Bull. Natl. Mus. Nat. Sci., Ser. B, 45(3), pp. 107–118, August 22, 2019

Taxonomic Reappraisal of Liparis japonica and L. makinoana(Orchidaceae)
Chie Tsutsumi*, Tomohisa Yukawa and Masahiro Kato
Department of Botany, National Museum of Nature and Science,
4–1–1 Amakubo, Tsukuba, Ibaraki 305–0005, Japan* E-mail: tsutsumi@kahaku.go.jp(Received 3 June 2019; accepted 13 July 2019)

Abstract Liparis japonica and L. makinoana have been nomenclaturally confused for a longtime. Based on a recent DNA sequencing and morphological study, these species appear to becomposed of three different entities. Here, we treat the three entities as L. makinoana emend. andtwo new species, L. suzumushi and L. longiracemosa, with neotypiﬁcation of L. makinoana.
Liparis japonica should be treated as a synonym of Malaxis monophyllos, based on the originaldescription of the former.
Key words : classiﬁcation, Japan, Liparis japonica, Liparis makinoana.

Introduction
Liparis Rich. is an orchid genus consisting ofover 320 terrestrial and epiphytic species (Pridgeon et al., 2005) and widely distributed in tropical and temperate regions of the world. In Japan,
17 species of Liparis are recorded (Yukawa,2015). Liparis japonica (Miq.) Maxim., L. makinoana Schltr., and L. hostifolia (Koidz.) Koidz.
ex Nakai generally form a species complex characterized by ovoid pseudobulbs, two deciduous,
ovate-elliptic leaves, absence of a projection at
the lip base, and a beaked anther cap (Yukawa,
2015; Takayama et al., in press). The taxonomic
treatments of the species complex, particularly of
L. japonica and L. makinoana, as well as the pictures in various publications are confusing(Iinuma and Makino, 1913; Maekawa, 1971;
Satomi, 1982; Hashimoto, 1990; Hashimoto etal., 1991).
Tsutsumi and Yukawa (2008) found three entities in Japan, named as L. japonica and L. makinoana and tentatively named them as Type I, II,
and III of the L. makinoana complex [Fig. 1. See
also Tsutsumi and Yukawa (2008) for the front

views of the ﬂowers]. The entities are distinct
from each other in their ﬂower size and nucleic
acid sequences in the nuclear ribosomal ITS
region (Tsutsumi and Yukawa, 2008). A RADSeq analysis also suggested the distinct nature of
the three entities (unpublished). These results
suggest that the three entities should be treated as
distinct species. The other species of the L. makinoana complex, L. hostifolia, from the subtropical Bonin Islands, has several unique characters:a broadly ovate labellum, short dormancy, andﬂowering time from November to March(Takayama et al., in press). Therefore, we haveexcluded it from this reappraisal.
Liparis japonica was originally described as
Microstylis japonica by Miquel (1866), based ona specimen collected in Japan by H. Bürger.
Later, it was recombined with Liparis by Maximowicz (1886). The type specimen was supposed to have been deposited in the Naturalis (L)in Leiden, the Netherlands, but it has not beenfound there (Ohba et al., 2005; A. Schuiteman,pers. comm.). Liparis makinoana was describedby Schlechter (1919). Its type specimen, collected from Hokkaido, Japan, had been deposited

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Fig. 1. Liparis makinoana complex. A, B: Liparis suzumushi (Type I in Tsutsumi and Yukawa, 2008). C, D:
Liparis makinoana (Type II). E, F: Liparis longiracemosa (Type III).

Table 1. Taxonomic treatments of the Liparis makinoana complex in previous and present studies.
Previous treatments

Tsutsumi and
Yukawa(2008)

Present study

L. makinoana sensu Ohwi (1953, 1984), Satomi (1982),
Type I
L. suzumushi
Nakajima (2012)
L. makinoana sensu Maekawa (1971)
Types I and II L. makinoana and L. suzumushi
L. makinoana sensu Nevski (1935), Lee (2002), Eﬁmov (2010) not treated
not treated
L. liliifolia sensu Iinuma and Makino (1913), Hashimoto (1990), Type I
L. suzumushi
Inoue (2016)
L. japonica sensu Satomi (1982), Nakajima (2012)
Type II
L. makinoana
L. japonica sensu Hashimoto (1990), Inoue (2016)
Types II and III L. makinoana and L. longiracemosa
L. japonica sensu Iinuma and Makino (1913), Ohwi (1953,
Type IIIL. longiracemosa1984), Maekawa (1971)
L. japonica sensu Nevski (1935), Kitagawa (1939), Lee (2002), not treated
not treated
Vakhrameeva (2008), Eﬁmov (2010)

in the herbarium of Botanischer Garten und Botanisches Museum, Berlin-Dahlem (B), Germany,but it is thought to have been destroyed during
the World War II (cf. Hiepko, 1987), and no
duplicate was located in any herbaria (Yukawa
and Ohba, 1995). To clarify the nomenclatural
confusions, the protologues of L. japonica and L.
makinoana should be examined carefully, partly
because the type specimens are not available.
Few diagnostic characters are given in the protologue of L. japonica, but we consider it is iden-

tical to Malaxis monophyllos (L.) Sw., as noted
below. Although Type I of the L. makinoana
complex is usually treated as the species L. makinoana, morphological comparison of the three
entities suggests that the protologue of L. makinoana does not agree with Type I, but does agreewith Type II, which is usually called as L. japonica (Table 1). On the contrary, both Type I andType III have no valid scientiﬁc name. We havethoroughly checked descriptions of Liparis taxa
from Japan and adjacent regions. However, weTaxonomic reappraisal of Liparis japonica and L. makinoana

did not ﬁnd any taxa assignable to Types I and
III.Therefore we describe them here as new spe-

109

cies, L. suzumushi and L. longiracemosa, respectively.

Key to Liparis makinoana complex in Japan
1. Flowering in spring to early summer (May to July); labellum ovate or broadly ovate, slightly revolute, with inconspicuous purple veins
2. Flowering in May to June; labellum 14–17 mm long, 11–15 mm wide .................. L. suzumushi (1)2. Flowering in June to July; labellum 8–12 mm long, 5–9 mm wide
3. Labellum 9–12 mm long, 6–9 mm wide; ﬂowers somewhat sparse on inﬂorescence............................................................................................................................L. makinoana (2)3. Labellum 8–10 mm long, 5–7 mm wide; ﬂowers sparse on inﬂorescence ...L. longiracemosa (3)1. Flowering in winter to spring (November to March); labellum broadly ovate, strongly recurved,
with dark purple veins ....................................................................................................... L. hostifolia

Taxonomic treatment
1. Liparis suzumushi Tsutsumi, T.Yukawa et M.
Kato, sp. nov.
(Fig. 2)
Similar to L. makinoana and L. longiracemosa, but has different ﬂowering period (May toJune), shorter inﬂorescence (10–25 cm long), andlarger labellum (14–17 mm long, 11–15 mmwide).
Type. Japan. Shizuoka: Gotenba, Nakabata,
eastern foot of Mt. Fuji, ca. 600 m alt., 8 May
1976, A. Takahashi & F. Konta 10873 (TNS
01281158).
Liparis makinoana auct. non Schltr.: Ohwi, Fl.
Jap.: 379 (1953); Maekawa, Wild Orchids Jap.:
327, pl. 126 (1971) p.p.; Satomi in Satake et al.,Wild Flower Jap. Herb. Pl. 1: 219, pl. 197-2(1982); Nakajima, Ill. Jap. Orchids: 240, pl.
61–12 (2012).
Liparis liliifolia auct. non (L.) A.Rich. ex
Lindl.: Iinuma & Makino, Somoku-Dzusetsu, 3rd
ed. 4: 1231, pl. 67 (1913); Inoue in Iwatsuki etal., Fl. Jap. 4b: 287 (2016).
Japanese name. Suzumushi-so.
Pseudobulb ovoid, 1–3 cm long. Leaves 2,
ovate-elliptic, obtuse or subacute, 10–20 cm long,
2–6(–9) cm wide, conduplicate, glossy, glabrous,
margin entire or somewhat undulate, green; petiole 3–10 cm long, winged. Inﬂorescence termi-

nal, racemose; scape 10–25 cm long, with 4–16
ﬂowers; axis glabrous, ridged, green. Bractovate, acute, 1–5 mm long, green. Pedicellate
ovary clavate, twisted, 12–19 mm long, purplish.
Dorsal sepal linear-lanceolate, subacute, occasionally slightly revolute, erect or somewhat
recurved, 13–16 mm long, 3.5–4 mm wide, green
to purplish. Lateral sepals obliquely ovate or
obliquely lanceolate, subacute, somewhat revolute, 13–16 mm long, 3.5–4 mm wide, greenish.
Petals falcate, linear, obtuse, strongly revolute,pendulous,
sometimes
slightly
twisted,
13–16 mm long, 0.5–1.0 mm wide, purplish.
Labellum entire or minutely erose, broadlyovate, clawed, strongly recurved near base,obtuse or truncate, apiculate at apex, 14–17 mm
long, 11–15 mm wide, purplish or greenish-purple. Column terete, incurved, with rounded wings
in distal part, dilated at base, with shallow
groove at base on ventral side, 5.5–7 mm long,green, pale green on ventral surface, purple atbase; pollinia 4 in 2 pairs, waxy, yellow; anthercap with beaked apex, green.
Distribution. Japan (southern Hokkaido, Honshu, Shikoku), Korea, Russia.
Ecology. Terrestrial, in semi-open deciduous
forests or in semi-open secondary conifer forests.
Flowering season. May to June.
Etymology. It is named after an insect species,
Meloimorpha japonica (“Suzumushi” in Japa-

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Fig. 2. Liparis suzumushi Tsutsumi, T.Yukawa & M.Kato. A: Habit. B: Flower. C: Labellum. D: Dorsal sepal. E:
Petal. F: Lateral sepal. G: Column. H: Anther cap. Scale bars＝3 cm (A), 3 mm (B–G), 1 mm (H). Reproduced
from illustration of ʻL. makinoanaʼ in Nakajima (2012) with permission.

nese), because the lip of the ﬂower somewhat
resembles the wings of male M. japonica.
Notes. Liparis suzumushi is named for Type Iof the L. makinoana complex, which has been
wrongly identiﬁed as L. makinoana. This species
is distinct from the other members of the com-plex in having a different ﬂowering season (Mayto June versus June to July), fewer ﬂowers on ashort inﬂorescence (versus more ﬂowers on along inﬂorescence), and a larger labellum
14–17 mm long and 11–15 mm wide (versus＜13 mm long and ＜10 mm wide).

Taxonomic reappraisal of Liparis japonica and L. makinoana

This species has sometimes been referred to as
L. liliifolia (L.) A.Rich. ex Lindl. (Iinuma andMakino, 1913; Hashimoto, 1990; Hashimoto andKanda, 1991; Inoue, 2016). Liparis liliifolia is
distributed in North America and is phylogenetically distinct from the L. makinoana complex(Tsutsumi et al., 2007; Lee et al., 2010), although
they are morphologically similar. Therefore, we
treat L. suzumushi as a species distinct from L.
liliifolia.
Plants identiﬁed as ʻL. makinoanaʼ also havebeen recorded from Korea and Russia (Nevski,
1935; Lee, 2002). Since we did not examined thematerials from these areas and further analyses
are needed to conﬁrm the distribution of thisentity.
Other specimens examined: JAPAN. Hokkaido. Hidaka: Shizunai-cho, Livestock Farm of
Hokkaido Univ., 04-rinpan, 9 June 1998, H.
Takahashi et al. (SAPS 009837); Shizunai-cho,Livestock Farm of Hokkaido Univ., 02-rinpan,
30 May 2000, H. Takahashi (SAPS 009835);
Shizunai-gun, Shizunai-cho, Misono, Livestock
Farm, Fac. of Agr., Hokkaido Univ., 2-rinpan, 24
May 1995, H. Takahashi & T. Sato 17817 (SAPS
009838). —Iburi: Kojyohama, 26 May 1951, S.
Kono (TI); Tomakomai-shi, Experimental Forestof Hokkaido University, 412–421-rinpan, 31
May 1990, H. Takahashi et al. 10173 (SAPS
009840); Tomakomai, 29 June 1915, Y. Kudo &T. Yoshimi (SAPS 005678). —Ishikari: Chitose, 3June 1975, S. Kurosawa & Y. Tateishi (TI). —Oshima: Hakodate, 3 June 1916, F. C. Greatrex(SAPS 009846); Kamiiso-gun, Kamiiso-cho,near Kamiiso dam, along small valley, 1 June
1992, H. Takahashi et al. 11637 (SAPS 009839).
—Shiribeshi: 21 April 1931, K. Tagashi 21758(SAPS 009842). Akita. Kitaakita-gun, Takanosumachi, 10 June 1969, R. Kumikawa (TNS
255710); Oga, Honzan, 18 June 1933, S. Muramatsu (TI); Ugo, Hachimori, 27 May 1932, S.
Muramatsu (TI). Miyagi. Shiraishi-shi, Hachimoriyama, ca. 450 m alt., 14 May 1978, T. Sasaki& Y. Yamamoto (TUS 45989). Niigata. Nishikubiki-gun, Ichiburi, 31 May 1960, collecter
unknown (KYO); Sanjo-shi, 23 May 1943, Naka-

111mura (TUS 168196). Ibaraki. Kuji-gun, Kegano-mura, 25 May 1935, H. Tsurumachi (TUS
179702); Naka-gun, Yamagata-mahi, cliff in
Ryujin river, 5 May 1969, collecter unknown(TNS 253959); Naka-gun, Yamagata-machi,
along Ryujin-gawa River, 26 May 1935, H. Tsurumachi (TUS 169489). Kanagawa. Around Mt.
Tanzawa, Yozukugawa, Nyudodana, 3 May
1956, H. Kanai (TI); Hakone, 12 April 1953, K.
Segawa (TI); Mt. Hakone, 15 May 1925, S.
Muramatsu (TI); Mt. Hakone (Mt. Kami), 3 June
1953, J. Haginiwa JH021248 (TNS 971248);
Ashigara, C. Tsutsumi (TNS 8505372–4). Shizuoka. Sunto-gun, Subashiri-mura, 30 May
1931, K. Segawa (TI); Suntou-gun, Oyama-cho,
Nakajima, 400 m alt., 20 May 1998, E. Aihara(TNS 01281159); Gotenba-shi, Yaga, 18 May
1969, K. Inoue (TNS 281001). Ehime. Nii-gun,Nakahagi-mura, beginning of June 1930, H.
Yamamoto (KYO). Kochi. Kitakawa-mura, 17
May 1888, K. Watanabe (TNS 62606); Yuzuhara-mura, en route from Tanono to Onogahara,
800 m alt., 24 May 1959, G. Murata & T. Shimizu (KYO).
2. Liparis makinoana Schltr.

(Fig. 3)
Liparis makinoana Schltr. in Repert. Spec.
Nov. Regni Veg. Beih. 4 [Orch. Sin.-Jap. Prodr.]:
63 & 200 (1919); Maekawa in Wild Orchids Jap.:
327, pl. 126 (1971) p.p.
Neotype designated here. Japan. Hokkaido:
near Hakodate (Nodafu in Momijiyama-cho), inmossy, wet place [Japonica, insula Jesso, circa
Hakodate, Nodafu, in muscosis udis], Dr.
Albrecht, 22 June or 4 July 1861 (LE 01016040),
specimen at left bottom (Fig. 4, arrow).
Liparis japonica auct. non (Miq.) Maxim.:
Satomi in Satake et al., Wild Flower Jap. Herb.
Pl.1: 219, pl. 197-1 (1982); Nakajima, Ill. Jap.
Orchids: 234, pl. 61-06 (2012); Inoue in Iwatsuki
et al., Fl. Jap. 4b: 286 (2016), p.p.
Japanese name. Seitaka-suzumushi-so.
Pseudobulb ovoid, 1–3 cm long. Leaves 2,
ovate-elliptic, obtuse or subacute, 7–20 cm long,
2–6 cm wide, conduplicate, glossy, glabrous,

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Fig. 3. Liparis makinoana Schltr. A: Habit. B: Flower. C: Labellum. D: Dorsal sepal. E: Petal. F: Lateral sepal.
G: Column. H: Anther cap. Scale bars＝3 cm (A), 3 mm (B–G) or 1 mm (H). Reproduced from illustration ofʻL. japonicaʼ in Nakajima (2012) with permission.

margin entire or somewhat undulate, green; petiole 3–10 cm long, winged. Inﬂorescence terminal, racemose, 10–35 cm long, with 4–30 ﬂowers; axis glabrous, ridged, green. Bract ovate,
acute, 1–5 mm long, green. Pedicellate ovary clavate, twisted, 8–13 mm long, green to purplish.
Dorsal sepal linear-lanceolate, subacute, occasionally slightly revolute, erect or somewhat
recurved, 9–12 mm long, 2–2.5 mm wide, green.
Lateral sepals obliquely ovate or obliquely lanceolate, subacute, somewhat revolute, 8–11 mm
long, 2–3 mm wide, green. Petals falcate, linear,obtuse, strongly revolute, pendulous, sometimes
slightly twisted, 9–12 mm long, 0.5–1 mm wide,
purplish. Labellum entire or minutely erose,
ovate, clawed, strongly recurved in basal part,obtuse, apiculate at apex, 9–12 mm long, 6–9 mm
wide, purplish or greenish-purple. Column terete,
incurved, with rounded wings in distal part,
dilated at base, with shallow groove at base on
ventral side, 4–5 mm long, green, pale green on
ventral surface, purplish in basal part; pollinia 4
in 2 pairs, waxy, yellow; anther cap with beaked
apex, green.

Taxonomic reappraisal of Liparis japonica and L. makinoana

113

Fig. 4. Herbarium sheet designated as neotype of Liparis makinoana by Eﬁmov (2010), in which plant marked
by arrow is here designated as neotype of Liparis makinoana.

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Distribution. Japan (Hokkaido, Honshu),
China, Korea, Russia.
Ecology. Terrestrial, in semi-open secondary
conifer forests or in semi-open deciduous forests.
Flowering season. June to July.
Notes. Liparis makinoana was described by
Schlechter (1919), based on the type specimen(destroyed during World War II) collected in
Hokkaido, where L. suzumushi and L. longiracemosa (see below) also occur. Most of the characters in the protologue were unclear for identiﬁcation purposes, but the labellum size (1.2 cm long,
8 mm wide) is similar to Type II of the L. makinoana complex.
Eﬁmov (2010) designated a herbarium sheet,
LE 01016040, collected near Hakodate, Hokkaido, as the neotype. The sheet has plural plants.
The plant at the bottom left has ﬂower characters, such as scape length, ﬂower number, andlabellum size, similar to those of Type II, and isdesignated here as neotype of L. makinoana
according to Art. 9.17 of the current International
Code of Nomenclature for algae, fungi, andplants (Turland et al., 2018). The central large
plant is L. koreojaponica Tsutsumi, T.Yukawa,
N.S.Lee, C.S.Lee et M.Kato and the remaining
small plants at the right side and the top left arethought to be L. makinoana or L. suzumushi.
Liparis makinoana in the present concept has
been referred to ʻL. japonicaʼ in several previous
literature on Japanese ﬂora (Table 1). ʻLiparis
japonicaʼ mentioned in some ﬂoristic treatments
in China, Korea, and Russia (Nevski, 1935; Lee,
2002; Chen et al., 2009, as a synonym of L. elongata Fukuy.), along with L. koreana (Nakai)
Nakai ex W. T. Lee described from the northernpart of Korean Peninsula, is also similar to L.
makinoana, and further analyses are needed todecide their taxonomic treatment.
Other specimens examined: JAPAN. Hokkaido. Hidaka: Mt. Apoi, 9 July 1933, H. Hara
(TI); Mt. Horoman, 11 July 1933, H. Hara (TI);
Saru-gun, Monbetsu-cho, Satohira, 7 July 1963, Y.
Takahashi (KYO). —Iburi: Abuta-gun, Touyamura, Tomioka, 450–620 m alt., 29 June 1991, Y.
Tateishi et al. (TUS 155386). —Ishikari: Mt.

Teine, 5 July 1912, S. Hayakawa (TI). —Kamikawa: Tohma-cho, 310 m alt., 9 July 2007, C. Tsutsumi et al. (TNS 766277); Kamuikotan, June
1913, H. Koidzumi (TNS 416409). —Kushiro:
Shitakara-mura, Setsuri, June 1895, S. Ito (SAPS
009862); Akan-gun, Tsurui-mura, 30 m alt., 20
July 1997, K. Takita (TNS 01091437); Akkeshi, 5July 1917, Y. Kudo (TUS 26552). —Oshima:
Hakodate, 7 July 1918, F. C. Greatrex (SAPS
009864); Yunokawa, 4 July 1916, F. C. Greatrex(SAPS 009855); Yunokawa, near Hakodate, 25
June 1921, I. Yamamoto (SAPS 009853); near
Kamiiso, 25 July 1915, F. C. Greatrex (SAPS
009824). Iwate. Kamiihei-gun, Kurihashi-mura,
Y. Kudo (KYO). Miyagi. Kurihara-gun, Tsukidate-cho, date unknown, collecter unknown (TNS
01064095). Gunma. Sanpei-pass, near Oze, 17
July 1946, G. Nakai 2717 (KYO). Tochigi. Shiohara, Shonoyu, 23 July 1887, K. Nemoto (TNS
32642). Kanagawa. Mt. Tanzawa, 9 July 1961, J.
Haginiwa JH021249 (TNS 971249); Hakonesagamiguchi, 23 July 1935, A. Kojima (TNS 702484).
Yamanashi. Oshino, 5 July 1980, J. Satoh (TI).
Shizuoka. Mt. Fuji, Subashiriguchi, 6-goume, 4August 1908, I. Amamiya (TNS 16992). Nagano.
Komoro-machi, Goka, Mt. Morizumi, 31 August
1933, K. Misaizu (KYO); Minamisaku-gun,
Kawakami-mura, Azusayama, 16 July 1976, J.
Haginiwa JH021250 (TNS 971250).
3. Liparis longiracemosa Tsutsumi, T.Yukawa et
M.Kato, sp. nov.
(Fig. 5)
Similar to L. makinoana, but different in termsof smaller labellum, the wider sepals, and thesparser ﬂowers on the inﬂorescence.
Liparis japonica (Miq.) Maxim. sensu Maximowicz in Bull. Acad. Imp. Sci. Saint-Pétersbourg 31: 102 (1886), p.p.; Maximowicz in Mel.
Biol. 12: 544 (1887), p.p.; Iinuma & Makino,
Somoku-Dzusetsu, 3rd ed. 4: 1231, pl. 67(1913); Ohwi, Fl. Jap.: 379 (1953); Maekawa,
Wild Orchids Jap.: 331, pl. 128 (1971); Inoue inIwatsuki et al., Fl. Jap. 4b: 286 (2016), p.p.
Type: Kitaakita-shi, Anitotorinai, Akita Pref.,Japan. 530 m alt., 23 June 2008. C. Tsutsumi, K.

Taxonomic reappraisal of Liparis japonica and L. makinoana

115

Fig. 5. Liparis longiracemosa Tsutsumi, T.Yukawa & M.Kato. A: Habit. B: Flower. C: Labellum. D: Dorsal
sepal. E: Petal. F: Lateral sepal. G: Column. H: Anther cap. I: Pollinia. Scale bars＝3 cm (A), 3 mm (B–G) or
1 mm (H–I). Reproduced from illustration of ʻLiparis sp.ʼ in Nakajima (2012) with permission.

Miyoshi, M. Taguchi & Kon CT1608 (holotype:TNS 778816).
Distribution. Japan (Hokkaido, Honshu, Shikoku, Kyushu).
Ecology. Growing in semi-open, somewhat
damp places in deciduous or mixed coniferdeciduous forests.
Flowering season. June to July.
Japanese name. Akita-suzumushi-so.
Etymology. The species is so named as it has along inﬂorescence.
Pseudobulb ovoid, 1–3.5 cm long. Leaves 2,
ovate-elliptic, obtuse or subacute, 10–20 cm long,

2–6 (–9) cm wide, conduplicate, glossy, glabrous, margin entire or somewhat undulate,green; petiole 3–10 cm long, winged. Inﬂorescence terminal, racemose, 15–40 cm long, with
4–40 ﬂowers; axis glabrous, ridged, green. Bractovate, acute, 1–5 mm long, green. Pedicellate
ovary clavate, twisted, 10–20 mm long, purplish.
Dorsal sepal linear-lanceolate, subacute, occasionally slightly revolute, erect or somewhat
recurved, 8–11 mm long, 2.5–3.5 mm wide, purplish. Lateral sepals obliquely ovate or obliquely
lanceolate, subacute, somewhat revolute,
8–11 mm long, 2.5–3.5 mm wide, purplish. Petals

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falcate, linear, obtuse, strongly revolute, pendulous, sometimes slightly twisted, 9–12 mm long,
0.5–1.0 mm wide, purplish. Labellum entire orminutely erose, ovate, recurved at base, obtuse,
apiculate at apex, 8–10 mm long, 5–7 mm wide,
dark purple, purplish, or green. Column terete,
incurved, with rounded wings in distal part,
dilated at base, with shallow groove at base on
ventral side, 4–5 mm long, green, pale green on
ventral surface, purplish in basal part; pollinia 4
in 2 pairs, waxy, yellow; anther cap with beaked
apex, green to purplish.
Notes. Liparis longiracemosa is similar to L.
makinoana, but distinguishable by the smaller
labellum (8–10 mm×5–7 mm vs. 9–12 mm×
6–9 mm in L. makinoana), wider sepals(8–11 mm×2.5–3.5 mm vs. 8–11 mm×2–3) mm,
and sparser ﬂowers on the inﬂorescence whencompared with the latter. Flower color varies
from green to dark purplish.
Other specimens examined: JAPAN. Hokkaido. Hidaka: Shizunai-gun, Shizunai-cho.
Misono, Livestock Farm, Fac. Of Agr., Hokkaido
University, 12-rinpan, 13 July 1996, H. Takahashi 20548 (SAPS 009767); Shizunai-gun, Shizunai-cho, Misono, Livestock Farm, Hokkaido
University, 12-rinpan, 1 July 1998, H. Takahashi
25064 (SAPS 009765); Shizunai-gun, Shizunaicho, Misono, Livestock Farm, Hokkaido University, 12-rinpan, 1 July 1998, H. Takahashi 25064(SAPS 009766). —Iburi: Hayakita, 16 June
1956, K. Ito & A. Umezawa (SAPS 009763);
Tomakomai, 18 July 1994, M. Nakajima (TI);
Tomakomai, Experimental Forest of Hokkaido
University, 31 July 1933, H. Hara (TI); Yuufutsu-gun, Hobetsu-cho, 29 July 1991, K. Ito(SAPS 009769); Outside of Toyoura-cho, 25
June 1990, Y. Goda 797 (KYO); Mt. Usu, 23 July
1939, I. Yamamoto (KYO). —Kamikawa: C. Tsutsumi (TNS 8505378–81). —Shiribeshi: Around
Ezofuji, Hangetsu lakeside, 15 July 1951, Y.
Kuwabara (SAPS 009783); Souya, Sarufutsu,
26-rinpan, July 1931 (SAPS 009784). Aomori.
Higashitsugaru-gun, Tairadate, Yunosawa, 28
July 1959, H. Hara (TI). Akita. 13 June 1983,
Yoshie Hanei (TNS 9504246–7); Kitaakita-shi,

Anitotorinai, 530 m alt., 23 June 2008, C. Tsutsumi et al. (TNS 778815). Miyagi. Mt. Daito, 6
June 1959, H. Ohashi 1980 (TI). Niigata.
Itoigawa-shi, near Tsukimizu-no-ike, 6 June
1971, S. Iwano 14714 (TUS 146016); Toyama.
Kurobe canyon, 9 June 1960, N. Fukuoka(KYO). Fukui. National forests of Kurokawa, 10
June 1939, K. Hirobe (KYO). Nagano. Shimoina-gun, Urugi-mura, Mt. Chausu, 1200 m
alt., 13 June 1957, T. Yamazaki (TI). Shiga.
Shiga-gun, Mt. Hira, 11 June 1944, G. Nakai(KYO); Mt. Watamuki, 19 June 1937, C. Hashimoto (KYO). Kyoto. Ayabe-shi, Okukanbayashi,
Wasadani, ca. 300 m alt., 14 June 1980, G.
Murata 41065 (KYO); Miyadzu-shi, Sugiyama,
13 June 1997, S. Tsugaru et al. (KYO); Hosonomura, Mt. Ryugadake, 29 July 1933, M. Tagawa(KYO). Osaka. Mt. Kongo, 14 June 1931, Z.
Tashiro (KYO). Hyogo. Mt. Rokko, 23 June
1929, Z. Tashiro (KYO). Kagawa. Mt. Otaki, 26
June 1932, Z. Tashiro (KYO). Ehime. Kawanouchi, 21 July 1915, H. Yamamoto (KYO); Syusogun, Ishine-mura, Yasuitani (KYO). Kochi.
Agawa-gun, Nanokawa-mura, Shimonanokawa,
K. Watanabe (TI); Nanogawa, 30 June 1887, K.
Watanabe (TNS 62607); Nanokawa-mura, 7 June
1888, K. Watanabe (TI). Miyazaki. Gokase-cho,
Mt. Shiraiwa, Yukawa 07-21 (TNS 8505608).
Interpretation of Liparis japonica
Liparis japonica was described as Microstylis
japonica by Miquel (1866) based on a specimen
with fruits only, collected in Japan by H. Bürger.
The type specimen has not been found (Ohba etal., 2005; A. Schuiteman, pers. comm.). We reexamined the protologue and found it to be identical to Malaxis monophyllos (L.) Sw. (1) The
description “labellum apice 3 apiculis incrassates?” (3 apiced labellum) is a signiﬁcant diagnostic character of M. monophyllos, whereas it is
not evident in the L. makinoana complex. (2)The description “An M. diphyllae Lindl. L. c. p.
19 afﬁnis?” suggests an afﬁnity of this entity to
Microstylis diphyllos, which is currently treatedas conspeciﬁc with Malaxis monophyllos. Therefore, we conclude that Microstylis japonica is a

Taxonomic reappraisal of Liparis japonica and L. makinoana

synonym of Malaxis monophyllos, as given
below.
Maximowicz (1886) transferred Microstylis
japonica to Liparis with citations of Miquel(1866), Iinuma (1874), and Franchet and Savatier(1875). The specimen cited by Franchet andSavatier (1875), which is deposited in Muséum
National dʼHistoire Naturelle, Paris, France (P),is L. krameri Franch. et Sav., and the illustrationof Iinuma (1874) is assignable to L. longiracemosa. Therefore, L. japonica sensu Maxim. is aheterogeneous assemblage of multiple Liparis
species.
Malaxis monophyllos (L.) Sw. in Kongl. Vetensk.
Acad. Nya Handl. 21: 234 (1800).
Microstylis japonica Miq., Ann. Mus. Bot.
Lugduno-Batavi 2: 203 (1866), syn. nov. —Liparis japonica (Miq.) Maxim. in Bull. Acad.
Imp. Sci. Saint-Pétersbourg 31: 102 (1886), excl.
cit. & descr.; Maximowicz in Mel. Biol. 12: 544(1887), excl. cit. & descr.
Acknowledgments
We thank P. Eﬁmov, M. Goto, H. Hongo,
X.-H. Jin, Kon, C.-S. Lee, N.-S. Lee, A. Maeda,
T. Minamitani, K. Miyoshi, H. Nakayama, K.
Oikawa, A. Okuda, K. Sakamoto, M. Taguchi, H.
Takeuchi, K. Tanaka, K. Watanabe, C.-K. Yang,and Y. Yamashita for help in our ﬁeld trips andplant sampling. We also thank T. Kurosawa andH. Nagamasu for useful comments, M. Nakajima
and Bunʼichi Sogo Shuppan for the illustrations,and K. Suzuki for cultivating the living plants.
We are grateful to the directors and curators of
SAPS (H. Takahashi), TUS (K. Yonekura), KYO(H. Nagamasu), and TI (A. Shimizu) for allowing us to study the herbarium specimens. This
study was partly supported by the National
Museum of Nature and Science research grant
entitled “Integrated Analysis of Natural History
Collections for Conservation of Highly Endangered Species”.

117

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