J. Jpn. Bot. 90: 179–191 (2015)

Variation in sibiricum ()

Hiroyoshi Ohashi

Herbarium TUS, Botanical Garden, Tohoku University, Sendai, 980-0862 JAPAN E-mail: [email protected]

(Accepted on 9 January 2015)

Morphological variation of (L.) Pennell and V. japonicum (Nakai) T. Yamaz. with their infraspecific taxa were examined in herbarium specimens. Morphological differences used for delimitation of these taxa fall within a continuous range of variation. Veronicastrum sibiricum is recognized here as a single polymorphic species composed of five forms including two new combinations: f. glabratum (Nakai) H. Ohashi and f. humile (Nakai) H. Ohashi. Veronicastrum sibiricum var. yezoense, V. sibiricum var. zuccarinii, Veronicastrum japonicum, and V. japonicum var. australe were indistinct from V. sibiricum var. sibiricum. A bibliographic enumeration is presented for each name. Each lectotype was designated for sibirica var. glabra Nakai, Veronica sibirica var. japonica f. latifolia Nakai, and Veronica sachalinensis T. Yamaz.

Key words: Lectotypification, morphological variation, Plantaginaceae, polymorphic species, Veronica, Veronicastrum japonicum, Veronicastrum sibiricum.

Veronicastrum sibiricum (L.) Pennell This paper aims to reevaluate taxonomic status (Plantaginaceae) is known in Japan, China, of V. sibiricum and its related taxa based on Korea, E. Siberia and Far Eastern Russia. The examination of herbarium specimens. species shows a wide range of morphological variation mainly in degree of hairiness on the Taxonomic history of Veronicastrum rachis of inflorescences and the surfaces of sibiricum in Japan , length of the pedicel or relative length Thunberg (1784) recorded Veronica virginica between the pedicel and the calyx, shape of the L. from Nagasaki, Osaka and others in Japan apex of corolla-lobes, and number of leaves with Japanese names “Iaponice: Ireisin, vulgo at a node of the stem. The species had been Kwugai So; it. Sobi, vulgo Toranoo”. This is the divided into many taxa by Japanese taxonomists first record of the in botany, although it had (Nakai 1912, 1943, Koidzumi 1930, Hara 1940, been recorded by Terazima-Ryôan as “Kukaisô” 1949, Yamazaki 1952, 1957, 1982, 1990, 1993, in “Wakan-Sansai-Zue 46” with an illustration Yonekura 2012), but these taxa had not been published in 1712 (Nakai 1943). Rafinesque adopted by Ohwi (1953, 1965a, 1965b) and (1836) regarded the Japanese Veronica virginica Ohwi and Kitagawa (1992). Also, the species as different from North American species and was recognized as a single species without named it Eustaxia japonica Raf. Steudel (1841) infraspecific taxa in floras of China (Chin and referred it to Veronica japonica (Rafn.) Steudel. Hong 1979, Hong et al. 1998), Korea (W. T. Nakai (1912) treated Veronica virginica L. Lee 1996, Kim and Choi 2007) and Russia. conspecific with V. sibirica L. and V. virginica

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Fig. 1. Veronicastrum sibiricum from Mt. Daisen, Tottori Pref. (A. Kimura s.n. 10 Sep. 1925. TUS 230541). June 2015 The Journal of Japanese Botany Vol. 90 No. 3 181 sensu Thunb. He separated Veronica virginica pedicel as long as or longer than the calyx. Hara into three taxa: the typical taxon [var. virginica] (1940) recognized Veronicastrum sibiricum in N. America, var. sibirica (L.) Nakai in China, [var. sibiricum] as it has glabrous rachides of Far East Russia (“Dahuria, Amur, and Sachalin”) inflorescences and scarcely pubescent leaves. and Hokkaido, and var. japonica Nakai in Japan He recognized var. japonicum (Nakai) H. Hara (Honshu, Shikoku and southern Hokkaido). as a local variety of var. sibiricum and described Nakai (1912) named “Kukaisô” (= “Kwugai it having slightly longer pedicels, pubescent So” in Thunberg) var. japonica, but treated V. inflorescence-rachides and almost glabrous virginica sensu Thunb. as a synonym of var. leaves. Collectively Yamazaki (1993) separated sibirica. Nakai’s treatment is dubious, because V. japonicum from V. sibiricum by length of the V. virginica sensu Thunb. was recorded from pedicel: 1.5–3 mm long at anthesis in the former, “Jeso”, not Honshu as “Nippon tota” in the sense whereas 0–1 mm long in the latter. of Nakai. He overlooked Veronica japonica The pedicel of V. sibiricum var. zuccarinii (Rafn.) Steudel for a synonym of var. japonica is described as 1.5 mm long (Yamazaki 1993); Nakai. Hara (1940) adopted Veronicastrum hence, the pedicel-length is continuous between sibiricum for the Japanese species and divided it V. sibiricum and V. japonicum. In fact, V. into three varieties: V. sibiricum var. japonicum japonicum has pedicels (1–)1.5–3 mm long, (Nakai) H. Hara, var. zuccarinii (Koidz.) H. whereas V. sibiricum has pedicels 0–1.5(–2) mm Hara, and var. yezoense H. Hara. He corrected long in var. yezoense. Moreover, the pedicels Veronica virginica sensu Thunb. and Veronica elongate during and after anthesis. The pedicel- japonica (Rafn.) Steudel to be synonymous with length is not effective as a diagnostic character Veronicastrum sibiricum var. japonicum (Nakai) between V. japonicum and V. sibiricum. H. Hara. Nakai (1943) treated again Veronica Veronicastrum sibiricum has glabrous sibirica and split it into 10 taxa. Yamazaki inflorescence-rachides (Hara 1940). Density (1952, 1957, 1981, 1990, 1993) had eagerly of hairiness on the inflorescence-rachides in V. studied these taxa under Veronicastrum. He japonicum varies continuously from scarcely recognized the problematic “Kwugai So” (= V. puberulent (e.g., Miyagi Pref. Sasaki 93-0546- virginica sensu Thunb.) complex as it includes 1 & -2. TUS 329632, TUS 329628) to densely two species with each two varieties and two pubescent. A specimen referred to V. japonicum forms: Veronicastrum japonicum (Nakai) T. from Mt. Daisen, Tottori Pref. (A. Kimura s.n. Yamaz. with [var. japonicum], [f. japonicum], 10 Sep. 1925. TUS 230541), has the almost f. album (Sugim.) T. Yamaz., var. australe (T. glabrous central rachis and puberulent lateral Yamaz.) T. Yamaz., [f. australe (T. Yamaz.) T. rachides (Fig. 1 right plant). Yamaz.], f. albiflorum (Akasawa) T. Yamaz., and Yamazaki (1993) adopted number of leaves var. humile (Nakai) T. Yamaz., and V. sibiricum per node for another diagnostic character (L.) Pennell [var. sibiricum], var. yezoense H. between V. japonicum and V. sibiricum. He Hara var. zuccarinii (Koidz.) H. Hara (Yamazaki noted 5 or 6-verticillate in V. japonicum against 1993). 7- or 8-verticillate in V. sibiricum in the key, but actually in the description of each species Veronicastrum sibiricum and V. japonicum he described (5–)7–8(–10) in var. yezoense and Nakai (1912) characterized Veronica (4–)5–6(–7) in var. zuccarinii of V. sibiricum, virginica var. sibirica (L.) Nakai as it has whereas (4–)5–6(–8) in V. japonicum. A sessile or subsessile flowers with the pedicel recent description of V. sibiricum published shorter than the calyx, whereas var. japonica in China (Chin and Hong 1979, Hong et al. Nakai as it has pedicellate flowers with the 1998) indicates that the leaves in whorls of 182 植物研究雑誌 第 90 巻 第 3 号 2015 年 6 月

Fig. 2. Holotype of Veronica sibirica var. yezoensis f. glabrata Nakai (Hokkaido. Prov. Ishikari, Sapporo. 7 Jul. 1891. Tokubuti s.n. TI). June 2015 The Journal of Japanese Botany Vol. 90 No. 3 183

Fig. 3. Holotype of Veronica sibirica var. humilis Nakai (Japan. Honshu. Prov. Ômi, in summa montis Ibukiyama. 1 Aug. 1881. Ryôkichi Yatabe s.n. TI). 184 植物研究雑誌 第 90 巻 第 3 号 2015 年 6 月

4–6, glabrous or sparsely hirsute and with Veronica sibirica var. humilis Nakai (Fig. multicellular hairs on both surfaces. This 3) was described in 1943. It has smaller stems Chinese form may be referable to V. japonicum (30–60 cm tall) including inflorescences 7–10 in the sense of Yamazaki (1993). cm long and leaves that are oblong or narrowly Yamazaki (1993) added other distinctions elliptic with acute or acuminate apex (4–6 cm that the apex of corolla-lobes is acute in V. long, 1.5–3 cm wide) with sparcely puberulent sibiricum var. sibiricum, obtuse in var. yezoense, or sometimes glabrous on both surfaces and subacute in V. japonicum. The apex is (Yamazakai 1993). The rachis of inflorescences usually acute in these taxa and shape of the is glabrous or pubescent. The pedicels are ca. corolla-lobes is useless for a diagnostic character. 2 mm long in anthesis and 2.5–3 mm long in Veronicastrum japonicum is apparently fruit. It has been recorded as endemic to the conspecific withV. sibiricum. summit of Mt. Ibuki in Shiga Prefecture (Nakai 1943, Yamazaki 1993). This form is found in Infraspecific taxa of Veronicastrum Mts. Higashi-akaishi in Besshiyama-son, Ehime sibiricum and V. japonicum Prefecture and Mt. Tsurugi in Higashi-iyayama- Hara (1940) described Veronicastrum son, Tokushima Prefecture, Shikoku (e.g., M. sibiricum var. yezoense from Hokkaido and Yamanaka 926. TUS 10321. Fig. 4). Veronica northern Honshu as a densely pubescent form sibirica var. humilis appears to be a form of V. sibiricum in having pubescent leaves confined in limestone or olivine areas in these and inflorescence-rachides. Nakai (1943) habitats. characterized Veronica sibirica with glabrous Veronicastrum japonicum var. australe (T. adaxial surface of leaves and var. glabra Nakai Yamaz.) T. Yamaz. (Fig. 5) is characterized with glabrous abaxial -surface and “var. in having glabrous inflorescence-rachides typica” [= var. sibirica] hirsute at least on nerves. (Yamazaki 1993), but is identical with var. He divided Veronica sibirica var. yezoensis (H. sibiricum. Hara) Nakai into three forms: f. glabrata Nakai Veronica sibirica var. humilis and (Fig. 2) with glabrous abaxial leaf-surfaces Veronicastrum japonicum var. australe are except the pubescent midrib, f. pubescens Nakai apparently continuous through intermediate with entirely appressed pubescent abaxial leaf- forms in Shikoku. Such specimens as, for surfaces, and f. latifolia with appressed pilose example, Ehime Pref., Mt. Gongen. 10 Aug. abaxial leaf-surfaces. Except in V. sibirica 1959. M. Yamanaka s.n. (TUS 10255), Mt. var. glabra the abaxial leaf-surface varies Higashi-akaishi. 9 Aug. 1950. M. Yamanaka continuously in hairiness between var. sibirica s.n. (TUS 54470), and Gongen-o in Mt. and var. yezoensis. Veronicastrum sibiricum var. Higashiakaishi-yama, Sekikawa-mura. 9 Aug. yezoense is distinguishable from var. sibiricum 1950. T. Yamanaka 5463 (TUS 409752) have only by the hairy inflorescence-rachis. longer inflorescences than those of typical var. Veronicastrum sibiricum var. zuccarinii humile and narrowly elliptic leaves, but (Koidz.) H. Hara is known in Kyushu and are smaller (3–40 cm tall) with small leaves southern Korea (Yamazaki 1993). It was 5–7 cm long on the middle stem. A similar distinguished from V. sibiricum var. sibiricum form intermediate between var. humile and var. in having densely pubescent adaxial surface japonicum was collected in Honshu, Tottori of leaves, because var. sibiricum has leaves Pref., Mt. Daisen (A. Kimura s.n. 10 Sep. 1925. uniformly to sparsely pubescent on the adaxial TUS 230541) (Fig. 1, the left plant). This is surface. Var. zuccarinii is indistinguishable from about 55 cm tall with an inflorescence 7.5 cm var. sibiricum. long and a puberulent rachis of inflorescence. June 2015 The Journal of Japanese Botany Vol. 90 No. 3 185

Fig. 4. Veronicastrum sibiricum f. humile (Nakai) H. Ohashi, new to Shikoku (Ehime Pref.: Mt. Gongen- yama. 30 Jul. 1952. M. Yamanaka 926. TUS 10321). 186 植物研究雑誌 第 90 巻 第 3 号 2015 年 6 月

Fig. 5. Holotype of Veronicastrum sibiricum var. australe T. Yamaz. (Japan. Shikoku. Tokushima Pref., Mt. Tsurugisan, 1800 m alt. 29 Jul. 1949. T. Akasawa s.n. TI). June 2015 The Journal of Japanese Botany Vol. 90 No. 3 187

It is regarded here as a form of Veronicastrum Jun. 1933. Akagi-Tuyosi s.n. TI. “Typus florum” sibiricum. designated by Nakai is selected here as lectotype for the name]. – V. sibirica L. f. glabra (Nakai) Key to the forms of Veronicastrum Kitag., Neolin. Fl. Manshur.: 574 (1979). sibiricum Veronica virginica L. var. zuccarinii Koidz. in Veronicastrum sibiricum is recognized here Bot. Mag. (Tokyo) 44: 112 (1930). – V. sibirica as a single species with four forms. They are var. zuccarinii (Koidz.) Nakai f. zuccarinii divided as in the following key: (Koidz.) Nakai in J. Jap. Bot. 19: 10 (1943), ut “f. 1. Plants 40–70 cm tall; usual leaves less than typica Nakai” [Lectotype (designated by Nakai 6 cm long, elliptic or ovate, rarely narrowly 1943, Yamazaki 1952): JAPAN. “Kyushu. Prov. elliptic, apex acute to acuminate ..... f. humile Hizen: in promontorio Nomosaki. Pierot. Hb. 1. Plants (60–)80–150 cm tall; usual leaves more Lugd. Batavi” (fide Nakai), “No. 908.233-460- than 6 cm long, narrowly elliptic or narrowly 412” (fide Yamazaki). L]; Makino & Nemoto, ovate, apex acuminate to caudate ...... 2 Nippon Shokubutsu Sôran (Fl. Jap.) ed. 2, 1079 2. Inflorescence-rachides glabrous...... 3 (1931). – Veronicastrum sibiricum var. zuccarinii 2. Inflorescence-rachides pubescent...... 4 (Koidz.) H. Hara in J. Jap. Bot. 16: 160 (1940), 3. Corolla purple ...... f. sibiricum & Enum. Spermat. Jap. 1: 282 (1949); T. Yamaz. 3. Corolla white ...... f. albiflorum in J. Jap. Bot. 27: 64 (1952), & in J. Fac. Sci. 4. Corolla purple ...... f. glabratum Univ. Tokyo sect. 3, Bot., 7(2): 132 (1957), & 4. Corolla white ...... f. album in Satake & al., Wild Flowers Jap. Herb. 3: 107 (1981), & in K. Iwats. & al., Fl. Jap. 3a: 346 Veronicastrum sibiricum (L.) Pennell, Acad. (1993); Yonek. in J. Jap. Bot. 65: 241 (1990). – Nat. Sci. Philadelphia Monogr. 1: 321, in obs. Veronica sibirica var. zuccarinii (Koidz.) Nakai (1935); H. Hara in J. Jap. Bot. 16: 159 (1940), & in J. Jap. Bot. 19: 9 (1943). Enum. Spermat. Jap. 1: 281 (1949); T. Yamaz. in Veronica sibirica var. zuccarinii f. J. Fac. Sci. Univ. Tokyo sect. 3, Bot., 7(2): 132 angustifolia Nakai in J. Jap. Bot. 19: 10 (1943) (1957); Ohwi, Fl. Jap. ed. Engl.: 803 (1965), & [Holotype: COREA (Tyôsen). Prov. Kôkai, ed. rev.: 1203 (1965); T. L. Chin & D. Y. Hong Kumiho. R. G. Mills 4360. TI]. in Fl. Reipubl. Popularis Sin. 67(2): 248, f. 69 Veronicastrum sibiricum var. australe T. (1979); Ivanina in Kharkev., Pl. Vasc. Or. Extr. Yamaz. in J. Jap. Bot. 27: 64 (1952) [Holotype: Soviet. 5: 308 (1991); Ohwi & Kitag., New Fl. JAPAN. Shikoku. Tokushima Pref., Mt. Jap. ed. rev.: 1342 (1992); W. T. Lee, Lineam. Tsurugisan, 1800 m alt. 29 Jul. 1949. T. Fl. Kor. 1: 1017 (1996), & Standard Illust. Kor. Akasawa s.n. TI], & in J. Fac. Sci. Univ. Tokyo Pl.: 325, fig. 1949 (1996); D. Y. Hong & al., sect. 3, Bot., 7(2): 132 (1957). – Veronicastrum Fl. China 18: 61 (1998). – Veronica sibirica L., japonicum (Nakai) T. Yamaz. var. australe (T. Sp. Pl. ed. 2, 2: 12 (1762) [Type not designated. Yamaz.) T. Yamaz. in J. Jap. Bot. 65: 242 (1990) Original material: Herb. Linn. No. 26.1 (LINN). & in K. Iwats. & al., Fl. Jap. 3a: 346 (1993). Jarvis, Order out of Chaos: 920. 2007]; Nakai Nom. Jap.: Shiberia Kugaisô シベリアクガイソ in J. Jap. Bot. 19: 5 (1943); Ohwi, Fl. Jap.: 1043 ウ（ツクシクガイソウ，ナンゴククガイソウ）． (1953). – Veronica virginica L. var. sibirica (L.) Distr.: Japan (Honshu, Shikoku, and Nakai in Bot. Mag. (Tokyo) 26: 170 (1912). Kyushu), Korea, China, Mongolia, and Russia f. sibiricum (E. Siberia, Dahuria, Amur, Ussuri). Veronica sibirica var. glabra Nakai in J. Jap. Bot. 19: 5 (1943) [Lectotype (designated here): f. albiflorum Akasawa CHINA (Manchoukuo). Prov. Chilin: Pâtaokêu. Veronicastrum sibiricum var. japonicum f. 188 植物研究雑誌 第 90 巻 第 3 号 2015 年 6 月 albiflorum Akasawa in Bull. Kochi Women’s Wild Flowers Jap. Herb. 3: 107 (1981), comb. Coll. 2: 109 (1955). – V. japonicum f. albiflorum nud.]. – Veronicastrum japonicum (Nakai) T. (Akasawa) T. Yamaz. in K. Iwats. & al., Fl. Jap. Yamaz. in J. Jap. Bot. 65: 241 (1990), & in K. 3a: 347 (1993). Iwats. & al., Fl. Jap. 3a: 346 (1993); H. Ohashi Nom. Jap.: Shirobana Nangoku Kugaisô シロ & al., New Makino Illust. Fl. Jap. 684, fig. 2736 バナナンゴククガイソウ． (2008). Distr.: Japan. Shikoku. Veronicastrum sibiricum (L.) Pennell var. yezoense H. Hara in J. Jap. Bot. 16: 161 (1940) f. glabratum (Nakai) H. Ohashi, comb. nov. [Holotype: RUSSIA. Sakhalin. Toyohara. 28 [Fig. 6] Jul. 1936. H. Hara s.n. TI], & Enum. Spermat. Veronica sibirica var. yezoensis f. glabrata Jap. 1: 282 (1949); T. Yamaz. in K. Iwats. & al., Nakai in J. Jap. Bot. 19: 6 (1943) [Holotype: Fl. Jap. 3a: 346 (1993). – Veronica sibirica var. JAPAN. Hokkaido. Prov. Ishikari, Sapporo. 7 yezoensis (H. Hara) Nakai in J. Jap. Bot. 19: 6 Jul. 1891. Tokubuti s.n. TI]. (1943), ut ‘var. yezoensis Hara’. – Veronicastrum Veronica virginica Thunb., Fl. Jap.: 20 sibiricum subsp. yezoense (H. Hara) T. Yamaz. (1784), non L. (1753) [The specimen not existing in J. Jap. Bot. 65: 241 (1990). in UPS (fide Nakai 1943)]; Matsumura, Ind. Pl. Veronica sibirica var. yezoensis f. pubescens Jap. 2(2): 574 (1912). – Leptandra virginica (L.) Nakai in J. Jap. Bot. 19: 6 (1943) [Holotype: Nutt., Gen. N. Amer. Pl. 1: 7 (1818), p.p., incl. pl. RUSSIA. Sakhalin (Karafuto). Toyohara. 5 Aug. ex Japonia. – [Leptandra sibirica Nutt., Gen. N. 1923. Sawada-Taketarô s.n. TI]. Amer. Pl. 1: 7 (1818), nom. inval.]: Makino in J. Veronica sibirica var. yezoensis f. latifolia Jap. Bot. 1(2): (31) (1916), & Illust. Fl. Nippon Nakai in J. Jap. Bot. 19: 7 (1943), nom. illeg., 139, f. 417 (1940). – Eustachya japonica Raf. in non V. sibirica var. japonica f. latifolia Nakai Ann. Gen. Sci. Phys. 6: 97 (1820). – Leptandra [Holotype: JAPAN. Hokkaido. Prov. Osima, japonica (Raf.) Raf., Med. Fl. 2: 21 (1830). – Hakodate. 15 Aug. 1899. Matumura-Zinzô (J. Veronica japonica (Raf.) Steud., Nomencl. Bot. Matsumura) s.n. TI]. ed. 2, 2: 757 (1841); Siebold & Zucc. in Abh. Veronica sibirica var. japonica f. latifolia Akad. Muench. 4(3): 143 (1846). Nakai in J. Jap. Bot. 19: 7 (1943), nom. illeg., Veronica virginica L. var. japonica Nakai in non V. sibirica var. yezoensis f. latifolia Nakai Bot. Mag. (Tokyo) 26: 170 (1912) [Lectotype [Lectotype (designated here): JAPAN. Honshu. (designated by Yamazaki 1990): JAPAN. Prov. Mutsu (Aomori Pref.), Iwaya-Kwannon Honshu. Nagano Pref., Yatsugatake, Umagaeshi. (Kinashi-Nobutarô Aug. 1911. TI)]. 17 Aug. 1902. Y. Yabe s.n. TI]; Makino & Veronica sachalinensis T. Yamaz. in J. Jap. Nemoto, Nippon Shokubutsu Sôran (Fl. Jap.) Bot. 27: 62 (1952) [Lectotype (designated ed. 2, 1079 (1931). – Veronica sibirica L. var. here): RUSSIA. Sakhalin (Saghalia). Anbetu. 1 japonica (Nakai) Nakai in Cot. Sem. Hort. Bot. Aug. 1936. H. Hara, no. 1. TI]. Yamazaki (1952) Univ. Tokyo: 33 (1914); Furumi in Bot. Mag. designated two types for the name, of which (Tokyo) 30: 121 (1916); Nakai in J. Jap. Bot. 19: “typus florum” was selected here as lectotype. 7 (1943). – Veronicastrum sibiricum (L.) Pennell Veronica sibirica var. japonica f. rosacea var. japonicum (Nakai) H. Hara in J. Jap. Bot. Kitag. in J. Jap. Bot. 55: 268 (1980) [Holotype: 16: 160 (1940), & Enum. Spermat. Jap. 1: 281 JAPAN. Prov. Shinano, prope Yunomaru. 27 (1949); T. Yamaz. in J. Jap. Bot. 27: 64 (1952) Aug. 1972. M. Kitagawa s.n. TI]． & in J. Fac. Sci. Univ. Tokyo sect. 3, 7(2): 132 Veronicastrum borissovae (Czerep.) Soják (1957). – [Veronicastrum sibiricum subsp. in Sborn. Nár. Muz. Praz. ser. B, 39: 59 (1983) japonicum (Nakai) T. Yamaz. in Satake & al., [Type: Veronica sachalinensis Boriss. (1955) June 2015 The Journal of Japanese Botany Vol. 90 No. 3 189

Fig. 6. Makinoʼs illustration of “Kugaisô”, Leptandra sibirica Nutt. (=Veronicastrum sibiricum f. glabratum) from The Journal of Japanese Botany 1: (32) (1916).

[Fl. URSS 22: 496, t. 23, f. 3 (1955)], non T. sibiricum var. japonicum f. albiflorum Hayashi Yamaz. (1952)]; H. Ohashi & al., New Makino in Bull. Gov. For. Exp. Stat. no. 170: 83 (1964), Illust. Fl. Jap. 684, fig. 2735 (2008); Yonek., non Akasawa (1955) [Type: JAPAN. Honshu. Enum. Vasc. Pl. Jap.: 192 (2012). Nagano Pref., Mt. Yatsu-nishi. 6 Jul. 1964. Y. Nom. Jap.: Kugaisô クガイソウ（ エゾクカイソ Hayashi s.n. Herb. Govt. For. Exp. Stat.]. – ウ）． V. japonicum (Nakai) T. Yamaz. f. album (T. Distr.: Japan (Hokkaido, Honshu) and Russia Shimizu) T. Yamaz. in K. Iwats. & al., Fl. Jap. (Sakhalin). 3a: 346 (1993), as “(Sugim.) T. Yamaz.” V. sibiricum subsp. japonicum (Nakai) T. f. album T. Shimizu Yamaz. f. albiflorum N. Yonez. in J. Phytogeogr. V. sibiricum (L.) Pennell var. japonicum & Taxon. 32: 126, photo (1984), non Akasawa (Nakai) H. Hara f. album T. Shimizu in J. (1955) [Holotype: JAPAN. Honshu. Yamanashi Phytogeogr. & Taxon. 37: 120 (1989). – V. Pref., Mt. 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大橋広好：クガイソウ（オオバコ科）の変異 クガイソウはクカイソウとも呼ばれていた．牧野 と し て Veronicastrum japonicum (Nakai) T. Yamaz. (1916) は和名クガイソウの語源を「くがいさうハ九蓋 とした．一方，大井 (1953, 1965a, 1965b)，大井・北 草ノ義ナリ．其葉莖上ニ層ヲナス．少キハ八九層多キハ 川 (1992) は ク ガ イ ソ ウ を Veronica sibirica L.， 後 に 十二三層ニ至ル．故ニ此ノ名アリ．くかいそうハ九階草 Veronicastrum sibiricum (L.) Pennell とみなし，シベリ ノ義ナリ．亦能ク其實際ヲ顕ハスノ名ナリ」と解説した アクガイソウと区別せず，また種内分類群を認めていな （牧野富太郎 1916. 小學讀本ノくがいさう．植物研究雑 い． 誌 1(2): (29)–(33), Fig. 6）．と こ ろ が Nakai (1943, p. 山崎 (1952) は当初クガイソウを次のように考えてい 11) はクカイソウは寺島良安和漢三才圖繪 46 (1712) や た．「輪生葉を持つクガイソウの類は東亜に１種であり， 飯沼慾斎草木圖説 (1856) にあり，クガイソウは岩崎灌 かなり形態的変化があってシベリアクガイソウ，ツクシ 園本草圖譜巻 29 (1838) からなので，クカイソウが（優 クガイソウ，ナンゴククガイソウ，クガイソウと変種と 先権があり）正しいとした（平凡社東洋文庫 527 寺島 して区別される．大陸から本州にかけて逐次的に変化 良安「和漢三才図会」17 島田勇雄・竹島淳夫・樋口元 しており，北海道・樺太にあるエゾクガイソウは他の 巳（訳注）1991 の 238 ページには「威霊仙」のふりが 変種に比べて変化の程度は大きい」．後に山崎 (1990) と なに「くがいそう」があり，解説の始めに「九蓋（くがい） Yamazaki (1993) ではこの見方を変更して，シベリアク 草［いま俗にこういう］」とあるが，「くかいそう」はない． ガイソウとクガイソウを別種とし，シベリアクガイソウ 圖繪 (1712) の原本で確認していないが，クカイソウが はツクシクガイソウとエゾクガイソウに，クガイソウは 中井の間違いとは考え難い）．さらに，Thunberg (1784) イブキクガイソウとナンゴククガイソウの 2 変種を区 はクガイソウの記載に ‘Tobi, vulgo Toranoo. Kaempf. 分した． Am. ex Fasc. V. p.886’ と引用していることからすると， しかし，これまでのクガイソウ類相互の間の区別は変 Kaempfer（1690–92 年に日本滞在）によれば，和漢三 種以上のランクで識別できるほど明瞭とは考えられな 才圖繪 46 (1712) より前に「Tobi または虎の尾」と呼 い．山崎 (1952) のいう逐次的形態変化は連続的変異と ばれていたらしい．現在の和名ではクガイソウに落ち着 みることができ，気温差に関連したクライン的変異と いていると思われるので，以下ではクガイソウで統一す 思われる．著者はクガイソウを 1 種と見る大井 (1953, る． 1965) の見解が正しいと考える．中国，韓国，ロシア クガイソウの最初の学名は Thunberg (1784) に よ でも１種 Veronicastrum sibiricum としている．中国で る．Thunberg はこの植物を北アメリカ原産の Veronica V. sibiricum としている種を記載 (Chin and Hong 1979, virginica L. に 当 て， 和 名 に ‘Ireisin’（ イ レ イ シ ン， Hong et al. 1998) から判断するに Yamazaki (1993) の 漢 名「 威 霊 仙 」 よ り ） ま た は ‘Kwugai So’（ ク ガ イ V. japonicum に当たる形である． ソウ）などを記録し，長崎，大阪などに生えるとし 以上の点から，クガイソウを 1 種とし，この中にい た． 中 井（1943 p. 9 脚 注 ） に よ れ ば，Thunberg は くつかの品種を認めて，学名，和名（異名），分布を次 Veronica virginica L. を大阪で目撃して備忘録に記 のようにまとめた． 録 し た も の で，UPS に は 標 本 が な く，「く が い そ う Veronicastrum sibiricum (L.) Pennell クガイソウ（広義） ノ如何ナル形態ノモノヲ見タカ判ラヌ・・」という． f. sibiricum シベリアクガイソウ（ナンゴククガイソ Rafinesque (1836) は，Thunberg の記録した植物は北 ウ，ツクシクガイソウ）．分布：本州，四国，九州（阿 アメリカの Veronica virginica とは異なるとして，新 蘇山系），朝鮮，中国，東シベリア，極東ロシア． 名 Eustaxia japonica Raf. を 用 い た．Steudel (1841) f. albiflorum Akasawa シロバナナンゴククガイソウ． は こ れ を Veronica japonica (Raf.) Steudel. に 変 更 し シベリアクガイソウの白花品．分布：四国． た．Siebold and Zuccarini (1846) は こ の 学 名 で 日 本 f. glabrata (Nakai) H. Ohashi クガイソウ．この学名に か ら ク ガ イ ソ ウ を 記 録 し た． し か し，Nakai (1912) 対する和名は原記載ではエゾクカイソウであるが，クガ は ク ガ イ ソ ウ を Veronica virginica の 新 変 種 と し て， イソウを採用する．分布：本州，北海道，サハリン． Veronica virginica var. japonica Nakai と命名し，1914 f. album T. Shimizu シロバナクガイソウ（シロバナエ 年に Veronica sibirica L. シベリアクガイソウの変種 ゾクガイソウ）．クガイソウの白花品．分布：北海道， に移した．その後，クガイソウはシベリアクガイソウ 長野県． の 変 種 (Nakai 1914, 1943, Furumi 1916, Hara 1940, f. humile (Nakai) H. Ohashi イブキクガイソウ．分布： Yamazaki 1952, 1947) またはその亜種（山崎 1981）と 滋賀県（伊吹山），愛媛県（東赤石山系），徳島県（剣山）． され，さらに山崎 (1990), Yamazaki (1993) は独立種 （東北大学植物園津田記念館）