J. Jpn. Bot. 87: 1–21 (2012)

Systematic Studies of Asian Aconitum (Ranunculaceae) XIV. Four New Species of Subgenus Lycoctonum from , Northern

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 September 10, 2011)

Four new species of Aconitum subgenus Lycoctonum are described from Hokkaido, northern Japan. Aconitum asahikawaense described from the serpentine areas of and its neighboring areas is characterized by sepals with dark brownish- purple spots at apex, long-projecting helmet beaks and strigose carpels. Aconitum hiroshi- igarashii described from the Niseko zone, southwestern Hokkaido, is characterized by blue-violet flowers, pedicels strigose with curved hairs and entire nectary labia. Aconitum ikedae also described from the Niseko zone is characterized by violet flowers, pedicels villous with spreading hairs and glandular hairs and irregularly serrate nectary labia. Aconitum umezawae described from Oshima Peninsula, southwestern Hokkaido, is delineated by yellowish-white flowers, pedicels villous with spreading hairs, thicker and shorter nectary spurs, bracteoles positioned below the middle of the pedicels and irregularly serrate nectary labia. As a result eight species in total are recognized in subgenus Lycoctonum from Hokkaido, northern Japan and they are all endemic to Japan. A key for the eight species is provided here.

Key words: Aconitum asahikawaense, Aconitum hiroshi-igarashii, Aconitum ikedae, Aconitum umezawae, Hokkaido, new species, Niseko zone, subgenus Lycoctonum.

Aconitum gigas H. Lév. & Vaniot f. bicolor Lycoctonum plants with blue-violet flowers grow Kadota & Umezawa (Ranunculaceae) was wild mainly in the Niseko zone, southwestern described from Lake Kokkuri-ko, Rankoshi- Hokkaido. The 'Niseko zone' is a region in cho, Shiribeshi Subpref., Hokkaido, northern which Rankoshi-cho, the type locality of A. Japan (Kadota 2001, 2006a, 2006b, Umezawa gigas f. bicolor, Niseko-cho and Kucchan-cho 2009). At that time this plant was considered a are included. variant with two-toned-flowers (blue-violet and During 2006 to 2010 conclusive field white) within the species A. gigas because blue- examinations were executed for subgenus violet-flowered species in subgenus Lycoctonum Lycoctonum plants in the Niseko zone and of the genus Aconitum had never been reported its surrounding region. This clarified that two from Hokkaido. However, after the publication species with blue-violet flowers are present in of this 'form', it has been gradually known that the area. The two new species are considered

—1— 2 植物研究雑誌 第 87 巻 第 1 号 2012 年 2 月 to belong to ser. Umbrosa (Steinb. ex Nakai) Also after the publication of A. gigas f. Kadota instead of ser. Volubilia (Steinb. ex bicolor, subgenus Lycoctonum plants having Nakai) Tamura & Lauener since both have sepals with dark brownish-purple spots at cylindrical helmets and thick and short nectary apex have been collected in serpentine areas spurs. The first species is characterized by of Asahikawa and its adjacent region, central having pedicels villous with patent hairs and Hokkaido. Later field explorations revealed that glandular hairs and the second is delineated by the plants represent a distinct species which is pedicels strigose with curved hairs. As a result discriminated from A. gigas by the pubescence of the field examinations the two new species of carpels and the position of bracteoles on were frequently found sympatrically and many the pedicels as well as the flower color. This natural hybrid derivatives were also observed in is described here as the fourth new species, populations. Investigating the holotype specimen A. asahikawaense, which is regarded as a of A. gigas f. bicolor, it turned out that the ultrabasicosaxophyte (Toyokuni 1955). individual is a hybrid between the two species In conclusion, eight species are recognized in question. Actually the Kokkuri-ko population in the genus Aconitum subgenus Lycoctonum in proved to consist of the two species and a hybrid Hokkaido, northern Japan. The eight species are swarm between them. In conclusion the two enumerated below. species are here described as A. ikedae and A. hiroshi-igarashii, respectively and the name A. Enumeration of species of subgenus gigas f. bicolor is ascribed to a hybrid between Lycoctonum from Hokkaido the two species. Subgenus Lycoctonum (DC.) Peterm.: In 2008 and 2009 fieldwork was also Tamura & Lauener in Notes Roy. Bot. done in Mt. Obira-yama [or Mt. Ôhira-yama], Gard. Edinb. 37: 431 (1979) – Tamura, Nat. Shimamaki-mura, Shimamaki-gun, Shiribeshi Pflanzenfam. ed. 2,17aIV : 278 (1995) – Kadota Subpref., southwestern Hokkaido, Japan. During in K. Iwats. & al., FL. Jpn. IIa: 268 (2006). the fieldwork plants having yellowish-white Sect. Lycoctonum: Tamura & Lauener in flowers and pedicels covered with a mixture of Notes Roy. Bot. Gard. Edinb. 37: 118 (1978) – spreading hairs and curved hairs as well as the Tamura, Nat. Pflanzenfam. ed. 2, 17aIV: 280 plants belonging to A. gigas were obtained there. (1995) – Kadota in K. Iwats. & al., Fl. Jpn. IIa: This situation was quite similar to that observed 268 (2006). in the Mashike Mountains; the discovery of Ser. Umbrosa (Steinb. ex Nakai) Kadota in A. mashikense (Kadota 2001). In other words K. Iwats. & al., Fl. Jpn. IIa: 268 (2006). the presence of an undescribed species having Ser. Lycoctonia Tamura & Lauener in Notes pedicels villous with spreading hairs was Roy. Bot. Gard. Edinb. 37: 451 (1979), p.p. – expected because the pedicels of A. gigas are Tamura, Nat. Pflanzenfam. ed. 2, 17aIV: 284 covered with only curved hairs. The third new (1995), p.p. species with yellowish-white flowers and villous pedicels has been additionally found in Oshima 1. Aconitum asahikawaense Kadota, sp. nov. Peninsula, southwestern Hokkaido, based on [Figs. 1–2] the intense field examinations. This species Haec species ab Aconito gigate sepalis apice is described as A. umezawae here. Aconitum fusco-purpuleo-punctatis, carpelis et folliculis umezawae is distinguished from A. mashikense vulgo glabris et bracteolis positis prope basi by having faint flower color, thick and short pedicellorum differt. nectary spurs and roundish, irregularly serrate Type: JAPAN. Hokkaido. Kamikawa nectary labia. Subpref., Asahikawa-shi, Shinkai, the Kamui February 2012 Journal of Japanese Botany Vol. 87 No.1 3

Fig. 1. Type of Aconitum asahikawaense Kadota (JAPAN. Hokkaido. Kamikawa Subpref., Asahikawa-shi, the Kamui (Horonai) Mountains, along the Oroen-gawa river, serpentine area, 29 June 2011, K. Horie s.n., TNS 1028851, holotype). 4 植物研究雑誌 第 87 巻 第 1 号 2012 年 2 月

(Horonai) Mountains, along the Oroen-gawa and short, 1–2 mm long, incurved to straight; River, serpentine area, alt. 370 m, 29 June 2011, labium 1 mm long, subentire; claw 8–9 cm mm K. Horie s.n. (TNS 1028851–holotype, Fig. 1; long, nearly straight. Stamens glabrous, anthers TNS 1028852–1028853 –isotype). 0.5 mm long, filaments 4 mm long, staminal A subscapose, perennial herb, 0.7–1.0 m tall. teeth absent or present. Carpels 3, more or less Root 0.5–1 cm in diameter at the ground surface strigose with rough-surfaced, curved hairs on the level, branched, rhizomatous. Stem rather dorsal side or rarely glabrous. Follicles 15–18 slender, terete, suberect to declining, branched mm long, divergent; seeds black, 2.5 mm long, from the middle part, sparingly strigose with 1.5 mm wide, trigonous, lamellate transversely, rough-surfaced, curved hairs in the distal part. not alate. Basal leaves persistent at anthesis. Blades of Japanese name: Kamui-reijinsô (nom. nov.). basal leaves roundish reniform in outline, 12–22 新和名：カムイレイジンソウ cm long, 14–24 cm wide, medially 7–9-lobed Distribution: Asahikawa and its surrounding to 2–3 cm from the base, glabrous or sparingly region, cenral Hokkaido, northern Japan (Fig. 9, sericeous with straight, long, golden, rough- triangle). Endemic to Japan. surfaced, adpressed hairs along the veins on Additional specimens examined: JAPAN. Hokkaido. the abaxial side, deeply to shallowly cordate at Hokkaido, Kamikawa Subpref., Kamikawa Subpref., base; middle lobe obovate-rhombic, 6–11 cm Nakagawa-gun, Wassamu-cho, Seiwa, a serpentine area, long, 5–9 cm wide, acute, laciniate; laciniae alt. 200 m, 25 June 2006, fl., K. Horie s.n. (TNS 757746– narrowly ovate, 5–7 mm wide, acuminate; 757750). Asahikawa-shi, Kamuikotan, Kamui-iwa, along a trail, 31 May 2008, fl., H. Igarashi s.n. (TNS 774976, petioles 30–40 cm long, glabrous, 4-angled, 774980–774981); Kamuikotan, Kamui-iwa, a serpentine hollow. Cauline leaves similar to basal in shape, area, alt. 200 m, 14 June 2010, fl., K. Horie s.n. (TNS similarly pubescent to the basal, diminishing in 1107380–1107383); Kamuikotan, alt. 158 m, 20 June 2008, size, petiolate to sessile. Inflorescence terminal, fr., Y. Kadota 091131–091133 (TNS 735731–735733); racemose, 15–30 cm long, 12–24-flowered, Kamui-cho, Harushinai Tunnel, 1 May 2008, fl., H. Igarashi (TNS 774977–774978); Kamui-cho, Harushinai, in indeterminate condition. Pedicel of the alt. 114 m, 20 June 2008, fl., Y. Kadota 091111–091121 lowermost flower 3–8.5 cm long, arched to (TNS 735507–735515); Kamuikotan, along the Kamui Dai incurved, strigose with rough-surfaced, curved 3-sen-gawa river, 31 May 2008, fl., H. Igarashi s.n. (TNS hairs throughout; bracteoles 2, linear, 1–2 mm 774979); Kamuikotan, alt. 180 m, 14 June 2010, K. Horie s.n. (TNS 1119941–1119942); Kamuikotan, serpentine area, long, less than 0.5 mm wide, situated near the alt. 180 m, 24 June 2011, K. Horie s.n. (TNS 1028854– base of the pedicel. Flowers in June to July, dull 1028856); Shinkai, the Kamui (Horonai) Mounatins, Mt. yellow and brownish purple-spotted at apex Kamui-yama, along a tributary of Kamui Dai-1-sen-gawa in each sepal , 2.3–2.7 cm long, strigose with river, a serpentine area, alt. 220 m, 20 June 2010, fl., K. rough-surfaced, curved hairs on the abaxial Horie s.n. (TNS 1107387–1107389); the Kamui (Horonai) Mounatins, along the upstream of Oichanunpe-gawa, side; helmet cylindrical, 17–23 mm long, 10–14 serpentine area, alt. 375 m, 29 June 2011, K. Horie s.n. mm wide, 13–15 mm high, rounded at apex, (TNS 1028810–1028812); Mt. Kamui-yama, along the with relatively long beak 3–5 mm long, the line upstream of the Oichanunpe-gawa river, a serpentine area, between the base and the beak oblique (ca. 15º); alt. 550 m, 22 June 2010, fl., K. Horie s.n. (TNS 1107390– 1107392); Mt. Kamui-yama, along the upstream of the Ino- lateral sepals distorted obovate, 7–10 mm long, gawa river, a serpentine area, alt. 230 m, 14 June 2010, 5–8 mm wide, densely hirsute with long, rough- fl., K. Horie s.n. (TNS 1107384–1107386); Mt. Arashi- surfaced, flexuous hairs (pollen-collecting hairs) yama, serpentine area, alt. 260 m, 1 July 2011, K. Horie s.n. at the aperture on the adaxial side. Nectaries (TNS); Arashiyama, Mt. Chikabumi-yama – Mt. Hanmen- glabrous, yellowish-white; blade 2–3 mm long, yama, a serpentine area, alt. 220 m, 18 June 2010, K. Horie s.n. (TNS 1119938–1119940). Fukagawa-shi, Irumuchippu, 1 mm wide, not inflated and tapering to the alt. 449 m, 20 June 2008, fl., Y. Kadota 091101–091103 spur, tubular part 1 mm long; spur rather thick (TNS 735522–735524). February 2012 Journal of Japanese Botany Vol. 87 No.1 5 C Kadota. A. Habit. At Oroen-gawa, the Kamui (Horonai) Mountains, Asahikawa, central Hokkaido, Japan, on 29 June 2011. Courtesy of of Courtesy 2011. June 29 on Japan, Hokkaido, central Asahikawa, Mountains, (Horonai) Kamui the Oroen-gawa, At Habit. A. Kadota. asahikawaense asahikawaense

Aconitum Dr. K. Horie. B. Flower. C. Nectary. B–C. Harushinai, Asahikawa, central Hokkaido, on 29 June 2011. Asahikawa, central Hokkaido, Japan, on 20 June 2008. June 20 on Japan, Hokkaido, central Asahikawa, 2011. June 29 on Hokkaido, central Asahikawa, Harushinai, B–C. Nectary. C. Flower. B. Horie. K. Dr. A B Fig. 2. 2. Fig. 6 植物研究雑誌 第 87 巻 第 1 号 2012 年 2 月

Note: Aconitum asahikawaense is 和名：エゾレイジンソウ distinguished from A. gigas H. Lév. & Vaniot by Distribution: From southwestern to northern the sepals with brownish-purple spots at apex, and eastern Hokkaido, northern Japan (Kadota usually glabrous carples and bracteoles situated 2011, map. 120). Endemic to Japan. near the base of the pedicels. Representative specimens examined: JAPAN. Plants with purplish stems and/or leaves Hokkaido. Ochotsk Subpref., Sahri-gun, Shari-cho, Shiretoko Peninsula, Mt. Unabetsu-dake, alpine meadow, purplish beneath (e.g., Viola yubariana and V. 10 Aug. 1980, K. Sato 1146 (SAPS 14026); Kiyosato- sacchalinensis f. alpina (Violaceae) and Veronica cho, Mt. Shari-dake, alt. 850–1200 m, 24 July 1983, H. schmidtiana subsp. senanensis var. yezoalpina Takahashi 4462 (SAPS 17284). -gun, Tsubetsu- (Scrophulariaceae)), are reported among cho, Aioi, 19 June 2006, H. Takahashi & Y. Takashima Japanese ultrabasicosaxophytes (Toyokuni 32934 (SAPS 19838). Kitamis-shi, On'netô, 16 July 1937, M. Tatewaki & T. Yamanaka 28177 (SAPS 19818); Tokoro, 1955). Aconitum asahikawaense is similarly River Hiyoshi-gawa, 3 June 2008, H. Igarashi s.n. (TNS considered to be an example of this kind of 774973, 774975). Monbetsu-gun, Engaru-cho, Shirataki, ultrabasicosaxophytes due to the presence of the Kami-Shiyûbetsu, 4 June 2008, H. Igarashi s.n. (TNS dark brownish-purple spots at the apex of sepals. 774972). Nemuro Subpref., Shibetsu-gun, Shibetsu-cho, 21 July 1884, K. Miyabe s.n. (SAPS 19825); Shibetsu-cho, NE side of Mt. Kunbetsu-dake, upstream of Sakimui River, 2. Aconitum gigas H. Lév. & Vaniot in along forest road, 18 June 2007, H. Takahashi & al. 33599 Bull. Bot. Soc. Fr. 53: 389 (1906) – Tamura & (SAPS 17578). Subpref., Kutcharo [Kawakami- Lauener in Notes Roy. Bot. Gard. Edinb. 37: 458 gun, Teshikaga-cho, Kussharo], Biborogoye, lakeside, (1979), incl. pl. Hokkaido –Tamura in Kitam. & 24 Aug. 1917, K. Miyabe s.n. (SAPS 19823); Kutcharo, Pontô, 21 Aug. 1917, K. Miyabe s.n. (SAPS 19821); Murata, Col. llust. Herb. Pl. Jpn. II: 209 (1980), Kutcharo-kohan, 13 Aug. 1915, K. Miyabe s.n. (SAPS incl. pl. Hokkaido; in Satake & al., Wild Flow. 19822); Teshikaga-cho, N side of Lake Kussharo, Mt. Jpn. II: 63, pl. 58-4 (1982), excl. pl. Honshu; in Mokoto-yama, lower elevation of Gozo-zawa, alt. 380–420 Ohwi (Kitag.), New Fl. Jpn.: 707 (1992), excl. m, 8 July 1988, H. Takahashi 8637 (SAPS 19778). Akan- gun, Akan-cho, S side of Lake Akan, alt. 420–460 m, 15 pl. Sakhalin – Kadota in K. Iwats. & al. , Fl. Jpn. June 1990, H. Takahashi 10309 (SAPS 19779). Kushiro- IIa: 268 (2006); in H. Ohashi & al., New Illust. shi, Akan-cho, Mt. O-Akan-dake, 1 June 1974, Y. Goda 46 Fl. Jpn.: 135 (2008); in M. Kato & Ebihara, End. (TNS 141633); Akan-cho, Panketô, alt. ca. 450 m, 13 July Sp. Jpn.: 55, 290, map 120 (2011) – Umezawa, 2009, U. Umezawa s.n. (TNS 739580–739581). Ashoro- Alp. Fl. Hokkaido 240 (2009) – Lycoctonum gun, Ashoro-cho, On'netô, 13 July 2008, S. Umezawa s.n. (TNS–photo). Soya Subpref., Esashi-gun, Esashi-cho, 2 gigas (H. Lév. & Vaniot) Nakai in J. Jpn. Bot. July 1971, M. Yamanoi 110 (TNS 319420); Utanobori- 13: 406 (1937); in Bull. Natn. Sci. Mus., Tokyo cho [currently Esashi-cho], Mt. Poronupuri, 2 July 1989, no. 32: 15 (1953) – Sugim., Keys Herb. Pl. Jpn. Y. Kadota s.n. (TNS 643507). Hama-Tonbetsu-cho, I: 154 (1978), incl. pl. Hokkaido. Usotan, 27 June 2006, S. Umezawa s.n. (TNS 768268). Kamikawa Subpref., -shi, Mt. Kudo-san, alt. 400 Type: JAPAN. Insula Yezo, in sylvis Sirafu m, 3 June 2000, T. Shibata s.n. (TNS 698263–698267). [Hokkaido, Shiribeshi Subpref., Abuta-gun, Nakagawa-gun, Nakagawa-cho, Kamiji, 9 June 1976, J. Niseko-cho, Hirafu, cf. Kadota 2006b], 28 July Haginiwa JH 000210 (TNS 950210). Bifuka-cho, at the 1905, U. Faurie 6934 (P 201588–holotype !, entrance of Matsuyama Mire, 6 July 2007, S. Umezawa Kadota 2006, fig. 4A; BM, KYO 119, 861, TI– 07070611 (TNS 772847). Kamikawa-gun, Aibetsu-cho, 4 July 1982, M. Hara 5358 (SAPS 19754). Teshio Subpref., isotype !). Nakagawa-gun, Abeshunai, 22 June 1905, S. Ninouye Aconitum tatewakii Miyabe var. vegetum in s.n. (SAPS s.n.); in the vicinity of Otoineppu, Hokkaido Trans. Sappro Nat. Hist. Soc. 14(2): 72 (1935). Univ. Nakagawa Exper. forest, alt. 100–400 m, 23 June Type: JAPAN. Yezo [Hokkaido]. Prov. Ishikari, 1978, G. Murata & al. 38401 (SAPS 19836). Uryû-gun, Horokanai-cho, 10 June 1959, T. Takahashi 1717 (TNS Jôzankei, 1896, K. Miyabe & Y. Tokubuchi s.n. 305284); Horokanai-cho, Moshiri, Uryû Exper. Forest of (SAPS–holotype !). Hokkaido Univ., alt. 300–400 m, 15 June 1978, G. Murata Japanese name: Ezo-reijinsô (Miyabe 1935). & al. 37779 (TNS 395785–395786); Horokanai-cho, Mt. February 2012 Journal of Japanese Botany Vol. 87 No.1 7

Santô-zan, alt. 990 m, 24 July 2011, S. Umezawa s.n. (TNS crassioribus brevioribus et foliis basalibus 1139248–1139250). Sorachi Subpref., Uryu-gun, Numata- grandioribus; ab A. gigate floribus violaceis cho, N of Lake Horopiri, SE of Tappu-tôge pass, 26 June 2005, H. Takahashi & S. Miyazawa 31841 (SAPS 19761); differt. Uryû-cho, 26 June 1894, Y. Jin s.n. (SAPS 19840). Ishikari Type: JAPAN. Hokkaido. Shiribeshi Subpref., -shi, Hôheikyo –– Mt. Sapporo-dake, Subpref., Niseko-cho, Mt. Konbu-dake, alt. 413 360–900 m, 13 June 1983, H. Takahashi 4155, 4165 (SAPS m, 23 June 2009, fl., Y. Kadota 093121 (TNS 19766–19767). Iburi Subpref., Shiraoi-gun, Shiraoi-cho, 1101916–holotype; Fig. 3). Morino, Oboshi-zawa forest road, 13 Aug. 2006, K. Oita 3113 (SAPS s.n.). Shiribeshi Subpref., Shimamaki-gun, A subscapose, perennial herb, 0.8–1.2 m tall. Shimamaki-mura, Mt. Obira-yama (Ôhira-yama), alt. 280 Root 0.5–2 cm in diameter at the ground surface m, 25 June 2009, fl., Y. Kadota 093151, 093154 (TNS level, branched, rhizomatous. Stem robust, 1101918–1101920, 1101923). terete, suberect to declining, branched from Note: The type locality of A. gigas, 'Sirafu' the lower part, sparingly strigose with rough- (sometimes erroneously spelled as 'Jirafu'), has surfaced, curved hairs in the distal part. Basal been unclear because a locality so-pronounced leaves persistent at anthesis. Blades of basal is not known in Hokkaido. However, according leaves roundish reniforme in outline, 22–28 to the itinerary of U. Faurie the locality is cm long, 26–36 cm wide, medially 7–9-lobed now regarded as Hirafu which is located in to 2.5–3.5 cm from the base, glabrous on both Kucchan-cho, Abuta-gun, Shiribeshi Subpref., sides or sparingly sericeous with long, golden, southwestern part of Hokkaido, the Niseko zone rough-surfaced, adpressed hairs along the veins (Matsui pers. comm.; cf. Kadota 2006b). on the abaxial side, deeply cordate at base; A wide range of variation in the length of middle lobe obovate-rhombic, 9.5–14 cm long, nectary spurs among populations from the 11–12 cm wide, acute, laciniate; teeth narrowly eastern part of Hokkaido is known and plants ovate, 3–5 mm wide, acute; petioles 22–45 having long spurs are sometimes found there. cm long, glabrous, 4-angled, hollow. Cauline These plants were frequently attributed to A. leaves similar to basal in shape, diminishing umbrosum (Korsh.) Kom. sensu Kadota (2006a, in size, petiolate to sessile. Inflorescence 2006b) from Honshu (denoted as the Honshu terminal, racemose, (20–)25–45 cm long, ca. materials below), central Japan based on the 30–40-flowered, in indeterminate condition. difference of spur length (e.g., Tamura 1980, Pedicel of the lowermost flower 4.5–11 cm long, 1982, Tamura and Lauener 1979). However, the incurved to arched, strigose with rough-surfaced Honshu materials are characterized by longer, curved hairs throughout; bracteoles 2, linear, tubular parts of nectary blades and longer labia ca. 1 cm long, ca. 1 mm wide, situated below (Kadota 2006b) and are significantly different the middle of the pedicel. Flowers in June, dull from either A. gigas or A. umbrosum of Sakhalin pale reddish violet, 2–3 cm long, glabrous on and East Siberia. The plants with relatively the abaxial side; helmet broadly cylindrical, ca. long spurs from eastern Hokkaido have shorter, 22 mm long, 11–12 mm wide, ca. 14 mm high, tubular parts of nectary blades and should be rounded at apex, with short beak 2–3 mm long, thus included in A. gigas. The Honshu materials the line between the base and the beak oblique should be treated as another distinct species. (ca. 45º); lateral sepals distorted obovate, ca. This issue will be discussed elsewhere. 1 cm long, 7–8 mm wide, hirsute with long, smooth-surfaced patent hairs (pollen-collecting 3. Aconitum hiroshi-igarashii Kadota, sp. nov. hairs) at the aperture on the adaxial side. [Figs. 3–4] Nectaries glabrous, yellowish-white; blade 4 mm Differt ab Aconito pterocaule cassibus long, 2 mm wide, not inflated and tapering to cylindricis brevirostris, calcaribus nectariorum the spur, tubular part ca. 1 mm long; spur thick 8 植物研究雑誌 第 87 巻 第 1 号 2012 年 2 月

Fig. 3. Type of Aconitum hiroshi-igarashii Kadota (JAPAN. Hokkaido. Shiribeshi Subpref., Niseko-cho, Mt. Konbu-dake, 23 June 2009, Y. Kadota 093121, TNS 1101916, holotype). February 2012 Journal of Japanese Botany Vol. 87 No.1 9

A B C

Fig. 4. Aconitum hiroshi-igarashii Kadota. A. Habit. B. Flower. C. Nectary. At Mt. Konbu-dake, the Niseko zone, Niseko-cho, western Hokkaido, Japan, on 25 June 2009. 10 植物研究雑誌 第 87 巻 第 1 号 2012 年 2 月

Fig. 5. Type of Aconitum ikedae Kadota (JAPAN. Hokkaido, Shiribeshi Subpref., Abuta-gun, Niseko-cho, Mt. Konbu-dake, 23 June 2009, Y. Kadota 093135b, TNS 1102951, holotype). February 2012 Journal of Japanese Botany Vol. 87 No.1 11 and short, 2 mm long, slightly incurved; labium vs. villous with spreading hairs and glandular 1 mm long, entire; claw ca. 1 cm mm long, hairs) and the margin of nectary labia (entire vs. slightly incurved. Stamens glabrous, anthers irregularly, minutely toothed). 0.5 mm long, filaments 4 mm long, staminal teeth absent. Carpels 3, glabrous. Follicles 1–1.5 4. Aconitum ikedae Kadota, sp. nov. cm long, divergent; seeds 2.5 mm long, 2 mm [Figs. 5–6] wide, black, trigonous to broadly ellipsoidal, Differt ab Aconito hiroshi-igarashii pedicellis transversely lamellate, not alate. villosis cum pilis patentibus et eis glandulatis et Japanese name: Konbu-reijinsô (nom. nov.). labiis nectariorum minute irregulariter serratis. 新和名：コンブレイジンソウ Type: JAPAN. Hokkaido, Shiribeshi Distribution: The Niseko zone and Toyoura, Subpref., Abuta-gun, Niseko-cho, Mt. Konbu- southwestern Hokkaido, northern Japan (Fig. 9, dake, alt. 511 m, 23 June 2009, Y. Kadota diamond). Endemic to Japan. 093135b (TNS 1102951–holotype; Fig. 5). A subscapose, perennial herb, 0.7–1.2 m tall. Additional specimens examined: JAPAN. Hokkaido. Shiribeshi Subpref., Isoya-gun, Rankoshi-cho, Mt. Root 0.5–1 cm in diameter at the ground surface Horomui-dake, alt. 272 m, 26 June 2009, fl., Y. Kadota level, branched, rhizomatous. Stem robust, 093161–093165 (TNS 1101938–1101944); Rankoshi-cho, terete, suberect to declining, branched from Mt. Horonai-dake, alt. 267 m, 26 June 2009, fl., Y. Kadota the middle part, sparingly villous with straight, 093187 (TNS 1101958–1101961). Abuta-gun, Niseko-cho, rough-surfaced, adpressed hairs in the distal Mt. Konbu-dake, 26 June 2005, I. Ikeda s.n. (TNS 755929); Niseko-cho, Mt. Konbu-dake, 2 July 2005, fl., H. Igarashi part. Basal leaves persistent at anthesis. Blades s.n. (TNS 755932); Niseko-cho, Mt. Konbu-dake, alt. 510 of basal leaves roundish reniform in outline, m, 18 June 2008, fl., Y. Kadota 091013–091025, 091026 18–24 cm long, 21–32 cm wide, medially (TNS 734889, 734891–734903); Niseko-cho, Mt. Konbu- 7–9-lobed to 2.5–3 cm from the base, sparingly dake, alt. ca. 500 m, 19 June 2008, fl., Y. Kadota 091073– 091074, 091082, 091084–091086 (TNS 735019–735021, sericeous with straight, long, golden, rough- 735023, 735030–735032); Niseko-cho, Mt. Konbu-dake, surfaced, adpressed hairs along the veins on alt. 413 m, 23 June 2009, fl., Y. Kadota 093121, 093123– the abaxial side, deeply cordate at base; middle 093127 (TNS 1101908–1101912, 1101915–1101917); lobe obovate-rhombic, 10–16 cm long, 8–13 cm Niseko-cho, Mt. Konbu-dake, alt. 511 m, 23 June 2009, wide, acute, laciniate; laciniae narrowly ovate, fl., Y. Kadota 093138–093142 (TNS 1140651–1140658); Niseko-cho, Mt. Konbu-dake, alt. 374 m, 26 June 2009, 4–6 mm wide, acuminate; petioles 20–55 cm fl., Y. Kadota 093186 (TNS 1101962–1101964). Iburi long, glabrous, 4-angled, hollow. Cauline leaves Subpref., Abuta-gun, Toyoura-cho, Rebunge, 4 June similar to basal in shape, similarly pubescent 1972, fl., M. Hara 5356 (SAPS 19624; heavily damaged to the basal, diminishing in size, petiolate to by insects); Toyoura-cho, Rebunge, 29 July 1971, fr., M. Hara 5357b (SAPS 19756). Oshima Subpref., Kameda- sessile. Inflorescence terminal, racemose, 17–30 gun, Nanae-cho, Narukawa, alt. 440 m, 21 June 2010, fl., Y. cm long, ca. 20–30-flowered, in indeterminate Kadota 1006501 (TNS 1028889–1028891). condition. Pedicel of the lowermost flower Etymology: The specific epithet is dedicated 3.5–7 cm long, slightly incurved, villous with to Mr. Hiroshi Igarashi, Chitose-shi, Hokkaido, rough-surfaced, curved hairs throughout and who contributed to the discovery of this new additionally with a few glandular hairs in the species. distal part; bracteoles 2, linear, ca. 5 mm long, ca. Note: Aconitum hiroshi-igarashii is 0.5 mm wide, situated below the middle of the characterized by having pale blue-violet flowers, pedicel. Flowers in June, dull pale reddish violet, cylindrical helmets, pedicels strigose with rough- ca. 2.5 cm long, glabrous on the abaxial side; surfaced curved hairs and thick but short nectary helmet broadly cylindrical, 20–25 mm long, 11– spurs and is different from A. ikedae in the 12 mm wide, 15–18 mm high, rounded at apex, pedicel pubescence (strigose with curved hairs with short beak 2–3 mm long, the line between 12 植物研究雑誌 第 87 巻 第 1 号 2012 年 2 月 C Kadota. A. Habit. B. Flower. C. Nectary. At Mt. Konbu-dake, the Niseko zone, Niseko-cho, western Hokkaido, Japan, on 24 June 2009. At Mt. Konbu-dake, the Niseko C. Nectary. A. Habit. B. Flower. Kadota. ikedae Aconitum Fig. 6. 6. Fig. A B February 2012 Journal of Japanese Botany Vol. 87 No.1 13 the base and the beak oblique (ca. 45º); lateral Hamamasu-gun, Hamamasu-mura, Mt. Kogane- sepals distorted obovate, 9 mm long, 6 mm yama, alt. 380 m, 17 June 2000, fl., S. Umezawa wide, hirsute with long, rough-surfaced flexuous 20104 (TNS 697735–holotype !). hairs (pollen-collecting hairs) at the aperture on Japanese name: Mashike-reijinsô (Kadota the adaxial side. Nectaries glabrous, yellowish- 2001). white; blade 4 mm long, 1 mm wide, not inflated 和名：マシケレイジンソウ and tapering to the spur, tubular part longer than Distribution: Mashike and Kabato 1 mm; spur thin and short, 3 mm long, slightly Mountains, central Hokkaido, northern Japan incurved; labium 1 mm long, irregularly and (Kadota, 2001, fig. 5). Endemic to Japan. minutely serrate; claw ca. 1 cm mm long, nearly Representative specimens examined: JAPAN. straight. Stamens glabrous, anthers 0.5 mm long, Hokkaido. Rumoi Subpref., Mashike-gun, Mashike-cho, Mt. Shokanbetsu-dake, 28 July 1933, I. Yamamoto 5361 filaments 4 mm long, staminal teeth absent or (SAPS 19848); Mt. Shokanbetsu-dake, 28 July 1983, H. present. Carpels 3, glabrous. Mature follicles and Takahashi 4532 (SAPS 19763); Mt. Shokanbetsu-dake, alt. seeds unknown. 1480 m, 18 July 1990, Y. Kadota 90102 (TNS 9024605); Japanese name: Niseko-reijinsô (nom. nov.). Mt. Shokanbetsu-dake, alt. 650 m, 6 Aug. 2000, S. 新和名：ニセコレイジンソウ Umezawa 20120 (TNS 754127–754128); Mt. Shokanbetsu- dake, alt. 300 m, 2 Aug. 2006, Y. Kadota s.n. (TNS Distribution: The Niseko zone and Toyoura, 757745). Sorachi Subpref., Kabato-gun, Shintotsukawa- southwestern Hokkaido, northern Japan (Fig. 9, cho, Mt. Pin'neshiri, alt. 380 m, 8 July 2000, S. Umezawa star). Endemic to Japan. 20119 (TNS 754129–754130); Mt. Pin'neshiri, alt. 950 m, Additional specimens examined: JAPAN. Hokkaido. 8 July 2000, S. Umezawa 20117 (TNS 754116–754119). Shiribeshi Subpref., Abuta-gun, Niseko-cho, Mt. Konbu- Uryû-gun, Uryû-cho, Uryû-numa mire, alt. 640–860 m, dake, alt. 510 m, 18 June 2008, Y. Kadota 091012 (TNS 8 July 1998, H. Takahashi 25154 (SAPS 19785); Uryû- 734904); Niseko-cho, Mt. Konbu-dake, alt. 511 m, 23 June chio, Mt. Shokanbetsu-dake, Uryû-numa mire, alt. 660 m, 2009, Y. Kadota 093135 (TNS 1028818–1028819); Niseko- 21 Aug. 2000, S. Umezawa 20122–20125 (TNS 754123– cho, Mt. Konbu-dake, alt. 374 m, 26 June 2009, Y. Kadota 754126); Uryû-numa mire, alt. 840 m, 28 July 2007, Y. 093184 (TNS 1101968). Isoya-gun, Rankoshi-cho, Lake Kadota 073211 (TNS 771083). Ishikari Subpref., Ishikari- Kokkuri-ko, alt. 576 m, 18 June 2008, Y. Kadota 091037 gun, Tôbetsu-cho, Mt. Kamuishiri-yama, alt. 380 m, 8 June (TNS 734906). Iburi Subpref., Abuta-gun, Toyoura-cho, 2000, S. Umezawa 20107 (TNS 697731, paratype of A. Rebunge, 27 June 1971, fl., M. Hara 5357a (SAPS 19755); mashikense). Hamamasu-gun, Hamamasu-mura [currently Toyoura-cho, Rebunge, 11 Aug. 1980, fr., M. Hara 5353 Ishikari-shi, Hamamasu-ku], Mt. Kogane-yama, alt. 380 (SAPS 19753). m, 17 June 2000, S. umezawa 20105–20106, 20112–20113 Etymology: The specific epithet is dedicated (TNS 697732–697733, 697739–697740, 754113, 754120); Mt. Kogane-yama, alt. 470 m, 17 June 2000, S. umezawa to Mr. Ikuro Ikeda, Niseko-cho, Hokkaido, who 20102–20103, 20110, 20114–20115 (TNS 697685, made a contribution to the discovery of this new 697741–697742, 754111–754112); Hamamasu-mura, species. Gryôchi, alt. 80 m, 17 June 2000, S. Umezawa 20101, Note: Aconitum ikedae is distinguished 20108, 20111 (TNS 697682–697687, 697737, 697738, from A. hiroshi-igarashii by having the pedicels 697743. villous with golden, rough-surfaced, spreading 6. Acontum soyaense Kadota in J. Jpn. Bot. hairs and the nectary labia with irregularly, 82(1): 41, figs. 1–2 (2007) – Umezawa, Alp. Fl. minutely serrate margin. Hokkaido 241 (2009). Type: JAPAN. Hokkaido. Soya Subpref., 5. Aconitum mashikense Kadota & Esashi-gun, Naka-Tonbetsu-cho, Machineshiri, Umezawa in J. Jpn. Bot. 76(1): 20, figs. 1, left, River Kikusui-gawa gorge, alt. 95 m, 17 June 2, 3 (2001) – Kadota in K. Iwats. & al., Fl. Jpn. 2006, Y. Kadota 063028 (TNS 757737–holotype IIa: 268 (2006) – Umezawa, Alp. Fl. Hokkaido !). 240 (2009). Japanese name: Sôya-reijinsô (Kadota 2007). Type: JAPAN. Hokkaido. Ishikari Subpref., 和名：ソウヤレイジンソウ 14 植物研究雑誌 第 87 巻 第 1 号 2012 年 2 月

Distribution: Soya, northern Hokkaido, san, Mt. Haku-un-dake, 19 July 1961, K. Inagaki northern Japan. Endemic to Japan. & al. s.n. (SAPT !). Specimens examined: JAPAN. Hokkaido. Soya Aconitum gigas H. Lév. & Vaniot var. Subpref., Esashi-gun, Naka-Tonbetsu-cho, Machineshiri, tatewakii (Miyabe) Toyok., in sched. (SAPS). River Kikusui-gawa gorge, alt. 95 m, 17 June 2006, fr., Y. Kadota 063031 (TNS 757744); at the same locality, Japanese name: Hidaka-reijinsô (nom. nov.). 31 July 2006, fr., Y. Kadota 063201–063204 (TNS 新和名：ヒダカレイジンソウ 757740–757743); at the same locality, 26 June 2007, fl. Distribution: Hidaka, Yubari and Ishikari & fr., Y. Kadota 071101–071104 (TNS 770971–770974); Mountain Ranges, central Hokkaido, northern Naka-Tonbetsu-cho, Mt. Pin'neshiri-dake, 27 June 2008, S. Japan. Endemic to Japan. Umezawa (TNS–photo). Note: Aconitum soyaense is well Representative pecimens examined: JAPAN. Hokkaido. Hidaka Subpref., Samani-gun, Samani-cho, characterized by having shallowly cylindrical Mt. Pin'neshiri, alt. 430 m, fl., 20 June 2006, Y. Kadota helmets with short beaks and with roundish s.n. (TNS 757783). Horoizumi-gun, Erimo-cho, Meguro, tops, carpels and follicles hirsute with ascending Lake Toyoni-ko, alt. 200 m, 20 June 2006, fl., Y. Kadota hairs, elongated, divaricate branches, smaller s.n. (TNS 757778–757782). Shizunai-gun, Mt. Petegari- seeds, inflated nectary blades, saccate nectary dake, alt. 1200–1730 m, 28 July 1970, H. Koyama & N. Fukuoka 3325 (TNS 272146). Saru-gun, Hidaka-cho, spurs and leaves sericeous with long, appressed along a tributary of the River Saru-gawa, alt. 800 m, 23 hairs beneath. The pedicels of A. soyaense Jult 2005, fl., Y. Kadota 053006–053026 (TNS 755862– (including the type specimen) are covered with 755880). Tokachi Subpref., Kamikawa-gun, Shimizu- a mixture of rough-surafced, curved hairs and cho, Mt. Tsurugisan, alt. 532 m, 25 June 2011, fl. & fr., S. Umezawa 1 (TNS 1138383–1138385); Mt. Memuro-dake, smooth-surfaced, spreading hairs. This kind of 22 July 1914, S. Nishida s.n. (SAPS 19803); Mt. Pibairo- pedicel pubescence strongly suggests that these dake, 28 July 1914, S. Nishida s.n. (SAPS 19802); Mt. plants were of hybrid origin and at least two Satsuna-dake, 24 Jult 1917, S. Nishida s.n. (SAPS 19798). undescribed species occur in northern Hokkaido. Sorachi Subpref., Sorachi-gun, Yubari-shi, Mt. Yubari- Kadota (2007) noted that A. soyaense was dake, 12 Aug. 1893, Y. Tokubuchi s.n. (SAPS 19804); Mt. Yubari-dake, 7 Aug. 1912, H. Yanagisawa & A. Hamana a ultrabasicosaxophyte, however, this species s.n. (SAPS 19795); Mt. Yubari-dake, an upper stream of was later reported from a non-serpentine area the River Epanaomandonashubetsu-gawa, a mire, 8 Aug. (Umezawa pers. comm.; Mt. Pin'neshiri-dake, 1912, H. Yanagisawa s.n. (SAPS 19800); Mt. Yubari-dake, cf. specimens examined). Mt. Mae-dake, alt. 1200 m, 26 July 1979, fl., Y. Kadota 6151 (TNS 905649–905650); Mt. Yubari-dake, 7 Aug. 1987, fl., Y. Kadota 14050 (TNS 643343–643345); Yubari- 7. Aconitum tatewakii Miyabe in Trans. shi, Mt. Ikushunbetsu-dake, 10 July 1931, T. Irinoda 12981 Sapporo Nat. Hist. Soc. 14(2): 71, fig. 5, a–b, (SAPS 22460). -shi, Garo-no-sawa, 3 June 1950, fl., (1935) – Lycoctonum tatewakii (Miyabe) Honda, M. Hemmi 1463 (TNS 116986, 141992). -shi, Nom. Pl. Jap. 97, 510 (1939) – Lycoctonum Mt. Ashibetsu-dake, 28–31 July 1915, H. Koidzumi s.n. (SAPS 22457); Mt. Kirigishi-yama, 28 June 1987, M. Hara gigas (H. Lév. & Vaniot) Nakai var. tatewakii 6868, 6869 (SAPS 19759, 19760); Mt. Kirigishi-yama, alt. (Miyabe) Nakai in Bull. Natn. Sci. Mus., Tokyo 900–1000 m, 26 July S45, M. Tohyama & al. s.n. (SAPS no. 32: 16 (1953). 19827). Kamikawa Subpref., Sprachi-gun, Minami- Type: JAPAN. Yezo [Hokkaido]. Prov. Furano-cho, Kanayama, along an upper stream of the River Tonashibetsu, alt. 400 m, 3 July 2007, fl., K. Horie s.n. Tokachi (Tokachi Subpref.), [Hidaka Mountains] (TNS 771080–771082). Yufutsu-gun, -mura, Mt. Satunai-dake, 27 July 1927, M. Tatewaki Nini-u, 22 June 2008, fl., H. Igarashi s.n. (TNS 774974). s.n. (SAPS–holotype !). Kamikawa-gun, Kamikawa-cho, Mts. Taisetsuzan, Mt. Aconitum gigas H. Lév. & Vaniot var. Kurodake, 27 July 1947, M. Tatewaki 35198 (SAPS 19806, taisetsuense Inagaki & Toyok. in J. Geobot. 19834); Kamikawa-cho, Sô-un-kyô, 13 June 1970, fl., T. Satow s.n. (TNS 280194); Kamikawa-cho, Aizan-kei, (Hokuriku) 10(3): 81, figs. 1–2 (1961). Type: alt. 1120 m, 26 July 1985, fl., K. Tsuchigura 1017 (TNS JAPAN. Hokkaido. Prov. Ishikari, Mts. Taisetsu- 695328); Kamikawa-cho, Mt. Niseikaushuppe, 9 Aug. February 2012 Journal of Japanese Botany Vol. 87 No.1 15

Fig. 7. Type of Aconitum umezawae Kadota (JAPAN. Hokkaido, Oshima Subpref., Futami-gun, Yakumo-cho, Mt. Oboko-dake, 21 June 2010, Y. Kadota 1006521, TNS 1028813, holotype). 16 植物研究雑誌 第 87 巻 第 1 号 2012 年 2 月

Fig. 8. Aconitum umezawae Kadota. A. Habit. B. Flower. C. Nectary. At Mt. Oboko-dake, Yakumo-cho, southwestern Hokkaido, Japan, on 21 June 2010. February 2012 Journal of Japanese Botany Vol. 87 No.1 17

1947, M. Tatewaki & J. Samejima 35406 (SAPS 19807, tinged with brownish violet (the Minami- 19829). Kato-gun, Kamishihoro-cho, Nukabira, fl., 22 June Sakkai population), 2.4–3.0 cm long, glabrous 1968, M. Kikuchi s.n. (TNS 222787); Kamishihoro-cho, Mt. Yuni-Ishikari-dake, 25 Aug. 1953, M. Tatewaki 42566 on the abaxial side; helmet cylindrical, 26–30 (SAPA 19832); Kamishihoro-cho, Mt. Nishi-Kumaneshi- mm long, 11–12 mm wide, 17–18 mm high, dake, 27 July 1985, fl. & y. fr., Y. Kadota 12325, 12340 rounded at apex, with short beak 1 mm long, the (TNS 905646–905647). line between the base and the beak oblique (ca. 45º); lateral sepals distorted obovate, 12–13 mm 8. Aconitum umezawae Kadota, sp. nov. long, 10–11 mm wide, hirsute with long, rough- [Figs. 7–8] surfaced, flexuous hairs (pollen-collecting hairs) Differt ab Aconito mashikensi caricalibus at the aperture on the adaxial side. Nectaries nectariorum brevioribus, labiis nectariorum glabrous, yellowish-white; blade 3 mm long, 2 rotundatis et minute irregulariter serratis, floris mm wide, not inflated and tapering to the spur, eborinis, folliculis parvis et bracteolis positis tubular part 1 mm long; spur thick and short, super media pedicellorum. 4–5 mm long, incurved; labium 1 mm long, Type: JAPAN. Hokkaido. Oshima Subpref., emarginate and irregularly and minutely serrate; Futami-gun, Yakumo-cho, Mt. Oboko-dake, alt. claw 9–10 cm mm long, nearly straight. Stamens 296 m, 21 June 2010, Y. Kadota 1006521 (TNS glabrous, anthers 0.5 mm long, filaments 5–7 1028813–holotype; Fig. 7). mm long, staminal teeth absent or present. A subscapose, perennial herb, 1–1.2 m tall. Carpels 3, glabrous. Follicles 7–10 mm long, Root 1–2 cm in diameter at the ground surface divergent; seeds black, 3 mm long, 2 mm wide. level, branched, rhizomatic. Stem robust, terete, Japanese name: Oshima-reijinsô (nom. nov.). erect to declining, branched from the middle part, 新和名：オシマレイジンソウ sparingly villous with rough-surfaced, spreading Distribution: Southwestern Hokkaido, hairs in the distal part. Basal leaves persistent northern Japan (Fig. 9, disc). Endemic to Japan. at anthesis. Blades of basal leaves roundish Additional specimens examined: JAPAN. Hokkaido. reniforme in outline, 20–39 cm long, 23–39 Oshima Subpref., -shi, Kakkumi-cho, Ôfuna cm wide, medially 9–11-lobed to 4–5 cm from forest road, alt. ca. 100 m, 27 June 2010, M. Sakai s.n. the base, densely sericeous with straight, long, (TNS 1108750–1108753). Kameda-gun, Nanae-cho, golden, rough-surfaced, adpressed hairs along Narukawa, alt. 440 m, 21 June 2010, Y. Kadota 1006502– 1006503 (TNS 1028892–1028897). Yamakoshi-gun, the veins on the abaxial side, deeply cordate Yakumo-cho, Mt. Oboko-dake, Ginzan-zawa – Oboko- at base; middle lobe obovate-rhombic, 13–15 zawa – Kanega-sawa, alt. 250–780 m, 22 July 1987, fr., cm long, 10–13 cm wide, acute, dentate; tooth H. Takahashi & al. 7223 (SAPS 19769); Mt. Oboko-dake, narrowly ovate, 5–7(–12) mm wide, acuminate; alt. 300 m, 6 Sept. 2009, fr., S. Umezawa 09090601– petioles 37–46 cm long, glabrous, 4-angled, 09090602 (TNS 741145, 741149); Mt. Oboko-dake, alt. 296 m, 21 June 2010, Y. Kadota 1006511–1006513 (TNS hollow. Cauline leaves similar to basal in shape, 1028821–1028832). Shiribeshi Subpref., Shimamaki- similarly pubescent to the basal, diminishing in gun, Shimamaki-mura, Mt. Obira-yama [Ôhira-yama], size, petiolate to sessile. Inflorescence terminal, alt. 740 m, 6 Aug. 2009, fr., Y. Kadota 093407 (TNS racemose, 33–47 cm long, ca. 20–40-flowered, 739538–739539). Isoya-gun, Rankoshi-cho, Mt. Horomui- dake, Tsubame-no-sawa gorge, alt. 386 m, 26 June 2009, in indeterminate condition. Pedicel of the fl., Y. Kadota 093167–093169 (TNS 1101945–1101950). lowermost flower 4.5–10 cm long, arched to Suttsu-gun, Kuromatsunai-cho, Minami-Sakkai, alt. 36 incurved, villous with rough-surfaced, spreading m, 25 June 2009, fl., Y. Kadota 093142–093147 (TNS hairs throughout, devoid of glandular hairs; 1101870–1101880). Shakon-gun, Shakotan-cho, Bikuni, bracteoles 2, linear, 8–17 mm long, 1.5–2 mm Kawakami, alt. 145 m, 22 June 2010, Y. Kadota 1006561– 1006567 (TNS 1028863–1028875). Yoichi-gun, Yoichi- wide, situated above the middle of the pedicel. cho, Yoichi dam, alt. 219 m, 22 June 2010, Y. Kadota Flowers in June, yellowish white, sometimes 1006577–1006578 (TNS 1028857–1028860). 18 植物研究雑誌 第 87 巻 第 1 号 2012 年 2 月

Fig. 9. Distribution of four Aconitum species. Triangle. A. asahikawaense. Diamond. A. hiroshi-igarashii. Star. A. ikedae. Disc. A. umezawae.

Etymology: The specific epithet is dedicated to Mr. Shun Umezawa, who is the discoverer of Key to the species of subgenus Lycoctonum this new Aconitum species. in Hokkaido Note: Aconitum umezawae is discriminated 1A. Flowers pale blue-violet from A. mashikense by having yellowish-white 2A. Pedicels strigose with curved hairs, flowers, bracteoles positioned above the middle nectary labia entire ... 3. A. hiroshi-igarashii of the pedicels, thick and short nectary spurs, 2B. Pedicels villous with spreading hairs and roundish, irregularly and minutely toothed glandular hairs, nectary labia irregularly nectary labia and short follicles. serrate ...... 4. A. ikedae At the Ôfuna population, Hakodate, 1B. Flowers yellow to yellowish-white southernmost Hokkaido, putative hybrids 3A. Pedicels villous with spreading hairs, between A. umezawae and A. gigas were leaves sericeous with long, adpressed hairs collected (H. Igarashi s.n., TNS 1108754– along the veins beneath 1108759) similar to the Mt. Obira-yama 4A. Flowers yellow, bracteoles positioned population. below the middle of the pedicels, nectary February 2012 Journal of Japanese Botany Vol. 87 No.1 19

spurs thick and long, nectary labia obovate 81–83. and slightly emarginate, follicles 1–2 cm Kadota Y. 2001. Systematic studies of Asian Aconitum (Ranunculaceae) VII. A new species and a new form long ...... … 5. A. mashikense of subgenus Lycoctonum from Hokkaido, Japan. J. Jpn. 4B. Flowers yellowish-white, bracteoles Bot. 76(1): 20–27. positioned above the middle of the Kadota Y. 2006a. Aconitum L. In: Iwatsuki K., Boufford pedicels, nectary spurs thick and short, D. E. and Ohba H. (eds.), Flora of Japan IIa: 267–285. nectary labia roundish and irregularly Kodansha Ltd., Tokyo. Kadota Y. 2006b. Ezo-Reijinsô-to-Ô-Reijinsô. Hoppô- serrate, follicles 1 cm long ...... Sansô no. 23: 105–114 (in Japanese)...... 8. A. umezawae Kadota Y. 2007. Systematic studies of Asian Aconitum 3B. Pedicels strigose with curved hairs (Ranunculaceae) XII. Aconitum soyaense, a new 5A. Flowers smaller, 2.5–3 cm long, nectary species of subgenus Lycoctonum from Hokkaido, northern Japan. J. Jpn. Bot. 82(1): 41–44. spurs 2–4 mm long, incurved to 180 Kadota Y. 2008. Aconitum L. In: Ohashi H., Murata J. and degrees or straight, follicles 1–2 cm long, Iwatsuki K. (eds.), New Makino's Illustrated Flora erect to ascending of Japan. pp. 135–141. The Hokuryukan, Tokyo (in 6A. Sepals with purplish-brown spots Japanese). at apex, carpels strigose with curved Kadota Y. 2010. Systematic studies of Asian Aconitum (Ranunculaceae) XIII. Species of Bhutan including a hairs or sometimes glabrous, bracteoles new scandent species. J. Jpn. Bot. 85(1): 8–24. positioned around the base of the Kadota Y. 2011. Aconitum L. In: Kato M. and Ebihara A. pedicels ...... 1. A. asahikawaense (eds.), Endemic Plants of Japan. A Book Series from 6B. Sepals devoid of purplish-brown the National Museum of Nature and Science No. 11. pp. 55–57. Tokai University Press, Hadano (in spots, carpels glabrous or rarely Japanese). sparingly strigose with curved hairs, Lauener L. A. and Tamura M. 1978. A synopsis of bracteoles positioned around the middle Aconitum Subgenus Paraconitum: I. Notes Roy. Bot. of the pedicels ...... 2. A. gigas Gard. Edinb. 37(1): 113–124. 5B. Flowers larger, 2–2.5 cm long, nectary Miyabe K. and Tatewaki M. 1935. Contributions to the flora of northern Japan VI. Trans. Sapporo Nat. Hist. spurs saccate, follicles ca. 1 cm long, Soc. 14(2): 69–86. ascending Nakai T. 1953. A new classification of Lycoctonum and 7A. Sepals shallowly cylindrical, obtuse Aconitum in Korea, Japan, and their surronding area. at apex, carpels hirsute with ascending Bull. Natn. Sci. Mus., Tokyo (32): 1–53. Sugimoto J. 1978. Aconitum L. and Lycotonum Fourrier. hairs ...... 6. A. soyaense Keys to Herbaceous Plants of Japan. I. Dicotyledonae. 7B. Sepals pyramidal to cylindrical, acute Revised and enlarged. pp. 134–140, 154–155. Inoue at apex, carpels glabrous 7. A. tatewakii Book Co., Tokyo (in Japanese). Tamura M. 1980. Aconitum L. In: Kitamura s. and Murata G., Coloured Illustrations of Herbaceous Plants of I wish to show my cordial thanks to Japan (Choripetalae). II: 207–213. Hoikusha, Osaka Messrs. Hiroshi Igarashi, Ikuro Ikeda and Shun (in Japanese). Umezawa and Dr. Kenji Horie for their guidance Tamura M. 1982. Aconitum L. In: Satake Y., Ohwi J., to the localities and their gifts of the Aconitum Kitamura S., Watari S. and Tominari T. (eds.), Wild species; to Prof. Dr. Hideki Takahashi for his Flowers of Japan. Herbaceous Plants (Including Dwarf Subshrubs). II: 61–67. Heibonsha, Tokyo (in Japanese). permission for specimen examination at the Tamura M. 1992. Aconitum L. In: Ohwi J. (Kitagawa M.), Herbarium of the Hokkaido University Museum New Flora of Japan Revised. pp. 705–715. Shibundo, (SAPS). Tokyo (in Japanese). Tamura M. 1995. Aconitum L. Die Natürichen Pflanzenfamilien Band 17aIV: 274–291. Duncker & References Humblot, Berlin. Inagaki K. and Toyokuni H. 1961. Symbolae Tamura M. and Lauener L. A. 1979. A synopsis of Taisetsusanenses (1). J. Geobot. (Hokuriku) 10(3): Aconitum Subgenus Lycoctonum: II. Notes Roy. Bot. 20 植物研究雑誌 第 87 巻 第 1 号 2012 年 2 月

Gard. Edinb. 37(3): 431–466. Umezawa S. 2009. Aconitum L. Alpine Flowers of Toyokuni H. 1955. On the ultrabasicosaxicolous flora of Hokkaido. pp. 230–241. The Hokkaido Shimbun Press, Hokkaido, Japan. (1). Hokuriku J. Bot. 4(4): 97–101. Sapporo (in Japanese).

門田裕一：アジア産トリカブト属植物（キンポウゲ科） の分類学的研究 XIV．北海道産レイジンソウ亜属の 4 新 種 北海道からトリカブト属レイジンソウ亜属レイ オシマレイジンソウは黄花のグループの 1 種で，花 ジンソウ節オオレイジンソウ列 Aconitum subgenus 梗に滑面開出毛が生える．花色と花梗の有毛性では， Lycoctonum sect. Lycoctonum ser. Umbrosa (Steinb. ex 増毛山地と樺戸山地に分布するマシケレイジンソウ A. Nakai) Kadota の 4 新種を記載した．それらは，青花の mashikense Kadota & Umezawa に似るが，花色が黄白 グループとしてコンブレイジンソウ Aconitum hiroshi- 色で，小苞は花梗の中部以上に付き，花弁の距が太く短 igarashii Kadota と ニ セ コ レ イ ジ ン ソ ウ A. ikedae く，花弁の唇部が円形で縁が不規則に切れ込む点で区別 Kadota の 2 種，黄花のグループとしてオシマレイジ される．本種は渡島半島から積丹半島にかけて分布す ンソウ A. umezawae Kadota とカムイレイジンソウ A. る．ニセコ地域から，札幌の西方に位置する余市山地に asahikawaense Kadota の 2 種である．いずれも北海道 かけての山域では，本種とエゾレイジンソウの自然雑種 の固有種で，夏緑林の林縁に生育する． と推定される個体が多数得られている． 青花のコンブレイジンソウとニセコレイジンソウは カムイレイジンソウも黄花の種群の一つで，花梗に粗 円筒形の上萼片（かぶと）をもち，嘴が短く，花弁の距 面屈毛があり，各萼片の先端部に紫褐色の斑紋があり， が太くかつ短いことで特徴付けられる．同様に淡い赤 上萼片の嘴が長く水平に突き出し，心皮に屈毛があるこ 紫色を帯びた青花を付けるレイジンソウ A. loczyanum とで特徴付けられる．本種は旭川市と周辺の蛇紋岩地域 Rapaics（ 四 国， 九 州 ） と ア ズ マ レ イ ジ ン ソ ウ A. にのみ生育し，萼片に見られる紫褐色の斑紋は蛇紋岩変 pterocaule Koidz.（本州）は先端がすぼまる狭円錐形 形の一例とみなすことができる． の上萼片をもち，嘴が長く，花弁の距が細くかつ長いも エゾレイジンソウのうち道東の集団（特に阿寒湖周辺 ので，レイジンソウ亜属レイジンソウ列 ser. Volubilia 地域）には花弁の距の長さに変異があり，中には距が (Steinb. ex Nakai) Tamura & Lauener に所属する別の 長いものがある．このような個体はこれまでしばしば 一群である． 本州産のオオレイジンソウ A. umbrosum (Korsh.) Kom. ニセコレイジンソウは花梗の全体に渡って滑面開出 （Kadota 2006a，門田 2006b）に当てられてきた．しか 毛が生え，先端部に滑面腺毛が生える．蘭越町（ニセコ しながら，エゾレイジンソウとオオレイジンソウの 2 山系西部の南面）とニセコ町（昆布岳の北面）及び豊浦 種は距の長さの違いのみで認識できるのではない．オオ 町（礼文華）に分布する． レイジンソウは花弁舷部の筒状部分が長く，かつ唇部も コンブレイジンソウはニセコレイジンソウに似るが， 長い点でエゾレイジンソウと異なっている．したがっ 花梗には粗面屈毛が生える点で区別できる．分布域はニ て，道東の距の長いものはエゾレイジンソウに含めるべ セコレイジンソウより少し広く，蘭越町，ニセコ町，豊 きであり，本州のオオレイジンソウとは異なる．なお， 浦町の他，渡島半島の七飯町にも分布する． オオレイジンソウの学名については検討の余地がある 北をニセコ山系に，南を昆布岳山系に挟まれた地域 ことを付記する． （ニセコ地域）では上記 2 種の雑種が多数得られてい こ れ ま で，Aconitum tatewakii Miyabe は Aconitum る．ハゴロモレイジンソウ A. gigas H. Lév. & Vaniot f. gigas H. Lév. & Vaniot の異名として扱われることが普 bicolor Kadota & Umezawa は，エゾレイジンソウ A. 通であった（例えば，Kadota 2006a など）．しかしながら， gigas の色変わりの品種として，蘭越町コックリ湖湖畔 A. tatewakii は長さ 2 cm ほどの小型の花を付け，花弁 から記載された (Kadota 2001)．ハゴロモレイジンソウ の距が嚢状となるもので，分布域も基準産地である日 のタイプ標本を改めて検討したところ，この個体はニセ 高山脈のほか，夕張山地と石狩山地に限られ，A. gigas コレイジンソウとコンブレイジンソウとの自然雑種で とは異なる実在の分類群と認められる．A. tatewakii の あることが分かった．また，実際に蘭越町の集団を調べ 和名は Miyabe (1935) により，エゾレイジンソウとさ た結果，上記 2 種の自然雑種とニセコレイジンソウか れた．「エゾレイジンソウ」という和名はこれが初出で ら成り，コンブレイジンソウに相当する個体は見出せな ある．しかし，現行の国内のほとんど全ての出版物で かった． は A. gigas にエゾレイジンソウ（あるいはエゾノレイ February 2012 Journal of Japanese Botany Vol. 87 No.1 21

ジンソウ）が用いられている．そこで，混乱を避けるた 規則な波状縁となる，袋果は長さ 1 cm ...... めに，エゾレイジンソウの和名は A. gigas に残し，A...... 8. オシマレイジンソウ tatewakii の和名には，本種が日高山脈から最初に認識 （函館市，七飯町， されたことと同山脈では実際に全域で本種が見られる 八雲町，島牧村，蘭越町，黒松内町，積丹町，余市町） ことから，新和名としてヒダカレイジンソウを用いるこ 3B. 花梗に屈毛が生える とを提唱したい． 5A. 花はより大型で長さ 2.5– 3 cm，花弁の距は 以上の結果，ここで記載した 4 新種の他に，エゾレ 長さ 2–4 mm，湾曲して約 180° まで屈曲す イジンソウ，ソウヤレイジンソウ A. soyaense Kadota， るか，あるいは直線的に伸びる，袋果は長さ ヒダカレイジンソウ，マシケレイジンソウの合計 8 種 1–2 cm，直立〜斜開する が北海道に分布することが明らかになった．これらの 8 6A. 各萼片の先端部に紫褐色の斑紋がある，心 種は次のように区別できる： 皮に屈毛が生えるが時に無毛，小苞は花梗 の基部につく ...... 1. カムイレイジンソウ 1A. 花は淡青紫色 （旭川市とその周辺の蛇紋岩地域） 2A. 花梗に屈毛が生える，花弁の唇部は全縁...... 6B. 萼片に紫褐色の斑紋はない，心皮は無毛， ...... 3. コンブレイジンソウ 稀に屈毛が散生し，小苞は花梗の中部につ （七飯町，蘭越町，ニセコ町，豊浦町） く ...... 2. エゾレイジンソウ 2B. 花梗に開出毛と腺毛が生える，花弁の唇部は不規 （ニセコ地域〜余市山地，道北，道東） 則な波状縁となる...... 4. ニセコレイジンソウ 5B. 花はより小型で長さ 2−2.5 cm，花弁の距は嚢 （蘭越町，ニセコ町，豊浦町） 状， 袋果は長さ約 1 cm，斜開する 1B. 花は黄色〜黄白色 7A. 上萼片は浅い円筒状，頂部は円頭，心皮に 3A. 花梗に開出毛が生えるが腺毛を欠く，葉の下面 斜上毛が密生する.....6. ソウヤレイジンソウ は 脈に沿って長い伏毛がある （道北の蛇紋岩地域） 4A. 花は黄色，小苞は花梗の中部以下につく，花 7B. 上萼片は三角状〜円筒状，頂部は鋭頭，心 弁の距は太く長い，花弁の唇部は倒卵形，先 皮は無毛...... 7. ヒダカレイジンソウ 端はわずかに凹む，袋果は長さ 1–2 cm ...... （日高，夕張，石狩山系） ...... 5. マシケレイジンソウ（増毛，樺戸山系） 4B. 花は黄白色，小苞は花梗の中部以上につく， （国立科学博物館植物研究部） 花弁の距は太く短い，花弁の唇部は円形，不