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 molecularphylogenetic analyses, several new identiﬁcations, new combinations, and new synonyms are proposed here.
Key words : Cephalanthera, Crepidium, Eastern Asia, Japan, Odontochilus, Orchidaceae, taxonomy,
Thrixspermum, Yoania

Recent reviews of Japanese Orchidaceae(Yukawa, 2015a, 2015b, in preparation) haverevealed some misidentiﬁcations and unrecognised synonyms of the taxa in Japan and adjacentregions. Furthermore, results of molecular phylogenetic analyses demonstrated the non-monophyly of several orchid genera and new combinations are thus needed to recover the monophyly.
Subsequent to Yukawa and Cribb (2014) andTang et al. (2015), the necessary treatment inaccordance with these ﬁndings is provided here.
Cephalanthera longifolia (L.) Fritsch.
Cephalanthera longifolia (L.) Fritsch. is a panEurasian species widely distributed across subarctic and temperate regions in Europe, Asia andNorth Africa from Ireland and Morocco in thewest to Korea in the east. Two species apparently
related to this species, namely, C. elegans Schltr.
and C. shizuoi F.Maek., were described from Japanese material. Taxonomic problems concerning
the two species were reviewed and discussed by
Yukawa et al. (2003) and Yukawa (2009).
Schlechter (1919) described Cephalanthera
elegans on the basis of a specimen collected by
K. Miyabe in Hakodate, southern part of Hokkaido Island in northern Japan. The holotype wasdeposited in Botanischer Garten und Botanisches

Museum Berlin-Dahlem, Zentraleinrichtung derFreien Universität Berlin (B) but almost all of
Schlechterʼs type collections in B were destroyed
by ﬁre in 1943. Yukawa and Ohba (1995) foundthat most of the duplicates of the type collectionsof Japanese orchid taxa described by Schlechterare deposited in the Herbarium, University ofTokyo (TI). However, they failed to locate any
type material of C. elegans in TI and the Herbarium, Hokkaido University Museum (SAPA)
where substantial parts of the Miyabe collection
are housed. I therefore examined characters of C.
elegans only on the basis of the protologue of thespecies and did not ﬁnd any morphological differences from C. longifolia.
Cephalanthera shizuoi was described byMaekawa (1936) on the basis of material collected by S. Hattori in Kanagawa Prefecture,eastern part of Honshu, the largest island ofJapan. The holotype deposited in TI and an illustration associated with the protologue perfectly
match with C. longifolia.
I also investigated herbarium specimens andliving plants referable to Cephalanthera longifolia from various regions of Japan. Although sizesof vegetative parts are variable among the individuals, characters of reproductive parts such as
the shape and colour of the perianth segments
and the morphology of the gynostemium are con-

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Tomohisa Yukawa

sistent. A notable exception is the uniform dwarf
habit in an isolated population around the summit of Mt. Tsurugi, Shikoku Island, western partof Japan. Even in this population, ﬂoral characters are identical to those of the other localities.
Besides, Sugaya (1959) emphasized the papillose
leaf, bract, and ovary of a specimen (A. Kimuras.n. collected on Mt. Moiwa, Hokkaido (TUS))
and suspected its conspeciﬁcity with C. shizuoi
in which both vegetative and reproductive organs
are glabrous except for the sparingly papillose
inﬂorescence (Maekawa, 1936). However, thisinterpretation is due to the overlooking of the
caducous nature of the papillae of C. longifolia
including C. shizuoi. This character thus cannot
be used as a diagnostic character of the species.
Further, comparisons with the Eurasian material of this species did not ﬁnd any morphological differences (Yukawa et al., 2003). The Eurasian material also showed great size variations
in vegetative parts and the contrasting uniformity
in reproductive parts. Consequently, it is reasonable to conclude that Cephalanthera longifolia isa variable species and distributed widely inJapan.
In Taiwan, Fukuyama (1938) described Cephalanthera alpicola Fukuy. from the Central
Mountains of the island, and the type specimen
and the protologue depict the characteristics of
C. longifolia. I further observed herbarium specimens and living plants collected in Taiwan andconcluded that these samples are identical to C.
longifolia. C. taiwaniana S.S. Ying, the otherTaiwanese Cephalanthera species, has beentreated as a synonym of C. alpicola in recent ﬂoristic works such as Su (2000) and Chen et al.
(2009). However, in the type material of C. taiwaniana, the foliage leaf is wider than C. longifolia and the sepals are shorter and wider thanthe petals (generally, sepals are longer and narrower than petals in C. longifolia), and the 5–7
lamellae on the epichile of the lip (3–5 in C. longifolia) are different. Further examinations ofalpine populations of C. longifolia in Taiwan areneeded to evaluate these variations.
Genetic diversity among Japanese, Taiwanese,and Eurasian samples identiﬁed as C. longifolia(including collections adjacent to type localities
of C. elegans, C. shizuoi, C. alpicola, and C. taiwaniana) was very low and these samples
formed a clade in relation to the other Cephalanthera species (M. Maki et al., unpublished data).
Consequently, both morphological and molecularevidence supported the view that the four entries,
C. elegans, C. shizuoi, C. alpicola, and C. taiwaniana are synonymous with C. longifolia.
Cephalanthera elegans and C. shizuoi aresometimes considered conspeciﬁc with C. erecta(Thunb.) Blume as noted by Ohwi (1965) and
Chen et al. (2009). However, C. erecta is distinct
from C. longifolia by morphological and molecular characters.
Cephalanthera longifolia (L.) Fritsch, Oesterr.
Bot. Z. 38: 81 (1888). [Fig. 1, A, B]
Basionym: Serapias helleborine var. longifolia
L., Sp. Pl.: 950 (1753). Neotype: based on
Oeder, Fl. Danica 3 (9): t. 506, f. media(1770).
Cephalanthera elegans Schltr., Repert. Spec.
Nov. Regni Veg. Beih. 4: 58 (1919), syn. nov.
Type: Japan, Hokkaido, Hakodate, Miyabe s.n.
(holotype B, destroyed).
Cephalanthera shizuoi F.Maek., in Nakai,
Iconogr. Pl. Asiae Orient. 1: 58 (1936). Type:Japan, Honshu, Kanagawa Pref., Kugenuma,
S.Hattori s.n. (holotype TI!).
Cephalanthera alpicola Fukuy., Bot. Mag.
(Tokyo) 52: 242 (1938), syn. nov. Type: Taiwan, Kwarenko, Mt. Gokwan-zan (Hohuanshan), 16 June 1935, K.Segawa s.n., Herb.
Fukuyama 5915 (holotype TAI, not located).
Taiwan, Kwarenko, Mt. Gokwan-zan, 16 June
1935, K.Segawa s.n. (lectotype KPM, here
designated). Although the holotype is not
located, a specimen with the identical collection data but without the collection number isdeposited in KPM (Inoue et al., 1998). This
specimen (KPM-NA0105517) is appropriate
for the lectotype of this species.
Specimens examined: Japan, Hokkaido: Asahikawa-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,Shiretoko Peninsula, along Uwaebetsu River, alt.
ca. 170 m, 2014-6-29, Akitomo Uchida s.n.
(TNS8505657); Noboribetsu-shi, Washibesu-cho,on a hill, 1973-7-21, Matsuji Hara s.n.
(TNS01013413); Noboribetsu-shi, Washibetsucho, 1974-6-12, Matsuji Hara s.n. (TNS01013414).
Honshu: Aomori Pref. Higashidori-mura, alt.

10 m, 2010-6-8, T. Kinoshita s.n. (TNS8504905);
Azumadake, T. Numata s.n. (TNS8504537);
Misawa-shi, Sabishiro, on the beach of Ogawarako Lake, 1977-5-29, Tatsuro Ohsawa s.n.
(TNS343232). Akita Pref. Akita-shi, Shimoshinjo, 2008-5-21, K. Miyoshi s.n. (TNS8501244).
Yamagata Pref. Mukaihara-Kanrin, 1930-6-2,
Haruki Okuyama s.n. (TNS39611); Yamadera,

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Tomohisa Yukawa

1931-6-14, Haruki Okuyama 24138 (TNS
291933). Ibaraki Pref. Tsukuba-shi, Universityof Tsukuba, 2008-5-7, Yuki Tsujita T08-4(TNS8500539); Tsukuba-shi, Namiki, 2008-5-6,
Yuki Tsujita T08-3 (TNS 8500541); Tsukuba-shi,
Amakubo, 2008-5-14, Tomohisa Yukawa 08-2
(TNS8501231). Tokyo Metropolis. Nishitamagun, Okutama-machi, Nippara, Ogawatani, 19715-27, Midori Miyamoto 495 (TNS01086553).
Chiba Pref. Chiba-shi, Mihama-ku, Saiwaimachi, 2005-4-28, M. Saito s.n. (TNS8501418).
Kanagawa Pref. Tsukui-gun, Sagamiko-machi,
near Sagami-ko Lake, alt. 200 m, 1969-5-6, Tetsuya
Kawasaki 6135 (TNS673113); Yokohama-shi,
Aoba-ku, Nara-cho, Tamagawa University, 20025-7, J. Yamazaki s.n. (TNS8503114); Yokohamashi, Aoba-ku, Nara-cho, Tamagawa University,
2003-5-7, J. Yamazaki s.n. (TNS8503107,
8503110); Yokohama-shi, Aoba-ku, Nara-cho,Tamagawa University, 2004-4-28, J. Yamazakis.n. (TNS8503113); Fujisawa-shi, Hatori, 19924-30 (TNS8503981). KYUSHU: Fukuoka Pref.
Kitakyushu-shi, Kokuraminami-ku, broad-leaved
deciduous forest, alt. 520 m, Kohji Tanaka s.n.
(TNS8500585).
Crepidium Blume
Results of phylogenetic analysis of tribe Malaxideae using macromolecular characters (Cameron, 2005) deﬁnitely showed the polyphyletic
status of the genus Malaxis s.l. To restore themonophyly of the genus, it is necessary to recognise Malaxis section Crepidium as an independent genus, Crepidium Blume. The following
endemic species of the Paciﬁc islands have notbeen transferred to Crepidium yet.
Crepidium boninense (Koidz.) T.Yukawa, comb.
nov.
Basionym: Microstylis boninensis Koidz., Bot.
Mag. (Tokyo) 32: 137 (1918). Type: Japan,
Bonin Isls., Insl. Chichishima, 30 September
1917, S.Nishimura 107 (holotype TI).
Crepidium alamaganense (S.Kobay.) T.Yukawa,comb. nov.

Basionym: Malaxis alamaganensis S.Kobay.,
Nat. Hist. Res. (Chiba), Special Issue, 1: 71(1994). Type: Northern Mariana Isls., Alamagan, ca. alt. 640 m, 9 June 1992, T.Ohba
CBM-BS-59238 (holotype CBM).
Odontochilus Blume
As pointed out by Ormerod (2002), the deﬁnition of Odontochilus Blume is problematic
because it seems that otherwise critical genericcharacters in tribe Cranichideae subtribe
Goodyerinae such as twisting of the column andconnation of stigma lobes appear to be inconsistent or variable and thus have less value ingeneric circumscription. Moreover, results ofmolecular phylogenetic analysis of subtribe
Goodyerinae revealed that Evrardianthe
Rauschert, Kuhlhasseltia J.J.Sm., Myrmechis
Blume, Pristiglottis Cretz. & J.J.Sm., and Vexillabium F.Maek. are nested within Odontochilus(T. Yukawa, unpublished). As a result, Odontochilus represents a polyphyletic entity, if Evrardianthe, Kuhlhasseltia, Myrmechis, Pristiglottis,
and Vexillabium are treated as independent genera. Consequently, it is appropriate to circumscribe Odontochilus in a broad sense to recover
the monophyletic status of this genus. The transfer of several species in Kuhlhasseltia, Myrmechis, Pristiglottis, and Vexillabium to Odontochilus is proposed here.
Odontochilus Blume, Fl. Javae Nov. Ser. 1: 66(1858–1859). Type: Odontochilus ﬂavescens(Blume) Blume.
Kuhlhasseltia J.J.Sm., Icon. Bogor. 4: t. 301(1910). Type: Kuhlhasseltia javanica J.J.Sm.,
syn. nov.
Myrmechis Blume, Coll. Orchid.: 76 (1859).
Type: Myrmechis gracilis (Blume) Blume,
syn. nov.
Vexillabium F.Maek., J. Jap. Bot. 11: 457 (1935).
Type: Vexillabium nakaianum F.Maek., syn.
nov.
Odontochilus aureus (J.J.Sm.) T.Yukawa, comb.
nov.

Orchid Flora of Japan

Basionym: Tubilabium aureum J.J.Sm., Bull.
Jard. Bot. Buitenzorg, sér. 3, 9: 446 (1928).
Odontochilus bakhimensis (D.Maity, N.Pradhan& Maiti) T.Yukawa, comb. nov.
Basionym: Myrmechis bakhimensis D.Maity,
N.Pradhan & Maiti, Acta Phytotax. Sin. 45:
321 (2007).
Odontochilus bilobuliferus (J.J.Sm.) T.Yukawa,comb. nov.
Basionym: Tubilabium bilobuliferum J.J.Sm.,
Bull. Jard. Bot. Buitenzorg, sér. 3, 10: 6(1928).
Odontochilus chalmersii (Schltr.) T.Yukawa,comb. nov.
Basionym: Zeuxine chalmersii Schltr., Bull.
Herb. Boissier, sér. 2, 6: 297 (1906).
Odontochilus chinensis (Rolfe) T.Yukawa,comb. nov.
Basionym: Myrmechis chinensis Rolfe, J. Linn.
Soc., Bot. 36: 44 (1903).
Odontochilus coerulescens (Schltr.) T.Yukawa,comb. nov.
Basionym: Cystopus coerulescens Schltr., Repert. Spec. Nov. Regni Veg. Beih. 1: 68 (1911).
Odontochilus drymoglossifolius (Hayata) T.
Yukawa, comb. nov.
Basionym: Myrmechis drymoglossifolia Hayata,Icon. Pl. Formosan. 6: 90 (1916).
Odontochilus elongatus (Blume) T.Yukawa,comb. nov.
Basionym: Cystopus elongatus Blume, Coll.
Orchid.: 84 (1859).
Odontochilus ﬁmbriatus (J.J.Sm.) T.Yukawa,comb. nov.
Basionym: Cystopus ﬁmbriatus J.J.Sm., Bull.
Dép. Agric. Indes Néerl. 10: 3 (1907).
Odontochilus ﬁssus (F.Maek.) T.Yukawa, comb.
nov.
Basionym: Vexillabium ﬁssum F.Maek., J. Jap.
Bot. 12: 91 (1936).
Odontochilus gilesii (Ormerod) T.Yukawa,comb. nov.
Basionym: Kuhlhasseltia gilesii Ormerod, Lindleyana 17: 207 (2002).
Odontochilus glaber (Blume) T.Yukawa, comb.
nov.

107

Basionym: Myrmechis glabra Blume, Coll.
Orchid.: 76 (1859). This species is a later synonym of Rhamphidia grandiﬂora Lindl. (J.
Proc. Linn. Soc., Bot. 1: 182 (1857)) but thetransfer of R. grandiﬂora generates a homonym of Odontochilus grandiﬂorus (Lindl.)
Hook.f.
Odontochilus gracilis (Blume) T.Yukawa, comb.
nov.
Basionym: Anoectochilus gracilis Blume, Bijdr.
Fl. Ned. Ind.: 413 (1825).
Odontochilus hatumetensis (J.J.Sm.) T.Yukawa,comb. nov.
Basionym: Cystopus hatumetensis J.J.Sm., Bull.
Jard. Bot. Buitenzorg, sér. 3, 10: 99 (1928).
Odontochilus integrus (Fukuy.) T.Yukawa,comb. nov.
Basionym: Pristiglottis integra Fukuy., Bot Mag.
(Tokyo) 50: 20 (1936).
Odontochilus japonicus (Rchb.f.) T.Yukawa,comb. nov.
Basionym: Rhamphidia japonica Rchb.f., Bot.
Zeitung (Berlin) 36: 75 (1878).
Odontochilus javanicus (J.J.Sm.) T.Yukawa,comb. nov.
Basionym: Kuhlhasseltia javanica J.J.Sm., Icon.
Bogor. 4: t. 301 (1910).
Odontochilus kinabaluensis (Carr) T.Yukawa,comb. nov.
Basionym: Myrmechis kinabaluensis Carr, Gard.
Bull. Straits Settlem. 8: 188 (1935).
Odontochilus luzonensis (Ames) T.Yukawa,comb. nov. et nom. nov.
Basionym: Cystopus philippinensis Ames, Orchidaceae 5: 35 (1915). The binominal O. philippinensis cannot be used for this entity because
this combination was used for another entity
based on Myrmechis philippinensis Ames. The
epithet “luzonensis” was derived from the type
locality of the species.
Odontochilus mindanaensis (Ames) T.Yukawa,comb. nov.
Basionym: Cystopus mindanaensis Ames, Orchidaceae 5: 34 (1915).
Odontochilus montanus (Schltr.) T.Yukawa,comb. nov.

108

Tomohisa Yukawa

Basionym: Anoectochilus montanus Schltr., Bot.
Jahrb. Syst. 39: 55 (1906).
Odontochilus muricatus (J.J.Sm.) T.Yukawa,comb. nov.
Basionym: Cystopus muricatus J.J.Sm., Orch.
Ambon: 13 (1905).
Odontochilus nakaianus (F.Maek.) T.Yukawa,comb. nov.
Basionym: Vexillabium nakaianum F.Maek., J.
Jap. Bot. 11: 458 (1935).
Odontochilus occultus (Blume) T.Yukawa,comb. nov.
Basionym: Cystopus occultus Blume, Coll.
Orchid.: 85 (1859).
Odontochilus papuanus (J.J.Sm.) T.Yukawa,comb. nov.
Basionym: Kuhlhasseltia papuana J.J.Sm., Nova
Guinea 12: 9 (1913).
Odontochilus pectiniferus (Schltr.) T.Yukawa,comb. nov.
Basionym: Cystopus pectiniferus Schltr., Repert.
Spec. Nov. Regni Veg. Beih. 1: 69 (1911).
Odontochilus perpusillus (Ames) T.Yukawa,comb. nov.
Basionym: Myrmechis perpusilla Ames, Schedul.
Orchid. 6: 15 (1923).
Odontochilus philippinensis (Ames) T.Yukawa,comb. nov.
Basionym: Myrmechis philippinensis Ames,
Orchidaceae 2: 64 (1908).
Odontochilus puberulus (Schltr.) T.Yukawa,comb. nov.
Basionym: Cystopus puberulus Schltr., Repert.
Spec. Nov. Regni Veg. Beih. 1: 70 (1911).
Odontochilus pubescens (Blume) T.Yukawa,comb. nov.
Basionym: Anoectochilus pubescens Blume,
Bijdr. Fl. Ned. Ind.: 412 (1825).
Odontochilus quadrilobatus (Schltr.) T.Yukawa,comb. nov.
Basionym: Cheirostylis quadrilobata Schltr.,
Repert. Spec. Nov. Regni Veg. 10: 11 (1911).
Odontochilus rajanus (J.J.Sm.) T.Yukawa,comb. nov.
Basionym: Kuhlhasseltia rajana J.J.Sm., Mitt.
Inst. Allg. Bot. Hamburg 7: 26 (1927).

Odontochilus seranicus (J.J.Sm.) T.Yukawa,comb. nov.
Basionym: Myrmechis seranica J.J.Sm., Bull.
Jard. Bot. Buitenzorg, sér. 3, 10: 95 (1928).
Odontochilus sibelae (Ormerod) T.Yukawa,comb. nov.
Basionym: Kuhlhasseltia sibelae Ormerod, Lindleyana 17: 209 (2002).
Odontochilus spicatus (Blume) T.Yukawa,comb. nov.
Basionym: Cystopus spicatus Blume, Coll.
Orchid.: 84 (1859).
Odontochilus tsukusianus (Masam.) T.Yukawa,comb. nov.
Basionym: Myrmechis tsukusiana Masam., Bot.
Mag. (Tokyo) 43: 250 (1929).
Odontochilus urceolatus (Tang & K.Y.Lang)
T.Yukawa, comb. nov.
Basionym: Myrmechis urceolata Tang &K.Y.Lang, Acta Phytotax. Sin. 34: 638 (1996).
Odontochilus whiteheadii (Rendle) T.Yukawa,comb. nov.
Basionym: Zeuxine whiteheadii Rendle, J. Bot.
34: 358 (1896).
Odontochilus yakushimensis (Yamam.) T.
Yukawa, comb. nov.
Basionym: Anoectochilus yakushimensis Yamam.,Bot. Mag. (Tokyo) 38: 131 (1924).
Thrixspermum pygmaeum(King & Pantl.) Holttum
A small Thrixspermum species was discovered
in a few localities of subtropical evergreen
broadleaf forests on Amami-oshima Island,
Ryukyu Islands by Hiroshi Yamashita in 1980.
This entity was ﬁrst reported by Hatusima (1986)
as T. pricei (Rolfe) Schltr. Subsequently, twomore binominals, T. formosanum (Hayata) Schltr.
and T. saruwatarii (Hayata) Schltr., have been
applied to the Amami populations. Re-examination of the living material revealed that the plants
should be referred to T. pygmaeum (King &Pantl.) Holttum rather than the three aforementioned species. T. pygmaeum is well demarcatedfrom T. saruwatarii by the ﬂower colour (cream

Orchid Flora of Japan109

yellow in the former, white with purplish tints inthe latter), an apical callus of the disc of the lip(absent in the former, present in the latter), and alonger disc of the lip (2 mm in the former, 3 mmin the latter). T. formosanum can be distinguishedby a white ﬂower, a well-developed, cylindric
spur, and successive and ephemeral ﬂowering.
As correctly pointed out by Lin and Hsu (1977),
T. pricei is a later synonym of T. formosanum.
Moreover, protologues of Thrixspermum laurisilvaticum (Fukuy.) Garay and T. xanthanthum
Tuyama, both described on the basis of Taiwanese specimens, match well with T. pygmaeumand I treat them as synonyms of T. pygmaeum.
Besides, specimens of T. saruwatarii fromArunachal Pradesh (Mao et al., 2010) also likely
represent T. pygmaeum. Since the identiﬁcationof these Thrixspermum species is confusing, reexamination is necessary for the records of these
species in East Asia.
So far, “Keitao-furan” has been used for theJapanese name of the Amami-oshima plants.
However, this name corresponds to Thrixspermum saruwatarii. To emphasise the afﬁnity ofthis species with T. japonicum (Miq.) Rchb.f., i.
e. “Kayaran” in Japanese, I hereby propose a newJapanese name “Amami-kayaran” for this entity.

Hatusima, Fl. Kagoshima Rev. ed.: 236(1986). Shimabuku, Check List Vascular Fl.
Ryukyu Islands: 702 (1990). Hatusima &Amano, Fl. Ryukyu 2nd ed.: 345 (1994).
Shimabuku, Check List Vascular Fl. Ryukyu
Islands Rev. ed.: 797 (1997).
Thrixspermum formosanum auct. non (Hayata)
Schltr., Hashimoto, Proc. 11 World Orch.
Conf.: 124 (1987). Masamune, Native Orch.
Nippon 7: 164 (1995).
Thrixspermum saruwatarii auct. non (Hayata)
Schltr., Takahashi, Wild Orch. Jap. 3: 162(1987). Hashimoto, Jap. Ind. Orch. Colour
Rev. ed.: 251 (1991).
Specimens examined: Japan, Ryukyu Isls.,
Amami-oshima Is., Kinsakubaru, 15 June 2004,
H. Yamashita s.n. ex hort. Tsukuba Bot. Gard.
(TNS8500407); same locality, 24 February 1990,
T. Hashimoto s.n. (TNS8503922, TNS9504630);
same locality, 27 March 1993, H. Yamashita s.n.
(TNS8504492); same locality, 1 April 1992, H.
Yamashita s.n. (TNS8504748); Ryukyu Isls.,
Amami-oshima Is., along Sumiyo-gawa River,
upper stream, alt. 194 m. March 2013, Y. Maeda
s.n. ex hort. Tsukuba Bot. Gard. (TNS8505800).

Thrixspermum pygmaeum (King & Pantl.) Holttum, Kew Bull. 14: 275 (1960). [Fig. 1, C, D]
Basionym: Sarcochilus pygmaeus King & Pantl.,
Ann. Roy. Bot. Gard. (Calcutta) 8: 207 (1898).
Type: Sikkim, Pemiongtsi (Pemayangtse),
Pantling 472 (holotype CAL).
Sarcochilus laurisilvaticus Fukuy., Bot. Mag.
(Tokyo) 52: 246 (1938), syn. nov. Type:
Taiwan, Praef. Sintiku, alt. ca. 600 m,
N.Fukuyama s.n., Herb. Fukuyama 6106
(holotype, not located).
Thrixspermum laurisilvaticum (Fukuy.) Garay,
Bot. Mus. Leaﬂ. 23: 207 (1972).
Thrixspermum xanthanthum Tuyama, J. Jap. Bot.
16: 523 (1940), syn. nov. Type: Taiwan, Praef.
Taihoku, Taiheizan, Kasinoki-daira, 2 April
1940, Y.Kobayashi s.n. (holotype TI!).
Thrixspermum pricei auct. non (Rolfe) Schltr.,

A new genus Yunorchis and its sole species Y.
pingbianensis were described by Z.J. Liu, G.Q.
Zhang and M. He Li in Zhang et al. (2015). Theyemphasised that Yunorchis is morphologicallydifferent from the other genera in tribe Calypsoeae by a combination of the subumbellate
inﬂorescence, the long pedicel, and the forwardhooked lip with reduced lateral lobes. Moreover,results of molecular phylogenetic analysis bythem showed that the new taxon occupies a sistergroup position to Dactylostalix Rchb.f. and/or
Calypso Salisb. However, morphological characters of Yunorchis ﬁt well with Yoania Maxim.
Molecular phylogenetic analysis incorporating
more taxa of Calypsoeae revealed the monophylyof Yunorchis and Yoania with strong support (J.
Freudenstein and T. Yukawa, unpublished). Consequently, it is appropriate to reduce Yunorchis as

Yoania Maxim.

110

Tomohisa Yukawa

a synonym of Yoania.
Yoania pingbianensis is closely related to Y.
prainii King & Pantl., a species from Sikkim.
The former has a more developed basal part ofthe lip and also exhibits purple-red in the abaxial
surface of the sepals while the latter presents
white ﬂowers. Furthermore, a central papillatepubescent callus of the lip and distant scales on
the rhizome characterise Y. pingbianensis. In
contrast, Y. prainii does not show a callus of the
lip and has characteristic, densely clothed, imbricating scales on the rhizome.
Descriptions and illustrations of Yoania prainii
from Ha Giang, Vietnam (Averyanov, 2011)
show that these plants were misidentiﬁed and are
actually referable to Y. pingbianensis, and thatthe distribution of the latter species extends into
the northern part of Vietnam.
Yoania Maxim., Bull. Acad. Imp. Sci. SaintPétersbourg, sér. 3, 18: 68 (1873). Type:
Yoania japonica Maxim.
Yunorchis Z.J.Liu, G.Q.Zhang & M.He Li, PLoS
ONE 10(4): e123382 (11) (2015), syn. nov.
Type: Yunorchis pingbianensis Z.J.Liu, G.Q.
Zhang & M.He Li
Yoania pingbianensis (Z.J. Liu, G.Q. Zhang &M. He Li) T.Yukawa & Freudenstein, comb.
nov.
Basionym: Yunorchis pingbianensis Z.J.Liu,
G.Q.Zhang & M.He Li, PLoS ONE 10(4):e123382 (11) (2015). Type: China, Yunnan,
Pingbian, in a forest, alt. 2100 m, 31 May
2013, Z. J. Liu 7103 (holotype NOCC).
Yoania prainii auct. non King & Pantl., Averyanov & Averyanova, Turczaninowia 3: 85(2000). Averyanov, Turczaninowia 14: 15–100(2011).
Acknowledgments
I would like to thank John Freudenstein,
Teruo Katsuyama, Masayuki Maki, and Yumi
Yamashita for sharing useful information, Yun-I
Lee, Yoshiyuki Maeda and Hiroshi Yamashita for
supplying the plant material, Takahiro Yagame

and Hiroshi Yamashita for providing thephotographs, and Hubert Kurzweil for criticalreading of the manuscript. This study is partly
supported by JSPS KAKENHI Grant Numbers
JP15H04417 and JP15K14442 and a research
grant from National Museum of Nature and Science.
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