Biogeography 9.
49–56. Aug. 20, 2007

Review of Japanese Records of a Grouper, Epinephelus amblycephalus(Perciformes, Serranidae), with New Specimens from Kagoshima
and Wakayama
Hiroyuki Motomura1*, Masahide Ito2, Hiromi Ikeda3, Hiromitsu Endo4,
Mizuki Matsunuma5 and Kiyotaka Hatooka6
1The Kagoshima University Museum, 1-21-30 Korimoto, Kagoshima, 890-0065 Japan
2Kasasa Fishery Cooperative Society,2-6772 Kataura, Kasasa, Minami-satsuma, Kagoshima, 897-1301, Japan
3 32-41 Akebono, Tanabe, Wakayama, 646-9837 Japan
4Laboratory of Marine Biology, Faculty of Science, Kochi University,
2-5-1 Akebono, Kochi, 780-8520 Japan
5Faculty of Fisheries, Kagoshima University,4-50-20 Shimoarata, Kagoshima, 890-0056 Japan
6Osaka Museum of Natural History,
1-23 Nagai Park, Higashi-sumiyoshi, Osaka, 546-0034 Japan

Abstract. Records of the Banded Grouper, Epinephelus amblycephalus (Perciformes: Serranidae), fromJapanese waters are reviewed on the basis of specimens examined and literature accounts, and descriptions
provided of three newly collected specimens from Kagoshima and Wakayama Prefectures. Although E.
amblycephalus has previously been thought not to occur in Japanese waters, eight individual records are now
known, five specimens having been deposited in museum collections. Distributional implications are discussed.
Key words: Serranidae, Epinephelus amblycephalus, distribution, Japan.

The Banded Grouper, Epinephelus amblycephalus (Perciformes: Serranidae), originallydescribed by Bleeker (1857) as Serranus amblycephalus from a single specimen from Ambon, theMoluccas Islands, Indonesia, is a medium-sized, primarily tropical grouper (attaining 50 cm total length)
distributed in the eastern Indian and western Pacific
Oceans, where it is usually found on offshore coraland rocky reefs (Heemstra and Randall, 1993, 1999).
Occurrences of the species around Japan have beenequivocal. Senou (2000) believed that it was unlikely to occur in Japanese waters and suggested that
two early records (Masuda, 1942; Katayama, 1960)*Corresponding author: motomura@kaum.kagoshimau.ac.jp

may have been based on specimens collected fromoverseas, prior to being landed at fish markets inJapan.
Recently, three individuals of E. amblycephalus,
collected from Japanese waters, were made directly
available to us from local fishermen. In addition, asurvey of relevant literature and fish held in Japaneseaquariums revealed both historical and aquarium
examples of the species collected from Japan.
Records of the species from Japan were reviewed
and all existing Japanese specimens examined. Distributional implications are also discussed.
Counts and measurements followed Heemstra
and Randall (1993, 1999), Allen and Robertson(1999) and Randall and Satapoomin (2000). Standard and total lengths are expressed as SL and TL,
respectively. Institutional abbreviations used in this− 49 −

Review of Japanese records of Epinephelus amblycephalus

paper are as follows: BSKU—Department of Biology, Faculty of Science, Kochi University, Japan;
FAKU—Department of Fisheries, Faculty of Agriculture, Kyoto University, Japan; HUMZ—Laboratory of Marine Zoology, Faculty of Fisheries,
Hokkaido University, Japan; KAUM—Kagoshima
University Museum, Japan; OMNH—Osaka Museum of Natural History, Japan; NMW—Naturhistorisches Museum, Wien (Vienna), Austria;
RMNH—National Museum of Natural History, Leiden, the Netherlands; URM—Department of Marine
Sciences, Faculty of Science, University of the
Ryukyus, Japan; and ZUMT—Department of Zoology, University Museum, University of Tokyo, Japan.
The following specimens recently collected fromJapanese waters were examined in this study: FAKU
93700, 340.5 mm SL, off Inami, Wakayama, 60 m
depth, long-line, 19 Nov. 2006, coll. by K. Hamamoto; KAUM-I. 821, 280.2 mm SL, north of Koejihama
Beach, Oura, Minami-satsuma, Kagoshima, 31˚27´N,
130˚13–14´E, 25–27 m depth, hand-line, 3 Oct. 2006,
coll. by F. Yamashita; and OMNH-P 13890, 270.0mm SL, off Shibushi, Kagoshima, 27 Apr. 2000,
coll. by the late M. Kumemura.

Results and Discussion
Historical records of Epinephelus amblycephalus
from Japanese waters — In their revision of the
Indo-Pacific groupers, Randall and Heemstra (1991)
examined a single specimen of E. amblycephalus(NMW 40812, 176 mm SL) collected from Yokohama, Japan, but they did not indicate any otherinformation on the specimen. According to H.
Wellendorf (NMW), the specimen was donated by
Dr Karl Haberer from Yokohama, Kanagawa Prefecture (probably in 1903). Most fishes landed in
Yokohama Market were collected outside Kanagawa
Prefecture (Yokohama Market web site). We
believe that the NMW 40812 specimen was also collected from overseas, prior to being landed at a fish
market in Yokohama.
In his review of Japanese fishes of the genus
Epinephelus, Masuda (1942) reported a single speci-

men of E. amblycephalus (as E. diacanthus amblycephalus), along with an illustration (pl. 5, uppermost fig.) and brief description. However, the locality of the specimen was unknown, although given as“probably Riukiu Islands (=Ryukyu Islands) or Formosa (=Taiwan)”. Because numerous marine fishes
collected from Taiwan had been landed at fish markets in Okinawa in the first half of the 1900s (T.
Yoshino, pers. comm.), and no subsequent records ofE. amblycephalus have been reported from the
Ryukyu Islands, Masuda’s specimen was most likely
to have been collected from Taiwan.
Masuda (1942) gave the specimen size (170 mm
TL), but did not indicate its whereabouts. However,during this study, we found a single specimen of E.
amblycephalus registered as “ZUMT 39148,
Epinephelus amblycephalus, locality unknown” inthe fish collection of the University Museum, University of Tokyo. Although there was no other information, the specimen ledger of the fish collection
indicated that specimens ZUMT 39146–39147 andZUMT 39149–39154 were collected in June ofShouwa 12 (=1937) and August of Shouwa 13(=1938), respectively, suggesting that ZUMT 39148
was collected between those dates. The specimen
measured 167.8 mm TL (134.0 mm SL), and had asimilar number and arrangement of spots on the dorsal head and body as those illustrated by Masuda(1942). Thus, it is probably the specimen used by
Masuda (1942).
Kamohara (1957) reported a single specimen (asE. diacanthus amblycephalus) from Kochi Prefecture, Shikoku, Japan as follows: “a single specimen,
122 mm long, was obtained by Mr. Kunio Amaoka,
a student of the Kochi University, at the Mimase
market, Kochi City”. Although the whereabouts had
not been given, we found a specimen (registered as
BSKU 7442) in the fish collection of Kochi University, the collection data (i. e. coll. by K. Amaoka at
Mimase Fish Market on 26 Aug. 1957) being consistent with that given by Kamohara (1957). This is thefirst specimen, based record of E. amblycephalus
confirmed from Japanese waters (Mimase Market
was probably for locally-caught fishes only). Kamo-− 50 −

Hiroyuki Motomura, Masahide Ito, Hiromi Ikeda, Hiromitsu Endo, Mizuki Matsunuma and Kiyotaka Hatooka

Fig. 1. Fresh specimens of Epinephelus amblycephalus collected from Japan, showing back spots
on fins disappearing with growth. a, OMNH-P 13890, 270.0 mm SL, off Shibushi, Kagoshima Pref. (photo by K. Hatooka, 2 days after collection); b, KAUM-I. 821, 280.2 mm SL,
north of Koejihama Beach, Oura, Minami-satsuma, Kagoshima Pref. (photo by H. Motomura); c, FAKU 93700, 340.5 mm SL, off Inami, Wakayama Pref. (photo by H. Ikeda).

hara (1957) also proposed a new Japanese name,
Kokuten-aohata, for the species.
In his catalog of fishes of Kochi Prefecture,
Kamohara (1958) included E. diacanthus ambly-

cephalus, without reference to specimens. There
being no additional specimens of E. amblycephalus
collected before 1958 in the fish collection of Kochi
University, this was probably based on his earlier− 51 −

Review of Japanese records of Epinephelus amblycephalus(Kamohara, 1957) record. Subsequently, Kamohara(1960) listed the species in a list of the shore fishesof Okinoshima and adjacent regions, Kochi Prefecture. Although specimen data was not included, therecord was probably based on a specimen (BSKU
8475, 89.5 mm SL) collected from Susaki, Kochi, on
17 April 1959, since the listed species included some“collected from Susaki” (see Introduction of Kamohara, 1960). Thus, BSKU 8475 appears to have been
the second example of E. amblycephalus collectedfrom Japanese waters.
Katayama (1960) described E. amblycephalus in
detail on the basis of a single specimen collectedfrom Yamakawa (correct spelling, Yamagawa; current name, Kaimon-cho, Ibusuki City), Kagoshima,
Kyushu, Japan, including an illustration (pl. 46).
The present whereabouts of the specimen is
unknown. The distribution of the species given by
Katayama (1960) included “Amboina [=Ambon,
Indonesia]; Riu Kiu Islands or Formosa [=theRyukyu Islands or Taiwan]; Japan (Yamakawa [=Kaimon], Kagoshima Prefecture)”, the first two
localities having been based on Bleeker (1857) and
Masuda (1942), respectively. Although Katayama(1960) did not refer to Kamohara’s (1957, 1958,
1960) publications and did not include Kochi in the
distribution of the species, he did use the Japanesename, Kokuten-aohata, proposed by Kamohara(1957).
In his revised catalog of fishes of Kochi Prefecture, Japan, Kamohara (1964) referred to his earlier
papers (above) and listed E. amblycephalus. However, no new specimens have been collected orreported from Kochi since Kamohara’s (1960)
record.
In the fishes of the Japanese Archipelago, Katayama (1984) briefly described E. amblycephalus andprovided a color photograph (pl. 114, taken by K.
Kyushin). However, the photograph is the same as
one reported by Kyushin et al. (1982), the specimen
having been collected from the South China Sea.
That specimen has been registered in the fish collection of the Hokkaido University Museum (HUMZ
87475, 05˚52.00´N, 114˚50.00´E, 111 m, 4 JulyFig. 2. Map of Japan (except for Hokkaido) and adjacentwaters. Numbers in this figure indicate capture
localities of specimens examined in this study(numbers in the order of collected date, oldest
first): 1, Kochi, Kochi Pref.; 2, Susaki, Kochi
Pref.; 3, Shibushi, Kagoshima Pref.; 4, Minamisatsuma, Kagoshima Pref.; and 5, Inami,
Wakayama Pref.

1975) (H. Imamura, pers. comm.).
Senou (1993, 2000) later referred to Masuda(1942) and Katayama (1964) and concluded that
there was little possibility that E. amblycephalus
occurred on the coast of Japan. In addition, Senou(2002) stated “Katayama (1960) reported thisspecies from Yamakawa of Kagoshima Prefecture,
but there has been no further firm record based on
specimens from Japan”. However, he overlooked
the two BSKU specimens from Kochi, which wereused in Kamohara’s (1957, 1960) lists.
Misaki (2000) reported a single individual of E.
amblycephalus (ca 60 cm TL) collected by longline
off Kushimoto, Wakayama Prefecture, at a depth ofca 40 m, on 4 September 2000. Subsequently kept at
Kushimoto Marine Park, Wakayama, the specimen
died in August, 2005 and was not retained (Y.
Kotera, pers. comm.).
However, an example of E. amblycephalus collected from off Tomita, Shirahama, Wakayama Prefecture on 16 October 1997 (H. Tanase, pers. comm.

− 52 −

* As “Kokuten-aohata”; **capture locality unknown, landed in Yokohama

Locality
SourceOne specimen (ZUMT 39148, 134.0 mm SL)
TaiwanOne specimen (BSKU 7442, 98.3 mm SL)
Kochi, Kochi Pref.
No specimens; quoted from Kamohara (1957)—
One specimen (BSKU 8475, 89.5 mm SL)
Susaki, Kochi Pref.
One specimen; present whereabouts and size unknown
Ibusuki, Kagoshima Pref.
No specimens; quoted from Kamohara (1957, 1958, 1960)—
One specimen (HUMZ 87475, 386 mm SL)
off Brunei, South China Sea
One specimen (NMW 40812, 176 mm SL)
Yokohama, Kanagawa Pref.**
No specimens; quoted from Masuda (1942) and Katayama (1960)——
No specimens; quoted from Masuda (1942) and Katayama (1960)
Kushimoto, Wakayama Pref.
One individual; kept in aquarium and died in Aug. 2005; specimen not retained
Shirahama, Wakayama Pref.
One individual; kept in aquarium and still alive as of Jan. 2007—
No specimens; quoted from Masuda (1942) and Katayama (1960)
Shibushi, Kagoshima Pref.
One specimen (OMNH-P 13890, 270.0 mm SL)
Minami-satsuma, Kagoshima Pref. One specimen (KAUM-I. 821, 280.2 mm SL)
Inami, Wakayama Pref.
One specimen (FAKU 93700, 340.5 mm SL)
Reference
Masuda (1942)
Kamohara (1957)
Kamohara (1958)
Kamohara (1960)
Katayama (1960)
Kamohara (1964)
Katayama (1984)
Randall & Heemstra (1991)
Senou (1993)
Senou (2000)
Misaki (2000)
Misaki (2000)
Senou (2002)
This study
This study
This study
Reported scientific name
E. diacanthus amblycephalus
E. diacanthus amblycephalus
E. diacanthus amblycephalus
E. diacanthus amblycephalus
E. amblycephalus
E. amblycephalus
E. amblycephalus
E. amblycephalus
E. amblycephalus
E. amblycephalus—*—*
E. amblycephalus
E. amblycephalus
E. amblycephalus
E. amblycephalus

Table 1. Epinephelus amblycephalus reported from Japanese waters (see text).

Hiroyuki Motomura, Masahide Ito, Hiromi Ikeda, Hiromitsu Endo, Mizuki Matsunuma and Kiyotaka Hatooka

in Misaki, 2000) and kept at the Kyoto University
Aquarium, is alive as of January 2007. Accordingly,
five reliable records of E. amblycephalus fromJapanese waters (one from Kagoshima, two from
Kochi and two from Wakayama) have been reported
prior to this study, two specimens being currently
retained as BSKU 7442 and 8475 (Table 1).
Newly collected specimens from Kagoshima andWakayama Prefectures — Recently, three specimensof E. amblycephalus have been collected fromJapanese waters (Fig. 1), from Shibushi, Kagoshima(Pacific coast side) on 27 April 2000, from Minamisatsuma, Kagoshima (East China Sea side) on 3
October 2006 and from Inami, Wakayama (Pacificcoast) on 19 November 2006 (Fig. 2). Counts andmeasurements of the three specimens are given in
Table 2, along with data for BSKU 7442 and 8475.
Morphological description based on all of theJapanese specimens — Dorsal profile of head slightly convex; nape and anterodorsal portion of bodywithout numerous pores; anterior and posterior nostrils subequal in size; posterior margin of maxilla
extending slightly beyond a vertical through posterior margin of eye (not confirmed in BSKU 7422
specimen as its mouth widely opened and head wasbent dorsally); middle part of lower jaw with 2 rowsof teeth; palatines with teeth; preopercular margin
with 4–6 slightly enlarged serrae posteroventrally
and 13–32 tiny serrae centrally, upper marginsmooth, without serrae in larger specimens; 2 opercular spines; interspinous dorsal-fin membranes
moderately incised; base of spinous portion of dorsal
fin longer than base of soft-rayed portion; pectoral
fins nearly symmetric, middle rays longest; base of
upper pectoral-fin rays joined by scaly skin flap tobody; caudal-fin margin rounded; lateral-line scaleswith single tubule. Coloration of fresh specimens(based on color photographs of KAUM-I. 821 and
FAKU 93700): head and body brownish to whitish,with dark brown bars; anterior portion of head with 4
dark brown bars radiating from eye, 2 of the 4extending onto upper-jaw lip; nape with a broad darkbrown saddle; body with 5 dark brown bars, first to
fourth bars extending onto ventral surface from dor-

− 53 −

Review of Japanese records of Epinephelus amblycephalus

Table 2. Counts and measurements of all existing specimens of Epinephelus amblycephalus collected fromJapanese waters.
Registration number
Locality
Collection dateStandard length (mm)Total length (mm)Dorsal-fin rays
Anal-fin rays
Pectoral-fin rays
Pelvic-fin rays
Lateral-line scalesLongitudinal scale series
Gill rakers% of standard length
Body depth
Body width
Head length
Snout length
Orbit diameter
Interorbital width
Upper-jaw length
Caudal-peduncle depth
Caudal-peduncle length
Predorsal length
Preanal length
Prepelvic length
First dorsal-fin spine length
Longest dorsal-fin spine length
Longest dorsal-fin soft ray length
First anal-fin spine length
Second anal-fin spine length
Third anal-fin spine length
Longest anal-fin soft ray length
Caudal-fin length
Pectoral-fin length
Pelvic-fin spine length
Longest pelvic-fin soft ray length

BSKU 7442
Kochi Pref.
Aug. 1957
98.3
122.1
XI, 15
III, 8
18
I, 5
52
97
7+16=23

BSKU 8475
Kochi Pref.
Apr. 1959
89.5
110.6
XI, 16
III, 8
19
I, 5
51
105
8+15=23

37.6
18.6
40.2
9.7
10.2
5.4
19.0
10.2
16.6
37.7
73.9
45.8
7.6
14.6 (4th)18.1 (4th)
6.7
14.1
12.4
17.8 (4th)
24.5
23.3
11.8
19.8** (2nd)

39.4
19.3
41.8
10.5
9.8
6.0
20.9
10.4
17.2
37.1
70.6
40.7
8.2
16.6 (4th)
18.0 (4th)
7.2
11.5
11.7
17.0 (4th)
23.7
19.6
12.7
21.6 (3rd)OMNH-P 13890 KAUM-I. 821* FAKU 93700
Kagoshima Pref. Kagoshima Pref. Wakayama Pref.
Apr. 2000
Oct. 2006
Nov. 2006
270.0
280.2
340.5
329.0
344.3
415.1
XI,15
XI, 15
XI, 14
III,8
III, 8
III, 8
18
19
18
I,5
I, 5
I, 5
57
–
50
115
–
114
7+16=23–
8+16=24
38.9
21.3
42.4
11.2
7.6
8.1
20.6
11.7
15.3
36.7
69.6
37.1
7.4
14.4 (4th)15.1 (4th)
1.9
10.5
11.2
20.1 (4th)
21.4
21.9
11.6
18.1 (3rd)

37.8
20.1
43.6
11.7
7.3
8.4
21.1
11.5
15.4
37.2
73.4
42.5
5.7**
13.8 (4th)15.4 (4th)
4.2
8.8
9.6
19.2 (4th)
22.9
23.8
12.5
21.8 (3rd)

38.0
19.4
42.2
11.3
6.6
8.0
19.7
11.3
14.5
37.3
71.7
40.2
6.2
13.0 (4th)15.2 (4th)
4.1
8.2
8.8
18.9 (4th)
21.9
22.9
11.5
19.8 (3rd)*Gill, internal organs and scales removed. **Tip slightly broken.

sal-fin membranes, fifth bar on caudal peduncle, narrowest; numerous black spots scattered mainly alongedges of bars and saddle; eye membrane yellowish;
maxillary groove vivid yellow; dorsal fin yellowishbrown; pectoral fin yellowish translucent; pelvic andanal fins dark brown basally, yellowish-brown distally; caudal fin brown distally, yellowish-brown distally, with a narrow yellow marginal band. Specimen

OMNH-P 13890 with black spots on all fins, characteristic of young stages of the species (Heemstra andRandall, 1993, 1999), but which eventually disappear with growth (see Fig. 1).
The above characters of the Japanese specimens
were completely consistent with those of E. amblycephalus given by Heemstra and Randall (1993,1999) and the holotype of S. amblycephalus (RMNH− 54 −

Hiroyuki Motomura, Masahide Ito, Hiromi Ikeda, Hiromitsu Endo, Mizuki Matsunuma and Kiyotaka Hatooka

5159, 241.5 mm SL, Ambon, Indonesia) examinedin this study. Although Heemstra and Randall(1993, 1999) described number of the dorsal-fin softrays as 15 or 16, the holotype and the Wakayama
specimen had 14 soft rays. In addition to thoseexamined in this study, all of the specimens reported
in the publications listed in Table 1 were identified
as E. amblycephalus, on the basis of their descriptions, illustrations and photographs.
Distribution and conclusions — The literatureand specimen survey revealed that a total of eightindividuals of E. amblycephalus have been collected
to date from Japanese waters, despite the considered
rarity or absence of the species in Japanese waters (e.
g. Senou, 2002). Of the eight individuals, five specimens are currently held in museum collections(Tables 1–2).
Epinephelus amblycephalus is distributed in theeastern Indian and western Pacific Oceans, where it
ranges from the Andaman Sea east to Fiji (Heemstra
and Randall, 1993, 1999). In Japanese waters, thespecies has been recorded from Kagoshima, Kochi
and Wakayama Prefectures, but has never beenfound in the Ryukyu Islands (Masuda’s specimen
was probably collected from Taiwan) or north of
Wakayama Prefecture. These three aforementioned
prefectures face the flow path of the Kuroshio Current running from Taiwan via the coast of China, but
not through the Ryukyu Islands. This suggests thatthe primary tropical epibenthic fish, E. amblycephalus, is transported by the Kuroshio Current to
the Pacific coast of southern Japan from Taiwan or
China, at egg or larval stages. The Japanese specimens varied from juveniles to adults (89.5–340.5mm SL), being collected throughout the year (Apr.,
Aug., Oct. and Nov.). Such indicates that E. amblycephalus can probably over-winter and attain adult
size in Japanese waters (at 340.5 mm SL, the
Wakayama specimen is close to maximum sizerecorded for the species). However, because so few
of this distinctly colorful grouper have been foundover the past 50 years in Japanese waters, being firsttime catches for all of the local fishermen involved,
it is unlikely that the species reproduces around

Japan.

Acknowledgments
We greatly appreciate Fukuyoshi Yamashita(Kasasa Fishery Cooperative Society, Japan), Mitsuo
Kumemura (Shibushi Fishery Cooperative Society,Japan) and Shunji Kaji (Shibushi Station, National
Center for Stock Enhancement, Fisheries Research
Agency, Japan) for opportunities to examine the
newly collected specimens of Epinephelus amblycephalus, and Tetsuji Nakabo (FAKU), Mayumi
Takayama, Yuriko Haraguchi, Nami Kita, Masatoshi
Meguro and Gouta Ogihara (KAUM) and Junko
Omori (Kagoshima Aquarium, Japan) for their curatorial assistance. We thank Kazuo Sakamoto(ZUMT) for a specimen loan, and Takeshi
Yamakawa (BSKU), Hisashi Imamura (Hokkaido
University Museum, Japan), Helmut Wellendorf(NMW), Tetsuo Yoshino (URM) and Yoshihiko
Kotera (Kushimoto Marine Park, Japan) for providing data and information on previously reported
specimens of E. amblycephalus. The first author
thanks Yuko Motomura (Kagoshima, Japan) for her
assistance, Martien J. P. van Oijen (RMNH) for
opportunity to examine the holotype of Serranus
amblycephalus and Graham S. Hardy (Ngunguru,
New Zealand), who read the initial manuscript andoffered helpful comments. 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) and for Young Scientists(B) by the Ministry of Education, Science, Sports
and Culture, Japan (19770067).

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—, 2002. 219 Serranidae. Groupers, basslets
and soapfishes. In Nakabo, T. (Ed), Fishes ofJapan with pictorial Keys to the Species (English
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Press, Tokyo.

− 56 −(Accepted February 21, 2007)