J. Jpn. Bot. 90: 115–118 (2015)

Resurrection of Saionia (Thismiaceae)

Hiroyoshi Ohashi

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

(Accepted on November 1, 2014)

Four species of Oxygyne (Thismiaceae or Brumanniaceae s.l.) are known in Cameroon in West Africa and Japan in East Asia. The first Japanese species of Oxygyne was attributed to a new genus Saionia in 1975 and 1976, but the genus was invalidly published. Together with two recently discovered species of Oxygyne, the three Japanese species are distinct from the type species of Oxygyne from Cameroon in having campanulate flowers with bluish perianth, patent perianth-lobes, deflexed stamens of which the anthers are positioned below the base of the filament, and appendages on the style. Saionia is adopted here as a new genus for the three Japanese species. The generic name Saionia Hatus. and its type Saionia shinzatoi Hatus. were validly published in this paper. Two new combinations are proposed: Saionia hyodoi (C. Abe & Akasawa) H. Ohashi and S. yamashitae (Yahara & Tsukaya) H. Ohashi.

Key words: Brumanniaceae, Cameroon, Japan, mycoheterotrophic , new combinations, Oxygyne, resurrection, Saionia, Thismiaceae, validated genus.

The genus Oxygyne was studied while on the invalid ‘Saionia shinzatoi’. Abe and preparing a revised edition of new ‘Wild Flowers Akasawa (1989) presented a clear illustration of Japan’ (Ohashi et al. in press) as a member of the rare saprophyte, and pointed out such of the family Thismiaceae, though the family differences of their species from O. triandra as is included in in the APG III “the style with three capitate stigmas at the apex system. Oxygyne was published by Schlechter and three clavate appendices on the upper lateral (1906) based on a single specimen of Oxygyne side” and “the perianth-lamellae transversely triandra collected in Cameroon in West Africa. rectangular and forming an annular taenia on the The first species ofOxygyne in Japan was found throat” (on page 163 in Abe and Akasawa 1989). in northern Okinawa Island in the Ryukyus and Oxygyne hyodoi has not been found again. On was named Saionia shinzatoi Hatus. (Hatusima the other hand, ‘O. shinzatoi’ has recently been 1976). However, Hatusima cited two different rediscovered by Yokota of the University of the specimens as holotype. Therefore, the generic Ryukyus at the original locality. and specific names were invalid. Oxygyne was The fourth species of Oxygyne was reported adopted first in Japan by Abe and Akasawa by Yahara and Tsukaya (2008) in Yakushima isl., (1989) for their new mycoheterotrophic plant, southern Kyushu. This species was described Oxygyne hyodoi found in Ehime Prefecture, and illustrated in detail and the flower structure Shikoku. They also made a new combination, was compared with ‘Oxygyne shinzatoi’. O. shinzatoi (Hatus.) Abe & Akasawa. The According to the protologue of Oxygyne and combination is invalid, because it was based O. triandra by Schlechter (1906), the flower

—115— 116 植物研究雑誌 第 90 巻 第 2 号 2015 年 4 月 is fundamentally different from the Japanese and Corrected)’ in 1975 as ‘Saionia Shinzatoi species. The flower of O. triandra is tubular or Hatusima, gen. et sp. nov.-pl. 2’, with a deep campanulate with erect perianth-lobes (Fig. description of the species in Japanese. Hatusima 1, H in the protologue), and the filament, though (1976) provided the description of the genus and it was described as “filamentis erectis brevibus” species in Latin. This fulfiled the condition for in the protologue, is erect then recurved as Art. 39.1 in ICN Melbourne Code, but the name illustrated in Fig. 1, M, hence the anther is was typified by the invalidly published Saionia placed almost equal to a position at the base shinzatoi Hatus., which was based on two of the filament. The three species of Oxygyne holotypes. The generic name is validated here by known in Japan have campanulate flowers valid publication of the type, Saionia shinzatoi with patent perianth-lobes and descending Hatus. in this paper. filaments of which the anther is below the base According to Hatusima (1976), “the generic of the filament. These species have distinct name was dedicated to SAI-ON (1682–1762) appendages on the style near the apex, but no who [was a politician of the Ryukyu Kingdom] appendages are described in O. triandra (Fig. firstly established the forestry regulations in 1, K in the protologue). The flowers have a Okinawa and published the famous book ‘The dark brown tube with orange yellow lobes in Eight Volumes of Forestry Administration’ about O. triandra, whereas the Japanese species have two hundred years ago.” bluish flowers basically. These characteristic Hatusima (1976) considered Saionia as differences in color, morphology and structure representing a new subtribe Saioniinae (as of flowers suggest that a different pollination Saioneae Hatus. in J. Geobot. 24(1): 2. 1976, syndrome exists between Oxygyne triandra and nom. invalid.) within the tribe Thismieae in the Japanese Oxygyne species. the family Burmanniaceae (Jonker 1938). He The chromosomes were examined in characterized the subtribe by lacking an annular ‘Oxygyne shinzatoi (nom. invalid.)’ (Tsukaya et band at the throat of the perianth and by having al. 2007). The species has unique numbers, 2n declined stamens from the geniculate portion = 18 (x = 9), in the Burmanniaceae (including of the inner perianth lobe (Fig. 2, F in the Thismiaceae) so far known. This fact indicates a protologue of ‘Saionia shinzatoi’) in comparison distinct systematic position of the species in the with Oxygyninae Jonker, a monotypic subtribe family, although no other chromosomal data are based on Oxygyne. These characteristics of known in Oxygyne. Saionia are insufficient to separate it from Based on the differences in floral morphology Oxygyne even at generic rank as pointed and probable pollination syndromes together out by Abe and Akasawa (1989). Tsukaya with the disjunct distribution between West et al. (2007) treated ‘Oxygyne shinzatoi’as African and Japanese Oxygyne, the Japanese belonging to the subtribe Oxygyninae, but did species should be treated as a different genus. not mention ‘Saioniinae’. On the basis of the They are listed below with their bibliographies: new characterization of Saionia adopted in the present study, Saionia and Oxygyne should be Saionia Hatus. [in J. Geobot. 24(1): 2 included within the subtribe Oxygyninae in the (1976), cum descr. Latin., nom. invalid.], gen. tribe Thismieae. nov. Type: Saionia shinzatoi Hatus. [validated below]. Saionia shinzatoi Hatus. [in J. Geobot. 24(1): 2, fig. 1, 2 (1976), cum descr. Latin. et 2 The generic name first appeared invalidly holotypis, nom. invalid.], sp. nov. Holotype: in Hatusima’s ‘Flora of the Ryukyus (Added JAPAN. Okinawa Pref., Isl. Okinawa, Yona, April 2015 The Journal of Japanese Botany Vol. 90 No. 2 117 in the forest of Pinus lutchuensis and Schima Japanese name: Hina-no-bonbori (Abe and wallichii subsp. liukiuensis (T. Shinzato s.n., 18 Akasawa 1989). Sep. 1974, RYU). [Oxygyne shinzatoi (Hatus.) C. Abe & Saionia yamashitae (Yahara & Tsukaya) H. Akasawa in J. Jap. Bot. 64: 163 (1989), comb. Ohashi, comb. nov. invalid.]. Oxygyne yamashitae Yahara & Tsukaya in Japanese name: Hoshizaki-shakujyô Acta Phytotax. Geobot. 59: 99, f. 1, 2A–B, 3A– (Hatusima 1975). F (2008) [Type: JAPAN. Kagoshima Pref., Yaku Saionia shinzatoi Hatus. was invalidly Island, along the western branch of Futamata published by Hatusima (1975) first in Japanese River (K. Fuse, H. Yamashita & H. Ikeda s.n., in his ‘Flora of the Ryukyus (Added and 24 Oct. 2007, FU–holo.; TI–iso.)]. Corrected)’ in 1975 with citation of a single Japanese name: Yaku-no-hinahoshi (Yahara specimen, “T. Shinzato s.n., 18 Sept. 1974”. and Tsukaya 2008). He (1976) published the name again with Latin description for valid publication, but the name I wish to thank Professor M. Yokota of the is invalid by designation of two holotypes “T. University of the Ryukyus for confirmation of Shinzato s.n., 20 Sept. 1972 and 18 Sep. 1974, the type of Saionia shinzatoi in RYU. Professors RYU”. These specimens were collected by J. Murata of The University of Tokyo and Shinzato at the same locality in 1972 and 1974, H. Nagamasu of Kyoto University for their respectively. They were shown in his paper comments on a manuscript of the present paper. (1976) as “1972 specimen in Fig. 1 C and 1974 in Fig. 1 A, B, and D” (Hatusima 1976, on page References 8 in Japanese). One of the specimens was cited Abe C. and Akasawa Y. 1989. A new species of Oxygyne by Shimabuku (1990) in his “Check List” as (Burmanniaceae) found in Shikoku, Japan. J. Jap. Bot. “Okinawa: Yona, leg. Shinzato s.n. 1974. RYU” 64(6): 161–164. Jonker F. P. 1938. A monograph of the Burmanniaceae. without any indication of the type of the name. Meded. Bot. Mus. Herb. Rijks Utrecht 51: 1–279. The citation is not regarded to be the typification Hatusima S. 1976. Two new species of Burmanniaceae of Saionia shinzatoi (Art. 40.6 in Melbourne from Japan. J. Geobot. 24(1): 2–10. Code). The description of Saionia shinzatoi by Schlechter R. 1906. Burmanniaceae africanae. Bot. Jahrb. Syst. 38: 137–143. Hatusima is clearly based on the 1974 specimen. Shimabuku K. 1997. Check List Vascular Flora of the Hatusima (1976) recorded that he examined Ryukyu Islands. Revised edition. Kyushu-daigaku the flower in the 1974 specimen. Therefore, Syuppankai, Fukuoka. the 1974 specimen is recognized here for the Tsukaya H., Yokota M. and Okada H. 2007. Chromosome holotype of S. shinzatoi Hatus. characteristics of Oxygyne shinzatoi (Burmanniaceae) and its phylogenetic significance. Acta Phytotax. Geobot. 58(2/3): 100–106. Saionia hyodoi (C. Abe & Akasawa) H. Yahara T. and Tsukaya H. 2008. Oxygyne yamashitae, a Ohashi, comb. nov. new species of Thismiaceae from Yaku Island, Japan. Oxygyne hyodoi C. Abe & Akasawa in J. Acta Phytotax. Geobot. 59(2): 97–104. Yokoyama J., Koizumi Y., Yokota M. and Tsukaya H. Jap. Bot. 64: 161, f. 1 (1989) [Type: JAPAN. 2008. Phylogenetic position of Oxygyne shinzatoi Shikoku. Ehime Pref., Minamiuwa-gun, (Burmanniaceae) inferred from 18S rDNA sequences. Nishiumi-cho, in evergreen forest (Masaharu J. Pl. Res. 121(1): 27–32. Hyodo s.n., 9 Oct. 1988, TI–holo.)]. 118 植物研究雑誌 第 90 巻 第 2 号 2015 年 4 月

大橋広好:ホシザキシャクジョウ属(タヌキノショクダ イ科)の復活 平凡社『日本の野生植物 草本』(佐竹義輔他 1982) 移したが,その組み合わせは国際命名規約に反して発 を改訂するためにタヌキノショクダイ科のヒナノボン 表された Saionia shinzatoi Hatus. (1976) に基づくもので ボリ属 Oxygyne を検討した結果,日本産の種は別属と 非合法である.2008 年に屋久島から 4 番目のヒナノボ してホシザキシャクジョウ属 Saionia とする見解に至っ ンボリ属が報告され,ヤクノヒナホシと名付けられた た.1972 年と 1974 年に沖縄島国頭村琉球大学農学部 (Yahara and Tsukaya 2008).この間にホシザキシャクジ 与那演習林で 3 個の雄蕊をもつヒナノシャクジョウ科 ョウが再々発見され,その染色体が調べられ (Tsukaya の腐生植物が発見され,1975 年に新属新種のホシザキ et al. 2007),DNA 系統解析が行われた (Yokoyama et al. シャクジョウとして初島住彦『琉球植物誌(追加・訂 2008). 正)』909 ページに日本語で発表された.この発表では Oxygyne triandra の 原 記 載( ホ ロ タ イ プ も BGBM この学名は国際命名規約第 39 条(日本植物分類学会国 で画像が公開され 際命名規約邦訳委員会(訳・編集)2014.国際藻類・ ている)と日本産 3 種の記載とを比較すると,花の色 菌類・植物命名規約(メルボルン規約)2012.日本語 および形態と構造が異なり,両者の間には花粉媒介機構 版.北隆館,東京)によって,正式に発表されていない の違いが推測される.これらの違いに極端な隔離分布 と見なされる.この学名 Saionia shinzatoi Hatus. は翌年 をも考慮して日本産種は Oxygyne とは別属と認めたい. ラテン語で再発表されたが,2 点の標本をホロタイプと 本論文で,非合法な属名 Saionia とホシザキシャクジ して引用した (Hatusima 1976).このためこの発表でも ョウの学名 Saionia shinzatoi を正式に発表した.ヒナ 国際命名規約第 8.1 条,第 9.1 条および第 40 条によって, ノボンボリの学名は Saionia hyodoi (C. Abe & Akasawa) 属名と種名は正式に発表されなかったと判定される. H. Ohashi,ヤクノヒナホシの学名は Saionia yamashitae 1988 年に愛媛県でも 3 個の雄蕊をもつ腐生植物が発見 (Yahara & Tsukaya) H. Ohashi と 変 更 す る. ま た, され,Abe and Akasawa はこれを Oxygyne Schltr. の新 Hatusima (1976) の設立したヒナノシャクジョウ科タヌ 種ヒナノボンボリとし,その属名をヒナノボンボリ属と キノショクダイ連 tribe Thismieae の新亜連 Saioniinae は 呼んだ.Oxygyne はカメルーンで発見された O. triandra 非合法に発表された属名に基づいたものであり,その名 に基づいて 1906 年に発表され,形容語のように 3 個の 称は非合法であるが,その内容を Oxygyninae と同一と 雄蕊をもつことを大きな特徴として,ヒナノシャクジ 見なした. ョウ科の新属とされたものであった.Abe and Akasawa (東北大学植物園津田記念館) (1988) はホシザキシャクジョウもヒナノボンボリ属に