Bull. Natn. Sci. Mus., Tokyo, Ser. A, 30(4), pp. 191–218, December 22, 2004

Ophiuroids (Echinodermata) Collected from the Oki Islandsin the Sea of Japan
Toshihiko Fujita1, Yoshiaki Ishida2, Takumu Kato3 and Seiichi Irimura4
1)

Department of Zoology, National Science Museum, Hyakunin-cho 3–23–1,
Shinjuku-ku, Tokyo 169–0073, Japane-mail: fujita@kahaku.go.jp
2)
Chitosegaoka Senior High School, Funabashi 3–18–1,
Setagaya-ku, Tokyo 156–0055, Japane-mail: y-ishida@msi.biglobe.ne.jp
3)
Osuwa-sou 201, Miyamae 369–5, Fujisawa-shi, Kanagawa 251–0014, Japane-mail: kato-takumu@mx3.ttcn.ne.jp
4)
Okusawa 1–49–1, Setagaya-ku, Tokyo 158–0083, Japane-mail: irimura@f2.dion.ne.jp

Abstract Ophiuroids were collected by dredging and skin diving from the shallow water of the
Oki Islands in the Sea of Japan. They comprise 29 species of 18 genera of 9 families. Ophiurodon
acanthophora is transferred to Ophioconis. Amphiura kandai, Ophionereis thryptica, and Ophioconis acanthophora are reported for the first time from the Sea of Japan.
Key words : Ophiuroidea, echinoderm, taxonomy, new combination, new record, Shimane Prefecture, Japan.

Introduction

Materials and Methods

The Oki Islands are located in the southernpart of the Sea of Japan, about 70 km off Honshu,
the main land of Japan, and composed of four big
islands and about 180 small islands. They arewarmed by the Tsushima Current, and the marine
fauna includes the animals which are typicallymore southerly in distribution; for example, reefbuilding corals were found near Matsushima Island (Miyaoka, 1997; see Figure 1 for its locality) and also off Dougo Island (T. Kato, unpublished data).
Ishida et al. (2001) compiled the ophiuroid
fauna of the Sea of Japan. However, the ophiuroid fauna of the Oki Islands has been neverstudied. One of the authors (TK) collected marine animals of the Oki Islands mainly from 1987
to 1995, some of which have been reported on(Kato, 1993, 1998). In this paper, we examine
ophiuroid specimens of Kato’s collection fromthe shallow waters of the Oki Islands.

Ophiuroid specimens of Kato’s collection arefrom the intertidal zone to up to the depth ofabout 130 m around south of Dougo Island and in
the strait between Dougo Island and Nakanoshima Island (Fig. 1). They were collected using
a dredge during cruises of the research vessels
Pagurus and Galathea of the Oki Marine Biological Station of Shimane University mainly from1987 to 1992, and by skin diving from intertidal
and shallow subtidal waters. Additional specimens were obtained with fishing gear. Material
studied also includes specimens dredged by
Takeo Yamauchi in 2002. Ophiuroids were fixed
in 5% seawater formalin or alcohol and preserved in 80% alcohol, although a few specimens
were dried for observation. Photographs weretaken of preserved specimens. The specimens aredeposited in the National Science Museum,
Tokyo (NSMT).
In the Taxonomic Account section, families
are arranged according to Smith et al. (1995).
The Japanese name, important synonymy, materi-

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Fig. 1.

Sampling localities at the Oki Islands in the Sea of Japan.

al examined, remarks, and geographic distribution are given for each species. Matsumoto(1917) and Irimura (1982) discuss the taxonomical determination of many species found in thisstudy. In Material examined, for each locality,
water depth, bottom nature, and specimen registration number are shown if available. Abbreviations for bottom sediment characteristics appearing in parentheses are as follows: G, gravel; M,
mud; R, rock; S, sand; Sh, shell.
Taxonomic AccountA total of 860 specimens determined in thisstudy comprise 29 species of 18 genera of 9 families.
Order Euryalida
Family Gorgonocephalidae
Astrocladus coniferus (Döderlein, 1902)[Japanese name: Seno-tedurumoduru]
Astrophyton coniferum Döderlein, 1902: 325–326.

Astrocladus coniferus: Matsumoto, 1917: 77–79, fig. 23.
Astrocladus dofleini Döderlein, 1910: 256; 1911: 41–46,
pl. 2 fig. 6, pl. 4 fig. 4–5, pl. 7 fig. 15–15b; 1927:
35–36, pl. 3 fig. 2–2a.

Material examined. Off Takugi; 120–130 m(R, G), 2 specimens (NSMT E-4793).
Remarks. The specimens have many tubercles on disk and arms. Astrocladus dofleini
Döderlein, 1910 was distinguished from A.
coniferus by having tubercles (Döderlein, 1911),
but later was synonymized with A. coniferus because many intermediate forms were found inJapanese waters (Matsumoto, 1917; Fujita &Kohtsuka, 2003). This is only species of Order
Euryalida in the present study, but several othereuryalid species have been reported from the Sea
of Japan (Ishida et al., 2001) and are probably
distributed in the study area.
Geographic range. This species is distributed
from the Sea of Japan to the Philippines, and alsoin India. Widely distributed in the Sea of Japan(see Ishida et al., 2001; present study). Korea(Rho & Shin, 1987; Yi & Irimura, 1987). PacificOphiuroids from the Oki Islands, Japan

coast of Japan (see Irimura, 1982) from Boso
Peninsula (Irimura, 2000) to Tanegashima Strait(H. L. Clark, 1911: 293, Astrophyton cornutum),
the Ogasawara Islands (Murakami, 1944), andthe Yaeyama Islands (Murakami, 1944). Philippines (Döderlein, 1927; Guille, 1981). India(Bomford, 1913). The type locality is Kagoshima
Bay, Japan (Döderlein, 1902).
Order Ophiurida
Family Ophiacanthidae
Ophiacantha aff. prionota H. L. Clark, 1911
(Fig. 2)
Ophiacantha prionota H. L. Clark, 1911: 213–215, fig.
99.

Material examined. Off Onbejima Island;
71 m, 1 specimen (NSMT E-4794). Locality nodata; 1 specimen (NSMT E-4795).
Remarks. One specimen (NSMT E-4795)
has 5 arms, and another one (NSMT E-4794) has
6 arms. These specimens resemble Ophiacantha
prionota, which has been so far reported from
Tanegashima Island, Japan (type locality; H. L.
Clark, 1911); however, the shape of dorsal andventral arm plates differs from those of O. prionota. The dorsal arm plates are triangular, wider
than long, and widely separated by lateral arm
plates (Fig. 2C, K). The ventral arm plates arehexagonal with rounded distal margin, and separated by lateral arm plates (Fig. 2E, M). The dorsal surface of disk has spines and granules onthin scales (Fig. 2B, J). There are three oral
papillae on each side of the jaw (Fig. 2D, L). The
tentacle pores sometimes have two scales at armbase (Fig. 2G), but generally there in only one,large, leaf-like thin scale (Fig. 2H, O). The arm
spines (Fig. 2F, N) are serrated and the uppermost one is usually the longest. Maximum number of arm spines is six.
Family Ophiuridae
Ophiura kinbergi (Ljungman, 1867)[Japanese name: Kushinoha-kumohitode]

Material examined.

Off Hobomi; 65 m, 1

193

specimen (NSMT E-4811). Between Oomorijima
Island and Shijikijima Island; 58 m, 1 specimen(NSMT E-4797); 58 m, 12 specimens (NSMT E4798). Off Onbejima Island; 50 m, 6 specimens(NSMT E-4813); 52 m, 2 specimens (NSMT E4799); 67 m (S, Sh), 9 specimens (NSMT E4804). East of Onbejima Island; 54 m, 48 specimens (NSMT E-4803); 60 m (Sh), 24 specimens(NSMT E-4806). Off Tsuma; 50 m, 5 specimens(NSMT E-4802). Off Uzunohana; 50 m (R, G,
S), 9 specimens (NSMT E-4796). South-southeast of Tsudo; 62–65 m (G), 6 specimens (NSMTE-4807). Off Shijikijima Island; 36°9N,
133°14E, 20–30 m (G, R), 2 specimens (NSMT
E-4999); 60 m (R), 5 specimens (NSMT E4801). East of Shijikijima Island; 15 m (G), 1
specimen (NSMT E-4809). Between MatsushimaIsland and Kanjima Island; 16–18 m (Sh), 2 specimens (NSMT E-4810). Off Kanbi; 38 m, 5 specimens (NSMT E-4805). Kamo Bay; 17 m, 5specimens (NSMT E-4819); 37 m, 4 specimens(NSMT E-4800); 38 m, 6 specimens (NSMT E4818); 53 m, 10 specimens (NSMT E-4815). At
the mouth of Kamo Bay; 1 specimen (NSMT E4812). Off Kamo Bay; 43 m, 1 specimen (NSMT
E-4816). Saigou Bay; (M), 2 specimens (NSMTE-4808). South of Dougo Island (no detailed locality data); 1 specimen (NSMT E-4817); 9 specimens (NSMT E-4814).
Geographical range. This species is distributed in the Sea of Japan and widely in Indo-West
Pacific (see A. M. Clark & Rowe, 1971). It is acommon species in the Sea of Japan (see Irimura,
1982; Ishida et al., 2001; Fujita & Kohtsuka,
2003; present study). The type locality is Sidney,
Australia (Ljungman, 1867).
Stegophiura sladeni (Duncan, 1879)[Japanese name: Aka-hako-kumohitode]

Material examined. At mouth of Nukadani
Bay; 57 m (S), 6 specimens (NSMT E-4820);
57 m (S), 1 specimen (NSMT E-4821). South ofDougo Island (no detailed locality data); 2 specimens (NSMT E-4822); 5 specimens (NSMT E4823); 2 specimens (NSMT E-4824).

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Toshihiko Fujita et al.

A

C

B

C-H

D

F

E
H

G
I

K

J

K-O

L

M

N
O

Fig. 2. Ophiacantha aff. prionota. A–H: NSMT E-4795 (disk diameter 2.6 mm). I–O: NSMT E-4794 (disk diameter 2.1 mm). A: Disk and arms, aboral view. B: Disk spines and granules, aboral view. C: Arm base, aboral view. D: Jaw. E: Arm base, oral view. F: Arm spines near arm base. G–H: Tentacle scales (arrows). I: Disk
and arms, aboral view. J: Disk spines and granules, aboral view. K: Basal arm, aboral view. L: Mouth frame.
M: Basal arm, oral view. N: Arm spines near arm base. O: Tentacle scales (arrow). A–O: Alcohol specimen.
Scales 2 mm (A, I), 0.5 mm (B–H, J–O).

Ophiuroids from the Oki Islands, Japan

Remarks. Two species of Stegophiura were
found in this study. Stegophiura sladeni was easily distinguished from S. vivipara by its dimorphic arm spines (Matsumoto, 1917).
Geographic range. This species is distributedin Korea and Japan. Widely distributed in the Sea
of Japan (see Ishida et al., 2001; present study).
Korea, the Yellow Sea (Yi & Irimura, 1987).
Japan, Hakodate (Koehler, 1922), Kinkasan(Matsumoto, 1918), Boso Peninsula (Irimura,2000), Sagami Sea (Matsumoto, 1917; Irimura,1982), Suruga Bay (Fujita et al., 1997), Kii
Peninsula (Murakami, 1963; Irimura, 1981), Tosa
Bay (Irimura, 1990), Kagoshima Bay (H. L.
Clark, 1911 as Ophiura stiphra), East China Sea(Irimura & Kubodera, 1998). The type locality is
Korea (Duncan, 1879).
Stegophiura vivipara Matsumoto, 1915

195

Japan, Boso Peninsula (Irimura, 2000), Sagami
Sea (type locality, Matsumoto, 1915, 1917; Murakami, 1942; Irimura, 1982), Suruga Bay (Fujita
et al., 1997), the Ogasawara Islands (Irimura &Tachikawa, 2003).
Ophiochalcis aspera Koehler, 1931
[Japanese name: Samehada-mame-kumohitode]

Material examined. At mouth of Nukadani
Bay; 57 m (S), 2 specimens (NSMT E-4841).
Kamo Bay; 53 m, 1 specimen (NSMT E-4842).
Geographic range. This species is distributedin Japan and Indonesia. The Sea of Japan,
Tsushima Strait (Ishida et al., 2001), the Oki Islands (present study). Pacific coast of Japan,
Boso Peninsula (Irimura, 2000), Suruga Bay (Fujita et al., 1997). Indonesia, Ambon (type locality, Koehler, 1931).

[Japanese name: Komochi-kumohitode]

Material examined. Off Hobomi; 18 m (S), 1
specimen (NSMT E-4833). Off Onbejima Island;
50 m, 3 specimens (NSMT E-4834); 52 m, 1
specimen (NSMT E-4839); 52 m, 2 specimens(NSMT E-4825); 60 m (S, Sh), 1 specimen(NSMT E-4830); 67 m (S, Sh), 3 specimens(NSMT E-4831); 67 m, 3 specimens (NSMT E4838). East of Onbejima Island; 54 m, 15 specimens (NSMT E-4829). Off Tsuma; 50 m, 1 specimen (NSMT E-4827); 50 m, 4 specimens(NSMT E-4828). Off Tsudo; 62 m, 2 specimens(NSMT E-4840). Off Shijikijima Island; 60 m
(R), 1 specimen (NSMT E-4826). At mouth of
Nukadani Bay; 57 m (S), 1 specimen (NSMT E4832). Kamo Bay; 53 m, 9 specimens (NSMT E4836). South of Dougo Island (no detailed locality data); 36 specimens (NSMT E-4835); 4 specimens (NSMT E-4837).
Remarks. Matsumoto (1917) reported thebrooding habit of this species.
Geographic range. This species is distributedin Korea and Japan. Korea, the Yellow Sea (Yi &Irimura, 1987). The Sea of Japan, Sado (Kogure,1999), Tsushima Strait (Ishida et al., 2001), theOki Islands (present study). Pacific coast of

Family Amphiuridae
Amphioplus (Amphioplus) asterictus
H. L. Clark, 1915
[Japanese name: Chûrippu-kumohitode]
Amphioplus asterictus H. L. Clark, 1915a: 252, pl. 7 fig.
9–11; Murakami, 1963: 174–175.
Amphioplus diacritus Murakami, 1943c: 225–227, fig. 1.

Material examined. Kamo Bay; 19 m, 1
specimen (NSMT E-4843). Saigou Bay; (M), 28
specimens (NSMT E-4844).
Remarks. Particularly in larger specimens
ventral arm plates within the disk have two lateral lobes and one medial lobe at the distal margin.
Amphioplus asterictus seems to be characterizedby tulip-shaped ventral arm plates like those ofAmphioplus diacritus Murakami, 1943 (see Murakami, 1943c: fig. 1), which was synonymized
with A. asterictus by Murakami (1963). The ventral arm plate of the holotype of A. asterictus (A.
M. Clark, 1965: fig. 5) lacks a distinctly tuliplike shape probably due to its small size (disk diameter 8 mm).
Geographic range. This species is distributedin Korea and Japan. Korea, the Yellow Sea (Yi &Irimura, 1987). The Sea of Japan, Notojima Is-196

Toshihiko Fujita et al.

land (Fujita & Kohtsuka, 2003), the Oki islands(present study). Pacific coast of Japan, Tokyo
Bay (type locality, H. L. Clark, 1915a), Sagami
Bay (A. M. Clark, 1965), Kii Peninsula (Murakami, 1943c; 1963).
Amphioplus (Lymanella) megapomus
H. L. Clark, 1911
[Japanese name: Miyadi-kumohitode]
Amphioplus megapomus H. L. Clark, 1911: 170–171; Irimura, 1981: 25, fig. 2.
Amphioplus miyadii Murakami, 1943c: 227–229, fig. 2.
?Amphiura laevis Lyman, 1874: 229, pl. 4 fig. 18–21.
?Amphioplus (Lymanella) laevis: A. M. Clark, 1970: 52,
53, 75.

Material examined. Kamo Bay; 10 m (M), 5
specimens (NSMT E-4845).
Remarks. The specimens agree well with thedescription of Amphioplus miyadii by Murakami(1943c). A. M. Clark (1970) suggested that A.
miyadii may be synonymous with Amphioplus
laevis (Lyman, 1874), but Irimura (1981) indicated that A. miyadii is a junior synonym of A.
megapomus H. L. Clark, 1911. A. M. Clark(1970) also suggested that A. megapomus is synonymous either with A. laevis or with Amphioplus andreae (Lütken, 1872), so both A. megapomus and A. miyadii may be possibly a synonymof A. laevis though further studies are required toclarify the synonymy.
Geographic range. This species is distributedin Korea and Japan. Korea, the Yellow Sea (Yi &Irimura, 1987). The Sea of Japan, Hakata Bay(Murakami, 1943c), the Oki Islands (present
study). Pacific coast of Japan, Kii-suido Channel(type locality, H. L. Clark, 1911), Kii Peninsula(Murakami, 1943c; Irimura, 1981), Beppu Bay(Murakami, 1943c).
Amphipholis squamata (Delle Chiaje, 1829)[Japanese name: Iso-komochi-kumohitode]

Material examined. Off Onbejima Island;
52 m, 1 specimen (NSMT E-4846); 71 m, 1 specimen (NSMT E-4847). Off Shijikijima Island;

65 m (G), 2 specimens (NSMT E-4848). East of
Shijikijima Island; 15 m (G), 1 specimen (NSMT
E-4849). Kamo Bay; subtidal, 1 specimen(NSMT E-4850).
Remarks. See A. M. Clark (1987) for thenomenclature of this species. Dupont (2002) discussed the possibility of sibling species.
Geographic range. This species is widelydistributed in warm and temperate areas in theworld, and has been recorded mainly from intertidal but from depths up to 1200 m (Paterson,1985). It is widely distributed in the Sea of Japan(see Irimura, 1982; Ishida et al., 2001; present
study).
Amphipholis kochii Lütken, 1872
[Japanese name: Suna-kumohitode]

Material examined. Kamo Bay; subtidal, 2specimens (NSMT E-4851); subtidal 3 specimens (NSMT E-4852).
Geographic ranges. This species is distributed in the Sea of Japan and coastal waters ofJapan. The Sea of Japan; Russia, Peter the Great
Bay, Tatar Strait (Djakonov, 1954); Korea, East
Sea (Shin & Rho, 1996); Japan, Sado (Irimura,1979), Notojima Island (Fujita & Kohtsuka,
2003), the Oki Islands (present study). Japan,
Mutsu Bay (Matsumoto, 1917, 1941), Misaki(Matsumoto, 1917), Izu (Murakami, 1942),
Amakusa (Murakami, 1944; Irimura, 1969). The
type locality is probably the Sea of Japan (“eastern Manchow” by Lütken, 1872).
Amphiura kandai Murakami, 1942
[Japanese name: Hakkou-kumohitode](Fig. 3)
Amphiura kandai Murakami, 1942: 11–12, fig. 3; Irimura
et al. , 2001: 313, 314.

Material examined. Off Onbejima Island;
67 m, 1 specimen (NSMT E-4853).
Remarks. Disk diameter is 2.1 mm. Arms are
six in number, 11.3 mm in length. Disk is circularand covered with many coarse, thin, imbricating
scales without any granules or spines (Fig. 3A).

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197

Fig. 3. Amphiura kandai. NSMT E-4853 (disk diameter 2.1 mm). A: Disk, aboral view. B: Arm base, aboral
view. C. Arm spines (arrows) at 6th and 7th arm segments (corresponding 3rd and 4th free arm segments).
Aboral side upward. D: Disk and basal arm, oral view. E: Mouth frame. F: Jaws and teeth, obliquely oral
view. A–B: Dry specimen, C, F: Alcohol specimen, D–E: SEM specimen. Scales 1 mm (A), 0.5 mm (B, D),
0.2 mm (C), 0.1 mm (E–F).

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Toshihiko Fujita et al.

Primary plates are indistinguishable. Radial
shields are more than twice as long as broad, separated from each other by a few intervening
scales except at their distal edge. Oral interradius
is covered with scales similar to aboral ones (Fig.
3D). Oral shields are almost circular or rhomboidal with rounded angles, about as long as
broad. Adoral plates are in contact or with asmall gap radially. Oral plates are small, and intheir distal part separated radially. There are twoor three oral papillae on each side of the oral
plate (Fig. 3D–F). The oral-most teeth are pentagonal with rather pointed edge; two other teeth
are rectangular with rounded edges (Fig. 3E–F).
Tentacle scale of oral tentacle pores is single androunded, on proximal edge of adoral plate (Fig.
3D–E). Dorsal arm plates are transversely oval orsomewhat triangular with rounded angles, muchwider than long, not in contact with each other(Fig. 3B). First ventral arm plates are fan-shaped,
slightly longer than broad. Following plates arepentagonal, longer than broad, with acute proximal angle, separated from each other by a narrowinterval except first a few plates (Fig. 3D). Lateral arm plates meet on both aboral and oral sides.
Arm spines are four on proximal 2–4 free arm
segments, and three on more distal segments.
They are conical, subequal, and similar in length
to the corresponding arm joint (Fig. 3C). Tentacle scales are two to each pore, but one in distal
part of arm (Fig. 3D). Color of specimen in alcohol is almost whitish, and light greenish yellow
color within body can be seen through dorsal
disc scales.
Some deviations from Murakami’s (1942) description are recognized. He described the radial
shields as completely separated, and the oral
shields as rhomboidal or triangular. He mentioned that the species has ‘very peculiar mouthparts’. There are no distinct paired infradental
papillae in the present specimen, and that seems
different from normal mouth structure of Amphiura species. Therefore, further studies are required to evaluate the systematic position of thisspecies.
Two small specimens (disk diameter 1.1–1.5mm) of Amphiura kandai were collected from
Shimoda (see Irimura et al., 2001), and one ofthem has three thick arms and three thin arms (T.
Fujita, unpublished data), which indicates thatthe species may be fissiparous although Murakami (1942) did not comment on that. Amphiura
velox Koehler, 1910 with a junior synonym ofAmphiura sexradiata Koehler, 1931 is the only
other 6-armed fissiparous Amphiura species (Devaney, 1974). Amphiura kandai is luminous (seeMurakami, 1942), and its luminous cells havebeen studied histologically (Kato, 1947; Okada& Kato, 1950).
Geographic range. This species was previously known only from Shimoda, Izu, Pacificcoast of Japan (type locality, Murakami, 1942;
Irimura et al., 2001). This is the first record fromthe Sea of Japan.
Amphiura (Amphiura) luetkeni Duncan, 1879
[New Japanese name: Rutsuken-suna-kumohitode](Fig. 4)

Material examined. Off Onbejima Island;
67 m, 1 specimen (NSMT E-4868); 67 m (S, Sh),
2 specimens (NSMT E-4863); 71 m, 2 specimens(NSMT E-4859). East of Onbejima Island; 54 m,
1 specimen (NSMT E-4858). Off Tsudo; 62 m, 1
specimen (NSMT E-4869). South-southeast of
Tsudo; 62–65 m (G), 3 specimens (NSMT E4865). Off Shijikijima Island; 36°9N, 133°14E,
20–30 m (G, R), 3 specimens (NSMT E-5004);
56 m, 1 specimen (NSMT E-4854); 60 m (R), 1
specimen (NSMT E-4855); 60 m (R), 3 specimens (NSMT E-4856); 60 m (R), 2 specimens(NSMT E-4857); 65 m (G), 15 specimens(NSMT E-4860); 65 m (G), 2 specimens (NSMT
E-4998); 65 m (G), 1 specimen (NSMT E-4861);
65 m (G), 1 specimen (NSMT E-4862). East of
Shijikijima Island; 15 m (G), 1 specimen (NSMT
E-4866). Off Kanbi; 38 m, 1 specimen (NSMT
E-4864). South of Dougo Island (no detailed locality data); 1 specimen (NSMT E-4867).
Remarks. This species is distinguished fromthe closely related and co-occurring species Amphiura micraspis by the form of arm spines. The

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199

Fig. 4. Amphiura (Amphiura) luetkeni. NSMT E-4998 (disk diameter 3.2 mm). A: Disk, aboral view. B: Basalarm, aboral view. C: Middle arm, aboral view. D: Mouth frame. E: Arm base, oral view. F: Middle arm, oral
view. G: Arm spines, basal arm, proximal side right. H: Arm spines, middle arm, obliquely distal view. I:
Arm spines, distal arm, ventro-distal view. A–I: Dry specimen. Scales 1 mm (A), 0.5 mm (B–I).

oral-most arm spine is not conspicuously longer
but rather wider than the others in A. luetkeni(Fig. 4H–I), while that of A. micraspis is very
long (Fig. 5G–I).
Geographic range. This species is distributedin the Sea of Japan and Indo-West Pacific. Korea(type locality, Duncan, 1879). The Sea of Japan,
the Oki Islands (present study). Japan, Amakusa(Murakami, 1944); Nagasaki (A. M. Clark &Rowe, 1971: pl. 13 fig. 6–7). The Philippines,
Sulu Sea (Koehler, 1905). Indonesia, Ambon (deLoriol, 1893 as Amphiura duncani), Banda Sea,
Java Sea, the Sangihe Islands (Koehler, 1905).
New Caledonia (Guille et al., 1986); Ceylon (H.
L. Clark, 1915b).

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Fig. 5. Amphiura (Amphiura) micraspis. NSMT E-4997 (disk diameter 3.7 mm). A: Disk, aboral view. B: Basalarm, aboral view. C: Middle arm, aboral view. D: Mouth frame. E: Basal arm, oral view. F: Middle arm, oral
view. G: Arm spines, basal arm, proximal side right. H: Arm spines, middle arm, proximal side right. I: Armspines, distal arm, proximal side left. A–I: Dry specimen. Scales 1 mm (A), 0.5 mm (B–I).

Amphiura (Amphiura) micraspis
H. L. Clark, 1911
[Japanese name: Chibitate-suna-kumohitode](Fig. 5)Material examined. Off Onbejima Island;
60 m (S, Sh), 1 specimen (NSMT E-4878); 71 m,
1 specimen (NSMT E-4874); 71 m, 7 specimens(NSMT E-4875); 71 m, 1 specimen (NSMT E-

4876); 67 m, 2 specimens (NSMT E-4881). Eastof Onbejima Island; 54 m, 3 specimens (NSMT
E-4873). Shijikijima Island; subtidal, 1 specimen(NSMT E-4882). Off Shijikijima Island; 56 m, 2
specimens (NSMT E-4871); 60 m (R), 27 specimens (NSMT E-4872); 65 m (G), 28 specimens(NSMT E-4877); 65 m (G), 2 specimens (NSMT
E-4997); 65 m (G), 15 specimens (NSMT E4879). South-southeast of Tsudo; 62–65 m (G), 9

Ophiuroids from the Oki Islands, Japan

specimens (NSMT E-4880). Tsutsuka Bay; subtidal, 1 specimen (NSMT E-4870).
Remarks. See above for the comparison ofthis species with Amphiura luetkeni.
Geographic range. This species is distributedin Korea and Japan. Korea (Yi, 1983). The Sea ofJapan, the Oki Islands (present study). Pacificcoast of Japan, Boso Peninsula (Irimura, 2000),
Suruga Bay (Fujita et al., 1997), Omaizaki (typelocality, H. L. Clark, 1911), Amakusa (Irimura,1969).
Ophiocentrus koehleri Gislén, 1926
[Japanese name: Igaguri-kumohitode](Fig. 6)
Ophiocentrus verticillatus: Matsumoto, 1917: 213–214,
fig. 59; Murakami, 1942: 20; Irimura, 1969: 42–43(non Ophiocnida verticillata Döderlein, 1896).
Ophiocentrus aculeatus: Koehler, 1922: 196–199, pl. 71
fig. 1–3, pl. 96 fig. 8 (non Ophiocentrus aculeatus
Ljungman, 1867)
Ophiocentrus koehleri Gislén, 1926:13; Koehler, 1931:
119; Guille and Wolff, 1984: 14–15, pl. 2 fig. A-B;
Liao & A. M. Clark, 1995: 204–205, fig. 100; Irimura,
2000: 403, fig. 23–46; Ishida et al., 2001: 4.

Material examined. Off Onbejima Island;
45 m, 1 specimen (NSMT E-4883). Off Shijikijima Island; 65 m (G), 1 specimen (NSMT E4884).
Remarks. Matsumoto (1917) reported two
specimens of Ophiocentrus verticillatus (Döderlein, 1896) from Sagami Bay, Japan. Gislén(1926) considered Matsumoto’s specimens not as
O. verticillatus but as Ophiocentrus vexator described by Koehler (1922) from the Philippines
based on the arm spines with thorny end. But
later workers (Murakami, 1942; Irimura, 1969)
used O. verticillatus for Japanese specimens according to Matsumoto (1917). Gislén (1926) established another species Ophiocentrus koehleri
for the specimen reported as Ophiocentrus aculeatus Ljungman, 1966 by Koehler (1922) from
Hong Kong, characterized by naked disk except
around radial shields, and, recently, Guille andWolff (1984) recognized Matsumoto’s (1917) O.
verticillatus as Ophiocentrus koehleri. Irimura

201(2000) and Ishida et al. (2001) adopted Guilleand Wolff’s (1984) view and determined Japanese Ophiocentrus specimens as O. koehleri.
The present specimens have no completely
naked area and disk scales covers the whole disk.
The larger specimen (NSMT E-4883, disk diameter 11.6 mm) has relatively large, thick scales
around radial shields which form brighter-color
area than the other part of disk where there aresmall and thin scales in the skin (Fig. 6A). The
smaller specimen (NSMT E-4884, disk diameter
2.9 mm) has uniform scaling and shows no large
difference in size of scales between around radial
shield and the other part of disk (Fig. 6D). Twoother NSMT specimens collected from Misaki(NSMT E-1683) and Enoshima (NSMT E-1258),
the same locality as Matsumoto’s (1917) specimens, were also examined. In the large specimen(NSMT E-1258, disk diameter 16.6 mm), relatively large scales are clearly seen from the surface around radial shields while at the other partof disk small scales are buried in the skin andonly spine-supporting scales are visible outsideof the skin (Fig. 6H). The smaller one (NSMT E1683, disk diameter 7.3 mm) has uniform disk
scales (Fig. 6I) similar to the smaller Oki specimen (NSMT E-4884). Thus, Japanese specimens
seem to have more or less scales on whole disk
but smaller scales tend to bury in the skin at center and interradius like as reported by Guille andWolff (1984). There is a contrastive difference inscaling, at least in large-sized specimens, between around radial shields and at the other partof disk, and this is enough to assign the specimens from the Oki Islands as well as Matsumoto’s (1917) and NSMT specimens from Sagami
Bay to O. koehleri distinguished from O. vexator
and O. verticillatus. In addition, the disk spinesof O. vexator (Koehler, 1922: pl. 62 fig. 1, 4–5;
Guille & Wolff, 1984: pl. 2 fig. C–D) and O. verticillatus (Döderlein, 1896: pl. 15 fig. 7–7a) seemto be longer and thinner, and more dense thanthose of O. koehleri (Fig. 6; Guille & Wolff,
1984: pl. 2 fig. A–B). Most of the disk scales
bear no spine in the Japanese O. koehleri specimens, while the almost all disk scales have spines

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Toshihiko Fujita et al.

Fig. 6. Ophiocentrus koehleri. A–C: NSMT E-4883 (disk diameter 11.6 mm). D–G: NSMT E-4884 (disk diameter 2. 9 mm). H: NSMT E-1258 (disk diameter 16.6 mm). I: NSMT E-1683 (disk diameter 7.3 mm). A: Disk
and basal arm, aboral view. B: Disk, oral view. C: Jaw. D: Disk, aboral view. E: Radial shield and basal arm.
F. Jaw. Spinelets on oral shields shown by arrows. G: Oral interadial. H: A part of disk, aboral view. I: Disk
and basal arm, aboral view. A–G: Alcohol specimen. Scales 2 mm (A–B, H–I), 1 mm (C–D), 0.5 mm (E–G).

Ophiuroids from the Oki Islands, Japan

203

Fig. 7. Ophiocentrus verticillatus. Holotype (disk diameter 7.5 mm). Ambon, Indonesia. Musée Zoologique
Strasbourg. A: Disk, aboral view. B: Arm spines of 15th arm segment. Scales 2 mm (A), 0.5 mm (B).

in the holotype of Ophiocentrus verticillatus(Fig. 7A, Musée Zoologique Strasbourg, disk diameter 7.5 mm). The disk spines of the present
specimens are mostly acute, but sometimes with
two or more terminals on their tip.
Guille and Wolff (1984) reported that the characteristics of arm spines are very similar between
O. koehleri and O. vexator. The maximum number of arm spines of the specimens from the Oki
Islands varies from 7 to 10. They are usuallymore or less flat, rarely cylindrical, with usuallythorny and sometimes blunt tips, and their lengthis subequal or the lower most one is sometimes
longer than the others. Matsumoto (1917) reported that arm spines of his specimens with thorny
end are different from flat lancet-form arm spinesof O. verticillatus (Döderlein, 1896: 287, pl. 14
fig. 2b), though, in fact, some of the arm spines
are a little thorny at the end (Fig. 7B). Koehler(1922) also described O. vexator which is very
close to O. verticillatus but distinguished from itby the arm spines. Guille and Wolff (1984) indicated that O. verticillatus is rather easily distinguished from O. vexator by the characteristics ofarm spines and that further detailed studies mayneed to compare O. aculeatus and O. koehleri
with O. vexator, but O. verticillatus should bealso compared together with them because ofvariation of the arm spines.
We should suggest one additional character of
O. koehleri which has not been mentioned in previous works. The oral shields of the smaller spec-

imen have a characteristic pair of spinelets at
both sides of the distal edge (Fig. 6F).
Geographic range. This species is distributedin Japan, China and Indonesia. The Sea of Japan,
Tsushima Strait (Ishida et al., 2001), the Oki Islands (present study). Japan, Boso Peninsula (Irimura, 2000), Sagami Bay, Enoshima and Misaki(Matsumoto, 1917), Izu (Murakami, 1942),
Amakusa (Irimura, 1969), the Ogasawara Islands(Irimura & Tachikawa, 2003). China, Hong Kong
(type locality; Koehler, 1922; Gislén, 1926; Liao
& A. M. Clark, 1995), Gulf of Tonkin (Liao & A,
M, Clark, 1995). Indonesia, Kei (Koehler, 1931),
Timor (Guille & Wolff, 1984).
Family Ophiotrichidae
Ophiogymna fulgens (Koehler, 1905)[Japanese name: Toge-tosaka-kumohitode]

Material examined. Off Shijikijima Island;
65 m (G), 1 specimen (NSMT E-4885).
Remarks. This species is usually epizoic onsoft corals (Irimura, 1982), although the host of
the present specimen is not known.
Geographic range. This species is distributed
from the Sea of Japan to Indonesia in the west
Pacific Ocean, and in the Indian Ocean. Korea(Rho & Shin, 1987; Shin & Rho, 1996). The Sea
of Japan, Korea Strait (H. L. Clark, 1911 as
Ophiothrix macrobrachia), the Oki Islands (present study). Pacific coast of Japan from Boso
Peninsula to off Kagoshima Bay (see Matsumoto,204

Toshihiko Fujita et al.

1917; Koehler, 1922; Irimura, 1982, 2000); theOgasawara Islands (Irimura & Tachikawa, 2003);
off the Goto Islands (H. L. Clark, 1911 as Ophiothrix macrobrachia); East China Sea (Irimura &Kubodera, 1998). Philippines (Koehler, 1922,1931). Indonesia (Koehler, 1931). Indian Ocean,south Arabia (H. L. Clark, 1939 as Placophiothrix phrixa), south Africa (Mortensen, 1933; A.
M. Clark & Courtman-Stock, 1976). The type
locality is the Kei Islands and the Paternoster
Islands (Tengah Archipelago), Indonesia(Koehler, 1905).
Ophiothrix exigua Lyman, 1874
[Japanese name: Naga-toge-kumohitode](Fig. 8)
Ophiothrix exigua Lyman, 1874: 226–237, pl. 4 fig.
24–26; Irimura, 1981: 28–29, pl. 1 fig. 4.
Ophiothrix (Ophiothrix) exigua: Irimura, 1982: 43–46,
fig. 28, pl. 8 fig. 2–6, pl. 10 fig. 1–3, 5, 6. Liao & A.
M. Clark, 1995: 247–248, fig. 131.
Ophiothrix marenzelleri Koehler, 1904: 103–105, figs.
76–78; Liao & A. M. Clark, 1995: 248–250, fig. 132.

Material examined. Between Oomorijima Island and Shijikijima Island; 58 m, 3 specimens(NSMT E-4891). Off Nagu; 67 m, 3 specimens(NSMT E-4887). Off Onbejima Island; 45 m, 1
specimen (NSMT E-4889); 52 m, 1 specimen(NSMT E-4892); 52 m, 2 specimens (NSMT E4924); 60 m (S, Sh), 3 specimens (NSMT E4900); 67 m (S, Sh), 1 specimen (NSMT E4901); 67 m, 8 specimens (NSMT E-4923); 71 m,
13 specimens (NSMT E-4895). East of Onbejima
Island; 54 m, 3 specimens (NSMT E-4894); 62 m
(R), 2 specimens (NSMT E-4886). South-southeast of Tsudo; 62–65 m (G), 1 specimen (NSMT
E-4905). Off Shijikijima Island; 36°9N, 133°14
E, 20–30 m (G, R), 4 specimens (NSMT E5002); 51 m, 6 specimens (NSMT E-4920); 60 m
(R), 11 specimens (NSMT E-4893); 65 m (G), 1
specimen (NSMT E-4896); 65 m (G), 13 specimens (NSMT E-4897); 65 m (G), 4 specimens(NSMT E-4898); 65 m (G), 24 specimens(NSMT E-4902); 70 m (R, S), 5 specimens(NSMT E-4888). East of Shijikijima Island; 13 m

(G), 4 specimens (NSMT E-4910); 15 m (G), 5
specimens (NSMT E-4906); 15 m (G), 8 specimens (NSMT E-4907); 15 m (G), 1 specimen(NSMT E-4908). Between Matsushima Island
and Kanjima Island; 16–18 m (Sh), 1 specimen(NSMT E-4909). Off Kanjima Island; 34 m, 1
specimen (NSMT E-4918); 34 m, 4 specimens(NSMT E-4919). Off Kanbi; 38 m, 1 specimen(NSMT E-4904). Kamo Bay; 22 m (seagrassbed), 2 specimens (NSMT E-4903); 53 m, 3
specimens (NSMT E-4916); 30 m (M), 1 specimen (NSMT E-4899). Off Kamo Bay; 43 m, 2
specimens (NSMT E-4917). Tsutsuka Bay; subtidal, 9 specimens (NSMT E-4890). South ofDougo Island (no detailed locality data); 1 specimen (NSMT E-4921); 1 specimen (NSMT E4922); 1 specimen (NSMT E-4911); 1 specimen(NSMT E-4912); 2 specimens (NSMT E-4913);
1 specimen (NSMT E-4914); 10 specimens(NSMT E-4915).
Remarks. A specimen (NSMT E-4914, disk
diameter 12.3 mm; Fig. 8) is very close to Ophiothrix marenzelleri Koehler, 1904, which was synonymized with O. exigua by Irimura (1981). Liaoand A. M. Clark (1995) recognized a clear distinction between these two species in China seas;in Japanese waters, however, a large variation in
disk armature is observed and many intermediate
forms were found according to Irimura (1982).
Geographic range. This species is distributedin the Sea of Japan and widely in Indo-west Pacific (see A. M. Clark & Rowe, 1971). It is acommon species in the Sea of Japan (see Irimura,
1982; Ishida et al., 2001; Fujita & Kohtsuka,
2003; present study). The type locality is thePhilippines (Lyman, 1874)
Family Ophiactidae
Ophiactis affinis Duncan, 1879
[Japanese name: Kusairo-chibi-kumohitode](Figs. 9–10)

Material examined. Off Shijikijima Island;
65 m (G), 1 specimen (NSMT E-4925). East of
Shijikijima Island; 15 m (G), 1 specimen (NSMT
E-4926); 15 m (G), 1 specimen (NSMT E-4927).

Ophiuroids from the Oki Islands, Japan

205

Fig. 8. Ophiothrix exigua. NSMT E-4914 (disk diameter 12.3 mm). A: Disk, aboral view. B: Basal arm, aboral
view. C: Mouth frame. D: Basal arm, oral view. E: Dorsal arm plate at middle part of arm, proximal side left.
F: Disk spines at central disk. G: Disk spines at disk edge. H: Arm spines near arm base. Eight spines present
on each side. I: Arm spines at middle part of arm. J: Arm spine. A–G: Alcohol specimen, H–I: Specimen
gently bleached to remove skin, J: SEM specimen. Scales 2 mm (A–D), 0.5 mm (E–G), 1 mm (H–I), 0.05 mm
(J).

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Toshihiko Fujita et al.

Fig. 9. Ophiactis affinis. NSMT E-4926 (disk diameter 1.1 mm). A: Disk and basal arms, aboral view. B: Disk
and basal arms, oral view. C: Arm spines. A–C: Alcohol specimen. Scales 0.5 mm (A–C).

Fig. 10. Ophiactis affinis. NSMT E-4926 (diskdiameter 1.1 mm). Jaw. Abbreviations. as: adoral shield, op: oral plate, opa: oral papillae, os:oral shield. Scale 0.1 mm.

Remarks. Three small specimens (disk diameter 0.85–1.3 mm) were collected. They have fivearms and a few spines on the disk margin (Fig.
9A). The oral shield has a distal lobe (Fig. 10),and this seems a characteristic feature of Ophiactis affinis (see Duncan 1879: fig. 24; Koehler
1922: pl. 63 fig. 5; Liao & A. M. Clark, 1995:
fig. 107b). Liao and A. M. Clark (1995: 214) reported the dorsal arm plates are elliptical andabout twice as broad as long, but those of thepresent specimens are not so wide but pentagonal in shape and as long as wide probably due totheir small size.
Geographic range. This species is distrib-

uted from the Sea of Japan to the Gulf of Thailand in the west Pacific Ocean, and in India. The
Sea of Japan, Korean Strait (type locality, Duncan, 1879; Koehler, 1922), Notojima Island (Fujita & Kohtsuka, 2003), the Oki Islands (present
study). Korea, the Yellow Sea (Yi & Irimura,1987). Japan, Boso Peninsula (Irimura, 2000),
Kii Peninsula (Irimura, 1981). China (Chang,1948; Liao & A. M. Clark, 1995). The Philippines, San Juanico Strait (Koehler, 1922). Gulf ofThailand (Koehler, 1931). India, Chilika Lake
(Koehler, 1898).
Ophiactis macrolepidota MarktannerTurneretscher, 1887
[Japanese name: Dairin-chibi-kumohitode](Fig. 11)
Ophiactis macrolepidota Marktanner-Turneretscher,1887: 298, pl. 12 fig. 12–13; A. M. Clark & Rowe,
1971: 104; Rowe & Gates, 1995: 379.
Ophiactis delicata H. L. Clark, 1915a: 260–261, pl. 11
fig. 9–10; A. M. Clark & Rowe, 1971: 104.
Ophiactis parva Mortensen, 1926: 123–124, fig. 12; A.
M. Clark & Rowe, 1971: 104.
Ophiactis acosmeta H. L. Clark, 1938: 262–264; A. M.
Clark & Rowe, 1971: 104.

Fig. 11. Ophiactis macrolepidota. A–B: Holotype (Naturhistorisches Museum Wien, Specimen No. T66/2,
10340; disk diameter 3.1 mm). C–D: NSMT E-4932 (disk diameter 3.6 mm). E–H: NSMT E-4935 (disk diameter 2.3 mm). A: Disk and basal arm, aboral view. B: Disk, oral view. C: Disk and basal arm, aboral view.
D: Disk and basal arm, oral view. E: Disk and basal arm, aboral view. F: Disk and basal arm, oral view. G:
Oral interradius with a few acute spines (arrows). H: Arm spines of between 2nd and 4th free arm segments(shown by 2, 3, 4). A–D: Semi-dry specimen, E–H: Dry specimen. Scales 1 mm (A–E, G), 0.5 mm (F, H).

Ophiuroids from the Oki Islands, Japan

207

208

Toshihiko Fujita et al.

Material examined. Off Onbejima Island;
67 m, 1 specimen (NSMT E-4943). Shijikijima
Island; subtidal, 3 specimens (NSMT E-4942).
Off Shijikijima Island; 36°9N, 133°14E, 20–
30 m (G, R), 11 specimens (NSMT E-5003);
51 m, 18 specimens (NSMT E-4941); 54 m, 1
specimen (NSMT E-4931); 56 m, 1 specimen(NSMT E-4929); 56 m, 1 specimen (NSMT E4930); 65 m, 3 specimens (NSMT E-4932); 65 m
(G), 2 specimens (NSMT E-4933). East of Shijikijima Island; 15 m (G), 2 specimens (NSMT
E-4934); 15 m (G), 1 specimen (NSMT E-4935);
15 m (G), 2 specimens (NSMT E-4936); 15 m
(G), 6 specimens (NSMT E-4937); 13 m (G), 2
specimens (NSMT E-4939). Between Matsushima Island and Kanjima Island; 16-18 m (Sh), 3
specimens (NSMT E-4938). Kamo Bay; subtidal,
6 specimens (NSMT E-4940). Tsutsuka Bay;
subtidal, 3 specimens (NSMT E-4928). ?Ophiactis macrolepidota: Kamo Bay; subtidal, 2 specimens (NSMT E-4944).
Remarks. This fissiparous species was described by Marktanner-Turneretscher (1887)
based on a specimen presumably collected from
Sydney, Australia. He reported that the oral interradius is free of scales; however, the holotype(Naturhistorisches Museum Wien, Specimen No.
T66/2, 10340) has rough scales (Fig. 11A–B).
Due to the inaccuracy of the original description,
O. macrolepidota may be known by other names,
as suggested by H. L. Clark (1946) and A. M.
Clark (1965). Recently Rowe (in Rowe & Gates,
1995) synonymized three Australian species with
O. macrolepidota, and suggested the possibilityof additional synonyms and cosmopolitan distribution of O. macrolepidota.
The present specimens have six arms except
one specimen with seven arms, two with fivearms, and one with only three arms probably collected soon after fission. Some variation was observed for the spinelets in oral interradius and thenumber of arm spines. Among a total of 66 examined specimens (disk diameter range 0.8–3.6mm), only six specimens have a small number of
acute spines in the oral interradius (Fig. 11F–G).
None of the specimens smaller than 1.7 mm indisk diameter have such spines. Forty-five specimens have segments with four-arm spine segments on at least one arm. Usually, the second
free arm segment has four arm spines and thefollowing segments have only three (Fig. 11H).
Eleven of the specimens smaller than 1.7 mm indisk diameter have no segment with four arm
spines.
Additionally, two specimens with five arms
(NSMT E-4944, disk diameter 1.5 and 1.1 mm)
are probably O. macrolepidota, and both of them
have several arm segments with 4 arm spines.
Geographic range. This species is distributedin Korea, Japan, Indonesia, and Australia. The
Sea of Japan, Tsushima (Matsumoto, 1917), theOki Islands (present study). Korea (Rho, 1979;
Yi & Irimura, 1987; Shin & Rho, 1996). Pacificcoast of Japan, Uraga Channel (Matsumoto,1917), Shimoda (Murakami, 1942), Jogashima
and Katsuyama (referred to Yogashima andKashiyama in Döderlein’s collection: A. M.
Clark, 1965), Kii Peninsula (Irimura, 1981).
Japan, Amakusa (Irimura, 1969). Indonesia,
Ambon (Döderlein, 1898). Australia (Rowe &Gates, 1995), Sydney (presumably type locality,
Marktanner-Turneretscher, 1887; A. M. Clark &Rowe, 1971).
Ophiactis savignyi (Müller & Troschel, 1842)[Japanese name: Chibi-kumohitode]

Material examined. Off Tsuma; 50 m, 1
specimen (NSMT E-4945). Shijikijima Island;
subtidal, 1 specimen (NSMT E-4946).
Remarks: This species is also fissiparous andnormally has 6 arms, but occasionally 5-arm
specimens are found (Liao & A. M. Clark, 1995).
It is distinguishable from Ophiactis macrolepidota usually by having two oral papillae though
Liao & A. M. Clark (1995) reported that somespecimens have only one oral papilla.
Geographic range. This species is distributedin the Sea of Japan and widely in temperate andtropical waters in the world (see A. M. Clark &Rowe, 1971). The Sea of Japan, Sado Island (Irimura, 1979), the Oki Islands (present study);

Ophiuroids from the Oki Islands, Japan

Korea (Rho, 1979; Rho & Shin, 1987). The type
locality is the Red Sea (Muller & Troschel, 1842
as Ophiolepis savignyi ).
Ophiopholis mirabilis (Duncan, 1879)[Japanese name: Madara-kumohitode]

Material examined. Off Onbejima Island;
67 m (S, Sh), 1 specimen (NSMT E-4948). Eastof Onbejima Island; 62 m (R), 3 specimens(NSMT E-4947). Off Tsudo; 62 m, 1 specimen(NSMT E-4951). South-southeast of Tsudo; 62–
65 m (G), 1 specimen (NSMT E-4949). South ofDougo Island (no detailed locality data); 1 specimen (NSMT E-4950).
Remarks. Four Ophiopholis species havebeen reported from the Sea of Japan: O. aculeata(Linnaeus, 1767), O. brachygenis H. L. Clark,
1911, O. japonica Lyman, 1879, and O. mirabilis(Ishida et al., 2001). Ophiopholis mirabilis are
distinguished from the other three by the presence of large supplemental plates on both side of209

arms (see Irimura 1990: 83, fig.).
Geographic range. This species is distributed
the Sea of Okhotsk, the Sea of Japan, the Yellow
Sea, and Pacific coast of northern Japan. The Seaof Okhotsk (Matsumoto, 1917; Djakonov, 1954).
Korea, Sondai Bay (type locality, Duncan, 1879
as Ophiolepis mirabilis), Cheju Island (Rho,
1979), the Yellow Sea (Yi & Irimura, 1987). The
Sea of Japan, Notojima Island (Fujita & Kohtsuka, 2003), the Oki Islands (present study). Pacificcoast of Japan from Hakodate and Mutsu Bay to
Sagami Bay (H. L. Clark, 1911; Matsumoto,
1917, 1941; Koehler, 1922; Irimura, 1982, 1990).
Family Ophionereididae
Ophionereis dubia (Müller & Troschel, 1842)[Japanese name: Amime-kumohitode](Fig. 12D)

Material examined. Off Onbejima Island;
71 m, 1 specimen (NSMT E-4960). Off Shijikijima Island; 65 m (G), 1 specimen (NSMT E-

Fig. 12. A–C: Ophionereis intermedia, D: Ophionereis dubia. A-B: NSMT E-4965 (disk diameter 5.1 mm), C:
NSMT E-4969 (disk diameter ca. 6 mm). D: NSMT E-4959 (disk diameter 5.9 mm). A: Disk and basal arms,
aboral view. B: Disk, oral view. C: Genital papillae (arrows). D: Genital slit without genital papillae. A–B:
Alcohol specimen, C–D: Dry specimen. Abbreviations. os: oral shield, gs: genital slit. Scales 2 mm (A),
0.5 mm (B–D).

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Toshihiko Fujita et al.

4961). East of Shijikijima Island; 15 m (G), 1
specimen (NSMT E-4962); 15 m (G), 1 specimen(NSMT E-4963). Tsutsuka Bay; subtidal, 3 specimens (NSMT E-4958); subtidal, 1 specimen(NSMT E-4959).
Geographic range. This species is distributedin the Sea of Japan and widely in Indo-west Pacific (see A. M. Clark & Rowe, 1971). Widelydistributed in the Sea of Japan (see Ishida et al.,
2001; Fujita & Kohtsuka, 2003). Pacific coast ofJapan in Boso Peninsula (Irimura, 2000) and themore southern and western area (see Irimura,1982). The type locality is the Red Sea (seeLyman, 1865).
Ophionereis intermedia A. M. Clark, 1953
[Japanese name: Nise-amime-kumohitode](Fig. 12A–C)
Ophionereis intermedia A. M. Clark, 1953: 90–92, fig.
12.

Material examined. Off Onbejima Island;
71 m, 3 specimens (NSMT E-4966). Southsoutheast of Tsudo; 62–65 m (G), 2 specimens(NSMT E-4968); 62–65 m (G), 1 specimen(NSMT E-4969). Off Shijikijima Island; 36°9N,
133°14E , 20–30 m (G, R), 2 specimens (NSMT
E-5000); 60 m (R), 1 specimen (NSMT E-4965);
65 m (G), 1 specimen (NSMT E-4967). East of
Shijikijima Island; 13 m (G), 1 specimen (NSMT
E-4970). Off Kanjima Island; 34 m, 4 specimens(NSMT E-4971). Kanbi; subtidal, 1 specimen(NSMT E-4964).
Remarks. This species is similar to
Ophionereis dubia, but differs from it by havingvery minute, acute genital papillae (Fig. 12C).
The scaling of the disk is very fine like O. dubia,
but is more distinct.
Geographic range. This species is distributedin Japan and Australia. The Sea of Japan, Notojima Island (Fujita & Kohtsuka, 2003), the Oki Islands (present study). Northern Japan, Tsugaru
Strait (type locality, A. M. Clark, 1953). Australia, Swain Reef (Kingston, 1980).

Ophionereis sasakii A. M. Clark, 1953
[New Japanese name: Sasaki-kumohitode](Fig. 13)
Ophionereis sasakii A. M. Clark, 1953: 75–77, fig. 4, pl.
3 fig. 1–2; Yi & Irimura, 1987: 130–131, fig. 7.
Ophionereis eurybrachiplax: Irimura, 1982: 72–73, fig. 44(non Ophionereis eurybrachiplax H. L. Clark, 1911:
173–175, fig. 78).
?Ophionereis eurybrachiplax: Matsumoto, 1918: 479–
480, fig. 2.

Material examined. Off Shijikijima Island;
65 m (G), 1 specimen (NSMT E-4972). Southsoutheast of Tsudo; 62–65 m (G), 1 specimen(NSMT E-4973).
Remarks. This species is close to
Ophionereis eurybrachiplax H. L. Clark, 1911,
but A. M. Clark (1953) described O. sasakii
based on a specimen from Hakodate. The present
specimens were determined to be O. sasakii by
the subovoidal shape of oral shield (Fig. 13C),
the presence of a second tentacle scale (Fig.
13D), and the presence of genital papillae thatare visible on the dorsal side of some arm bases(Fig. 13E–F) although often concealed by diskedge. If these two species are distinguishable, therecord of O. eurybrachiplax from Japanese waters by H. L. Clark (1911) seems doubtful. Irimura (1982) reported O. eurybrachiplax from Sagami Bay, central Japan. In his specimen (No. 321,
currently deposited in NSMT), which was re-examined, the shape of oral shield is not triangular(see Irimura 1982: fig. 44B), the second tentacle
scale is present, and the genital papillae was observed from the dorsal side though only one or
two at each arm base. Thus, the Sagami Bay
specimen must be conspecific with specimensfrom the Oki Islands. Some other specimens deposited in NSMT, collected from Sagami Bay, 70
m (NSMT E-1615, 1680) and from the Yellow
Sea (NSMT E-1950) are also conspecific to thepresent specimens. Original description of O. eurybrachiplax by H. L. Clark (1911) was based ona large specimen (disk diameter 24 mm). The
largest Japanese specimen examined (NSMT E1680), having a disk diameter of 16.7 mm, haswide dorsal arm plates like O. eurybrachiplax,

Ophiuroids from the Oki Islands, Japan

A

B

C

D

E

211

F

Fig. 13. Ophionereis sasakii. A–E: NSMT E-4972 (disk diameter 7.1 mm), F: NSMT E-4973 (disk diameter
10.5 mm) A: Whole body, aboral view. B: Whole body, oral view. C: Jaw. Scale 0.5 mm. D: Second tentacle
scales (arrows). Scale 0.5 mm. E: Genital papillae (arrows) viewed from aboral side. Scale 0.5 mm. F: Genital
papillae (arrows) viewed aboral-laterally. Scale 0.5 mm. A–F: Alcohol specimen. Scale divisions 1 mm
(A–B). Scales 0.5 mm (C–F).

but the shape of oral shield is not triangular, asecondary tentacle scale is present, and genital
papillae are partly visible from dorsal side. It’sdifficult to determine whether O. eurybrachiplax
from California and O. sasakii from Japan are
distinct species because of variation in visibilityof genital papillae from the dorsal side. It is necessary to evaluate the type specimens of O. eurybrachiplax to show the relationships between

these two species. However, for the present,
Japanese specimens should be identified as O.
sasakii. Such variation was also reported in Korean specimens (Yi & Irimura, 1987).
Geographic range. This species is reported
Japan and Korea. The Japan Sea, the Oki Islands(present study). Korea, the Yellow Sea (Yi & Irimura, 1987). Japan, Hakodate (type locality, A.
M. Clark, 1953), Sagami Bay (Irimura, 1982).

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Ophionereis variegata Duncan, 1879
[Japanese name: Sanmen-kumohitode]

Material examined. Off Onbejima Island;
67 m, 1 specimen (NSMT E-4977). East of Onbejima Island; 54 m, 2 specimens (NSMT E4975). Off Shijikijima Island; 36°9N, 133°14E,
20–30 m (G, R), 1 specimen (NSMT E-5001); 56
m, 1 specimen (NSMT E-4974). East of Shijikijima Island; 15 m (G), 1 specimen (NSMT E4976).
Remarks. This species is close to
Ophionereis porrecta Lyman, 1860, which is also
distributed in southern Japan (see A. M. Clark &Rowe, 1971). The specimens have fan-shaped
ventral arm plates (see A. M. Clark, 1953: fig. 7)
and one accessory lateral plate (see Matsumoto,
1917: fig. 93d) between the main lateral arm
plates, and O. variegata can be distinguishedfrom O. porrecta by these characteristics (A. M.
Clark, 1953).

A

D

B

Geographic range. This species is distributedin Japan and Australia. The Sea of Japan, Korean
Strait (type locality, Duncan, 1879), the Oki Islands (present study). Japan, Boso Peninsula (Irimura, 2000), the Goto Islands (Fujita, 1998), Kii(Irimura, 1981), Okinawa (Matsumoto, 1917 as
Ophionereis porrecta). Australia, Great Barrier
Reef (Rowe & Gates, 1995).
Ophionereis thryptica (Murakami, 1943)[New Japanese name: Mutsuude-amime-kumohitode](Fig. 14)

Material examined. Between Oomorijima Island and Shijikijima Island; 58 m, 2 specimens(NSMT E-4979). Off Onbejima Island; 52 m, 3
specimens (NSMT E-4980); 54 m, 2 specimens(NSMT E-4983); 54 m, 3 specimens (NSMT E4984); 67 m, 2 specimens (NSMT E-4990); 71 m,
1 specimen (NSMT E-4985). Off Shijikijima Island; 60 m (R), 7 specimens (NSMT E-4981); 60

CE
F

Fig. 14. Ophionereis thryptica. A: NSMT E-4981 (disk diameter 2.6 mm), B–E: NSMT E-4982 (disk diameter
3.0 mm), F: NSMT E-4988 (disk diameter 3.1 mm). A: Basal arm, aboral view. B: Supplemental dorsal arm
plate (right arrow) and secondary supplemental dorsal arm plate (left arrow). C: Disk, aboral view. D: Mouth
frame. E: Arm spines of 17th arm segment, distal view, oral side rightward. F: An ectoparasitic copepod. A,
F: Alcohol specimen, B–E: Dry specimen. Scales 0.5 mm (A, D–E), 0.1 mm (B), 1 mm (C, F).

Ophiuroids from the Oki Islands, Japanm (R), 1 specimen (NSMT E-4982); 65 m (G), 2
specimens (NSMT E-4986); 65 m (G), 30 specimens (NSMT E-4987); 65 m (G), 3 specimens(NSMT E-4988). Kamo Bay; 53 m, 1 specimen(NSMT E-4989). Tsutsuka Bay; Subtidal, 1 specimen (NSMT E-4978).
Remarks. The specimens agrees well withthe description by Murakami (1943a). There are
two six-armed Ophionereis species, O. hexactis
H. L. Clark, 1938 and O. sexradia Mortensen,
1936. The former has the middle arm spine
markedly elongated (A. M. Clark & Rowe, 1971:
fig. 56i) instead of subequal as in O. thryptica(Fig. 14E). Ophionereis sexradia has no small
secondary supplementary dorsal arm plate (Fig.
14B). Three specimens each have one unidentified ectoparasitic copepod (Fig. 14F).
Geographic range. This species had been
known from the Ogasawara Islands, Japan (Irimura & Tachikawa, 2003), Palau (type locality,
Murakami, 1943a as Ophiocrasis thryptica), Indonesia (Guille & Wolff, 1984), Madagascar(Cherbonnier & Guille 1978). This is the firstrecord from the Sea of Japan.
Family Ophiodermatidae
Ophiarachnella gorgonia (Müller &Troschel, 1842)[Japanese name: Toume-kumohitode]

Material examined.

Kanbi; subtidal, 1 speci-213

men (NSMT E-4952).
Geographic range. This species is distributedin the Sea of Japan and widely in Indo-west Pacific (see A. M. Clark & Rowe, 1971). Korea(Rho, 1979; Yi & Irimura, 1987). The Sea ofJapan, Sado (Irimura, 1979), Notojima Island(Fujita & Kohtsuka, 2003), the Oki Islands (present study). Pacific coast of Japan, Inubo-zaki
cape and more southern waters (see Irimura,
1982, 2000). The type locality is Mauritius(Müller & Troschel, 1842).
Ophiopsammus anchistus (H. L. Clark, 1911)[Japanese name: Menashi-kumohitode](Fig. 15)

Material examined. Off Onbejima Island;
67 m, 1 specimen (NSMT E-4955). Off Shijikijima Island; 60 m (R), 1 specimen (NSMT E4954); 70 m (R, S), 1 specimen (NSMT E-4953).
Remarks. This species was originally described as Pectinura anchista by H. L. Clark(1911) and was transferred to Ophiopsammus by
Vail & Rowe (1989). The present specimens(disk diameter 4.3–11.6 mm) have several granules between dorsal arm plates and lateral arm
plates, and the shape of dorsal arm plates ranges
from fan shape to rectangular shape in individuals of increasing size (Fig. 15A–C).
Geographic range. This species is distributedin Korea and Japan. Korea (Rho, 1979; Rho &Fig. 15. Ophiopsammus anchistus. Basal arm, aboral view. A: NSMT E-4953 (disk diameter 11.6 mm). B:
NSMT E-4954 (disk diameter 10.4 mm). C: NSMT E-4955 (disk diameter 4.4 mm). A–C: Alcohol specimen.
Scales 1 mm (A–C).

214

Toshihiko Fujita et al.

Shin, 1987). The Sea of Japan, Sado (Irimura,1979), Notojima Island (Fujita & Kohtsuka,
2003), the Oki Islands (present study). Pacificcoast of Japan, Boso Peninsula (Irimura, 2000),
Sagami Bay (Matsumoto, 1917; Irimura, 1982),
Tanabe Bay (Irimura, 1981), the Ogasawara Islands (Irimura & Tachikawa, 2003). Type locality
is off Suno-saki, off Goto Island, and Kagoshima
Bay, Japan (H. L. Clark, 1911).

Ophioconis acanthophora (Murakami, 1942)
new combination[Japanese name: Awahada-kumohitode](Fig. 16)
Ophiurodon acanthophora Murakami, 1942: 31–33, fig.
12.

Material examined. East of Onbejima Island;
54 m, 1 specimen (NSMT E-4956). South ofDougo Island (no detailed locality data); 1 speci-

Fig. 16. Ophioconis acanthophora. NSMT E-4956 (disk diameter 4.7 mm). A: Disk, aboral view. B: Disk
spinelet (arrow). C: Basal arm, aboral view. D: Mouth frame. E: Tentacle scales at arm base. F: Basal arm,
oral view. G: Arm spines near arm base, oral-lateral view. A–G: alcohol specimen. Scales 1 mm (A, C, G),
0.5 mm (B, D–F).

Ophiuroids from the Oki Islands, Japanmen (NSMT E-4957).
Remarks. The present specimens agree wellwith the description of Ophiurodon acanthophora Murakami, 1942, which was characterized by
non-serrated teeth, the presence of spinelets on
disk, and two tentacle scales (Murakami, 1942).
One specimen (NSMT E-4956, disk diameter 4.7mm) has only one spinelet on the disk (Fig. 16B),
there is no spinelet found on the disk of anotherspecimen (NSMT E-4957) though the dorsal disk
is so severely damaged. Tentacle scales are 4–5
in number for the first two tentacle pores and 2
for the following pores (Fig. 16E). The teeth havehyaline margin slightly serrated (Fig. 16D).
These characteristics suggest that this species is
at least congeneric, possibly conspecific, with
Ophiurodon cupidum (Koehler, 1905) reported
from southern Japan by Murakami (1943b).
The generic status of Ophiurodon Matsumoto,1915 is very doubtful (A. M. Clark, 1965; Baker& Devaney, 1981). A. M. Clark (1965) transferred Ophiurodon cupidum to Ophioconis andlikewise Ophiurodon acanthophora should be regarded as congeneric with Ophioconis cupida
Koehler, 1905.
Geographic range. This species had been reported only from the type locality, Izu, Japan(Murakami, 1942). This is the first record fromthe Sea of Japan.
Family Ophiolepididae
Ophioplocus japonicus H. L. Clark, 1911
[Japanese name: Nihon-kumohitode]

Material examined. Shijikijima Island; subtidal, 4 specimens (NSMT E-4996). Kanbi; subtidal, 2 specimens (NSMT E-4993). Kamo Bay;
subtidal, 2 specimens (NSMT E-4991); subtidal,
4 specimens (NSMT E-4995). Tsutsuka Bay;
subtidal, 5 specimens (NSMT E-4992). Inugu;
subtidal, 1 specimen (NSMT E-4994).
Remarks. This species is distributed in Korea(see Rho & Shin, 1987; Yi & Irimura, 1987) andJapan (see Irimura, 1982). It is very common
species especially in intertidal zone in the Sea ofJapan (see Irimura, 1982; Ishida et al., 2001; Fu-

215

jita & Kohtsuka, 2003).
Acknowledgments
We would like to express our sincere gratitude
to Drs. Akira Matsuno, Kohzoh Ohtsu and thestaff of Shimane University and Dr. Takeo Yamauchi of Hiroshima University for their generous support in collecting material and providing
their specimens and to Mr. Mitsufumi Torii andfishermen of Saigou Fishing Port for their help insampling. Dr. Gordon Hendler of the Natural
History Museum of Los Angeles County kindly
reviewed the manuscript and gave us many usefulsuggestions. The first author (TF) thanks also
Mme Elizabeth Lang of Musée Zoologique
Strasbourg and Dr. Helmut Sattman of Naturhistorisches Museum Wien for their kind help in
examining the specimens during the study of
Ludwig Döderlein’s collection supported by agrant-in-aid for International Scientific Research(Nos. 09041155 & 12575008) from the Japanese
Ministry of Education, Science, Sports, Culture
and Technology.
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