Bull. Natl. Mus. Nat. Sci., Ser. A, Suppl. 1, pp. 59–66, March 22, 2007

Lycenchelys ryukyuensis sp. nov. (Perciformes: Zoarcidae) from the
Okinawa Trough, Japan
Gento Shinohara1 and M. Eric Anderson2
1

Department of Zoology, National Museum of Nature and Science, 3–23–1 Hyakunin-cho,Shinjuku-ku, Tokyo 169–0073 Japan
E-mail: s-gento@kahaku.go.jp
2
SAIAB, Private Bag 1015, Grahamstown 6140, South Africa
E-mail: E.Anderson@ru.ac.za

Abstract A new zoarcid fish, Lycenchelys ryukyuensis, is described on the basis of 5 specimens(113.2–169.8 mm in standard length) collected from 991–1533 m in the Okinawa Trough near the
Ryukyu Islands, Japan. Although this species resembles Lycenchelys remissaria in having similar
meristic counts of fin rays, absence of interorbital and occipital pores, and body coloration, it differs from the latter in having 61 (rarely 62) suborbital pores, 44 preoperculomandibularpores, a single ventral lateral line, and the pelvic fin located at about mid-distance between the eye
and the pectoral fin base. These specimens are the southernmost records for Lycenchelys in thewestern North Pacific.
Key words : Zoarcidae, Lycenchelys ryukyuensis, Lycenchelys remissaria, Okinawa Trough,Western North Pacific.

The zoarcid genus Lycenchelys includes 59
deep-sea species (about 1/4 the total numbers ofvalid species in the family) and almost all species
inhabit soft bottoms from the upper bathyal toabyssal zones (Anderson, 1995; Anderson andFedorov, 2004) with some polar species from thecontinental shelves (see Møller and Jørgensen,
2000). The genus is distributed mainly in thenorthern hemisphere, but species also occur in
the South Pacific, southwestern Atlantic, subantarctic, and Antarctic waters. The genus ismainly characterized by the followings: 6–10
suborbital bones with 6–10 pores from the canal;
first dorsal fin ray pterygiophore associated withvertebrae 2–21, with 0–16 free pterygiophores;
weak palatopterygoid series; weak oral valve;presence of pseudobranch, usually pelvic fins,usually both vomerine and palatine teeth; scales,
pyloric caeca, and lateral line(s) present; and vertebrae 19306611885144 (Anderson,
1994; Shinohara and Matsuura, 1998). Ten Lycenchelys species are known from Japanese waters, and the southern limit of the distribution in

the western North Pacific is northern Honshu Island, Japan (Anderson and Imamura, 2002; Hatooka, 2002; Imamura et al., 2004).
In 2002, 5 specimens of Lycenchelys were collected during beam trawl surveys made by the
National Science Museum, Tokyo, and Ocean
Research Institute, the University of Tokyo fromthe Okinawa Trough near the Ryukyu Islands inthe East China Sea. This species is most similar
to Lycenchelys remissaria Fedorov, 1995 among
the Japanese Lycenchelys in lacking interorbital
and occipital pores. It differs from the latter inhaving the pelvic fin in a normal position, and we
describe it as a new species to science.
Methods for taking counts and measurements
follow Peden and Anderson (1978) and Anderson(1982). Vertebrae and other osteological elements were examined from radiographs. Terminology of head pores follows Anderson (1994).
Standard length (SL) and head length (HL) areused throughout.
The specimens examined are deposited in the
Department of Zoology, National Museum of

G. Shinohara and M. E. Anderson

Fig. 1. Lycenchelys ryukyuensis sp. nov., Okinawa Trough, East China Sea, Japan, holotype, NSMT-P 63965, male, 159.8 mm SL. Fresh specimen(top), line drawings (bottom)

60

Nature and Science (formerly National Science
Museum), Tokyo (NSMT), Laboratory of Marine
Biology, Kochi University (BSKU), Laboratoryof Marine Biology and Biodiversity, Hokkaido
University (HUMZ), and Zoological Institute,
Russian Academy of Sciences, St. Petersburg(ZIN).
Lycenchelys ryukyuensis sp. nov.
(New Japanese name: Ryukyu-hebigenge)(Figs. 1–2, Table 1)
Holotype. NSMT-P 63965, male, 159.8 mm SL, Okinawa Trough, East China Sea, 26°11.34N, 124°54.27E–
26°12.65°N, 124°55.47E, 1493–1533 m, 26 April 2002,
R/V Tansei-maru, beam trawl.
Paratypes. BSKU 57850–57851, 2 females, 113.2–
151.3 mm SL, Okinawa Trough, 26°15.10N, 125°17.22
E–26°13.85N, 125°18.43E, 991–995 m, 26 April 2002,
R/V Tansei-maru, beam trawl; NSMT-P 63966–63967, 2
females, 118.6–143.2 mm SL, collection data as same as
holotype.

Diagnosis. Vertebrae
222496102
118124; interorbital pore and occipital poresabsent; postorbital pores 3 (rarely 4); suborbital
pores 61 (rarely 62); preoperculomandibularpores 44; lateral line ventral, complete; dorsal
fin rays 110–114; anal fin rays 98–104; pelvic finbase about mid-distance between the eye andpectoral fin base.
Description. Counts and proportional measurements are shown in Table 1. Conditions ofholotype given first, followed by those ofparatypes in parentheses. Body long, its crosssection almost round at nape, elliptical at bothabdominal and caudal regions. Head short, its
dorsal profile gently convex. Scales absent onhead, nape, thorax, pectoral fin, and pelvic fin.
Scales present on body, abdomen, tail, on dorsal
and anal fins, extending to about half of theirheight. Eye ovoid, not entering dorsal profile ofhead. Interorbital space convex. Gill slit extending ventrally below the level of lower pectoral
base. Dorsal fin origin above middle of pectoral
fin. Opercular flap at upper margin of gill slit developed. Pectoral fin origin about level of body
midline. Posterior margin of pectoral fin slightly

New Eelpout of Lycenchelys from Okinawa Troughpointed, ray tips almost evenly rounded dorsally
but well exserted ventrally (Fig. 2). Pelvic fin
small and thin.
Mouth subterminal, upper jaw extending beyond anterior margin of pupil. Nostril tube small,
not reaching upper lip when depressed forward.
Snout rounded. Labial lobe of lower jaw weak.
Jaw teeth conical, sharp; upper jaw with 3 (2) anterior tooth rows, lower jaw with 3 (2–3) irregular rows; teeth in outer row in both jaws slender,
large. Two (2–4) vomerine teeth in small patch;
palatine teeth in a single row, 2 in left/1 in right(1–2/1).
Head pores well developed (Fig. 2). Two nasal
pores, anterior one set just in front of nostril
tube, the other situated dorsoposteriorly. Seven(7, rarely 8) suborbital pores which associatedwith 8 suborbital bones (NSMT-P 63966); 6
pores arising from ventral ramus of bone chain
under eye and 1 (1–2) from ascending ramus behind eye just posteroventral to first postorbital
pore [formula 61 (61, rarely 62 in left of
NSMT-P 63966)]. Three (3, rarely 4 in right of
BSKU 57851) postorbital pores. Eight preoperculomandibular pores (4 mandibular pores4
preopercular pores): 4 arising from dentary, one
from anguloarticular and 3 from preopercle. Interorbital and occipital pores absent. Lateral line
ventral, complete; of reduced superficial neuromasts from above posterior tip of opercle to near
tip of tail.
Dorsal fin origin associated with 7th (7th or8th) vertebra; no supraneurals. Anal fin origin associated with ultimate precaudal vertebra, with 3(2–4) pterygiophores inserted anterior to haemal
spine of first caudal vertebra; last anal pterygiophore associated with 4th (3rd or 4th) preuralvertebra, with 2 (2–3) anal rays articulating withhaemal spine. Caudal fin with 2 epural, 4 (4–5)

61

upper and 3 lower hypural rays.
Gill rakers degenerated, triangular. Pseudobranch filaments short. Pyloric caeca 2, nublike.
Color when fresh (in alcohol). Head, pectoralfin, and pelvic fin black (brown). Margin of vertical fins dark brown (brown). Body and tail grayish (light brown).
Etymology. The specific name is derived
from the collection locality.
Distribution. Known only from the Okinawa
Trough near Ryukyu Islands, East China Sea in
depths of 991–1533 m.
Comparison. In having no interorbital
pores, no occipital pores, 3 (rarely 4) postorbital
pores, pelvic fins, a single ventral lateral line, Lycenchelys ryukyuensis sp. nov. is similar to Lycenchelys aratrirostris Andriashev and Permitin,
1968 (Antarctica), Lycenchelys bellingshausenni
Andriashev and Permitin, 1968 (Antarctica),

Fig. 2. Head and anterior body of Lycenchelys
ryukyuensis sp. nov., holotype, NSMT-P
63965, male, 159.8 mm SL. Shaded area indicates squamation

Fig. 3. Alcohol preserved specimen of Lycenchelys remissaria, off Fukushima Prefecture, Japan, holotype, ZIN
50586, photographed by G. Shinohara (November 1999)

62

G. Shinohara and M. E. Anderson
Table 1. Comparison of counts and proportional measurements between Lycenchelys ryukyuensis sp. nov. andL. remissaria
Lycenchelys ryukyuensis sp. nov.

Counts
Dorsal fin rays
Anal fin rays
Pectoral fin rays
Pelvic fin rays
Caudal fin rays
Gill rakers
Vertebrae
Nasal pores
Suborbital pores
Postorbital poresInterorbital pores
Preopercular pores
Mandibular pores
Occipital pores% SL
Head length
Body height at pectoral fin base
Body height at anal fin origin
Predorsal length
Preanal length
Pectoral fin length
Gill slit length% HL
Upper jaw length
Pectoral fin length
Snout length
Eye diameter
Gill slit lengthInterorbital width
Pelvic fin length
Pectoral base/length ratio

Lycenchelys remissaria

Holotype

Paratypes(n4)

Holotype

Non-types(n4)Shinohara et al.
(1996)
(n7)

114
104
15
3
9
12
124
2
7
3
0
4
4
0

110–114
98–102
13–15
3
9–10
11–13
118–123
2
7–8
3–4
0
4
4
0

118a
104a
17
3
11a
13a
128a
2
8
3
0
4
5
0

115–119
102–107
15–17
3
9–10
12–16
123–127
2
8
3–4
0
4
4–5
0

–
–
16–17
–
–
–
122–130
–
8
3
0
4
5
0

14.3
5.1
5.8
20.2
31.4
8.8
4.8

14.3–14.7
5.5–5.8
4.7–5.7
18.7–19.7
30.9–32.3
8.3–10.8
4.8–5.6

14.1
6.5
7.2
17.9
29.8
8.3
5.0

14.7–16.2
6.0–7.5
5.3–8.4
19.5–20.9
31.7–34.3
8.8–11.3
5.2–5.7

–
–
–
–
–
–
–

38.6
61.8
29.0
18.4
33.3
3.5
14.9
0.34

30.0–39.9
56.5–73.5
24.7–29.6
18.4–20.6
33.3–38.0
3.0–4.4
16.1–18.8
0.23–0.31

37.2
58.9
27.3
24.8
35.5
4.3
19.2
–

29.6–37.4
59.4–69.7
23.5–30.3
20.9–23.5
32.5–38.9
4.9–6.1
12.2–18.3
0.31–0.39

36.9–41.7––
20.3–24.9–
3.2–3.7–––, no data.
aData from Fedorov (1995).

Lycenchelys folletti Anderson, 1995 (Gulf ofCalifornia), Lycenchelys lonchoura Anderson,
1995 (eastern South Pacific), and Lycenchelys
maoriensis Andriashev and Fedorov, 1986 (western South Pacific). The new species is different
from L. aratrirostris, L. bellingshauseni, L. folletti, and L. lonchoura in having higher counts ofvertebrae (118–124 in L. ryukyuensis vs. 101–110 in L. aratrirostris, 102–110 in L. bellingshauseni, 102–109 in L. folletti, and 107 in L.
lonchoura), dorsal fin rays (110–114 vs. 97–107,
94–103, 97–104, and 103), and anal fin rays

(98–104 vs. 79–89, 82–91, 83–92, and 86) (seeAnderson, 1990, 1995). Lycenchelys ryukyuensis
is further distinguished from L. aratrirostris in
having entirely dark pectoral fins (vs. only margin of pectoral fin dark in L. aratrirostris), from
L. bellingishauseni in having scales on posterior
parts of dorsal and anal fins (no scales on bothfins in L. bellingishauseni), from L. folletti inhaving both vomerine and palatine teeth (vs. noteeth on vomer and palatine in L. folletti), andfrom L. lonchoura in having 13–15 pectoral fin
rays and 2 pyloric caeca (18 and 0 in L. lon-

New Eelpout of Lycenchelys from Okinawa Troughchoura). The new species differs from L.
maoriensis in having 7–8 suborbital pores and noblack or brown variegations all along body andtail in males (vs. 6 and the variegations present in
the latter).
Except the position of pelvic fins and the lateral line configuration, L. ryukyuensis appears
close to L. remissaria, also known from Japan(Pacific side of Tohoku, northern Honshu Island)in having similar body color and meristic counts(122–130 vertebrae, 115–119 dorsal fin rays,
102–107 anal fin rays, 15–17 pectoral fin rays in
the latter; Table 1). The new species is distinguished from L. remissaria by having the pelvic
fin origin mid-distance between the eye and pectoral fin base (vs. nearer the eye), a single ventral
lateral line (vs. lateral line double, with mediolateral and ventral branches), 4 preopercular pores(vs. 5), and normally 7 suborbital pores (vs. 8).
In the western North Pacific, the southern limitof the distribution of Lycenchelys spp. has beennorthern Honshu Island (ca. 36°30N, 141°10E,
Lycenchelys squamosa Toyoshima, 1983, Lycenchelys makushok Fedorov, 1993, and L. remis-

63

saria; see Shinohara et al., 1996). The specimensof L. ryukyuensis extend the range of the genus
by about 1800 km.
Taxonomic notes on Lycenchelys remissaria.
The holotype of Lycenchelys remissaria has
scales in front of the dorsal fin origin and along
the dorsal fin base (Fedorov, 1995: fig. 2). Lycenchelys ryukyuensis, however, lacks them inthese parts. The 2 smallest specimens of L. remissaria (NSMT-P 49210, 87.6–102.6 mm SL)
have a naked anterior dorsal fin base which issimilar to the condition in L. ryukyuensis. Thus
the squamation of both the nape and dorsal finbase show an ontogenetic change in L. remissaria.
Key to Japanese Lychenchelys species.
Here we provide a key to the known species of
Lycenchelys found in Japanese waters. All occur
along the Okhotsk and Pacific coasts of Hokkaido and the Pacific of northern Honshu except the
new L. ryukyuensis. Other species from thesouthern Okhotsk and Kuril Islands may eventually be found in the area. We have tried to avoid
characters observable only by radiography, thus
the key requires specimens in good condition.

Key to Late Juveniles and Adults of Lycenchelys in Japanese Waters
1a. Interorbital and occipital pores absent                                              2
1b. Interorbital pore present; occipital pores present or absent                              3
2a. Pelvic fin base shifted forward, its origin at about a vertical through middle of preopercle; preoperculomandibular pores 9; suborbital pores 71; 2 lateral lines              L. remissaria
2b. Pelvic fin base at about a vertical through posterior margin of preopercle, mid-distance between
eye and pectoral base; preoperculomandibular pores 8; suborbital pores 61 or 62, 1 lateral line                                                                 L. ryukyuensis
3a. Preoperculomandibular pores 7–9; body depth >5% total length                          4
3b. Preoperculomandibular pores 10; body depth 3–4% total length                 L. makushok
4a. Pelvic fins present; 1 lateral line                                                   5
4b. Pelvic fins absent; 2 lateral lines                                        L. tohokuensis
5a. Suborbital pores 6                                                              6
5b. Suborbital pores 8–11                                                           7

64

G. Shinohara and M. E. Anderson

6a. Color more or less uniformly orange; preoperculomandibular pores 7; pectoral fin rays 13–16
                                                                  L. aurantiaca
6b. Color more or less uniformly dark brown; preoperculomandibular pores 8; pectoral fin rays
17–19                                                               L. squamosa
7a. Body dark brown, with 6–10 wide, white, vertical bands extending from the dorsal fin to midbody; pectoral fin rays 18–19                                         L. albomaculata
7b. Body and dorsal fin without white bands; pectoral fin rays 13–17                         8
8a. Body, tail and dorsal fin with many distinct dark blotches, sometimes on anal fin; total vertebrae
138–147                                                              L. maculata
8b. Body uniformly dark brown or black, no blotches; total vertebrae 118–138                  9
9a. Preoperculomandibular pores 8; postorbital pores 3–4; total vertebrae 122–138             10
9b. Preoperculomandibular pores 9; postorbital pores 5; total vertebrae 118–122                                                                               L. melanostomias
10a. Pelvic fin rays 3; nostril tube not reaching upper lip when depressed forward; total vertebrae
132–138                                                          L. hippopotamus
10b. Pelvic fin rays 2; nostril tube reaching upper lip or overhanging it; total vertebrae 122–131                                                                            L. rassi
Comparative materials. Lycenchelys albomaculata:

NSMT-P 69787(1), female, 203.0 mm SL, 44°9.565N,

NSMT-P 48684(1), 190.5 mm SL, male, 36–57.5N, 141°

144°52.673E–44°9.243N, 144°51.371E (Okhotsk Sea

41.46E–36°50.9N, 141°44.06E (Pacific off Fukushima

off Hokkaido, Japan), 491–495 m depths, 13 Sep. 2004,

Prefecture, Japan), 777–905 m depths, 5 Nov. 1995, R/V

R/V Tansei-maru, coll. by G. Shinohara and K. Suetsugu;

Watakata-maru, coll. by G. Shinohara; NSMT-P 49610

HUMZ 126197(1), 126199(1), female and male, 175.5–(2), 163.0–184.0 mm SL, males, 36°57.02N, 141°46.09

177.1 mm SL, 44°34.1N, 144°59.7E (Sea of Okhotsk),

E–36°56.22N, 141°45.35E (Pacific off Fukushima Pre-

1301 m depth, 26 May 1993; HUMZ 126212(1), female,fecture), 800–802 m depths, 16 Apr. 1996, R/V Tanshu-

187. mm SL, 44°21.2N, 144°51.8E (Sea of Okhotsk),

maru, coll. by G. Shinohara; NSMT-P 53659(1), 175.0

1068 m depth, 28 May 1993. Lycenchelys makushok:

mm SL, male, 36°57.08N, 141°50.8E–36°55.13N, 141°

NSMT-P 73403(1), 139.0 mm SL, male, 36°59.99N,

57.33E (Pacific off Fukushima Prefecture), 1001–1023

141°46.00E–36°58.97N,m, 16 Apr. 1996), R/V Tanshu-maru, coll. by G. Shino-

Fukushima Prefecture), 17 Apr. 1997, R/V Tanshu-maru,

hara; NSMT-P 61162(2), 186.0–190.0 mm SL, females,coll. by G. Shinohara; NSMT-P 61163(4), 136.0–149.0

37°1.36N, 141°38.96E–37°0.03N, 141°38.03E (Pacificmm SL, females, 37°1.36N, 141°38.96E–37°0.03N,

off Fukushima Prefecture), 500–501 m depths, 17 Apr.

141°38.03E (Pacific off Fukushima Prefecture), 500–

141°44.65E(Pacific

off1997, R/V Tanshu-maru, coll. by G. Shinohara. Ly-

501 m depths, 17 Apr. 1997, R/V Tanshu-maru, coll. by

cenchelys aurantiaca: NSMT-P 73401(4), 109.0–128.0

G. Shinohara; NSMT-P 73404(1), 90.0 mm SL, female,

mm SL, 2 females and 2 males, 37°54.69N, 141°58.44

36°26.45N, 141°12.13E–36°31.05N, 141°15.50E (Pa-

E–37°53.81N, 141°56.94E (Pacific off Fukushima Pre-cific off Ibaraki Prefecture, Japan), 785–794 m depths, 10

fecture), 21 Apr. 1997, R/V Tanshu-maru, coll. by G.

Nov. 1995, R/V Wakataka-maru, coll. by G. Shinohara.

Shinohara; NSMT-P 73402(1), 124.0 mm SL, male, 36°

Lycenchelys

59.99N, 141°46.00E–36°58.97N, 141°44.65E (Pacific126254(1), 126258(1), 126259(1), 149.2–216.4 mm SL, 3

off Fukushima Prefecture), 17 Apr. 1997, R/V Tanshu-

females and 2 males, 44°40.7N, 144°24.9E (Sea of

maru, coll. by G. Shinohara. Lycenchelys hippopotamus:

Okhotsk off Hokkaido, Japan), 1336 m depth, 8 Junerassi:

HUMZ

126251(1),126253(1),New Eelpout of Lycenchelys from Okinawa Trough1993. Lycenchelys remissaria: ZIN 50586(holotype),
male, 36°544N, 141°557E, 1020 m depth depth, 4 Feb.
1981, R/V Mys Dal’niy; NSMT-P 47278(1), female,262.1 mm SL, 38°10.5N, 143°7.6E–38°10.6N, 143°7.8
E (Pacific off Sendai, Miyagi Prefecture, Japan), 2021–

65

Kimura (Mie University) and 2 anonymous reviews for valuable comments. This study was partly supported by a research project entitled “Deepsea Fauna and Pollutants in Nansei Islands” (National Science Museum, Tokyo, 2001–2004).

2034 m depths, 20 May 1995, R/V Wakataka-maru, coll.
by T. Fujita; NSMT-P 48697(1), 277.7 mm SL, 36°28.43

Literature Cited

N, 141°24.18E–36°27.27N, 141°32.2E (Pacific off
Ibaraki Prefecture, Japan), 1464–1537 m depths, 10 Nov.
1995, R/V Wakataka-maru, coll. by G. Shinohara; NSMTP 49210(2), 87.6–102.6 mm SL, 36°45.41N, 141°43.16
E–36°45.06N, 141°42.78E (Pacific off Ibaraki Prefecture), 1408 m depth, 19 Apr. 1996, R/V Tanshu-maru,
coll. by G. Shinohara. Lycenchelys squamosa: NSMT-P
49609(1), 204.0 mm SL, male, 36°57.02N, 141°46.09
E–36°56.22N, 141°45.35E (Pacific off Fukushima Prefecture), 800–802 m depths, 16 Apr. 1996, R/V Tanshumaru, coll. by G. Shinohara; NSMT-P 49043(1), 218.0mm SL, female, 36°30.31N, 141°8.51E–36°26.04N,
141°7.12E (Pacific off Ibaraki Prefecture), 588–603 m
depths, 9 Nov. 1995, R/V Watakata-maru, coll. by G. Shinohara; NSMT-P 65692(1), 221.0 mm SL, female,
36°32.45N, 141°7.82E–36°29.09N, 141°4.44E (Pacificoff Ibaraki Prefecture), 748–789 m depths, 25 Oct. 2002,
R/V Watakata-maru, coll. by G. Shinohara; NSMT-P
49917(1), 195 mm SL, male, 37°29.3N, 142°15.6E–
37°28.9N, 142°13.8E (Pacific off Fukushima Prefecture), 810–823 m depths, 15 Apr. 1996, R/V Tanshumaru, coll. by G. Shinohara; NSMT-P 48696(1), 225 mm
SL, male, 36°57.5N, 141°41.46E–36°50.9N, 141°44.06E (Pacific off Fukushima Prefecture), 777–905
m depths, 5 Nov. 1995, R/V Watakata-maru, coll. by G.
Shinohara.

Acknowledgments
We are grateful to Y. Machida (BSKU), K.
Suetsugu (Ministry of the Environment), and T.
Kubodera (NSMT) for help in collecting specimens on the vessel, and H. Endo (BSKU) and H.
Imamura (Hokkaido University Museum) forloan of specimens. GS expresses sincere thanks
to A. Balushkin, V. V. Fedorov, and M. V.
Nazarkin (ZIN) for their help in examining theholotype of Lycenchelys remissaria during his
staying in St. Petersburg in 1999. Thanks go to S.

Anderson, M. E. 1982. Revision of the fish genera Gymnelus Reinhardt and Gymnelopsis Soldatov (Zoarcidae),with two new species and comparative osteology ofGymnelus viridis. National Museum of Natural Sciences, Publications in Zoology, (17): i–iv1–76.
Anderson, M. E. 1990. Zoarcidae. Pages 257–276 in O.
Gon and P. C. Heemstra, eds. Fishes of the Southern
Ocean. J. L. B. Smith Institute of Ichthyology, Grahamstown.
Anderson, M. E. 1994. Systematics and osteology of the
Zoarcidae (Teleostei:Perciformes). Ichthyological Bulletin of the J. L. B. Smith Institute of Ichthyology, 60:
1–120.
Anderson, M. E. 1995. The eelpout genera Lycenchelys
Gill and Taranetzella Andriashev (Teleostei: Zoarcidae)
in the eastern Pacific, with descriptions of nine new
species. Proceedings of the California Academy of Sciences, 49: 55–113.
Anderson, M. E. and V. V. Fedorov. 2004. Family Zoarcidae Swainson 1839. California Academy of Sciences
Annotated Checklists of Fishes, 34: 1–58.
Anderson, M. E. and H. Imamura. 2002. A new species of
Lycenchelys (Perciformes: Zoarcidae) from Pacific coastof northern Japan. Ichthyological Research, 49: 355–357.
Andriashev, A. P. and V. V. Fedorov. 1986. First discoveryof Zoarcidae in New Zealand waters. Voprosy Ikhtiologii, 26: 24–32. (In Russian.)
Andriashev, A. P. and Y. Y. Permitin. 1968. Two new
species of eelpouts of the genus Lycenchelys Gill
[Pisces, Zoarcidae] from bathypelagic depths in the
Scotia Sea (Antarctica). Voprosy Ikhtiologii, 51: 611–
618. (In Russian.)
Fedorov, V. V. 1993. Lycenchelys makushok sp.nova (Perciformes, Zoarcidae) from bathyal depths of the KurilKamchatka trench. Voprosy Ikhtiologii, 33: 133–136.
(In Russian.)
Fedorov, V. V. 1995. Lycenchelys remissaria sp.nova (Perciformes:Zoarcidae) from the bathyal region of the Pacific Ocean shores of Japan. Voprosy Ikhtiologii, 35:
130–133. (In Russian.)
Hatooka, K. 2002. Zoarcidae. Pages 1028–1044,
1581–1583 in T. Nakabo, ed. Fishes of Japan with Pictorial Keys to the Species, English edition. Tokai University Press, Tokyo.
Imamura, H., Y. Machida and S. Ohta. 2004. Lycenchelys

66

G. Shinohara and M. E. Anderson

melanostomias Toyoshima, 1983, a senior synonym of
L. brevimaxillaris Toyoshima, 1985 (Teleostei: Zoarcidae). Ichthyological Research, 51: 84–88.
Møller, P. R. and O. A. Jørgensen. 2000. Distribution andabundance of eelpouts (Pisces, Zoarcidae) off West
Greenland. Sarsia, 85: 23–48.
Peden, A. E. and M. E. Anderson. 1978. A systematic review of the fish genus Lycodapus (Zoarcidae) with description of two new species. Canadian Journal of Zoology, 56: 1925–1961.
Shinohara, G. and K. Matsuura. 1998. A new zoarcid, Lycenchelys aurantiaca, from the Pacific coast off northern Japan (Teleostei: Perciformes). Ichthyological Research, 45: 151–155.

Shinohara, G., H. Endo and K. Matsuura. 1996. Deepwater fishes collected from the Pacific coast of northern
Honshu, Japan. Memoier of the National Science Museum, Tokyo, (29): 153–185.
Toyoshima, M. 1983. Lycenchelys squamosus Toyoshima,sp. nov. Pages 144–147 in K. Amaoka, K. Nakaya, H.
Araya, and T. Yasui, eds. Fishes from the North-easternsea of Japan and the Okhotsk Sea off Hokkaido, Japan
Fisheries Resource Conservation Association, Tokyo.
Manuscript received 25 January 2006; revised 10 January2007; accepted 12 January 2007.
Associate editor: S. Kimura.