Taxonomic Notes on the Orchidaceae of Japan and Adjacent Regions
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Bull. Natl. Mus. Nat. Sci., Ser. B, 42(3), pp. 103–111, August 22, 2016 Taxonomic Notes on the Orchidaceae of Japan and Adjacent Regions Tomohisa Yukawa Department of Botany, Tsukuba Botanical Garden, National Museum of Nature and Science, Amakubo 4–1–1, Tsukuba, Ibaraki 305–0005, Japan (Received 27 May 2016; accepted 4 July 2016) Abstract In accordance with recent reviews of Japanese Orchidaceae and results of molecular phylogenetic analyses, several new identifications, new combinations, and new synonyms are pro- posed here. Key words : Cephalanthera, Crepidium, Eastern Asia, Japan, Odontochilus, Orchidaceae, taxonomy, Thrixspermum, Yoania Recent reviews of Japanese Orchidaceae Museum Berlin-Dahlem, Zentraleinrichtung der (Yukawa, 2015a, 2015b, in preparation) have Freien Universität Berlin (B) but almost all of revealed some misidentifications and unrecog- Schlechter’s type collections in B were destroyed nised synonyms of the taxa in Japan and adjacent by fire in 1943. Yukawa and Ohba (1995) found regions. Furthermore, results of molecular phylo- that most of the duplicates of the type collections genetic analyses demonstrated the non-mono- of Japanese orchid taxa described by Schlechter phyly of several orchid genera and new combina- are deposited in the Herbarium, University of tions are thus needed to recover the monophyly. Tokyo (TI). However, they failed to locate any Subsequent to Yukawa and Cribb (2014) and type material of C. elegans in TI and the Herbar- Tang et al. (2015), the necessary treatment in ium, Hokkaido University Museum (SAPA) accordance with these findings is provided here. where substantial parts of the Miyabe collection are housed. I therefore examined characters of C. elegans only on the basis of the protologue of the Cephalanthera longifolia (L.) Fritsch. species and did not find any morphological dif- Cephalanthera longifolia (L.) Fritsch. is a pan- ferences from C. longifolia. Eurasian species widely distributed across sub- Cephalanthera shizuoi was described by arctic and temperate regions in Europe, Asia and Maekawa (1936) on the basis of material col- North Africa from Ireland and Morocco in the lected by S. Hattori in Kanagawa Prefecture, west to Korea in the east. Two species apparently eastern part of Honshu, the largest island of related to this species, namely, C. elegans Schltr. Japan. The holotype deposited in TI and an illus- and C. shizuoi F.Maek., were described from Jap- tration associated with the protologue perfectly anese material. Taxonomic problems concerning match with C. longifolia. the two species were reviewed and discussed by I also investigated herbarium specimens and Yukawa et al. (2003) and Yukawa (2009). living plants referable to Cephalanthera longifo- Schlechter (1919) described Cephalanthera lia from various regions of Japan. Although sizes elegans on the basis of a specimen collected by of vegetative parts are variable among the indi- K. Miyabe in Hakodate, southern part of Hok- viduals, characters of reproductive parts such as kaido Island in northern Japan. The holotype was the shape and colour of the perianth segments deposited in Botanischer Garten und Botanisches and the morphology of the gynostemium are con- 104 Tomohisa Yukawa sistent. A notable exception is the uniform dwarf and Eurasian samples identified as C. longifolia habit in an isolated population around the sum- (including collections adjacent to type localities mit of Mt. Tsurugi, Shikoku Island, western part of C. elegans, C. shizuoi, C. alpicola, and C. tai- of Japan. Even in this population, floral charac- waniana) was very low and these samples ters are identical to those of the other localities. formed a clade in relation to the other Cephalan- Besides, Sugaya (1959) emphasized the papillose thera species (M. Maki et al., unpublished data). leaf, bract, and ovary of a specimen (A. Kimura Consequently, both morphological and molecular s.n. collected on Mt. Moiwa, Hokkaido (TUS)) evidence supported the view that the four entries, and suspected its conspecificity with C. shizuoi C. elegans, C. shizuoi, C. alpicola, and C. tai- in which both vegetative and reproductive organs waniana are synonymous with C. longifolia. are glabrous except for the sparingly papillose Cephalanthera elegans and C. shizuoi are inflorescence (Maekawa, 1936). However, this sometimes considered conspecific with C. erecta interpretation is due to the overlooking of the (Thunb.) Blume as noted by Ohwi (1965) and caducous nature of the papillae of C. longifolia Chen et al. (2009). However, C. erecta is distinct including C. shizuoi. This character thus cannot from C. longifolia by morphological and molecu- be used as a diagnostic character of the species. lar characters. Further, comparisons with the Eurasian mate- rial of this species did not find any morphologi- Cephalanthera longifolia (L.) Fritsch, Oesterr. cal differences (Yukawa et al., 2003). The Eur- Bot. Z. 38: 81 (1888). [Fig. 1, A, B] asian material also showed great size variations Basionym: Serapias helleborine var. longifolia in vegetative parts and the contrasting uniformity L., Sp. Pl.: 950 (1753). Neotype: based on in reproductive parts. Consequently, it is reason- Oeder, Fl. Danica 3 (9): t. 506, f. media able to conclude that Cephalanthera longifolia is (1770). a variable species and distributed widely in Cephalanthera elegans Schltr., Repert. Spec. Japan. Nov. Regni Veg. Beih. 4: 58 (1919), syn. nov. In Taiwan, Fukuyama (1938) described Ceph- Type: Japan, Hokkaido, Hakodate, Miyabe s.n. alanthera alpicola Fukuy. from the Central (holotype B, destroyed). Mountains of the island, and the type specimen Cephalanthera shizuoi F.Maek., in Nakai, and the protologue depict the characteristics of Iconogr. Pl. Asiae Orient. 1: 58 (1936). Type: C. longifolia. I further observed herbarium speci- Japan, Honshu, Kanagawa Pref., Kugenuma, mens and living plants collected in Taiwan and S.Hattori s.n. (holotype TI!). concluded that these samples are identical to C. Cephalanthera alpicola Fukuy., Bot. Mag. longifolia. C. taiwaniana S.S. Ying, the other (Tokyo) 52: 242 (1938), syn. nov. Type: Tai- Taiwanese Cephalanthera species, has been wan, Kwarenko, Mt. Gokwan-zan (Hohuan- treated as a synonym of C. alpicola in recent flo- shan), 16 June 1935, K.Segawa s.n., Herb. ristic works such as Su (2000) and Chen et al. Fukuyama 5915 (holotype TAI, not located). (2009). However, in the type material of C. tai- Taiwan, Kwarenko, Mt. Gokwan-zan, 16 June waniana, the foliage leaf is wider than C. longi- 1935, K.Segawa s.n. (lectotype KPM, here folia and the sepals are shorter and wider than designated). Although the holotype is not the petals (generally, sepals are longer and nar- located, a specimen with the identical collec- rower than petals in C. longifolia), and the 5–7 tion data but without the collection number is lamellae on the epichile of the lip (3–5 in C. lon- deposited in KPM (Inoue et al., 1998). This gifolia) are different. Further examinations of specimen (KPM-NA0105517) is appropriate alpine populations of C. longifolia in Taiwan are for the lectotype of this species. needed to evaluate these variations. Specimens examined: Japan, Hokkaido: Asa- Genetic diversity among Japanese, Taiwanese, hikawa-shi, Asahiyama, alt. 240 m, 1960-6-11, Orchid Flora of Japan 105 Fig. 1. Cephalanthera longifolia (L.) Fritsch from the habitat of Mt. Daisen, Tottori Pref., Honshu, Japan (A, B). A. Habit. B. Flowers. Thrixspermum pygmaeum (King & Pantl.) Holttum from the habitat of Amami Is., Kagoshima Pref., Ryukyu Isls., Japan (C, D). C. Habit. D. Flowers. (Photos A, B by T. Yagame; C, D by H. Yamashita) Kanji Tokura 953 (TNS695747); Shari-cho, 10 m, 2010-6-8, T. Kinoshita s.n. (TNS8504905); Shiretoko Peninsula, along Uwaebetsu River, alt. Azumadake, T. Numata s.n. (TNS8504537); ca. 170 m, 2014-6-29, Akitomo Uchida s.n. Mi sawa-shi, Sabishiro, on the beach of Ogawara- (TNS8505657); Noboribetsu-shi, Washibesu-cho, ko Lake, 1977-5-29, Tatsuro Ohsawa s.n. on a hill, 1973-7-21, Matsuji Hara s.n. (TNS343232). Akita Pref. Akita-shi, Shimo- (TNS01013413); Noboribetsu-shi, Washibetsu- shinjo, 2008-5-21, K. Miyoshi s.n. (TNS8501244). cho, 1974-6-12, Matsuji Hara s.n. (TNS01013414). Yamagata Pref. Mukaihara-Kanrin, 1930-6-2, Honshu: Aomori Pref. Higashidori-mura, alt. Haruki Okuyama s.n. (TNS39611); Yamadera, 106 Tomohisa Yukawa 1931-6-14, Haruki Okuyama 24138 (TNS Basionym: Malaxis alamaganensis S.Kobay., 291933). Ibaraki Pref. Tsukuba-shi, University Nat. Hist. Res. (Chiba), Special Issue, 1: 71 of Tsukuba, 2008-5-7, Yuki Tsujita T08-4 (1994). Type: Northern Mariana Isls., Ala- (TNS8500539); Tsukuba-shi, Namiki, 2008-5-6, magan, ca. alt. 640 m, 9 June 1992, T.Ohba Yuki Tsujita T08-3 (TNS 8500541); Tsukuba-shi, CBM-BS-59238 (holotype CBM). Amakubo, 2008-5-14, Tomohisa Yukawa 08-2 (TNS8501231). Tokyo Metropolis. Nishitama- Odontochilus Blume gun, Okutama-machi, Nippara, Ogawatani, 1971- 5-27, Midori Miyamoto 495 (TNS01086553). As pointed out by Ormerod (2002), the defini- Chiba Pref. Chiba-shi, Mihama-ku, Saiwai- tion of Odontochilus Blume is problematic machi, 2005-4-28, M. Saito s.n. (TNS8501418). because it seems that otherwise critical generic Kanagawa Pref. Tsukui-gun, Sagamiko-machi, characters in tribe Cranichideae subtribe near Sagami-ko Lake, alt. 200 m, 1969-5-6, Tetsu ya Goodyerinae such as twisting of the column and Kawasaki 6135 (TNS673113); Yokohama-shi, connation of stigma lobes appear to be inconsis- Aoba-ku, Nara-cho, Tamagawa University, 2002- tent or variable and thus have less value in 5-7, J. Yamazaki s.n. (TNS8503114); Yokohama- generic circumscription. Moreover, results of shi, Aoba-ku, Nara-cho, Tamagawa University, molecular phylogenetic analysis of subtribe 2003-5-7, J. Yamazaki s.n. (TNS8503107, Goodyerinae revealed that Evrardianthe 8503110); Yokohama-shi, Aoba-ku, Nara-cho, Rauschert, Kuhlhasseltia J.J.Sm., Myrmechis Tamagawa University, 2004-4-28, J. Yamazaki Blume, Pristiglottis Cretz. & J.J.Sm., and Vexill- s.n. (TNS8503113); Fujisawa-shi, Hatori, 1992- abium F.Maek. are nested within Odontochilus 4-30 (TNS8503981).