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Variation in Veronicastrum Sibiricum (Plantaginaceae)

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Variation in Veronicastrum Sibiricum (Plantaginaceae) J. Jpn. Bot. 90: 179–191 (2015) Variation in Veronicastrum sibiricum (Plantaginaceae) Hiroyoshi OHASHI Herbarium TUS, Botanical Garden, Tohoku University, Sendai, 980-0862 JAPAN E-mail: [email protected] (Accepted on 9 January 2015) Morphological variation of Veronicastrum sibiricum (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 Veronica 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 leaves, 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 plant 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 —179— 180 植物研究雑誌 第 90 巻 第 3 号 2015 年 6 月 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 leaf-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.
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