Bull. Natl. Mus. Nat. Sci., Ser. B, 34(1), pp. 1–16, March 21, 2008

Lichens of Akkeshi and its Adjacent Areas,
Northeastern Hokkaido, Japan
Takashi Shiba1, Kwang Hee Moon2 and Hiroyuki Kashiwadani3
1

Narashino City Highschool, 1–2–1 Higashi-narashino, Narashino-shi, Chiba 275–0001, Japan
E-mail: tshibafs@ybb.ne.jp
2
Department of Biological Resources, National Institute of Biological Resources Environmental Research Complex,
Gyoungser-dong, Ser-gu, Incheon 404–170, Korea
3
Department of Botany, National Museum of Nature and Science,
4–1–1 Amakubo, Tsukuba 305–0005, Japan

Abstract A total of 223 lichen taxa representing 66 genera are reported from Akkeshi and its adjacent regions. Among them, Lecanora kurokawae is described as new to science. Leptogium
tenuissimum is newly added to the Japanese lichen flora. Collema callopismum, Hypotrachina
incognita and Parmotrema tinctorum are new to Hokkaido. Results of phytogeographical analysis
show that the lichen flora of the present areas is characterized by high proportions of the Northeast
Asiatic (25.1%) and Pantemperate (39.5%) elements. On the other hand, no taxon belonging to
Bipolar, Pantropical, Indo-Malayan, Pacific, Circum-Pacific and Australian elements is found.
Key words : Akkeshi, Lecanora kurokawae, lichen, Kirittapu, Kushiro, phytogeographical elements.

Introduction

Materials and Methods

Lichens of Akkeshi and the adjacent areas will
be repoted in the present paper. This region includes Kushiro-cho and Hamanaka-cho (about 70
km east to west and 40 km north to south;
42°56– 43°14N, 144°18–145°13E, Fig. 1) andis situated on the Pacific Coast of Hokkaido.
Even though there is no high mountain, thelichen flora of the present region is significantly
notable from the phytogeographical point ofview, since the area is located in northeasternedge of Hokkaido as well as of Japan. Lichens
floras of certain mountains or areas have beenstudied quite well in Japan (Kashiwadani et al.,1987, 1993, 1996, 1998, 2000, 2002; Kurokawa
1957). However, phytogeographical analysis of
the floras in these areas is still insufficient. In thepresent paper, we will try to make a phytogeographical analysis in refernce to phytogeographical elements proposed by Kurokawa (2006).

About 2,000 lichen specimens were collected
at various sites in the present area. This area ismostly covered with natural or secondary forests
composed of deciduous or coniferous trees such
as Abies sachalinensis, Alnus japonica, Acermono, Betula ermanii, B. platyphylla, Fraxinus
lanuginosa, F. mandshurica, Larix leptolepis,
Magnolia kobus, Picea glehnii, Prunus sargentii,
Quercus crispula, Q. dentata, Salix bakko, Sorbus commixta, etc. There are also Bekanbeushi
and Kiritappu marshes and some rocky shores.
The morphology of lichens was observed under
dissecting microscope and light microscope withimmersion lens. Sections of thalli and apothecia
were prepared by hand-razor and mounted in
GAW. Sections of thalli of Lecanora species wereobserved under polarized lens to find glanular
crystals and those of apothecia were observed inconcentrated nitric acid (HNO3) and 10% potassium hydroxide (KOH) replaced for water(Brodo, 1984). Ascus structure was examined in2Takashi Shiba et al.

Lugol’s solution. Secondary products were studied by means of thin-layer chromatography (Culberson and Johnson, 1982), employing only solvent B system. Micro-crystal tests were alsomade, if necessary. All specimens cited in thepresent paper are preserved in the Lichen
Herbarium of the National Museum of Natureand Science (TNS).
Results
Through the morphological and chemical studies, 223 lichen taxa belonging to 66 genera havebeen recognized among the collections. In thefollowing list, they are enumerated in alphabetical order of scientific names, followed by habit,
collection site(s) and collector(s) with collection

number(s). In order to reduce the space, the following abbreviations are used for collection sites
and dates. Collectors are also abbreviated as follows: TS — T. Shiba, HK — H. Kashiwadani and
KM — K.H. Moon. Morphological and/or phytogeographical notes are given, if any. Numbers
after locality abbreviation refer to Fig. 1.
Aikappu (11) ... Around the Marine Laboratoryof Hokkaido University, Cape Aikappu,
Akkeshi, Akkeshi-gun, elevation 0 – 70 m. In
forests dominated by Acer, Abies, Alnus andQuercus. August 4 – 8, 2002 and July 31,
2003.
Ayamegahara (20) ... Ayamegahara pastureland,
Akkeshi, Akkeshi-gun. Elevation 5 – 20 m.
August 6, 2002 and July 30 – 31 and August 1,

Fig. 1. Akkeshi and its adjacent areas showing collection sights for the present study. Aikappu (11), Ayamegahara (20), Bekanbeushi-bridge (15), Bekanbeushi marsh (18), Chirippu to Akkeshi (21), Chomin-no-mori(16), Daikokujima (10), Fukushima (22), Horonai (1), Kiritappu (24), Kitoushi (2), Kokutaiji (12), Konohipark (14), Mabiro (19), Monshizu shurine (8), N shore of Kiritappu marsh (23), Ohbetsu (17), Ohoro (7),Ohta (9), Osonaeyama (13), Osyamai (5), Pontomari (4), Shiriba (6), Tomachise (3).

Lichens of Akkeshi

2003.
Bekanbeushi-bridge (15) ... Around the Bekanbeushi-bridge, Akkeshi, Akkeshi-gun, Exposed rocky area and mixed forest of Abies,
Alnus, Quercus and Salix. Elevation about
70 m. August 9, 2002.
Bekanbeushi marsh (18) ... Bekanbeushi marsh,
Akkeshi, Akkeshi-gun. Elevation 0 – 10 m.
July 27, 2003, September 11, 2003 and August
5, 2004.
Chirippu to Akkeshi (21) ... En route from
Chirippu to Akkeshi, Akkeshi-gun. Elevation
about 50 – 70 m. September 3, 1999.
Chomin-no-mori (16) ... Akkeshi, Akkeshi-gun.
Plantation of Quercus and Acer. Elevation
about 70 m. August 7, 2002 and July 28, 2003.
Daikokujima (10) ... Daikokujima Island, about
8 km off from Aikappu cape. Akkeshi,
Akkeshi-gun. Elevation about 0 – 50 m. July
31, 2003.
Fukushima (22) ... Vicinity of Fukushima,
Biwase, Hamanaka, Akkeshi-gun. Elevation
about 75 m. September 1, 1999.
Horonai (1) ... Around Horonai, Kushiro, Kushiro-gun. Elevation 0 – 100 m. August 7 andOctober 10 – 11, 2004.
Kiritappu (24) ... Cape Kiritappu, about 2.8 km
south of Hamanaka, Akkeshi-gun. Elevation
about 10 m. September 1, 1999.
Kitoushi (2) ... Around Kitoushi, Kushiro, Kushiro-gun. Elevation 0 – 100 m. August 10 and
October 11, 2004.
Kokutaiji (12) ... Around Kokutaiji temple, Honmachi, Akkeshi, Akkeshi-gun. Elevation 5 –20 m. August 4 – 6, 2002.
Konohi-park(14) ... Konohi-park,
Akkeshi,Akkeshi-gun. Elevation about 2 m. July 28,
2003.
Mabiro (19) ... En route from Akkeshi to Kiritappu, Mabiro, Akkeshi, Akkeshi-gun. Cultivatedforest of Abies sachalinensis. Elevation about
70 m. August 3 – 6, 2002, August 4 and October 10, 2004.
Monshizu shrine (8) ... Monshizu. Akkeshi,
Akkeshi-gun. Elevation about 10 m. August 7,
2002.

3

N shore of Kiritappu marsh (23) ... Hillside faced
to Kiritappu marsh, Hamanaka, Akkeshi-gun.
Elevation about 20 – 40 m. September 2, 1999.
Ohbetsu (17) ... Around Mizudori-kansatukan,
Ohbetsu, Akkeshi, Akkeshi-gun. Elevation
about 2 – 10 m. July 27 – 30, 2003.
Ohoro (7) ... Around Ohoro, Akkeshi, Akkeshigun. Elevation about 50 m. October 11, 2004.
Ohta (9) ... Vicinity of Ohta, Akkeshi, Akkeshigun. Cattle farms and pasture land, with scattered trees of Abies, Alnus, Fraxinus and Larix.
Elevation about 70 m. August 6, 2002.
Osonaeyama (13) ... Osonaeyama, Near Akkeshi
Great Bridge, Akkeshi, Akkeshi-gun. Small
hill with mixed forest of Acer, Abies, Alnus
and Quercus. Elevation 30 – 70 m. August 7,
2002 and August 2, 2003.
Osyamai (5) ... Around Osyamai, Kushiro, Kushiro-gun. Elevation 0 – 100 m. August 6, 2004.
Pontomari (4) ... Around Pontomari, Kushiro,
Kushiro-gun. Elevation 0 – 100 m. August 7
and October 10, 2004.
Shiriba (6) ... Cape Shiriba, Kushiro, Kushirogun. Elevation 0 – 100 m. August 6 and October
10, 2004.
Tomachise (3) ... Around Tomachise, Kushiro,
Kushiro-gun. Elevation 0 – 100 m. August 9
and October 10 – 11, 2004.
Enumeration of lichens1) Agonimia pacifica (H. Harada) Diederich.
Rare, over mosses on bark of Abies sachalinensis.
Mabiro, TS 1459; Tomachise, TS 2027.
The present species is also known from Central Honsyu (or main land of Japan).
2) Anaptychia isidiza (Zahlbr.) Kurok.
Occasional, on bark of Abies sachalinensis andBetula ermanii. Aikappu, TS 1422; Ohta, TS 396.
3) Anaptychia palmulata (Michaux) Vain.
Occasional, on bark of Quercus crispula and on
rocks. Aikappu, TS 814 and TS 1423; Shiriba,
HK 47529.
4) Anzia colpota Vain. Locally common,on bark of Alnus japonica, Abies, Quercus andAcer spp. Aikappu, TS 1426; Aikappu, HK
45836.

4

Takashi Shiba et al.

5) Anzia japonica (Tuck.) Müll. Arg.
Occasional, on bark of decayed wood. Fukushima, KM 4089; Shiriba, TS 1945.
6) Anzia opuntiella Müll. Arg. Common,on bark of Acer sp., Alnus japonica and Fraxinus
lanuginosa. Aikappu, TS 174 and TS 823.
7) Brigantiaea ferruginea (Müll. Arg.)
Kashiw. et Kurok. Occasional, on bark of Betula
ermanii, Fraxinus lanuginosa and Quercus
crispula usually with mosses. N shore of Kiritappu marsh, KM 4392; Aikappu, TS 809 and TS
826.
8) Bryoria asiatica (Du Rietz) Brodo et D.
Hawksw. Occasional, on bark of Acer mono.
Tomachise, TS 1997.
9) Bryoria confusa (D.D. Awasthi) Brodo
et D. Hawksw. Occasional, on bark. Shiriba, TS
1918.
10) Bryoria furcellata (Fr.) Brodo et D.
Hawksw. Rare, on bark of Abies sachalinensis.
Aikappu, HK 47513.
11) Bryoria trichodes Brodo et D. Hawksw.
Common, on bark of Abies sachalinensis andSalix subfragilis. Chirippu to Akkeshi, KM 4508;
Horonai, TS 2058; Kitoushi, HK 47413.
12) Calicium abietinum Pers. Occasional,
on decayed wood of Abies sachalinensis. N shoreof Kiritappu marsh, KM 4441; Tomachise, TS
2030.
13) Candelaria concolor (Dickson) B.
Stein. Occacional, on rocks. Kokutaiji, TS 410.
14) Candelariella vitellina (Ehrh.) Müll.
Arg. Occasional, on bark. Ohta, TS 395.
15) Cetrelia braunsiana (Müll. Arg.) W. L.
Culb. et C.F. Culb. Very common, on barks ofAbies sachalinensis, Alnus japonica, Betula
ermanii, Fraxinus lanuginosa and Sorbus
commixta. Chirippu to Akkeshi, KM 4453, KM
4475 and KM 4513; Fukushima, KM 4090 and
KM 4101; Aikappu TS 272; Tomachise, HK
47420.
16) Cetrelia cetrarioides (Delise ex Duby)
W.L. Culb. et C.F. Culb. Occasional, on bark.
Shiriba, TS 1951.
17) Cetrelia chicitae (W.L. Culb.) W.L.
Culb. et C.F. Culb. Common, on barks of Abiessachalinensis and Alnus hirsuta. N shore of Kiritappu marsh, KM 4433 and KM 4439; Aikappu,
TS 274 and HK 45835.
18) Cetrelia japonica (Zahlbr.) W.L. Culb.
et C.F. Culb. Occasional, on bark of Abiessachalinensis. Shiriba, TS 2033; Tomachise, HK
47418.
19) Cetrelia olivetorum (Nyl.) W.L. Culb. et
C.F. Culb. Rare, on bark of Alnus japonica. N
shore of Kiritappu marsh, KM 4382.
20) Cetreliopsis asahinae (M. Sato) Randlane et A. Thell. Occasional, on twigs of Fraxinus lanuginosa and Sorbus commixta. Bekanbeushi bridge, TS 828; Kitoushi, TS 1977.
21) Chaenotheca burunneola (Ach.) Müll.
Arg. Occasional, on bark. Shiriba, TS 1937.
22) Chrysothrix candelaris (L.) J.R. Laundon. Occasional, on bark of Larix leptolepis.
Kokutaiji, TS 2067.
23) Cladonia alpina (Asahina) Yoshim.
Rare, restricted on high moor. Bekanbeushi
marsh, TS 844.
24) Cladonia arbuscula (Wallr.) Rabenh.
subsp. beringiana Ahti. Restricted on high moor,
among mosses on soil. Bekanbeushi marsh, TS
1352.
25) Cladonia caespiticia (Pers.) Flörke.
Occasional, on ground. Horoto, HK 47503.
26) Cladonia chlorophaea (Flörke ex Sommerf.) Spreng. Occasional, among mosses onrocks. Bekanbeushi bridge, TS 1276.
27) Cladonia coniocraea (Flörke) Spreng.
Common, on trunk base of Betula ermanii, Fraxinus mandshurica or on soil. Aikappu, TS 835
and TS 1404.
28) Cladonia cornuta (L.) Hoffm. subsp.
cornuta. Common, on soil, decayed wood andtrunk base. Ohbetsu, TS 840.
29) Cladonia farinacea (Vain.) A.W. Evans.
Occasional, on rocks, Aikappu, TS 832.
30) Cladonia furcata Hoffm. Common, on
decayed wood and on soil. Fukushima, KM 4126
and KM 4165; Kiritappu, KM 4200.
31) Cladonia glauca Flörke. Occasional, on
rocks. Bekanbeushi bridge, TS 1873.
32) Cladonia grayi G.K. Merr. ex Sandst.

Lichens of Akkeshi

Occasional, on soil. Bekanbeushi marsh, TS 848.
33) Cladonia hondoensis Asahina. Locallycommon, on soil. Kiritappu, KM 4204.
34) Cladonia humilis (With.) Laundon.
Occasional, on soil. Aikappu, TS 849.
35) Cladonia kurokawae Ahti et Steenroos.
Occasional, over humus on soil. Ohbetsu, HK
45816.
36) Cladonia macilenta Hoffm. Occasional,
over humus on soil, trunk base of Abies sachalinensis and Betula ermanii. Aikappu, TS 854;
Mabiro, TS 834; Bekanbeushi marsh, HK 45852.
37) Cladonia ochrochlora Flörke. Common,on barks of Abies sachalinensis and Alnus japonica, on decayed woods, trunk base and overhumus on soil. Chirippu to Akkeshi, KM 450; N
shore of Kiritappu marsh, KM 4931 and KM
4404; Aikappu, TS 346.
38) Cladonia pleurota (Flörke) Schaer.
Locally common, on soil. Kiritappu, KM 4196.
39) Cladonia ramulosa (With.) Laundon.
Common, on trunk base of Larix leptolepis, onsoil and on rocks. Aikappu, TS 836 and HK
45829; Bekanbeushi bridge, TS 851.
40) Cladonia rei Schaer. Occasional, on
rocks and over humus on soil. Aikappu, TS 830;
Ohbetsu, HK 45821.
41) Cladonia scabriuscula (Delise in Duby)
Leight. Occasional, on rocks. Aikappu, TS 854.
42) Cladonia squamosa (Scop.) Hoffm. var.
squamosa. Occasional, on bark of Abies sachalinensis. Aikappu, HK 45811; Shiriba, TS 2001.
43) Coccocarpia erythroxyli (Spreng.) Swinscow et Krog. Rare, on bark of Acer mono.
Kitoushi, HK 47409.
44) Coccocarpia palmicola (Spreng.) Arvids.
et Gall. Occasional, on bark of Abies sachalinensis and on rocks. Mabiro, TS 964.
45) Collema callopismum A. Massal. Veryrare, on concrete base, Aikappu, TS 868 and HK
45807.
Collema callopismum is a saxicolous species
found in calcareous areas, being widely distributed in the Northern Hemisphere. It has been reported from Sweden, Scotland, England, France,
Portugal, Germany, Poland, Italy, Yugoslavia,

5Greenland, Canada and Japan (Degelius, 1954,1974). In Japan, however, it has been reported
from only two localities; Mt. Myouhougatakeand Mt. Buko, in Prov. Musashi, central Honshu(Degelius, 1954). In the present area, it was
found on concrete base of marine laboratory of
Hokkaido University, Akkeshi. This is the firstrecord for this species in Hokkaido, and is thenorthern most locality in Japan. The specimen
from the present area has non-isidiate and semicrustose thallus with pulvinate lobes, subparaplectenchymatous proper exciples, ellipsoid andsubmuriform ascospores 25 – 3210 – 17 m m insize. These characters coincide well with those ofauthentic or exsiccata specimens of C. callopismum preserved in TNS.
Other Japanese specimens examined. Honshu.
Prov. Musashi (Pref. Saitama): Mt. Buko,
Chichibu. On rock (limestone), elevation about
1,000 m, November 3, 1969, H. Shibuichi 3993
(TNS); Mt. Myouhougatake, on summit of Mt.
Myouhougatake, Mitsumine, Chichibu, on concrete, elevation about 1,332 m, June 25, 1967, S.
Kurokawa 67001 and 67002 (TNS).
46) Collema coccophyllum Nyl. Rare, onbark of Fraxinus mandshurica var. japonica.
Ohbetsu, HK 45805.
This is one of rare species of Collema distributed in East Asia, having been reported fromJapan, India and Thailand (Degelius, 1974). In
Japan, it was reported from two localities in Honshu by Degelius (1974) and Kashiwadani et al.
(1996). In the present survey, we found the third
locality for this species, where it grows on barkof Fraxinus mandshurica var. japonica at Ohbetsu, Akkeshi-gun. The specimen has membranousand fenestrate lobes, small apthecia to 1 mm indiam., paraplectenchymatous proper exciples andglobose to subglobose ascospores with 4 – 5 cells
8 – 107 – 8 m m in size. This is the northern
most locality for this species in Japan.
Other Japanese specimens examined. Honshu.
Prov. Etchu: Fukuno-cho, Higashitonami-gun, on
bark, March 15, 1930, M. Nishijima 20 (TNS).
Prov. Iwashiro (Pref. Fukushima) Mt. Yahazu,
Azuma Mts., Yama-gun, on bark of Fagus crena-

6

Takashi Shiba et al.

ta, elevation 1,400 – 1,500 m, May 31, 1995, H.
Kashiwadani 38418 (TNS); Wasezawa, Kitashiobara-mura, Yama-gun, on bark of Juglans mandshurica var. sachalinensis, elevation about 830
m, August 22, 1995, K.H. Moon. 468 (TNS).
47) Collema complanatum Hue. Occasional,
on barks of Alnus japonica and Salix subfragilis
or on rocks. Ohta, TS 297; Horoto, HK 47429.
48) Collema japonicum (Müll. Arg.) Hue. f.
japonicum. Occasional, on barks of Acer monoand Kalopanax pictus. Shiriba, TS 1921;
Kitoushi, TS 1975.
49) Collema leptaleum Tuck. var. leptaleum.
Rare, among mosses on bark of Picea glehnii.
Chirippu to Akkeshi, KM 4451.
50) Collema subflaccidum Degel. Occasional, on bark of Alnus japonica and Fraxinus
mandshurica var. japonica. Chirippu to Akkeshi,
KM 4454; Shiriba, TS 1625.
51) Dibaeis baeomyces (L. f. ) Rambold etHertel. On ground. Horoto, HK 47502.
52) Dimerella lutea (Dickson) Trevisan.
Occasional, among mosses on bark of Abiessachalinensis in shaded condition. Mabiro, TS
907; Mabiro, HK 47444; Horonai, TS 1822.
53) Dimerella pineti (Ach.) Vězda. Occasional, on bark of Picea glehnii. Chirippu to
Akkeshi, KM 4499; Shiriba, TS 1998.
54) Dolichousnea diffracta (Vain.) Articus.
Abundant, on bark of Abies sachalinensis.
Mabiro, TS 493; Kitoushi, HK 47416.
55) Dolichousnea longissima (Ach.) Articus. Occasional, on bark of Abies sachalinensis.
Aikappu, TS 296.
56) Dolichousnea trichodeoides (Vain.)
Articus. Occasional, on bark of Abies sachalinensis. Mabiro, TS 932.
The present species can be distinguished bybranches flattened with longitudinal-furrows on
the surface from D. longissima.
57) Evernia mesomorpha Nyl. Occasional,
on bark of Abies sachalinensis. Kokutaiji, TS
290.
58) Evernia prunastri (L.) Ach. Rare, onbark of Abies sachalinensis. N shore of Kiritappu
marsh, KM 4368; Kokutaiji, TS 409.

Although this species is widely distributed in
the Northern Hemisphere, it has been collected atonly few localities in Hokkaido (Provinces
Ishikari, Kitami and Kushiro).
59) Fissurina inabensis (Vain.) M. Nakan.
et Kashiw. in Nakanishi et al. Rare, on bark ofAbies sachalinensis. Shiriba, TS 1648.
This species is endemic to Japan, having beenreported from south of middle Honshu (Nakanishi, 1966).
60) Flavoparmelia caperata (L.) Hale. Frequent, on bark of Larix leptolepis. Kokutaiji, TS
485.
61) Graphis cognata Müll. Arg. Occasional,
on bark of Betula ermanii. Aikappu, TS 920.
62) Graphis rikuzensis (Vain.) M. Nakan.
Common, on barks of Abies sachalinensis, Phellodendron amurense and deciduous trees. Chirippu to Akkeshi, KM 4464; Kokutaiji, TS 414;
Mabiro TS 1357.
63) Graphis scripta (L.) Ach. Common, onbark of Abies sachalinensis. Aikappu, TS 164.
64) Graphis tenella Ach. Occasional, onbark of Abies sachalinensis. Fukushima, KM
4116; Aikappu, HK 46261; Kokutaiji, TS 1217.
65) Hafellina disciformis (Fr.) Marbach et
H. Mayrhofer (Marbach 2000). Very common, on
barks of Picea glehnii, Abies sp. and Quercus sp.
Fukushima, KM 4150; Kitoushi, HK 47404.
66) Heterodermia boryi (Fée) K.P. et S.R.
Singh. Very rare, on bark. Kitoushi, TS 1971.
67) Heterodermia diademata (Taylor) D.D.
Awasthi. Occasional, on bark of Quercus crispula. Aikappu, TS 422.
68) Heterodermia hypochraea (Vain.) Swinscow et Krog. Occasional, on bark of Abiessachalinensis. Aikappu, HK 45819; Horonai, TS
1820.
69) Heterodermia
hypoleuca(Muhl.)
Trevisan. Occasional, on bark of Alnus sp., Abies
sp. or rarely on rocks. Aikappu, TS 819 and HK
45828.
70) Heterodermia
isidiophora(Vain.)
Skorepa. Occasional, on bark of Fraxinus mandshurica var. japonica. N shore of Kiritappu
marsh, KM 4262.

Lichens of Akkeshi71) Heterodermia japonica (Sato) Swinscow et Krog. Rare, among mosses on bark ofAbies and on rocks. Shiriba, TS 2005.
72) Heterodermia microphylla (Kurok.)
Swinscow et Krog. Common, on barks of Betula
ermanii and Larix leptolepis and on rocks.
Chirippu to Akkeshi, KM 4473; Fukushima, KM
4085; Aikappu, TS 802.
73) Heterodermia obscurata (Nyl.) Trevisan. Rare, among mosses on bark of Salix sp.
Chirippu to Akkeshi, KM 4477.
74) Heterodermia propagulifera (Vain.) J.P.
Dey. Rare, among mosses on bark of Abiessachalinensis. N shore of Kiritappu marsh, KM
4227.
75) Heterodermia tremulans (Müll. Arg.) W.
L. Culb. Rare, once collected on bark of Betula
ermanii. Aikappu, TS 805.
76) Hypogymnia fragillima (Hillm.) Rass.
Occasional, on barks of Abies sachalinensis andSorbus commixta. Fukushima, KM 4103 and KM
4125; Shiriba, TS 1634.
77) Hypogymnia hokkaidensis Kurok. Veryrare, on bark of Picea sp. Shiriba, TS 1964.
This species is one of rare species in Japan,
having been collected at five localities in Hokkaido (Provinces Hidaka, Ishikari, Kushiro andTokachi).
78) Hypogymnia pseudoenteromorpha (Ach.)
Nyl. f. inactiva (Asahina) Asahina. Locally common, on barks of Abies sachalinensis anddecayed wood. Fukushima, KM 4901, KM 4124
and KM 4174.
79) Hypogymnia pseudophysodes (Asahina)
Rass. var. pseudophysodes. Occasional, on barkof Abies sachalinensis and rocks. N shore of
Kiritappu marsh, KM 4216; Aikappu, TS 1107
and TS 1138.
80) Hypogymnia pulverata (Nyl.) Elix.
Occasional, on bark of Picea glehnii. Fukushima,
KM 4148.
81) Hypogymnia submundata (Oxsner) Rass.
f. submundata. Occasional, on bark of Abiessachalinensis. Aikappu, TS 251.
82) Hypogimnia vitata (Ach.) Parrique f.
vittata. Occasional, on bark of Abies sachalinen-

7

sis. Fukushima, KM 4098; Mabiro, TS 1381.
83) Hypotrachyna incognita (Kurok.) Hale.
Rare, on bark of Prunus yezoensis. Kokutaiji,
HK 45806.
This species is distinguished from allied
species of the genus by having cylindrical isidia,
dichotomously branched rhizines and white
medulla containing gyrophoric and protolichesterinic acids. It is one of rare lichens in Japan,
having been collected at only three localities in
Nagano and Miyazaki Prefectures. This is thenorthern most locality for this species in Japan.
New to Hokkaido.
Other Japanese specimens examined. Honshu.
Prov. Shinano (Pref. Nagano): Mt. Takabottchi,
Shiojiri-city, on bark of Larix leptolepis, elevation about 1,600 m, October 29, 1986. H. Kashiwadani 23783 and 34428 (TNS), Kyusyu. Prov.
Hyuga (Pref. Miyazaki): Minamikata, Saito-city,
on bark of Quercus glauca, elevation 670 –765 m, February 23, 2004, K.H. Moon. 7346
(TNS); Shiratori, Ebino-shi, on trunk of Pinus
densiflora, elevation about 900 m, July 9, 1994,
H. Shibuichi 9546 (TNS).
84) Japewia subaurifera Muhr et Tonsberg.
Occasional, on bark of Picea sp. Aikappu, TS
1342.
85) Lecanora chionocarpa Hue. Occasional,
on bark of Fraxinus mandshurica. Ohbetsu,
HK45804.

Fig. 2. Lecanora kurokawae Shiba, K.H. Moon &Kashiw. sp. nov. Scale bar1 mm

8Takashi Shiba et al.

86) Lecanora kurokawae Shiba, K.H. Moon
& Kashiw. sp. nov. (Fig. 2)
Thallus ut in Lecanora cenisia sed subfarinose-sorediatus et acidum porphyrilicum continentus, discis epruinosis et excipulis marginalibus cum crystalis minimis vel granularis differt.
Type collection. Japan, Hokkaido, Prov. Kushiro: Takoiwa, Pontomari, Kushiro-gun. On rocks,
along coast; elevation about 2 m, October 10,
2004. T. Shiba 2055 (holotype in TNS).
Thallus saxicolous, whitish gray, consisting ofhemispherical or globose verrucae (0.2 – 0.4 mm
in diam., 0.2 – 0.5 mm high); verrucae covered
with gelatinous cortex, cortex prosoplectenchymatous 12 – 15 m m thick, gonidial layer subcontinuous, 35 – 40 m m thick, gonidia 6 – 8 m m indiam., upper medulla composed of densely intertangled hyphae, lower medulla composed ofloosely intangle hyphae, containing numerous
granular crystals, verrucae are completely filledwith HNO3 soluble crystals together with loose
hyphae towards the center but are composed ofdensely intangled hyphae and algal celles towards the cortex. Soredia subfarinose (0.02 –0.03 mm in diam.), whitish-gray, formed near the
top of verrucae (Fig. 2). Apothecia common, sessile, 1.0 – 1.3 mm in diam.; disc reddish brown,
epruinose, margins of apothecia concolorouswith the thallus; thalline margin containing small
crystals which are scattered in medulla as well as
inner cortex (allophana-type by Brodo, 1984);
crystals are soluble by HNO3 but insoluble by
KOH; epihymenium not granulate, brown (glabrata type by Brodo, 1984); hymenium 80 – 100 m mhigh; hypothecium 40 – 50 m m thick; proper
exciple 60 – 120 m m thick; asci clavate, 30 – 40
3 – 4 m m; ascospores ellipsoid, 17 – 185 – 6 m m.
Pycnidia not seen.
Chemical substances: atranorin (minor),chloroatranorin (minor), porphyrilic acid (major),
1a -hydroxyporphyrilic acid (trace).
Lecanora kurokawae is unique in Japanesesaxicolous species of Lecanora in having verrucose thalli with soredia. It resembles L. cenisia
Ach., a species widely distributed in North
America and Europe, because they both havesaxicolous habitat, verrucose thalli and similar
spores. However, it can be easily distinguishedfrom the latter by the presence of soredia, epruinose disc, and by the presence of porphyrilic
acid as a major chemical substance.
This species was collected on granites atwater’s edge along the coast. It covers widely onrock surface together with species of Acarospora
and Caloplaca.
Other specimen examined. Pontomari, HK
47436 (TNS).
87) Lecanora sibirica Müll. Arg. Frequent,
on bark of Quercus crispula. Aikappu, TS 159.
88) Leptogium asiaticum Jørg. Occasional,
on bark of Fraxinus lanuginosa. Kitoushi, TS
1981.
89) Leptogium azureum (Sw.) Mont. Occasional, on barks of Fraxinus mandshurica var.
japonica and Juglans mandshurica var. sachalinensis and on decayed wood. Chirippu-mura to
Akkeshi, KM 4456; N shore of Kiritappu marsh,
KM 4298, KM 4401 and KM 4409.
90) Leptogium burnetiae Dodge. Occasional, on bark of Acer mono. Aikappu, TS 893 andHK 45834.
91) Leptogium pedicellatum Jørg. Occasional, on barks of Acer mono and Fraxinus mandshurica. Aikappu, TS 355; Ohbetsu, HK 45803.
92) Leptogium saturunum (Ach.) Nyl.
Occasional, on bark of Quercus crispula. Aikappu, TS 496.
93) Leptogium tenuissimum (Dickson)
Körber. Rare, on soil. Pontomari, TS 2054;
Tomachise, HK 47497.
Although Leptogium tenuissimum is widelydistributed in the Northern Hemisphere, it has
never been reported from Japan. In the present
survey, we found this species at one locality
along the coast, where it grows among mosses onground. This is the first record for this species inJapan as well as in Eastern Asia.
Specimens from the present area have spreading lobes, forming grayish brown cushions; lobes
up to 1.5 mm wide, 10 – 12 mm high, elongate
and deeply incised with crenate margin; thulli arecomposed of paraplectenchymatous tissue with-

Lichens of Akkeshi

out fibrous hyphae. Although the Japanese specimen is sterile, it is quite identical with exsiccate
specimens kept in TNS.
Kashiwadani et al. (2002) reported Leptogium
subtile (Schrader) Torss. from Mt. Kasagatake in
Prov. Kohzuke. However, the specimen has characteristic features written above and can be better
identified with this species.
Other specimen examined. Honshu. Prov.
Kohzuke: (Pref. Gunma): around the summit of
Mt. Kasagatake, elevation 2,000 – 2,050 m, October 11, 2001, M. Inoue 30569 and 30570 (TNS).
Exsiccati examined. V. Räsanen: Lich. Fenn.
Exs. Mus. Bot. Univ. Helsink. ed. 116 (TNS). J.
P. Norrlin and W. Nylander: the same above, 356(TNS).
94) Leptogium trichophoroides P.M. Jørg. et
Wallace. Occasional, on bark of Quercus crispula. Aikappu, TS 496.
95) Lobaria adscripturiens (Nyl.) Hue. f.
adscripturiens. Occasional, on bark of Taxus
cuspidata. Shiriba, TS 1671.
96) Lobaria crassior Vain. Rare, on bark of
Alnus japonica. N shore of Kiritappu marsh, KM
4381.
97) Lobaria discolor (Bory) Hue. var.
discolor. Occasional, on barks of Quercus sp.
and Salix sp. Aikappu, TS 960.
98) Lobaria fuscotomentosa Yoshim. Occasional, on barks of Acer sp. and Quercus sp.
Aikappu, TS 328.
99) Lobaria japonica (Zahlbr.) Asahina.
Common, on bark of Quercus crispula. Aikappu,
TS 158.
100) Lobaria linita (Ach.) Rabenh. Occasional, on barks of Betula ermanii, Quercus
crispula and Salix sp. Aikappu, TS 169 and TS
962; Ohbetsu, TS 965.
101) Lobaria orientalis (Asahina) Yoshim.
Occasional, on bark of Fraxinus mandshurica
var. japonica. Ohbetsu, HK 45832; Mabiro, TS968.
102) Lobaria quercizans Michaux. Occasional, on bark of Acer mono. Chomin-no-mori, TS
184; Shiriba, TS 1672.
103) Lobaria retigera (Bory) Trevis. var.

9

retigera. Occasional, on bark. Shiriba, TS 1917.
104) Lobaria sachalinensis Asahina. Occasional, on bark of Acer mono. Kitoushi, TS 1968.
105) Lobaria spathulata (Inum.) Yoshim.
Occasional, on barks of Alnus japonica andQuercus crispula. Aikappu, TS 150, TS 909 andTS 1253.
106) Lobaria tuberculata Yoshim. Occasional, on bark of Alnus japonica. Aikappu, TS 326
and TS 966; Mabiro, HK 47445.
107) Lopadium disciforme (Flot.) Kulh.
Occasional, on bark of Abies sachalinensis.
Aikappu, HK 46260.
108) Loxospora ochrophaea (Tuck.) R.C.
Harris. Occasional, on barks of Alnus japonica
and Fraxinus lanuginosa. N shore of Kiritappu
marsh, KM 4400 and KM4426; Osyamai, TS
1693.
109) Megalospora tuberculosa (Fée) Sipman.
Occasional, on bark of Quercus crispula. Aikappu, TS 982.
110) Melanelixia huei (Asahina) O. Blanco,
A. Crespo, Divakar, Essl., D. Hawksw. et Lumbsh. Common, on barks of Fraxinus lanuginosaand Fraxinus mandshurica var. japonica. Aikappu, TS 974; Ohbetsu, HK 45849.
111) Melanohalea olivacea (L.) O. Blanco,
A. Crespo, Divakar, Essl., D. Hawksw. et Lumbsh. Rare, on bark of Salix subfragilis. Horoto,
HK 47522.
112) Menegazzia primaria Aptroot et al.
Very rare, once reported from Kushiro (Moon etal., 2006), on bark of Abies sachalinensis.
Horonai, HK 47459.
113) Menegazzia terebrata (Hoffm.) A. Massal. Common, on bark of Abies sachalinensis.
Aikappu, TS 267; Horoto, HK 47508.
114) Myelochroa entotheiochroa (Hue) Elixet Hale. Occasional, on barks of Alnus japonica
and Quercus crispula. Aikappu, TS 152 and TS
973; Osonaeyama, TS 971.
115) Myelochroa irrugans (Nyl.) Elix etHale. Occasional, on barks of Alnus japonica andFraxinus lanuginosa. Aikappu, TS 899; Kitoushi,
HK 47467.
116) Myelochroa leucotyliza (Nyl.) Elix et10Takashi Shiba et al.

Hale. Occasional, on barks of Abies sachalinensis and Quercus crispula. N shore of Kiritappu
marsh, KM 4334; Aikappu, TS 975.
117) Myelochroa metarevoluta (Asahina)
Elix et Hale. Occasional, on rocks and on bark of
Fraxinus lanuginosa. Konohi-park, TS 1021;
Ohbetsu, HK 45850.
118) Nephroma bellum (Spreng.) Tuck.
Occasional, on rocks. Aikappu, TS 983.
119) Nephroma resupinatum (L.) Ach. Occasional, on bark of Fraxinus lanuginosa. N shoreof Kirittapu marsh, KM 4396; Shiriba, TS 1684;
Tomachise, TS 2034.
120) Nephromopsis endocrocea Asahina.
Occasional, on barks of Abies sachalinensis andFraxinus lanuginosa. Aikappu, TS 157, TS 228
and TS 301.
121) Nephromopsis ornata (Müll. Arg.) Hue.
Occasional, on bark. Shiriba, TS 2008.
122) Nephromopsis rugosa Asahina. Occasional, on bark. Aikappu, TS 858.
123) Normandina pulchella (Borrer) Nyl.
Occasional, on bark of Alnus japonica. Aikappu,
TS 814b.
124) Ochrolechia akagiensis Yasuda. Occasional, on barks of Abies sachalinensis and Quercus crispula. Mabiro, TS 988; Shiriba, HK 47431.
125) Ochrolechia parellula (Müll. Arg.)
Zahlbr. Occasional, on rocks. Bekanbeushi
bridge, TS 1296.
126) Ochrolechia trochophora (Vain.) Oshio.
Common, on barks of Abies sachalinensis, Acer
sp. and Fraxinus lanuginosa. Aikappu, TS 991;
Mabiro, TS 989 and TS 990.
127) Ochrolechia yasudae Vain. Ooccasional,
on bark of Quercus crispula. Aikappu, TS 151.
128) Oropogon asiaticus Asahina. Rare, on
bark. Shiriba, TS 1624.
129) Pannaria lurida (Mont.) Nyl. subsp.
lurida. Occasional, on barks of Acer mono andAlnus japonica and on decayed wood. Aikappu,
TS 1034; Kitoushi, HK 47480; Monshizu, TS419.
130) Parmelia fertilis Müll. Arg. Common,on bark of Abies sachalinensis and on rocks.
Kokutaiji, TS 1020; Monsizu, TS 477.

131) Parmelia laevior Nyl. Common, on barkof Alnus japonica. Aikappu, TS 224 and TS1030.
132) Parmelia praesquarrosa Kurok. Veryfrequent, on barks of Alnus japonica, Betula
ermanii. Aikappu, TS 1019 and TS 1029;
Osonaeyama, TS 402.
133) Parmelia squarrosa Hale. Frequent, onbarks of Abies sachalinensis, Alnus japonica andPicea glehni. Aikappu, TS 269; Mabiro, TS 1371
and TS 1190; Bekanbeushi marsh, TS 1356.
134) Parmotrema chinense (Osbeck) Hale etAhti. Locally abundant, on twigs of Betula andon barks of Alnus japonica and Quercus
crispula. Kiritappu, Arakawa 1472 and Arakawa
1573.
135) Parmotrema clavuliferum (Räsänen)Streimann. Rare, on bark. Kokutaiji, TS 1228.
136) Parmotrema reticulatum (Taylor) M.
Choisy. Common, on barks of Abies sachalinensis, Acer mono and Betula ermanii and on rocks.
Aikappu, TS 1453 and TS 1455; Kokutaiji, TS
1233 and HK 45813.
137) Parmotrema tinctorum (Nyl.) Hale.
Very rare, on bark of Abies sachalinensis. N
shore of Kiritappu marsh, KM 4234.
This species is widely distributed in tropical totemperate areas in the world. In Japan, it has
been recorded from south of Aomori Prefecture
but has not been reported from Hokkaido. In thepresent survey, we found this species at hillsideof Kiritappu marsh. This is the northern most
locality for this species in Japan.
138) Peltigera collina (Ach.) Schrad. Rare,on bark of Quercus crispula. Shiriba, HK 47515.
139) Peltigera didactyla (With.) J.R. Laundon. Occasional, on ground along trail. Pontomari, TS 1731.
140) Peltigera dilacerata (Gyeln.) Gyeln.
Occasional, on soil or on trank base. Aikappu,
TS 163 and TS 1069.
141) Peltigera dolichorrhiza Nyl. Occasional,
on soil. Pontomari, TS 1881.
142) Peltigera horizontalis (Huds.) Baumg.
Occasional, on soil. Aikappu, TS 1069.
143) Peltigera leucophlebia (Nyl.) Gyelnik.

Lichens of Akkeshi

Occasional, on soil in mixed forest dominated byAcer, Abies, Alnus and Quercus. Aikappu, TS
1070.
144) Peltigera praetextata (Flörke) Vain.
Occasional, on soil. Aikappu, TS 1068.
145) Peltigera rufescens (Weiss.) Humb.
Occassional, on soil. Aikappu, TS 1096.
146) Pertusaria carneopallida (Nyl.) Anzi ex
Nyl. Rare, on bark of Sorbus commixta. Fukushima, KM 4189.
147) Pertusaria commutata Müll. Arg. Occasional, on bark of Abies sachalinensis and on
rocks. N shore of Kiritappu marsh, KM 4247;
Aikappu, TS 155 and HK 45825.
148) Pertusaria composita Zahlbr. Occasional, on bark of Betula ermanii. Aikappu, TS 1062.
149) Pertusaria corallina (L.) Arnold. var.
corallina. Occasional, on bark of Betula ermanii.
Aikappu, TS 972.
150) Pertusaria ferruginea (Müll. Arg.)
Oshio. Rare, on bark of Abies sachalinensis.
Fukushima, KM 4716.
151) Pertusaria lactea (L.) Arnold. (Räsänen
1940a, Oshio 1968). Occasional, on bark ofAbies sachalinensis and on rocks. Mabiro, TS
1124; Aikappu, TS 1044.
152) Pertusaria
leucosoraNyl.
var.
leucosora. Common, on barks of Abies sachalinensis, Alnus japonica and Betula ermanii.
Aikappu, TS 1132 and TS 1133, Mabiro, TS1061.
153) Pertusaria mendax Müll. Arg. Locallycommon, on bark of Abies sachalinensis. N shoreof Kiritappu marsh, KM 4222.
154) Pertusaria multipuncta (Turner) Nyl.
Occasional, on bark of Alnus japonica. Aikappu,
TS 262.
155) Pertusaria oshioi J.C. Wei. Locallycommon, on barks of Abies sachalinensis andAlnus japonica. N shore of Kiritappu marsh, KM
4316 and KM 4318.
156) Pertusaria pertusa (L.) Tuck. Occasional, on bark of Fraxinus lanuginosa. Osonaeyama,
TS 1059.
157) Pertusaria pustulata (Ach.) Duby. Rare,on bark of Abies sachalinensis. Chrippu to

11

Akkeshi, KM 4458.
158) Pertusaria quartans Nyl. Rare, on
rocks, Aikappu, HK 45826.
159) Pertusaria subcomposita Oshio. Occasional, on bark of Alnus japonica. Aikappu, TS
462; Daikokujima, TS 1055.
160) Pertusaria subfallens Vain. Occasionalon bark of Abies sachalinensis. Aikappu, TS1035.
161) Pertusaria sublaeviganda Vain. Occasional, on bark of Betula ermanii. Aikappu, TS1145.
162) Pertusaria submultipuncta Nyl. Occasional, on rocks. Aikappu, TS 1038.
163) Pertusaria variolina Nyl. Occasional,
on barks of Quercus crispula and Salix sp. andon rocks. Osonaeyama, TS 953; Ohbetsu, TS1375; Horonai, TS 1786.
164) Pertusaria velata (Turner) Nyl. Occasional, on rocks. Aikappu, TS 1268 and HK
45818.
165) Phaeographis fuscodisca M. Nakan.
Rare, on bark of Abies sachalinensis. Aikappu,
HK 46270.
166) Phaeophyscia hirtuosa (Krempelh.)Essl. Occasional, on bark of Fraxinus
lanuginosa. Aikappu, TS 1142.
167) Phaeophyscia hispidula (Ach.) Moberg.
Occasional, on bark of Salix subfragilis. Horoto,
HK 47501.
168) Phaeophyscia imbricata (Vain.) Essl.
Occasional, on bark of Quercus crispula. Aikappu, TS 1378.
169) Phaeophyscia limbata (Poelt) Kashiw.
Occasional, on rocks. Aikappu, TS 334; Kokutaiji, TS 1092.
170) Phaeophyscia rubropulchra (Degel.)Moberg. Occasional, on bark. Osonaeyama, TS
1135.
171) Phaeophyscia trichophora (Hue) Essl.
Occasional, on bark of Salix sp. Aikappu, TS
1281; Ohbetsu, HK 45847.
172) Physcia aipolia (Ehrh. ex Humb.) Furnr.
Rare, on bark of Alnus japonica. N shore of Kiritappu marsh, KM 4347.
173) Physcia caesia (Hoffm.) Hampe in12Takashi Shiba et al.

Furnr. Occasional, on concrete floor. Aikappu,
HK 45808.
174) Physcia dubia (Hampe) Lettau. Occasional, on bark of Larix leptolepis. Ohta, TS 187.
175) Physcia phaea (Tuck.) Thomson. Occasional, on bark of Alnus japonica. Daikokujima,
TS 1085.
176) Physcia stellaris (L.) Nyl. Occasional,
on barks of Alnus japonica and Salix sp. Kokutaiji, 1093, Daikokujima, TS 1159; Ohbetsu, HK
45848 and TS 1110.
177) Physciella melanchra (Hue) Essl. Common, on bark of Abies sachalinensis and on
rocks. Aikappu, HK 45810 and HK 45820.
178) Physconia detersa (Nyl.) Poelt. Occasional, on rocks. Osonaeyama, TS 1141.
179) Physconia grumosa Kashiw. et Poelt.
Occasional, on bark of Quercus crispula. Aikappu, TS 1109; Kitoushi, HK 47477.
180) Physconia hokkaidensis Kashiw. Occasional, on bark. Aikappu, TS 1892.
181) Platismatia interrupta W.L. Culb. et C.
F. Culb. Occassional, on bark of Abies sachalinensis. Aikappu, TS 287; Mabiro TS 490.
182) Polychidium dendriscum (Nyl.) Henssen.
Occasional, among mosses on bark. Shiriba, TS
1924.
183) Porpidia albocaerulescens (Wulfen)
Hertel et Knoph. var. albocaerulescens. Occasional, on rocks. Aikappu, TS 1270.
184) Pseudocyphellaria crocata (L.) Vain.
Occasional, on bark of Salix sp. Shiriba, TS 1931.
185) Punctelia borreri (Sm.) Krog. Occasional, on bark of Salix sp. Ohta, TS 207.
186) Punctelia rudecta (Ach.) Krog. Occasional, on bark. Kokutaiji, TS 1072.
187) Pyrenula japonica Kurok. Common, onbark of Abies sachalinensis. Aikappu, TS 248.
188) Pyxine sorediata (Ach.) Mont. Occasional, on decayed wood and bark of Acer mono.
Aikappu, TS 1077.
189) Ramalina commixta Asahina. Rare, onbark of Fraxinus lanuginosa. Horonai, HK
47464.
190) Ramalina conduplicans Vain. Common,on barks of Abies sachalinensis and Salix sub-fragilis, and on rocks. Aikappu, HK 45817;
Kitoushi, HK 47411; Ohoro, HK 47424.
191) Ramalina dilacerata (Hoffm.) Hoffm.
Occasional, on bark of Quercus dentata. Aikappu, TS 3912b.
192) Ramalina geniculata Hook. et Tayl. var.
globiformis Räsänen. Rare, on bark of Salix subfragilis. Kitoushi, HK 47410.
193) Ramalina hokkaidensis Kashiw. Occasional, on barks of Abies sachalinensis, and Salix
subfragilis. Ohoro, HK 47426 and HK 47430;
Shiriba, TS 1262.
194) Ramalina pertusa Kashiw. Occasional,
on barks of Alnus japonica and Salix subfragilis.
Kiritappu, HK 47510; Ohoro, HK 47423.
195) Ramalina pollinaria (Wester.) Ach.
Occasional, on rocks. Bekanbeushi bridge, TS407.
196) Ramalina roesleri (Hochst.) Nyl. Occasional, on bark of Abies sachalinensis. Aikappu,
HK 45833; Kitoushi, HK 47478.
197) Ramalina shinanoana Kashiw. Rare, on
bark. Kokutaiji, TS 993.
198) Ramalina sinensis Jatta. Occasional, onbark of Sorbus commixta. Ohbetsu, HK 45822.
199) Ramalina subbreviuscula Asahina.
Occasional, on concrete basses. Daikokujima, TS
1001.
200) Sticta fuliginosa (Dickson) Ach. Occasional, on bark of Fraxinus lanuginosa.
Tomachise, TS 2036 and HK 47490.
201) Sticta nylanderiana Zahlbr. Occasional,
on bark of Taxus cuspidata. Shiriba, TS 1670.
202) Sticta weigelii (Ach.) Vain. Rare, on
bark. Shiriba, TS 1913.
203) Sticta wrightii Tuck. Occasional, onbark of Abies sachalinensis. Aikappu, TS 283.
204) Sticta yatabeana Müll. Arg. Occasional,
on bark of Acer mono. Shiriba, TS 1909.
205) Tephromela atra (Huds.) Hafellner. Frequent, on bark of Betula ermanii. Aikappu, TS
1331.
206) Thelotrema lepadinum (Ach.) Ach.
Rare, on bark of Betula ermanii. Kitoushi, HK
47473.
207) Thelotrema nipponicum Tat. Matsumo-

Lichens of Akkeshi

to. Occasional, on bark of Abies sachalinensis.
Shiriba, TS 1645.
208) Thelotrema similans Nyl. Occasional,
on bark of Alnus japonica. Daikokujima, TS
1053.
209) Thelotrema subtile Tuck. Occasional, onbark of Abies sachalinensis. Mabiro, HK 47478.
210) Tuckneraria pseudocomplicata (Asahina) Randlane et Saag. Occasional, on bark of
Quercus crispula. Pontomari, TS 1740.
211) Usnea cornuta Körb. subsp. cornuta.
Rare, on bark of Abies sachalinensis. Fukushima,
KM 420.
212) Usnea dasaea Stirt. Occasional, on barkof Abies sachalinensis. Mabiro, TS 923.
213) Usnea fragilescens Lynge. Rare, on barkof Abies sachalinensis. Mabiro, TS 921.
This species has also been known from Western Europe and British Columbia. It is characterized by inflated branchs and excavated soraria
with granular soredia.
214) Usnea glabrata (Ach.) Vain. Occasional, on bark of Abies sachalinensis. Kitoushi, TS
1703.
215) Usnea mutabilis Stirton. Occasional, onbark of Abies sachalinensis. Mabiro, 925.
216) Usnea pangiana Stirton. Occasional, onbark of Abies sachalinensis. Aikappu, TS 229.
217) Usnea pygmoidea (Asahina) Y. Ohmura.
Rare, on bark of Salix subfragilis. Kitoushi, HK
47414b.
218) Usnea rubicunda Stirton. Occasional,
on bark of Abies sachalinensis. Aikappu, HK
47512.
219) Usnea rubrotincta Stirton. Occasional,
on bark of Abies sachalinensis. Aikappu, HK
45827.
220) Usnea subfloridana Stirton. Occasional,
on bark of Abies sachalinensis. Kokutaiji, TS
483.
221) Usnea wasmuthii Räsänen. Occasional,
on bark of Abies sachalinensis. Kokutaiji, TS
484.
222) Xanthoria fallax (Hepp) Arnold. Rare,on rocks. Chirippu to Akkeshi, Arakawa 1360.
223) Xantoria mandschurica (Zahlbr.) Asahi-

13na. Occasional, on rock (conclete). Aikappu, HK
45809.
Phytogeographical features
As shown above, 223 taxa of lichens have beenrecognized. They certainly belong to different
phytogeographical elements composing Japaneselichen flora. With regard to lichen flora of Japan,
various phytogeographical elements have been
proposed by Jørgensen (1983), Moon (1999),
Kashiwadani et al. (2000), Harada (2001) andKurokawa (2006). Among them, phytogeographical elements proposed by Kurokawa (2006) seemto be most reliable. According to him, 16 elements are recognized as composing lichen floraof Japan. In the present area, ten of them are recognized as shown below and taxa belonging eachof those elements are also listed in the following
list, referring their world distributions known atpresent.
1. Holarctic Element
Only one species (0.4%), Cladonia alpina,
belongs to this element. Even though the present
area situated in northeastern most Hokkaido, it isquite natural that the proportion of holarctic
element is so low, since ther is no high mountain.
2. Alpine Element
One species (0.4%), Cladonia farinacea,
belongs to the present element.
3. Boreal Element
According to Kurokawa (2006), lichens distributed in boreal zone in the Northern andSouthern Hemispheres belong to the present element. The following 31 taxa (13.9%) are foundin the present area: Agonimia pacifica, Bryoria
confusa, B. furcellata, B. trichodes, Cetrelia
cetrarioides, Cladonia glauca, C. grayi, C. rei,Dibaeis baeomyces, Dolichousnea longissima,
Evernia mesomorpha, Hafellia disciformis,
Hypogimnia pseudophysodes var. pseudophysodes,
H. submundata f. submundata, H. vitata, Japewia
subaurifera, Leptogium tenuissimum, Lobaria

14

Takashi Shiba et al.

linita, Melanoholea olivacea, Nephromopsis
rugosa, Pertusaria lactea, P. multipuncta, P.
pertusa, Physcia aipolia, P. dubia, P. phaea,
Ramalina roesleri, Sticta weigelii, Usnea
fragilescens, U. subfloridana and Xanthoria
fallax.
4. Beringian element
One species (0.4%), Cladonia arbuscula
subsp. beringiana belongs to the present element.
5. Eastern Asiatic-North American Disjunctive element
As shown by Kurokawa (2006), Anaptychia palmulata, Cetrelia chicitae, Heterodermia hypoleuca,
Lobaria quercizans, Loxsospora ochrophaea, Myerochroa metarevoluta, Ochrolechia trichophora, O.
yasudae and Parmelia sqarrosa belong to the present element. The following three species foundin the present area can be also concidered tobelong to the present element; Cladonia humilis,
Hypogimnia pseudoenteromorpha and Phaeophyscia rubropuchra. In total, 12 taxa (5.4%)
belong to the present element.
6. Northeast Asiatic Element
When he proposed 16 phytogeographical elements for the Japanese lichen flora, the following
18 taxa, which also occur in the present area,
were enumerated as belonging to the present element. They are Anzia japonica, A. opuntiella,
Brigantiaea ferruginea, Cetreliopsis asahinae,
Collema complanatum, Dolichousnea diffracta,
Heterodermia microphylla, L. sachalinensis,
Melanelixia huei, Myerochroa irrugans, M. leucotyliza, Nephromopsis endcrocea, Parmelia fertilis, P. laevior, P. praesqarrosa, Platismatia
interrupta, Pyrenula japonica and Xanthoria
mandshurica. On the other hand, the following
taxa found in the present area can be also considered to belong to the present element; Anaptychia
isidiza, Anzia colpota, Calicium abietinum,
Cetrelia japonica, Cladonia kurokawae, Evernia
prunastri, Hypogymnia fragillima, H. pulverata,
Lecanora chionocarpa, L. sibirica, Lobaria
japonica, Lobaria orientaris, L. tuberculata,

Menegazzia primaria, Oropogon asiaticus,
Peltigera didactyla, P. dolichorrhiza, Pertusaria
composita, P. oshioi, P. quertans, P. subcomposita, P. subfallens, P. submultipuncta, Phaeophyscia imbricata, Physconia grumosa, Ramalina commixta, R. geniculata var. globiformis, R.
pertusa, R. polinaria, R. shinanoana, R. subbreviuscula, Sticta wrightii, Sticta yatabeana, Tuckneraria pseudocomplicata, Usea dasaea, U. pangiana, U. pygmoidea and U. rubrotincta. Thus,
56 taxa (25.1%) occurring in the present area
belong to the Northeast Asiatic element.
7. Pantemperate Element
Among the taxa occurring in the present area,
the following 10 taxa were reported as membersof the present element by Kurokawa (2006);
Candelaria concolor, Chaenotheca brunneola,
Cladonia furcata, C. scabriuscula, C. squamosavar. squamosa, Flavoparmelia caperata, Menegazzia terebrata, Porpidia albocaerulescens var.
albocaerulescens, Punctelia borreri and Sticta
fuliginosa. The following 78 taxa, occurring in
the present area, can be added to this element;
Candelariella vitellina, Cetrelia olivetorum,
Chrysothrix candelaris, Cladonia caespiticia,
C. chlorophaea, C. coniocraea, C. cornuta subsp.
cornuta, C. hondoensis, C. macilenta, C.
ochrochlora, C. pleurota, C. ramulosa, Coccocarpia erythroxyli, C. palmicola, Collema callopismum, C. cocophyllum, C. japonicum f. japonicum, C. leptaleum var. leptaleum, C. subflaccidum, Dimerella lutea, D. pineti, Dolichousnea
trichodeoides, Graphis scripta, G. tenella, Heterodermia boryi, H. diademata, H. isidiophora,
H. japonica, H. obscurata, H. propagulifera, H.
tremulans, Hypotrachina incognita, Leptogium
azureum, L. burrnetiae, Lobaria crassior, Lopadium disciforme, Megarospora tuberculosa, Myerochroa entotheiochroa, Nephroma respinatum,
Normandina pulchella, Pannaria lulida, Parmotrema chinensis, P. clavriferum, P. reticulatum,
P. tinctorum, Peltigera collina, P. dilacerata, P.
horizontaris, P. leucophlebia, P. praetextata, P.
rufescens, Pertusaria carneopallida, P. corallina,
P. leucosora, P. pustulata, P. variolina, P. velata,

Lichens of Akkeshi

Phaeophyscia hirtuosa, P. hispidula, P. trichophora, Physcia caesia, P. stellaris, Physciellamelanchra, Physconia deterusa, Polychidium
dendriscum, Pseudocyphellaria crocata, Puncteria rudecta, Pyxne sorediata, Ramalina dilacerata, R. sinensis, Tephromela atra, Terotrema lepadium, T. subtile, Usnea cornuta subsp. cornuta,
U. glabrata, Usnea mutabilis, U. rubicunda andU. wasmuthii. In total, 88 taxa (39.5%) belong to
the Panteperate element.
8. Sino-Japanese Element
Among taxa listed above, the following 14
taxa (6.3%) can be considered to the Sino-Japanese element. They are Bryoria asiatica, Cetrelia
braunsiana, Heterodermia hypochraea, Leptogium asiaticum, L. pedicellatum, L. saturunum,
L. trichophoroides, Lobaria adscripturiens f.
adscripturiens, L. fuscotomentosa, L. spathulata,
Nephromopsis ornata, Phaeophyscia limbata,
Ramalina conduplicans and Sticta nylanderiana.
Among them, only one species, Bryoria asiatica,
was treated as a member of Sino-Japanese element by Kurokawa (2006).
9. Endemic Element
At present, following 17 taxa (7.6%) found in
the present area are considered to be endemic toJapan: Fissurina inabensis, Graphis cognata, G.
rikuzensis, Hypogimnia hokkaidensis, Lecanora
kurokawae, Nephroma bellum, Ochrolechia
akagiensis, O. parellula, Pertusaria commutata, P.
ferruginea, P. mendax, Pertusaria sublaeviganda,
Phaeophyscia fuscodisca, Physconia hokkaidensis, Ramalina hokkaidensis, Thelotrema nipponicum and T. similans. As pointed out by
Kurokawa (2006), some of them are not neccessarily endemic to Japan and may be found in theadjacent areas such as China, Korea and eastern
Siberia in future.
10. Paleotropical Element
Two taxa (0.9%), Lobaria discolor var. discolor and L. retigera var. retigera, belong to the present element, as stated by Kurokawa (2006).

15

The lichen flora of Akkeshi and the adjacentarea is composed of the Holarctic, Alpine, Boreal, Beringian, Northeastern Asiatic-North American Disjunctive, Northeast Asiatic, Pantemperate, Sino-Japanese, Endemic and Paleotropicalelements. It is characterized by high proportionsof the Northeast Asiatic (25.1%) and Panteperate(39.5%) elements. Rather high propotion of
Northeastern Asiatic-North American Disjunctive element is probably related to the geographical location of the present area which is situatedon the midway of the southwards migration of
Arcto-Tertiary flora. On the other hand, it is also
noteworthy that none of the taxa belonging to the
Bipolar, Pantropical, Indo-Malayan, Pacific, Circum-pacific and Austrarian elements is found in
the present area.
Acknowledgements
We wish to express our sincere gratitude to Dr.
S. Kurokawa of the Botanic Gardens of Toyama,
Japan, for his valuable guidance and critical reading. My deep appreciation is also expressed to
Mr. H. Shibuichi, Dr. Y. Ohmura of the National
Instutute for Environmental Studies who helped
us identifying species of Pertusaria and Usnea
respectively. We also thank to Dr. M. Nakanishi
of Okayama University of Science, who gave us
valuavle advices for the family Graphidaceae, toDr. J. Elix of Australian National University Canberra for the determination of chemical substances of Lecanora kurokawa and to Dr. H.
Mukai of Marine Laboratory of Hokkaido University for kind help to our field works. We also
thank to Mr. T. Shibuya of Akkeshi Waterfall
Observation Center, who carried out fildworks
with us and helped us in varous ways. This study
was partly supported by a Grant-in Aid provided
by town office of Akkeshi-cho.
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