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.

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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. tanzawana, the On the investigation through the whole outside of the perianth tube is covered with geographical range, we concluded that velvety hairs, the limb is densely patterned by Aristolochia kampteri var. tanzawana should dark purple striate, and the throat is spotted be recognized as a morphologically and by dark purple leopard brindle (Fig. 4: 1–12). geographically distinct species from var. Some plants have the entirely black-purple kaempferi, although each of the two varieties is limb. In comparison with var. kaempferi, var. paraphyletic due to past geographical isolation tanzawana has significantly larger size of and subsequent secondary gene flow as well as flowers (Table 1), whose perianth tube is not other taxa of the A. kaempferi group according much constricted at a utricle (“f/g” in Table 1). to the molecular phylogeographic analyses Thus, A. kaempferi var. tanzawana found to be (Watanabe et al. 2006, 2008). clearly distinguished from var. kaempferi based In the literature survey, it was revealed that on flower morphology. Aristolochia lineata Duch. ex Decne., which was known only from a drawing of Japanese Geographical distribution collections of M. Delessert (see the digital image Based on the detailed fieldwork and in herbarium MPU; http://www.herbier-mpu. herbarium specimens, the geographical org/zoomify/zoomify.php?fichier=MPU018703) distribution of Aristolochia kaempferi var. and was treated as a synonym of A. kaempferi kaempferi and var. tanzawana was figured out var. kaempferi (Murata 2006), seems to have (Fig. 5). Var. kaempferi is widely distributed similar floral characteristics (i.e., dotted throat, from the Kanto region and the Pacific side densely striate on the whole of the limb, and 156 植物研究雑誌 第 89 巻 第 3 号 2014 年 6 月

Fig. 3. Floral variations of Aristolochia kaempferi var. kaempferi in front view. 1. Chiba (K582). 2. Tokyo (K290). 3. Kanagawa (K665). 4. Shizuoka (K764). 5. Aichi (K786). 6. Aichi (K787). 7. Kyoto (K576). 8. Wakayama (K564). 9. Wakayama (K795). 10. Mie (K801). 11. Mie (K799). 12. Mie (K569). 13. Kochi (K158). 14. Ehime (K377). 15. Ehime (K336). 16. Tokushima (K370). 17. Hyogo (K363). 18. Okayama (K398). 19. Hiroshima (K401). 20. Yamaguchi (K325). 21. Fukuoka (K305). 22. Fukuoka (K618). 23. Nagasaki (K444). 24. Nagasaki (K606). 25. Saga (K432). 26. Saga (K443). 27. Kumamoto (K69). 28. Miyazaki (K549). 29. Kagoshima (K301). 30. Kagoshima (K453). Figure in parentheses is sample reference number (see Appendix 1). Scar bar = 5 mm. perianth tube not much constricted to a utricle) lineata that the whole of plant is glabrous and to var. tanzawana. Because we cannot evaluate smooth. Therefore, we regard A. lineata as a the hairs on the nerves of abaxial side of leaves synonym of A. kaempferi var. kaempferi. in the drawing and the detailed locality is not In the examination of herbarium specimens, indicated, we depended on the description of A. two specimens in MAK from the Ise-Shima June 2014 The Journal of Japanese Botany Vol. 89 No. 3 157

Fig. 4. Floral variations of Aristolochia kaempferi var. tanzawana (= A. tanzawana, 1–12) and putative hybrids (13–15) in front view. 1. Ibaraki (K258). 2. Ibaraki (K260). 3. Kanagawa (K96). 4. Kanagawa (K669). 5. Kanagawa (K649). 6. Yamanashi (K757). 7. Yamanashi (K768). 8. Yamanashi (K721). 9. Shizuoka (K532). 10. Shizuoka (K747). 11. Aichi (K780). 12. Aichi (K775). 13. Kanagawa (K750). 14. Yamanashi (K756). 15. Shizuoka (K686). Figure in parentheses is sample reference number (see Appendix 1 and “Representative specimens examined”). Scar bar = 5 mm.

Table 1. Size variation in the floral characters ofAristolochia kaempferi var, kaempferi and var. tanzawana (=A. tanzawana), using 96 and 44 individuals, respectively. Var. kaempferi Var. tanzawana t-test (p) Average ± SD (mm*) Average ± SD (mm*) Height of perianth (a) 26.73 3.07 36.61 3.75 0.00 Height of limb (b) 16.77 2.13 22.35 2.56 0.00 Width of perianth mouth (c) 7.04 1.28 9.56 1.71 0.00** Height of perianth mouth (d) 5.21 1.28 7.08 1.52 0.00 Height of utricle (e) 8.19 1.53 10.63 1.78 0.00 Width of utricle mouth (f) 3.96 0.58 6.72 1.10 0.00** Width of utricle (g) 7.77 1.45 10.52 1.13 0.00 Width of limb (h) 20.04 3.16 25.23 3.32 0.00 Length of utricle (i) 17.85 2.63 23.51 2.83 0.00 Length of ovary (j) 10.01 1.71 11.35 1.43 0.00 Length of terminal pedicel (k) 26.41 7.21 31.79 6.47 0.00 Length of based pedicel (l) 3.44 4.22 3.55 4.05 0.89 Length of bract (m) 5.36 1.59 5.72 1.60 0.25 Constriction of perianth tube (f/g) 0.52 0.08 0.64 0.09 0.00 *Except for the value of “f/g”. **The variances were unequal in both samples by the F-test. 158 植物研究雑誌 第 89 巻 第 3 号 2014 年 6 月

Fig. 5. Geographical distribution of Aristolcohia kaempferi (black circles) and A. tanzawana (= A. kaempferi var tanzawana, gray squares). Open circles are indicated plants, which are probably A. kaempferi but these flowers were not confirmed. Another species distributed in the Japanese mainland, A. shimadae, is also included by stars in this map (see Appendix 2). region (J. Umemura s.n., 14 May 1893, and K. selectae plantarum, quas in Japonia, t. 49 (1791). Uno 22684), were annotated as “Aristolochia [Fig. 3, 1–30] simaensis Hiyama” (the meaning of the species Aristolochia lineata Duch. ex Decne. in Rev. epithet is probably the regional name) by Dr. Hort., ser. IV 3: 284 (1854). Type: JAPAN. A K. Hiyama in 1960, but this name was not drawing in Japanese collections of M. Delessert published. Plants in the natural populations of (MPU081703, digital image!). the Ise-Shima region have a whitish perianth Aristolochia kaempferi Willd. var. longifolia tube and a limb with fine reddish purple striae in Franch. & Sav. in Enum. Pl. Jap. 1: 419 (1875) the upper part and reddish purple reticulation in & 2: 485 (1878). Type: JAPAN. Savatier n. 2719 the lower part (Fig. 3: 6, 12), and a fine reddish (P–isotype, digital image P!). purple pedicel that is sometimes longer than the Aristolochia kaempferi Willd. var. pallescens plants in other regions (data not shown). We Nakai in Rep. Veg. Isl. Shikajima, Iyo: 38 do not recognize these of plants as a distinct (1927), in Japanese; Nakai in Bot. Mag. (Tokyo) taxonomic group at this time, because the range 43: 439 (1929). Lectotype (designated here, ICN of morphological variation is not discrete from Art.8.3): JAPAN. Shikajima, prov. Iyo (Ehime), total variation range of other A. kaempferi. T. Nakai s.n., 9 June 1927 (TI–lectotype [fls.]!, TI–isoletotype [no fls.]!). Taxonomic treatment Woody, twining perennial climbers 2–10 m Aristolochia kaempferi Willd. in Sp. Pl. 4: long or more. Young branches pale green, terete, 152 (1805). Type: JAPAN. E. Kaempfer, Icones pubescent; old branches gray, terete, striate. Leaf June 2014 The Journal of Japanese Botany Vol. 89 No. 3 159 lamina cordate, ovate to narrowly ovate, base Kochi Prefs.) and Kyusyu (Fukuoka, Oita, Saga, cordate, sometimes with round lateral lobes at Nagasaki, Kumamoto, Miyazaki and Kagoshima base, apex obtuse or acuminate, or angustate in Prefs.) to Tanegashima and Yakushima Islands, the middle lobe with auricules at base, to 3–18 in altitude 0 m to 1,000 m. Except in Chiba, cm long, 4–20 cm wide, herbaceous to leathery, Kanagawa, Shizuoka and Mie (the Ise-Shima) adaxial surface green or light green and sparsely Prefs. and south Kyusyu, the flowering plants pubescent, abaxial surface grayish green and of the species are rare. In some areas, cf. Osaka, shortly hairy; nerves prominent on abaxial central part of the Kii peninsula and Yakushima surface, with appressed hairs; petiole 1.5–9 Island, some specimens of juveniles were cm long. Flowers April to June, solitary in axil collected, but flowering plants have not been of prophylls of lateral branches; pedicel light observed. The species was recorded in Nagano green or rarely fine reddish purple, 18.5–41.5 Pref., central Honshu (Shimizu 1997), but mm long; bract usually at base of pedicel, ovate, we have confirmed neither living plants nor 3.5–7 mm long. Perianth tube U-shaped, 2–3 herbarium specimens. cm , 5–10.5 mm wide at utricle, upper 1/3 more Representative specimens examined: JAPAN. Chiba: slenderer (3.5–4.5 mm wide), outside whitish Minami-Boso, K. Watanabe & T. Ohi-Toma K642 (TI); Kimitsu, Mt. Kanou, without collector name, May 1881 yellow to creamy, pubescent, inside greenish (TNS). Tokyo: Izu-Oshima Island, T. Ohi-Toma & T. Ideno yellow with purple markings, smooth, and throat K656 (TI). Kanagawa: Kamakura, K. Watanabe K162 dotted reddish to dark purple; mouth round (TI); Oiso, K. Watanabe-Toma & T. Ohi-Toma K645 (TI); to depressed ovate, 3.5–5.5 mm high, 5.5–8 Kiyokawa, K. Watanabe-Toma & T. Ohi-Toma K658, K659 mm wide; limb broadly obovate, shallowly (TI); Yamakita, Yaga, K. Watanabe-Toma & T. Ohi-Toma K663 (TI); Hadano, Kominoge, K. Watanabe-Toma & T. trilobed, incurved apically or not, 16–23 mm Ohi-Toma K691, K692 (TI); between Mt. Takatori and Mt. high, 13.5–30.0 mm wide in front view, smooth, Nenbutsu, K. Watanabe-Toma & T. Ohi-Toma K694 (TI); yellowish with fine reddish to dark purple striae, Odawara, Kuno, K. Watanabe-Toma & T. Ohi-Toma K698 sometimes densely striate or reticulate in the (TI); Ashigara, K. Watanabe-Toma & T. Ohi-Toma K699 (TI); Hakone, Gora, K. Watanabe-Toma & T. Ohi-Toma lower part, rarely entire black-purple or whitish K701 (TI); Hadano, Imaizumi, K. Watanabe-Toma & T. yellow without striae. Stamens 6, adnate to Ohi-Toma K734 (TI); Mt. Takamatsu, T. Ohi-Toma K741 style column, anthers oblong; stigmatic lobes (TI). Yamanashi: Mt. Kanga-dake, T. Ohi-Toma K754 3, triangular; ovary inferior. Capsules May to (TI). Shizuoka: Amagi, T. Ideno D-518 (TI); Shuzenji, K. August, pubescent, cylindrical to narrowly Watanabe-Toma & T. Ohi-Toma K680 (TI); Gotenba, T. Ohi-Toma K574 (TI); Otome Pass, K. Watanabe-Toma & ellipsoid, with 6 developed ridges, 2.5–5 cm T. Ohi-Toma K676 (TI); Kan-nami, K. Watanabe-Toma long, dehiscent from apex. Seeds flat, narrowly & T. Ohi-Toma K681 (TI); Fuji, Kuwasaki, K. Watanabe oval, 5 mm long. K497, K500 (TI); Fuji, Kitamatsuno, T. Ohi-Toma K752 Japanese name: Ôba-uma-no-suzu-kusa (TI); Fujinomiya, Hoshiyama, K. Watanabe K590 (TI); Fujinomiya, Mt. Myojo, K. Watanabe-Toma & T. Ohi- (Ôba-muma-no-sudzu-kusa; Matsumura in Toma K740 (TI); Fujinomiya, Mt. Shiratori, K. Watanabe- Nippon Shokubutsumeii, p. 18, 1884). Toma & T. Ohi-Toma K745 (TI); Shizuoka, Makigaya, 和名：オオバウマノスズクサ（松村） K. Watanabe K90 (TI); Shizuoka, Kanbara, K. Watanabe- Chromosome number: 2n = 32 (Sugawara Toma & T. Ohi-Toma K705 (TI); Mt. Takakusa, K. Watanabe K589 (TI); Shizuoka, Yuyama, K. Watanabe- and Murata 1992). Toma K689 (TI); Shizuoka, Yui, K. Watanabe-Toma & T. Distribution: JAPAN (endemic). Pacific side Ohi-Toma K736, K737 (TI); Fujieda, Okabe, K. Watanabe- and the western part of Honshu (Chiba, Tokyo, Toma & T. Ohi-Toma K749 (TI). Aichi: Inuyama, K. Kanagawa, Yamanashi, Shizuoka, Aichi, Mie, Watanabe-Toma & T. Ohi-Toma K791 (TI). Mie: Mt. Asama, K. Watanabe K408 (TI); Shima, Kawachi, J. Kyoto, Nara, Wakayama, Hyogo, Okayama, Umemura s.n., 14 May 1893 (MAK); Toba, K. Uno 22684 Hiroshima, and Yamaguchi Prefs.), through (MAK); Ise, Tashiro-dani, G. Murata 19506 (MAK); Shikoku (Kagawa, Tokushima, Ehime and Ise, Ouchiyama-mura, S. Suzuki s.n., 4 Jun 1933 (TUS). 160 植物研究雑誌 第 89 巻 第 3 号 2014 年 6 月

Osaka: Mt. Izumi-Katsuragi, S. Fujii 7678, no fls. (OSA); Aristolochia tanzawana (Kigawa) Watanabe- Mt. Inunaki, C. Satonaka s.n., 8 Nov. 1987, no fls. (OSA); Toma & Ohi-Toma, stat. nov. [Fig. 4, 1–12] Izumi-sano, Kaminogou, S. Nakanishi s.n., 7 July 1962, no fls. (OSA); Mt. Imori, S. Nakanishi s.n., 24 June 1962, no Aristolochia kaempferi Willd. var. tanzawana fls. (OSA).Nara : Mt. Ikoma, Kurondo Pond, N. Morimoto Kigawa in Bull. Kanagawa Pref. Mus. Nat. 3548-2 (OSA); Shimo-Kitayama, Zenki, K. Kawabata Sci. 18: 17 (1989). Type: JAPAN. Honshu. 10370, no fls. (KYO). Wakayama: Nachi-Katsuura, K. Kanagawa Pref., Hadano, Mt. Nabewari, 11 Watanabe K128 (TI); Jyujyo Pass, K. Watanabe K138 (TI); June 1988, S. Kigawa s.n. (KPM-NA 1000103– Kushimoto, A. Yamamoto 7259 (WMNH); Mt. Ishido, A. Yamamoto 10774 (WMNH); Hiokigawa, T. Goto 52721 holotype!). (WMNH); Kitayama, Takehara, M. Hori s.n., 2 June 1951 Woody, twining perennial climbers to 1 m (OSA); Oto, Yasukawa-keikoku, K. Seto 53240 (OSA); long or more, sometimes scandent perennial Mt. Negoro, K. Watanabe-Toma & T. Ohi-Toma K794, vines. Young branches pale green, terete, no fls. (TI); Kozagawa, Takinohai, K. Watanabe-Toma & pubescent; old branches gray, terete, striate or T. Ohi-Toma K796, no fls. (TI). Hyogo: Sumoto, Inohara, K. Watanabe et al. K364 (TI); Kamigori, Manshoin green on scandent vines. Leaf lamina cordate, Temple, K. Watanabe-Toma & T. Ohi-Toma K713, no fls. ovate to narrowly ovate, triangular, base cordate, (TI); Tatsuno, Shobu-dani, T. Ono & al. 16745 (SHO). sometimes with round lateral lobes at base, apex Okayama: Mt. Hattoji, K. Watanabe-Toma & T. Ohi-Toma obtuse or acuminate, or angustate in the middle K712, no fls. (TI). Hiroshima: Miyajima Island, S. Mukai lobe, auricules at base, to 3 18 cm long, 4 16 s.n., 2004 (HIRO-MY). Yamaguchi: Ohuchi, without – – collector name, no.167, 14 May 1892 (TI); Yashiro- cm wide, herbaceous to leathery, adaxial surface jima Island, K. Oka 4343 (TNS). Kagawa: Shodo-shima green or light green and densely pubescent, Island, Mt. Hoshigajo, S. Mitani s.n., 20 June 1982 (TI). abaxial surface grayish green and hairy; nerves Tokushima: Mt. Kumosa, K. Watanabe K578 (TI); Mugi, prominent on abaxial surface, with spreading Ohshima Island, Y. Ibaragi & al. 280403006 (TUS). Kochi: Ohikiwari Pass, K. Watanabe K381, no fls. (TI); Nakatosa, hairs; petiole 1.5–10 cm long. Flowers April Kogusa, H. Kakegawa FOK-076185 (MBK), Y. Kurahashi to July, solitary or a few in axil of prophylls of FOK-614763 (MBK); Umaji, Tochi-dani, M. Furusawa & lateral branches; pedicel light green, 20–45 mm al. FOK-077026 (MBK); Gohoku, Takadaru, N. Inagaki & long; bract usually at base of pedicel, ovate, al. FOK-055665 (MBK). Fukuoka: Nakagawa, Minami- 4–7.5 mm long. Perianth tube U-shaped, 3–4.5 hata dam, K. Watanabe K620, no fls. (TI); Mt. Gozen, K. Watanabe K310 (TI). Oita: Kamae, M. Kuwashima cm tall, 8–13 mm wide at utricle, upper 1/3 33257 (OSA). Saga: Tara, Nakayama Campground, K. slightly slender (4.5–8.5 mm wide), outside Yonekura & R. Hirano 98738 (TUS). Nagasaki: Mt. whitish yellow to creamy, velvety-pubescent, Shokan, K. Watanabe K440 (TI). Kumamoto: Mt. Shodai, inside greenish yellow with purple markings, K. Watanabe K73 (TI); Yashiro, Izumi, Y. Koga 12671 smooth, and throat spotted with dark purple (KYO). Miyazaki: Mt. Ohkue, H. Funakoshi 1813 (TI); Kushima, Ichigi, N. Maruyama s.n., 6 April 1948 (TNS); leopard brindle; mouth round to depressed ovate Kijo, Shiinoki, K. Watanabe K49, no fls. (TI).Kagoshima : near the center of the limb, 4–10 mm high, Kirishima, K. Watanabe K42 (TI); Isaku Pass, T. Sugawara 7–14.5 mm wide; limb round to broadly obovate Isaku3 (TI); Ohdomari, S. Hatusima & S. Sako 26891 (TI); in front view, shallowly trilobed, 18–31 mm Yamakawa, K. Watanabe K59 (TI); Yakushima Island, Oko fall, K. Watanabe K179, K180, no fls. (TI). high, 19.5–32.5 mm wide in front view, smooth, Note: Some synonyms of the species recognized whitish to greenish yellow with dark purple striae in Flora of China (Huang et al. 2003) should represent or sometimes entirely black-purple. Stamens 6, distinct taxa, i.e., A. shimadae Hayata in Japan and adnate to style column, anthers oblong; stigmatic Taiwan, A. dabieshanensis C. Y. Cheng & W. Yu and A. lobes 3, triangular; ovary inferior. Capsules May heterophylla Hemsley in central China, and A. mollis Dunn in southeastern China, A. neolongifolia J. L. Wu & to August, pubescent, cylindrical to narrowly Z. L. Yang in western China. In addition, plants identified ellipsoid, with 6 undeveloped ridges, 3.5–6 cm as A. kaempferi in eastern China (Quan 1992) should be long, dehiscent from apex. Seeds flat, narrowly recognized as new taxa (Ohi-Toma et al. in prep.). oval, 5 mm long. Japanese name: Tanzawa-umanosuzukusa June 2014 The Journal of Japanese Botany Vol. 89 No. 3 161

(Kigawa 1988). appressed. Perianth tube 2–3 cm tall, outside 和名：タンザワウマノスズクサ（城川） pubescent. much constricted above the utricle. Chromosome number: 2n = 32 (the plant Throat of perianth tube dotted with reddish to from Mt. Ogasa, Shizuoka Pref., identified as A. dark purple. Perianth limb usually yellow with onoei in Sugawara and Murata 1992). fine reddish to dark puplish striae, sometimes Distribution: JAPAN (endemic). Honshu densely striate or reticulate on in the lower part, (Ibaraki, Kanagawa, Yamanashi, Shizuoka, rarely entire black-purple or whitish yellow Aichi, and Gifu Prefs.), at an altitude of 100– without striae ...... A. kaempferi 1,500 m. In Gifu, only juveniles without flowers 2b. Hairs on nerves of the abaxial side of leaves were recently found. In Kigawa (1989), one spreading. Perianth tube 3–4.5 cm tall, outside locality in Nagano was recorded on their figures, velvety-pubescent, not much constricted above but we could not confirm their specimens. the utricle. Throat of perianth tube spotted Representative specimens examined: JAPAN. Ibaraki: with dark purple leopard brindle. Perianth limb Mt. Tsukuba, K. Watanabe K255, K257, K258, K259, usually whitish to greenish yellow with dark K260, K261 (TI); Ishioka, Shibauchi, K. Watanabe-Toma & T. Ohi-Toma K690 (TI). Kanagawa: Mt. Nabewari, K. purple striae or sometimes entire black-purple ... Watanabe K92, K95, K96 (TI); Mt. Sekiro, K. Watanabe ...... A. tanazawana K120 (TI); Sagamihara, Aone, K. Watanabe K322 (TI); Matsuda, T. Sugawara A2734 (TI); Nishi-tanzawa, K. Aristolochia kaempferi × A. tanzawana Watanabe-Toma & T. Ohi-Toma K673 (TI). Yamanashi: [Fig. 4, 13–15]. Tatsunotsuka Pass, K. Watanabe K593 (TI); Nijumagari Pass, K. Watanabe K652, K653 (TI); Nishikatsura, K. Plants with their intermediate floral Watanabe-Toma & T. Ohi-Toma K702, no fls. (TI); Mt. morphology, which are rarely found along Kenashi, K. Watanabe-Toma & T. Ohi-Toma K744 (TI). the eastern border of the distribution range of Shizuoka: Shizuoka, Oma, K. Watanabe-Toma & T. Ohi- A. kaempferi and A. tanzawana, are putative Toma K717, no fls. (TI); Mt. Ogasa, T. Sugawara A2735, hybrids. The floral morphology is variable, the voucher of chromosome number (TI); K. Watanabe K535, K536 (TI); Mt. Tonmaku, K. Watanabe K169, no and several types are observed. The plants are fls. (TI); Umegashima, K. Watanabe K431 (TI); Fujieda, genetically heterozygous of the parental species Hongo, K. Watanabe-Toma & T. Ohi-Toma K765 (TI); on the tentative genotyping by nuclear DNA Sumata valley, K. Watanabe-Toma & T. Ohi-Toma K771, genes, although the authors are still investigating no fls. (TI); Mt. Fudo, F. Konta & S. Kusaka SK41 (TNS); Makinohara, Saimyoji, K. Watanabe K542, K543 (TI). the detail by multi-locus analyses and artificial Aichi: Toyota, Yotsuya, K. Watanabe K298, K299, K300 crossings. (TI); Toei, Shimoda, K. Yoshida 920473 (TNS). Gifu: Representative specimens examined: JAPAN. Tokyo: Kani, T. Ohi-Toma et al. K788, no fls. (TI). Mt. Takao, T. Ohi-Toma K797 (TI). Kanagawa: Mt. Sekiro, K. Watanabe K125 (TI); Mt. Oyama, K. Watanabe- Key to the species of the Arisaema kaempferi Toma & T. Ohi-Toma K647 (TI); T. Makino s.n., 1906 (MAK); Omote-tanzawa, K. Watanabe-Toma & T. Ohi- group in the Japanese mainland, Toma K664 (TI); Mt. Nabewari, K. Watanabe K287, K289 excluding Nansei Islands: (TI); Mt. Takatori, K. Watanabe-Toma & T. Ohi-Toma 1a. Throat of perianth tube yellow without dots. K693 (TI); Yamakita, Yaga, K. Watanabe-Toma K707 (TI); Perianth limb usually dark purple without striae, Mt. Yagura, K. Watanabe-Toma & T. Ohi-Toma K700 (TI); Sagamihara, Aone, K. Watanabe-Toma & T. Ohi-Toma lobes of limbs recurved, obtriangular in front K320, K750 (TI). Yamanashi: Sanokawa, K. Watanabe- view ...... A. shimadae Toma & T. Ohi-Toma K704 (TI); K. Watanabe-Toma & T. 1b. Throat of perianth tube with reddish to dark Ohi-Toma K779-01~08 (TI); Nanbu, K. Watanabe-Toma purple dots or spots . Perianth limb usually & T. Ohi-Toma K756 (TI). Shizuoka: Shibakawa, K. yellow or greenish yellow with striae, rarely Watanabe-Toma & T. Ohi-Toma K718 (TI); Mt. Kenashi, K. Watanabe-Toma & T. Ohi-Toma K719 (TI); Nihondaira, entire black-purple or whitish yellow without K. Watanabe-Toma K686, K687 (TI); Fujieda, K. striae, broadly obovate or round in front view ...2 Watanabe-Toma & T. Ohi-Toma K766, K767, K776 (TI). 2a. Hairs on nerves of the abaxial side of leaves 162 植物研究雑誌 第 89 巻 第 3 号 2014 年 6 月

We would like to thank the curators of Watanabe K., Kajita T. and Murata J. 2006. Chloroplast TI, KPM, KYO, MAK, SHO, TNS, and DNA variation and geographical structure of the Aristolochia kaempferi group (Aristolochiaceae). Am. TUS for allowing us to examine herbarium J. Bot. 93: 442–453. specimens, to S. Mukai (HIRO-MY), N. Tanaka Watanabe K., Ohi-Toma T. and Murata J. 2008. Multiple (MBK), M. Yokogawa (OSA) and A. Naito hybridization in the Aristolochia kaempferi group (WMNH) for providing digital images, to T. (Aristolochiaceae): evidence from reproductive Ideno and K. Mizunashi for cultivating plants isolation and molecular phylogeny. Am. J. Bot. 95: 885–896. in Koishikawa Botanical Garden, and to T. Watanabe-Toma K., Murata J. and Ohi-Toma T. 2012. Sugawara, T, Shiraiwa, Y. Ito, Y. Yamaguchi, Morphological and ecological differences between K. Sasamura, Y. Yamashita, S. Iio and H. Suga Aristolochia kaempferi var. kaempferi and var. for providing materials and supporting our tanzawana (Aristolochiaceae). J. Jpn. Bot. 87: 67-70 (in Japanese with English abstract). field work. This work was partly supported by Sasakawa Scientific Research Grant from The Appendix 1. Japan Science Society to K.W. on 2003–2004, List of plants examined for floral characters. Their Yamaguchi Encourage Scholarship Foundation voucher specimens are deposited in TI. for TO on 2011–2012, and Fujiwara Natural Aristolochia kaempferi var. kaempferi Chiba: Tohnosho, K. Watanabe-Toma & T. Ohi- History Foundation for TO on 2012. Toma K582; Minami-Boso, K. Watanabe & T. Ohi- Toma K643. Tokyo: Mt. Takao, K. Watanabe K290. References Kanagawa: Yabitsu Pass, K. Watanabe-Toma & T. Huang S. M., Kelly L. M. and Gilbert M. G. 2003. Ohi-Toma K665; Mt. Nenbutsu, K. Watanabe-Toma Aristolochiaceae. In: Wu C. Y. Raven P. H. and Hong & T. Ohi-Toma K697; Mt. Takatori, K. Watanabe- D. Y. (eds), Flora of China 5: 246–269. Science Press, Toma K708. Shizuoka: Kawazu, K. Watanabe K560; Beijing, and Missouri Botanical Garden Press, St. Nishi-Izu, K. Watanabe K561; Shimoda, K. Watanabe Louis. K572, K573; Higashi-Izu, T. Ohi-Toma K588; Susono, Hwang S.-M. 1981. Materials for Chinese Aristolochia. K. Watanabe K77; Mt. Ashitaka, K. Watanabe K81, Acta Phytotax. Sin. 19: 222–231. K83, K85, K88; Fuji, K. Watanabe K494; Fujinomiya, Kigawa S. 1988. Aristolochia. Flora of Kanagawa Torinami, K. Watanabe-Toma & T. Ohi-Toma K738; 1988, Japan. Supplement. pp. 5-6. Flora-Kanagawa Fujinomiya, Inako, T. Ohi-Toma K753; Shizuoka, Association, Yokohama (in Japanese). Makigaya, K. Watanabe K89; Shizuoka, Yuyama, K. Kigawa S. 1989. On the two new varieties and a new Watanabe K430; Shizuoka, Ubume, K. Watanabe-Toma form of the plants from Kanagawa Prefecture. Bull. K682; Mt. Ryuso, K. Watanabe-Toma K688; Shimizu, Kanagawa Pref. Mus. Nat. Sci. 18: 11–22. K. Watanabe-Toma & T. Ohi-Toma K706; Mt. Ma J.-S. 1989. A revision of Aristolochia Linn. from E. & Hamaishi, K. Watanabe-Toma & T. Ohi-Toma K731; S. Asia. Acta Phytotax. Sin. 27: 321-364 (in Chinese Fujieda, Kitagata, K. Watanabe-Toma & T. Ohi-Toma with English abstract). K748; Mt. Bikuishi, K. Watanabe-Toma & T. Ohi- Murata J. 2006. Aristolochia. In: K. Iwatsuki, D. E. Toma K764; Mt. Ushio, K. Watanabe-Toma & T. Ohi- Boufford, and H. Ohba (eds.), Flora of Japan IIa: Toma K778; Fujieda, Kami-yabuta, K. Watanabe-Toma 366–368. Kodansha, Tokyo. & T. Ohi-Toma K783. Aichi: Nagoya, T. Ohi-Toma Ohi-Toma T., Sugawara T., Murata H., Wanke S., Neinhuis et al. K784; Owari-Asahi, T. Ohi-Toma et al. K785; C. and Murata J. 2006. Molecular phylogeny of Komaki, T. Ohi-Toma et al. K786; Kasugai, T. Ohi- Aristolochia sensu lato (Aristolochiaceae) based on Toma et al. K787. Mie: Mt. Yokoyama, K. Watanabe sequences of rbcL, matK, and phyA genes, with special K567, K569, K570; Toba, K. Watanabe K571; Minami- reference to differentiation of chromosome numbers. Ise, Y. Ito & Y. Yamaguchi K581; K. Watanabe-Toma Syst. Bot. 31: 481–492. & T. Ohi-Toma K800, K801; Nomisaka Pass, T. Ohi- Quan L. 1992. Aristolochiaceae. In: Wang J.-X. (ed.), Toma K789; Ise, Tashiro-dani, K. Watanabe-Toma & Flora of Zhejiang 2: 134–144. Zhejiang Science ad T. Ohi-Toma K798, K799; Mt. Asama, K. Watanabe- Technology Publishing House, Hangzhou (in Chinese). Toma & T. Ohi-Toma K802; Taiki, H. Murata K805. Shimizu T. 1997. Flora of Nagano Prefecture. Shinano Kyoto: Minami-Yamashiro, K. Watanabe K576. Mainichi Press, Nagano (in Japanese). Wakayama: Nachi-Katsuura, K. Watanabe K129; Sugawara T. and Murata J. 1992. Chromosome numbers of Yasukawa Valley, K. Watanabe et al. K564, K565; eight species of Aristolochia (Aristolochiaceae) from Kushimoto, K. Watanabe-Toma & T. Ohi-Toma K795. East Asia. Acta Phytotax. Geobot. 43: 27–30. Hyogo: Sumoto, Inohara, K. Watanabe et al. K362, June 2014 The Journal of Japanese Botany Vol. 89 No. 3 163

K363. Okayama: Mt. Kumayama, K. Watanabe K657; near Lake Shoji, T. Ohi-Toma K721; Mt. K398. Hiroshima: Hatsukaichi, Shikigaoka Park, K. Kanyudo, T. Ohi-Toma K751; near Lake Shibire, Watanabe K401. Yamaguchi: Mt. Eboshi, K. Watanabe K. Watanabe-Toma & T. Ohi-Toma K703, K758, K325, K326. Tokushima: Naka, Kita-Nishiu, K. K759; Mt. Minobu, K. Watanabe-Toma & T. Ohi- Watanabe K370; Mt. Kumosa, K. Sasamura K579, Toma K755; Mt. Shinoi, T. Ohi-Toma K757; Dosaka K. Watanabe K580. Ehime: Mt. Seta, K. Watanabe Pass, K. Watanabe-Toma & T. Ohi-Toma K760; near K335, K336; Mt. Sasayama, K. Watanabe K377, K378; Lake Yamanaka, K. Watanabe & T. Ohi-Toma K768. Nishiumi, Nakadomari, K. Watanabe K380. Kochi: Shizuoka: Mt. Akiba, K. Watanabe K165, K167; Mt. Tosashimizu, Izuta, K. Watanabe K158. Fukuoka: Tonmaku, K. Watanabe K176; Mt. Ogasa, K. Watanabe Fukuoka, Shiiba, K. Watanabe K304; Fukuoka, K532, K537; Shimada, Kikugawa, K. Watanabe-Toma Itaya, K. Watanabe K619; Nakagawa, Sefuri Dam, K709; Kakegawa, Nissaka, K. Watanabe-Toma K710; K. Watanabe K618; Mt. Shaka, K. Watanabe K305. Mt. Chiba, K. Watanabe-Toma K711; Omaezaki, Oita: Misentao Pass, K. Watanabe K311, K315. Saga: K. Watanabe-Toma & T. Ohi-Toma K727; Fujieda, Karatsu, Kashiharu Moor, K. Watanabe K432; Mitsuse Nishikata, K. Watanabe-Toma & T. Ohi-Toma K747; Pass, K. Watanabe K443. Nagasaki: Mt. Shokan, K. Umegashima, T. Ohi-Toma K763; Mori, K. Watanabe- Watanabe K441, K442; Mt. Tara, K. Watanabe K444, Toma & T. Ohi-Toma K792. Aichi: Minami-Chita, K447; Seihi, near Saikai Bridge, K. Watanabe K606. K. Watanabe K556; Mt. Sanage, K. Watanabe K297, Kumamoto: Mt. Shodai, K. Watanabe K69, K70, K577; Owari-Asahi, T. Ohi-Toma K775; Mt. Kichijo, K72, K75. Miyazaki: Mt. Osuzu, K. Watanabe K50, T. Ohi-Toma K780; Taketoyo, T. Ohi-Toma K782. K54; Mt. Wanitsuka, T. Ohi-Toma K549; Hyuga, K. Watanabe K186, K187. Kagoshima: Kirishima, K. Appendix 2. Watanabe K43, K44; Mt. Takakuma, K. Watanabe List of the voucher specimens of Aristolochia K637; Mt. Kaimon, K. Watanabe K58; Yamakawa, shimadae with flowers in Fig. 5. K. Watanabe K61, K62, K65; Nejime, F. Shimozono Hyogo: Arima, near Sennenji Temple, K. Watanabe K301; Tanegashima Island, H. Murata K453. & T. Shiraiwa K356 (TI); Arima, Mizunashi River, K. Watanabe & T. Shiraiwa K357, K358 (TI); Mt. Rokko, Aristolochia kaempferi var. tanzawana = A. tanzawana K. Watanabe & T. Shiraiwa K365, K366, K367, K368, Ibaraki: Mt. Tsukuba, K. Watanabe K102, K103, K369 (TI); Inagawa, Ibuchi, T. Ushijima 8965 (SHO). K104, K105, K263. Kanagawa: Mt. Nabewari, K. Nagasaki: Sasebo, Kugurugi, K. Watanabe K437 Watanabe K97; Mt. Tanzawa, K. Watanabe-Toma & (TI); Goto Islands, Fukue-jima, I. Hiura s.n., 7 June T. Ohi-Toma K668, K669, K670; Sagamihara, Aone, 1969 (OSA); Hirado Island, Noko, K. Watanabe K617 T. Ohi-Toma K761; Fujino, K. Watanabe-Toma & T. (TI); Tsushima Island, Kami-Tsushima, K. Watanabe Ohi-Toma K649, K762; Ohara, T. Ohi-Toma K770; & T. Ohi-Toma K562 (TI). Saga: Tara, Nakayama Yamakita, Nakagawa, K. Watanabe-Toma & T. Ohi- Campground, K. Yonekura & R. Hirano 98733 (TUS). Toma K671. Yamanashi: Ochi Pass, K. Watanabe Kumamoto: Mt. Shodai, T. Hamada 15 (KYO, TUS). K116, K118; Abe Pass, T. Ohi-Toma & K. Sasamura

大井・東馬哲雄 a, 渡邉・東馬加奈 a, b, 邑田裕子 c, 邑田 a 仁 ：日本におけるオオバウマノスズクサとタンザワウ マノスズクサ（ウマノスズクサ科）の形態的変異 オオバウマノスズクサ Aristolochia kaempferi Willd. の他にも花形態（サイズ・形・色・模様）で明確に区別 は中国大陸にもあるとされていたが，日本固有種であ でき，さらに分布域が重ならないことが明らかになっ り，本州（関東以西）・四国・九州に分布する．本種は た．これらを踏まえてタンザワウマノスズクサを種のラ 形態変異が大きいが，葉裏中肋に開出毛を持つことで変 ン ク と し，A. tanzawana (Kigawa) Watanabe-Toma & 種タンザワウマノスズクサ var. tanzawana Kigawa が Ohi-Toma とする． 認識されていた．本研究において，A. kaempferi の分布 （a 東京大学大学院理学系研究科附属植物園， 域全域を対象に形態変異と生育場所の検討を行ったと b 武蔵高等学校中学校， ころ，var. kaempferi と var. tanzawana は葉裏中肋の毛 c 摂南大学薬学部）