ISSN 1346-7565 Acta Phytotax. Geobot. 71 (2): 163–169 (2020) doi: 10.18942/apg.201918

A New Peloric Form of erecta () from Japan

1,* 2 3,† 3,4,‡ Hiroshi Hayakawa , Kenji Suetsugu , Shohei Fujimori , Takuto Shitara , Jun 5 6 Yokoyama and Tomohisa Yukawa

1Museum of Natural and Environmental History, Shizuoka, 5762 Oya, Suruga-ku, Shizuoka 422-8017, Japan. *[email protected] (author for correspondence); 2Department of Biology, Graduate School of Science, Kobe University, 1-1 Rokkodai, Nada-ku, Kobe, Hyogo 657-8501, Japan; 3Graduate School of Life and Environmental Sciences, University of Tsukuba, 1-1-1 Tennodai, Tsukuba, Ibaraki 305-8572, Japan; 4Iriomote Sta- tion, Tropical Biosphere Research Center, University of the Ryukyus, 870 Uehara, Taketomi-cho, Yaeyama-gun, Okinawa 907-1541, Japan; 5Faculty of Science, Yamagata University, Kojirakawa, Yamagata 990-8560, Japan; 6Tsukuba Botanical Garden, National Museum of Nature and Science, 1-1, Amakubo 4, Tsukuba, Ibaraki 305-0005, Japan; †Present address: Department of Horticulture, Kochi Prefectural Makino Botanical Garden, 4200-6 Godai- san,, Kochi 781-8125, Japan; ‡Present address: Institute of Agriculture, Tokyo University of Agriculture and Technol- ogy, 3-5-8 Saiwai, Fuchu-shi, Tokyo 183-8509, Japan

A new peloric form of (Thunb.) Blume (Orchidaceae) is reported from Hokkaido, Honshu, and Shikoku, Japan. Cephalanthera erecta f. pelorica Hiros. Hayak. & Suetsugu has an incon- spicuous lip and stigmas positioned at the apex of the column. The flowers of C. erecta f. pelorica re- semble those of the probable peloric form of C. erecta (i.e., C. erecta var. oblanceolata N. Pearce & P. J. Cribb, C. nanlingensis A. Q. Hu & F. W. Xing) and C. longifolia (L.) Fritsch. f. conformis Suetsugu & Hiros. Hayak. However, C. erecta f. pelorica can be distinguished from C. erecta var. oblanceolata, which has a swollen organ on the abaxial surface of the column apex (vs. no swollen organ); C. nanlin- gensis, which has two staminodes (vs. five staminodes), andC. longifolia f. conformis, which has elliptic, coriaceous leaves (vs. lanceolate, papyraceous leaves).

Key words: Cephalanthera, lip, peloria,

Orchidaceae, one of the largest families of an- cal explorations in Japan (Suetsugu 2013, Hay- giosperms with over 26,000 species in 880 gen- akawa et al. 2014, 2016b, Suetsugu & Hayakawa era (Stevens 2001 onwards), exhibits consider- 2019). Peloric mutants of other taxa have also able morphological and ecological variation been observed to self-pollinate (e.g., Hayakawa et among the species. The specialized orchid lip dif- al. 2014, Suetsugu 2015). We therefore suspect fers from other perianth segments and aids in out- that additional self-pollinating peloric taxa might crossing by pollinators. The lip of peloric mu- be found in further surveys. tants, however, is undifferentiated and closely re- Cephalanthera Rich. comprises approximate- sembles the petals and/or sepals. Such flowers ex- ly 15 mixotrophic or fully mycoheterotrophic hibit radial symmetry unlike the normally zygo- species that occur throughout Europe, northern morphic flowers. Peloric mutants may be less Africa, eastern Asia, southeast Asia, and western common in nature because of the absence of a North America (Luer 1975, Satomi 1982, Yuka- specialized lip and their reduced reproductive wa 2015a). Seven peloric taxa have been reported success via outcrossing. Peloric mutants may in the genus (Pearce et al. 2001, Hu et al. 2009, therefore become extinct before being discovered Chen et al. 2010, Jin et al. 2011, Xiang et al. 2012, by botanists (Hayakawa et al. 2016a). We have, Hayakawa et al. 2014, 2016b, Suetsugu & Hay- however, observed self-pollination among newly akawa 2019). discovered peloric orchids during recent botani- Cephalanthera erecta (Thunb.) Blume, a 164 Acta Phytotax. Geobot. Vol. 71

Fig. 1. Cephalanthera erecta f. pelorica. A-B: Habitat of C. erecta f. pelorica. A: Tsukuba, Ibaraki Prefecture. B: Utsunomiya, Tochigi Prefecture. C–F: Column and anther of C. erecta f. pelorica. C: Takao, Tokyo Metropolitan. D: Yokoyama, Kanagawa Prefecture (K. Suetsugu s.n., 2/V/2015, KYO-holotype). E: Utsunomiya, Tochigi Prefecture (H. Hayakawa & M. Takashima s.n., 8/V/2015). F: Tsukuba, Ibaraki Prefecture (S. Fujimori F130506b, 6/V/2013, TNS). Arrows and white and black arrowheads indicate stigmas, staminodes, and pollinia, respectively. leafy mixotrophic orchid in the Himalaya, China, but did not formally propose a scientific name for Taiwan, the Korean peninsula, Japan (Hokkaido, them. Takashima & Kurihara (2016) also report- Honshu, Shikoku, and Kyushu), and the Kuril Is- ed peloric mutants of C. erecta from Japan and lands (Yukawa 2015a, b), has been reported to be identified them as C. erecta var. oblanceolata N. self-pollinating (Tanaka 1965). In Japan, Yanaga- Pearce & P. J. Cribb described from Bhutan. wa (1982) reported and illustrated peloric mu- During our botanical explorations, we found tants of Cephalanthera erecta from Japan and six populations of peloric mutants of C. erecta gave them the Japanese name "Yabitsu-Ginran”, (Figs. 1 & 2). We observed three sympatric popu- June 2020 Hayakawa & al. A New Peloric Form of Cephalanthera erecta 165

Fig. 2. Distribution of Cephalanthera erecta f. pelorica in Japan. Circles and squares indicate sites of C. erecta f. pelorica from herbarium and field surveys, respectively. Gray and black squares indicateC. erecta f. pelorica populations with and with- out C. erecta f. erecta, respectively. Figure edited from map generated by MapMap v.6.0 software (Kamada 2005) . lations of peloric and typical forms of Cephalan- has 3–5 inconspicuous staminodes. The five sta- thera erecta in Abiko City, Tsukuba City, and Os- minodes differ in size among populations of the hino Village (Fig. 2). The other three populations peloric mutants (Fig. 1). of peloric mutants were not sympatric with the The Japanese peloric forms are sometimes typical form. From a survey of 21 herbaria and sympatric with the typical form. They also differ our field survey, we discovered 39 specimens of morphologically from the peloric forms of other peloric mutants of C. erecta from 17 prefectures taxa of Cephalanthera (See Taxonomic note). in Japan (Fig. 2, Appendix). The peloric mutants Thus, the Japanese peloric mutant of C. erecta are mainly on the Pacific Ocean side of eastern should be formally recognized. Based on our Japan and in the Tsurugi Mountains and Okaya- findings, we conclude that the Japanese mutant is ma Prefecture in western Japan (Fig. 2). a form of C. erecta, which we here name Cepha- Specimens of the peloric mutant have been lanthera erecta (Thunb.) Blume f. pelorica. collected in Japan from 1928 until the present and most have been identified as the typical form of Cephalanthera erecta (Thunb.) Blume f. C. erecta or C. longifolia (L.) Fritsh. The lip of pelorica Hiros. Hayak. & Suetsugu, forma nov. the peloric mutant is similar to the dorsal sepal or —Figs. 1, 3, 4(A). lateral petals and the lateral sepals resemble the Typus. Yokoyama, Kanagawa Prefecture, Japan, Ken- dorsal sepal. The lateral sepals of the typical form ji Suetsugu s.n., 2/V/2015 (holo-, KYO) are lanceolate (Figs. 3 & 4). The peloric mutant also has stigmas positioned at the apex of the col- Herbs, terrestrial, mixotrophic, 5–40 cm tall, umn (not on the adaxial surface) and five stami- erect. Leaves 3–6, alternate, ovate-lanceolate, nodes around the column apex. The typical form 3–9 cm long, 1–4 cm wide, apex acute-subacute, 166 Acta Phytotax. Geobot. Vol. 71

Fig. 3. Illustration of Cephalanthera erecta f. pelorica (S. Fujimori F130506b, 6/V/2013, TNS). A: Ventral view of column. B: Clockwise from top: dorsal sepal, lateral petal, lateral sepal, lip. Bar = 1mm. Drawn by S. Fujimori. veins 3–5, glabrous. Dorsal sepal oblanceolate, var. erecta (Pearce et al. 2001), because popula- 7−9 mm long, apex obtuse. Lateral sepal oblan- tions of C. erecta var. erecta in Yunnan, China ceolate 2–3 mm wide, apex obtuse. Lateral petals are in the distribution limit of the western side of oblanceolate, 2–3 mm wide, apex obtuse. Lip ob- C. erecta var. erecta (Chen et al. 2010) and it has lanceolate, petal-like, entire, unlobed, apex ob- yet to be collected from Bhutan. tuse. Stigmas positioned at column apex. Stami- Based on the description and illustration by nodes 5. Floral bracts lanceolate, acute, lower flo- Pearce et al. (2001) and our observation of the ho- ral bracts foliaceous, lanceolate, acute. Flowers lotype (H. Kanai et al. s.n., 10/V/1967, TI), C. white, May to June. erecta var. oblanceolata has a swollen organ on the abaxial surface of the column apex (Fig. 5). Japanese name. Yabitsu-Ginran (Yanagawa Cephalanthera erecta f. pelorica lacks a swollen 1982) organ. Takashima & Kurihara (2016) reported C. Distribution. Hokkaido, Honshu, and Shi- erecta var. oblanceolata from Utsunomiya, Toch- koku, Japan, and probably South Korea (See be- igi Prefecture, Japan. We collected a peloric mu- low) tant at the same locality (Fig. 2) and identified it as C. erecta f. pelorica [Fig. 1(B & E)]. Cepha- Taxonomic note. Cephalanthera erecta var. lanthera erecta var. oblanceolata was reported oblanceolata, C. nanlingensis A.Q. Hu & F.W. from South Korea (Lee et al. 2009), but the illus- Xing, and C. longifolia (L.) Fritsch. f. conformis tration does not show a swollen organ on the col- Suetsugu & Hiros. Hayak. also have floral mor- umn. The South Korean peloric mutant is similar phologies similar to Cephalanthera erecta f. pe- to C. erecta f. pelorica rather than to C. erecta lorica. var. oblanceolata. A peloric mutant of C. erecta var. oblanceo- Cephalanthera erecta var. oblanceolata, re- lata was reported from Bhutan (Pearce et al. ported by Subedi et al. (2018) from Nepal may be 2001). Its distribution is distinct from C. erecta f. conformis (See be- June 2020 Hayakawa & al. A New Peloric Form of Cephalanthera erecta 167

Fig. 4. Perianth segments of Cephalanthera erecta. A: C. erecta f. pelorica (K. Suetsugu s.n., 2/V/2015, KYO-holotype). B: C. erecta f. erecta. Photographed by T. Shitara.

pelorica in having non-foliaceous floral bracts and two staminodes. Cephalanthera longifolia f. conformis is a pe- loric mutant from Japan (Hokkaido), Bhutan, and probably Nepal (see above) (Suetsugu & Hayaka- wa 2019). Cephalanthera longifolia f. longifolia can be distinguished from C. erecta f. erecta by having a non-protruding spur (vs. protruding spur), lanceolate, coriaceous leaves (vs. elliptic, papyraceous) (Yukawa 2016, Yokota et al. 2016, Suetsugu & Hayakawa 2019). Peloric mutants of C. longifolia and C. erecta are similar in their flowers because of the non-specialized lip, but they can be distinguished by leaf shape and tex- ture (Suetsugu & Hayakawa 2019). Fig. 5. Column of Cephalanthera erecta var. oblanceolata (H. Kanai et al. s.n., 10/V/1967, TI-holotype). A: Abaxi- al surface, B: Adaxial surface. Arrowhead indicates swollen organ on abaxial surface of column apex. We thank the curators of the AKPM, CBM, Ehime Pre- fectural Science Museum, FKSE, Fukui City Museum of Natural History, HYO, INM, KPM, KSNHM, KURA, KYO, MAK, MBK, OSA, SMNH, SPMN, TI, TNS, low). We therefore believe that Cephalanthera TOCH, TOYA, and TUS herbaria for herbarium access. erecta var. oblanceolata is only in Bhutan. We thank Dr. M. Takashima for guiding us in Tochigi Cephalanthera nanlingensis is a peloric Prefecture, Messrs. T. Kaneko, R. Kurashige, S. Nemoto, from the Nanling Mountains, China (Hu et al. Y. Yamashita, H. Yamaji for providing useful information regarding botanical explorations, and the Tochigi Prefec- 2009). Although C. nanlingensis has not been tural Office and Tochigi Central Park for permitting our found to be sympatric with C. erecta (Hu et al. botanical exploration. We would like to thank Editage 2009), Chen et al. (2010) hypothesized that it is (www.editage.jp) for editing the English language of our probably derived from C. erecta. Cephalanthera manuscript. We would like to thank two anonymous re- nanlingensis is distinguished from C. erecta f. viewers for their constructive comments on the paper. 168 Acta Phytotax. Geobot. Vol. 71

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Received July 18, 2018; accepted October 15, 2019 June 2020 Hayakawa & al. A New Peloric Form of Cephalanthera erecta 169

Appendix. Specimens examined. Japan: Hokkaido: H. Yamaji & H. Akutagawa 2024 Takashima s.n., 8/V/2015 (TOCH); Saitama: R. (TUS), S. Kigawa 80042 (KPM), Y. Goda 68 (KYO); Morimoto s.n. (MBK0101071) (MBK); Gunma: K. Aomori: K. Yonekura 13253 (TUS); Akita: H. Yoneda Masuda s.n., 12/May/2007 (KYO), K. Masuda s.n., 26/ 34881 (AKPM), H. Yoneda 34882 (AKPM), J. Takada May/2007 (KYO); Chiba: T. Kaneko s.n., 28/IV/2018 32406 (AKPM), M. Hatayama 44883 (AKPM), unknown (SPMN); Yamanashi: K. Suzuki 3782 (MAK); K. s.n., 1970 (TI); Iwate: T. Kosegawa & T. Takahashi s.n. Suetsugu KS282 (TNS); Tokyo: T. Yamazaki 771 (TI), T. (TNS01206782) (TNS), H. Kudo & A. Hiratsuka 268 Yamazaki s.n., 5/V/1982 (TI), J. Yamazaki 9518325 (TNS), (KYO) , F. Matsumoto 4272 (MBK); Miyagi: S. Saito 423 M. Toda s.n. (MBK0105687) (MBK); Kanagawa: S. (TI), J. Haginiwa JH002934 (TNS), H. S. Ogura 2811 Yanagawa s.n. (KPM-NA0294035) (KPM), K. Suetsugu (TUS); Fukushima: H. Sase 58-115 (FKSE), N. Sakurai s.n., 2/V/2015 (KYO-holotype, OSA-isotype); Okayama: s.n. (FKSE33074) (FKSE), Y. Endo s.n. (FKSE83315) S. Kariyama 53219 (KURA); Tokushima: K. Kondo s.n., (FKSE); Ibaraki: S. Fujimori F130506b (TNS); Tochigi: 5/VI/1928 (TI); Kochi: M. Watanabe et al. s.n. T. Noguchi & J. Oda 1440 (TOCH), H. Hayakawa & M. (MBK0090169) (MBK)