Species Diversity 17: 7–14
25 May 2012

First Records of the Two-tone Goatfish, Upeneus
guttatus, from Japan, and Comparisons with U. japonicus(Perciformes: Mullidae)
Hiroyuki Motomura1, Masahiro Yamashita1, Masahide Itou2,
Yuriko Haraguchi1 and Yukio Iwatsuki3
1 The Kagoshima University Museum, 1-21-30 Korimoto, Kagoshima 890-0065, Japan

E-mail: motomura@kaum.kagoshima-u.ac.jp

2 Kasasa Fishery Cooperative Society, 2-6772 Kataura, Kasasa, Minami-satsuma, Kagoshima 897-1301, Japan
3 Division of Fisheries Sciences, Faculty of Agriculture, University of Miyazaki,

1-1 Gakuen-kibanadai-nishi, Miyazaki 889-2192, Japan(Received 11 October 2011; Accepted 11 January 2012)
Upeneus guttatus (Day, 1868) (Perciformes: Mullidae) is recorded from Japan for the first time on the basis of 21 specimens (57.8–139.5 mm standard length) collected from Kagoshima Prefecture, southern Japan. These specimens also represent the first records of U. guttatus from East Asia and the northernmost records in the western Pacific Ocean; this species
has not previously been recorded north of the Philippines. The Japanese specimens of U. guttatus, described herein in detail,and 14 specimens from the Indo–West Pacific are compared with 33 specimens of U. japonicus (Houttuyn, 1782).
Key Words: Teleostei, Actinopterygii, Upeneus japonicus, Kagoshima, distribution.

Introduction
Upeneus Cuvier, 1829 (Perciformes: Mullidae) was recently reviewed by Uiblein and Heemstra (2010), who recognized 26 valid species in the genus, including their four
new species from the western Indian Ocean. Subsequently,
Uiblein and Heemstra (2011a, b) described U. randalli andU. seychellensis as new species from the Persian Gulf andSeychelles Bank respectively. Uiblein and Heemstra (2010)
defined four major species groups within Upeneus using a
combination of the numbers of dorsal-fin spines, gill rakers
and pectoral-fin rays, and caudal-fin coloration.
Upeneus guttatus (Day, 1868), widespread Indo–West Pacific species, was included in the Upeneus japonicus species
group, which is characterized by having seven dorsal-fin
spines (Uiblein and Heemstra 2010, 2011b). In the northwestern Pacific Ocean, U. guttatus had been recorded only
from the Philippines until an ichthyological survey of southern Japan yielded 21 specimens from the East China Sea andPacific Ocean sides of Kagoshima Prefecture at depths ofless than 40 m. These specimens are described herein as thefirst records of U. guttatus from Japan and the northernmost
records of this species in the western Pacific Ocean. Detailed
comparisons of U. guttatus with a co-occurring species, U.
japonicus (Houttuyn, 1782), are also made.

Material and Methods
Counts and measurements generally follow Randall andKulbicki (2006) and Uiblein and Heemstra (2010). Standard
length is abbreviated as SL. Osteological characters were
examined from radiographs taken from six specimens ofUpeneus guttatus. The formula for the configuration of theanterior neural spines and anterior dorsal fin pterygiophores
follows Ahlstrom et al. (1976). The presence of a swimbladder was confirmed by dissection of the abdomen on theright side. The ‘Description’ section below is based on Japanese specimens of U. guttatus.
Specimens examined in this study have been deposited in
the Australian Museum, Sydney (AMS); Bishop Museum,
Honolulu (BPBM); Fisheries Research Laboratory, Mie University, Shima (FRLM); Kagoshima University Museum, Kagoshima (KAUM); Muséum National d’Histoire Naturelle,
Paris (MNHN); National Museum of Nature and Science,
Tsukuba (NSMT); Seikai National Fisheries Research Institute, Nagasaki (SNFR); Museum of Zoology, University ofMichigan, Ann Arbor (UMMZ); and Museum Support Center, Smithsonian Institution National Museum of Natural
History, Suitland (USNM).
Full counts and measurements of the following 23 specimens (74.7–139.0 mm SL) of Upeneus japonicus weretaken: Japan — KAUM–I. 388, 95.8 mm SL, 19 Apr. 2006,
KAUM–I. 7015, 110.7 mm SL, 9 June 2007, KAUM–I. 9847,
104.8 mm SL, 7 May 2008, east of Sakinoyama, Kataura, Kasasa, Minami-satsuma, Kagoshima, 31°25′44″N,
130°11′49″E, set net, 27 m, M. Itou; KAUM–I. 3128,
139.0 mm SL, off Kouzaki-yama, Kataura, Kasasa, Minamisatsuma, Kagoshima, 31°26′00″N, 130°10′05″E, set net,
36 m, Y. Masuda, 20 Apr. 2007; KAUM–I. 9212, 90.9 mm
SL, 9 Apr. 2009, KAUM–I. 13819, 90.6 mm SL, 28 Jan. 2009,© 2012 The Japanese Society of Systematic Zoology

8

Hiroyuki Motomura et al.

1 km southwest of Kawajiri Fishing Port, Kaimon-kawajiri,
Ibusuki, Kagoshima, 31°10′N, 130°32′E, set net, 40 m, G.
Ogihara and T. Yoshida; KAUM–I. 5947, 82.6 mm SL, 8
Aug. 2007, KAUM–I. 7440, 80.5 mm SL, 10 Dec. 2007, off
Chiringa-jima island, Ibusuki, Kagoshima Bay, 31°16′38″N,
130°40′18″E, set net, 25 m, Orita Fishery; KAUM–I. 331,
83.4 mm SL, Uchinoura Bay, Kimotsuki, Kagoshima,
31°17′N, 131°05′E, set net, 40 m, M. Yamada, 6 Apr. 2006;
KAUM–I. 792, 94.0 mm SL, 1 km north of Izashiki Port,
Sata, Minami-osumi, Kagoshima, 31°05′N, 130°41′E, set
net, 30–40 m, M. Chikuchishin, 25 Sep. 2006; KAUM–I.
9614, 89.8 mm SL, Meitsu, Nango, Miyazaki, 31°32′44″N,
131°22′53″E, H. Motomura, 26 Apr. 2008; KAUM–I.
12623, 107.6 mm SL, off Urato, Mimase, Kochi, 33°29′N,
133°34′E, trawl, 100–200 m, G. Ogihara and M. Meguro,
9 Mar. 2008; NSMT-P 45177, neotype of Mullus japonicus,
96.5 mm SL, off Futo Harbor, east of Izu Peninsula, Shizuoka, 15 m, K. Okamoto et al., 12 Mar. 1993. China — SNFR
12061, 5, 109.0–128.5 mm SL, Yellow Sea, 9 Feb. 1997. Taiwan — KAUM–I. 39239, 74.7 mm SL, off Kaohsiung, trawl,
H. Ho and M. Matsunuma, 3 July 2011; KAUM–I. 39614,
115.5 mm SL, KAUM–I. 39615, 123.4 mm SL, KAUM–I.
39616, 119.3 mm SL, KAUM–I. 39617, 110.9 mm SL, off
Keelung, M. Meguro and M. Matsunuma, 13 July 2011.
Selected counts (i.e., gill rakers) and measurements (i.e.,barbel and pectoral-fin lengths) of the following 10 specimens (96.0–137.3 mm SL) of U. japonicus from Japan weretaken for comparative purposes: KAUM–I. 21033, 115.1 mm
SL, off Kouzaki-yama, Kataura, Kasasa, Minami-satsuma,
Kagoshima, 31°26′00″N, 130°10′05″E, set net, 36 m, M. Itou,
17 Apr. 2009; KAUM–I. 18956, 103.7 mm SL, 1 km southwest of Kawajiri Fishing Port, Kaimon-kawajiri, Ibusuki, Kagoshima, 31°10′N, 130°32′E, set net, 40 m, G. Ogihara andT. Yoshida, 15 Apr. 2009; KAUM–I. 21228, 114.4 mm SL, off
Chiringa-jima island, Ibusuki, Kagoshima Bay, 31°16′38″N,
130°40′18″E, set net, 25 m, Orita Fishery, 8 July 2009;
KAUM–I. 30949, 137.3 mm SL, KAUM–I. 30956, 127.4 mm
SL, Shibushi Bay, Shibushi, Kagoshima, 31°38′N, 131°14′E,
trawl, 70–100 m, G. Ogihara, M. Yamashita, and Y. Ohashi,
8 July 2010; UMMZ 14279, 5, 96.0–128.3 mm SL, market in
Kobe, Hyogo, Japan, 1922.
Upeneus guttatus (Day, 1868)[New standard Japanese name: Akane-himeji](Figs 1A, B, 2, 3; Tables 1–2)
Upeneoides guttatus Day, 1868: 938 [type locality: Madras(currently Chennai), India].
Upeneus bensasi (not of Temminck and Schlegel 1843): Day
1875: 121, pl. 30, fig. 5 (Madras, India); Lachner 1954: 509(in part; Philippines).
Upeneus crosnieri Fourmanoir and Guézé, 1967: 52, fig. I/c(type locality: Mitsio, Pracel Bank, Madagascar).
Upeneus sp. 1: Gloerfelt-Tarp and Kailola 1984: 215, lowermost right fig. (southwestern Indonesia).
Upeneus guttatus: Randall and Kulbicki 2006: 301, figs 3–4(Indo–West Pacific); Uiblein and Heemstra 2010: 42, pls
1C, 3C (Indian Ocean); Uiblein and Heemstra 2011b:

647, figs 1B, 2–3 (Indian Ocean).
Japanese specimens examined. 21 specimens, 57.8–
139.5 mm SL: KAUM–I. 5492, 116.7 mm SL, 24 July 2007,
KAUM–I. 7819, 57.8 mm SL, 7 July 2007, KAUM–I. 9636,
84.4 mm SL, 26 Apr. 2008, KAUM–I. 9845, 100.1 mm SL,
KAUM–I. 9846, 89.3 mm SL, 7 May 2008, KAUM–I. 10542,
91.3 mm SL, 29 May 2008, KAUM–I. 11885, 80.5 mm
SL, KAUM–I. 11887, 88.3 mm SL, 5 June 2008, KAUM–
I. 9847, 104.8 mm SL, KAUM–I. 11894, 91.5 mm SL, 10
June 2008, KAUM–I. 12842, 128.4 mm SL, 30 June 2008,
KAUM–I. 12844, 107.5 mm SL, 9 June 2008, KAUM–
I. 13067, 119.2 mm SL, 9 Sep. 2008, KAUM–I. 24229,
82.1 mm SL, 23 May 2009, KAUM–I. 24422, 74.4 mm SL,
22 July 2009, KAUM–I. 24423, 84.3 mm SL, 12 June 2009,
KAUM–I. 30431, 135.6 mm SL, 1 June 2010, KAUM–
I. 36056, 125.1 mm SL, 2 Sep. 2010, east of Sakinoyama,
Kataura, Kasasa, Minami-satsuma, Kagoshima, 31°25′44″N,
130°11′49″E, set net, 27 m, M. Itou; KAUM–I. 11011,
114.1 mm SL, 28 July 2008, KAUM–I. 21032, 139.5 mm SL,
17 Apr. 2009, KAUM–I. 21035, 79.9 mm SL, 26 May 2009,
off Kouzaki-yama, Kataura, Kasasa, Minami-satsuma, Kagoshima, 31°26′00″N, 130°10′05″E, set net, 36 m; KAUM–I.
6999, 61.5 mm SL, Uchinoura Bay, Kimotsuki, Kagoshima,
31°17′N, 131°05′E, 15 Feb. 2007, set net, 40 m, M. Yamada.
Comparative material from the Indo–West Pacific. 14
specimens, 96.3–133.9 mm SL: AMS I. 25, syntype of Upeneoides guttatus, 104.4 mm SL, Madras (Chennai), India,
13°01′N, 80°03′E, F. Day; BPBM 33440, 115.4 mm SL,
Mafia Channel, Tanzania, bottom trawl, 10–50 m, F. Nansen, Nov. 1982; BPBM 33815, 107.3 mm SL, Chesterfield
Islands, New Caledonia, R/V Alis, 20 Aug. 1988; BPBM
39470, 117.2 mm SL, Belep Islands, New Caledonia, trawl,
35 m, M. Kulbicki, R/V Vauban, 10 July 1986; FRLM 26495,
133.9 mm SL, Bitung, Sulawesi, Indonesia, S. Kimura and T.
Peristiwady, 23 Oct. 2000; FRLM 29978, 96.3 mm SL, FRLM
30073, 110.8 mm SL, Phuket, Thailand, S. Kimura et al., 24
Sep. 2003; MNHN 1965-17, holotype of Upeneus crosnieri,
127.5 mm SL, Mitsio, Pracel Bank, Madagascar, 45 m, A.
Crosnier; USNM 306102, 3, 102.8–129.0 mm SL, Ras Binnah, Somalia, 11°16′18″N, 51°15′36″E, 57–59 m, G. Small,
16 Nov. 1989; USNM 395448, 3, 106.9–115.2 mm SL, off Isle
Umm, Massawa, Eritrea, Red Sea, 15°09′N, 40°31′E, 35 m,
L. Knapp, 19 Sep. 1971.
Description. Counts and measurements, expressed aspercentages of SL, are given in Table 1. Frequency distributions of gill-raker counts are given in Table 2.
First dorsal-fin spine longest; all dorsal-fin soft rays
branched. Anal-fin spine extremely short; first anal-fin
soft ray unbranched and spinous, segmented in outer half.
Mouth small, ventral and oblique; posterior margin of maxilla extending beyond a vertical through anterior margin oforbit; posterior margin of maxilla membranous and convex;
upper-jaw length less than half head length. Posterior tip ofdepressed barbel reaching (rarely slightly extending beyond)a vertical through preopercular margin. Band of small nodular teeth in each jaw, about 2 or 3 rows of teeth at front,
becoming about 6–8 rows centrally. Vomer with a few vil-

The goatfish Upeneus guttatus from Japan

9

Fig. 1. Color photographs of fresh specimens of (A–B) Upeneus guttatus and (C–D) U. japonicus. A–B: KAUM–I. 11011, 114.1 mm SL, Kagoshima, Japan; C–D: KAUM–I. 9212, 90.9 mm SL, Kagoshima, Japan.

liform teeth posteriorly, no teeth anteriorly. Palatines withbroad band of villiform teeth. Teeth present on ectopterygoids. Tongue fused to floor of mouth. Anterior nostril ashort, vertically oval opening with dorsal edge of opening
below level of ventral margin of orbit. Posterior nostril vertically oval opening, dorsal edge of opening at same hight asventral margin of pupil. Length of longest gill raker on firstgill arch subequal to that of longest gill filaments. Opercular spine at level of middle of eye, spine tip not reaching
opercular margin. Scales finely ctenoid; body scales extending onto bases of second dorsal, anal, and caudal fins. First
dorsal-fin origin above fourth pored lateral-line scale. Second dorsal-fin origin anterior to anal-fin origin. Pectoral fin
extending beyond a vertical through last spine base of first10

Hiroyuki Motomura et al.

Fig. 2. Color variations of caudal fins of Upeneus guttatus from Kagoshima, Japan. A, KAUM–I. 7819, 57.8 mm SL; B, KAUM–I. 11885,
80.5 mm SL; C, KAUM–I. 24423, 84.3 mm SL; D, KAUM–I. 13067, 119.2 mm SL.

Fig. 3. Relationships of (A) barbel length and (B) pectoral-fin length to standard length in Upeneus guttatus (closed symbols: stars fromJapan, squares from Indo–West Pacific) and U. japonicus (open circles).

dorsal fin. Posterior tip of depressed pelvic fin extending
slightly beyond a vertical through pectoral-fin tip, but notreaching to anus. Formula for configuration of supraneuralbones, anterior neural spines, and anterior dorsal pterygiophores 0/0/0+1/1+1/1/1/1/1/. Vertebrae 10+14. Upper
series of procurrent caudal-fin rays 7 or 8, lower series 7;
segmented caudal-fin rays 10+9; branched caudal-fin rays
7+6. Swimbladder present.
Color when fresh (based on color photographs of 19 fresh
specimens): Head and body reddish dorsally, whitish pink
ventrally, sometimes with faint longitudinal stripe from eye
to caudal-fin base. Barbels yellow. First dorsal fin pale redanteriorly, translucent with poorly defined reddish spots
posteriorly. Second dorsal fin translucent with 4 irregular
red bars. Pectoral fin translucent yellow. Pelvic and anal finstranslucent with indistinct reddish spots scattered on rays.
Caudal-fin upper lobe with 4 or 5 red bars reaching to fin
margin and indistinct red bar close to fin base (not reaching to margin). Caudal-fin lower lobe with 4–9 reddish bars
or spots on ventral margin of lobe (bars and spots distinct

when young, becoming more irregular with growth); upper half of lobe uniformly reddish without spots or stripes.
Posterior margin of caudal fin white, dark sub-marginally.
In preserved specimens, reddish bars and spots faded andbarely discernible.
Distribution. Widely distributed in the Indo–West Pacific, ranging from the east coast of Africa and the Red Sea
east to southern Japan, the Philippines, and New Caledonia(Uiblein and Heemstra 2011; this study). The present specimens from Kagoshima represent the first records of U. guttatus from Japan and also from East Asian waters.
Remarks. Upeneus guttatus was originally described by
Day (1868) as Upeneoides guttatus from Madras (currently
Chennai), India. Day’s description was based on multiple
specimens, although he did not provide registration numbers or the number of type specimens. Three syntypes arerecognized in museum collections, i.e., AMS I.25, BMNH
1868.5.14.11, and BMNH 1975.9.30.19 (dry) (Eschmeyer1998).
Uiblein and Heemstra (2010, 2011b) redescribed U. gut-

2

I, 7

Anal-fin rays

Scales above lateral line

6.7–9.1 (7.7)

7.2–8.0 (7.6)

10.2–11.7 (11.2)

Orbit diameter

Interorbital width

Upper-jaw length

29.2–32.7 (30.8)

63.7–65.7 (61.4)

20.6–22.4 (21.4)

17.8–22.4 (19.8)

16.0–20.0 (17.5)

12.5–16.6 (14.2)

7.1–9.6 (8.8)

Pre-first dorsal-fin length

Pre-pelvic-fin lengthPre-anal-fin length

First dorsal-fin spine length

Second dorsal-fin spine length

Third dorsal-fin spine length

Fourth dorsal-fin spine length

First dorsal-fin soft ray length

14.0–16.0 (15.1)

34.9–37.3 (36.2)

Caudal-peduncle length

Second dorsal-fin soft ray length

9.5–10.7 (10.1)

21.2–24.9 (23.0)

Caudal-peduncle depth

7.9–9.2 (8.5)

10.4–12.5 (11.3)

Snout length

16.3–20.2 (18.4)

26.8–30.5 (28.3)

Head length

Barbel length

13.3–15.8 (14.5)

Cheek depth

23.1–26.3 (24.7)

15.0

8.7

12.4

15.4

18.8
—

61.7

32.2

36.5

23.8

10.5

18.7

8.6

11.9

7.7

6.8

11.9

28.8

13.3

23.7

6+16=22

Body width3

4–5 (4)

5

2

29

I, 7

I, 5

14

VII, 8

104.4

India
Syntype of U. guttatus
AMS I. 25

5–7 (6)+16–18 (17)=22–24 (23)

5–5.5 (5.5)

Body depth

Measurements (% of SL)

Gill rakers (upper+lower = total)

Scale rows between dorsal fins

Scales below lateral line

28–29 (29)I, 5Pelvic-fin rays

Pored lateral-line scalesVII, 913–14 (13)Pectoral-fin rays

57.8–139.5

Dorsal-fin rays

CountsJapan (Kagoshima)
Non-types
n=21

13.9

13.5–16.0 (14.6)

7.4–9.9 (8.5)

11.3–13.7 (12.6)—
9.1

17.2–21.2 (19.8)
13.8–18.1 (16.5)——

19.6–23.5 (22.1)

63.7–68.6 (65.6)

28.6–32.9 (30.9)

34.7–36.8 (36.1)

20.6–23.5 (22.1)

9.7–11.4 (10.4)

15.4–19.9 (17.9)

8.5–9.8 (9.1)

10.7–12.2 (11.3)

6.8–8.4 (7.6)

6.5–8.1 (7.1)

10.7–11.9 (11.4)

26.2–30.0 (28.1)

12.4–15.6 (14.3)

22.9–25.6 (24.2)

65.1
—

32.8

35.5

26.1

10.1

18.4

9.0

11.2

7.2

6.1

12.2

28.4

15.4

24.4

7+18=25

3–4 (4)—
6–7 (6)+15–18 (17)=21–25 (24)2
5–5.5 (5)—28–29 (29)

I, 7

I, 5

13–15 (13)VII, 996.3–133.9

Indo–West PacificNon-types
n=12

30
—

I, 7

I, 5

14

VII, 9127.5

Madagascar
Holotype of U. crosnieri
MNHN 1965-17

Upeneus guttatus

3–4 (3)

5–5.5 (5.5)

2

29–30 (29)

I, 7

I, 5

13–15 (14)VII, 974.7–139.0

7.3–10.4 (9.1)
13.6–16.3 (15.0)—

10.9–15.4 (13.5)

14.7–18.6 (16.9)

16.8–21.8 (19.4)

17.7–23.9 (21.2)

30.7–33.2 (31.9)

60.0–66.1 (62.7)

34.7–38.8 (36.4)

21.0–25.9 (23.7)

8.4–11.1 (10.1)

18.9–25.1 (22.1)1

7.8–9.6 (8.6)

10.1–11.9 (10.9)

7.0–8.4 (7.5)

7.2–9.5 (8.3)

10.6–13.1 (11.6)

14.9–31.3 (28.4)

14.0–17.7 (15.6)

22.6–26.3 (24.5)

12.6
—

17.0

18.0

31.3
—

37.1
—

23.7

10.2

20.5

8.6

10.5

7.9

7.4

11.2

29.4

13.7

24.9

7+18=25

—

—

29
—

I, 7

I, 5

14

VII, 996.5

East Asia
Non-types
n=22

6–7 (7)+17–20 (18)
=24–27 (25)1

Upeneus japonicus
Japan
Neotype of M. japonicus
NSMT-P 45177

Counts and measurements of specimens of Upeneus guttatus and U. japonicus, expressed as percentages of standard length with means in parentheses.

Standard length (SL, mm)

Table 1.

The goatfish Upeneus guttatus from Japan11

0.5–1.6 (1.1)

8.4–10.5 (9.1)

12.9–15.1 (14.4)

Anal-fin spine length

First anal-fin soft ray length

Second anal-fin soft ray length

13.0–16.4 (15.0)

19.1–22.5 (20.3)

Pelvic-fin spine length

Longest pelvic-fin ray length

—

18.5

14.3

1

1

12

24N

22S

8

15

7

6
24

4S
4

17

16
16N

6

18

Lower limb

11

19

19.5

14.5

20
1

1

21

5S

22

18.5–21.7 (20.1)

14.2–16.1 (14.9)

19.1–21.4 (20.1)

13.9–18.1 (16.1)

—
18.2

—
18.9

6.8–9.7 (8.0)
23.2–30.2 (27.6)

7.9–10.4 (9.0)
13.3–15.7 (14.8)—

9.3

0.7–1.5 (1.0)

5.7–8.8 (7.5)

6.1–7.3 (6.6)

7.7—

S Includes data for syntype of Upeneoides guttatus; N includes data for neotype of Mullus japonicus.

U. japonicus

5Upper limb

6.7
7.5
—
15.0
—

14.0

8.2

1.0

7.6

Frequency distribution of gill-raker counts of Upeneus guttatus and U. japonicus.

U. guttatus

Table 2.

1 Based on 32 specimens.

14.6–18.1 (16.5)

19.6–22.6 (20.7)

Caudal-concavity length

Pectoral-fin length

25.4–29.8 (28.2)

Caudal-fin length

7.2–9.4 (8.2)

6.5-8.5 (7.3)

Last anal-fin soft ray length

6.2–7.9 (7.0)

Eighth dorsal-fin soft ray length

Ninth dorsal-fin soft ray length

13

23

—

7

14

24

Total

19.7

13.9

24.5

25
2
15N

8

26

17.5–22.5 (20.0)

12.0–16.3 (14.1)

21.3–26.0 (23.5)1

11.7–15.1 (13.1)

22.5–30.7 (25.6)

7.4–10.0 (8.4)

—
8.7
—

7.4–11.2 (9.5)
12.9–16.3 (14.6)—

6.0–8.4 (7.3)
0.5–1.1 (0.9)

5.8–8.1 (7.0)

7.6
—

7.5

1

27

12
Hiroyuki Motomura et al.

The goatfish Upeneus guttatus from Japan

tatus on the basis of Indian Ocean specimens and distinguished it from other members of the Indo-Pacific U. japonicus species group by having a reddish body without adistinct lateral stripe, 12–14 (mostly 13) pectoral-fin rays,
28–31 lateral-line scales, 23–25 gill rakers, the color patternof the caudal-fin lower lobe, and several morphometric features. Characters of the present specimens from Kagoshima,
Japan, agree with those of U. guttatus given by Uiblein andHeemstra (2010, 2011b), with the exception of the gill raker
count (22–24, but only one specimen with 22 rakers; Table
2).
Upeneus guttatus is similar to a co-occurring species inJapanese waters, U. japonicus, in sharing the presence ofteeth on the ectopterygoids (Kim and Nakaya 2002; thisstudy) and a reddish body coloration. The two species havebeen confused; Day (1875) and Lachner (1954) mistakenly reported U. guttatus as a junior synonym of U. bensasi (Temminck and Schlegel, 1843) (currently regarded asa junior synonym of U. japonicus; see Randall et al. 1993).
Uiblein and Heemstra (2010) mentioned that U. guttatus
differs from U. japonicus in having “fewer gill rakers, shorterbarbels, shorter pectoral fins, and bars present along ventral
margin of lower caudal-fin lobe in fresh fish”. However, they
did not give concrete values of meristics and morphometrics
of U. japonicus. Detailed comparisons between the two species in this study show that Uiblein and Heemstra’s (2010)
purported differences between the two species are accurate.
Upeneus guttatus has 5–7 (mode 6) gill rakers on the upper
limb, 15–18 (17) rakers on the lower limb, and 21–25 (24)rakers in total whereas U. japonicus has 6 or 7 (7), 17–20(18), and 24–27 (25), respectively (Table 2). Barbels of U.
guttatus tend to be shorter [15.4–20.2% of SL (mean 18.2%)]
than those of U. japonicus [18.9–25.1% (22.1%)] (Table 1;
Fig. 3A). While the posterior tips of the depressed barbelsof U. japonicus extend well beyond a vertical through thepreopercular margin, those of U. guttatus usually just reachthe margin. Pectoral-fin length of U. guttatus [18.2–22.6% of
SL (mean 20.4%)] is also shorter than that of U. japonicus[21.3–26.0% (23.5%)] (Table 1; Fig. 3B).
Coloration of the lower caudal-fin lobe of U. guttatus differs from that of U. japonicus when the fish are fresh. The
ventral half of the lower lobe of U. guttatus is whitish withirregular red bars and/or poorly defined spots (bars andspots distinct when young, becoming more irregular withgrowth; Fig. 2), and the upper half of the lobe is dark red
without any markings (Figs. 1A, B, 2A–D). The ventral andposterior margins of the lower lobe of U. japonicus are white
without red bars or spots and the remaining part of the lobe
is uniformly red with a narrow black margin posteroventrally (Figs. 1C, D).
Although Uiblein and Heemstra (2011b) showed geographic variation in morphology and color patterns of U.
guttatus within the Indian Ocean, detailed population data
for this species in the western Pacific Ocean are unavailable and cannot therefore be compared with data from theJapanese population. All the Japanese specimens examinedin this study had yellow barbels, whereas specimens fromthe Indian Ocean had white (2 specimens) and yellow (1313specimens) barbels (Uiblein and Heemstra 2011b). Examination of color photographs of 19 specimens taken whenfresh shows that the upper caudal-fin lobe of Japanese U.
guttatus has 4 or 5 red bars reaching to the fin margin andan indistinct red bar close to the fin base (not reaching to
the margin) (Figs. 1A, B, 2). In contrast, the upper lobe of U.
guttatus from the Gulf of Suez has 3 and 1 bars respectively,
and in fish from the Indian Ocean it has 4 and 1 bars respectively (Uiblein and Heemstra 2011b). The number of reddish
bars on the ventral margin of the caudal-fin lower lobe inJapanese U. guttatus is 4–9 whereas that of conspecifics fromthe Seychelles Bank is 2–5, and 5–8 in fish from the Gulf ofSuez and Chennai (Uiblein and Heemstra 2011b). There issome possibility that the Japanese specimens of U. guttatus
represent a separate species or a distinct geographic population of U. guttatus; further research on specimens from thewestern Pacific is required.

Acknowledgements
We thank M. McGrouther, A. Hay, and S. Reader (AMS);
A. Suzumoto (BPBM); S. Kimura (FRLM); R. Causse, P.
Pruvost, and P. Béarez (MNHN); G. Shinohara (NSMT);K. Hoshino (SNFR); D. Nelson (UMMZ); and J. Williams(USNM) for opportunities to examine comparative materials. We are especially grateful to volunteers and students of
KAUM for their curatorial assistance and to staff of the Kagoshima Aquarium for providing some specimens of U. guttatus. We greatly appreciate comments on the manuscript by
G. Yearsley (Hobart, Australia). This study was conducted aspart of the Kagoshima Fish Diversity Project (KAUM).

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