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Aristolochiaceae) in Japan J. Jpn. Bot. 89: 152–163 (2014) Morphological Variations of Aristolochia kaempferi and A. tanzawana (Aristolochiaceae) in Japan a, d, a, b, d Tetsuo OHI-TOMA *, Kana WATANABE-TOMA , c a Hiroko MURATA and Jin MURATA aBotanical Gardens, Graduate School of Science, the University of Tokyo, Bunkyo-ku, Tokyo, 112-0001 JAPAN; b Current address: Musashi High School and Junior High School, Nerima-ku, Tokyo, 176-8535 JAPAN; c Faculty of Pharmaceutical Sciences, Setsunan University, Hirakata, Osaka, 573-0101 JAPAN d Equal contribution *Corresponding author: [email protected] (Accepted on February 4, 2014) Aristolochia kaempferi Willd. is widely distributed in Honshu (Kanto and westwards), Shikoku and Kyushu in Japan, and shows large morphological variation. Within the species, several infraspecific taxa have been recognized, but only one variety, var. tanzawana Kigawa, is recently accepted based on the spreading hairs on the nerves of the abaxial side of leaves. Based on an extensive investigation through the whole geographical range of A. kaempferi, we found the differences in the flower morphology and the segregation of distribution between var. kaempferi and var. tanzawana. Here, A. tanzawana (Kigawa) Watanabe-Toma & Ohi-Toma is newly recognized in the rank of species. Key words: Aristolochia kaempferi, Aristolochia tanzawana, Aristolochiaceae, var. kaempferi, var. tanzawana. The genus Aristolochia L. sensu lato species). (Aristolochiaceae) includes over 400 species One of the monophyletic groups of “Isotrema” from temperate to tropical regions worldwide. is known as the Aristolochia kaempferi group In the molecular phylogeny, the genus is divided consisting of six taxa from Japan through Taiwan into two lineages each with two subgroups, to the mainland of China (Watanabe et al. 2006), one with “Aristolochia” and “Pararistolochia” i.e., A. kaempferi Willd. var. kaempferi and and another with “Isotrema” and “Endodeca” var. tanzawana Kigawa, A. liukiuensis Hatus., (Ohi-Toma et al. 2006). Of them, “Isotrema“ is A. shimadae Hayata, A. cucurbitifolia Hayata, characterized by having a gynostemium with and A. mollissima Hance. Of the members, three segments, anthera paired on the outer A. kaempferi var. kaempferi, which is widely surface of each gynostemium segment, a trilobed distributed in Honshu (Kanto and westwards), perianth, and an apically dehiscent capsule. This Shikoku and Kyushu in Japan (Murata 2006), group is diversified mainly in East Asia (over has often been recognized also in China 50 species) and Central America (nearly 15 including Taiwan (cf. Hwang 1981, Huang et al. —152— June 2014 The Journal of Japanese Botany Vol. 89 No. 3 153 and var. tanzawana based on detailed fieldwork around the Kanto region (Watanabe-Toma et al. 2012). After that we continued and expanded the investigation of the morphological variations of the varieties to the whole geographical range from the Kanto region to Shikoku and Kyushu, we come to the conclusion that var. tanzawana should be recognized at the rank of species, as A. tanzawana. Fig. 1. Floral characters measured. a. Height of perianth. b. Materials and Methods Height of limb. c. Width of perianth mouth. d. Height of perianth mouth. e. Height of utricle (in front view). As the proportion of flowering plants of f. Width of utricle mouth. g. Width of utricle (in front Aristolochia kaempferi var. kaempferi and view). h. Width of limb. i. Length of utricle. j. Length var. tanzawana is generally very low in most of ovary. k. Length of terminal pedicel. l. Length of natural populations, we needed to conduct basal pedicel. m. Length of bract. detailed field investigations over a decade. Nevertheless, if flowering plants could not be 2003), although Ma (1989) recognized Chinese obtained, we cultivated juvenile plants from plants as different. This taxonomic confusion the populations several years until flowering. A is due to the difficulties of finding flowering total 140 flowering plants from 101 locations plants in natural populations and figuring out could be observed (Appendix 1). Their voucher the 3D shape and coloring of the flowers from specimens were deposited in the herbarium TI, herbarium specimens, even though flower the University of Tokyo. morphology has important diagnostic characters The obtained plants were primarily identified for the genus. Based on our recent examinations as var. kaempferi or var. tanzawana based on of literatures, herbarium specimens, and living the hairs on the nerves of the abaxial side of plants in natural populations, A. kaempferi leaves. These varieties have flowers consisting becomes evident to be endemic to Japan. of a U-shaped perianth tube and a trilobed- Under A. kaempferi, several infraspecific limb with a perianth mouth (Fig. 1), but size, taxa have been recognized based on leaf shape and coloring of floral parts are largely variation in Japan and China, but most of them variable. In order to examine the morphological are not accepted in recent floras (Huang et variation, for one fresh flower per individual, 13 al. 2003, Murata 2006). In the recent flora of floral characters (Fig. 1, a–m) were measured Japan (Murata 2006), only one variety, var. with a digital slide caliper and a ruler, and tanzawana Kigawa, which is endemic to Japan, coloring of limb and throat were observed. A has been accepted based on the spreading hairs pairwise comparison for the floral measurements on the nerves of the abaxial side of leaves, between var. kaempferi and var. tanzawana was perianth limb with nigrescent tiger brindle and conducted. If F-test indicated unequal variances dark purple dots inside perianth mouth, but the of the two samples, t-test with a correction for floral characters have not been emphasized as the unequal variances was conducted. Here, diagnostic characters (Kigawa 1989, Murata because the plants with intermediate floral 2006). Recently, we showed the differences morphology between the varieties are rarely of their flower morphology (size, shape and found along the border of their distribution range color), and the segregation of the geographical (Watanabe-Toma et al. 2012, see Taxonomic distribution between A. kaempferi var. kaempferi treatment), these morphological intermediates 154 植物研究雑誌 第 89 巻 第 3 号 2014 年 6 月 Fig. 2. Leaf variations of Aristolochia kaempferi. A. Var. kaempferi. 1. Shizuoka (K500). 2. Fukuoka (K304). 3–5. Kochi (K158). B. Var. tanzawana (= A. tanzawana). 1. Shizuoka (K542). 2. Ibaraki (K255). 3. Shizuoka (K167). 4. Kanagawa (K669). Figure in parentheses is sample reference number (see Appendix 1 and “Representative specimens examined”). Scale bar = 5 cm. were not included in the comparison. at base, and obtuse or acuminate apex, or is Herbarium specimens in Japanese herbaria slender in the middle lobe, auriculate at base. (TI, HIRO-MY, KPM, KYO, MAK, MBK, The leaf shape variation is often observed OSA, SHO, TNS, TUS, and WMNH) were within an individual. Even without flowers, examined to check leaf variation and the plants having narrowly ovate leaves with geographical distribution. auriculate base were sometimes identified as A. shimadae (= A. onoei Franch. & Sav. ex Results and Discussion Koidz. and A. kaempferi Willd. var. trilobata Variation of leaves and plant sizes Franch. & Sav.), although the species is clearly For all examined samples, Aristolochia distinguished from A. kaempferi based on the kaempferi var. kaempferi and var. tanzawana flower morphology (Murata 2006, see Key to were clearly distinguished by the hairs on nerves the species). Because plants having such leaves of abaxial side of leaves. In both varieties, the are sometimes observed in both species, it is leaf shape shows large variation (Fig. 2); leaf difficult to distinguish between A. shimadae and lamina is cordate, ovate to narrowly ovate, A. kaempferi based on only leaf shape. triangular with cordate or round lateral lobes Flowering plants of var. kaempferi are a June 2014 The Journal of Japanese Botany Vol. 89 No. 3 155 large woody climber with a long underground of central Honshu through the Setouchi side stem. On the other hand, flowering plants of of Chugoku region, Shikoku and Kyushu var. tanzawana shows large size variation; large to Tanegashima and Yakushima Islands. woody climbers (over 10 m) at lowlands in the Around the Kanto and Izu regions, the Ise- western part of the range (western Shizuoka and Shima region (Mie Pref.) of the eastern part Aichi Prefs.) and small vine like herbs (under of the Kii Peninsula, and the southern part of 30 cm) having several flowers in the mountain Kyusyu (Miyazaki and Kagoshima Prefs.), region of southern Yamanashi Pref. moderate numbers of flowering plants were observed, but few flowering plants could be Floral variation found in other regions. The geographical range The flower of Aristolochia kaempferi of var. tanzawana is from mountain regions of var. kaempferi generally have a perianth tube Kanagawa, Yamanashi, and Shizuoka Prefs. constricted to a utricle, a yellowish limb with (but not in Mt. Fuji), through lowlands in a fine reddish to dark purplish striae, and fine western part of Shizuoka Pref. to Aichi Pref. and reddish to dark purple colored or densely dotted a southern part of Gifu Pref., and is distantly throat. The lower lobe of the limb is little longer isolated around Mt. Tsukuba (Ibaraki Pref.) in than upper lobes and incurved apically or not. the north Kanto region. The geographical ranges The shape of the perianth mouth varies form of the varieties are mostly segregated, although round to depressed ovate. The coloring of limb, the varieties rarely live in intimate proximity to perianth mouth, and throat are largely varied each other (cf. around mountains in the Kanto (Fig. 3), for example, the whole or the basal lobe distinct and around the northwestern part of of a limb is sometimes densely patterned by dark Aichi Pref.). purple striae, and in rare cases a limb is entirely black-purple or whitish yellow without striae. Taxonomic consideration For the flowers of var.
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