Bull. Natl. Mus. Nat. Sci., Ser. B, 35(2), pp. 57–62, June 22, 2009

A Taxonomic Reappraisal of Neottia (Orchidaceae) from East AsiaTomohisa Yukawa1, Takahiro Yagame2 and Nam Sook Lee3
1Tsukuba Botanical Garden, National Museum of Nature and Science, Amakubo, Tsukuba, 305–0005 Japan
2
Orchid Museum Takamori, Takamori, Shimoina, Nagano, 399–3107 Japan3
Department of Life Sciences, Ewha Womans University, Seoul, 120–750 Korea

Abstract Two taxonomic problems in the genus Neottia (Orchidaceae) are elucidated. (1) N. kiusiana T. Hashim. et Hatus. and N. hypocastanoptica Y. N. Lee are rare and obscure entities previously known only from the type collections. We examined new material referable to these species
in Japan and Korea and concluded that they are conspecific. Neottia kiusiana becomes a correctname due to the priority of publication. (2) We found that Archineottia japonica M. Furuse shouldbe transferred to the genus Neottia. However, the combination N. japonica was already used for adifferent species. We therefore named N. furusei T. Yukawa et T. Yagame for this entity.
Key words : Japan, Korea, Neottia, new combination, Orchidaceae, taxonomy.

Identification of two obscure species of Neottia
The genus Neottia Guett. comprises about 10species distributed widely in the temperate zoneof Eurasia and adjacent regions. The genus is circumscribed by a combination of characters, suchas 1) achlorophyllous habit, 2) densely crowded,
fleshy, short roots, 3) an erect stem covered withclasping sheaths, 4) an elongated column with asuberect anther, 5) two, soft, mealy pollinia, and6) a sensitive rostellum. In Japan, the following
taxa are recognized: N. asiatica Ohwi, N. inagakii Yagame, Katsuy. et T. Yukawa, N. kiusiana
T. Hashim. et Hatus., N. nidus-avis (L.) Rich. var.
nidus-avis, and N. nidus-avis (L.) Rich. var.
mandshurica Kom.
Among them, Neottia kiusiana is an obscureentity previously known only from the type collection. The species was described in 1991 on thebasis of a specimen collected in Kagoshima Prefecture, southern part of Kyushu Island, Japan(Hashimoto and Hatusima, 1991).
Subsequently, Neottia hypocastanoptica Y. N.
Lee (Lee, 2002) was described from Jeju Island,Korea. Although the author did not compare thenew species with N. kiusiana in the protologue,
the description suggests its close affinity with N.

kiusiana. We compared several flowering samples collected at the type locality of N. hypocastanoptica with N. kiusiana.
Furthermore, we found two more Japanesecollections possibly referable to N. kiusiana. One
is a herbarium specimen collected by Daisuke
Shimizu in Chiba Prefecture, southeastern part ofHonshu Island, Japan. The other is herbariumand spirit material collected by Jun-nosukeÔhara in Aichi Prefecture, central part of Honshu
Island, Japan.
We compared morphological characters ofthese specimens. Figure 1 shows a flowering
plant from the type locality of Neottia hypocastanoptica in Jeju Island. Both vegetative and reproductive characters are consistent among thespecimens observed except for the shape ofepichile of lip (Fig. 2). However, we found thischaracter variable even among individuals in thesame population. As shown in Figures 1G and
2D, two individuals of Jeju population exhibit
different shape of epichile lobe: one is rectangular and the other is transversely rectangular. With
respect to the type material of N. kiusiana, although the holotype has a transversely obliquerectangular epichile lobe (Fig. 1I in Hashimotoand Hatusima, 1991), the relevant part of the iso-58

T. Yukawa, T. Yagame and N. S. Lee

Fig. 1. Neottia kiusiana T. Hashim. et S. Hatus. A. Habit. B. Flower, side view. C. Lip and column, side view. D.
Dorsal sepal. E. Petal. F. Lateral sepal. G. Lip. H. Column, from below. I. Column, side view. J. Anther cap.
Drawn from N. S. Lee & C. S. Lee D-241 (A) and N. S. Lee & C. S. Lee L-016 (B–J: topotype of N. hypocastanoptica) by M. Nakajima. Scale bar3 cm (A), 3 mm (B–G), or 1 mm (H–J).

A Taxonomic Reappraisal of Neottia from East Asia

59

Fig. 2. Variations of lip in Neottia kiusiana T. Hashim. et S. Hatus. Drawn from Y. Ôhira s.n. (A: isotype of N.
kiusiana), J. Ôhara s.n. (B), D. Shimizu s.n. (C), and Yukawa 08-9 (D: in the vicinity of the type locality of N.
hypocastanoptica) by M. Nakajima. Scale bar 1 mm.

type is oblong (Fig. 2A). This variation of lip indicates that taxonomic subdivision on the basisof epichile shape is not appropriate. Consequently, we concluded that all of the material used inthis study belongs to the same single species.
Since the publication of N. kiusiana precedes thatof N. hypocastanoptica, the former becomes acorrect name.
These findings shed more lights on the geographical and ecological features of the species.
The distribution range is from Jeju Island in thewest to Kyushu Island, central and southeasternparts of Honshu Island in the east. These records
also indicate very rare, scattered distribution ofthe species. Hashimoto and Hatusima (1991) did
not provide ecological information on the type
locality of Neottia kiusiana except for its occurrence below 100 m above sea level where it is assumed to develop warm-temperate evergreen
broad-leaved forest. The habitat in Jeju Island isa similar forest type dominated by Castanopsis
trees at 300–330 m altitude high (Lee, 2002; T.

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T. Yukawa, T. Yagame and N. S. LeeYukawa, unpublished). Ôhara (1967) recordedthat the Aichi plant was collected in a forest
dominated by Quercus spp., Litsea coreana H.
Lév., Daphniphyllum macropodum Miq., and Aucuba japonica Thunb. at 500 m altitude high. The
collector’s notes on the sheet of Chiba material
stated that the habitat is a humus-rich forest floor
on the northeast slope where the vegetation iscomposed of Castanopsis sieboldii (Makino)
Hatus. ex T. Yamaz. et Mashiba, Quercus glauca
Thunb., Camellia sasanqua Thunb., and Illicium
anisatum L.
Therefore, the vegetation type associated withthis species is likely confined to warm-temperate
evergreen broad-leaved forest. By contrast, Neottia nidus-avis, a closely related species to N. kiusiana, inhabits subarctic coniferous forest andcool temperate deciduous broad-leaved forest.
The vegetation type is crucial for establishmentof Neottia species because they represent obligate mycoheterotrophy and at least, N. nidus-avis
is known to obtain nutrients from Sebacina spp.,
a group of tree ectomycorrhizal fungi, nutritionally dependent on particular tree species (McKendrick et al., 2002; Selosse et al., 2002). In thisrespect, habitat differentiation between the twospecies infers a shift of mycorrhizal fungi between them.
Neottia kiusiana is most closely related to N.
inagakii, but they can separate each other by thefollowing combinations of characters; 1) the lipof N. kiusiana is longer and wider than that of N.
inagakii; 2) the lip of N. kiusiana is inverselytrapeziform in outline and 2-lobed, whereas thatof N. inagakii is oblong in outline and 2- or 3lobed; 3) the epichile is well-developed andopens flat in N. kiusiana but is ill-developed andfolded adaxially in N. inagakii; 4) the anther ofN. kiusiana bears a short filament, while that ofN. inagakii does not have a filament.
Neottia kiusiana T. Hashim. et S. Hatus.,
Ann. Tsukuba Bot. Gard. (10): 41 (1991). TYPE:
JAPAN: Kyushu, Kagoshuma Pref., Satsumagun, Tsuruda-cho, Tsuruda, south from Tsuruda
Dam, 13 May 1991, Yutaka Ôhira s. n. (TNS–holotype).
Neottia hypocastanoptica Y. N. Lee, Fl. Korea,
ed. 4: 1167 (2002), syn. nov. TYPE: KOREA:
Jeju Is., Mt. Halla, alt. 300 m, 2001, Y. Lee, Y.
Shin et K. Lee s. n. (Korean Plant Research Institute–holotype).
Neottia nidus-avis (L.) Rich. var. mandshurica
auct. non Kom.; Ôhara, Shokubutsu KenkyûShûroku 11: 48 (1967).
Plant terrestrial, achlorophyllous, glossy lightbrown, 6–21 cm tall. Roots fascicled, ageotropic,
facing upward, cylindrical–slightly fusiform, up
to 2.7 cm long, 2–2.3 mm in diameter. Stemerect, terete, glabrous, ivory white, 3.5–4 mm indiameter, dark brown when dried. Sheaths 3–5,foliaceous, spathulate, subacute, glabrous, tubular, membranaceous, upper ones much longer
than lower ones, up to 25 mm long. Inflorescenceterminal, erect, racemous, with 10–28 flowers,4.6–13 cm long, rachis sparsely glandular hairy.
Floral bracts membranaceous, lowermost ones
much longer than flowers, gradually diminishingin upper ones of which shorter than ovaries, linear, ovate-lanceolate, obtuse to subacute, 1- to 4nerved, 3–20 mm long, 1.8–12 mm wide. Flowers
brownish ivory, porrect. Pedicel and ovary
suberect in flowering, 6–15 mm long, 3–4.5 mm
in diameter; pedicel terete, glaborous; ovary
clavate-obovoid, glandular hairy, longer thanpedicel except for the lowermost. Dorsal sepaloblong-obovate, obtuse, cucullate, 1-nerved,3.5–4 mm long, 2.2–2.7 mm wide when spreadout. Lateral sepals obliquely oblong-obovate, obtuse, strongly cupped, cucullate, 1-nerved,
3.2–4.3 mm long, 2.1–2.4 mm wide when spreadout. Petals obliquely obovate-ovate, obtuse, 1nerved, 3.5–5.2 mm long, 2.2–2.7 mm wide. Lip
inversely trapeziform in outline, anteriorly 2lobed, 5.2–8 mm long, 5.5–7.5 mm wide when
spread out; hypochile rectangular-inversely
trapeziform, shallowly concave at base, adaxiallypurple-dotted; epichle 2-lobed, lobes transversely
oblique-rectangular, rectangular, or oblong. Column cylindrical-slightly conical, 1.7–3.5 mm
long; stigma lamellate, erect in flowering, 2- to 3lobed, inveresely trapeziform-transversely rectan-

A Taxonomic Reappraisal of Neottia from East Asia

gular in outline; rostellum erect in flowering, oblong-narrowly triangular, obtuse. Anther erectdecumbent in flowering with a short, often insignificant filament.
Japanese name: Tsukushi-sakane-ran.
Flowering period: Mid-May to June.
Distribution: Japan: Honshu (Chiba Pref.,Aichi Pref.), Kyushu (Kagoshima Pref.); Korea:
Jeju-do, Seogwipo-si.
Habitat: Floor of warm temperate evergreen
broad-leaved forest.
Other specimens examined: JAPAN, Honshu:
Chiba Pref., Mt. Kiyosumiyama, alt. ca. 430 m[sic], 15 June 1958, D. Shimizu s. n. (TNS307293); Aichi Pref., Higashikamo-gun, Asahimura, Ikuma, alt. ca. 500 m, June 1966, J. Ôharas. n. (AICH, TNS 287241); KOREA, Jeju-do:
Seogwipo-si, Seon-dol, 19 May 2004, N. S. Lee& C. S. Lee D-241 (Plant Systematics Laboratory of Ewha Womans University); Seogwipo-si,Don-nae-ko (topotype of N. hypocastanoptica),
19 May 2004, N. S. Lee & C. S. Lee L-016(Plant Systematics Laboratory of Ewha WomansUniversity); Seogwipo-si, Seon-dol, alt. 330 m,
25 May 2008, Yukawa 08-9 (TNS 8501018).
Nomenclatural problem ofArchineottia japonica
Chen (1979) emphasized primitive status of aterminal stigma, an erect stamen with its free filament, and absence of clinadrium and rostellumshared by several Neottia species in line with his
phylognetic hypothesis of Orchidaceae. He thus
segregated these species from Neottia and established a new genus Archineottia. Subsequently,
Furuse and Chen (1988) found a new species exhibiting the above-mentioned characters andnamed Archineottia japonica M. Furuse.
Hashimoto and Hatusima (1991) suspected theindependent status of Archineottia because a filament, an important diagnostic character of thegenus, is also found in several Neottia species.
Inoue and Ikegami (1997) and Imai and Inoue(1998) adopted this view and proposed a new
combination, Neottia japonica (M. Furuse) K.

61Inoue. Molecular phylognetic analyses of tribe
Neottieae also confirmed the inclusion of Archineottia into Neottia (Yukawa and Yagame, unpublished data). Unfortunately, the combination
Neottia japonica was already used by Szlachetko(1995) for an entity on the basis of a different
type, thereby rendering N. japonica (M. Furuse)
K. Inoue illegitimate as a later homonym. A new
name is therefore needed for the latter species
when we treat this in the genus Neottia.
Neottia furusei T. Yukawa et Yagame, nom.
nov.
Archineottia japonica M. Furuse in Acta Phytotax. Sin. 26: 69 (1988). TYPE: JAPAN: Honshu, Nagano Pref., Kami-ina-gun, Hase-mura,Todai, alt. 1200 m, on limestone rocky slope inbroad-leaved deciduous forest, 4 September
1977, Miyoshi Furuse 12452 (PE–holotype).
Holopogon japonicus (M. Furuse) S. C. Chenin Acta Phytotax. Sin. 35: 180 (1997).
Neottia japonica (M. Furuse) K. Inoue in Acta
Phytotax. Geobot. 49: 199 (1998), non Neottiajaponica (Blume) Szlach. in Fragm. Florist.
Geobot., Suppl. 3: 117 (1995).
The species epithet refers to the late Mr.
Miyoshi Furuse, a prominent plant collector, who
discovered and described this species.
Acknowledgments
We would like to thank Shunsuke Serizawa,
Kyung-Seo Lee, and Won-Bok Kim for providing
the plant material, Teruo Katsuyama for sharing
his observations, and Mutsuko Nakajima for
preparing excellent illustrations. This study ispartly supported by a Grant-in-Aid to Scientific
Research from the Japan Society for Promotionof Science (No. 17370033) and a research grantfrom the National Museum of Nature and Science for T. Yukawa, and a research grant fromthe Korea Research Foundation Grant fundedby the Korean Government (KRF-2007-031C00003) for N. S. Lee.

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T. Yukawa, T. Yagame and N. S. Lee

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