J. Jpn. Bot. 91: 79–83 (2016)

Correct Name of the Japanese Endemic Species Pertya trilobata ()

Tetsuo Ohi-Toma* and Jin Murata

Botanical Gardens, Graduate School of Science, the University of Tokyo, Bunkyo-ku, Tokyo, 112-0001 JAPAN *Corresponding author: [email protected]

(Accepted on October 14, 2015)

For the name of one of the Japanese endemic species of Pertya sect. Macroclinidium (Asteraceae) or the Macroclinidium, the adjective trilobus has been used as the specific epithet, as P. triloba or M. trilobum. However, this adjectival epithet should be treated as an orthographical variant of trilobatus used in M. trilobatum Makino, because they are confusingly similar epithets based on the same type. Thus, its correct name under Pertya is P. trilobata (Makino) Makino based on M. trilobatum, which has priority as the validly published name. Concurrently, on the basis of cytological, morphological and preliminary molecular evidences, P. ×koribana (Nakai) Makino & Nemoto, which has been known as a putative hybrid between P. trilobata and P. robusta, is to be treated as P. trilobata having unlobed leaves.

Key words: Asteraceae, Macroclinidium trilobum, Macroclinidium trilobatum, Macroclinidium koribanum, nomenclature, Pertya ×koribana, Pertya triloba, Pertya trilobata.

Pertya Sch. Bip. (Asteraceae) consists of nested in Pertya (Ohi-Toma unpublished). ca. 25 species from Sino-Japanese region to For the name of one of the Japanese endemic Thailand and Afghanistan (Katinas et al. 2008, perennial herbs, the adjective trilobus has Gao and Hind 2011). Most of them, which are been used as a specific epithet: Pertya triloba shrubs or subshrubs, are classified into section (Makino) Makino (Kitamura 1981, Yonekura Pertya. Three Japanese endemic perennial 2012), Pertya triloba (Makino) Matsum. herbs constitute sect. Macroclinidium (Maxim.) (Koyama 1995), Macroclinidium trilobum Makino (Kitamura 1938, 1940, Koyama 1995). (Makino) Makino (Katinas et al. 2008), or This section has often been recognized as an M. trilobum Makino (Freire 2012). However, independent genus Macroclinidium Maxim. this adjectival epithet should be treated as an (Makino 1898a, 1900b, Nakai 1912, Nemoto orthographical variant of trilobatus used in M. 1936, Hara 1952, Suzuki 1970, Katinas et al. trilobatum Makino (1894). Here, we review the 2008). Recently, Freire (2012) indicated the nomenclature of the taxon (Tables 1–3), and separation of Macroclinidium from Pertya based showed that its correct name is Pertya trilobata on morphological cladistic analysis. However, (Makino) Makino (1900a: 144) based on M. a preliminary molecular phylogeny indicated trilobatum. that Macroclinidium is not monophyletic and is

—79— 80 植物研究雑誌 第 91 巻 第 2 号 2016 年 4 月 Makino Makino Makino (Makino) Matsum. (Makino) Makino and Nemoto Nemoto and Makino 1931) (1925, trilobatum M. P. triloba P. M. trilobum trilobum M. P. triloba P. Koyama (1995) Koyama Makino Makino Makino Makino Makino (Makino) Matsum., [comb. superfl.] [comb. Matsum., (Makino) Franch. Makino Franch. P. triloba P. fauriei P. P. triloba P. P. triloba P. M. trilobum M. Freire (2012) Freire Makino P. fauriei P. M. trilobatum M. Makino (1900b) Makino M. trilobum M. triloba A. (Makino) Makino (Makino) M. trilobum M. P. triloba P. Kitamura (1981) Kitamura Makino Makino Franch. Makino Makino (Makino) Makino (Makino) Makino, [basionym] Makino, M. trilobum M. P. fauriei fauriei P. M. trilobatum M. Makino (1900a) Makino triloba P. A. triloba A. Makino A. triloba A. Katinas et al. (2008) al. et Katinas trilobum M. (Makino) Makino (Makino) Franch. Makino Makino Makino M. trilobum M. P. fauriei P. P. triloba P. Ohwi (1953, 1965a, 1965b) 1965a, (1953, Ohwi M. trilobatum M. A. triloba A. M. trilobum M. Makino (1898b) Makino Makino Makino, Makino, Makino Franch. Makino Makino, [nom. seminud.] [nom. Makino,

Makino ] Franch. Makino, [nom. nud.] [nom. Makino, Makino (Makino) Matsum. (Makino) Makino, Makino, [nom. nud.] [nom. trilobum M. A. triloba A. P. fauriei P. P. triloba P. Kitamura (1940) Kitamura P. fauriei fauriei P. P. triloba P. P. triloba P. M. trilobatum M. A. triloba A. Hara (1952), Suzuki (1970) Suzuki (1952), Hara M. trilobum M. nom. nud. nom. [ A. triloba A. M. trilobatum M. Makino (1898a) Makino . The boldface is used for the accepted name and others are its synonyms its are others and name accepted the for used is boldface The . Makino, Makino, Makino Pertya. Pertya. Pertya. Makino, Makino, = = Makino Franch. = = = = Makino P. P. Makino . The boldface is used for the accepted name and others are its synonyms its are others and name accepted the for used is boldface The . P. P. P. P.

, , , , , , ] Makino, Makino, M. trilobatum M. nud.] [nom. trilobum M. A. triloba A. nud.] [nom. P. fauriei P. Kitamura (1938) Kitamura P. triloba P. Nemoto (1936) Nemoto M. trilobum M. Macroclinidium trilobum Macroclinidium nom. nud. nom. Pertya triloba Pertya A. triloba A. [ M. trilobatum M. Makino (1894) Makino Macroclinidium Macroclinidium Macroclinidium ======M. M. M. M. M. M. , , Makino , , , , Makino Makino Makino ] Makino Makino Franch. Makino, Makino, Makino (Makino) Matsum. (Makino) Franch. Ainsliaea Ainsliaea ======nom. nud. nom. P. fauriei P. A. A. P. triloba P. M. trilobatum M. A. triloba A. Table 3. Treatments as as Treatments 3. Table Nakai (1912) Nakai M. trilobum M. A. A. A. triloba A. [ Makino (1892) Makino M. trilobum M. P. fauriei P. M. trilobatum M. A. triloba A. A. A. Table 2. Treatments as as Treatments 2. Table Matsumura (1902) Matsumura P. triloba P. Table 1. Changes in treatments by T. Makino. The boldface is used for the accepted name and others are its synonyms synonyms its are others and name accepted the for used is boldface The Makino. T. by treatments in Changes 1. Table April 2016 The Journal of Japanese Botany Vol. 91 No. 2 81

Pertya trilobata (Makino) Makino Makino, whose basionym is A. triloba, Freire Makino (1892) proposed Ainsliaea triloba (2012) did not regard either A. triloba nor M. Makino, whose Japanese name is “Oyari- trilobatum as the basionym of M. trilobum. haguma,” as a new species candidate without a However, Makino (1898b) listed A. triloba and description, diagnosis, or reference to a previous M. trilobatum with their references as synonyms publication. In it, he cited two specimens (Iwaki, of M. trilobum and cited three specimens, of T. Makino s.n., and Iwashiro, T. Makino s.n.), which two are the syntypes of M. trilobatum. which are kept in the herbarium TI. Two years Therefore, M. trilobatum and M. trilobum are later, when Makino (1894) validly published the treated as orthographical variants, because new species providing a description in Japanese, they are confusingly similar names based on he adopted Macroclinidium trilobatum Makino the same type (Art. 61.5 of the ICN, McNeill rather than A. triloba. However, Hara (1952) 2012). M. trilobatum in Makino (1894: 302) has and Suzuki (1970) regarded M. trilobatum as priority as the validly published name. Here, the a nomen seminudum because the description specimen, which Freire (2012) designated as the was given in Japanese only. Katinas et al. lectotype of M. trilobum from three specimens (2008) recognized A. triloba (1892) as a validly cited in Makino (1898b), is one of the syntypes published name, but it was not accompanied of M. trilobatum. Therefore, the lectotype of M. by a description. Recently, when Freire (2012) trilobatum was designated by Freire (2012). reviewed the genus Macroclinidium, she When Makino (1900a) treated additionally mentioned that A. triloba in Makino Macroclinidium as a section of the genus Pertya, (1892) is a nomen nudum and that M. trilobatum he published P. triloba Makino as nomen novum. in Makino (1894) is apparently a synonym of A. After Makino (1898b, 1900a), many authors triloba, probably overlooking the description in used P. triloba Makino or P. triloba (Makino) Japanese. Makino and included A. triloba Makino, M. According to the mention in Makino (1894), trilobum and/or M. trilobatum in the synonymic he accepted M. trilobatum, not A. triloba, as list (Tables 1–2). Of them, Kitamura (1938, a new species (Art. 36.1 and 38.1 of the ICN, 1940) regarded A. triloba and M. trilobatum McNeill 2012). In addition, when Makino as nomina nuda. However, P. triloba is to be (1898a) published a diagnosis of M. trilobatum treated as an orthographical variant, because Makino in both English and Japanese, he Makino (1900a) cited A. triloba Makino, M. cited Makino (1894) as the protologue of the trilobatum Makino, M. trilobum Makino with species. Here, although no type was designated their references as synonyms. Therefore, Pertya or definitely indicated in the protologue, the triloba Makino (1900a: 144) is based on M. two specimens, which were associated with A. trilobatum (1894: 302), and its correct name is to triloba in Makino (1892), are to be the syntypes be P. trilobata (Makino) Makino (1900a: 144). of M. trilobatum. Matsumura (1902) published Pertya triloba On the other hand, only four months after (Makino) Matsum. (Table 2). Koyama (1995) Makino (1898a), Makino (1898b) published adopted the name and cited M. trilobum Makino Macroclinidium trilobum Makino with a as a synonym. However, P. triloba (Makino) detailed description in English based on “Illustr. Matsum. is a later isonym of P. triloba (Makino) Flor. Jap. I. ined. n. 12, tab LXXI”, whose plate Makino (Art. 6, Note 2 of the ICN, McNeill with Japanese description was published by 2012). Makino (1934). After that, M. trilobum has often been recognized (Table 3). Although Katinas Pertya koribanum et al. (2008) recognized M. trilobum (Makino) As putative hybrids related to P. trilobata, 82 植物研究雑誌 第 91 巻 第 2 号 2016 年 4 月

P. ×koribana (Nakai) Makino & Nemoto and putative hybrid is to be treated as P. trilobata P. ×suzukii Kitam. are known (cf. Koyama having unlobed leaves. 1995, Yonekura 2012, Freire 2012). Of them, P. ×suzukii is morphologically and genetically Taxonomic treatment confirmed as a natural hybrid between P. Pertya trilobata (Makino) Makino in trilobata and P. rigidula (Miq.) Makino (Ohi- Bot. Mag. (Tokyo) 14: 144 (1900), ‘triloba’; Toma unpublished). However, it is not supported Matsum. in Bot. Mag. (Tokyo) 16: 18 (1902), that P. ×koribana is a natural hybrid. isonym, ‘triloba’. [≡ Ainsliaea triloba Makino Nakai (1912) published Macroclinidium in Bot. Mag. (Tokyo) 6: 55 (1892), nom. nud.] ≡ koribanum Nakai, whose Japanese name is Macroclinidium trilobatum Makino in Bot. Mag. “Sendai-haguma,” based on capitula with one (Tokyo) 8: 302 (1894). ≡ M. trilobatum Makino flower and ovate-acuminate leaves. Makino and in Bot. Mag. (Tokyo) 12: 78 (1898), ‘trilobum’. Nemoto (1925) treated it as Pertya koribana Lectotype (designated by Freire 2012): JAPAN. (Nakai) Makino & Nemoto. Makino (1933) Iwaki, Yanagibashi, 16 Aug. 1890, T. Makino published it as P. triloba Makino var. koribana s.n. (TI!). Syntype: JAPAN, Iwaki, Moniwa, 3 (Nakai) Makino, whose Japanese names are Sept. 1890, T. Makino s.n. (TI!). “Sendai-haguma” or “Maruba-oyarihaguma.” = Pertya fauriei Franch. in Mém. Herb. Kitamura (1938) follows Makino (1933), Boiss. 14: 2, t.1 (1900). Lectotype (designated but he mentioned, “If the species is a hybrid by Freire 2012): JAPAN. Yamagata, U. J. Faurie between P. robusta Beauverd and P. triloba 4289 (P [P00703240, digital image!]; iso-, P Makino, the name may be P. koribana (Nakai) [P00703239, digital image!; P00703241, digital Makino et Nemoto.” Honda (1939) treated it image!], KYO [KYO00081206, digital image!]). as M. trilobum Makino var. koribanum (Nakai) Syntype: JAPAN. Sendai, U. J. Faurie 6741 (P, Honda. Kitamura (1940), Ohwi (1953, 1965a, not seen). 1965b), and Koyama (1995) regarded P. = Macroclinidium koribanum Nakai in koribana (Nakai) Makino & Nemoto as a hybrid Bot. Mag. (Tokyo) 26: 249 (1912), syn. nov. between P. robusta and P. trilobata. Hara (1952), ≡ P. koribana (Nakai) Makino & Nemoto, Fl. Suzuki (1970) and Freire (2012) treated it as M. Japan, ed. 2 (Makino & Nemoto): 83 (1925), ×koribanum Nakai. pro hybr. ≡ P. triloba Makino var. koribana The putative hybrid resembles Pertya (Nakai) Makino in J. Jap. Bot. 8: 46 (1933). ≡ trilobata in capitula with one flower, but M. trilobum Makino var. koribanum (Nakai) the hybrid having leaves ovate to elliptic is Honda, Nom. Pl. Jap.: 510 (1939). Type: distinguished from P. trilobata having 3-lobed JAPAN. Sendai, Mt. Taihakusan, Oct. 1901, K. leaves. Although P. trilobata is rarely sympatric Kôriba s.n. (TI–holotype!). with P. robusta, P. trilobata having unlobed leaves are often observed in populations of P. References trilobata (Suzuki 1975, Ohi-Toma personal Freire S. E. 2012. Systematics of the Japanese observation). Based on the chromosome Macroclinidium and ×Macropertya (Asteraceae, Pertyoideae, Pertyeae) and their phylogeny inferred numbers of the putative hybrid (2n = 28) and from morphology. Syst. Bot. 37: 554–572. morphology, Suzuki (1975) did not support that Gao T. G. and Hind D. J. N. 2011. Pertya (Asteraceae). it is a natural hybrid between P. robusta (2n = In: Wu Z. Y., Raven P. H. and Hong D. Y. (eds.), Flora 26) and P. trilobata (2n = 28). In addition, a of China 20–21: 27–31. Science Press, Beijing and Missouri Botanical Garden Press, St. Louis. preliminary molecular analysis does not support Hara H. 1952. Enumeratio Spermatophytarum that it is a hybrid between these species (Ohi- Japonicarum, Part 2. Iwanami Shoten, Tokyo. Toma, unpublished). It is concluded that the Honda M. 1939. Nomina Plantarum Japonicarum. April 2016 The Journal of Japanese Botany Vol. 91 No. 2 83

Sanseido, Tokyo (in Japanese). shokubutsu-Soran publishing Association, Tokyo (in Katinas L., Pruski J., Sancho G. and Tellería M. C. 2008. Japanese). The subfamily Mutisioideae (Asteraceae). Bot. Rev. Makino T. and Nemoto K. 1931. Flora of Japan, 2nd ed. 74: 469–716. Shunyodo Shoten, Tokyo (in Japanese). Kitamura S. 1938. Les Mutisieae du Japon (II) J. Jap. Bot. Matsumura J. 1902. Some rare in Japan. Bot. Mag. 14: 377–378 (in Japanese). (Tokyo) 16: 17–18. Kitamura S. 1940. Compositae Japonicae, pars secunda. McNeil J., Barrie F. R., Back W. R., Demoulin V., Greuter Mem. Coll. Sci. Kyoto Imp. Univ., Ser. B, Biol. 15: W., Hawksworth D. L., Heredeen P. S., Knapp S., 285–447. Marhold K., Prado J., Prud’homme van Reine W. Kitamura S. 1981. Pertya. In: Satake Y., Ohwi J., Kitamura F., Smith G. E., Wiersema J. H. and Turland N. J. S., Watari S. and Tominari T. (eds.), Wild Flowers of (eds.) 2012. International Code of Nomenclature for Japan, Herbaceous plants (including dwarf subshrubs) algae, fungi, and plants (Melbourne Code): Adopted III: 209–210. Heibonsha, Tokyo (in Japanese). by the Eighteenth International Botanical Congress Koyama H. 1995. Pertya. In: Iwatsuki K., Yamazaki T., Melbourne, Australia, July 2011. Regnum Vegetabile Boufford D. E. and Ohba H. (eds.), Flora of Japan IIIb: 154. Königstein: Koeltz Scientific Book. http://www. 165–167. Kodansha, Tokyo. iaapt-taxon.org/nomen/main.php. Makino T. 1933. A contribution to the knowledge of the Nakai T. 1912. Notulae ad plantas Japoniae et Koreae VI. Flora of Nippon (Continued from p. 40). J. Jap. Bot. 8: Bot. Mag. (Tokyo) 26: 247–250. 43–46. Nemoto K. 1936. Flora of Japan Supplement. Shunyodo Makino T. 1934. Nihon shokubutsu zusetsu shû. Shoten, Tokyo (in Japanese). Seibundo-Sinkosha, Tokyo (Japanese). Ohwi J. 1953. Flora of Japan. Shibundo, Tokyo (in Makino T. 1892. Notes on Japanese Plants, XV. Bot. Mag. Japanese). (Tokyo) 6: 45–56. Ohwi J. 1965a. Flora of Japan, rev. ed. Shibundo, Tokyo (in Makino T. 1894. Miscellaneous notes on the plants of “Yo- Japanese). jo-syo-oku”, no. 10. Bot. Mag. (Tokyo) 8: (300)–(305) Ohwi J. 1965b. Flora of Japan, Eng. ed. Smithsonian (in Japanese). Institution, Washington, D.C. Makino T. 1898a. Contributions to the study of the Flora of Suzuki M. 1970. Biosystematic studies on Macroclinidium Japan, V. Bot. Mag. (Tokyo) 12: 192–196. (Compositae–Mutisieae) I. Geographical distribution Makino T. 1898b Plantae Japonenses novae vel minus and ecology. Bull. Fac. Educ., Ibaraki Univ., Nat. Sci. congnitae (Continued from p. 74). Bot. Mag. (Tokyo) 20: 233–259. 12: 77–82. Suzuki M. 1975. Species of Macroclinidium (Compositae) Makino T. 1900a. Plantae Japonenses novae vel minus and their relationship. Biol. Sci. (Seibustu-Kagaku) 27: congnitae (Continued from p. 56). Bot. Mag. (Tokyo) 180–188. 14: 144. Yonekura K. and Murata J. 2012. An Enumeration of the Makino T. 1900b. Plate XLI, Oyari-haguma. Phan. Pter. Vascular Plants of Japan. Hokuryukan, Tokyo (in Jap. Icon. 1: t. 45 (in Japanese). Japanese). Makino T. and Nemoto K. 1925. Flora of Japan. Nippon-

大井・東馬哲雄，邑田 仁：日本固有種オヤリハグマ（キ ク科）の正名 キク科コウヤボウキ属カシワバハグマ節（またはカシ ボウキ属のもとでは，もっとも早く正式に発表された ワバハグマ属）の日本固有種オヤリハグマは，最近では， Macroclinidium trilobatum Makino に 基 づ く Pertya Pertya triloba または Macroclinidium trilobum のよう trilobata (Makino) Makino が正名である．また，オヤ に，形容詞 trilobus を種形容語とする学名が用いられて リハグマとカシワバハグマの雑種とされていたセンダ いる．しかし，これらは，類似した形容詞 trilobatus を イハグマ (P. ×koribana) は，染色体数や外部形態，予備 種形容語とする M. trilobatum Makino と「同じタイプ 的な DNA 解析においては，雑種とする積極的な根拠は に基づいた混同しやすい類似の学名」であるため，両 無く，オヤリハグマの無裂葉型として扱うのが適当であ 者は正字法上の異形体として扱われる（国際藻類・菌 る． 類・植物命名規約 第 61.5 条）．したがって，コウヤ （東京大学大学院理学系研究科附属植物園）