国立科博専報，（44），2006年3月28日 Mem. Natn. Sci. Mus., Tokyo, (44), March 28, 2006

The Genus Ramalina (Ascomycotina: Ramalinaceae) in Taiwan

Hiroyuki Kashiwadani1, Kwang Hee Moon2 and Ming-Jou Lai3

1 Department of Botany, National Science Museum, Tokyo, 4–1–1 Amakubo, Tsukuba, Ibaraki, 305–0005 Japan E-mail: [email protected] 2 Laboratory of Mycology, Biological Sciences-Systematics and Ecology, College of Natural Sciences, Seoul National University, San 56–1, Sillim 9 dong, Gwanak-gu, Seoul, 151–742, Korea E-mail: [email protected] 3 Department of Landscape Architecture, Tunghai University, P.O. Box 834, Taitung, Taiwan E-mail: [email protected]

Abstract. The genus Ramalina in Taiwan is taxonomically revised. Among the 12 species re- ported, R. inclinata is a species newly described. Ramalina litoralis, R. pollinaria, R. subpollinaria, and R. shinanoana are new to Taiwan. Ramalina geniculata and R. subgeniculata are excluded from the lichen flora of Taiwan. Key words: Ramalina, Ramalina inclinata, lichens, Taiwan.

Introduction Materials and Methods The Ramalina flora of Taiwan is in general The present study is based primarily on about poorly known. The first study for the genus of 300 specimens of Ramalina collected in Taiwan Taiwan was made by Zahlbruckner (1933) who by the authors and housed in the herbarium of the reported following two species and four varieties, National Science Museum, Tokyo (TNS). Vari- R. calicaris (L.) Röhl., R. calicaris var. japonica ous type specimens preserved in other herbaria Hue, R. farinacea var. multifida Ach., R. fari- were also examined. In addition, about 60 speci- nacea var. pendulina Ach. R. geniculata var. oli- mens loaned to H. Kashiwadani from the Nation- vacea Müll.Arg., and R. pumila Mont. Asahina al Museum of Natural Science, Taichung (TNM), (1938, 1939) added four species, R. asahinana were also studied. The collections in Taiwan by Zahlbr., R. geniculata Hook. & Tayl., R. interme- the authors were primarily carried out during diella Vain., and R. subgeniculata Nyl. Kashi- their field studies for the Natural History Re- wadani (1986, 1987, 1988) recorded additional searches of the Island Arcs in the Western Pacific three species, R. hossei Vain., R. nervulosa organized by the National Science Museum, (Müll.Arg.) des Abb., and R. conduplicans Vain. Tokyo. The secondary products of the specimens At present, therefore, nine species and four vari- examined were determined by thin layer chro- eties are known from the present area. matography (TLC) using the amended proce- The purpose of the present study is to provide dures of Culberson and Johnson (1982). To eval- a revision of the genus in Taiwan, with special uate anatomical variation within the thallus and emphasis on the morphological and chemical apothecia, sections were cut by hand with a razor variations in the scope of comparison with the blade. These were mounted in GAW solution, Japanese specimens. A key to all the species of and the anatomy and spores were examined with Ramalina known at present in Taiwan is also pro- standard light microscopic procedures. vided. 162 H. Kashiwadani, K. H. Moon and M.-J. Lai

Korea, and Himalayas (Kashiwadani, 1986). Two Taxonomy chemical races were found in the specimens from Ramalina conduplicans Vain., Ann. Soc. Zool. Taiwan; the chemical race 1, containing ho- Bot. Fenn., 1: 35. 1921. (Fig. 1a, b) mosekikaic acid aggregates and the race 3, con- Ramalina conduplicans is characterized by the taining salazinic acid as a major chemical sub- corticolous habit, shrubby thallus growing from a stance (Kashiwadani, 1986). The race 3 is more narrow holdfast, solid flattened or canaliculated common than the race 1 in this area. branches, punctiform and ellipsoid pseudo- This species was reported from Taiwan under cyphellae, cracked chondroid tissue of branches, Ramalina calicaris var. japonica Hue (Asahina, fusiform ascospores of 12–144–5 mm in size, 1939) or R. subcomplanata (Nyl.) Kashiw. and the presence of depsides and/or depsidones (Kashiwadani, 1986). In Taiwan, it usually grows such as sekikaic, homosekikaic, divaricatic, and on bark of trees (very rarely on rocks) in moun- salazinic acids. tainous regions at elevations between 800 and Ramalina conduplicans is very variable mor- 2950 m. phologically and chemically. The holotype of this Representative specimens examined. Race species is a fragmental specimen having dor- 1. Prov. Chiayi: Mt. Tsu-Tson-San, Mt. Ali, el- siventral main lobes with sparse secondary evation about 2300–2900 m, December 31, 1963, branches tapering towards the apices. Most of the S. Kurokawa 162 (TNS); Mt. Chien-San, Mt. specimens from Taiwan have narrow and canalic- Shin-Kao-San, elevation about 3100–3300 m, ulated main branches with tapering side branches January 1, 1964, S. Kurokawa 273 and 274 (Fig. 1a). Asahina (1939) reported such morpho- (TNS). Prov. Hsinchu: 3 km E of Talulindao, type under Ramalina calicaris (L.) Röhl. var. Kuanwu, on bark of Alnus formosanum, 1950 m japonica Hue. However, specimens with wider alt., October 18, 1994, C.K. Lin 4205 (L1079, and dorsiventral lobes and flattened distal TNM). Prov. Hualien: Mt. Nan-Fu-Ta-San, ele- branches (Fig. 1b) are also found from Taiwan. vation about 1500–2400 m, January 19, 1964, S. These morphological differences found in this Kurokawa 940 and 1048 (TNS); Mt. Nan-Fu-Ta- species are consecutive and do not seem to have San, on twigs of Osmanthus bioritsuensis, eleva- a taxonomic value. tion about 2400–2600 m, January 20, 1964, S. This species is widely distributed in eastern Kurokawa 1051 (TNS); Guaniuan, Shioulin Asia, having been collected in China, Japan, Hshiang, on bark of Alnus japonica, elevation

Fig. 1. Morphological variation of Ramalina conduplicans Vain. a, Narrow and canaliculated main branches with tapering side branches (H. Kashiwadani no. 45535, TNS); b, dorsiventral lobes with flattened distal branches (H. Kashiwadani 43901, TNS). Scale bars2 mm. Genus Ramalina in Taiwan 163 about 2300 m, March 12, 2004, H. Kashiwadani uary 20, 1964, S. Kurokawa 1052 (TNS); Tayul- 46935 (TNS). Prov. Nantou: en route from ing, Hsiulin-gun, elevation about 2700 m, August Tungpushe to Lolo, on bark of Quercus sp., ele- 2, 1985, H. Shibuichi 8069 (TNS); Yangtoushan, vation 1605 m, March 16, 1963, S. Nakanishi Hsiulin, on rotten log, 2025–3035 m alt., Nov. 12655 (TNS); Keitau, October 30, 1933, H. Ma- 19, 2000, C.K. Lin 7019 (L 2543, TNM). Prov. suda s.n. (TNS); Tayuling, Jenai, on rock, eleva- Taichung: Anmashan, Tungshih, on tree, alt. tion about 2600 m, August 2, 1985, K. Yoshida 2665 m, June 1, 2001, C.K. Lin 7257 (L2742, 7056 (TNS); the same locality, on bark of Prunus TNM); en route from Suyen to Mt. Nanhutashan, yedoensis, elevation about 2540 m, March 5, Hopin, on bark of Salix sp., elevation about 1991, H. Kashiwadani 43967 (TNS); along route 2000 m, Nov. 9, 1989, K. Yoshida 9884 and 9891 820, Hohuanshan Mts., Jen-Ai Hsiang, on bark, (TNS); Keitau, 1935, Y. Asahina s.n. (TNS); en elevation about 2590 m, March 9, 2003, H. route from Ssuyuan to Tochiatun Shan, Mt. Nan- Kashiwadani 45535 (TNS); ca 10 km SE from huta Shan, Hoping, elevation 1900–2250 m, No- Lishan, along route 8, Hohuanshan Mts., Jen-Ai vember 13, 1989, H. Kashiwadani 35762 and Hsiang, on rock along road, elevation about 35803 (TNS); Hoping, on Salix sp., elevation 2100 m, March 10, 2003, H. Kashiwadani 45692 about 2250 m, November 14, 1989, H. Kashi- (TNS); ca 3 km W of Kueniynag, Jen-Ai Hsiang, wadani 35737 (TNS). Prov. Nantou: en route on bark of Salix sp., elevation about 2780 m, from Lolo to Tuikuan, on bark, elevation 1870 m, March 8, 2003, H. Kashiwadani 45561 and March 7, 1963, S. Nakanishi 12749 and 12938 45556 (TNS). Prov. Taichung: en route from (TNS); Lisan, January 17, 1964, S. Kurokawa Suyen to Mt. Nanhutashan, Hopin, on bark of 882 (TNS); Tayuling, Jenai, on rock, elevation Salix sp., elevation about 2000 m, Nov. 9, 1989, about 2600 m, August 2, 1985, K. Yoshida 7155 K. Yoshida 9888 (TNS); en route from Ssu-yuan (TNS); along route 820, Hohuanshan Mts., Jen- to To-chia-tun Shan, Mt. Nanhuta Shan, Hoping, Ai Hsiang, on bark, elevation about 2590 m, on bark of Salix sp., elevation 1900–2250 m, No- March 9, 2003, H. Kashiwadani 45536 (TNS); ca vember 9, 1989, H. Kashiwadani 35766 and 3 km W of Kueniynag, Jen-Ai Hsiang, on bark of 35763 (TNS); en route from Tochiatun Shan to Salix sp., elevation about 2780 m, March 8, 2003, Sheen-ma-jenn Shan, Mt. Nanhuta Shan, Hop- H. Kashiwadani 45554 and 45563 (TNS). Prov. ing, elevation 2500–2900 m, November 11, 1989, Taitung: Mt. Lachialachiaerh, elevation about H. Kashiwadani 35908 and 35948 (TNS); 1800 m, January 22, 1965, S. Kurokawa 2496 Guguan Hot Spring, Hopen Hsiang, on rock, ele- (TNS); Mt. Wunitoparu, elevation about 1900 m, vation 800–900 m, March 6, 2003, H. Kashi- January 23, 1965, S. Kurokawa 2596 (TNS); wadani 45604 (TNS); Tashueshan Forest Park, Raisha, January 5, 1926, Y. Asahina F 288 and F above Hsiaoshuehshan Hostel (upper parking 290 (TNS); ca 2 km E of Kwansan Yako, Haitu- lot), on road to summit of Mt. Tashueshan, an-gun, on bark of Alnus sp., elevation about Hopen, on branch of Acer sp., elevation 2800 m, 2350 m, March 1, 1991, H. Kashiwadani 43940 November 29, 2002, H. Kashiwadani 45318 (TNS); en route from Tentzu to Kwansan Yako, (TNS); the same locality, on branch of Berberis Haituan-gun, on bark of Pinus sp., elevation sp., elevation 2600–2800 m, November 29, 2002, about 2580 m, February 28, 1991, H. Kashi- H. Kashiwadani 45308 and 45064 (TNS). wadani 43901 and 43909 (TNS). Race 3. Prov. Chiayi: Mt. Alisan. July 14, 1935, M. Ogata 928 (TNS); Mt. Tsu-Tson-San, Ramalina dilacerata (Hoffm.) Hoffm., Herbari- Mt. Ali, elevation about 2200–2600 m, January um vivum, sive collection plantarum siccarum, 4, 1964, S. Kurokawa 497 (TNS). Prov. Hualien: caesareae universitatis Mosquensis: 451. 1825. Mt. Nan-Fu-Ta-San, on twigs of Osmanthus Ramalina geniculata var. globiformis Räsänen, bioritsuensis, elevation about 2400–2600 m, Jan- J. Jpn. Bot., 16:88. (1940). Type collection: 164 H. Kashiwadani, K. H. Moon and M.-J. Lai

Japan, Honshu, Prov. Rikuzen, Matsushima, race 1. April 20, 1919, E. Ochiai s.n (herb. A. Yasuda This species is widely distributed in the North- 623)–holotype in H!, isotype in TNS! (tlc: usnic ern Hemisphere, having been reported from acid, sekikaic acid, and 4-O-demethylsekikaic Canada, Nordic countries in Europe, Japan, and acid). Syn. nov. Korea. The distribution now includes Taiwan, Ramalina dilacerata is characterized by the where it grows on bark in mountainous regions at corticolous habit, shrubby and fistulose thallus a elevation from 1600 to 2900 m. growing from a narrow holdfast, inflated and Representative specimens examined. Race more or less pellucid thalli with sparse side 1. Prov. Tainan: Toroyen, Mt. Arisan, July 14, branches tapering towards the apices, sparse and 1935, M. Ogata s.n. (TNS). small fenestrations, discontinuous medullary hy- Race 2. Prov. Ilan: Piyanan Pass, elevation phae with isolated bundles of hyphae (Kashi- 1600–1900 m, January 16, 1964, S. Kurokawa wadani & Moon 2002), shortly fusiform as- 844 (TNS). Prov. Taichung: en route from Ssu- cospores of 12–14(16)4–5 mm in size, and the yuan to To-chia-tun Shan, Mt. Nanhuta Shan, presence of sekikaic acid without accompanying Hoping, on bark of Alnus japonica, elevation homosekikaic or divaricatic acids as major chem- 1900–2250 m, November 9, 1989, H. Kashi- ical substances. wadani 35822 (TNS); en route from Ssu-yuan to Historical review for this species has been well To-chia-tun Shan, Mt. Nanhuta Shan, Hoping, on arranged by Krog and James (1977). The type bark of Pinus sp., elevation about 2500 m, No- specimen of this species from Germany (MW) vember 10, 1989, H. Kashiwadani 35987 (TNS); was not studied by the present authors. However, en route from To-chia-tun Shan to Sheen-ma- specimens from Taiwan have characteristic fea- jenn Shan, Mt. Nanhuta Shan, Hoping, elevation tures as written above, which coincide well with 2500–2900 m, November 11, 1989, H. Kashi- those of the lectotype of R. calicaris f. minuscula wadani 35946 (TNS); Tashueshan Forest Park, Nyl. (R. dilacerata; Type: Fellman, Lich. Arct. above Hsiaoshuehshan Hostel, on road to summit 57, BM!) or exsiccata specimens by Räsänen of Mt. Tashueshan, Hopen, on branch of Acer (Lich. Fenn. Exs. 34, TNS) and Santesson (Lich. sp., elevation 2800 m, November 29, 2002, H. Sel. Scand. Exs. 189, TNS). Kashiwadani 45317 and 45319 (TNS); the same Räsänen (1940), when he studied Japanese locality, on branch of Berberis sp., elevation lichens mainly collected by A. Yasuda, described 2600–2800 m, November 29, 2002, H. Kashi- one variety, Ramalina geniculata var. globiformis. wadani 45066 (TNS). Prov. Nantou: along route The type specimen of this taxon preserved in H 820, Hohuanshan Mts., Jen-Ai Hsiang, on bark, and TNS are well fertile and have fistulose elevation about 2590 m, March 9, 2003, H. thalli without soredia, terminal apothecia, dis- Kashiwadani 45533 and 45537 (TNS); ca 3 km continuous medullary hyphae, ascospores of 12– W of Kueniynag, Jen-Ai Hsiang, on bark of Salix 154–5 mm, and produces sekikaic and 4-O- sp., elevation about 2780 m, March 8, 2003, H. demethylsekikaic acids as major chemical sub- Kashiwadani 45559 and 45560 (TNS). stances. Therefore, the name is now reduced to a synonym of the present species, belonging to the Ramalina hossei Vain., Ann. Soc. Zool.-Bot. chemical race 1. Fenn. Vanamo, 1: 36. 1921. As reported by Kashiwadani and Inoue (1993) Ramalina hossei is characterized by solid and for Japanese specimens, two chemical races, the narrow (up to 1.5 mm wide) branches growing sekikaic acid and norstictic acid () containing from a common holdfast, punctiform pseudo- race (Race 1) and the divaricatic acid containing cyphellae, finely divided terminal branches form- race (Race 2), are found also in Taiwan, where ing nodular branchlets, absence of soredia, the race 2 is apparently more common than the cracked chondroid tissue, shortly fusiform as- Genus Ramalina in Taiwan 165 cospores of 14–164–5 mm in size, and the pres- ence of divaricatic acid as major chemical sub- stances (for specimens in Taiwan). Ramalina hossei might be confused with R. conduplicans, because they both have similar solid branches, punctiform pseudocyphellae, and produce homosekikaic or divaricatic acids as chemical substances. However, it can be distin- guished from the latter by the common holdfast and the presence of terminal branches with nodu- lar and isidioid branchlets. For distinctions be- tween R. hossei and R. shinanoana, see under the latter species. Fig. 2. Holotype of Ramalina inclinata Kashiw., When Kashiwadani (1988) first reported this K. H. Moon & M. J. Lai [H. Kashiwadani species from Taiwan, he recognized homosekika- (46998) & K. H. Moon, TNS]. Scale bar2 mm. ic acid containing race from the present area. Re- examination of specimens, however, reveals that the chemical data cited by him came from frag- 46938 (TNS). Prov. Taitung: en route from ments of R. conduplicans mixed in the herbarium Tentzu to Kansan, Haituan-gun, on bark of Salix packets. All the specimens of this species in Tai- sp., elevation about 2450 m, February 28, 1991, wan so far studied belong to the divaricatic acid H. Kashiwadani 43880 (TNS). containing race. This species has been reported from Bhutan, Ramalina inclinata Kashiw., K.H.Moon & M.- Thailand, and Taiwan. In Taiwan, it is very com- J.Lai, sp. nov. (Figs 2, 3a) mon on tree trunks (Alnus, Salix, Tsuga, etc.) or Thallus ut in Ramalina litoraris sed ramis sub- rarely on rocks along trail in mountainous re- cylindricis, sorediis et isidiis destitutis et acidiis gions at elevations between 1500 and 2900 m. sekikaicis et norsticticis continentibus differt. Representative specimens examined. Prov. Type collection. Taiwan, Prov. Pintung (Pin- Taichung: en route from Tochiatun Shan to tung Co.), Mt. Da Jian Shu Shan, Henchung Sheenmajenn Shan, Mt. Nanhuta Shan, Hoping, (21°57N, 120°48E), on rocks, elevation about elevation 2500–2900 m, November 11, 1989, H. 300 m, March 8, 2004, H. Kashiwadani (no. Kashiwadani 35912 (TNS); the same locality, on 46998–holotype in TNS) and K.H. Moon. rock, 2500 m, November 10, 1989, H. Kashi- Chemistry (TLC). Usnic and sekikaic, 4- wadani 35986 (TNS). Prov. Nantou: Along route O-demethylsekikaic and norstictic acids. 820, Hohuanshan Mts., Jen-Ai Hsiang, on rock, Description. Thallus saxicolous, up to 3 cm elevation about 2590 m, March 9, 2003, H. long, decumbent, sparingly and irregularly Kashiwadani 45528 and 45534 (TNS); ca 10 km branched, growing from a narrow holdfast. Lobes SE from Lishan, along route 8, Hohuanshan greenish yellow, solid, up to 1.2 mm wide, cylin- Mts., Jen-Ai Hsiang, on rock along road, eleva- drical, slightly flattened near base, main branches tion about 2100 m, March 10, 2003, H. Kashi- with tapering side branches. Soralia absent. wadani 45687 and 45698 (TNS). Prov. Chiayi: Pseudocyphellae rare, narrowly ellipsoid, flat. Mt. Tsutson-san, Mt. Ali, elevation 2300–2900 Cortex of lobes indistinct, 10–12 mm thick; chon- m, December 31, 1963, S. Kurokawa 161 and droid tissue smooth, continuous; medulla loose, 170 (TNS). Prov. Hualien: Guaniuan, Shioulin with embedded strands of chondroid tissue. Hshian, on bark of Alnus japonica, elevation Apothecia rare, lateral; disc more or less convex, about 2300 m, March 12, 2004, H. Kashiwadani without white margin; thalloid exciple smooth, 166 H. Kashiwadani, K. H. Moon and M.-J. Lai

Fig. 3. Cross section of branches. a, Medulla with separated chondroid tissue without cracks of R. inclinata; b, medulla with separated chondroid tissue with cracks of R. litoralis. Scale bars100 mm. lacking pseudocyphellae; ascospores short- Ramalina litoralis Asahina, J. Jpn. Bot., 15: 220. fusiform, 12–153.5–4.5 mm. Pycnidia not seen. 1939. (Fig. 3b) Ramalina inclinata resembles R. litoralis, a Ramalina litoralis is distinguished from allied saxicolous species growing on seaside rocks in species by the saxicolous habit, caespitose thal- eastern Asia, as both species have similar lus, more or less dorsiventral and solid branches branches with ellipsoid pseudocyphellae. Howev- with irregularly thickened branchlets ending in er, it can be distinguished from the latter by the nodules, absence of soredia, sparse shortly linear absence of soralia and the presence of norstictic pseudocyphellae, fusiform ascospores, 10–13 acid together with sekikaic acid as major chemi- 3–4(5) mm, and the presence of sekikaic (never cal substances. In addition, the chondroid tissue accompanying homosekikaic acid) or divaricatic of this species is smooth and the medulla has acids as major chemical substances. separated chondroid tissue (Fig. 3a); whereas the When Kashiwadani (1987) revised this species chondroid tissue of the latter is distinctly cracked for Japanese specimens, he reported two chemi- and does not spread into the medullary hyphae cal races in this species. In Taiwan only the di- (Fig. 3b). varicatic acid containing race is found. A similar anatomy for branches was reported This species might be confused with dwarf by Østhagen and Krog (1976) from Ramalina form of R. peruviana. However, it can be easily pluviariae Krog & Østh. and R. scopulorum var. distinguished from the latter by the absence of nematodes Nyl., both the Canary Islands’ taxa, soredia. In addition, R. litoralis lacks ho- which differ in having striate pseudocyphellae mosekikaic acid which is constantly produced in and distinct cortex and in producing protocetraric the latter species. or salazinic acids. This species has been known from Japan and At present this species is known only from Korea (Kashiwadani, 1987; Kashiwadani et al., several places nearby the type locality near the 2002). The distribution range now includes Tai- top of steep congromelatic rocks together with R. wan where only two collections are made on sea- pollinaria. side rocks. Specimens examined. Prov. Pintung: Mt. Specimens examined. Prov. Taipei: Shrmen, Da Jian Shu Shan, Henchung, on rocks, elevation Shrmen-gun, on rock along coast, elevation 200–300 m, March 8, 2004, H. Kashiwadani about 5 m, March 11, 2005, H. Kashiwadani (46813, 16816, 46824, TNS) and K.H. Moon; the 47387 (TNS). Prov. Taitung: Landao, Lanyu Is- same locality, K. Yoshida 9557 and 9558 (TNS). land, on rock along the coast, October 26, 1989, H. Kashiwadani 35731 (TNS); the same locality, on rock, elevation 20 m, October 26, 1989, K. Genus Ramalina in Taiwan 167

Yoshida 9463 (TNS); Hongtou-cun, Lanyu, on Chiayi: Kagi, December 30, 1933, Y. Asahina rocks along the coast, October 24, 1989, H. s.n.* (TNS); the same locality and date, Y. Kashiwadani 35688 (TNS); Lanyi Island, June, Asahina F291 (pr. p., TNS). Prov. Kaohsiung: 1936, S. Sasaki s.n. (TNS). Kural, elevation 250 m, S. Kurokawa (TNS); the same locality, December 30, 1933, Y. Asahina Ramalina nervulosa (Müll.Arg.) des Abb., s.n. (TNS). Prov. Pintung: Mt. Dajianshu-shan, Bull. Inst. Fr. Afr. Noire, 14: 25. 1952. Henchung, on branches of Berberis sp., elevation This species is characterized by the corticolous 300 m, March 8, 2004, H. Kashiwadani 46824 habit, shrubby or subpendulous branches from a (TNS); the same locality, May 12, 1985, M.J. Lai narrow holdfast, dorsiventral and sparingly 17031 (TNS). branched thallus often with fenestrations, striate pseudocyphellae, marginal farinose soralia, Ramalina pertusa Kashiw., Mem. Natn. Sci. cracked chondroid tissue, broadly fusiform as- Mus., 18: 102. 1985. cospores of 12–164–5 mm, and the presence of Ramalina pertusa is easily distinguished from homosekikaic or divaricatic acids as major chem- other corticolous species of Taiwanese Ramalina ical substances. by shrubby and fistulose thallus with orbicular to When Kashiwadani (1986) revised this species elongate openings on the lower surface, more or from Japan and Taiwan, he reported two chemi- less continuous medullary hyphae tightly at- cal races, the homosekikaic and sekikaic acids tached onto chondroid layer, broadly ellipsoid as- containing race and the divaricatic acid contain- cospores of 12–154–5(6) mm in size, and the ing race. In Taiwan, the former is apparently production of evernic and obtusatic acids as more common than the latter (specimen with an major chemical substances. asterisk in the following list). The specimen Ramalina pertusa is a polymorphic species. (Asahina F291, pr. p., TNS) reported under Ra- Even in a single specimen, fertile branches usual- malina farinacea var. multifida Ach. by ly have sparse side branches whereas most of the Zahlbruckner (1933) is now identified with the sterile branches frequently form cylindrical and homosekikaic acid containing race of R. nervu- tapering side branches. This species resembles R. losa. dilacerata, which differs in having discontinuous Ramalina nervulosa resembles R. peruviana in clusters of medullary hyphae together with em- having similar branches with soredia. However, bedded strands of chondroid tissue (Kashiwadani this species can be distinguished from the latter & Moon, 2002). by having wider branches (1.5–3.0 mm wide) Ramalina pertusa was described from Japan with farinose soredia and occasional fenestra- and has not been recorded from other countries. tions and by shortly fusiform ascospores. Rama- The present study, however, reveals that this lina peruviana has narrower branches (less than species is also common in Taiwan, where it 1.2 mm wide) with granular soredia, isidioid grows on bark of trees in mountainous regions branchlets, and narrowly fusiform ascospores. In from 800 to 2900 m high. Asahina (1938) first re- addition, the latter species never forms fenestra- ported this species from Taiwan under R. sub- tion on branches. geniculata Nyl. This species is widely distributed in tropical Representative specimens examined. Prov. and subtropical regions in the world including Ilan: Taipingshan, along the road to Santieh wa- Japan and Taiwan (Kashiwadani, 1986; Stevens, terfall, on tree, elevation 1800 m, August 2, 1997, 1987). In Taiwan, however, it is rather rare, being C.K. Lin 6089 (L 1942, TNM). Prov. Hualien: collected on twigs of shrubs in the lowland south Guaniuan, Shioulin Hshiang, on bark of Alnus of Chiayi Province at a elevation below 300 m. japonica, elevation about 2300 m, March 12, Representative specimens examined. Prov. 2004, H. Kashiwadani 46988 and 46939 (TNS); 168 H. Kashiwadani, K. H. Moon and M.-J. Lai

Tayuling, Hsiulin-gun, on rocks, elevation about branches, narrowly elongated pseudocyphellae, 2700 m, August 2, 1985, H. Shibuichi 8069b marginal or laminal soralia with granular soredia (TNS); Mt. Nan-Fu-Ta-San, elevation about and tiny branchlets, smooth chondroid tissue, 2400–2600 m, January 20, 1964, S. Kurokawa narrowly fusiform ascospores of 15–203–4 mm 1045 and 1046 (TNS); the same locality, on bark, in size, and the presence of homosekikaic and elevation 1500–2400 m, January 19, 1964, S. sekikaic acids together with ramalinoric acid. Kurokawa 941 (TNS); Yangtoushan, Hsiulin, on This species might be confused with R. nervu- rotten log, 2025–3035 m, Nov. 19, 2000, C.K. losa in Taiwan, because these species have simi- Lin 7040 (L2564, YNM). Prov. Taichung: en lar branches with soralia. However, it can be dis- route from Suyen to Mt. Nanhutashan, Hopin, on tinguished from the latter by the narrow and ir- bark of Salix sp., elevation about 2200 m, Nov. 9, regularly thickened branches, soralia with granu- 1989, K. Yoshida 9885 (TNS); the same locality, lar soredia and branchlets, and narrowly fusiform on bark of Alnus japonica, elevation 1900–2250 ascospores; the lobes are dorsiventral even in the m, November 9, 1989, H. Kashiwadani 35870 distal branches and soralia does not produce (TNS); the same locality, on bark of Pinus sp., branchlets in R. nervulosa. In addition, speci- elevation about 2500 m, November 10, 1989, H. mens kept in the herbarium for long time turn Kashiwadani 35989 (TNS); en route from To- into reddish brown in R. peruviana, whereas they chia-tun Shan to Sheen-ma-jenn Shan, Mt. Nan- turn into ocher yellow in the latter species. It also huta Shan, Hoping, on bark of Salix sp., elevation resembles R. dissecta which differs in having 2500–2900 m, November 11, 1989, H. Kashi- punctiform pseudocyphellae and shortly fusiform wadani 35911 (TNS); Mt. Wunitoparu, elevation ascospores. about 1900 m, January 23, 1965, S. Kurokawa This species was reported under R. farinacea 2594 (TNS); Tashueshan Forest Park, above var. pendulina Ach. by Zahlbruckner (1933) and Hsiaoshuehshan Hostel (upper parking lot), on R. intermediella Vain. by Asahina (1939). A road to summit of Mt. Tashueshan, Hopen, on specimen (Asahina F291, pr. p., TNS) reported branch of Prunus sp, elevation 2600–2800 m, under Ramalina farinacea var. multifida Ach. by November 29, 2002, H. Kashiwadani 45316 Zahlbruckner (1933) is also identified with R. pe- (TNS); the same locality, on bark of Pinus mor- ruviana. risonicola, elevation 2550–2580 m, November Ramalina peruviana is occasionally collected 30, 2002, H. Kashiwadani 45324 (TNS). Prov. elevations below ca 1000 m in Taiwan, but it is Nantou: Along route 820, Hohuanshan Mts., locally abundant especially on the trunks of Jen-Ai Hsiang, on rock, elevation about 2590 m, planted Cocos nucifera in southern Taiwan March 9, 2003, H. Kashiwadani 45529 (TNS). (Hualien and Taitung Provinces). Prov. Kaohsiung: Mt. Nanfong, elevation 800– Specimens examined. Prov. Taipei: Okase- 1300 m, February 7, 1965, S. Kurokawa 2839 ki, June 1903, U. Faurie 295 (TNS). Prov. (TNS). Prov. Taitung: en route from Tentzu to Hualien: Taroko Gorge, Shioulin Hsiang, on Kwansan Yako, Haituan-gun, on bark, elevation bark of Zelkova serrata, elevation about 720 m, about 2580 m, March 1, 1991, H. Kashiwadani March 11, 2004, H. Kashiwadani 46972 (TNS); 43921 (TNS). Tienhsiang, Shulin-gun, on bark of Prunus persi- ca, elevation about 950 m, March 3, 1991, H. Ramalina peruviana Ach., Lich. Univ.: 1599. Kashiwadani (43961 and 44005, TNS) and K.H. 1810. Moon. Prov. Taichung: Keitau, 1935, T. Masuda Ramalina peruviana is characterized by tufted s.n. (TNS). Prov. Chiayi: Kagi, December 30, thallus growing from a common holdfast, narrow 1933, Y. Asahina F 291 and Asahina s.n. (TNS). (less than 1.5 mm wide) branches becoming Prov. Kaohsiung: Kural, December 30, 1933, Y. terete or irregularly thickened at the distal Asahina s.n. (TNS). Prov. Taitung: ca 5 km N of Genus Ramalina in Taiwan 169

Tai Yuan, Dongho Hshiang, on bark of Cocos nu- in montane forests higher than 1500 m in Taiwan. cifera, elevation about 270 m, March 11, 2004, Ramalina pumila resembles R. pertusa which Kashiwadani (46997, TNS) and K.H. Moon. differs in having larger perforations (up to 3– 62–4 mm in size) and in producing evernic and Ramalina pollinaria (Westr. ex Ach.) Ach., obtusatic acids. Lich. Univ.: 608. 1810. Specimens of this species from Taiwan had Ramalina pollinaria is easily distinguished been misidentified with R. geniculata var. oli- from other species of Ramalina in Taiwan by the vacea Müll.Arg. (Zahlbruckner, 1933) or R. saxicolous habit, shrubby and solid thallus with geniculata Hook.f. & Tayl. (Asahina, 1938). terminal or laminal soralia with granular soredia, Specimens examined. Race 1. Prov. Hualien: and the production of evernic and obtusatic acids Taroko Gorge, Shioulin Hshiang, on branches of as chemical substances. Ficus sp., elevation about 150 m, March 11, This is one of the well known species of Ra- 2004, H. Kashiwadani 46995 and 46996 (TNS); malina widely distributed in the world. Although Tienhsiang, Shulin-gun, on branches of Malus this species has been reported from China, Japan, torgingo, elevation about 950 m, March 3, 1991, and Korea in eastern Asia (Kashiwadani & H. Kashiwadani 4402 (TNS). Prov. Kaohsiung: Moon, 2003), it has never been known from Tai- Pongari-sha, January 7, 1926, Y. Asahina F289 wan. In fact, only one collection was made dur- (TNS); Shanping, elevation about 750 m, Febru- ing our field study, but it must has been over- ary 6, 1965, S. Kurokawa 2735 (TNS). looked in Taiwan, since it is very common like Race 2. Prov. Taipei: Tamusui (Tansui), weeds in other areas. June 1903, U. Faurie (KYO, TNS). Specimen examined. Prov. Hualien: Taroko Gorge, Shioulin Hshiang, on rock, elevation Ramalina shinanoana Kashiw., Bull. Natn. Sci. about 720 m, March 11, 2004, H. Kashiwadani Mus. Ser. B, 12: 122. 1986. (Fig. 4) 46976 (TNS). When Kashiwadani (1986) described this species, he reported two chemical races, the ho- Ramalina pumila Mont., Ann. Sci. Nat. Bot. Ser. mosekikaic acid race (Race 1) and the acid defi- 2, 20: 356. 1843. cient race (Race 2). However, all specimens list- Ramalina pumila is characterized by the corti- ed below have divaricatic acid and can be regard- colous habit, hollow thallus with a narrow hold- ed as a new chemical race for R. shinanoana. fast, partly flatted to more or less inflated branch- These three chemical races are indistinguishable es with sparse cylindrical branchlets on main regarding the morphology of thalli, apothecia, branches, sparse smaller perforations (up to 1–2 and ascospores. The Race 2 was not found 1 mm in size), continuous medullary hyphae among the collections from Taiwan. without forming bundles of hyphae, shortly Ramalina shinanoana resembles R. hossei, a fusiform ascospores of 12–14(1–15)(3.5)4– commonly found species in mountainous regions 5 mm in size, and the production of sekikaic acid of Taiwan. However, it can be distinguished from (Race 1) or divaricatic acid (Race 2) as a major the latter by the thallus growing from a narrow chemical substance. holdfast and the presence of soralia. Although Ramalina pumila might be confused with R. this species was reported from Japan and China dilacerata in having similar branches and in pro- (Kashiwadani, 1986), the distribution now in- ducing same chemical substances. However, it cludes Taiwan, where it grows on barks or twigs can be distinguished from the latter by the con- of trees at elevations between 2300 m and 2800 tinuous medullary hyphae tightly attached to the m. chondroid layer of thallus. In addition, R. pumila Specimens examined. Prov. Hualien: Mt. is a lowland species whereas R. dilacerata grows Nan-Fu-Ta-San, on bark, elevation about 2400– 170 H. Kashiwadani, K. H. Moon and M.-J. Lai

Fig. 4. Ramalina shinanoana Kashiw. (H. Kashiwadani 45531, TNS). Scale bar3 mm.

Fig. 5. Morphological variations of Ramalina sinensis Jatta. a, Palmate and loosely tufted lobes (H. Kashi- wadani 46933, TNS); b, elongate, marginally outgrown lobes (H. Kashiwadani 46940, TNS). Scale bars10 mm for a; 3 mm for b.

2600 m, January 20, 1964, S. Kurokawa 1047c ticolous habit, palmate and flattened lobes with and 1115 (TNS). Prov. Taichung: Tashueshan distinctly dorsiventral appearance, absence of Forest Park, above Hsiaoshuehshan Hostel (upper soredia, longitudinally orientated anastomosing parking lot), on road to summit of Mt. Tashue- ridges of chondroid tissue, depressed and reticu- shan, Hopen, on branch of Berberis sp., elevation lately arranged pseudocyphellae on the lower 2600–2800 m, November 29, 2002, H. Kashi- surface of lobes, straight, fusiform ascospores of wadani 45067, 45307, and 45309 (TNS). Prov. 13–15(17)5–6(7) mm in size, and the absence Nantou: along route 820, Hohuanshan Mts., Jen- of chemical substances in the medulla, although Ai Hsiang, on rock, elevation about 2590 m, small amount of salazinic acid is occasionally March 9, 2003, H. Kashiwadani 45531 (TNS). found. Ramalina sinensis is very variable in lobe Ramalina sinensis Jatta, Nouv. G. Bot. Ital., 9: shape and branching. Although specimens with 462. 1902. (Fig. 5a, b) palmate and loosely tufted lobes (Fig. 5a) are Ramalina sinensis is characterized by the cor- commonly found in Taiwan, those forming elon- Genus Ramalina in Taiwan 171

Fig. 6. Ramalina subpollinaria Nyl. (H. Kashiwadani 35761, TNS). Scale bar1 mm. gate, marginally outgrown lobes are also found 1993, C.K. Lin 3289 (L 0708, TNM); Tayuling, (Fig. 5b). In the latter case, it can be confused Jenai, on rock, elevation about 2600 m, August 2, with R. fraxinea, a species widely distributed in 1985, K. Yoshida 7158 (TNS). boreal regions, which differs in having branches without dorsiventral appearance, the ellipsoid Ramalina subpollinaria Nyl., Bull. Soc. Linn. pseudocyphellae, and the distinctly curved as- Normand. Ser. 2, 4: 125. 1870. (Fig. 6) cospores. Ramalina subpollinaria is recognized by its This cosmopolitan species was first reported small solid thallus growing from a narrow hold- from Taiwan by Asahina (1939) under the name fast, terminal soralia with farinose soredia, and Ramalina asahinana Zahlbr. It is rather com- the presence of homosekikaic and sekikaic acids, monly found on trees in mountainous regions in or salazinic acid. Taiwan. Kashiwadani (1987) reported two chemical Specimens examined. Prov. Chiayi: Mt. races of this species from Brazil, the plants from Anfuta-shan, on bark, elevation 1500–2400 m, the present area are the homosekikaic acid aggr. January 19, 1964, S. Kurokawa 939 (TNS); the race lacking salazinic acid. This is the first report same locality, on twigs of Osmanthus bioritsuen- for the species outside South America. sis, 2400–2600 m, January 20, 1964, S. Specimens examined. Prov. Taichung: en Kurokawa 1049 (TNS). Prov. Hualien: Guani- route from Su-yuan to Tochiatun Shan, Mt. Nan- uan, Shioulin Hshiang, on bark of Alnus huta Shan, Hoping, on rock, elevation about 2000 japonica, elevation about 2300 m, March 12, m, November 9, 1989, H. Kashiwadani 35865 2004, H. Kashiwadani 46933 and 46940 (TNS). (TNS); the same locality, on bark of Salix sp., el- Prov. Nantou: Tantalindao, Hsini, on branch of evation about 1900 m, November 9, 1989, H. Alnus formosana, alt. 2200 m, September 25, Kashiwadani 35761 (TNS). 172 H. Kashiwadani, K. H. Moon and M.-J. Lai

Key to the Ramalina species of Taiwan terete; pseudocyphellae indistinct, shortly 1. Thallus hollow...... 2 linear...... 11 – Thallus solid...... 4 11. Branches more or less cylindrical, tapering, 2. Perforations common, conspicuous, more never producing nodules; center of medulla than 1/2 of branches in width; evernic and (in cross section) with strands of chondroid obtusatic acids present ...... R. pertusa tissue...... R. inclinata – Perforations rare, small, less than 1/3 of – Branches irregular in thickness, often with branches in width; sekikaic acid or divaricat- isidioid nodules; center of medulla (in cross ic acid present...... 3 section) with interwoven hyphae...R. litoralis 3. Medullary hyphae dense, continuous, tightly attached to the cortex; coastal lowland Acknowledgement species...... R. pumila – Medullary hyphae sparse, discontinuous, The authors wish to express their sincere forming bundles of chondroid tissue in thanks to Dr S. Kurokawa in Toyama for his criti- medulla; montane species...... R. dilacerata cal reading of the manuscript. They are most 4. Soralia present...... 5 grateful to the curator of H, KYO, S, TNM, and – Soralia absent ...... 10 TUR for the loan of specimens of Ramalina. 5. Thallus less than 3 cm long, sparingly Thanks are extended to Dr Y. C. Tsai of National branched ...... 6 Yang-Ming Medical College, Taipei, Mr R.-R. – Thallus more than 3 cm long, repeatedly Liao, Taichung, and Mr D. Wan, Taipei, for their branched ...... 7 kind help during our lichenological field survey 6. Soredia granular; evernic and obtusatic acids in Taiwan. This study was supported in part by a present...... R. pollinaria Grant-in Aid for Scientific Research from the – Soredia farinose; homosekikaic and sekikaic Ministry of Education, Science, Sports and Cul- acids present...... R. subpollinaria ture of Japan to H. Kashiwadani (nos. 02640642 7. Main branches up to 1.5 mm wide, growing and 17570089). from a common holdfast ...... 8 – Main branches 1.5–3.0 mm wide, growing References from a narrow holdfast...... 9 8. Pseudocyphellae punctiform; distal branches Asahina, Y., 1938. Ramalina-Arten aus Japan (I). Journal finely divided, ending in nodular isidioid of Japanese Botany, 14: 721–730. branchlets...... R. hossei Asahina, Y., 1939. Ramalina-Arten aus Japan (II). Jour- nal of Japanese Botany, 15: 205–223. – Pseudocyphellae shortly linear; distal Culberson, C. & A. Johnson, 1982. Substitution of methyl branches tapering but never forming isidioid ter.-butyl ether for diethyl ether in the standardized branchlets...... R. peruviana thin-layer chromatographic method for lichen products. 9. Soredia farinose, marginal; pseudocyphellae Journal of Chromatography, 238: 483–487. shortly linear; maritime species Kashiwadani, H., 1986. Genus Ramalina (Lichens) in Japan (2). On Ramalina pacifica Asah. and its allies...... R. nervulosa Bulletin of the National Science Museum, Series B, 12: – Soredia granular, terminal or laminal; 117–125. pseudocyphellae ellipsoid; montane species . Kashiwadani, H., 1987. Genus Ramalina (Lichens) in ...... R. shinanoana Japan (3). R. exilis Asahina and its allies. Bulletin of 10. Thallus corticolous, branches more or less the National Science Museum, Series. B, 13: 133–140. canaliculated to flat; pseudocyphellae dis- Kashiwadani, H., 1988. Ramalina hossei Vain. found in Bhutan and Formosa. Bulletin of the National Science tinct, punctiform to ellipsoid Museum, Series. B, 14: 129–133...... R. conduplicans Kashiwadani, H. & M. Inoue, 1993. The lichens of Kushi- – Thallus saxicolous, branches cylindrical or ro marsh, Hokkaido, Japan. Memor of the National Sci- Genus Ramalina in Taiwan 173

ence Museum, Tokyo, (26): 53–66. Fennoscandia and the British Isles. Norwegian Journal Kashiwadani, H. & K. H. Moon, 2002. A new of interest- of Botany, 24: 15–43. ing species of the genus Ramalina (Ascomycotina: Ra- Østhagen, H. & H. Krog, 1976. Contribution to the lichen malinaceae) from Korea and Japan. Bulletin of the Na- flora of the Canary Islands. Norwegian Journal of tional Science Museum, Series B, 29: 91–95. Botany, 23: 221–242. Kashiwadani, H. & K. H. Moon, 2003. A note on two Räsänen, V., 1940. Lichenes ab A. Yasuda et aliis in species of Ramalina (Ascomycotina) in Eastern Asia. Japonia collecti (I). Journal of Japanese Botany, 16: Bulletin of the National Science Museum, Series B, 28: 81–98. 1–6. Stevens, G. N., 1987. The lichen genus Ramalina in Aus- Kashiwadani, H., K. H. Moon, M. Inoue, G. Thor & Y. S. tralia. Bulletin of the British Museum (Natural History), Kim, 2002. Lichens of the Cheju Island, Republic of Botany Series, 16: 107–223. Korea. I. The Macrolichens. National Science Museum Zahlbruckner, A., 1933. Flechten der Insel Formosa II. Monograph, (22): 115–135. Feddes Repertorium Specierum Novarum Regni Veg- Krog, H. & P. W. James, 1977. The genus Ramalina in etabilis, 33: 22–68.

台湾のカラタチゴケ属（子嚢菌亜綱，カラタチゴケ科）

柏谷博之・文 光喜・頼 名洲

台湾産カラタチゴケ属の分類学的研究を行った．同地域からは12種が報告され，そのうち Rama- lina inclinata は新種であり，R. litoralis, R. pollinaria, R. subpollinaria および R. shinanoana は台湾新産 である．Ramalina geniculata と R. subgeniculata は台湾産地衣相からは除外された．