Bull. Natn. Sci. Mus. , Tokyo, Ser. B, 23(2) , pp. 43- 58, June 24, 1997

Little Known Crustose Lichens Bearing LecideineBlack Apothecia from Japan
Masakane Inoue
Biological Institute, College of Education, Akita University,
Akita, 010 Japan
Abstract Eight new, or little known crustose lichens are reported. The genera
Arthrorhaphis, Cecidonia and Epilichen are new to Japan. Epilichen scabrosus andRhizocarpon hensseniae are new additions to the flora of Asia. Arthrorhaphis
citrinella , Cecidonia umbonella and TrapeZia mooreana are new additions to theflora of Japan. D escriptions including taxonomic and chemical data are provided . These are also provided for Rhizocarpon hochstetteri, Trapeliopsis jlexuosa
and Tremolecia atrata , which are the species previously little known in Japan.
Key words: lichens, taxonomy, Japan, Arthrorhaphis, Cecidonia , Epilichen ,Rhizocarpon, Trapelia , Trapeliopsis, Tremolecia.

In the course of my investigations on Japanese Lecideaceae (s. lato-sensu
Zahlbruckner 1926) . I have been detecting the several new, or a little known
crustose lichens being characterized by lecideine black apothecia. I will present
some taxonomic notes on them.
In the list specimens examined, the locality name is followed by the citationof altitude, herbarium number and the date collected. The specimens listed here
are preserved in the herbarium of Akita University and in the herbarium of the
National Science Museum, Tokyo (TNS) .
1.

The genera which are new to JapanARTHRORHAPHIS Th. Fr. , Acta reg. Soc. Sci, ups., ser. Ill, 3: 203- 204,1860.
Type species: Arthrorhaphis .flavovirescens (Dicks.) Th. Fr.
2) CECIDONIA Triebel & Rambold, Nova Hedw. 47: 279- 309, 1988.
Type species: Cecidonia umbonella (Nyl.) Triebel & Rambold
3) EPILICHEN Clem. ex Hafellner, Nova Hedwigia 30: 676, 1978.
Type species: Epilichen scabrosus (Ach.) Clem. ex Hafellner
1)

Enumeration of species
Arthrorhaphis citrinella (Ach.) Poelt(Pl. I: a)
Bestimmungsschliissel Europaischer Flechten 126 ( 1969). - Lichen citrinellus Ach., Kongl. vetensk. acad. nya handl. 135 ( 1795) . Type: Sweden, -lecto2.

1)

44

Masakane Inouetype (Obermayer, 1994) in H-Ach 262, vidi. -Bacidia citrinella (Ach.) Branth &Rostr., Bot. Tidsskr. 3: 235 (1869).
Thallus irregularly bullate-areolate forming patches reached to 2 cm indiameter; areolae with roughened surface, in parts sorediate, crowded, or at times
subscattered, bright yellow with green tinge. Hypothallus indistinct.
Apothecia 0.3--D.5 mm in diameter, adnate, solitary or congregated, black;disc naked, concave to plane, with a thick margin. Excipulum 7o-90 f.lm thick,dark brown externally becoming gradually paler internally, but not colorless;hyphae subradiating, irregularly entangled, 2- 4 f.lm thick, leptodermatous tomesodermatous (Fig. 1). Epithecium colorless or pale. Hymenium crystalline,
6o-70 f.lm high. Subhymenium colorless or pale, 2o-40 f.lm high. Hypothecium
pale brown, reaching to 100 f.lm high; hyphae more or less perpendicular. Paraphyses slender, 1.5-2 f.lm thick, coherent, branched, anastomosed; apices not
swollen. Asci clavate, thick-walled apically, 55-65 X 6- 8 f.lm. Spores acicular,
35-60 X 2- 3 f.lm, 8- 10 septate.
Reaction: thallus & medulla P - , K - , KC - , C - . Chemical substances:rhizocarpic acid, an unidentified substance (TLC).
Habitat: on humus, mosses, Nostoc and rocks (lava) in subalpine and alpine

Fig. 1. Cross-section of an excipulum of Arthrorhaphis citrinella (M. Inoue no. 16031),
X 100.

Little Known Crustose Lichens Bearing Lecideine Apothecia

45

regions.
Range: Japan; China (Himalaya, Obermayer 1996); Nepal (Langton, Obermayer 1996) ; Malaysia (Mt. Kinabalu, Sipman 1993); Europe (Poelt & Vezda
1977; Scandinavia, Santesson 1993); Canada (British Columbia, Noble et al.
1987, Obermayer 1994); Subantarctic (Marion Island, Lindsay 1976).
Anatomically the Japanese representatives resemble the lectotype of A.
citrinel/a, which, however, differs in having wider apothecia (0.8--0.9 mm wide in
the type).
A. citrinella is easily recognized by acicular multi-septate spores and thebright yellow thallus bearing black lecideine apothecia, which looks like a yellow
Rhizocarpon.
This species was previously known from Asia (China, Nepal and Malaysia)
including North America, Europe and the Subantarctic. However, the range has
now extended to Japan.
Specimens examined. HONSHU. Pref. Akita: Mt. Moriyoshi, 1270m, M.
Inoue no. 16026 (8 Oct., 1982), 1280m, M. Inoue no. 22975 (23 Sept., 1984),
1400m, M. Inoue no. 16031 (8 Oct., 1982); Mt. Kurikoma, 1420m, M. Inoueno. 16735 (22 Aug., 1983). Pref. Iwate: Mt. Iwate, 650m, M. Inoue no. 22042(1 Sept., 1991), 1270m, M. Inoue no. 21823 (1 Sept. , 1991), 1350m, M. Inoueno. 21932 (1 Sept., 1991); Mt. Yakeishi, 1450m, M. Inoue no. 16652 (26 Aug.,
1983), 1500m, coli. K. Sasaki, M. Inoue no. 22978 (11 Aug., 1982) . Pref.
Yamagata: Mt. Chokai, 1370m, M . Inoue no. 16916 (21 Aug., 1984) .
An additional specimen examined. Sweden, Jemtlem, coli. S. Almquist, s. n.
(in 1868), as Arthrorhaphis flavovirescens , TNS .
Cecidonia umbonella (Nyl.) Triebel & Rambold(PI. I: b)
Nova Hedw. 47:284 (1988) . - Lecidea umbonella Nyl., Flora 49 : 372 (1866).
Type: Scotland, Perthshire, Ben More, - holotype in H (H-Nyl. 15535a), vidi.
Thallus subdeterminate, forming islets in thalli of Lecidealactea, irregularly
cracked-areolate, chalky or glaucous white; medulla I + violet.
Apothecia 0.2--0.5 mm in diameter, subimmersed, black, with a thick entire
margin; disc with central umbo. Excipulum developed, dark brown; hyphae
leptodermatous, radiating, conglutinating, 2- 3 f.lm thick. Epithecium darkbrown. Hymenium 7G-90 f.liD high. Subhymenium colorless, reaching 50 f.1m
high; hypothecium not obvious. Paraphyses coherent, anastomosed, 2- 2.5 f.1m
thick; apices not swollen. Asci clavate, 6G-75 X 15- 25 f.1ID. Spores ellipsoid withobtuse ends, 11- 13 X 7- 9 f.lm; walls thin.
Reaction: thallus P + yellow, K + yellow, then red, KC - , C -. Chemical
substances: norstictic acid (TLC) .
Habitat: on the thallus of Lecidea lactea growing on rocks.
Range: Japan; Europe, Russia (Caucasus), U.S.A. (Colorado) , Spitzbergen2)

46

Masakane Inoue

Fig. 2. Cross-section of an apothecium showing "central umbo" of Cecidonia umbonella(M. Inoue no. 12410), X 100.

& Greenland (Triebel & Rambold 1988); Alaska (Hertel 1991) .
C. umbonella is distinctive by the chalky or glaucous-white thallus on thethallus of other lichens and by the subimmersed apothecial disc with central umbo
(Fig. 2) . This species is known in Japan only from one gathering, but the material
is well developed. Since the species is very characteristic, the identification as thisspecies seems justifiable.
This species was previously known only from Europe, North America, Arctic
and western Asia. The range now extends to eastern Asia.
Specimen examined. HONSHU. Pref. Yamanashi: Mt. Ainotake near Mt.
Kitadake, 3150m, M. Inoue no. 12410 (18 Aug., 1978) .
Additional specimens examined. Norway, Hordaland, 1350m, H . Hertel no.
15667 (coli. in 1974), HIRO; Harfedalen Prov., 920m, R . Santesson no. 31148,
Aug., 1984 (Moberg, Lich. sel. Exsic. Upsal. no. 15, TNS) . Austria, Carinthia,
2200m, coli. H. Hertel, A. Vezda & V. Wirth, July 1978 (Vezda, Lich. sel. Exsic.
no. 1591, TNS) .
Epilichen scabrosus ( Ach.) Clem. ex Hafellner(PI. I: c)
Nova Hedwigia 30: 677 (1978) . -Lecidea scabrosa Ach., Meth. Lich. 48( 1803) . Type: Europe ?, leg. Swartz, - isotype in UPS, non vidi. - Buel/ia
scabrosa (Ach.) A. Massal., Geneac. Lich. , 20 (1854) .
Thallus yellow with green tinge, polished, subsquamulose, on the thalli of
Baeomyces placophyllus in juvenile, then free-living. Hypothallus indistinct.
3)

Little Known Crustose Lichens Bearing Lecideine Apothecia

47

Apothecia reaching to 1.0 mm in diameter, appressed-adnate, immarginate,convex, black. Excipulum thin, 3Q-50 f.lm thick, dark brown internally andbecoming gradually paler externally; hyphae radiating, mesodermatous to pachydermatous, irregularly entangled, 4- 6 f.lm thick (Fig. 3). Epithecium brownish
green. Hymenium 10Q-120f.1m high. Subhymenium 5-10/.lm high, colorless.
Hypothecium dark brown, reaching to 250 f.lm thick; hypae irregularly arranged.
Paraphyses slender, 1.5- 2 f.lm thick, coherent, branched, anastomosed. Asciclavate, 8Q-100 X 15- l8f.1m, with thin thollus. Spores brown, 1-septate, 12- 15X
15-18 f.lm.
Reaction: thallus P + brick red, K + yellow, KC - , C - . Chemical substances: rhizocarpic acid, stictic acid, an unidentified substance (TLC).
Habitat: on Baeomyces placophyllus in juvenile stage and later on humus.
Range: Japan; Europe, northern Russia & North America (Hafellner 1978);Greenland (Th. Fries 1860, as Buellia scabrosa).
In general appearance E. scabrosus is most reminiscent of saxicolous
Tephromela aglaea (lnoue 1988, as Lecidea aglaea) because of the yellow thallus
with immarginate black apothecia, but is easily distinguished by its two-celledspores and peculiar habitat; on the thalli of Baeomyces during juvenile stage.
Although this species was previously known from Europe, Northern Russia,
Greenland and North America, the range now includes eastern Asia.
A specimen examined. HONSHU. Pref. Iwate: Mt. Yakeishi, 1450m, M.
Inoue no. 16651 (26 Aug., 1983).
Additional specimens examined. Norway, "Finmark, Aldiok", leg. Fries Th.

Fig. 3.

Cross-section of an excipulum of Epilichen scabrosus (M. Inoue no. 16651), X 100.

48

Masakane Inouein 1857, in H , s. n. Tirol? (without precise locality) , Arnold ex. no. 97 b, TNS .
Rhizocarpon hensseniae Brodo(Pl. I: d)Bibl. Lichenol. 38: 29- 35 (1990). Type: Canada. British Columbia: Queen
Charlotte Islands, Graham Island, Brodo no. l0264, holotype in CANL, non vidi.
Thallus contiguous, thin to medium, irregularly cracked-areolate, withnarrow fissures, violet-gray or paler, surrounded by blackish hypothalline lines;
areolae plane, somewhat polished; medulla I - ; cephalodia gray with violet tinge,
hemispherical, intermixed with areolae.
Apothecia black, adnate or appressed-adnate, up to 1.5 mm or at times to 2.0
mm wide, moderately constricted at the base; margin thin or indistinct; discepruinose, flat to slightly convex. Epithecium brown, rarely with green tinge.
Excipulum narrow, 3G-40 ,urn thick, brown in external part, and becoming paler
towards the center; hyphae perpendicularly radiating, paraplectenchymatous.
Hymenium (11G-)13G-160(- 180),um high. Subhymenium up to 200,um high,dark brown, with irregularly shaped cavities. Hypothecium indistinct. Paraphyses coherent, anastomosed; apices slightly swollen. Asci clavate, llG--145 X
26- 43 ,urn. Spores colorless at first, then becoming brown, !-septate, with a halo,32- 42 X 13- 20 ,urn.
Reaction: thallus & medulla P + reddish, K + yellow, KC - , C - . Chemical
substances: stictic acid (TLC).
Habitat: on non-calcareous rocks in subalpine and alpine regions.
Range: Japan; northwest coast of North America (Brodo 1990) .
R. hensseniae is closely related to R. hochstetteri and R. cinnereovirens(Muller Arg.) Vain. because of having !-septated spores, brown or brownish grayareolated thallus (rhizocarpic acid is deficient) and producing stictic acid or
norstictic acid. However, R. hensseniae can be easily distinguished from them
mainly by the cephalodiate thallus, which is a quite exceptional character within
the genus Rhizocarpon.
This species was previously known from North America. However, therange is now extend to Japan.
Amygdalaria subdissentiens (Nyl.) Mas. Inoue & Brodo, which is known
from the northwest coast of North America (Brodo & Hertel 1987), Japan andSiberia (Inoue, 1984), appears to have a distribution similar to R. hensseniae.
Brodo ( 1990) assumed that R. hensseniae, like some other lichens from theCharlotte, survived the Wisconsin glacier and speciated in refugia on thatarchipelago. R. hennsseniae and A. subdissentiens, of which vertical distribution
range is rather higher regions in Japan, appears to have important phytogeographical implications.
Specimen examined. HOKKAIDO. Prov. Kamikawa: Mt. Midori in Mts.
Daisetsu, 2070m, M. Inoue no. 8555 (8 Aug., 1974). Prov. Rumoi: Mt.

4)

Little Known Crustose Lichens Bearing Lecideine Apothecia

49

Shokanbetsu, 1440m, M. Inoue no. 8264 (21 Aug., 1974); 1470m, M. Inoue no.
8293 (21 Aug., 1974). HONSHU. Pref. Yamagata: Mt. Chokai, 1360m, M.
Inoue no. 16923 (20 Aug., 1984); Mt. Ita in Mts. Asahi, 1640m, M. Inoue no.
10448 (12 Aug., 1975) ; Mt. lide, 1780m, M. Inoue no. 10902 (9 Aug., 1975).
Pref. Miyagi: Mt. Zao, 1570m, M. Inoue no. 10691 (21 Aug., 1975); Mt.
Kurikoma, 1570m, M. Inoue no. 10359 (27 Aug., 1975). Pref. Nagana: Mt.
Goryu, 2640m, M. Inoue no. 5539 (30 July, 1973); Mt. Renge in Mts. Hida, 2790
m, M. Inoue no. 5011 (18 Aug. , 1973); Mt. Noguchigoro in Mts. Hida, 2770m,
M. Inoue no. 5891 (19 Aug., 1973) & 2910m, M. Inoue no. 12731 (18 Aug.,
1979); Mt. Mitsudake in Mts. Hida, 2810m, M . Inoue no. 5965 (20 Aug., 1973);
Mt. Momisawa in Mts. Hida, 2750m, M. Inoue no. 4831(21 Aug., 1973); Mt.
Yari in Mts. Hida, 3110m, M . lnoue no. 4912 (21 Aug., 1973); Mt. Hoken in
Mts. Kiso, 2710m, M. lnoue no. 6940 (10 Aug., 1973); Mt. Ontake, 2950m, M.
Inoue no. 4754 (7 Aug., 1973).
An additional specimens examined. U.S.A. Alaska, Juneau, ea. 700m,
O' Clair, R . M. , s.n. (det. Brodo, I. M.) , Nat. Herb. Canada: CANAL no. 92425 .
Canada, Queen Charlotte Isls., Graham Isl. , 18G-210m, Brodo, I. M . no. 26959(Nat. Herb. Canada: CANAL no. 98879); Moresby lsl. , Brad, I. M. no. 23677(Nat. Herb. Canada: CANAL no. 107157) .
Rhizocarpon hochstetteri (Korb.) Vain.
(PI. 11: a)
Acta Soc. Fauna et Flora Fenn. 53: 332 (1922). - Catil/aria hochstetteri
Korb., Parerg. Lich. 195 (1861). Type: Germany, non vidi.-Lecidea col/udens
Nyl. , Flora 53: 38 (1870) ; Nyl., Lich. Jap. 82 (1890) , -Rhizocarpon massalongi
f. colludens (Nyl.) Zahlbr. , Cat. Lich. Univ. 4: 336 (1927) . Type: Finland,"Holmiae", leg. W. Nylander in 1852 - lectotype (1. M. Brodo 1992 in Nylander
Herbarium in H), in H (H-Nyl. 10047), vidi.
Thallus irregularly areolate, brown with grey tinge, somewhat polished;
areolae dispersed on a black hypothallus, or in part, contiguous, surrounded bya conspicuous, extensive, fimbriate-radiating, dendritic hypothalline margin;
medulla I - .
Apothecia reaching to 1.3(- 1.5) mm in diameter, black, appressed-adnate,variable in shape, round or at times a few small ones clustered in groups; discconcave, epruinose; margin prominent, flexuose. Excipulum brown, medium inthickness, or often well developed, reaching to 130 f.J.m thick, brown to darkbrown; hyphae radiating. Epithecium brown. Hymenium 8G-100 f.J.m high.
Subhymenium colorless or pale, 15- 20 f..l.m high. Hypothecium brown, reaching
to 200 f.J.m high. Paraphyses slender, 2- 2.5 f.J.m thick, well branched, anastomosed;
apices not swollen. Asci clavate, 7G-90 X 15- 22 f..l.m. Spores 1-septate, colorless,very often brownish when mature, 2G-25 X 1G-14 f..l.m.
Reaction: thallus & medulla P + reddish, K + yellow, KC - , C - . Chemical
5)

50

Masakane Inoue

substance: stictic acid (TLC).
Habitat: on non-calcareous rocks in subalpine and alpine regions.
Range: Japan; Russian Arctic (Andreev et al. 1996); Europe (Santesson
1993, Purvis et al. 1992); North America (Noble et al. 1987).
Unfortunately I have not reexamined the type of R. hochstetteri. However,I did examine the lee to type of Lee idea colludens which Lamb ( 1940) proposed asa synonym of R. hochstetteri. The Japanese representatives agree well with thetype both anatomically and morphologically. However they lack lichen substances.
R. hochstetteri is distinct due to one-septated spores and a greyish brownthallus, of which areolae are dispersed on a black hypothallus and surrounded bya fimbriate-radiating dendritic protothallus.
Nylander (1890) reported this species (as Lecidea colludens) from Japan,
Mt. Fuji, but I could not find a voucher specimen in Nylander Herbarium at
Helsinki University.
Representative specimens examined. HOKKAIDO. Prov. Soya: Mt. Rishiri,
1700m, M. Inoue no. 8107 (18 Aug. , 1974). Prov. Kamikawa: Mts. Daisetsu,
1860m, M. Inoue no. 8694 (10 Aug. , 1974). HONSHU. Pref. Akita. Mt. Chokai,
1600m, M . Inoue no. 15877 (12 Aug., 1982). Pref. Yamagata. Mt. Chokai,
1400m, M. Inoue no. 16418 (14 Aug. , 1983); 2235m (summit) , M . Inoue no.
16493 (6 Aug., 1983). Pref. Toyama. Mt. Shirouma, 2650m, M . Inoue no. 14146(21 Aug., 1980). Pref. Nagano. Mt. Yari, 2590m, M. Inoue no. 4801 (22 Aug.,1973).
Additional specimens examined. Tirol, Stubaier Alpen, ea. 1200m, leg. M.
Steiner, Poelt's Lich. Alp. 65 (Aug., 1958), in TNS.
Trapelia mooreana (Carroll) P . James(PI. II: b)
In Hertel, Herzogia 3: 404- 405 (1975) . -Lecidea mooreana Carron, Nat.
Hist. Rev. 6: 529 (1859) . Type: Ireland, non vidi.
Thallus medium to thick, whitish with yellow-brown tinge, bullate-areolate;areolae contiguous or, at times, dispersed, surface smooth and polished. Hypothallus indistinct.
Apothecia adnate, reaching to 1.0 mm diameter, markedly constricted at thebase, variable in shape, black or blackish brown; disc concave, sparingly pruinose;
margin thin to medium in thickness, sometimes flexuose, paler than the disc, attimes concolorous to the thallus. Excipulum well developed 6G-80(- 100) ,urn
thick, consisting of parathecium and amphithecium, brown it the margin, colorless within, gradually brownish internally, arachnoid; hyphae radiating, 2.5- 3 ,urn
thick, leptodermatous. Epithecium pale brown. Hymenium 1D0--130 ,urn high.
Hypothecium indistinct, subhymenium distinct, reaching to 150 ,urn high, colorless. Paraphyses slender, 1.5- 2 ,urn thick, richly branched, not thickened at the6)

Little Known Crustose Lichens Bearing Lecideine Apothecia

51

apices. Asci clavate, 9G-120 X 15- 20 JLm, with thin tholus, J - . Spores ( 13- ) 1517 X 6- 8 JLm, with somewhat acute ends.
Reaction: thallus & medulla P - , K - , C + orange. Chemical substance:
gyrophoric acid (TLC).
Habitat: on non-calcareous rocks in subalpine and alpine regions.
Range: Japan; Indonesia (Hertel 1977); U .K. (Coppins & James 1984);
Europe (Hertel 1969, as TrapeZia torelii); Canada (Hertel 1969, as T. torelii);
U.S.A. (Minnesota & Wisconsin, Fink 1910, as Lecidea brujeriana); Venezuela,
Cuba & Costa Rica (Hertel 1970, as Trapelia torelii); Argentina (Hertel 1969, as
TrapeZia torelii) ; Tasmania (Hertel 1987 & 1989) .
T. mooreana is easily recognized by the morphology of the thallus, welldeveloped excipulum being arachnoid (Fig. 4 ) , richly branched slender paraphyses and a production of gyrophoric acid.
This species was previously known from Southeast Asia, Europe, North &South America and Tasmania. However, the range has now extended to East
Asia.
Specimens exmined. HONSHU. Pref. Akita: Hachimantai, 1580m, M.
Inoue no. 24352 (26 July, 1988); Mt. Akitakoma, 1540m, M. Inoue no. 15943 &
15945 (24 July, 1981); Mt. Moriyoshi, 1270m, M. Inoue no. 22097 (20· July,
1990), 1410m, M. Inoue no. 16030 (8 Oct., 1982), 1420m, M. Inoue no. 16021(8 Oct., 1982); Mt. Chokai, 1250m, M. Inoue no. 16534 (14 Aug., 1983); Mt.
Kurikoma, 1230m, M . Inoue no. 16780 (22 Aug. , 1983), 1400m, M. Inoue no.
16708 (22 Aug., 1983) . Pref. Yamagata: Mt. Chokai, 1680m, M . Inoue no.
16881 (20Aug., 1984); Mt. Gassan, 1610m, M. Inoueno. 10822 (24Aug., 1975).

Fig. 4.

Cross-section of an excipulum of Trapelia mooreana (M. Inoue no. 21792), X 100.

52

Masakane Inoue

Pref. Miyagi: Mt. Kurikoma, 1550m, M. Inoue no. 10371 (26 Aug. , 1975) ; Mt.
Zao, 1570m, M . Inoue no. 10693 (21 Aug., 1975). Pref. Nagano: Mt. Naeba,
2145 m, M. Inoue nos. 372-374 (6 Aug., 1972) ; Mt. Asama, 2350m, M. Inoue no.
11329 (23 Aug., 1976); Mt. Norikura, 2430m, M. Inoue no. 12988 & 13002 (14Aug., 1979) .
Additional specimens examined. Ecuador, Zamora-Chinchipe, 2700 m, K .
Kalb. Lich. Neotrop. no. 450, TNS . Brazil, Sanpaulo, K . Kalb., Lich. Neotrop.
no. 248, TNS. Norway, Dammedalane, leg. J. J. Havaas (Sept., 1949), as Biatora
brujeriana , TNS . Scotland, Ardnamurchan Peninsula, ea. 230 m, coli. P. James s.
n. ( 11 July, 1966), as Leciea lopadioides, HIRO. Italy, Rhatische Alpen, Herb. E.
Kernstock, s. n. (coli. in 1892), as Biatora brujeriana f. deplanata , w.
7)

Trapeliopsis ftexuosa (Fr.) Coppins & P. James(PI. II: c)
Lichenologist 16: 258 (1984) . -Biatora.flexuosa Fr., Sched. crit. lich. suec.
part 2 (fasc. 8) : 11 (1826) . Type: Sweden, Fries, Lich. Suec. exs. no. 221. lectotype in UPS, non vidi; isolectotype in M , vidi.
Thallus effuse, bullate-areolate or irregularly areolate; areolae dispersed, or inpart, contiguous, thickness medium, surface more or less polished, livid-gray orgray with green tinge, slightly convex, 0.2-{).3 mm wide; medulla I - . Hypothallus indistinct. Apothecia 0.3-{).5 mm wide, between or partly on the areolae,
adnate, moderately constricted at the base, sometimes several small ones clusteredin groups; disc dark brown to blackish brown, epruinose, plane or concave;margin prominent, flexuose, paler than the disc, or at times concolorous. Excipulum 3o-50 pm thick, brown at the margin, paler within, I - , composed ofthin-walled subradiating hyphae with inflated cell-lumina; hyphae 4- 5 pm thick.
Epithecium pale brown. Hymenium 4o-60 pm high. Subhymenium 3o-40 p mhigh, colorless. Hypothecium reaching 80 pm high, consisting of loosely interwoven hyphae, colorless. Thecium I + pale bluish then turning reddish brown.
Paraphyses slender, 2- 2.5 pm thick, well branched, anastomosed; apices slightly
or not thickened. Asci clavate, 35- 50 X 7- 10 pm, tholus prominent, I + faintly
blue. Spores 8- 9 X 4- 5(6) pm, ellipsoid, colorless.
Reaction: thallus P - , K - , KC - , C - ; medulla P - , K - , C + red.
Chemical substances: gyrophoric acid (TLC).
Habitat: on a carbonized stump burnt by a forest fire and on decayed wood.
Range: Japan; Europe (Schreiner & Hafellner 1992; U . K., Coppins & James1984; Norway, T0nsberg 1992) ; North America (Egan 1987; Alaska, Talbot et al.
1992; British Columbia, Noble et al. 1987); Australia (Weber & Wetmore 1972,
as Lee idea .flexuosa).
Trapeliopsis Hertel & Schneider originally contained only + !- squamulose
taxa (Schneider 1979), but Hertel (1981) and Coppins & P. James (1984) placedsome crustose taxa, like the present species, in this genus. I am inclined to be ofLittle Known Crustose Lichens Bearing Lecideine Apothecia

53

the same opinion as the latter.
Inoue (1994) reported T. flexuosa as a new addition to the flora of Japan andalso of Asia without any descriptions.
The Japanese specimen differs from the type, in which apothecia are up to
0.4 mm wide and the hymenium 35- 45 f.J.m high, only by the somewhat wider
apothecia and higher hymenium. But morphological-, anatomical- and chemical
features as well as a peculiar habitat on "burnt lignum" of the Japanese specimen
are very similar to those of the type and other European representatives.
Diagnostic features of T. jlexuosa are: apothecia with flexuose margin which
is prominent and paler than the disc, excipulum composed of hyphae with inflated
cell-lumina, highly branched and anastomosing paraphyses, rather smaller sporesfor within the lecideoid lichens and a chemistry of producing gyrophoric acid.
External features, especially the apothecia with pale flexuose margin anddispersed bullate-areolate thallus of T. flexuosa , are most reminiscent of somespecies of Trapelia or Fuscidea. However, this species is easily distinguished by
the hyphal structure of excipulum and by the well-branched anastomosing paraphyses.
The saxicolous Trapeliopsis subconcolor (Anzi) Hertel*, which was reported
from Formosa (Taiwan), Nepal and Burma by Hertel (1977, as Trapelia subconcolor) , is easily distinguished from T. flexuosa by the wider apothecia (reaching
1. 6 mm wide) and longer spores (1 o--15 X 4- 5 f.l.m) .
Many reports mentioned that T. flexuosa was growing on old wood, stumps,
logs, and at times on decayed wood. But it is noteworthy that a Japanese
specimen ofT. flexuosa , like the type collection, is growing on a carbonized stump
burnt by a forest fire and on decayed wood. The slow growth of lichens islegendary. But the growth in T. flexuosa seems rapid because this lichen
established in its niche before carbonized layers of wood peeled off. Th. Fries(1874) and Tuckerman (1888) also reported T. flexuosa from the same habitat innorthern Europe and North America respectively.
T. jlexuosa has a world-wide distribution, but its occurrence in Japan seems
to be rather rare; it is known only for one locality in Kyushu, Southern Japan.
Specimen examined. KYUSHU. Pref. Miyazaki: Mt. Ichifusa, 1410m, on
burnt lignum in beech (Fagus crenata) wood, leg. M. lnoue no. 10309 (23 March,
1975) .
Additional specimens examined. Sweden, "Sodermanland", leg. S. Ahlner,
s.n. (22.VII.1951), TNS; "Bohuslan", leg A. H . Magnusson no. 24564 (23.VII.

*

Trap eliopsis subconcolor (Anzi) Hertel, Herzogia 5: 460 (19 81) . - Biatora subconcolorAnzi, Comment. Soc. Critt. Ita!. 1: 151 (1862) . Type: Italy, non vidi.- L ecidea ochrolechioides
Zahlbr. , Feddes Repert. 33: 38 (1 933). Type: Taiwan, leg. Y. Asahina F 155, - holotype in w,
vidi; - isotype in T NS, vidi. - L eeidea obsessa Zahlbr. Feddes Repert. 33: 38 ( 1933). Type:
Taiwan, leg. Y. Asahina F 140, - holotype in w, vidi; - isotype in TNS, vidi. -54

Masakane Inoue1955) , TNS .
Tremolecia atrata (Ach.) Hertel(PI. II: d)
Khumbu Himal 6: 351 (1977) . - Gyalecta atrata Ach. , Kong!. Vetensk.
Akad. Nya Hand!. 229 (1808). Type: Norway, "Finmarkia", 9.VI,1802, G.
Wahlenberg, - lectotype in UPS, vidi.
Based on the collection from the summit of Mt. Fuji Hertel (1977) added
T. atrata to the lichen flora of Japan and gave a precise description; it was the only
locality previously known. However, this species is apparently not rare in Japan,
where it frequently grows in alpine regions.
Habitat: on non-calcareous rocks in alpine regions.
Range: Japan; Himalaya, Armenia, Tanzania, Canary isls., Europe, Iceland,
Spitsbergen, Greenland, Russian Arctic region (Novaja Zemlja, Zemlja Frantsa
losifa, Cukotski Pen.), North America, Venezuela, Chile, Tasmania & New
Zealand & Marion is. (Hertel 1977); South America (Colombia; Argentina,
Hertel 1989); Australia (Rambold 1989) ; Subantarctic region (Kerguelen is.,King George is., South Georgia is. & Antarctic Peninsula, Hertel 1984 & 1987,
Jacobsen & Kappen 1988, Inoue 1993) .
Specimens examined. HOKKAIDO. Prov. Rumoi : Mt. Syokanbetsu, 1320m, M. Inoue no. 8304 (21 Aug., 1974, forming tiny patches with Fuscidea
submollis M . Inoue) . HONSHU. Pref. Toyama: Mt. Shirouma- Mt. Hachidake,
2500m, M. Inoue no. 14164 (21 Aug. 1980) . Pref. Nagana: Yamada Pass near
Mt. Shirane, 2060m, M. lnoue no. 1764 (20 July, 1972) & 2100m, M. Inoue no.
24745 (2 June, 1984); Mt. Yari, 3120m, M. Inoue no. 4934 (21 Aug., 1973); Mt.
Otensho, 2930m, M . Inoue no. 25618 (22 Aug., 1973); Mt. Chogatake, 2600m,
M . Inoue no. 14305 (15 Aug., 1980), 2650m, M. Inoue no. 14319 (15 Aug.,
1980). Pref. Yamanashi: Mt. Kitadake, 2950m, 3030m, 3050m alt. & 3140m,
M. Inoue nos. 12165, 12245, 12202 & 12215 respectively (17 Aug., 1978) ; Mt.
Ainotake, 3150m, M . Inoue no. 12364 (18 Aug., 1978).
Additional specimens examined. Sweden, Harjedalen, 900 m, coli. R. Santesson no. 27393 (Vezda, Lich. sel. Exsic. no. 1536), TNS. Portugal. "Beira Alta",
1150 m, coli. CN. Tavares, 5 Aug., 1962 (Tavares, Lich. Lusitaniae sel. Exsic.
no. 154), as Lecidea dicksonii, TNS . Novaja Zemlja, Salbugtan, coli. Lynge (24Aug. , 1921), as Lecidea dicksonii, TNS . Greenland, Ferslew Pynt, 2- 3 m, E . S.
Hansen, Lich. Groenland. Exsic. no. 311, TNS; Angmassalik, E. S. Hansen, Lich.
Groenland. Exsic. no. 290, TNS.
8)

Acknowledgements
I wish to express my hearty gratitude to the directors and curators of thefollowing herbaria who kindly sent me type and authentic specimens on loan:

Little Known Crustose Lichens Bearing Lecideine Apothecia

55

Prof. Dr. T. Ahti (H), Dr. I. M. Brodo & P.-Y. Wong (CANAL), Prof. Dr. H.
Hertel (M), Dr. H. Kashiwadani (TNS) and Dr. R. Moberg (UPS). This study
was partly supported by a Grant-in-Aid for Scientific Research from the Ministryof Education, Science, Sports and Culture of Japan to M. Inoue, no. 05640782. I
am grateful to Dr. Dianne Fahselt of the Univ. Western Ontario for revising theEnglish text.

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Explanation of plates
Plate I. a. Arthrorhaphis citrinella (Ach.) Poelt (M. Inoue no. 16031) . b. Cecidonia
umbonella (Nyl.) Triebel & Rambold, on the thallus of Lecidea lactea bearing wider
apothecia (M. Inoue no. 12410). c. Epilichen scabrosus (Ach.) Clem. ex Hafellner (M.
Inoue no. 16651) . d. Rhizocarpon hensseniae Brodo (M. Inoue no. 4754) . The ruler isgraduated in 0.5 mm units.
Plate II. a. Rhizocarpon hochstetteri (Kiirb.) Vain. (M. Inoue no. 8694). b. Trapelia
mooreana (Carroll) P. James (M. Inoue no. 21792). c. Trapeliopsisflexuosa (Fr.) Coppins& P. James (M. Inoue no. 10309) . d. Tremolecia atrata (Ach.) Hertel (M. Inoue no.
12165) . The ruler is graduated in 0.5 mm units.

Little Known Crustose Lichens Bearing Lecideine Apothecia

Plate I

57

58

Masakane Inoue

Plate 11