Bull. Natl. Mus. Nat. Sci., Ser. B, 45(3), pp. 119–129, August 22, 2019

Taxonomic Status of Lecanorchis suginoana (Tuyama)
Seriz. (Orchidaceae)
Michihisa Takashima1,2,*, Yumi Yamashita3, Akihiko Kinoshita3,4,
Mineo Ohta5, and Tomohisa Yukawa3
1

Tochigi Prefectural Museum, 2–2 Mutsumicho, Utsunomiya, Tochigi, 320–0865 Japan
2
Ryokusei-kenkyusho, 2–40–10 Kojimacho, Chofu, Tokyo, 182–0026 Japan
3
Tsukuba Botanical Garden, National Museum of Nature and Science,
4–1–1 Amakubo, Tsukuba, Ibaraki, 305–0005 Japan
4
Kyushu Research Center, Forestry and Forest Products Research Institute,
4–11–16 Kurokami, Kumamoto, 860–0862 Japan
5
Yuzen-sha, 1285–13 Takagosho, Kakegawa, Shizuoka, 436–0031 Japan* E-mail: m-takashima@ryokusei-ri.co.jp(Received 13 May 2019; accepted 26 June 2019)

Abstract Lecanorchis suginoana (Tuyama) Seriz. has been interpreted as a variety of L. japonica Blume, a variety of L. kiusiana Tuyama, or an independent species. To re-evaluate the taxonomic status of this entity, macromolecular, morphological, and ecological characters were investigated. Divergence of nucleotide sequences supported independent status of three close relatives,viz., L. suginoana, L. kiusiana, and L. japonica. Furthermore, the results of molecular phylogenetic analysis did not support infraspeciﬁc placement of L. suginoana in L. japonica. Moreover,ﬂower colour, qualitative characters of labellum and column, quantitative characters of perianth
lobes and column, ﬂowering time, and associated vegetation types clearly distinguish L. suginoana
from L. kiusiana. These differences corroborated the independent species status of L. suginoana
rather than the infraspeciﬁc position of L. kiusiana.
Key words : Japan, Lecanorchis, molecular analysis, morphological analysis, Orchidaceae, taxonomy.

Introduction
The genus Lecanorchis Blume is a terrestrial,
achlorophyllous orchid genus in which about 25
species are distributed throughout temperate totropical regions in East to Southeast Asia andextending to New Guinea (WCSP, 2019). This
group represents holomycoheterotrophy in theirnutritional mode (Motomura et al., 2010) anddepends upon carbon nutrition from particulargroups of ectomycorrhizal fungi belonging to
Basidiomycota (Okayama et al., 2012).
Lecanorchis japonica Blume var. suginoana
Tuyama is one of the northernmost distributed
members of the genus (Yukawa, 2015a, 2015b;
Suetsugu et al., 2018). In the original descrip-

tion, Tuyama (1982) pointed out close morphological afﬁnities of this entity to L. japonica var.
japonica and emphasized similarity of the shapeof apical part of the column between them.
In his revisionary work of the genus, Hashimoto(1990) proposed a new combination, L. kiusiana
Tuyama var. suginoana (Tuyama) T.Hashim. He
demonstrated common characters between thisentity and L. kiusiana var. kiusiana, i.e., rather
small plant, lip with marginal papilla and conspicuous branchlets of disc-hairs, ovary with anapparent ring-like excrescence below the calyculus, and rather short fruit. Subsequently, Serizawa (2005) proposed to recognize it as an independent species L. suginoana (Tuyama) Seriz.
He emphasized that this entity can be distin-Michihisa Takashima et al.

120

guished from L. kiusiana var. kiusiana by several
characters such as light yellow hairs on the discof the labellum, earlier ﬂowering time, andnearly truncate apical part of the column. However, these conﬂicting taxonomic views have notbeen tested so far. In this study we re-evaluate
the taxonomic status of this entity by use of morphological, macromolecular, and ecological characters.
Materials and Methods
From the outset, we tentatively treated thethree taxa as independent species, viz., Lecanorchis suginoana, L. kiusiana, and L. japonica(Fig. 1). Materials used in the DNA analysis arelisted in Table 1. Besides, in the course of thisstudy, populations of L. suginoana were found in
Tochigi Prefecture, eastern part of Honshu, Japan

Fig. 1.

and Ehime Prefecture, Shikoku, Japan (see Taxonomic Treatment). They are the ﬁrst records of
distribution in these areas. Total DNA of plants
was extracted from fresh or silica-gel-dried samples with the DNeasy Plant Mini Kit (Qiagen,
Valencia, California, USA) following the manufacturerʼs instructions. Nucleotide sequenceswere determined by amplifying the internal transcribed spacer (ITS) regions of the 18S–26S
nuclear ribosomal DNA via the polymerase chain
reaction (PCR) from a total DNA extract. Experimental methods follow those described in Topiket al. (2005) and Yukawa et al. (2005) in which
the primer information of the sequenced region is
also shown. Voucher specimens were depositedat the Department of Botany, National Museumof Nature and Science (TNS).
The ITS sequences were aligned using MUSCLE implemented in MEGA 7.0.14 (Kumar et

Flowering individuals of Lecanorchis suginoana (A), L. kiusiana (B) and L. japonica (C).

Table 1.
Species
Lecanorchis ﬂavicans
Lecanorchis japonica

Lecanorchis kiusiana

Lecanorchis suginoana

Materials used for DNA sequencing. All voucher specimens are deposited at TNS
Locality
Japan, Kagoshima Pref., Amami-oshima Is.
Japan, Kagoshima Pref., Amami-oshima Is
Japan, Shikoku, Ehime Pref., Uwajima-shi
Japan, Honshu, Gifu Pref., Mizunami-shi
Japan, Honshu, Tochigi Pref., Mashiko-machi
Japan, Honshu, Miyagi Pref., Shiroishi-shi
Japan, Honshu, Fukui Pref., Echizen-choJapan, Shikoku, Ehime Pref., Uwajima-shi
Japan, Honshu, Shizuoka Pref., Kakegawa-shi
Japan, Honshu, Tochigi Pref., Mashiko-machi
Korea, Jeju Is.
Japan, Honshu, Shizuoka Pref., Kakegawa-shi
Japan, Honshu, Ibaraki Pref., Mito-shi
Japan, Honshu, Tochigi Pref., Yaita-shi

Voucher
T. Yukawa 09-21
H. Yamashita s. n.
S. Hyodo 17864
H. Nakagawa s. n.
M. Takashima s. n.
M. Takashima s. n.
T. Wakasugi & H. Enomoto 51886
S. Hyodo 17862
M. Takashima s. n.
M. Takashima s. n.
S. Matsumoto 03-171
T. Yukawa 14-10
H. Uchiyama s. n.
M. Takashima s. n.

Taxonomic status of Lecanorchis suginoanaal., 2016). All insertions/deletions were coded as
missing data in the phylogenetic analysis of ITS.
Lecanorchis ﬂavicans Fukuy. was chosen as an
outgroup taxon on the basis of the results of phylogenetic analysis of the whole genus (unpublished). The evolutionary history was inferred byusing a model based Bayesian approach and theMaximum Likelihood (ML) method. A Bayesian
search was performed using MrBayes 3.2 (Ronquist et al., 2012) and using the program Kakusan 4 (Tanabe, 2011) to select the ʻbest-ﬁtʼ modelof evolution under the Akaike information criterion. Two separate runs were carried out using
the GTR＋I＋Γ model in each analysis. The
Markov chains were run for 1 million generations and the ﬁrst 10 thousand runs were burned
in. ML analysis was conducted using MEGA andthe Kimura 2-parameter model (Kimura, 1980)
was chosen as the best ﬁtting model for our dataset. Bootstrap analysis (Felsenstein, 1985) with
1000 replicates was used to assess the relativerobustness of support for all branches.
Since the results of molecular phylogenetic
analysis demonstrated that Lecanorchis suginoana occupied a sister position of L. kiusiana andwas distantly related to L. japonica (see Resultsand Discussion), morphological analyses were
conﬁned to the former two taxa. Observation ofmorphological characters was based on living
plants, dried herbarium specimens, and spiritpreserved specimens listed in the section “Specimens examined”. Detailed measurements ofquantitative morphological characters were carried out from the ﬂowers collected in Yaita andKakegawa populations for L. suginoana, andMashiko and Kakegawa populations for L. kiusiana. Yaita and Mashiko are situated in the eastern part of Honshu, Japan and Kakegawa is situated in the central part of Honshu, Japan. A
single ﬂower was taken from each individual and63 samples were collected in total. Ten characters, viz., length of dorsal sepal, length of lateral
sepal, length of petal, width of petal, length oflabellum, width of labellum, length of column,
and width of column, were measured for eachsample. Studentʼs t-test was performed to evalu-

121

ate differences of these characters between the
two taxa. Moreover, we conducted canonical discriminant analysis and cluster analysis (Wardʼs
method) of the two taxa by using the same data
set of morphological characters. All statisticalanalyses were performed using Bell Curve for
Excel ver. 3.00 (Social Survey Research Information Co., Ltd., Tokyo, Japan).
We further carried out detailed phenologicalsurveys of Lecanorchis suginoana and L. kiusiana in the four populations used for morphological analyses (Yaita and Kakegawa populations
for L. suginoana; Mashiko and Kakegawa populations for L. kiusiana). Five characters, viz.,
number of individuals, number of buds, numberof ﬂowers in full bloom, and number of wilted
ﬂowers, were recorded for the plants in eachquadrat. In Yaita (L. suginoana habitat) and
Mashiko (L. kiusiana habitat), ca. 47 km aparteach other, 11 consecutive surveys were conducted in May and June 2012. In Kakegawa, thetwo taxa formed populations less than 20 m apart.
Surveys were conducted in 17 and 30 May 2015.
Furthermore, we recorded vegetation in association with L. suginoana and L. kiusiana at theabove-mentioned four habitats. Methods forrecording vegetation composition followed
Braun-Blanquet (1964).
Results and Discussion
Macromolecular characters
In ca. 760 base pairs of ITS, three samples ofLecanorchis suginoana from different localities
had identical sequences. Four samples of L. kiusiana had identical ITS sequences and two nucleotide substitutions discriminated these two taxa.
In Kakegawa, populations of L. suginoana and L.
kiusiana are less than 20 m apart. It is likely that
reproductive isolation between these two taxa
occurs because no trace of hybridization was
detected in the nuclear nucleotide sequences
investigated for these nearly sympatric populations. Two ITS phylotypes demarcated by a single substitution were recognized in seven samples of L. japonica. Both L. suginoana and L.

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Michihisa Takashima et al.

Fig. 2. Phylogenetic relationships of Lecanorchis suginoana, L. kiusiana and L. japonica inferred from nuclear
ribosomal ITS sequences. The tree is rooted by an outgroup taxon L. ﬂavicans. The tree was constructed using
the maximum likelihood method and the topology was the same as the Bayesian inference analysis. Numbers
at the nodes indicate Bayesian posterior probabilities (PP) and bootstrap percentages (BS) shown as PP/BS.

kiusiana were separated from L. japonica by ﬁve
or six substitutions. Phylogenetic analysis of ITS
showed that L. suginoana and L. kiusiana make asister group relationship and these two taxa andL. japonica further form a clade (Fig. 2). These
results coincide with Okayama et al. (2012) anddemonstrate an independent taxonomic status of
L. suginoana. Further, the phylogenetic relationships did not support an infraspeciﬁc placementof L. suginoana in L. japonica as proposed byTuyama (1982).
Morphological characters
Our observations revealed that Lecanorchis
suginoana is distinguishable from L. kiusiana by
the following combination of qualitative morphological characters: yellowish-brown sepals
and petals, yellow labellum hair; scattered, long,
unicellular branchlets on labellum hair; weakly
recurved labellum apex; and truncate column
apex (Fig. 3, Table 2). These diagnostic characters were consistent among all the examined individuals and previous descriptions of the two taxa
(e.g., Tuyama, 1982; Sugino, 1985; Hashimoto,1990; Serizawa, 2005) except for L. suginoana f.

ﬂava Seriz., an anthocyanin defective ﬂower
form, having bright yellow sepals and petals.
Besides, Lecanorchis suginoana and L. kiusiana shared papillose labellum margins and unicellular branchlets on labellum hair, while thesecharacters were not evident in L. japonica and its
close relatives. Consequently, they are synapomorphies of the former two taxa and endorse
their sister relationship revealed by the molecularphylogenetic analysis.
Regarding quantitative morphological characters, Studentʼs t-test showed that length of dorsal
sepal, length of lateral sepal, length of petal,
width of petal, length of labellum, width of labellum, length of column, and width of labellum
were signiﬁcantly different between Lecanorchis
suginoana and L. kiusiana (p＜0.01; Table 2).
These differences indicated that L. suginoana has
longer perianth lobes, wider petal, narrowerlabellum, longer column, and apically wider column than L. kiusiana. Results of the canonicaldiscriminant analysis and the cluster analysis byusing quantitative morphological characters also
clearly separated these two taxa (Figs. 4, 5).

Taxonomic status of Lecanorchis suginoana

123

Fig. 3. Morphological comparison of Lecanorchis suginoana and L. kiusiana. Lateral view of labellum and column, L. suginoana (A) and L. kiusiana (B). Dorsal view of column, L. suginoana (C) and L. kiusiana (D).
Marginal part of labellum, L. suginoana (E) and L. kiusiana (F). Labellum hair of L. suginoana (G) and L. kiusiana (H).
Table 2.

Comparison of morphological characters in Lecanorchis suginoana and L. kiusiana
Character

Colour of petals and sepals
Shape of column apex
Colour of labellum hair
Length of branchlets on labellum hair (μm)
Density of branchlets on labellum hair
Lateral view of labellum
Length of dorsal sepal: mean±SD (mm)Width of dorsal sepal: mean±SD (mm)Length of lateral sepal: mean±SD (mm)Width of lateral sepal: mean±SD (mm)Length of petal: mean±SD (mm)Width of petal: mean±SD (mm)Length of labellum: mean±SD (mm)Width of labellum: mean±SD (mm)Length of column: mean±SD (mm)Width of column: mean±SD (mm)L. suginoana
yellowish brown
truncateyellow
50–70
lax in mid to apical part
weakly recurved
15.8±0.64* (N＝33)
3.14±0.25 (N＝33)15.0±0.57* (N＝34)2.88±0.16 (N＝34)15.4±0.55* (N＝34)
3.76±0.24* (N＝34)13.9±0.48* (N＝34)
6.49±0.37* (N＝34)9.27±0.22* (N＝34)2.17±0.10* (N＝34)

L. kiusiana
pale dull yellow
3 lobedreddish purple
10–30
dense in apical partstrongly recurved
14.3±1.12* (N＝29)3.32±0.50 (N＝29)13.9±0.90* (N＝29)2.81±0.29 (N＝29)13.7±0.93* (N＝29)3.54±0.34* (N＝28)13.4±0.53* (N＝29)8.37±0.66* (N＝29)8.46±0.42* (N＝29)
1.96±0.20* (N＝28)* denotes signiﬁcance at the 1% level.

Ecological characters
Figure 6 shows that ﬂowering of Lecanorchis
suginoana at Yaita population started on 1 Juneand reached the peak on 8 June. The last ﬂowersof the season were observed on 18 June. Flowering of L. kiusiana at Mashiko population (47 km
southward from Yaita) started on 8 June andreached the peak on 18 June. The last ﬂowers of
the season were observed on 24 June. This observation showed that the ﬂowering period of L.

suginoana was about one week earlier than thatof L. kiusiana in the eastern part of Honshu,
Japan, although about 10 days overlap period
existed. In contiguous populations of these two
taxa in Kakegawa, central part of Honshu, Japan,
ﬂowering of L. suginoana passed its peak on 22
May, while L. kiusiana just started ﬂowering at
the same date. On 30 May, ﬂowering of L. suginoana already ended, while L. kiusiana justpassed its peak (Table 3). These observations

124

Michihisa Takashima et al.

Fig. 4. Canonical discriminant analysis of 10 quantitative morphological characters of Lecanorchis suginoana
and L. kiusiana ﬂowers with the following four predeﬁned groups: □ L. suginoana, Yaita population; ■ L.
suginoana, Kakegawa population; △ L. kiusiana, Mashiko population; ▲ L. kiusiana, Kakegawa population.
Percentages of total variance explained by the functions are given in parentheses. The analysis showed that thetwo taxa can be discriminated in 100% hit ratio.

demonstrated that earlier ﬂowering of L. suginoana was likely to be a stable character.
Habitat differentiation between Lecanorchis
suginoana and L. kiusiana is also noticeable(Fig. 7). Table 4 showed that two L. suginoana
populations are associated with warm-temperate
deciduous forest dominated by Quercus serrata
Murray in the tree layer. In contrast, two L. kiusiana populations are in association with warmtemperate evergreen forest dominated by Castanopsis sieboldii (Makino）Hatus. ex T.Yamaz.
et Mashiba subsp. sieboldii or C. cuspidata(Thunb.) Schottky in the tree layer. A similar
habitat differentiation between these two
Lecanorchis taxa from the standpoint of associated vegetation types was recorded by Sugino
and Suzuki (1987).
Okayama et al. (2012) investigated mycorrhizal fungal diversity in 10 Lecanorchis taxa andfound that they are symbiotic with several ectomycorrhizal fungi in Basidiomycota such as Lactarius (Russulaceae), Russula (Russulaceae),
Atheliaceae, and Sebacina (Sebacinaceae). Since
these fungal groups also associate with Fagaceae
trees such as Quercus and Castanopsis and form

ectomycorrhizae, tripartite symbioses likely
establish among these organisms. Obviously,
existence of appropriate fungal partners at habitats is indispensable for sustaining the life of
Lecanorchis that completely depends upon nutrition to its symbiotic fungi. Okayama et al. (2012)
found that different sets of fungi dominantly colonized with L. suginoana and L. kiusiana. They
showed that the former species dominantly associated with Lactarius sp.1 and the latter mainly
associated with Lactarius sp. 2. These ﬁndings
indicate that habitat differentiation between the
two Lecanorchis taxa is caused by different
mycobiont communities between Quercus andCastanopsis forests, although sampling density
in that study was not sufﬁcient to substantiate
this hypothesis.
Conclusions
Divergence of ITS sequences supported the
independent status of Lecanorchis suginoana, L.
kiusiana, and L. japonica. Moreover, the resultsof molecular phylogenetic analysis demonstrated
that L. suginoana occupied a sister position of L.
kiusiana and was distantly related to L. japonica.

Taxonomic status of Lecanorchis suginoana

125

Fig. 5. Tree diagram generated by the cluster analysis of 10 quantitative morphological characters of Lecanorchis
suginoana and L. kiusiana ﬂowers using Wardʼs method. Height indicates the Euclidean distance. The results
showed that the basal branching clusters correspond to the two taxa.

Thus, the infraspeciﬁc placement of L. suginoana
in L. japonica was not supported. Moreover,ﬂower colour, qualitative characters of labellum
and column, quantitative characters of perianth
lobes and column, ﬂowering time, and associatedvegetation types clearly distinguish L. suginoana
from L. kiusiana. These differences corroborated
the independent species status of L. suginoana
rather than the infraspeciﬁc position of L. kiusiana.

Taxonomic Treatment
Lecanorchis suginoana (Tuyama) Seriz., Bunrui
5: 38 (2005)[Figs. 1, 3]
Basionym: Lecanorchis japonica Blume var.
suginoana Tuyama, J. Jpn. Bot. 57: 211 (1982).
TYPE: JAPAN, Honshu, Shizuoka Pref., Haruno-cho, Sugi, 8 June 1980, T. Tuyama s. n. (TI).
Lecanorchis kiusiana Tuyama var. suginoana(Tuyama) T.Hashim., Ann. Tsukuba Bot. Gard. 9:18 (1990).
Japanese name: Enshu-muyouran.

Michihisa Takashima et al.

126
120

Number of flowers

100

80

60

40

20

0

25 May 1 June 6 June

8 June 10 June 13 June 15 June 18 June 21 June 24 June 27 June
Date

Fig. 6. Flowering period of Lecanorchis suginoana (Yaita population) and L. kiusiana (Mashiko population).
These two populations are ca. 47 km apart and represent similar meteorological conditions. Black bars andgray ones indicate the number of ﬂowers in full bloom of L. suginoana and L. kiusiana respectively.
Table 3. Flowering stage of Lecanorchis suginoana and L. kiusiana at two adjacent habitats in Kakegawa, central
part of Honshu, Japan
Date

Taxon

22 May 2015

30 May 2015

L. suginoana

Number of buds
Number of ﬂowers in full bloom
Number of wilted ﬂowers
Total number of ﬂowers
Number of individuals

0
59
90
149
35

0
0
149
149
35

L. kiusiana

Number of buds
Number of ﬂowers in full bloom
Number of wilted ﬂowers
Total number of ﬂowers
Number of individuals

132
19
0
151
35

8
52
80
140
38

Flowering period: May to June.
Distribution: JAPAN: Honshu (Niigata Pref.,Tochigi Pref., Ibaraki Pref., Saitama Pref., Tokyo
Metro., Shizuoka Pref., Aichi Pref., Gifu Pref.,Shiga Pref., Nara Pref., Kyoto Pref., Osaka
Pref.); Shikoku (Ehime Pref., Kochi Pref.);
Kyushu (Miyazaki Pref.); TAIWAN. Records of
distribution were reviewed in Suetsugu et al.
(2018).
Specimens examined: JAPAN. Honshu: Tochigi Pref. Nasushiobara-shi, Shimo-ohnuki, alt.
270 m, 11 June 2018, T. Noguchi 180611-01(TNS1311685); Yaita-shi, Yamada, alt. 255 m, 10
June 2010, T. Ito s. n. (TOCH167982); Yaita-shi,
Yamada, 5 June 2012, M. Takashima s. n.
(TNS8505115); Yaita-shi, Arai, alt. 260 m, 10
June 2010, T. Ito s. n. (TOCH167983); Yaita-shi,
Higashi-izumi, alt. 260 m, 8 June 2012, N. Hoshi& M. Takashima s. n. (TOCH192869). Ibaraki
Pref. Takahagi-shi, Daitakahagi, 6 June 1990, M.
Suzuki 21838 (INM); Hitachi-ohmiya-shi, Mt.
Torinoko-san, alt. 430 m, 11 June 2005, H.
Ogura, Y. Maruyama & A. Narishima s. n.
(TOCH160932); Hitachi-ohmiya-shi, Torinoko,

Taxonomic status of Lecanorchis suginoana

127

Fig. 7. Habitats of Lecanorchis suginoana and L. kiusiana. A: L. suginoana, Yaita population. B: L. suginoana,
Kakegawa population. C: L. kiusiana, Mashiko population. D: L. kiusiana, Kakegawa population.

alt. 450 m, 16 June 2013, H. Uchiyama & H.
Fujita s. n. (TNS01184364); Mito-shi, Kawadacho, alt. 30 m, 31 May 2013, H. Uchiyama s. n.
(TNS8505120, 01184365). Shizuoka Pref. Shuchi-gun, Haruno-cho, Sugi, 28 June 1981, T.
Tuyama s. n. (TI1326096, TI1328095); Shuchigun, Haruno-cho, Sugi, 15 June 1986
(TNS8504021); Shuchi-gun, Haruno-cho, 15
June 1986, K. Kanda & Y. Hanei s. n.
(TNS8504481); Shuchi-gun, Haruno-cho, Sugi,
15 June 1986, Y. Hanei s. n. (TNS9504068,
9504069); Shimada-shi, Shitoro, alt. 170 m, 22
May 2015, M. Takashima & M. Ohta s. n.
(TNS1311686); Kakegawa-shi, Higashiyama, alt.
210 m, 22 May 2015, M. Takashima & M. Ohta
s. n. (TNS1311687, 1311688). Gifu Pref. Tajimishi, Higashimachi, 24 May 2013, C. Tsubaki, K.
Nomura & R. Nomura s. n. (GPM-B918390);
Gifu-shi, Tsubakibora, 30 May 2005, R. Naruses. n. (GPM-B756065); Gifu-shi, Hora, 21 May

2004, H. Minoura s. n. (GPM-B756066); Mizunami-shi, Inazu-machi, Kozato, alt. 250 m, 12
June 2006, M. Murase & S. Nagata s. n. (GPMB756993). Aichi Pref. Kota-cho, 26 March 1992,
K. Inami s. n. (TNS8504035); Okazaki-shi,
Kamisobumi-cho, alt. 120-150 m, 25 May 2005,
S. Fujii s. n. (TNS772431); Nagoya-shi, Chikusaku, Higashiyama, 5 April 1984, Y. Sawa 1411
(TNS01013786). Kyushu: Miyazaki Pref. Koyugun, Kawaminami-cho, Ohuchi, alt. 270 m, 1
June 2004, T. Miamitani s. n. (TNS9519671).
f. ﬂava Seriz., Bunrui 5: 38 (2005).
TYPE: JAPAN, Honshu, Aichi Pref., Seto-shi,
Kaisho-cho, alt. ca. 200 m, 29 May 2000, S. Serizawa 76752 (AICH).
Specimens examined: JAPAN, Honshu:
Shizuoka Pref. Kakegawa-shi, Higashiyama,
alt. 255 m, 18 June 2014, T. Yukawa 14-10(TNS8505619). Gifu Pref. Mizunami-shi, Hiyo-Michihisa Takashima et al.

128

Table 4. Vegetation associated with Lecanorchis suginoana and L. kiusiana populations
L. suginoana

Taxon
Locality

Yaita

Kakegawa 1

L. kiusiana
Mashiko

Kakegawa 2

—
SE
SW
S
0°
20°
25°
10°
14 m
15 m
20 m
17 m
85%
90%
95%
90%
30%
35%
15%
15%
0%
25%
10%
0%
30%
15%
3%
1%
200 m
210 m
200 m
210 m
15 m×15 m 15 m×10 m 20 m×20 m 15 m×15 m

Slope aspect
Slope degree
Canopy height
Canopy coverage (%)
Subtree layer coverage (%)
Shrub layer coverage (%)
Herb layer coverage (%)
Altitude
Quadrat size
Taxon

Dominance・Sociability

Dominance・Sociability

Castanopsis sieboldii (Makino) Hatus. ex T.Yamaz. et Mashiba subsp. sieboldii
Castanopsis cuspidata (Thunb.) Schottky
Quercus serrata Murray
Cryptomeria japonica (L. f.) D.Don
Camopy layer Clethra barbinervis Siebold et Zucc.
Cerasus jamasakura (Siebod ex Koidz.) H. Ohba
8 m–
Carpinus laxiﬂora (Siebold et Zucc.) Blume
Quercus sessilifolia Blume
Quercus salicifolia Murray
Ilex rotunda Thunb.
Trachelospermum asiaticum (Siebold et Zucc.) Nakai var. asiaticum

・
・
4・4
・
2・1
・
・
・
・
・
＋

・
・
4・4
・
3・3
2・2
2・1
・
・
・
・

5・5
・
・
・
・
・
・
1・1
1・1
1・1
・

・
4・4
2・2
2・1
・
・
・
・
・
・
・

Eurya japonica Thunb. var. japonica
Clethra barbinervis Siebold et Zucc.
Machilus japonica Thunb.
Lyonia ovalifolia (Wall.) Drude var. elliptica (Siebold et Zucc.) Hand.-Mazz.
Cryptomeria japonica (L.f.) D.Don
Alnus ﬁrma Siebold et Zucc.
Castanopsis sieboldii (Makino) Hatus. ex T.Yamaz. et Mashiba subsp. sieboldii
Quercus glauca Thunb. var. glauca
Camellia japonica L. var. japonica
Cleyera japonica Thunb.
Ilex integra Thunb.
Ilex chinensis Sims

・
2・1
・
2・1
2・1
・
・
・
・
・
・
・

3・3
2・2
・
・
・
1・1
・
・
・
・
・
・

1・1
・
・
・
・
・
1・1
1・1
1・1
1・1
2・2
・

・
・
2・2
・
・
・
・
1・1

Shrub layer
1–4 m

Eurya japonica Thunb. var. japonica
Cleyera japonica Thunb.
Hydrangea hirta (Thunb.) Siebold et Zucc.
Pieris japonica (Thunb.) D. Don ex G. Don
Camellia japonica L. var. japonica
Pleioblastus chino (Franch. et Sav.) Makino
Lindera umbellata Thunb.
Quercus glauca Thunb. var. glauca
Quercus salicina Blume
Quercus salicifolia Murray
Padus grayana (Maxim.) C. K. Schneid.
Wisteria ﬂoribunda (Wild.) DC.
Ilex integra Thunb.
Callicarpa mollis Siebold et Zucc.

・
・
・
・
・
・
・
・
・
・
・
＋・2
・
・

2・2
・
1・2
2・2
・
1・2
1・2
・
・
・
1・1
・
・
・

1・2
2・2
・
・
1・2
・
・
1・1
＋
1・1
・
・
＋
＋

・
・
・
・
・
・
・
・
・
・
・
・
・
・

Herb layer
0–1 m

Rhododendron kaempferi Planch. var. kaempferi
Lecanorchis kiusiana Tuyama
Lecanorchis suginoana (Tuyama) Seriz.
Dryopteris erythrosora (D. C. Eaton) Kuntze
Disporum smilacinum A. Gray
Hydrangea hirta (Thunb.) Siebold et Zucc.
Sasa ramosa (Makino) Makino
Helonias orientalis (Thunb.) N. Tanaka
Aucuba japonica Thunb. var. japonica
Trachelospermum asiaticum (Siebold et Zucc.) Nakai var. asiaticum
Pseudosasa japonica (Siebold et Zucc. ex Steud.) Makino ex Nakai
Pleioblastus chino (Franch. et Sav.) Makino
Toxicodendron tricocarpum (Miq.) Kuntze.
Ilex crenata Thunb.
Pertya glabrescens Sch. Bip. ex Nakai
Cymbidium goeringii (Rchb. f.) Rchb.f. var. goeringii
Platanthera minor (Miq.) Rchb.f.

3・3
・
1・2
・
・
・
1・2
・
＋・2
・
・
・
＋
・
・
・
・

2・2
・
1・2
1・2
1・2
1・2
・
1・2
・
・
・
＋・2
＋
＋
＋
＋
＋

・
1・2
・
＋
＋
・
・
・
＋
＋・2
＋・2
・
＋
＋
＋
＋
・

・
1・2
・
・
・
・
・
・
・
・
・
・
・
＋
・
・
＋

Subtree layer
4–8 m

1・1
1・1
1・1

Other companions. Running No. 1: Cryptomeria japonica (L.f) D.Don＋, Chamaecyparis obtusa (Siebold et Zucc.) Endl.＋, Quercus serrata
Murray＋, Marus toringo (Siebold) Siebold ex de Vriese＋, Tripterospermum japonicum (Siebold et Zucc.) Maxim.＋, Hydrangea paniculata
Siebold＋, Smilax nipponica Miq.＋, No. 2: Osmunda japonica Thunb.＋, Ardisia crenata Sims＋, No. 3: Illicium anisatum L. var. anisatum＋,
Quercus glauca Murray＋, Quercus sessilifolia Blume＋, Quercus myrsinifolia Blume＋, Castanopsis sieboldii (Makino）Hatus. ex T.Yamaz. et
Mashiba subsp. sieboldii＋, Laurocerasus spinulosa (Siebold et Zucc.) C.K.Schneid.＋, Camellia japonica L. var. japonica＋, Eurya japonica
Thunb. var. japonica＋, Cleyera japonica Thunb.＋, Ardisia japonica (Thunb.) Blume var. japonica＋, Ilex rotunda Thunb.＋, Viburnum erosum
Thunb.＋, Osmanthus heterophyllus (G.Don.) P.S.Green＋, Ophiopogon japonicus (Thunb.) Ker Gawl. var. umbrosus Maxim.＋, No. 4: Ilexpedunculosa Miq.＋, Callicarpa japonica Thunb.＋, Callicarpa mollis Siebold et Zucc.＋.

Taxonomic status of Lecanorchis suginoana

shi-cho, Kitano, 5 June 2013, H. Nakagawa s. n.
(TNS8505118). Shikoku: Ehime Pref. Saijo-shi,
Tanbara-cho, Myoga, Yokoumi, 31 May 2015, S.
Hyodo 17881 (TNS8505758).
Acknowledgments
We would like to thank Tatsunari Noguchi,
Takayuki Shibata, Hidehiko Shimizu, Toshio
Tashiro and Yuki Ueno for sharing their observations, Shoji Hyodo, Hisao Nakagawa, Haruo
Uchiyama, Takao Wakasugi, and Hiroshi
Yamashita for providing samples, Naoto Hoshi
for giving useful comments on the vegetation
survey, and Stephan Gale for critical reading of
the manuscript. We are also grateful to Hiroshi
Ikeda (TI), Akiko Shimizu (TI), Atsushi Ebihara(TNS), Nobuyuki Tanaka (TNS), Miki Kani
(Gifu Prefectural Museum), Chika Suyama (Gifu
University), Kazuo Obata (INM) and Mihoko
Uzawa (INM) for allowing us to examine specimens. This study is partly supported by JSPS
KAKENHI (Grant Numbers JP18H02500 and
JP18K06400) and by a National Museum of
Nature and Science research grant entitled “Integrated analysis of natural history collections forconservation of highly endangered species” to T.
Yukawa and Y. Yamashita.
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