H. Motomura and K. Matsuura (eds) Fishes of Yaku-shima Island
National Museum of Nature and Science, Tokyo, 10 March 2010

New records of a triplefin, Enneapterygius leucopunctatus,
from southern Japan (Perciformes: Tripterygiidae)
Hiromitsu Endo1*, Eri Katayama1, Mina Miyake1,2 and Kimi Watase1,3
1

Laboratory of Marine Biology, Faculty of Science, Kochi University, 2-5-1 Akebono-cho, Kochi 780-8520, Japan
Minami-hanahata, Adachi, Tokyo 121-0062, Japan3
Ikeshinden High School, 2097-1 Ikeshinden, Omaezaki, Shizuoka 437-1612, Japan*
Corresponding author: e-mail: endoh@kochi-u.ac.jp
2

Abstract Enneapterygius leucopunctatus Shen, 1994 (Tripterygiidae), known only from Taiwan,
is recorded for the first time from Yaku-shima Island, Kagoshima Prefecture, and Tosa Bay, Kochi
Prefecture, southern Japan. Seventeen Japanese specimens (25.2–36.1 mm in standard length) wereidentified as E. leucopunctatus based on the following combination of characters: second dorsalfin spines 12–13; third dorsal-fin soft rays 9–11; anal-fin soft rays 17–19; mandibular pore formulamostly 4+1+4; pored and notched lateral-line scales 15–18+16–19; simple dermal flap of anterior
nostril; 2 white oblique bands on shoulder girdle; 8 dark saddle markings and 5–6 small ring-likewhite markings on lateral body; inconspicuous sexual dichromatism.
Key words: Tripterygiidae, Enneapterygius leucopunctatus, Yaku-shima Island, Tosa Bay, Japan.

Introduction
The blennioid family Tripterygiidae is composed of 32 genera and 164 species, inhabitingcoral reefs and rocky substratum in tropical tocold seas (Fricke, 2009). The genus Enneapterygius Rüppell, 1835, the largest group in the family, is diagnosed by having a discontinuous lateralline with 6–22 pored scales anteriorly and 13–27
notched scales posteriorly; the following combination of fin rays—dorsal: III, IX–XVI, 7–12; anal: I,
15–22; pelvic: I, 2; and a body with ctenoid scales
except for naked head, operculum, abdomen, andpectoral-fin base (the last two areas covered withcycloid scales in some species) (Fricke, 1997;
Holleman, 2005; Chiang and Chen, 2008). In thewestern and central Pacific, 46 species of the genus are known (Meguro and Motomura, 2010). Of
4 species described by Shen (1994), Chiang andChen (2008) regarded E. rubicauda Shen, 1994 asa junior synonym of E. flavoccipitis Shen, 1994,
and E. hsiojenae Shen, 1994 and E. leucopunctatus Shen, 1994 as valid species, not junior synonyms of E. vexillarius Fowler, 1946 as asserted

by Fricke (1997). Around Japan, 15 species havebeen recorded (e.g., Shimojyo and Hayashi, 2000;
Hayashi, 2002; Yoshigou and Yoshino, 2002;
Shibukawa et al., 2004; Motomura et al., 2005).
During surveys of fish faunas along the coastsof Tosa Bay and Yaku-shima Island, many specimens of Enneapterygius were collected fromrocky shores and tide-pools to 4 m depth. Some ofthese were identified as E. leucopunctatus basedon counts of the fin rays and lateral-line scales,
the mandibular pore pattern, and distinctive markings and coloration. Herein, we report this species
from Japan for the first time.
Materials and methods
The specimens examined are deposited in
the following institutions: Academy of Natural
Sciences, Philadelphia, USA (ANSP); Laboratory of Marine Biology, Kochi University, Japan(BSKU); and Kagoshima University Museum,Japan (KAUM). Counts and measurements follow Hubbs and Lagler (1958). Measurements
were made to the nearest 0.1 mm with dividers
9H. Endo et al.

Fig. 1. Enneapterygius leucopunctatus from Yaku-shima Island, Kagoshima Prefecture (A, C: KAUM–I. 21838, male, 25.3mm SL; B, D: KAUM–I. 21837, female, 26.4 mm SL). Fresh (A, B) and preserved (C, D) conditions.

and digital calipers under a binocular microscope.
Terminology of the head lateral-line canals andmandibular pore formula follow Fricke (1997).
Standard length and head length are abbreviatedas SL and HL respectively. Fin rays and vertebrae
were counted from radiographs. Observationof dentition, cephalic sensory tubes and pores,
scales, and genital papillae were made using cyanine solution. To determine the sex, the gonads ofeach specimen were observed by partial dissection
on the right side of the abdomen.

10

Enneapterygius leucopunctatus Shen, 1994
[New Japanese name: Hakuten-hebigimpo](Figs. 1, 2; Table 1)
Enneapterygius sp. 3: Shen, 1993: 491, 777, pls.
166-1, 166-2.
Enneapterygius leucopunctatus Shen, 1994: 12,
fig. 8 (type locality: Taiwan); Chiang andChen, 2008: 196, fig. 4i.
Enneapterygius vexillarius (not of Fowler, 1946):
Fricke, 1997: 355 (in part); Randall and Lim,
2000: 632 (list; in part).

Enneapterygius leucopunctatus from Japan

Materials examined. 17 specimens. Tosa
Bay, Kochi (15 specimens): BSKU 55655, female, 35.6 mm SL, Shirano-hana, Tosa City, 4
Sept. 2001; BSKU 56266, female, 33.7 mm SL,
Shirano-hana, Yokonami Peninsula, Tosa City, 2
Nov. 2001; BSKU 58044, female, 34.6 mm SL,
Shirano-hana, Yokonami Peninsula, Tosa City, 23
May 2002; BSKU 65433, female, 28.9 mm SL,
Karyougo, Nahari Town, 17 July 2003; BSKU
69833, female, 34.2 mm SL, Shioya Beach, Tei,

Konan City, 9 Apr. 2004; BSKU 72857, female,29.7 mm SL, Shirano-hana, Tosa City, 18 Sept.
2004; BSKU 73022, female, 28.8 mm SL, Ijiri,Tosa City, 5 June 2004; BSKU 73108, female,26.2 mm SL, BSKU 73109, female, 26.6 mm SL,
Shioya Beach, Tei, Konan City, 19 May 2004;
BSKU 73162, female, 29.2 mm SL, BSKU 73163,
female, 36.1 mm SL, Gyoudo-misaki, Muroto
City, 26 May 2004; BSKU 73296, female, 30.1mm SL, BSKU 73307, male, 27.4 mm SL, BSKU

Table1. Counts and measurements of Enneapterygius leucopunctatus from Japan and Taiwan. Mode of counts and mean of
proportions in parentheses.
Locality
SL (mm)Second dorsal-fin rays
Third dorsal-fin rays
Anal-fin soft rays
Pectoral-fin rays
Principal caudal-fin rays
Pored latera-line scales
Notched lateral-line scales
Transverse scale rows
Madibular pore formula
In SL
Head length
Body depth
Predorsal length
Dorsal-fin baseAnal-fin baseCaudal-peduncle length
In HL
Snout length
Orbit diameter
Postorbital length
Upper-jaw length
In % of SL
Head length
Body depth
Body width
Predorsal length
Dorsal-fin baseAnal-fin baseCaudal-peduncle length
Caudal-peduncle depth
1st spine length of 1st dorsal fin
1st spine length of 2nd dorsal fin
1st ray length of 3rd dorsal fin
Pectoral-fin length
2nd ray length of pelvic fin
Anal-fin spine length
1st ray length of anal fin
Caudal-fin length
In % of HL
Snout length
Orbit diameter
Postorbital length
Upper-jaw length

This study
Yaku-shima Island
Tosa Bayn=2
n = 1525.2–26.4
26.2−36.1
XIII
XII–XIII10
9–11(10)18–19
17−19 (18)
3+6+7
3–5+3–7+6−7(3+6+7)ii+9–10+ii
ii+9+ii
15–17
16−18 (18)19
16−18 (17)
2.5–3.5+1+6.5 2.5−3+1+5.5−6.5(2.5+1+5.5)
4+1+4
4–5+1+4–6(4+1+4)

Shen (1994)
Taiwann = 9 (incl. 3 types)
26.6–34.8
XIII–XIV (XIII)
9–10
18–20
2–5+5−8+6–8(3+6+7)–
17–19 (18)
17–19−

Chiang and Chen (2008)
Taiwann=6
23.7–29.1
XIII
9–11
18
–

4+1+4

4+1+4

–
17−18
17
−

3.2−3.3
4.5−4.6
3.8−3.9
1.5
2.3
8.4−8.8

3.1−3.6 (3.3)
4.2−5.0 (4.6)
3.7−4.1 (3.8)
1.4−1.5 (1.5)
2.2−2.5 (2.4)
7.5−8.7 (7.9)

3.2–3.4
4.6–5.0
3.8–4.0
1.5–1.6
2.5–2.6
8.1–8.5

—
—
—
—
—
—

3.0−3.3
3.3
1.8−2.0
2.8−2.9

2.9−3.6 (3.2)
2.8−3.6 (3.1)
2.0−2.5 (2.2)
2.4−2.8 (2.6)

3.5–3.8
3.2–3.6
2.1–2.2
2.7–3.1————

30.7–31.1
21.7−22.1
18.3−21.3
25.8−26.2
67.1−68.1
43.0−44.4
11.3−11.9
7.5−8.4
10.6−11.1
16.2−16.7
17.6−18.0
31.6−34.5
22.7−25.0
6.9−7.3
10.8−12.5
23.4−25.8

27.7−31.9 (30.4)
20.2−23.7 (21.8)
14.7−20.2 (17.9)
24.2−27.2 (26.3)
65.6−69.7 (67.4)
40.6−43.8 (42.5)
11.4−13.4 (12.7)
7.7−10.1 (9.2)
8.1−12.9 (11.2)
13.8−17.7 (15.8)
14.6−20.2 (17.5)
28.6−34.9 (32.7)
18.3−24.8 (22.2)
5.7−8.5 (7.1)
8.8−14.8 (10.6)
20.6−26.2 (24.3)

—
—
—
—
—
—
—
—
—
—
—
—
—
—
—
—

27.3−31.0 (29.1)
20.6−22.4 (21.3)
20.4−21.6 (20.8)
24.9−28.0 (26.4)—
37.2−45.3 (41.4)
8.9−12.8 (10.1)
8.7−10.3 (9.4)
8.7−10.9 (9.8)
13.1−15.5 (14.2)—
28.8−32.8 (30.8)
22.2−25.4 (23.7)———

30.3−33.8
29.9−30.3
51.2−54.3
34.3−36.0

27.0−34.7 (31.1)
28.1−36.2 (32.8)
39.8−51.1 (46.5)
35.3−41.3 (38.3)

—
—
—
—

25.4−33.2
29.5−34.7—
34.4−38.2

11

H. Endo et al.

73308, female, 30.2 mm SL, BSKU 73309, female, 29.4 mm SL, Kiragawa Town, Muroto City,
22 Apr. 2004.
Yaku-shima Island, Kagoshima: KAUM–I.
21837, female, 26.4 mm SL, KAUM–I. 21838,
male, 25.2 mm SL, 30º16′03″N, 130º24′48″E,
Kurio, west of Kamazeno-hana, hand-net, 0–4 m
depth, coll. by KAUM fish team, 30 July 2009.
Diagnosis. A species of Enneapterygius withthe following combination of characters: second
dorsal-fin spines 12–13 (rarely 14); anal-fin softrays 17–19 (rarely 20); pored lateral-line scales
16–19; notched lateral-line scales 17–19; 1–6
rows of small cycloid scales on boundary between
abdomen and sides; simple, unbranched dermal
flap of anterior nostril; simple dermal flap on
orbit; 1 teeth row on prevomer; mandibular pore
formula usually 4+1+4 (4–5+1+4–6); body palereddish brown in male, pale yellowish brown infemale; 2 oblique white markings on shoulder
girdle, upper one anterior to pectoral-fin base andlower one above pelvic-fin base; 8 dark saddle
markings on body; 1 row of 5–6 small, ring-likewhite markings laterally on body; 1 blackish bandwith posterior white margin on caudal peduncle.
Description. Counts and measurements areshown in Table 1. Body moderately elongate andcompressed posteriorly. Head moderate in size, its
length about 3.3 in SL. Snout moderate in length,
dorsal contour somewhat gently sloping. Maxilla
reaching posteriorly to vertical lines between
anterior rims of eye and pupil. Anterior nostril
forming short tube with an unbranced dermal flap.
Supraorbital cirrus slender, unbranched. Occipitalbranch of supratemporal canal complex, dividing
into two large branches (united in some specimens) and some sub-branches with pores laterally
and distally (Fig. 2).
First dorsal fin much lower than second. All
dorsal- and anal-fin soft rays simple. Pectoral
fin with uppermost rays simple, middle rays
branched, and lowermost rays simple. Posterior
end of pectoral fin below tenth to thirteenth spinesof second dorsal fin. Caudal fin relatively longwith 13 principal rays: upper and lowermost 2
rays simple, other rays branched. Conical teeth on
both jaws, much enlarged in outer row, sparse in
inner row. Prevomer and palatines toothed, single
12

row in the former. Gill-rakers on first arch 1+5–6.
Body covered with ctenoid scales except
for head, pectoral-fin base, and ventral side ofabdominal region from isthmus to anus. Lateralline discontinuous, composed of anterior pored
scale row and posterior notched scale row, with
one scale row between them. Anterior lateral line
extending posteriorly to vertical line at twelfth or
thirteenth spine of second dorsal fin. Posterior lateral-line origin below last pored scale, extending
posteriorly to caudal-fin base. Longitudinal scalerows below first pored lateral line 30–36. 1–6
small cycloid scale rows located between naked
area of abdomen and ctenoid scale rows on lateralbody. Vertebrae 9–10+24–26.
Color when fresh (Fig. 1A, B). No melanisticcoloration in male. Ground color in male: headreddish-brown dorsally, pale reddish to yellowishbrown laterally and ventrally; body pale reddishbrown, abdomen whitish, and ventral side above
anal fin yellowish. Ground color in female: headand body pale reddish to yellowish-brown dorsally and laterally; cheek and ventral side of head,
abdomen, and ventral side above anal-fin basewhitish. A pair of brown stripes on snout between
upper lip and eye. Suborbitals white. Some whitespots on cheek. A large dark brown blotch on
opercular region. Two oblique white bands (ringlike in some specimens) on shoulder girdle; upper
blotch anterior to lowermost pectoral-fin base andlower blotch above pelvic-fin base. Dark brown
blotch partly covered with upper white blotch anterior to pectoral-fin base. Pectoral-fin base whitewith dark blotch, with white spots anteriorly. A
dark brown band dorsally on nape to middle offirst dorsal-fin base. Eight saddle markings dorsally and laterally on body dark to blackish brown(each pair closely arrayed forming an M-shapelaterally); dorsal interspace between saddle markings whitish to yellowish. A row of 6 white ringlike markings on lateral body, obscure in somespecimens. A broad band on posterior caudal
peduncle blackish to dark brown. Narrow whiteband on caudal-fin base. Fin spines and soft rays
reddish-orange to yellow, partly whitish to translucent, forming stripes, except for white pelvic-fin
rays. Fin membranes generally translucent. First
dorsal-fin membrane with white, reddish-orange

Enneapterygius leucopunctatus from Japan

A

B

C

Fig. 2. Head lateral-line system of Enneapterygius leucopunctatus (KAUM–I. 21837, female, 8.2 mm HL, 26.4 mm SL,
Yaku-shima Island, Kagoshima Prefecture). (A) dorsal view, (B) lateral view, and (C) ventral view. Scale bar indicates
3 mm. Drawn by E. Katayama.

13

H. Endo et al.

Fig. 3. Enneapterygius vexillarius from Aguni-jima Island,
Okinawa Prefecture, Japan (holotype, ANSP 72068,
30.1 mm SL).

and blackish pigments; second dorsal-fin membrane with two dark longitudinal bands on marginand near base, three oblique white bands generally interrupted; third dorsal-fin membrane with 3
to 4 oblique reddish-oragne bands. Anal-fin rays
reddish-orange basally, yellowish distally. Caudalfin rays pale yellow in male, light grayish with awhitish transverse band on middle in female; 1 to
3 upper and lowermost rays reddish with stripes(absent or faint in some specimens). Pectoral-fin
rays pale yellow in male, pale reddish-orange infemale, with a large white blotch near middle ofbase, white spots on middle of rays, and a small
red blotch on 3 to 4 uppermost rays near base; itsbase whitish.
Color in alcohol (Fig. 1C, D). Ground colorof head and body cream to yellowish-cream.
White, yellow, and reddish-orange coloration in
fresh specimens fades in preservative except thetransverse band on the caudal fin in female becomes translucent. Red, brown, and dark brown
to blackish coloration in fresh remain as melanophores: band on upper lip and snout, two blotcheson opercular and anterior to pectoral-fin base, 3
to 4 uppermost pectoral-fin rays near base, dorsal
band on nape and anterior of first dorsal fin, 8
saddle markings on body, caudal peduncle band,
first dorsal fin, and two longitudinal bands on
second dorsal fin. In addition, melanophores present partly on underside of head with unpigmented
area, isthmus, and anterior to pelvic-fin base; head
pigmentation somewhat dense in male but sparsein female.
Distribution. In Japan, known from rocky
shores at depths of 0–4 m in Tosa Bay, Kochi
Prefecture, and Yaku-shima Island, Kagoshima
Prefecture; also Taiwan (Shen, 1994; Chiang andChen, 2008).
14

Remarks. In the western and central Pacific,
the present specimens belong to the E. etheostomus species-group (Fricke, 1997) being characterized by having: a large body size (33–54 mm
SL); high numbers of second dorsal-fin spines(12–16), anterior lateral-line scales (15–23) andtotal scale rows (32–39); parts of the anterior headand body dark in males; males with a high firstdorsal fin; and posterior lateral-line series continuing 2 rows below the anterior series. This speciesgroup includes 6 species: E. etheostoma (Jordan
and Snyder, 1902), E. vexillarius, E. miyakensis
Fricke, 1987, E. leucopunctatus, E. hsiojenae, andE. senoui Motomura, Harazaki and Hardy, 2005.
As mentioned above, Chiang and Chen (2008)
regarded E. leucopunctatus and E. hsiojenae as
valid species, clearly different from E. vexillarius
by fresh coloration, the mandibular pore pattern(4+1+4 in E. leucopunctatus, 3+1+3 in E. hsiojenae and E. vexillarius), and the counts of second
dorsal-fin spines (14 in E. hsiojenae, 13 in E.
leucopunctatus, and 16 in E. vexillarius). Further,
their molecular analysis of 11 Enneapterygius species from around Taiwan indicated that E. hsiojenae and E. leucopunctatus are separated fromeach other (E. hsiojenae forms a clade with E. elegans), although E. vexillarius was not included in
the analysis. We confirmed that the holotype of E.
vexillarius (ANSP 72068) has 16 second dorsalfin spines, 14 pored lateral-line scales, 16 notchedlateral-line scales, 2+1+4 transverse scale rows,
no dermal flap above eye (but probably damaged),
and maxillary end slightly beyond vertical line atanterior rim of pupil (Fig. 3). In addition, Fowler(1946) noted that E. vexillarius showed the following markings and coloration: anal fin reddish,with 10 transverse dark bars, most distinct alongbase of fin; caudal fin red, with 5 irregular transverse gray bars; ventral body uniformly red.
The present specimens are easily separablefrom E. etheostoma and E. miyakensis by theshape of the anterior nostril’s dermal flap (unbranched vs. 2–3 branches in other two species),
the number of transverse scale rows above anterior lateral line (2.5–3 vs. 5–6), overall markingsand coloration, and nuptial coloration in male(inconspicuous vs. melanistic: Senou, 2009),
from E. senoui by the counts of second dorsal-

Enneapterygius leucopunctatus from Japanfin spines (12–13 vs. 14–15), pored lateral-line
scales (15–18 vs. 20–21), overall markings andcoloration, and nuptial coloration (vs. melanistic
in male and yellow anal fin in female: Motomura
et al., 2005), from E. vexillarius and E. hsiojenae
by the mandibular pore pattern (usually 4+1+4
vs. 3+1+3) and overall markings and coloration(Fowler, 1946; Shen, 1994; Chiang and Chen,
2008). Further, our specimens clearly differ fromE. vexillarius by the count of second dorsal-fin
spines (12–13 vs. 16) and pored lateral-line scales(15–18 vs. 14).
Our specimens conform to E. leucopunctatusin having 12–13 second dorsal-fin spines (13 in
15 specimens), 9–11 third dorsal-fin rays (9 or 10
in 15 specimens), 17–19 anal-fin soft rays (18 in
12 specimens), usually 3+6+7 pectoral-fin rays,
15–19 pored lateral-line scales, 16–19 notchedlateral-line scales, and usually a 4+1+4 mandibular pore pattern (Table 1; Fig. 2). In addition, 2
white bands on the shoulder girdle, 6 ring-likemarkings and 8 dark brown saddle-like markings
on the body, a dark band on the caudal peduncle,
other white and blackish stripes on fin membranes, and inconspicuous dichromatism agree
well with the color photographs of E. leucopunctatus by Shen (1994). We tentatively regard somedifferences of counts and proportions among thespecimens from Taiwan (Shen, 1994; Chiang andChen, 2008), Yaku-shima Island and Tosa Bay asvariation within the species. Although the rangeof snout length in Japanese specimens is different
from that in the original description (2.9–3.6 in
HL vs. 3.5–3.8), the photographs of two paratypes
and the present materials suggest that the snout
proportions do not significantly differ from eachother (Fig. 1; Shen, 1994: fig. 8). Further, therange in Japanese specimens is almost identicalwith that in Chiang and Chen (2008) (27.0–34.7% HL vs. 25.4–33.2) (Table 1). The wide rangeof some proportional characters, however, maysuggest that the materials include more than one
species.
Enneapterygius leucopunctatus has been previously recorded only from Taiwan: Wen-tz-keng(type locality, east of Keelung, in northern Taiwan), Chenggong (central east coast) and Fengchui-sha (southern Taiwan) at depths of 2–12 m(Shen, 1994; Chiang and Chen, 2008). Hence, thepresent specimens represent the first records fromoutside Taiwan. Although the species’ known
range is extended to Yaku-shima Island and Tosa
Bay, southern Japan, its occurrence around the
Ryukyu Islands and Amami-ohshima Islands isstill unknown. If this gap between Taiwan andYaku-shima Island is confirmed by further investigation, the distributional pattern of E. leucopunctatus may support a hypothesis of the Kuroshio
Current acting as a barrier related to the endemism of the fish fauna around the Ryukyu Islands
as proposed by Senou et al. (2006).
The new Japanese name of E. leucopunctatus,“Hakuten-hebigimpo”, meaning “white spotted triplefin”, is based on the male specimen of
KAUM–I. 21838 collected from Yaku-shima Island, Kagoshima.
Comparative material examined. Enneapterygius vexillarius: ANSP 72068, holotype, 30.1mm SL, 39.0 mm in total length, Aguni-jima Island, Okinawa Islands, 27 July 1945.
Acknowledgments
We are very grateful to D. Didier-Dagit, M. H.
Sabaj, and K. Luckenbill (ANSP) for examining
and taking the photograph of the holotype of E.
vexillaris, and to H. Motomura, M. Meguro and
KAUM fish team (Kagoshima University Museum), K. Shibukawa (Nagao Natural Environment Foundation, Tokyo), and S. Harazaki (MoriTo-Umi, Kagoshima) for providing information
on Japanese triplefins. We also thank G. Yearsley(Hobart, Australia) for editing the English text.
This study was supported in part by Grant-in-Aids
for Scientific Research (A) by the Japan Society for the Promotion of Science, Tokyo, Japan(19208019).
Literature cited
Chiang, M.-C. and I.-S. Chen. 2008. Taxonomic review and molecular phylogeny of the triplefin
genus Enneapterygius (Teleostei: Tripterygiidae)
from Taiwan, with description of two new species.
The Raffles Bulletin of Zoology, Supplement, (19):183–201.
Fowler, H. W. 1946. A collection of fishes obtained in
the Riu Kiu Islands by Captain Ernest R. Tinkham
A. U. S. Proceedings of the Academy of Natural
Sciences of Philadelphia, 98:123–218.

15

H. Endo et al.

Fricke, R. 1997. Tripterygiid fishes of the western andcentral Pacific with descriptions of 15 new species, including an annotated checklist of world
Tripterygiidae (Teleostei). Koeltz ScientificBooks, Königstein. ix + 607 pp.
Fricke, R. 2009. Systematics of the Tripterygiidae(triplefins). Pages 31–67 in R. A. Patzner, E. J.
Gonçalves, P. A. Hastings and B. D. G. Kapoor(eds.). The Biology of Blennies. Science Publishers, Enfield.
Hayashi, M. 2002. Tripterygiidae. Pages 1077–1086,
1590–1591 in T. Nakabo (ed.). Fishes of Japan
with pictorial keys to the species, English edition.
Tokai University Press, Tokyo.
Holleman, W. 2005. A review of the triplefin fish genus Enneapterygius (Blennioidei: Tripterygiidae)
in the western Indian Ocean, with descriptionsof four new species. Smithiana, Publications in
Aquatic Biodiversity, Bulletin, (5):i–ii + 1–25 +pls.1–2.
Hubbs, C. L. and K. F. Lagler. 1958. Fishes of theGreat Lakes region. Cranbrook Institute of Science, (26):i–xiii + 1–213, pls. 1–44.
Meguro, M. and H. Motomura. 2010. First records
of a triplefin (Tripterygiidae), Enneapterygius
hemimelas, from Japan. Pages 1–8 in H. Motomura and K. Matsuura (eds.). Fishes of Yaku-shimaIsland – A World Heritage island in the Osumi
Group, Kagoshima Prefecture, southern Japan.
National Museum of Nature and Science, Tokyo.
Motomura, H., S. Harazaki and G. S. Hardy. 2005. A
new species of triplefin (Perciformes: Tripterygiidae), Enneapterygius senoui, from Japan with a
discussion of its in situ colour pattern. Aqua, Journal of Ichthyology and Aquatic Biolology, 10(1):
5–14.

16

Randall, J. E. and K. K. P. Lim. 2000. A checklist of
the fishes of the South China Sea. The Raffles
Bulletin of Zoology, Supplement, (8):569–667.
Senou, H. (ed.). 2009. Sea fishes of Japan. Yama-Kei,
Tokyo. 544 pp.
Senou, H., K. Matsuura and G. Shinohara. 2006.
Checklist of fishes in the Sagami Sea with zoogeographical comments on shallow water fishes
occurring along the coastlines under the influenceof the Kuroshio Current. Memoirs of the National
Science Museum, Tokyo, (41):389–542.
Shen, S.-C. 1993. Tripterygiidae. Pages 489–492,
776–777 in S.-C. Shen, S.-C. Lee, K.-T. Shao, H.K. Mok, C.-H. Chen and C.-T. Chen (eds.). Fishesof Taiwan. Department of Zoology, National Taiwan University, Taipei.
Shen, S.-C. 1994. A revision of the tripterygiid fishes
from coastal waters of Taiwan with descriptionsof two new genera and five new species. Acta
Zoologica Taiwanica, 5(2):1–32.
Shibukawa, K., T. Suzuki and M. Hosokawa. 2004.
First record of a triplefin, Enneapterygius cheni,
from Japan (Perciformes: Tripterygidae). Izu Oceanic Park Diving News, 15(3):2–6 (In Japanesewith English abstract).
Shimojyo, A. and M. Hayashi. 2000. Seven new records of tripterigid fishes from the coastal watersof Japan. Science Reports of Yokosuka City Museum, (47):39–58.
Yoshigou, H. and T. Yoshino. 2002. First records oftwo blennioid fishes, Enneapterygius rhabdotus
and Blenniella interrupta from Japan. Izu Oceanic
Park Diving News, 13(7):2–4.