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New or Noteworthy Species of Lichens Found in Akita Prefecture, Japan

Hiroyuki Kashiwadania, André aptrootb, Minoru naKanishic, Kwang Hee Moond,*, Masakane inouee and Osanori sasaKif

aDepartment of Botany, the National Museum of Nature and Science, 4-1-1, Amakubo, Tsukuba, 305-0005 JAPAN;

bABL Herbarium, G.v.d.Veenstraat 107, NL-3762 XK Soest, THE NETHERLANDS;cFaculty of Education, Hiroshima University, 1-1-1 Kagamiyama, Higashihiroshima, 739-8524 JAPAN;dNational Institute of Biological Resources, Gyoungser-dong, Seo-gu, Incheon, 404-170 KOREA;

eFaculty of Education and Human Studies, Akita University, 1-1, Tegata Gakuen-machi, Akita, 010-8502 JAPAN;

fShizenkagakuchosajimusho Co., 102, Yachizoi, Tomaki, Daisen, 014-0044 JAPAN*Corresponding author: lichenkr@korea.kr

(Accepted on February 16, 2015)

Five species of pyrenocarpous lichens and one species of the genus Graphis are reported from Akita Prefecture, northern Japan, and their distribution ranges are presented. Among them, Graphis fulvescens M. Nakan., Kashiw. & K. H. Moon is new to science. It resembles G. kousyuensis M. Nakan., but it differs in having smaller spores and concealed discs. Agonimia opuntiella (Buschardt & Poelt) Vĕzda and Porina dolichophora (Nyl.) Müll. Arg. are new records for Japan. Additional localities are presented for Julella sericea (A. Massal.) Coppins, Pyrenula nitidella (Flörke ex Schaer.) Müll. Arg., and Pyrenula tokyoensis (Müll. Arg.) H. Harada, which have been found only in a few localities in Japan.

Key words: Agonimia opuntiella, Akita, Graphis fulvescens, Japan, Julella sericea, new record, new species, Porina dolichophora, Pyrenula nitidella, Pyrenula tokyoensis.

J. Jpn. Bot. 90: 227–234 (2015)

As a part of an investigation of the lichen flora in Akita Prefecture, northern Japan, we conducted field work in Higashinaruse-mura, Ogachi-gun from November 2012 to November 2013. The investigated area is located in the south-east of the prefecture, at elevations between 530 and 620 meters (Fig. 1). It is roughly covered by second-growth mixed forest where deciduous broad-leaved trees are commonly found (e.g., Acer distylum, A. micranthum, A. pictum subsp. mayrii, Aesculus turbinata, Alnus hirsuta var. sibirica, Betula ermanii, Cornus controversa, Fagus crenata,

Ilex macropoda, Quercus crispula, and Tilia maximowicziana) mixed with scattered Pinus parviflora var. pentaphylla and Thujopsis dolabrata. Planted Cryptomeria japonica is also found.

Among lichens collected in the present area, the following new or noteworthy species were found: Agonimia opuntiella (Buschardt & Poelt) Vĕzda, Graphis fulvescens M. Nakan., Kashiw. & K. H. Moon, Julella sericea (A. Massal.) Coppins, Porina dolichophora (Nyl.) Müll. Arg., and Pyrenula tokyoensis (Müll. Arg.) H. Harada. Graphis fulvescens is described as a new species.

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Agonimia opuntiella and Porina dolichophora are new records for Japan. P. tokyoensis, a species found only in the Kanto and Chubu regions, central Japan, is reported here as the northern most locality for this species.

The specimens used for the present study are kept in the herbarium of the National Museum of Nature and Science, Tsukuba (TNS).

1) Agonimia opuntiella (Buschardt & Poelt) Vĕzda in Lich. Rar. Exs. 33: 4, no. 330 (1997).

[Fig. 2A]During the present survey, Kashiwadani and

Sasaki found a tiny specimen of this species among mosses (Boulaya mittenii) on bark of Quercus crispula, at an elevation of 548 m. It has a greenish gray squamulose thallus composed of slightly convex lobes (0.20–0.40 mm long, 0.08–0.10 mm wide) covered with hyaline hairs; lobes totally composed of paraplectenchymatous tissue with papillate cells on both upper and lower cortices. In addition, it produces no chemical substance. Although it is a sterile specimen, these characteristic features coincide well with those found in an exsiccata specimen of Agonimia opuntiella (Lendemer, Lich. East. North Amer. Exs. 137, TNS).

Agonimia opuntiella is widely distributed in the world, having been reported from Australia, Europe, Central and North America, South America and East Asia including Papua New Guinea, Korea, and Taiwan (Aptroot 2003,

2011, Moon and Aptroot 2009). In Japan, it has never been reported before; it appears to have been overlooked because of its small size and the almost unnoticeable color of the thallus.

Specimen examined. JAPAN. Honshu. Prov. Ugo (Pref. Akita): Ainomatazawa, Higashinaruse-mura, Ogachi-gun, on bark of Quercus crispula, elevation 548 m, November 6, 2013, H. Kashiwadani 51327 (TNS).

2) Graphis fulvescens M. Nakan., Kashiw. & K. H. Moon, sp. nov. [Figs. 2B–E]

MycoBank No.: MB 811513.Species similar to Graphis kousyuensis, but

differs by the concealed disc, the 4(–6)-spored asci and the smaller spores, 32–35 × 12–13 µm in size.

Thallus corticolous, pale yellowish brown, opaque, corticated, 0.3–0.5 µm thick. Apothecia (Figs. 2B, C) simple or sparsely branched, up to 3 mm long, 0.24 mm wide; lirellae erumpent; labia entire, with lateral thalline margin; disc concealed, but more or less narrowly open in part; disc epruinose; exciples carbonized laterally approximately 50–75% of the total, 40–60 µm thick (Fig. 2D); hymenium clear, I−, 100–110 µm high. Asci 4–6-spored; spores hyaline (I+ blue), muriform, kidney-shaped (Fig. 2E), with 1–2 locules in one row, 28–30 × 10–12 µm in size.

Type collection: JAPAN. Honshu. Prov. Ugo (Akita Prefecture): Ainomatazawa, Higashinaruse-mura, Ogachi-gun, on bark of Acer amoenum var. matsumurae, elevation 548 m, November 6, 2013, H. Kashiwadani 51323 (TNS–holotype).

Chemistry: no chemical substance.Graphis fulvescens is characterized by simple

or sparsely branched lirellae with entire labia that have a lateral thalline margin (the deserpens morph sense Lücking et al. 2009, Figs. 2B & 2C), concealed disc, exciples carbonized laterally approximately 50–75% of the total (Fig. 2D), clear hymenium without inspersion, 4–6-spored asci with kidney-shaped muriform spores with 8–10 rows of 3–5 locules (Fig. 2E), 28–30 × 10–12 µm in size, and by the absence

Fig. 1. Position of Higashinaruse-mura, Akita Pref., Japan.

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of chemical substance.In general appearance, it resembles G.

kousyuensis (Horik. & M. Nakan.) Lücking, a species described from Japan (Type collection: JAPAN. Honshu, Prov. Kai, Mt. Mizugaki, August 5, 1953, S. Kurokawa 521101 HIRO–holotype; TNS–isotype!); however, it is clearly distinguished from the latter by the concealed discs, 4–6-spored asci, and smaller and kidney-shaped spores as cited above. Although G. kousyuensis has concealed discs, they are often open and produce 1-spored asci containing larger spores with 18–20 rows of 2–6 locules, 50–60 × 18–20 µm in size. It might be confused with G. alpestris (Zahlbr.) Staiger, a species reported from China, as they both show similar carbonization in the exciples; however, the latter species differs in having open discs with pruina and much larger spores with 25–30 rows of 2–6 locules, 100–120 × 25–28 µm in size.

This species is found at two localities in Akita and Shiga Prefectures, Honshu. At the first locality, it grows on the bark of Acer amoenum var. matsumurae together with Dicranum viride at an elevation of about 550 m. At the second locality in Shiga Prefecture, it grows on the bark of Quercus crispula at elevations between 1070 and 1120 m.

Other specimen examined: JAPAN. Honshu. Prov. Ohmi (Pref. Shiga): Mt. Kanakuso-dake, Takayama-cho, Nagahama-city, on bark of Quercus crispula, elevation 1070–1120 m, June 10, 2014, H. Kashiwadani (no. 51430, TNS) & K. Kobayashi.

3) Julella sericea (A. Massal.) Coppins in Lichenologist 24: 367 (1992). [Fig. 2G]

This species belongs to Thelenellaceae and is characterized by having a thin crustose thallus, whitish gray in color; black and solitary perithecia with dark brown involucrellum, 0.3–0.6 mm in diameter; branched and anastomosed paraphyses; and ellipsoid and submuriform spores, with 5–7 rows of 1–2 locules, 13–18 × 8–10 µm in size.

Julella sericea is widely distributed in the Northern Hemisphere including Europe, Canada,

and North America. In Japan, the occurrence of this species was reported only once from Hokkaido by Kashiwadani et al. (2009). During the present survey it was found on bark of Acer.

Specimen examined. JAPAN. Honshu. Prov. Ugo (Pref. Akita): Korogeyama, Higashinaruse-mura, Ogachi-gun, on bark of Acer distylum, elevation 530–600 m. July 3, 2013, H. Kashiwadani 51136 (TNS).

4) Porina dolichophora (Nyl.) Müll. Arg. in Bot. Jahrb. 6: 400 (1885). [Figs. 2F, 3A–B]

Basionym. Verrucaria dolichophora Nyl. in Annls Sci. Nat., Bot. 20: 245 (1863).

Type collection: NOVA GRANATA (Colombia). Villeta, on bark, A. Lindig 2591 (H–lectotype!, designated by McCarthy 1993).

The specimen from the present area has the following diagnostic characters: thallus crustose, superficial, pale fawn, smooth or slightly cracked, matt; perithecia solitary, prominent, hemispherical, not constricted at base (Fig. 3A), approximately 70 to 80% covered by thallus at the total, 0.24–0.30 mm in diameter; involucrellum thin, dark brown, determinate at the top (Fig. 3B, iv); true exciples pale yellow, 40–60 µm thick; paraphyses simple, not anastomosed; spores colorless, filiform, straight to slightly curved, with 25–35 transverse septa, 95–112 × 3–5 µm in size (Fig. 2F). No chemical substance detected by TLC.

The perithecia of the present species appear to be rather variable in size. McCarthy (1993) reported larger perithecia (to 0.83 mm in diameter) in the lectotype; however, he also observed smaller perithecia in a syntype (Lindig 2582, H!). The results of taxonomic study by Aptroot including type specimens revealed that the present specimen is identical with Porina dolichophora.

Porina dolichophora is distinct from all other Japanese species of Porina in its filiform spores with 16–21 transverse septa. It resembles P. kennedyensis P. M. McCarthy, a species described from Australia (McCarthy 2008), in having acicular spores with 11–19

230 植物研究雑誌　第 90巻　第 4号 2015年 8月

Fig. 2. A. Agonimia opuntiella (Buschardt & Poelt) Vĕzda, general view, H. Kashiwadani 51327 (TNS). B–E. Graphis fulvescens M. Nakan., Kashiw. & K. K. Moon, H. Kashiwadani 51323 (holotype, TNS), general view (B), close up of lirellae (C), cross section of an apothecium (D) and submuriform spores (E). F. Porina dolichophora (Nyl.) Müll. Arg., kidney-shaped spore, H. Kashiwadani 51078 (TNS). G. Julella sericea (A. Massal.) Coppins, general view, H. Kashiwadani 51136 (TNS). Scales: A = 200 µm, B, C and F = 1 mm, D = 100 µm, E = 10 µm.

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Fig. 3. A–B. Porina dolichophora (Nyl.) Müll. Arg., H. Kashiwadani 51078 (TNS), general view (A), cross section of a perithecium with narrow involucrellum (iv) at apex (B). C–D. Pyrenula nitidella (Flörke ex Schaer.) Müll. Arg. H. Kashiwadani 51319 (TNS), general view (C), cross section of a perithecium, showing red pigments at base (D, arrow). E–G. Pyrenula tokyoensis (Müll. Arg.) H. Harada, H. Kashiwadani 51003 (TNS), general view (E), ascospores (F), a part of an accurate drawing of P. tokyoensis on a piece of paper by late Dr. Y. Asahina (G; drawn from a specimen collected at Yatsumi, Chiba Pref., Y. Asahina s.n., TNS). Scales: A, C and E = 1 mm, B = 100 µm, D = 50 µm, F = 10 µm.

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septa and corticolous habit; however, the latter species differs by the dark brown involucrellum extending to half way down along the sides.

This is the first report for the species outside the New World. In Japan, it grows on bark of Acer pictum subsp. mayrii in more or less shady conditions.

Specimen examined: JAPAN. Honshu. Prov. Ugo (Pref. Akita): Tokusazawa, Higashinaruse-mura, Ogachi-gun, on bark of Acer pictum subsp. mayrii, elevation 455–500 m, July 2, 2013, H. Kashiwadani 51078 (TNS).

5) Pyrenula nitidella (Flörke ex Schaer.) Müll. Arg. in Bot. Jahrb. 6: 414 (1885). [Figs. 3C–D]

Pyrenula nitidella is characterized by single perithecia with an apical ostiole (Fig. 3C), absence of pigments in the thallus (UV−), 3-septate spores with separated terminal lumina from exospore wall, clear hamathecium and the presence of red pigments (K+ purple, Fig. 3D, arrow) in the hamathecium.

The occurrence of this species in Japan was first reported by Kashiwadani et al. in 2009 based on specimens collected in Hokkaido and central Honshu (Pref. Fukushima). In the present area it grows on barks of Acer and Fagus. In Japan, therefore, this species may grow in deciduous forests, at elevations between approximately 500 to 950 m. This is the third record for the species in Japan.

Specimens examined: JAPAN. Honshu. Prov. Ugo (Pref. Akita): Akataki, Higashinaruse-mura, Ogachi-gun, on bark of Fagus crenata, elevation 496 m, November 5, 2013, H. Kashiwadani 51319 (TNS); Akagawa, Higashinaruse-mura, Ogachi-gun. On bark of Acer pictum subsp. mayrii; elevation 503 m, November 5, 2013, H. Kashiwadani 51397 (TNS).

6) Pyrenula tokyoensis (Müll. Arg.) H. Harada in Harada, Okamoto & Yoshimura, Lichenology 2: 132 (2004). [Figs. 3E–G]

This species is characterized by corticolous habit, 3–6 radially aggregated perithecia with a shared ostile; perithecia are erumpent, covered by thallus or almost exposed (Fig. 3E), pale brown needle-shaped spores with 11–15 transverse septa, 50–70 × 4.5–6 µm in size (Fig.

3F), and by the absence of chemical substance. Pyrenula tokyoensis, an endemic species

of Japan, was first described from Japan by Müller (1892) under the name Pyrenastrum tokyoense Müll. Arg., on the basis of a specimen collected in Tokyo [Type collection: JAPAN. Prov. Musashi (Tokyo Metrop.), Tokyo (without locality detail), May 24, 1891, R. Yatabe 244 (G–holotype; TNS–isotype!)]. It was well revised taxonomically by several authors (Asahina 1931, Harada 2008 and Vainio 1921; 3F) and has been reported from a few localities in central Honshu, Japan (Chiba and Shizuoka Prefectures and Tokyo Metropolitan).

In the present area, it was also found on bark of Alnus hirsuta var. sibirica, at an elevation of 550 m. In addition, one specimen collected at Mt. Taisetsu in Hokkaido and two specimens collected at Hikone and Kohka cities in Shiga Prefecture are also identified with this species. They grow on barks of Alnus japonica and Quercus serrata. The present records show that it is widely distributed from Hokkaido to Shiga Prefecture in Japan, probably at lower elevations but possibly overlooked.

Specimen examined: JAPAN. Hokkaido, Prov. Ishikari: Mt. Daisetsu (without precise locality), August 10, 1932, F. Fujikawa s.n. (TNS). Honshu. Prov. Ugo (Pref. Akita): Tokusazawa, Higashinaruse-mura, Ogachi-gun, on bark of Alnus hirsuta var. sibirica at elevation 550 m, July 1, 2013, H. Kashiwadani 51003 (TNS). Prov. Kazusa (Pref. Chiba): Yatsumi, December 23, 1923, Y. Asahina s.n. (TNS). Prov. Ohmi (Pref. Shiga): Kohyama, Shigaraki-cho, Kohka-city, on bark of Quercus serrata, elevation about 320 m, June 10, 2014, H. Kashiwadani 51441 (TNS); Sasao, Hikone-city, on bark of Alnus japonica, elevation about 220 m, December 10, 2014, H. Kashiwadani 51539 (TNS).

We wish to express our sincere thanks to Dr. H. Higuchi, National Museum of Nature and Science, Tsukuba for his identification of mosses. Thanks are extended to Dr. S. Y. Yoon, Baekseok University, Cheonan city for her corrections of our English manuscript. This study was partly supported by a grant to K. H. Moon from the National Institute of Biological

August 2015 TheJournal of Japanese Botany Vol. 90 No. 4 233

Resources, Korea (NIBR 201401105) funded by the Ministry of Environment of the Republic of Korea.

ReferencesAptroot A. 2003. Pyrenocarpous lichens and related non-

lichenized ascomycetes from Taiwan. J. Hattori Bot. Lab. 93: 155–173.

Asahina Y. 1931. Materials for a lichen flora of Japan I. Rep. Saito Gratitude Found. 12. Sendai, 94 pp + 22 figs.

Harada H. 2008. Taxonomic notes on pyrenocarpous lichens in Japan (5). Further notes on Pyrenula concatervans and P. tokyoensis. Lichenology 7(2): 169–172 (in Japanese).

Kashiwadani H., Aptroot A. and Moon K. H. 2009. New pyrenocarpous lichens in Japan, with the resurrection of

the genus Trypetheliopsis for Musaespora. Bibliotheca Lichenologica 99: 247–258.

McCarthy P. M. 1993. Saxicolous Species of Porina Müll. Arg. (Trichotheliaceae) in the Southern Hemisphere. Bibliotheca Lichenologica, 52. 134 pp. J. Cramer, Berlin.

McCarthy P. M. 2008. A new species of Porina (Porinaceae) from Queensland. Austr. Lichen. 62: 29–31.

Moon K. H. and Aptroot A. 2009. Pyrenocarpous lichens in Korea. Bibliotheca Lichenologica 99: 297–314.

Müller J. A. 1892. Lichenes Yatabeani. Nuovo Giorn. Bot. Ital. 24: 182–202.

Nylander W. 1863. Florae Novo-Granatensis. Annals Sci. Nat., Bot., sér. 4, 20: 228–256.

Vainio W. 1921. Lichenes insularum Philippinarum III. Ann. Acad. Sci. fenn., Ser. A 15 (no. 6): 209. 1921.

柏谷博之 a，A. Aptrootb，中西　稔 c，文　光喜 d，井上正鉄 e，佐々木長徳 f：秋田県で見つかった地衣類の新種と希少種　筆者等は秋田県雄勝郡東成瀬村で 2012–2013年に地衣類の現地調査を実施した．採集した標本中に 1新種をはじめ，日本新産種や希種を認めたので報告する．　1) Agonimia opuntiella (Buschardt & Poelt) Vĕzda（ケツブゴケ，新称）：日本産ツブゴケ属にはマユゴケとツブゴケの 2種が報告されているが，本種は第三の種である．地衣体は微小な鱗片状で新鮮なものは黄緑色，標本庫に保管すると淡褐色となる．裂片は長さ 0.20–0.40 mm，幅 0.08–0.10 mm，表面に淡色の毛を密生するので，他の 2種とは容易に区別できる．地衣体内部は偽柔組織となり，皮層の分化は見られない．細胞に特異な乳頭状突起を持つ．　本種は世界中に広く分布し，アジアでは韓国，台湾，パプアニューギニアに産することが知られているが，日本からは報告されていなかった．地衣体が非常に小さく目立たないので，見過ごされている可能性が高い．今回の調査中，海抜 485 mのミズナラ樹幹上でチャボスズゴケと共に生育していた．　2) Graphis fulvescens M. Nakan., Kashiw. & K. H. Moon, sp. nov.（ウゴノシロミモジゴケ，新称）：本種の主な特徴は次の通りである．地衣体は樹皮生，淡褐色，リレラ (lirella)は基物から突出し，不規則に分枝する．ラビア (labia)は中部まで地衣体で被われ，線状痕 (striation)はない．盤は閉じる．果殻は側部の上部50–75%が炭化するが，底部が炭化することはない．子嚢層は顆粒状物質を欠く．子嚢中に 4(–6)個の胞子を生じる．胞子は無色，腎臓形，石垣状多室，8–10本の異隔壁 (distosepta)と 3–5個の小腔室 (locule)を持つ，大

きさは 28–30 × 10–12 µm．地衣成分は検出されない．　本種はコウシュウシロモジゴケ Graphis kousyuensis に似ているが，後者は盤が開き，子嚢は 2個の胞子を生じ，胞子は 50–60 × 18–20 µmと大きいので区別できる．また，中国産の G. alpestris (Zahlbr.) Staigerも果殻の形状が本種に似ているが，子嚢には 1個の胞子を生じ，胞子は 100–120 × 25–28 µmと大きく，異隔壁も 25–30本と多いので区別できる．　調査地内では海抜 550 m付近のヤマモミジの樹幹にタカネカモジゴケと共に生育していた．また，滋賀県金糞山の 1100 m付近のミズナラ樹幹上で採集された 1標本も本種と同定された．　3) Julella sericea (A. Massal.) Coppins（ミチノクコアミゴケ，新称）：本種は Thelenellaceaeの固着地衣類で散生する被子器を持つ．地衣体は樹皮生，灰白色でほとんど発達しない．被子器は黒色，孔口は頂生．子嚢に8個の胞子を生じる．胞子は無色，石垣状多室，5–7本の異隔壁と 1–2個の小腔室を持つ，13–18 × 8–10 µm．地衣成分は検出されない．　本種は北半球に広く分布するが，日本からは北海道から一度だけ報告がある (Kashiwadani et al. 2009)．今回の調査では，530–600 m付近のヒトツバカエデ樹幹上に生育するのを確認した．　4) Porina dolichophora (Nyl.) Müll. Arg.（オガチホルトノキゴケ，新称）：本種の主な特徴は次の通りである．地衣体は樹皮生，淡褐色．被子器は散生し地衣体上に突出し，上部近くまで地衣体で被われる．半球状で基部は収縮しない．孔口は頂生．インボルクレールム

234 植物研究雑誌　第 90巻　第 4号 2015年 8月

(involucrellum) は孔口の周辺のみに生じ，褐色．果殻は淡褐色，側糸は単一で融合しない．子嚢に 8個の胞子を生じる．胞子は無色，長針形で少し曲がる，23–35本の横隔膜がある，95–112 × 3–5 µm．地衣成分は検出されない．　本種は南米のコロンビアで記載された種であるが，その後報告はない．しかし，インボルクレールムは孔口付近にのみ生じ，淡褐色の果殻，単一で融合しない側糸，針状で 16–21本の横隔膜を持つ胞子 (55–75 × 3–4 µm) などの特徴はタイプ標本とも形態的によく一致する．調査地では海抜 455–500 m付近のアカイタヤ樹幹上に生育していた．　5) Pyrenula nitidella (Flörke ex Schaer.) Müll. Arg.（サネゴケモドキ，新称）：地衣体 (UV−)は緑褐色で色素を含まない，平滑．被子器は独立して生じ，円錐形で径 0.5 mm以下，上部は黒色のインボルクレールムを裸出する．子嚢果内菌糸系は顆粒を含まない．胞子は褐色，長さ 21–25 × 5–8 µm，4室，両端の小腔室は外壁から離れている．子嚢果の底部に赤色色素（K+紫色）を持つ． 　本種は P. pseudobufonia (Rahm) R. C. Harrisに似ているが，後者は地衣体が UV+橙色，子嚢果内菌糸系に顆粒を含み，子嚢果には赤色色素を欠き，胞子両端の小腔室は外壁に密着する等の特徴で区別できる．

　本種の国内での分布は明らかではないが，これまでに北海道と福島県からの報告がある (Kashiwadani et al. 2009)．調査地内では最も普通に出現するサネゴケ属地衣の一つである．　6) Pyrenula tokyoensis (Müll. Arg.) H. Harada（ホシミゴケ）：本種は 1個の孔口を共有する子嚢果が放射状に数個集まる特有の形状をなし，線状で 11–15本の横隔膜を持つ胞子 (50–70 × 4.5–6 µm)を形成するので他のサネゴケ属の種とは容易に区別できる．　本種は日本特産種で，東京都，千葉県，静岡県といった本州中部から報告されている．今回の調査では，550 m付近でヤマハンノキの樹幹に生育することが確認された．また，北海道大雪山，滋賀県彦根市と甲賀市のハンノキやコナラ上で採集された標本も本種と同定できる．これまでの記録は比較的限られた地域であったが，北海道から滋賀県に至る，低地のハンノキ属やコナラに着生して広く分布している可能性がある．

（a国立科学博物館植物研究部，bオランダ・Soest市 ABLハーバリウム，

c広島大学教育学部，d韓国・国立生物資源館，e秋田大学教育文化学部，f㈱自然科学調査事務所）