Systematic Studies of Asian Saussurea Asteraceae) VI. A ...268 植物研究雑誌第88 巻第5 号 2013 年10 月 Fig. 1. Type of Saussurea katoana Kadota (JAPAN, Honshu, Yamagata

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Systematic Studies of Asian Saussurea (Asteraceae) VI.A New Species and a New Combination from Hokkaido

and Three New Species from Honshu, Japan

Yuichi Kadota

Department of Botany, National Museum of Nature and Science,4–1–1, Amakubo, Tsukuba, 305-0005 JAPAN

E-mail: [email protected]

(Accepted on May 11, 2013)

A new species, Saussurea yubarimontana Kadota and a new combination, S. uryuensis (Kadota) Kadota from Hokkaido, and three new species, S. katoana Kadota, S. yanagitae Kadota and S. yuki-uenoana Kadota from Honshu, Japan are described. S. yubarimontana from Mt. Yubari-dake, central Hokkaido is different from S. riederi Herder var. yuparensis Kitam. [= S. riederi var. yezoensis (Maxim.) Kitam.] in the subscapose habit and mucronate, 8-seriate involucral phyllaries. Saussurea uryuensis (Kadota) Kadota is proposed as a new combination based on the comparison between S. yubarimontana and S. kudoana Kitam. & Tatew. A key is also proposed to the Saussurea kudoana group. S. katoana from Tobishima Island, Yamagata Pref. differs from S. hosoiana Kadota in the absence of basal leaves at anthesis, longer peduncles, 10-seriate involucral phyllaries and wider, dentate stem-wings. Saussurea yanagitae from Niigata Pref. is different from S. sagitta Franch. in the larger and pendulous habit, larger, thicker, slightly lustrous cauline leaves, more capitula arranged in a corymb, smaller, densely arachnoid involucres and shorter florets. S. yuki-uenoana from Abukuma Mountains, Miyagi Pref. is unique in gross morphology and is characterized by the plant height, 12-seriate involucral phyllaries, smaller but more capitula in a dense corymb, narrowly cylindrical involucres, ovate, coriaceous cauline leaves and strongly ribbed stem without wings.

Key words: Abukuma Mountains, Hokkaido, Japan, Mt. Arasawa-dake, Mt. Yubari-dake, new species, Saussurea katoana, Saussurea uryuensis, Saussurea yanagitae, Saussurea yubarimontana, Saussurea yuki-uenoana, Tobishima Island, Tohoku.

J. Jpn. Bot. 88: 267–285 (2013)

This is part of a systematic study of East Asian Saussurea (Asteraceae) (Kadota 1987, 2004, 2007, 2008, 2009, 2010, 2011a, 2011b, 2011c, 2012).

Tobishima Island is a small island (2.7 square kilometers in area) floating on the Japan Sea and lies at ca. 40 km NW from Sakata-shi, Yamagata Prefecture, Honshu, northern Japan. In 2009 some Saussurea plants were collected

and brought to the Herbarium, Department of Botany, National Museum of Nature and Science (TNS) by Mr. Shin-ei Kato, Tsuruoka, Yamagata Prefecture (Fig. 1). Saussurea muramatsui Kitam. is known to be distributed in the mainland of Yamagata Prefecture. However, the Tobishima plants are significantly different from S. muramatsui. In 2011 a mass collection of Saussurea from the island was

268 植物研究雑誌　第 88巻　第 5号 2013年 10月

Fig. 1. Type of Saussurea katoana Kadota (JAPAN, Honshu, Yamagata Pref., Sakata-shi, Tobishima Island, 29 Sept. 2009, Shin-ei Kato 1, TNS 741157, holotype).

October 2013 TheJournal of Japanese Botany Vol. 88 No. 5 269

additionally made by Mr. Kato. As a result of the examination it is clarified that the Saussurea plants from Tobishima Island represent an undescribed species. This new species is here named as Saussurea katoana.

In October 2010 unusual Saussurea plants (Fig. 9) were obtained in Marumori-machi, Igu-gun, Miyagi Prefecture, northern Honshu by Mr. Yuki Ueno, Shiroishi, Miyagi Prefecture. These plants are unique in gross morphology among the species of the genus. They are characterized by the plant height (up to 1.8 m high) and 12-seriate involucral phyllaries. In response to the finding I made field examinations in Marumori-machi in October 2011 under the guidance of Mr. Ueno. As a result it turned out that the Saussurea plants represent an undescribed species. It is described as a new species, Saussurea yuki-uenoana.

In 2011 a Saussurea specimen was given to the Herbarium TNS by Mr. Hiromitsu Yanagita, Ojiya, Niigata Prefecture. The specimen was a gathering from cliff in Mt. Arasawa-dake, Uonuma-shi, Niigata Prefecture, central Honshu, Japan. This Saussurea plant is similar to S. sagitta Franch. but the former is significantly different from the latter in several morphological characters. Upon my request Mr. Yanagita and his colleagues made repeated return trips to in the mountain and they could successfully collected good specimens and took pictures of the Saussurea plants. These plants are especially characteristic of the pendulous habit from the rocky cliff (Fig. 4). Consequently the Saussurea is described as a new species, Saussurea yanagitae.

It is known that Saussurea plants grow in serpentine areas of the foot of Mt. Yubari (Yûpari)-dake, Sorachi Subprefecture, central Hokkaido, northern Japan (Fig. 6). These plants have ʻdimorphicʼ cauline leaves (larger, lower cauline vs. smaller, upper cauline leaves) and 8-seriate involucral phyllaries. The plants in question have been considered to be S. riederi Herder var. yuparensis Kitam. (Nosaka 1959, 1961, Umezawa 2009, p. 33, 2010, cover photo,

Horie 2010). However, the type specimens of S. riederi var. yuparensis (Fig. 7) are characterized by having ʻmonomorphicʼcauline leaves (diminishing gradually in size from the lower to the upper parts) and 5-seriate involucral phyllaries. Agreeing with the statement of Toyokuni (1960), the name S. reideri var. yuparensis should be consequently reduced to be a synonym of S. riederi var. yezoensis (Maxim.) Kitam. Actually S. riederi var. yezoensis is commonly found in the alpine zone of Mt. Yubari-dake (including serpentine areas). Therefore the plants from the serpentine areas in the foot of Mt. Yubari-dake are ascribed to a new species, Saussurea yubarimontana.

Taxonomic treatment1) Saussurea katoana Kadota, sp. nov. [Figs. 1–2]

Saussurea katoana is distinguished from S. hosoiana Kadota in the absence of basal leaves at anthesis, the subscapose habit, 10-seriate involucral phyllaries, longer peduncles and dentate wings of the stem and from S. nipponica Miq. subsp. hokurokuensis Kitam. in thicker leaves, fewer capitula, larger involucres and the dentate stem wings.

Type: JAPAN. Honshu. Yamagata Pref., Sakata-shi, Tobishima Island, Arasaki – Kashiwagiyama, alt. ca. 50 m, under scattered Pinus thunbergii woods, 29 September 2009, Shin-ei Kato 1 (TNS 741157–holotype; Fig. 1).

A robust, subscapose, herbaceous perennial, 45–65 cm tall. Rhizome oblique to horizontal, 1–2 cm in diameter, with cord-like roots. Stem erect, striate, prominently winged, densely pubescent with brownish multicellular hairs in the upper half, glabrous in the lower half, 1–5 times branched in the upper part; wings 8–10 mm wide, coarsely dentate. Basal leaves withered at anthesis. Cauline leaves abruptly diminishing in size from the middle part of the stem. Lower cauline leaves coriaceous, dimly lustrous, broadly ovate, 20–30 cm long, 14–23 cm wide, dentate, sparingly pubescent with brownish

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Russian).

Fig. 2. Saussurea katoana Kadota from Tobishima Island, Sakata-shi, Yamagata Pref., northern Honshu, Japan. A. Habit. B. Achene with pappus. C. Stamen. D. Floret. E. Opened corolla with stamens. F. Capitulum. G. Juvenile involucre. Drawn by S. Domon.

＊

＊

B

CD

E

A

F

G


multicellular hairs on both sides, deeply cordate at base, mucronate at apex: petioles 6–18 cm long, winged in the apical part, vaginate at base, deccurent to the stem, not amplexicaul, almost glabrous. Upper cauline leaves narrowly ovate, 1–4.5 cm long, 1–2.5 cm wide, shortly petiolate or sessile. Flowering September to October. Capitula 5–10, arranged in a loose corymb; peduncles 0.5–1 cm long, divaricate, densely pubescent with grayish multicellular hairs. Capitula 1–3 in an axially corymb. Involucres campanulate, greenish, 10–15 mm in diameter (in vivo), 12–20 mm in diameter (in sicco), 14–17 mm long, more or less arachnoid; phyllaries 10-seriate; outer phyllaries ovate, 5–6 mm long, acuminate with recurved tips; middle phyllaries narrowly ovate, 8 mm long, acute with recurved tips; inner phyllaries narrowly ovato-lanceolate, 11 mm long, acute, erect; bracts ovate, 5 mm long, mucronate; setae 8–9 mm long. Corollae pale purplish violet, 10–15 mm long; lobes 2–5 mm long; throats 2 mm long; tubes 6–8 mm long; anthers 5–6 mm long, deep bluish purple; tails 2 mm long. Pappi 2-whorled, inner 10–12 mm long, outer 3 mm long, whitish. Achenes 5 mm long, glabrous, light gray-brownish purple.

Japanese name: Tobishima-tôhiren.新和名：トビシマトウヒレンDistribution: Tobishima Island, Sakata-shi,

Yamagata Pref., Honshu, Japan (endemic). Habitat: Growing under sparse Pinus

thunbergii woods near the seashore.Additional specimens examined: JAPAN. Honshu.

Yamagata Pref., Sakata-shi, Tobishima Island, Arasaki – Mt. Kashiwagi-yama, alt. ca. 50 m, 29 Sept. 2009, S. Kato 2, 3 (TNS 741158, 741160); Tobishima Island, Takamori-jinja shrine, alt. ca. 60 m, 29 Sept. 2009, S. Kato 2 (TNS 741161–741165); Tobishima Island, Arasaki, on the floor of Pinus thunbergii plantation, alt. ca. 10 m, 27 Sept. 2011, S. Kato s.n. (TNS 1155518–1155531).

Note: Saussurea muramatsui is clearly distinguished from S. katoana Kitam. by having herbaceous leaves, 6-seriate involucral phyllaries and patent to ascending outer and middle phyllaries. Saussurea muramatsui is distributed in Yamagata as well as Akita and

Aomori Prefectures and usually grows in and along summer-green woods of inland mountains. However, S. katoana occurs under sparse Pinus thunbergii woodlands in maritime areas.

2) Saussurea yanagitae Kadota, sp. nov. [Figs. 3–4]

Saussurea yanagitae clearly differs from S. sagitta Franch. in having the pendulous habit, larger, thicker, dimly lustrous, slightly sagittate cauline leaves, smaller, densely arachnoid involucres, more capitula arranged in a corymb and shorter florets.

Type: JAPAN. Honshu. Niigata Pref., Uonuma-shi, Yunotani, Mt. Arasawa-dake, Maegura, alt. 1165 m, 1 September 2012, Hiromitsu Yanagita, Emiko Yanagita, Sadayoshi Mezaki & Yoko Nedzu s.n. (TNS 1158501–holotype, Fig. 3).

A herbaceous perennial, 50–70 cm tall. Rhizome oblique to horizontal, ca. 0.5 cm in diameter, with string-like roots. Stem pendulous, striate, narrowly winged, smooth, 3–4 times branched in the apical part. Basal leaves withered. Cauline leaves gradually diminishing in size from the middle part of the stem. Lower cauline leaves coriaceous, dimly lustrous, narrowly ovate, 13–15 cm long, 3.5–4 cm wide, denticulate, glabrous on both sides, cuneate to rounded or shallowly cordate at base, slightly sagittate, acuminate at apex: petioles 5–8 cm long, winged in the apical part, deccurent to the stem, not amplexicaul, glabrous. Middle cauline leaves similar to the upper in shape; petioles 0–2 cm long, amplexicaul. Upper cauline leaves narrowly ovato-lanceolate, 4–9 cm long, 0.5–1 cm wide, sessile and amplexicaul. Flowers in August to September. Capitula 5, arranged in a compact corymb; peduncles 1–5 mm long in the terminal corymb, ascending at an acute angle, pubescent with brownish multicellular hairs. Capitula 3–4 in an axially corymb. Involucres narrowly cylindrical, 3.3–3.7 mm in diameter (in vivo), 5–6 mm in diameter (in sicco), 7.5–9 mm long, densely arachnoid; phyllaries 6-seriate;

272 植物研究雑誌　第 88巻　第 5号 2013年 10月

Fig. 3. Type of Saussurea yanagitae Kadota (JAPAN. Honshu. Niigata Pref., Uonuma-shi, Yunotani, Mt. Arasawa-dake, Maegura, alt. 1165 m, 1 Spet. 2012, H. Yanagita & al. s.n., TNS 1158501, holotype).


Fig. 4. Saussurea yanagitae Kadota. A. Habitat. B–C. Habit. D. Upper part of plant. E. Inflorescence. Photo by H. Yanagita & al. on 1 Sept. 2012 at Mt. Arasawa-dake, Uonuma-shi, Niigata Pref., Honshu, Japan.

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Fig. 5. Type of Saussurea yubarimontana Kadota (JAPAN, Hokkaido, Sorachi Subpref., Yubari-shi, Mt. Yubari-dake, Kaneopetsu-gawa, Sorachi Subpref., Yubari-shi, Mt. Yubari-dake, Kaneopetsu-gawa, alt. 530 m, K. Horie s.n., TNS 1109688, holotype).


outer phyllaries broadly ovate, 2 mm long, mucronate with brownish purple tips less than 1 mm long; middle phyllaries ovate, 5–6 mm long, cuspidate with brownish purple tips ca. 1 mm long; inner phyllaries lanceolate, ca. 8 mm long, acute; bracts ovate, 3 mm long, acuminate; setae 4 mm long. Corollae pale purplish violet, 9 mm long; lobes 3 mm long; throats 1 mm long; tubes 5 mm long; anthers 4 mm long, deep bluish purple. Pappi 2-whorled, inner 6–7 mm long, outer 2 mm long, whitish. Mature achenes unknown. Immature achenes 1.5 mm long, glabrous, ivory.

Japanese name: Arasawa-tôhiren.新和名：アラサワトウヒレンDistribution: Mt. Arasawa-dake, Uonuma-

shi, Niigata Pref., central Honshu, Japan (endemic).

Habitat: On the rocky cliff of mountains in the alpine zone (Fig. 4, A–B).

Note: Saussurea yanagitae is very unique in having the pendulous habit among the Japanese species of Saussurea.

Saussurea sagitta is also endemic to Honshu, Japan and is disjunctly distributed in the Tohoku and Chubu districts (Kadota 2011a). The range of S. yanagitae is located between the Tohoku and Chubu districts and is interposed by the ranges of S. sagitta.

3) Saussurea yubarimontana Kadota, sp. nov. [Figs. 5–6]

Saussurea yubarimontana is discriminated from S. kudoana Tatew. & Kitam. var. uryuensis Kadota [= S. uryuensis (Kadota) Kadota] in having longer involucres, shorter apices of outer and middle involucral phyllaries, narrowly ovate leaf blades with cuneate to shallowly cordate bases and non-deccurent petioles. Saussurea kudoana is clearly different from S. yubarimontana in the presence of persistent, thick, dark brownish green, dimly lustrous basal leaves at flowering time.

Type: JAPAN. Hokkaido, Sorachi Subpref., Yubari-shi, Mt. Yubari-dake, Kaneopetsu-gawa,

in serpentine area, alt. 530 m, K. Horie s.n. (TNS 1109688–holotype, Fig. 6).

A slender, herbaceous perennial, 50–95 cm tall. Rhizome horizontal to oblique, up to 1.5 cm in diameter, with string-like roots. Stem suberect to slightly declining, striate, not winged, smooth, 1–4 times branched in the apical part or simple. Basal leaves withered or sometimes persistent at anthesis. Cauline leaves abruptly diminishing in size from the middle part of the stem. Lower cauline leaves subcoriaceous, narrowly ovate to ovate, 8–13 cm long, 2–6.5 cm wide, serrate, glabrous on both sides, ciliate with multicellular hairs along the margin, cuneate to rounded or shallowly cordate at base, acuminate at apex: petioles 4–8 cm long, winged in the apical part, also winged in the basal part and almost not deccurent to the stem, not amplexicaul, glabrous or sparingly ciliate with multicellular hairs along the wings. Upper cauline leaves clearly smaller than the lower, narrowly ovato-lanceolate, 3–5 cm long, 0.5–1.5 cm wide, serrate, acuminate at apex, shortly petioled and not amplexicaul. Flowers in August to September. Capitula (3–) 5–8, arranged in a compact corymb; peduncles 0–9 mm long in the terminal corymb, ascending at an acute angle, glabrous and smooth. Capitula 2–4 in an axially corymb. Involucres narrowly cylindrical, 5–6 mm in diameter, (11–)13–15 mm long, sparingly arachnoid along the margin of phyllaries; phyllaries 8-seriate; outer phyllaries broadly ovate, 5 mm long, cuspidate with brownish purple tips 1 mm long; middle phyllaries ovate, 5–8 mm long, cuspidate with brownish purple tips 1–2 mm long; inner phyllaries lanceolate, ca. 10 mm long, obtuse; bracts narrowly ovate, 4–8 mm long, acuminate; setae 5 mm long. Corollae purplish violet, 11 mm long; lobes 4 mm long; throats 1 mm long; tubes 5–6 mm long; anthers 4 mm long, deep reddish purple. Pappi 2-whorled, inner 8–9 mm long, outer 3 mm long, whitish. Achenes ca. 5 mm long, glabrous, ivory tinged deep purple.

Japanese name: Yûbari-tôhiren.新和名：ユウバリトウヒレン

276 植物研究雑誌　第 88巻　第 5号 2013年 10月

A

B C

Fig. 6. Habit and involucres of Saussurea yubarimontana Kadota (A–B) and involucres of S. riederi Herder var. yezoensis (Maxim.) Kitam. (C). A. Habit. B. Involucre. At Kaneopetsu-gawa, Mt. Yubaridake, Yubari-shi, Hokkaido, northern Japan on 9 Aug. 2010. C. Involucre. At. the summit area, Mt. Yubaridake, on 10 Aug. 2010.


Distribution: Mt. Yubari-dake, Sorachi Subpref. and Mt. Bôzu-yama, Iburi Subpref., Hokkaido, Japan (endemic). In serpentine areas.

Additional specimens examined: JAPAN. Hokkaido. Iburi Subpref., Yufutsu-gun, Mukawa-cho, Hobetsu, Mt. Bôzu-yama, alt. 240 m, in serpentine area, 19 Aug. 1979, K. Horie 6 (TNS 684255, 684257–684258); Mt. Bôzu-yama, alt. 250 m, in serpentine area, on the floor of a scattered Picea glehnii forest, 19 Aug. 1979, K. Horie s.n. (TNS 1109705–1109706); Mt. Bôzu-yama, alt. 250 m, in serpentine area, 7 Sept. 1980, K. Horie s.n. (TNS 1109704).

Note: The plants from Hobetsu cited as ʻS. riederi var. yuparensisʼ by Nosaka and Horie (1993, 1994) should be ascribed to S. yubarimontana.

In Mt. Yubari-dake, Hokkaido, habitat segregation of S. yubarimontana and S. riederi var. yezoensis was clearly observed: S. yubarimontana was found in serpentine areas of the lower part of the mountain, while, S. riederi var. yezoensis grew in meadows of the alpine zone.

4) Saussurea uryuensis (Kadota) Kadota, stat. nov.

Saussurea kudoana Tatew. & Kitam. var. uryuensis Kadota in Mem. Natn. Sci. Mus. Tokyo no. 20: 87, figs. 1, 2A (1987); in Kato & Ebihara, Endem. Pl. Jap.: 148 (2011); Horie & al., Wild Pl. Serpent. Areas Asahikawa: 39 (2012).

Type: JAPAN. Hokkaido. Kamikawa Subpref., Shibetsu-shi, Mt. Inuushibetsu-dake, on serpentine soils, 31 August 1986, Y. Kadota 13638 (TNS–holotype).

Japanese name: Uryû-tôhiren (Kadota 1987).和名：ウリュウトウヒレン（門田）Distribution: Kamikawa and Soya Subprefs.,

Hokkaido, Japan (endemic). On serpentine, grassy slopes.

Specimens examined: JAPAN. Hokkaido. Soya Subpref., Esashi-gun, Nakatonbetsu-cho, Kikusui-gawa, 27 Aug. 1989, J. Haginiwa 6961–6962, 6968–6969 (TNS 956969, 956962, 956968, 981061); Kikusui-gawa, alt. 200 m, 6 Sept. 1998, K. Horie 1 (TNS 684271–684272). Kamikawa Subpref., Shibetsu-shi, Onʼnebetsu, Mt. Inuushibetsu-dake, alt. 300 m, 3 Aug. 1990, K. Horie s.n.

(TNS 1162396); Onʼnebetsu, Mt. Inuushibetsu-dake, alt. 500 m, 3 Aug. 1990, K. Horie 3 (TNS 684265–684267). Uryû-gun, Horokanai-cho, Mt. Shiratori-yama, alt. 180 m, 8 Aug. 1991, K. Horie 2 (TNS 684268–684270). Kamikawa-gun, Kenbuchi-cho, ʻKakushi-gen-yaʼ, Sept. 1916, H. Koidzumi 65135 (TNS 178959; Saussurea tesioensis H. Koidz., nom. nud.). Wassamu-cho, Fukuhara, Kakurei field, alt. 180 m, 5 Sept. 1987, K. Horie 4 (TNS 684262–684264). Asahikawa-shi, Etanbetsu. Maruyama, alt. 520 m, 7 Aug. 1994, K. Horie 5 (TNS 684259–684261); Etanbetsu. Maruyama, alt. 500 m, 20 Aug. 2010, K. Horie s.n. (TNS 1168993–1168994). Fukagawa-shi, in the vicinity of Takadomari dam, 19 Aug. 1997, S. Umezawa 97002 (TNS 651825); Takadomari, alt. 200 m, 30 July 2007, Y. Kadota 073321–073323 (TNS 771068–771070).

Note: Since the publication of S. kudoana var. uryuensis (Kadota 1987) a considerable amount of specimens has accumulated in the Herbarium TNS. As a result of the examination of these materials it turns out that this entity should be recognized as a distinct species as in the following key.

Kadota (1987) denoted that involucral phyllaries of both S. kudoana and S. uryuensis were 6-seriate. However, it should be corrected that the two species have 8-seriate involucral phyllaries as shown in Kadota (1987; figure 2).

The plants from Nuporomapporo, Horokanai, Shibetsu and Wassma cited as ʻS. riederi var. yuparensisʼ by Nosaka and Horie (1993, 1994) should be ascribed to S. uryuensis.

Saussurea kudoana Tatew. & Kitam. in Acta Phytotax. Geobot. 4: 76 (1935); Kitam. in Mem. Coll. Sci. Kyoto Imp. Univ. ser. B, 13 [Comp. Jap. I]: 170, pl. 22, f. 1 (1937); H. Hara in Bot. Mag. (Tokyo) 52: 125 (1938); Ohwi, Fl. Jap.: 1227 (1953); Kadota in Mem. Natn. Sci. Mus. Tokyo (20): 86, f. 2, B (1987) – Saussurea riederi Herder f. kudoana (Tatew. & Kitam.) Ohwi, Fl. Jap. rev. ed.: 1227 (1932); Ko. Ito, Alp. Fl. Hokkaido: 15, pl. 38 (1981) – Saussurea riederi subsp. kudoana (Tatew. & Kitam.) Kitam. in Kitam. & Murata, Col. Illust. Herb. Pl. Jap. 1: 27 (1980) & in Satake & al., Wild Fl. Jap. 1: 223 (1981) – Saussurea riederi var. kudoana (Tatew. & Kitam.) T. Shimizu, New Alp. Fl. Jap. 1: 25, 289 (1982); T. Yamaz. in T. Yamaz., Alp.

278 植物研究雑誌　第 88巻　第 5号 2013年 10月

Fig. 7. Type of Saussurea riederi Herder var. yuparensis Kitam. (JAPAN, Hokkaido, Mt. Yubari-dake, no date, G. Koidzumi s.n., KYO). Right inset shows involucre of the right plant. Lectotypified by S. Smirnov in 2006.


Fl. Jap.: 80 (1985). Type: JAPAN. Hokkaido. Hidaka Subpref., Mt. Apoi-dake, 17 August 1912, K. Kondo s.n. (SAPS–holotype !).

Japanese name: Hidaka-tôhiren (Tatewaki and Kitamura 1935).和名：ヒダカトウヒレン（館脇，北村）Distribution: Mt. Apoi-dake and Mt.

Horoman-dake, along the river Nikanbetsu-gawa, Hidaka Subpref., Hokkaido, Japan (endemic). On serpentine, rocky slopes.

Representative specimens examined: JAPAN. Hokkaido, Hidaka Subpref., Samani-gun, Samani-cho, Mt. Apoi-dake, 24 Aug. 1933, H. Hara s.n. (KYO); Mt. Apoi-dake, 31 July 1935, S. Kitamura s.n. (KYO); Mt. Apoi-dake, alt. 800 m, 14 Aug. 1935, K. Miyabe & K. Hino s.n. (KYO); Mt. Apoi-dake, 26 July 1941, S. Okuyama s.n. (TNS 68874); Mt. Apoi-dake, 9 Sept. 1986, Y. Kadota 13864 (TNS 692604–692605, 692694, 692697–692700). Samani-cho, Mt. Horoman-dake, alt. 300 m, 21 Aug. 1966, M. Hotta 16838 (KYO); Mt. Horoman-dake, alt. 700 m, 21 Aug. 1966, M. Hotta 16849 (KYO); along the river Horoman-gawa, 29 Aug. 1985, J. Haginiwa 16950–16952 (TNS 966950–966952). Horoizumi-gun, Erimo-cho, Rusaki, along the river Nikanbetsu-gawa, 5 Sept. 2005, C. Komai s.n. (TNS 755778).

Key to the Saussurea kudoana group1A. Basal leaves persistent at anthesis; leaves coriaceous, dark purple-brownish green, dimly lustrous; leaf blades narrowly ovate, rounded to cordate at base; involucres ca. 10 mm long; involucral phyllaries brownish purple to the middle part; outer and middle phyllaries acuminate with suberect tips less than 1 mm long .................................................... S. kudoana1B. Basal leaves usually deciduous or sometimes persistent at anthesis; leaves subcoriaceous, bright yellowish green, not lustrous; involucral phyllaries brownish purple marginally or apically, or sometimes entirely greenish; outer and middle phyllaries with ascending to slightly recurved tips 1–3 mm long ............................... 22A. Blades of lower cauline ovate to widely ovate, cordate at base; involucres 10–12 mm long, frequently arachnoid throughout the surface; apices of the outer and middle phyllaries 2–3 mm long ................................... S. uryuensis2B. Blades of lower cauline narrowly ovate,

shallowly cuneate to rounded or shallowly cordate at base; involucres (11–)13–15 mm long, arachnoid only along the margin of the phyllaries; apices of the outer and middle phyllaries 1–2 mm long ........ S. yubarimontana

As already stated the name S. riederi Herder subsp. yezoensis (Maxim.) Kitam. var. yuparensis Kitam. is synonymous with S. riederi var. yezoensis. A systematic treatment of S. riederi var. yezoensis is summarized as follows.

Saussurea riederi Herder in Bull. Soc. Nat. Mosc. 41(2): 35 (1869); Maxim. in Bull. Acad. Sci. St.-Pétersb. 19: 518 (1874); Franch. in Bull. Herb. Boiss. 5: 544 (1897); Hultén, Fl. Kamtch. 4: 209, f. 9 (1930); Kom., Fl. Pen. Kamtsch. 3: 178 (1930); Nakai in Bot. Mag. (Tokyo) 45: 521, in clavi (1931); Kitam. in Acta Phytotax. Geobot. 4: 7 (1935) & Mem. Coll. Sci. Kyoto Imp. Univ. ser. B, 13 [Comp. Jap. 1]: 170 (1937); H. Hara, Enum. Sperm. Jap. II: 248 (1952); Ohwi, Fl. Jap.: 1228 (1953); Lipsch., Genus Saussurea: 244 (1979); Vorosch., Opred. Rast. Sovjet. Dalyn. Vostoka: 578 (1982); Barkalov in Kharkev., Pl. Vasc. Or. Extr. Soviet. 6: 292 (1992). Lectotype (Lipschicz 1979): RUSSIA. Kamtschatka, no date, J. G. v. Rieder (LE!).

var. yezoensis Maxim. in Mel. Biol. 9: 343 (1874); Franch. in Bull. Herb. Boiss. 5: 544 (1897) – Nakai in Bot. Mag. (Tokyo) 45: 521 (1931); Kitam. in Acta Phytotax. Geobot. 1: 117 (1931); H. Hara, Enum. Sperm. Jap. II: 248 (1952); Ohwi, Fl. Jap.: 1227 (1953); Kadota in Mem. Natn. Sci. Mus. Tokyo no. 20: 88 (1987); Barkalov in Kharkev., Pl. Vasc. Or. Extr. Soviet. 6: 292, in nota (1992) – Saussurea riederi Herder subsp. yezoensis (Maxim.) Kitam. in Comp. Jap. 1: 171 (1937) & in Kitam. & al., Col. Illust. Herb. Pl. Jap. I: 27 (1980) & in Satake & al., Wild Fl. Jap. III: 223, pl. 204-3 (1981); H. Koyama, Fl. Jap. IIIb: 158 (1995). Lectotype (Lipschicz 1979): JAPAN. Hokaido. ʻOshima, Hakodate, Schuro, ad declivia siccis sat frequensʼ, 23 Aug. – 4 Sep. 1861, C. J.

280 植物研究雑誌　第 88巻　第 5号 2013年 10月

Fig. 8. Type of Saussurea yuki-uenoana Kadota (JAPAN, Honshu, Miyagi Pref., Igu-gun, Marumori-machi, Hippo, Waseda, alt. ca. 490 m, 24 Sept. 2010, Yuki Ueno 46668, TNS 1025461, holotype).


Fig. 9. Habit of Saussurea yuki-uenoana Kadota. Right top corner inset shows capitulum. At Waseda, Hippo, Marumori-machi, Igu-gun, Miyagi Pref., Honshu, Japan on 3 Oct. 2011.

282 植物研究雑誌　第 88巻　第 5号 2013年 10月

Maximowicz 712 (LE!). Saussurea riederi subsp. yezoensis (Maxim.)

Kitam. var. yuparensis Kitam. in Mem. Coll. Sci. Kyoto Imp. Univ. ser. B, 13 [Comp. Jap. 1]: 173 (1937), syn. nov.; Lipsch., Genus Saussurea: 245 (1979) – Saussurea riederi var. yezoensis Maxim. f. yuparensis (Kitam.) Nosaka in J. Fac. Sci. Hokkaido Univ., ser. 5, Bot. 9: 242 (1974) – Saussurea riederi subsp. kudoana (Tatew. & Kitam.) Kitam. var. yuparensis Kitam.: H. Koyama, Fl. Jap. IIIb: 158 (1995) – Saussurea kudoana Tatew. & Kitam. var. yuparensis (Kitam.) Yonek in J. Jpn. Bot. 80: 331 (2005). Type: JAPAN. Hokkaido, Ishikari, Mt. Yubari, no date, G. Koidzumi s.n. (holotype–KYO; Fig. 7).

Saussurea nambuana Koidz. in Bot. Mag. (Tokyo) 29: 115 (1915). Type: JAPAN. Prov. Rikuchu [Iwate Pref.], Mt. Hayachine-san, no date, G. Koidzumi s.n. (holotype–KYO!).

Japanese name: Nagaba-kita-azami (Koidzumi 1916, ʻnagaha-kita-azamiʼ).和名：ナガバキタアザミ（ナガハキタアザミ，

小泉）ユウバリキタアザミ（北村）

5) Saussurea yuki-uenoana Kadota, sp. nov. [Figs. 8–9]

Saussurea yuki-uenoana is characterized by having narrowly cylindrical involucres, shortly recurved, 10–12-seriate involucral phyllaries, smaller but more capitula in dense, compact corymbs, coriaceous, ovate cauline leaves and strongly ribbed stem without wings. Saussurea sendaica Franch. differs from S. yuki-uenoana in having campanulate involucres, 8-seriate involucral phyllaries, long patent outer and middle phyllaries and simple, clearly winged stems.

Type: JAPAN. Honshu. Miyagi Pref., Igu-gun, Marumori-machi, Hippo, Waseda, alt. ca. 490 m, 1.8 m in height, 24 September 2010, Yuki Ueno 46668 (TNS 1025460–1025461–holotype; TUS–isotype; Fig. 8).

A sturdy, herbaceous perennial, 70–180 cm tall. Rhizome oblique to horizontal, 1–2 cm in

diameter, with cord-like roots. Stem erect, striate, usually not winged, or sometimes narrowly winged, glabrous, 5–10 times branched from the lower part; wings, if present, ca. 1 mm wide, entire. Basal leaves usually withered at anthesis. Cauline leaves gradually diminishing in size. Lower cauline leaves coriaceous, not lustrous, ovate, 11–20 cm long, 9–15 cm wide, coarsely dentate, sparingly pubescent with brownish multicellular along midribs, cordate at base, acute to acuminate at apex: petioles 7–16 cm long, deccurent and ribbed or very narrowly winged, not vaginate nor amplexicaul, almost glabrous. Upper cauline leaves narrowly ovate, 4–11 cm long, 1–5 cm wide, with short petioles 0.5–1 cm long. Flowers in September to October. Capitula 4–5, arranged in a compact corymb; peduncles 0.5–0.8 cm long, ascending, densely pubescent with grayish multicellular hairs. Capitula 1–5 in an axially compact corymb. Involucres narrowly cylindrical, purplish, 5–8 mm in diameter (in vivo), 8–12 mm in diameter (in sicco), 13–17 mm long, arachnoid; phyllaries 11–12-seriate; outer phyllaries broadly ovate, 3 mm long, acute with recurved, purplish tips; middle phyllaries narrowly ovate, 8 mm long, acuminate with recurved, purplish tips; inner phyllaries narrowly ovato-lanceolate, 11 mm long, acuminate with erect, purplish tips; bracts narrowly ovate, 2–3 mm long, acuminate; setae 6–7 mm long. Corollae pale purplish violet, 10–14 mm long; lobes 4 mm long; throats 1 mm long; tubes 5–8 mm long; anthers 4 mm long, deep bluish purple; tails 1 mm long. Pappi 2-whorled, inner 10–11 mm long, outer 3 mm long, whitish. Achenes 4 mm long, glabrous, light gray-brownish purple.

Japanese name: Abukuma-tôhiren.新和名：アブクマトウヒレンDistribution: Marumori-machi and Shiroishi-

shi, Miyagi Pref., Honshu, Japan (endemic). Habitat: Growing in sparse deciduous woods

or herbal stand.Additional specimens examined: JAPAN. Honshu.

Miyagi Pref., Igu-gun, Marumori-machi, Hippo, Waseda,


alt. ca. 490 m, 24 Sept. 2010, Y. Ueno 46665–46667, 46669 (TNS 1025459, 1025462–1025464, TUS); Marumori-machi, Hippo, Waseda, alt. 505 m, 3 Oct. 2011, Y. Kadota 111001–111008 (TNS 1155584–1155594). Shiroishi-shi, Fukuoka, Yatsumiya, Zawo Mountains, Mt. Fubô-san, alt. 981 m, 15 Sept. 2008, Y. Kadota 085426–085432 (TNS 777436–777442).

Note: Plants resembling Saussurea yuki-uenoana occur in Fukushima Prefecture, northern Honshu, Japan (Kadota 2012). However, no voucher specimens have been kept in the Herbaria KYO and TNS at present Further studies are needed for identification of the plants.

I would like to show my sincere thanks to Mr. Shin-ei Kato for his help and presents of specimens of S. katoana from Tobishima Island; to Mr. Shôzô Domon for his beautiful line drawing of S. katona; to Mr. Yuki Ueno for his guidance to the locality of S. yuki-uenoana and his presents of specimens and photos of this species; Dr. Kenji Horie for his guidance to the localities of S. yubarimontana and S. uryuensis and his presents of specimens and photographs of both species; to Mr. Shun Umezawa for his useful information on S. riederi and S. yubarimontana; to Messrs. Hiromitsu Yanagita and Sadayoshi Mezaki and Mmes. Emiko Yanagita and Yoko Nedzu for their kind field survey in Mt. Arasawa-dake and their presents of specimens of S. yanagitae; to Dr. S. Smirnov for discussion about species of the genus Saussurea; to Dr. Hideki Takahashi (SAPT), Dr. Hidetoshi Nagamasu and Dr. Minoru N. Tamura (KYO) and Dr. I. Grubov (LE) for their kind permission of the examination of Saussurea specimens.

Literatures citedBarkalov V. Yu. 1993. Saussurea. In: Charkevicz

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Kadota Y. 1987. A new variety of Saussurea kudoana Tatew. & Kitam. (Asteraceae) from Hokkaido, Japan. Mem. Natn. Sci. Mus., Tokyo (20): 83–90.

Kadota Y. 2004. A new species of Saussurea (Asteraceae) from Honshu, Japan. J. Jpn. Bot. 79: 235–240.

Kadota Y. 2007. Systematic studies of Asian Saussurea (Asteraceae) I. Saussurea kubotae, a new species from western Japan. J. Jpn. Bot. 82: 259–265.

Kadota Y. 2008. Systematic studies of Asian Saussurea (Asteraceae) II. Two new species from northern Japan. J. Jpn. Bot. 83: 284–294.

Kadota Y. 2009. Systematic studies of Asian Saussurea (Asteraceae) III. Saussurea fuboensis, a new species from the southernmost part of Tohoku District, northern Japan. J. Jpn. Bot. 84: 177–183.

Kadota Y. 2011a. Saussurea. In: Kato M. and Ebihara A. (eds.), Endemic Plants of Japan. A Book Series from the National Museum of Nature and Science No. 11. pp. 147–148, maps 1544–1574. Tokai University Press, Hadano (in Japanese).

Kadota Y. 2011b. On the entity of Saussurea triptera Maxim. var. mikurashimensis Kitam. (Asteraceae) from Mikurajima Island, Idzu Islands, Japan (Systematic studies of Asian Saussurea (Asteraceae) IV). Mem. Natl. Mus. Nat. Sci., Tokyo no. 47: 411–415.

Kadota Y. 2011c. Systematic studies of Asian Saussurea (Asteraceae) V. Two new species from eastern Hokkaido, Japan and Sakhalin, Russia. J. Jpn. Bot. 86: 197–204.

Kadota Y. 2012. Notes on Saussurea and Parasenecio (Asteraceae) from Fukushima Prefecture, Japan. Flora Fukushima no. 28: 1–11 (in Japanese).

Kitamura S. 1935. Les Saussurées du Japon; leur classification et leur distribution. Acta Phytotax. Geobot. 4: 1–14.

Kitamura S. 1937. Compositae Japonicae. Pars prima. Mem. Coll. Sci., Kyoto Imp. Univ., ser. B, 13: 140–212.

Kitamura S. 1980. Saussurea. In: Kitamura S., Murata G. and Hori M., Coloured Illustrations of Herbaceous Plants of Japan (Sympetalae), revised edition. pp. 24–29, figs. 7–8, pl. 7. Hoikusha Publishing Co. Ltd., Osaka (in Japanese).

Kitamura S. 1981. Saussurea. In: Satake Y., Ohwi J., Kitamura S., Watari S. and Tominari T. (eds.), Wild Flowers of Japan. Herbaceous Plants (including Dwarf Subshrubs). 3: 220–224, pls. 201–207. Heibonsha Ltd., Tokyo (in Japanese).

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Moskva (in Russian).Nosaka S. 1959. A preliminary report of the phanerogam

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門田裕一：アジア産トウヒレン属（キク科）の分類学的研究 VI．北海道産の 1 新種と 1 新組合わせ及び本州産の 3 新種　キク科トウヒレン属において，北海道から 1新種ユウバリトウヒレン Saussurea yubarimontana Kadotaと1新組合わせ S. uryuensis (Kadota) Kadota，本州から 3新種，トビシマトウヒレン S. katoana Kadota，アラサワトウヒレン S. yanagitae Kadota，アブクマトウヒレン S. yuki-uenoana Kadotaを報告した．1)トビシマトウヒレン（基準産地：山形県酒田市飛島）　トビシマトウヒレンは，沿海地のクロマツ林床に生え，葉の質が厚い点ではムツトウヒレン S. hosoiana Kadotaに似る．しかし，ムツトウヒレンは花期に根生葉が生存し，茎は花茎状となり，頭花の柄が短く，総苞片が 8列で，茎の翼が幅狭くかつ全縁であることで区別される．ホクロクトウヒレン S. nipponica Miq. hokurokuensis Kitam. [= S. nipponica Miq. var. hokurokuensis (Kitam.) Ohwi, S. hokurokuensis Kitam., in sched.]もトビシマトウヒレンに似るが，トビシマトウヒレンは葉の質がさらに厚く，頭花が少数個つき，総苞がより大型で，茎の翼に鋸歯があることなどで区別される．山形県を始めとして，秋田県，青森県の内陸山地（青森県では沿海地にも分布）にはトガヒゴタイS. muramatsui Kitam.が分布するが，これは総苞片が 6列で，総苞外片と中片は開出ないし斜上し，葉が草質である点で区別される．　種形容語はこの新種トウヒレンの実態の解明に貢献された加藤信英氏への献名である．　トビシマトウヒレンは山形県酒田市飛島の固有種である． 2)アラサワトウヒレン（基準産地：新潟県魚沼市荒沢岳）　アラサワトウヒレンはヤハズトウヒレン S. sagitta Franch.に似ているが，①植物体はより大型で岩壁から懸垂し，②茎葉は大型で革質，鈍い光沢があり，基部は

わずかにヤハズ形となり，③頭花はより多数で 3–5個が散房状につき，④総苞はより小型で密にクモ毛に被われ，⑤小花がより小さい点で区別される．　種形容語は発見者の柳田宏光氏への献名である．　高山帯の岩壁に生育し，茎が懸垂するアラサワトウヒレンの性状は日本産トウヒレン属でも特異なものである．形態的に似たヤハズトウヒレンは本州の日本海側多雪地の高山風衝草原に生える．ヤハズトウヒレンは本州の固有種で東北地方と中部地方に隔離的に分布する．アラサワトウヒレンの分布域は中部地方の北部に限られ，ヤハズトウヒレンの分布域に挟まれるかたちになる．3)ユウバリトウヒレン（基準産地：北海道夕張市夕張岳，カネオペツ川）　夕張岳の山麓部の蛇紋岩地域にはトウヒレン属植物が分布することは以前から知られており，ユウバリキタアザミ S. riederi Herder subsp. yezoensis (Maxim.) var. yuparensis Kitam.と考えられてきた（例えば，Nosaka 1959，1961，梅沢 2009，2010など）．しかしながら，ユウバリキタアザミのタイプ標本を検討した結果，これは花期に根生葉を欠くこと，茎葉が茎の下部から上部に向かって次第に小さくなること，総苞片が 5列である点で，ナガバキタアザミ S. riederi var. yezoensis (Maxim.) Kitam.に含まれることが明らかになった．ユウバリトウヒレンは茎葉が中部から急に小型となり，総苞片が 8列であることで特徴付けられる，ヒダカトウヒレン S. kudoana Tatew. & Kitam.に近縁な植物である．ユウバリトウヒレンはヒダカトウヒレンとは異なり，葉は黒褐色を帯びた緑色ではなく鮮やかな黄緑色で，総苞片の先端が短く尖ることなどで明瞭に区別される．　ユウバリトウヒレンは夕張岳の山麓部の他，勇払郡むかわ町穂別の坊主山に分布する．これらの産地はいずれ


も蛇紋岩崩壊地である．4)ウリュウトウヒレン（基準産地：北海道士別市犬牛別岳）　ウリュウトウヒレンは，下部の茎葉の葉身が卵形で基部が心形，総苞が長さ 10–12 mmとやや短いこと，そして総苞外片と中片の先端部が長く伸びることで，ユウバリトウヒレンと区別される．また，ウリュウトウヒレンは花期に根生葉が生存しないこと，茎葉は常に鮮緑色で光沢がないこと，総苞外片と中片の先端がより長く伸びることなどでヒダカトウヒレンと異なる．このため，ウリュウトウヒレンを独立種 S. uryuensis (Kadota) Kadotaとみなすことが適当であることが分かった．ウリュウトウヒレンは旭川以北の道北地方の超塩基性岩地に広く分布する．ヒダカトウヒレン群 3種の検索表を本文中に示した．5)アブクマトウヒレン（基準産地：宮城県伊具郡丸森町）

　アブクマトウヒレンは形態的に特異な種で，次のような特徴をもっている．①総苞は狭筒形，②総苞片は11〜 12列，③総苞外片と中片の先端は短く反曲し，④茎はよく分枝して散房状の腋生花序を多数付け，⑤下部の茎葉は革質で，卵形，粗い鋸歯があり，湾入はなく，基部は心形，⑥茎には鋭い稜があるか極く狭い翼がある．日本産のトウヒレン属としては大型で，高さは1.8 mに達する．この地域に多いセンダイトウヒレン S. sendaica Franch.は，総苞が鐘形で，総苞片が 8列，総苞外片と中片の先端が長く開出ないし反曲し，茎に翼が発達し分枝が少ない点などでアブクマトウヒレンから区別できる．　種形容語は発見者の上野雄規氏への献名である．　アブクマトウヒレンは宮城県南部と福島県の阿武隈山地に分布する．（国立科学博物館植物研究部）

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Systematic Studies of Asian Saussurea Asteraceae) VI. A ...268 植物研究雑誌 第88 巻 第5 号 2013 年10 月 Fig. 1. Type of Saussurea katoana Kadota (JAPAN, Honshu, Yamagata

Systematic Studies of Asian Saussurea Asteraceae) VI. A ...268 植物研究雑誌第88 巻第5 号 2013 年10 月 Fig. 1. Type of Saussurea katoana Kadota (JAPAN, Honshu, Yamagata