. 20 : 67 ( 1884). Type: Japan , Kyushu , Pref. Nagasaki ,Nagasaki, leg. A. C. Maingay, s. n., - isotype in FH, vidi. - Lecidea nagasakensis
Nyl. , Lich. Jap. 62 ( 1890). Type: Japan , Kyushu, Pref. Nagasakiz Nagasaki, leg. E.
Almquist, - holotype in H (H-Nyl. 20933), vidi; - isotype inS, vidi. - Lecidea adpressula Mi.ill.Arg. , Nuov. Giorn. Bot. Ita!. 22: 126 (1891 ). Type: Japan , "Enoura" in
July 1886, no. 153, - holotype in G, non vidi; - isotype in TNS, vidi. - Lecidea
tapetiformis Zahlbr. in Handel-Mazzetti , Symbol. Sinic. 3: 107 ( 1930). Type: China,
Setshwan, 2125 malt. , leg. Handel-Mazz. no. 2699, - lectotype in W , - isolectotype

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Masakane Inoue

in WU, vidi. - Lecidea setschwanensis Zahlbr., in Handel-mazzetti , Symbol. Sinic.
3: 108 (1930). Type: China, Setschwan, 1450m alt. , leg. Handel-Mazz. no . 1170, holotype in _W , vidi; - isotype in WU , vidi.- Lecidea kelungana Zahlbr. , Feddes
Repertorium 33: 36 (1933). Type: Taiwan, Kelung, leg. U. Faurie no . 257 , vidi ; isolectotype in KYO, vidi . - Lecidea lentigerella Zahlbr. , Feddes Repertorium 33:
36 ( 1933). Type: Taiwan, Pahiran, leg. U. Faurie no. 277, - holotype in W , vidi; isotype in KYO, vidi.
Thallus effuse, continuous, white with yellow or brown tinge , rimose to irregularly rimose-areolate; areolae minute, plane. Hypothallus black, or sometimes indistinct.
Apoth~cia minute, up to 0.4(- 0.5) mm in diameter, immersed at the juvenilestage, later appressed-adnate or subimmersed, sometimes almost level with thalline
surface, usually with a distinct margin , not constricted basally, dark brown to blackish-brown; disc epruinose, plane or somewhat convex in old apothecia. Excipulum
brown with dark blue-green tinge , 30- 50 J.lm thick, composed of radiating hyphae;
hyphae 4-7 J.lm thick, pachydermatous. Epithecium pale brown or greenish brown.
Hymenium 40- 60(- 70) J.lm high, colorless. Subhymenium (50- )60- 80(- I 00) J.lm
high , with irregularly shaped cavities, colorless. Paraphyses simple or with a few
branches, lax, 2- 3 J.lm thick; apices slightly capitate, 4- 6 J.lm thick. Asci clavate,35- 55 X I 0- 15 J.lm ; tholus 4- 6 J.lm thick, I+ blue. Spores with obtuse ends, 8- 13(- 15) X 5- 8(- 10) J.lm.
Reaction: thallus & medulla P-, K+ yellow (faintly), KC- , C + orange. Chemical substances: atranorin , arthothelin , thuringione
Habitat: on non-calcareous rocks in the lowlands.
Range: Japan; China, India, Indonesia, Korea, and Taiwan; eastern Australia(Rambold 1989), South Africa (Knoph 1990); U.S.A. , Mexico, Puerto Rico ,Venezuela & Brazil (Knoph & Leuckert 1994).
The diagnostic features of this species include the rimose to irregularly rimoseareolate thallus , minute apothecium which is adnate or subimmersed, colorless subhymenium, and the production of atranorin and xanthones (C + ).
Zahlbruckner (1930, 1933) proposed four new species from Asia; Lecidea
kelungana and L. lentigerella from Taiwan, L. setschwanensis and L. tapetiformis
from China. After a detailed and careful examination of type specimens, it can be
now stated that they all belong to one species despite the divergence in diagnostic
statements, as Hertel ( 1977) correctly concluded. Superficial examination of the materials may be certainly the reason why Lecidella enteroleucella was redescribed asthese different species. For example, in the type specimens of L. tapet(formis, hymenia were observed to be less than 70 J.lm high, though they were stated by
Zahlbruckner to be 90- 100 J.lm . He stated, that for L. setschwanensis the spore dimensions were 9- 11 X 3-4 J.lm , but according to my re-examination of the type specimen, spores are larger, measuring I 0- 12 X 5- 7 J.lm.

Japanese Species o f Lecide/la (11 )147m

35001--------------~~-------------

3000 1 - - - - - - - - - - l
2500 1----------------l2000 f-,------T""Tnr--1
1500

I

Fi g. 2.

Distribution of Lecidella enteroleucella (Nyl. ) Hertel.

Lecidella enteroleucella is closely related to Lecidella viridans (Fiotow) Korb. ,which is one of well-known species from both Europe and America. I have not had aopportunity to re-examine any syntypes of L. viridans, but from an examination ofEuropean specimens (Heiderberg, leg. Ahles, Hepp 726, in TNS; Italy, Vinschgau ,coli. M. Inoue & H. Hertel , M. Inoue no. 11959 (19 Sept., 1977); "Sudeten" , Herb.
Korb. s. n., in W) and from some descriptions given by Korber ( 1855), Lighton( 1879), Jatta ( 1911 ), Poelt & Vezda ( 1981 ), Knoph ( 1990), and Thomson ( 1997) it
seems clear that L. enteroleucella is distinguished from L. viridans , in which thethallus is minutely granulate, by the plane areolae (Fig. l-4a, 4b ).
Hertel & Leuckert (1969), Knoph & Leuckert (1994), Knoph et al. (1995b) andThomson ( 1997) reported that L. viridans did not produce atranorin and my analysis
proves that above mentioned european representatives also do not. In contrast, Japanese L. enteroleucella produces atranorin .
Lecidella enteroleucella is common from seacoast to the lowlands of Japan, andis not known from Hokkaido situated in northern Japan (Fig. 2).
Specimens examined. HONSHU. Pref. Aomori: Kusoudomari coast, Wakino-

148

Masakane lnouesawa, Shimokita Pen. , 1- 2 m, M. lnoue no . 25609 ( 18 Oct., 1996). Pref. Akita :Nyudo Point, Oga Pen. , 20m, M. lnoue no. 16853 (5 Aug. , 1983), M. lnoue no.
24384 & 24385 (27 March, 1994); Monzen. Oga Pen ., 0- 5 m, M. lnoue no. 16614( 15 May, 1983). Pref. Yamagata : Fukura coast, Yuza-machi , 0- 5 m, M. lnoue no.
16605 (I June, 1983). Pref. Nagano: Mt. Akarusan , Saku-shi , 850 m, M. lnoue no .
11280 (13 Aug. , 1976). Tokyo Metropolis: The Imperial Palace, coli. G. Thor no.
14908 (20 June, 1995), TNS . Pref. Shizuoka: Kisegawa River, Sunto-gun, coli. Y.
Asahina no. 154 (14 Aug ., 1929), TNS ; Enoura, Sunto-gun, coli. Y. Asahina , s. n. (5Jan. , 1937), TNS; Johgasaki coarst, 2- 5 m, M . lnoue no . 11440 (25 Aug. , 1976); Kumomi , Matsuzaki-machi , 50 m, M. lnoue no . 11438 (25 Aug. , 1976); Sunpu Ruins ofa Castle in Shizuoka City, coli. S. Kurokawa no. 72006-8 (8 Feb., 1972), TNS. Pref.
Hyogo: Nunobiki Fall , Kobe-shi , 150 m, M. lnoue no. 11784 & 11785 (8 May, 1977).
Pref. Shimane: Hinomisaki Point, I m, H. Miyawaki no. 5219 (31 Dec. , 1981 ). Pref.
Yamaguchi: Coast of Susa Bay, 1- 2 m, M . lnoue no . 7449, 7466 & 7469 (9 June,
1973). Pref. Hiroshima : Shiwa-machi , 270 m , M . lnoue no . 7000 (8 March , 1974);
Togouchi , Hiroshima-shi , 80 m, M . lnoue no. 12059 (25 Apr., 1978); ltsukushima
Is!. , I m, M. lnoue no . 3444 (26 Apr. , 1973) & many. SHIKOKU. Pref. Kagawa :(Shodo-shima Is!.) Myokenzaki , 2-3 m, M. lnoue no . 26077 ( 18 Dec. , 1998), Kaburazaki, I 0 m, M. lnoue no. 26091 ( 18 Dec. , 1998), Kurosaki , 0- 1 m, M . Inoue no.
26121 ( 19 Dec. , 1998), Mt. Toun-san, 300 m, M. lnoue no. 26009 ( 17 Dec. , 1998).
Pref. Tokushima: Takegashima Is!. , Shishikui-machi , I- Sm, M. lnoueno. 11241 (19July, 1976). Pref. Ehime : Yuge isl , Inland sea, 3m, M. lnoue no. 7189 (11 Nov. ,1973). Pref. Kochi: Isa , Hata-gun , coli. F. Fujikawa, s. n. (25 Aug., 1931), TNS;
Mizushiri , Aki-gun , Sm , M . lnoue no . 11245 (19 July, 1976); Muroto Point, 2- !0m,
M. lnoue no. 11234 (19 July, 1976); Seiryuji temple , Tosa-shi , !Om, coli. H. Matsui(M. lnoue no. 11753, 23 Apr. , 1977). KYUSHU. Pref. Nagasaki : Gongen-iwa, Ooseto-machi , 110 m, M. lnoue no. 9458 (8 Dec., 1974), Uchigo , Ooseto-machi , M. Inoueno. 9437 (8 Dec., 1974); Matsushima Is!, Ooseto-machi, 3 m, M. lnoue no. 9299 (7Dec. , 1974); Kozera, Minamimatsuura-gun, 3 m, H. Miyawaki no. 4848 (15 Sept. ,1981). Pref. Kagoshima: Miyanoura, 1- 2 m, M. Inoue no. 10055 & 10063 (25March, 1975); Tashiro near Anbo, 1- 3 m, H. Kashiwadani no . 38577 (27 Oct. , 1994),
TNS.
7) Lecidella euphorea (Fii:irke) Hertel(Fig. 1-5)
Lichenologist 12: I 07 ( 1980). - Lee idea sabuletorum var. euphorea Fli:irke,
Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten Naturk. 3: 311( 1808). Type: Austria, Salzburg, coli. Wittmann in 21 Sept. , 1985, (SZU no . 4161 ),by Knoph & Leuckert (1999) , non vidi . - Lecidea euphorea (Fii:irke) Nyl , Mem.
Soc. Sci . Nat. Cherbourg 5: 126 (1858).
Thallus thin, effuse, irregularly rimulosed, or rarely subleprosed, whitish to darkgray; medulla 1-. Hypothallus indistinct.

Japanese Species of Lecidella (I! )149

Apothecia up to 0.8 mm in diameter, adnate , black, moderately constricted at thebase; disc epruinose, plane with a prominent entire margin , at times becoming somewhat convex with an obliterated margin. Excipulum 40- 60 J.lm thick, dark blue-green
externally, the interior part golden-brown or reddish brown; hyphae perpendicularly
radiating, 6- 8 J.lm thick , pachydermatous. Epithecium blue-green or paler. Hymenium 60- 90 J.lm high , not inspersed. Subhymenium golden-brown, up to 10 J.lm high,with irregularly shaped cavities. Paraphyses simple or slightly branched, lax ,1.5- 2.5 J.l.m thick; apices not clavate or slightly. Asci clavate, 50- 70X 12- 15 J.lm ; tholus thick , I + intensive blue. Spores with obtuse ends, I 0- 15 X 6- 9 J.lm.
Reaction : thallus & medulla P+ yellow (faint) , K + yellow, KC- , C-. Chemical substances: atranorin and unidentified minor constituents (TLC). (Knoph &Leuckert (1999) reported atranorin , aotearone , and 2,5, 7-trichloro-3-0-methylnorlichexanthone as major compounds from L. euphorea.)
Habitat: on bark in mountain regions .
Range: Japan ; Europe (Poelt & Vezda, 1981 ); Svalbard (Eivebakk & Hertel ,1997); Sardinia, Italy (Nimis & Poelt, 1987); India (Schubert & Klement, 1966);
Mongol (Huneck et al. , 1987); American arctic region (Thomson , 1997).
Knoph & Leuckert ( 1999) correctly proposed conservation of the name Lee idea
euphorea with a conserved type .
Lecidella euphorea is closely related to L. elaeochroma, which is one of thewell-known lichens in both the Northern and the Southern Hemispheres. The moderate-sized apothecia with prominent margin and the blue green epithecium as well thegolden-brown or reddish brown subhymenium are common characters in these two
corticolous lichens. However, L. euphorea differs in that the rimulose thallus (Fig. 15) is C-negative reaction and it lacks a black hypothalline line. This species is not as
common as L. elaeochroma in Japan.
This species was previously known from India and Mongol in Asia, however, therange now extended to Japan .
Specimens examined. HOKKAIDO. Prov. Tokachi : Mt. Tsurugi , 560 m, M.
lnoue no . 8211 (28 July, 1974). HONSHU. Pref. Akita, Mts. Shirakami, 650 m, M.
lnoue no . 22409 (14 Sept. , 1993). Pref. Nagano , Mts. Yatsu, 890m , M. Inoue no . 864(22 July, 1972). Additional specimens examined. Canada, British Columbia, Northern Rocky Mountains , leg. Brodo in 1977 (Brodo , Lich . Can. Exsic. no. 188, TNS).
8) Lecidella sendaiensis (Zahlbr.) Knoph & Leuckert(Fig. 1-6)
Bib I. Lich. 68: 131 ( 1997). - Lee idea sendaiensis Zahlbr. , Ann . Mycol. 14: 52(1916). Type: Japan, Honshu , Pref. Miyagi , Sendai , on bark of Tilia mique/iana, leg.
A. Yasuda no . 126, June 10, 1913 , - holotype in W , vidi . - Lecidea pruni Vain. ,Bot. Mag. , Tokyo 32 : 159 ( 1918). Type: Japan , Honshu, Pref. Miyagi , Sendai, onbark of Prunus sp. , leg. A. Yasuda no . 94, June I 0, 1913 , - isotype in TNS , vidi. Lee idea tiliae Vain. , Bot. Mag., Tokyo 32: 161 ( 1918). Type: Japan , Honshu, Pref.

150

Masa kane lnoue

Miyagi, Sendai, on bark of Tilia miqueliana, leg. A. Yasuda no. 93 , June 10, 1913,
holotype in TUR (H- Vain. 24216), vidi ; - isotype in TNS, vidi. - Lecidea exigua
auct. non Chaub.; Mull.Arg. , Flora 63 : 484 ( 1879).
Thallus subdeterminate, thin , continuous or in part evanescent, rimulose , plane,tartareous, ash-white with yellow tinge , surrounded by a blacki sh hypothalline line,
and sometimes forming a mosaic; medulla I- .
Apothecia minute, up to 0.3(- 0.4) mm in diameter, adnate, moderately constricted at the base; disc brown to blackish-brown, epruinose, plane with a prominent
entire margin at the juvenile stage, then becoming convex with an obliterated margin ;margin somewhat paler than the disc . Excipulum 30- 50 f..lm thick, pale brown or concolorous to the epithecium externally, the interior part colorless; hyphae perpendicularly radiating, 3-4 f..lm (4- 5 .urn in outer most) thick, with a thick wall. Epithecium
pale brown with green tinge . Hymenium 50- 70(- 80)f..lm high . Subhymenium reaching to I 00 f..lm high , colorless or rarely pale brown , with irregularly shaped cavities.
Paraphyses simple or rarely branched, lax , 1.5-2.5 f..lm thick ; apices moderately
swollen, 3-4 f..lm thick. Asci 45- 65 X I 0- 18 f..lm , clavate; tholus rather thin , I+ blue.
Spores with obtuse ends, 9-13(- 15) X 6- 8(- I 0) f..lm .
Reaction: P- , K + yellow, KC- , C-; medulla P-, K + ye llow (faintly), KC-,
C+ orange. Chemical substances: atranorin , thiophanic acid (+ I-), and an unidentified minor constituent (+ I-).
Habitat: on bark in the lowlands.
Range : known only from Japan and Korea.
Lecidella enteroleucella , which is the sax icolous assemblage and resembles L.
sendaiensis in terms of thallus morphology and apothecial size, is distinguished from
L. sendaiensis by somewhat wider apothecia which are appressed-adnate or subimmersed and by different xanthones (arthothelin and thuringione) .
Lecidella sendaiensis and L. tiliae were proposed by Zahlbruckner ( 1916) andVainio ( 1918) respectively based on Japanese materials which A. Yasuda collected.
Reexamination of the syntypes of both species, however, revealed that they are basedon the same collections (the same origin, with a citation of the same locality, collecting date and substrate ), and that the two species are actually identical with each other.
Vainio ( 1918) proposed L. pruni and L. tiliae in the same paper, but they should
be assigned to one species from both morphological and chemical points of view, despite the diverging statements in their diagnoses. Vainio ( 1918) described the sporesof L. pruni as !-septated, but reexamination of the type specimen revealed they aresimple. They could appear otherwise due to having 2 oil drops or, alternatively, aplasmatic cell structure.
Muller ( 1879) reported Lee idea exigua Chaub. from Japan. But the Japanesecollection cited is referable to L. sendaiensis since morphological and chemical features are similar.
Specimens examined. HOKKAIDO. Prov. Soya: Rishiri Is., M. Oshio no. 12915

Japanese Species of Lecidella (I I)

151( 17 Aug. , 1965). Prov. Kushiro : Kushiro Marsh, 5- 10 m, M . Inoue no. 2417 1 & 4others (4- 6 Sept., 1992). Prov. lburi : Mt. Usu , H. Kashiwadani no. 14166 (11 Nov.,
1977), TNS. HONSHU. Pref. Akita: Lake side of Towada-ko, 400 m , M. Inoue no.
16589 & 16593 (Oct. , 1981); Oga Pen. (Mt. Shinzan), coli. K. Sasaki (M. lnoue no.
25202 ; Aug. , 1991); Mt. Akitakoma, 650m , M. lnoue no . 14833 (9 May, 1982);
Tooshi Hieghts, Nigaho, 200m , M. Inoue no. 24523 (14 Aug., 1981). Pref. Ishikawa:Kanazawa-shi , 90 m , coli. H. Kashiwadani no. 39585 ( 11 Dec., 1995), TNS. Pref.
Nagano : Utsukushigahara, 1700 m , M. lnoue no . 24636 (20 May, 1976); Mt. Anpeiji,
1250 m, M. lnoue no. 13339 (5 Aug. , 1979), 960 m , M . lnoue no . 13528 (9 Aug .,1979); lida-shi , 550 m , M. lnoue no . 14483 (7 Aug ., 1980). Pref. Shizuoka : M ito
Seashore, Tagata-gun, coli. Y. Asahina s. n. (5 Jan ., 1931 ), TNS. Pref. Shiga : Mt.
Ryozen, 285 m, M . Oshio no. I 0746 ( 16 June, 1965); Hikone-shi , 150m, M. lnoue
no. 9218 (26 June , 1974). Pref. Hyogo: " J Kouno", coli. A . Henon in 1876, asLecidea exigua Chaub. (Miill.Arg. , 1879), G . Pref. Hiroshima: Yasaka Gorge , I 00 m ,M . lnoue no . 11765 (29 Apr., 1977); Koshigahara near Yasaka Gorge, 750 m, M.
lnoue no. 12437 (18 June, 1978); Mt. Oomine near Yasaka Gorge, 800m, M . lnoue
no . 12595 (Nov., 1977); Mt. Kanmuri , 860 m, M. lnoue no. 11846 (28 May, 1977).
SHIKOKU. Pref. Kagawa: Shodo-shima Is!., around Emon-no-taki, 270 m, M. lnoue
no. 26026 (18 Dec. , 1998). KYUSHU. Pref. Ooita : Mt. Kuju , 950m, M. lnoue no.
24240 & 24241 (7 May, 1989). Pref. Nagasaki: Nishisonogi Pen. (Se ikai) , I 00 m, M.
Nakanishi no . 9699 (4 Aug. , 1963); Nishisonogi Pen. (Ooseto) , 20m , M. lnoue no.
9253 (7 Dec., 1974); Mt. Nagaura, 560 m, M. lnoue no. 9695 (9 Dec., 1974); Mt.
Tohmi , 210 m , M. lnoue no. 9276 & 9277 (7 Dec. , 1974). Pref. Miyazaki: Mt. lshido,
850m , M. lnoue no. 10149 ( 22 March , 1975). Pref. Kagoshima: Mt. Takakuma, coli.
M . Oshio no. 683 (15 Apr. , 1965); Yakusima Is. (Nagata) , I m, M. lnoue no. 9981
(30 March , 1975), Yakushima Is. (Isso) , 30 m , M. lnoue no. I 0075 (25 March , 1975).
Additional specimen examined. Korea. Prov. Kyonggi: Mt. Pukhan-san, Koyang city,
about 780 m. M. lnoue no. 25659 (24 Aug. , 1997), TNS.

Acknowledgements
I wish to express my heartily gratitude to the directors and curators of the following herbaria who kindly sent me type and authentic specimens on loan : Drs. T.
Ahti & 0. Vitikainen of Helsinki University (H), Dr. F. Ehrendorfer of Wien University (WU) , Dr. H. Kashiwadani of the National Science Museum, Tokyo (TNS) , Dr.
J. Miege of the Conservatoire et Jardin Botaniques, Geneve (G), Dr. D. H. Pfister of
Farlow Herbarium (FH), Dr. H. Riedle of the Natiirhistorisches Museum , Wien (W),
Dr. R. Santesson of the Museum of Natural History, Stockholm (S) , and Dr. J.
Takeuchi of Kyoto University (KYO) . Thi s study was partly supported by a Grant-in
Aid for Scientific Research from the ministry of Education , Science, Sports and Culture of Japan to M. lnoue no . 05640782 and no. I 0640676. Last but not least, I152

Masakane lnoue

heartily thank Prof. Dr. Dianne Fahselt of the University of Western Ontario who
kindly read the manuscript and gave critical suggestions.

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