Species Diversity 18: 193–213
25 November 2013
DOI: 10.12782/sd.18.2.193

Three species of Western Pacific Chromis193

Redescriptions of Chromis notata (Temminck and Schlegel,1843) and C. kennensis Whitley, 1964 with the
Description of a New Species of Chromis(Perciformes: Pomacentridae)
Hiroki Iwatsubo1 and Hiroyuki Motomura2,3
1 Graduate School of Fisheries, Kagoshima University, 4-50-20 Shimoarata, Kagoshima 890-0056, Japan
2 The Kagoshima University Museum, 1-21-30 Korimoto, Kagoshima 890-0065, Japan

E-mail: motomura@kaum.kagoshima-u.ac.jp
3 Corresponding author(Received 5 March 2013; Accepted 18 November 2013)
Two species of Chromis, C. notata (Temminck and Schlegel, 1843) and C. kennensis Whitley, 1964, are redescribed, andthe description of a new species, C. yamakawai sp. nov., is provided. Two nominal species that were previously regarded asvalid, C. flavomaculata Kamohara, 1960 and C. miyakeensis Moyer and Ida, 1976, are synonymized with C. notata. Chromis
villadolidi Jordan and Tanaka, 1927 is confirmed as a junior synonym of C. notata. Geographic variation of C. notata is discussed. Two populations previously treated as C. flavomaculata (regarded as a junior synonym of C. notata in this study) include two geographically disjunct species: C. kennensis, previously regarded as a junior synonym of C. flavomaculata, fromthe southwestern Pacific Ocean, and C. yamakawai from the northwestern Pacific Ocean.
Key Words: Western Pacific Ocean, damselfishes, synonymy, taxonomy, Chromis miyakeensis, Chromis villadolidi,
Chromis flavomaculata.

Introduction
Fishes of the genus Chromis Cuvier, 1814, the most speciose genus in the family Pomacentridae, are common oncoral and rocky reefs throughout tropical and warm temperate seas (Allen 1991). Eighty-eight species of Chromis
are currently recognized as valid (Allen 1991; Randall and
McCosker 1992; Randall 1994; de Moura 1995; Randall
2001; Allen and Randall 2004; Lecchini and Williams 2004;
Allen and Erdmann 2005, 2008, 2009; Senou and Kudo
2007). The genus is characterized by having the following:
an elliptical and moderately deep body, its depth 53–62%
of standard length; a short snout; biserial teeth including anouter row of enlarged conical teeth; a fully scaled head, except for the tip of the snout and the area around the nasal
openings; a dorsal fin with 12–15 spines and 10–15 soft rays;
an anal fin with two spines and 10–14 soft rays; a caudal fin
with 15 principal rays and five or six procurrent rays, thefirst two or three of the latter being spiniform; and less than
30 scale rows in the longitudinal series (Randall et al. 1981;
Allen 1991).
Chromis notata notata (Temminck and Schlegel, 1843),
originally described as Heliases notatus, and C. notata
miyakeensis Moyer and Ida, 1976, originally described as
C. miyakeensis, have been regarded as two valid subspecies by many authors (e.g., Ida 1984; Aonuma and Yoshino
1993, 2000). Randall et al. (1981) were the first to treat C.
miyakeensis as a subspecies, albeit somewhat tentatively.
They characterized it as having a deep body, but stated that
more specimens should be examined to assess the validity ofthis feature. Since Randall et al. (1981), no taxonomic treatment of either C. notata notata or C. notata miyakeensis has
been reported.
We examined a large number of specimens of C. notata
notata from a wide distributional range and concluded thatthey represented three indistinct geographic populations,with C. miyakeensis being a junior synonym of C. notata.
Chromis villadolidi Jordan and Tanaka, 1927 was also confirmed in this study to be a junior synonym of C. notata.
Chromis flavomaculata Kamohara, 1960 (type locality:
Kochi, Japan), closely related to C. notata (see Randall etal. 1981), has been regarded as a valid species with an antiequatorial distribution in the northwestern and southwestern Pacific Ocean (Randall et al. 1981, 1997; Randall 2005;
Allen and Erdmann 2012). Examination of the holotype of
C. flavomaculata in this study showed it to be identical with
C. notata, and the two anti-equatorial populations were
found to differ from each other in several morphologicalfeatures. The northern population, previously identified as
C. flavomaculata, is herein described as a new species. The
name Chromis kennensis Whitley, 1964 (type locality: Kenn
Reef, Coral Sea), previously regarded as a junior synonym of
C. flavomaculata (e.g., Randall et al. 1981; Allen 1991; Randall 2005; Allen and Erdmann 2012), should be applied tothe southern population.
This paper provides redescriptions of C. notata (as a senior synonym of C. villadolidi, C. miyakeensis, and C.
© 2013 The Japanese Society of Systematic Zoology

194

H. Iwatsubo and H. Motomura

flavomaculata) from East Asia and C. kennensis (a resurrected name from the southwestern Pacific Ocean), and thedescription of C. yamakawai sp. nov. from the northwestern
Pacific Ocean.

Materials and Methods
Counts and measurements follow Hubbs and Lagler(1947); observations were made on the left side of the body
if possible. Measurements of the basal widths of the pelvic
fin axillary scale and the scale between the pelvic fin bases
are shown in Fig. 1. Standard length is abbreviated as SL.
Gill rakers were counted on the first gill arch. Osteological characters were confirmed from X-ray photographs (33specimens of C. notata, five of C. kennensis, and nine of C.
yamakawai). Counts and proportional measurements, as
percentages of SL, of the specimens of Chromis notata are
given in Tables 1, 2, and 5, and those of C. kennensis and C.
yamakawai are given in Table 4. Frequency distributions of
selected meristic characters of the three species are presented in Table 3.
The specimens examined in this study are deposited at
the Australian Museum, Sydney (AMS); Bishop Museum,
Honolulu (BPBM); Department of Biology, Faculty of Science, Kochi University, Kochi (BSKU); California Academy of Sciences, San Francisco (CAS); Kyoto University

Fig. 1.
Measurements of basal widths of scales. A, pelvic fin axillary scale, lateral view; B, scale between bases of pelvic fins, ventral view.

Museum, Kyoto (FAKU); Fisheries Research Laboratory,
Mie University, Shima (FRLM); Hiwa Museum for Natural History, Shobara (HMNH); Hokkaido University Museum, Hakodate (HUMZ); Kagoshima University Museum,Kagoshima (KAUM); Kanagawa Prefectural Museum of
Natural History, Odawara (KPM); Kochi Prefectural Kochi
Nishi Senior High School, Kochi (KSHS; currently deposited at NSMT); Division of Fisheries Sciences, University ofMiyazaki, Miyazaki (MUFS); National Museum of Natureand Science, Tsukuba (NSMT); Osaka Museum of Natural History, Osaka (OMNH); Naturalis Biodiversity Center, Leiden (RMNH); Seikai National Fisheries ResearchInstitute, Nagasaki (SNFR); Churashima Research Center,
Okinawa Churashima Foundation, Motobu (formerly University of the Ryukyus: URM); Museum Support Center of
the National Museum of Natural History, Smithsonian Institution, Suitland, MD (USNM); Western Australian Museum, Welshpool (WAM); Humboldt-Universität, Museum
für Naturkunde, Zoologisches Museum, Berlin (ZMB); andUniversity Museum, University of Tokyo, Tokyo (ZUMT).
Chromis notata (Temminck and Schlegel, 1843)[Standard Japanese name: Suzumedai][English name: Pearl-spot Chromis](Figs 2–4, 6, 7, 12A–D; Tables 1–3, 5)
Heliases notatus Temminck and Schlegel, 1843: 66 (type locality: Nagasaki, Japan).
Heliastes notatus: Günther 1862: 63 (China; Sea of Japan).
Chromis notatus: Jordan and Snyder 1902: 599 (Tokyo; Misaki; Enoshima; Kobe; Wakanoura; Hiroshima; Tsushima;
Nagasaki, Japan); Schmidt 1930: 66 (in part; Kominato
and Daikuma, Amami-oshima island; Itoman, Okinawa);
Okada 1938: 200 (in part; Honshu and Shikoku, Japan);Kamohara 1957: 38 (in part; middle part of Japan to
Ryukyu Islands; Korea; China); Kamohara 1960: 4 (Chiba
and Niigata, Japan; Korea; China); Masuda et al. 1975: 96,285, fig. D on p. 96 (southern Japan).
Chromis villadolidi Jordan and Tanaka, 1927: 387, pl. 34, fig.
1 (type locality: Sea of Japan between Tsushima and Fukuoka).
Chromis notata: Kamohara 1930: 483 (Kashiwa-jima island,
Kochi, Japan); Tanaka 1931: 31 (southern Japan); Boeseman 1947: 71 (Nagasaki, Japan); Kamohara 1963: 34, pl.
34, fig. 1 (Chiba and Niigata, Japan; Korea; East China
Sea); Masuda et al. 1980: 96, 285, fig. D on p. 96 (southern Japan); Allen 1991: 77, 240, unnumbered fig. (southern Japan, including Ryukyu Islands; Taiwan; China);
Kuiter and Debelius 2006: 530, unnumbered fig. (Japan);
Yoshino and Senou 2008: 265, unnumbered figs (Akita
and southwards in Japan, including Izu and Ryukyu
islands; Korean Peninsula; China; Taiwan); Kato 2011b:
70 (Akita and southwards; Izu Islands; Ryukyu Islands;
Korean Peninsula; China; Taiwan); Allen et al. 2012: 80,
unnumbered fig. (southern Japan to Taiwan).
Chromis flavomaculatus Kamohara, 1960: 5, pl. 1, fig. 2 (typelocality: Susaki, Kochi, Japan).
Chromis miyakeensis Moyer and Ida, 1976: 189, fig. 1 (typeThree species of Western Pacific Chromis195

Table 1.

Counts and measurements, expressed as percentages of standard length (SL), of type specimens now identified as Chromis notata.
C. notata
Lectotype
RMNH.PISC.
895

Standard length (mm)Counts
Dorsal fin rays
Anal fin rays
Pectoral fin rays
Pelvic fin rays
Spiniform caudal fin rays
Principal caudal fin rays
Tubed lateral line scales
Pored scales on caudal peduncle
Scale rows in longitudinal series
Scale rows above/below lateral line
Circumpeduncular scales
Gill rakers
Measurements (% SL)
Head length
Snout length
Body depth
Body width
Orbit diameter
Interorbital width
Caudal peduncle depth
Caudal peduncle length
Upper jaw lengthPre-dorsal-fin length
Spinous dorsal fin base length
Soft-rayed dorsal fin base length
1st dorsal fin spine length
2nd dorsal fin spine length
3rd dorsal fin spine length
5th dorsal fin spine length
7th dorsal fin spine length
Last dorsal fin spine length
1st dorsal fin soft ray length
Longest dorsal fin soft ray length
Last dorsal fin soft ray lengthPre-anal-fin length
Anal fin base length
1st anal fin spine length
2nd anal fin spine length
1st anal fin soft ray length
Longest anal fin soft ray length
Last anal fin soft ray length
Caudal fin length
Caudal concavity
Pectoral fin length
Pre-pelvic-fin length
Pelvic fin spine length
1st pelvic fin soft ray length
Last pelvic fin soft ray length
Basal width of pelvic fin axillary scale
Basal width of scale between pelvic fin bases

Paralectotypes
RMNH.PISC.
895
Nagasaki, JapanZMB 2792

C. villadolidi

C. miyakeensis

Holotype

Holotype

FMNH 59185

ZUMT 53957

Tsushima, Japan

Miyake-jima, Japan

86.8

69.2

88.8

100.8

126.3

XIII, 13II, 10
19
I, 5
2
8+7
18
9
28
4/12
14
8+23=31

XIII, 12II, 10
19
I, 5
2
8+7
18
9
28
4/11
14
8+22=30

XIII, 13II, 10
19
I, 5
2
8+7
18
9
26
4/11
14
9+21=30

XII, 12II, 10
19
I, 5
2
8+7
18
8
30
5/11
14
9+23=32

XIII, 13II, 11
20
I, 5
2
8+7
18
8
27
4/11
14
—

29.5
7.4
47.8
16.7
9.8
10.6
14.9
17.7
10.0
35.9
47.0
16.8
8.2
12.0
14.3
15.1
—
11.5
—
—
10.6
64.1
20.3
—
—
—
—
—
—
—
33.3
41.5
14.1
31.7
13.9
—
—

30.9
8.4
43.5
14.0
11.4
10.5
14.7
18.9
10.8
40.0
45.5
15.5
9.8
—
15.9
—
15.9
12.7
—
—
9.2
62.1
20.7
7.8
18.5
18.8
21.0
8.8
—
—
31.5
40.3
17.1
31.4
15.0
—
—

30.3
7.9
43.5
17.2
10.6
10.0
14.3
16.9
9.7
37.3
47.5
16.8
9.3
12.2
14.5
15.4
16.0
12.6
—
23.5
9.6
71.6
20.9
7.1
17.1
—
—
12.0
34.5
17.1
—
42.0
16.0
28.5
15.2
—
—

29.4
7.9
39.2
16.2
11.0
10.4
14.2
17.6
10.7
37.7
43.3
14.1
10.1
12.4
16.9
18.2
16.5
14.3
16.6
21.7
6.9
63.9
18.8
7.2
18.3
20.2
20.7
8.4
33.5
15.9
32.8
41.9
16.1
29.2
15.9
—
—

29.8
9.3
50.4
21.0
8.4
11.3
16.3
18.4
9.9
38.2
46.6
17.7
8.9
11.2
14.6
17.5
17.6
15.8
17.7
27.6
7.4
60.9
22.2
6.3
19.0
22.4
24.9
9.9
17.7
35.8
29.9
41.6
15.0
24.7
15.6
2.2
3.5

196

H. Iwatsubo and H. Motomura

Table 2. Counts and measurements, expressed as percentages of standard length (SL), of non-type specimens of Chromis notata. Modes ormeans in parentheses.
Chromis notata

Standard length (mm)Counts
Dorsal fin rays
Anal fin rays
Pectoral fin rays
Pelvic fin rays
Spiniform caudal fin rays
Principal caudal fin rays
Tubed lateral line scales
Pored scales on caudal peduncle
Scale rows in longitudinal series
Scale rows above / below lateral line
Circumpeduncular scales
Gill rakers
Measurements (% SL)
Head length
Snout length
Body depth
Body width
Orbit diameter
Interorbital width
Caudal peduncle depth
Caudal peduncle length
Upper jaw lengthPre-dorsal-fin length
Spinous dorsal fin base length
Soft-rayed dorsal fin base length
1st dorsal fin spine length
2nd dorsal fin spine length
3rd dorsal fin spine length
5th dorsal fin spine length
7th dorsal fin spine length
Last dorsal fin spine length
1st dorsal fin soft ray length
Longest dorsal fin soft ray length
Last dorsal fin soft ray lengthPre-anal-fin length
Anal fin base length
1st anal fin spine length
2nd anal fin spine length
1st anal fin soft ray length
Longest anal fin soft ray length
Last anal fin soft ray length
Caudal fin length
Caudal concavity
Pectoral fin length
Pre-pelvic-fin length
Pelvic fin spine length
1st pelvic fin soft ray length
Last pelvic fin soft ray length
Basal width of pelvic fin axillary scale
Basal width of scale between pelvic fin bases

Sea of Japann

Seto Inland Sea
plus East China
Sean

Pacific coast ofJapan plus
southern Chinan

Izu Islandsn20.4–113.8

44

33.7–110.5

82

26.7–122.3

71

65.9–128.8

5

XIII–XIV (XIII),
12–14 (13)II, 10–12 (11)
18–20 (19)
I, 5
2
8+7
16–19 (17)
8–9 (8)24–28 (26)
4/11–12 (11)14
7–9 (8)+20–24(22)=28–33 (31)42

XIII, 12–14 (13)75

XIII, 11–14 (13)70

XIII, 12–13 (12)

5

42
42
42
42
42
35
23
24
33
29
33

II, 10–12 (11)
16–20 (19)
I, 5
2
8+7
16–19 (17)
7–10 (8)24–27 (26)
4/11–12 (11)14
7–10 (9)+20–24(20)=27–33 (28)75
II, 10–12 (11)75
17–20 (19)
75
I, 5
75
2
75
8+7
73
16–21 (17)61
7–10 (8)66
23–28 (26)
70 4–5 (4)/11–12 (11)72
14
74 7–10 (9)+19–24(21)=27–33 (29)71
II, 10–11 (11)
69
19
71
I, 5
71
2
71
8+7
68
17–18 (17)
61
8–9
61
26–28 (27)
64 4–5 (4)/11–12 (11)67
1462
8–9+21–22=
29–31

5
5
5
5
5
5
4
5
5
5
2

40
39
43
43
39
39
42
39
39
38
38
38
37
38
38
38
38
35
28
30
34
38
39
36
38
35
35
36
28
26
35
39
38
28
33
22
30

28.3–33.3 (30.8)
6.7–9.4 (8.0)
41.1–50.7 (45.5)
15.4–23.4 (18.2)
8.4–11.6 (9.8)
9.4–12.1 (10.6)
13.3–15.9 (14.4)
11.5–20.4 (16.9)
8.4–11.3 (9.7)
30.6–41.0 (37.3)
41.9–51.9 (46.0)
12.8–20.3 (15.7)
7.0–12.0 (9.4)
9.2–16.6 (13.6)
12.7–18.9 (15.7)
14.0–19.5 (16.8)
13.7–19.3 (16.5)
11.1–18.0 (14.0)
12.2–21.3 (15.6)
20.6–31.5 (24.0)
4.5–14.5 (7.6)
58.0–74.8 (63.8)
17.3–22.4 (20.2)
4.9–8.5 (6.8)
14.8–21.0 (18.2)
14.1–22.9 (19.3)
17.2–28.1 (21.2)
5.0–12.5 (8.1)
14.0–40.6 (33.9)
12.7–39.4 (19.6)
27.1–42.0 (31.1)
28.1–48.1 (42.1)
13.7–18.8 (16.1)
15.3–33.2 (29.1)
11.0–17.3 (13.3)
1.5–3.2 (2.1)
2.0–3.7 (2.7)

73
73
82
73
73
73
73
73
73
73
73
73
72
73
72
71
72
73
69
72
71
73
73
73
73
72
70
72
71
70
73
72
73
68
73
31
34

71
70
70
70
69
69
71
69
67
69
69
69
69
69
69
69
69
67
65
65
68
70
65
67
70
67
64
67
63
61
70
70
70
69
68
44
48

5
5
5
5
5
5
5
5
5
5
5
5
5
5
5
5
5
5
3
4
5
5
5
5
5
3
4
4
3
4
5
5
5
5
5
4
5

28.9–33.9 (31.2)
6.5–9.3 (7.7)
37.8–49.9 (44.0)
14.1–24.6 (17.2)
8.9–13.2 (10.5)
9.3–11.4 (10.4)
12.5–15.7 (14.1)
9.5–20.6 (17.8)
8.8–12.1 (9.9)
34.5–40.2 (37.7)
41.1–49.0 (44.9)
11.3–18.0 (14.8)
7.4–12.5 (9.9)
10.9–17.9 (14.4)
12.4–19.7 (16.5)
14.5–21.0 (17.4)
13.6–19.7 (16.6)
9.7–16.1 (13.4)
10.7–18.0 (14.6)
18.8–26.0 (22.3)
5.0–10.2 (7.4)
58.5–71.5 (66.0)
16.3–21.9 (19.6)
5.2–11.3 (7.3)
16.1–22.1 (18.4)
14.1–23.1 (18.6)
15.6–24.5 (19.9)
5.8–10.8 (8.0)
28.7–38.4 (33.6)
12.0–21.5 (16.2)
20.1–33.2 (29.2)
38.1–47.6 (42.2)
13.7–18.7 (16.2)
21.7–30.9 (26.9)
10.7–15.5 (13.0)
1.5–2.6 (2.0)
2.0–3.6 (2.7)

28.5–37.3 (31.2)
5.7–9.5 (7.9)
42.0–51.5 (48.0)
15.4–23.4 (19.5)
8.4–11.3 (10.1)
9.3–11.8 (10.6)
11.2–15.8 (14.8)
9.8–20.0 (15.3)
8.0–11.1 (9.7)
33.8–46.3 (38.4)
43.9–51.9 (47.0)
12.6–20.4 (15.6)
4.4–12.5 (9.1)
9.5–17.8 (13.3)
11.9–18.2 (15.0)
13.3–20.1 (16.2)
11.2–19.2 (15.8)
9.4–18.8 (13.3)
10.5–22.2 (15.3)
18.6–33.1 (24.1)
4.2–9.9 (7.1)
59.0–73.4 (63.3)
16.6–23.6 (19.9)
4.5–10.2 (6.3)
12.7–21.7 (17.6)
14.1–25.8 (18.6)
17.3–26.1 (21.4)
4.5–11.0 (7.4)
28.4–43.9 (36.5)
10.8–26.1 (19.4)
17.0–35.0 (30.7)
38.2–49.9 (42.6)
10.9–19.3 (16.2)
17.8–33.0 (28.9)
9.7–16.0 (12.6)
1.5–2.6 (2.0)
2.2–3.7 (2.8)

30.0–32.3 (31.2)
7.4–8.3 (7.8)
45.8–52.3 (49.6)
17.0–20.0 (18.4)
9.0–12.1 (10.2)
9.7–11.2 (10.5)
14.4–15.9 (14.9)
12.1–19.5 (17.7)
8.9–10.6 (9.5)
36.1–39.2 (38.3)
36.8–51.3 (45.8)
14.6–16.5 (15.5)
8.8–11.4 (9.9)
13.0–17.0 (14.5)
15.0–18.5 (16.8)
17.0–19.7 (18.2)
16.9–18.7 (17.5)
13.6–15.9 (14.9)
15.6–21.2 (17.8)
22.0–26.1 (24.4)
6.4–8.7 (7.5)
58.5–64.1 (63.1)
19.1–22.1 (20.9)
5.9–8.5 (7.2)
17.5–23.4 (20.1)
17.9–23.1 (20.2)
19.7–25.2 (22.1)
6.4–9.1 (7.7)
33.6–36.1 (35.0)
11.5–17.6 (14.6)
31.5–35.5 (32.9)
40.5–45.6 (43.0)
16.8–19.3 (17.6)
20.7–34.1 (27.9)
12.0–16.5 (13.8)
1.9–2.5 (2.1)
2.3–3.5 (2.8)

Three species of Western Pacific Chromis197

Table 3.

Frequency distributions of selected counts of Chromis notata, C. kennensis, and C. yamakawai sp. nov.
n

C. notataSea of Japan
Seto Inland Sea plus East China Sea
Pacific coast of Japan plus southern China
Izu Islands
C. kennensis
C. yamakawai sp. nov.

42
78
70
5
22
105
n

C. notataSea of Japan
Seto Inland Sea plus East China Sea
Pacific coast of Japan plus southern China
Izu Islands
C. kennensis
C. yamakawai sp. nov.

42
78
69
5
16
96
n

C. notataSea of Japan
Seto Inland Sea plus East China Sea
Pacific coast of Japan plus southern China
Izu Islands
C. kennensis
C. yamakawai sp. nov.

35
75
68
5
17
78
n

C. notataSea of Japan
Seto Inland Sea plus East China Sea
Pacific coast of Japan plus southern China
Izu Islands
C. kennensis
C. yamakawai sp. nov.

24
68
61
5
12
35
n

C. notataSea of Japan
Seto Inland Sea plus East China Sea
Pacific coast of Japan plus southern China
Izu Islands
C. kennensis
C. yamakawai sp. nov.

33
72
64
5
14
94

Dorsal fin soft rays
11

12

13

14

—
—
1
—
5
24

10
10
12
3
16H
80H

30
61L
54
2
1
1

2
7
3
—
—
—

Anal fin soft rays

n

42
78
71
5
22
105

10

11

12

11
7L
2
1
6
13

27
65
56
3
15H
89H

4
6
13
1
1
3

Pectoral fin rays
16

17

18

19

20

21

—
2
—
—
—
—

—
3
1
—
1
2

9
10
6
—
—
4

32
59L
59
5
12H
67H

1
4
3
—
3
22

—
—
—
—
—
1

Tubed lateral line scales
15

16

17

18

19

20

21

—
—
—
—
1
—

4
7
8
—
2
2

19
40
30
3
6H
14

9
25L
24
2
6
26H

3
3
4
—
2
33

—
—
1
—
—
2

—
—
1
—
—
1

Scale rows in longitudinal series
21

22

23

24

25

26

27

28

—
—
—
—
—
4

—
—
—
—
—
—

—
—
1
—
—
2

2
3
1
—
—
1

2
8
15
—
2
4

10
37
34
1
9H
16H

9
18
6
3
1
8

1
2L
4
1
—
—

Scale rows above lateral line
3

4

5

—
—
—
—
14H
94H

33
72L
61
4
—
—

—
—
3
1
—
—

n

33
72
64
5
14
91

Scale rows below lateral line
9

10

11

12

—
—
—
—
13H
83H

—
—
—
—
1
8

27
57
63
4
—
—

6
15L
1
1
—
—

198

Table 3.

H. Iwatsubo and H. Motomura

ContinuednC. notataSea of Japan
Seto Inland Sea plus East China Sea
Pacific coast of Japan plus southern China
Izu Islands
C. kennensis
C. yamakawai sp. nov.

33
76
62
2
18
91
n

C. notataSea of Japan
Seto Inland Sea plus East China Sea
Pacific coast of Japan plus southern China
Izu Islands
C. kennensis
C. yamakawai sp. nov.

33
76
62
2
18
91

Gill rakers on upper limb

Gill rakers on lower limb

6

7

8

9

10

19

20

21

22

23

24

25

—
—
—
—
—
2

5
24
6
—
2
41

17
36L
22
1
10
34H

11
16
32
1
6H
14

—
—
2
—
—
—

1
—
4
—
—
—

8
21
10
—
—
6

8
17
30
1
1
10

7
21
10
1
3
27

9
15L
7
—
6H
34H

—
2
1
—
7
13

—
—
—
—
1
1

Total gill rakers
27

28

29

30

31

32

33

34

—
10
4
—
—
5

3
16
8
—
—
6

6
13
20
1
1
14

10
15
16
—
3
28

11
14L
5
1
4
27H

2
7
7
—
8H
10

1
1
2
—
1
1

—
—
—
—
1
—

H including data for holotypes of Chromis kennensis and C. yamakawai. L including data for lectotype of Chromis notata.

locality: Igaya Bay, Miyake-jima island, Japan).
Chromis notata notata: Randall et al. 1981: 231, fig. 15 (central Japan to Hong Kong); Kinoshita 1988: 561 (Akita andsouthwards in Sea of Japan; Chiba and southwards along
Pacific coast of Japan; East China Sea); Aonuma andYoshino 1993: 810, unnumbered fig. (Akita and southwards in Sea of Japan; Chiba and southwards along Pacific coast of Japan; East China Sea); Masuda and Kobayashi
1994: 222, fig. 1 (Akita and southwards in Sea of Japan;
Chiba and southwards along Pacific coast of Japan; East
China Sea); Allen 2000: 626 (South China Sea); Ao­numa
and Yoshino 2000: 925, unnumbered fig. (Akita andsouthwards in Sea of Japan; Chiba and southwards along
Pacific coast of Japan; East China Sea); Hirata et al. 2001:
70, unnumbered fig. (Iburi, Kochi, Japan); Ao­numa andYoshino 2002: 925, unnumbered fig. (Akita and southwards in Sea of Japan; Chiba and southwards along Pacific coast of Japan; East China Sea); Konishi 2007: 238,
unnumbered fig. (Akita and southwards in Sea of Japan;
Chiba and southwards along Pacific coast of Japan; East
China Sea); Nishida et al. 2007: 70 (Fukutsu, Fuku­oka,
Japan); Masuda 2008: 391 (Nagahama, Maizuru, Kyoto,
Japan); Araga 2009: 440, unnumbered fig. (Akita andsouthwards in Sea of Japan; Chiba and southwards along
Pacific coast of Japan; East China Sea); Motomura et al.
2010: 149 (Yaku-shima island, Kagoshima, Japan); Sakai
et al. 2010: 12 (Osaki-kami-shima island, Hiro­shima,
Japan); Shimizu et al. 2010: 45 (off Kurahashi-jima island,
Seto Inland Sea, Japan); Takagi et al. 2010: 78, unnumbered fig. (Ehime, Japan); Tsuji et al. 2010: 10 (Nanao Bay,
Noto Peninsula, Japan).
Chromis notatus notatus: Ida 1984: 192, pl. 185, fig. D (Akita
and southwards in Sea of Japan; Chiba and southwards
along Pacific coast of Japan; East China Sea); Araga et
al. 1985: 252, unnumbered fig. (Akita and southwards in
Sea of Japan; Chiba and southwards along Pacific coast ofJapan; East China Sea); Kinoshita 1988: 559, 561, unnumbered fig. (Akita and southwards in Sea of Japan; Chiba
and southwards along Pacific coast of Japan; East China
Sea).
Chromis notata miyakeensis: Randall et al. 1981: 233, fig. 16(Miyake-jima island, Izu Islands, Japan); Aonuma andYoshino 1993: 810, unnumbered fig. (Miyake-jima island, Izu Islands, Japan); Masuda and Kobayashi 1994:
221, figs 7, 8 (Miyake-jima island, Izu Islands; Izu Peninsula, Japan); Aonuma and Yoshino 2000: 925, unnumbered fig. (Miyake-jima island, Izu Islands; Izu Peninsula,
Japan); Aonuma and Yoshino 2002: 925, unnumbered fig.
(Miyake-jima island, Izu Islands; Izu Peninsula, Japan).
Chromis notatus miyakeensis: Ida 1984: 192, pl. 185, fig. E(Izu Islands).
Lectotype. RMNH.PISC. 895 (1 of 2 specimens),
86.8 mm SL, Nagasaki, Japan, P. F. von Siebold [lectotype
designated by Boeseman (1947)].
Paralectotypes. RMNH.PISC. 895 (1 of 2 specimens),
69.2 mm SL, ZMB 2792, 88.8 mm SL, same data as lectotype.
Other type material examined. BSKU 8482, holotype of C. flavomaculata, 84.8 mm SL, Susaki, Kochi, Japan,
19 May 1959; FMNH 59185, holotype of C. villadolidi,
101.0 mm SL, Sea of Japan between Tsushima and Fuku­
oka, D. Jordan and S. Tanaka; ZUMT 53957, holotype of C.
miyakeensis, 126.3 mm SL, Igaya Bay, Miyake-jima island,
Izu Islands, Japan, J. Moyer and H. Ida, October.
Non-type material examined. Most specimens fromfishing ports were obtained from fishermen; although thesite of capture of such specimens is unknown, in most cases
it should not be far distant from the port. The geographical categorization adopted here reflects the distribution ofmorphologically distinct populations, as is discussed afterwards. SEA OF JAPAN (44 specimens, 20.4–113.8 mm
SL)—AKITA: KAUM-I. 34909, 109.2 mm SL, KAUM-

Three species of Western Pacific Chromis199

Table 4. Counts and measurements, expressed as percentages of standard length (SL), of Chromis kennensis and C. yamakawai sp. nov.
Modes or means in parentheses.
Chromis kennensis

Standard length (mm)Counts
Dorsal fin rays
Anal fin rays
Pectoral fin rays
Pelvic fin rays
Spiniform caudal fin rays
Principal caudal fin rays
Tubed lateral line scales
Pored scales on caudal peduncle
Scale rows in longitudinal series
Scale rows above / below lateral line
Circumpeduncular scales
Gill rakers
Measurements (% SL)
Head length
Snout length
Body depth
Body width
Orbit diameter
Interorbital width
Caudal peduncle depth
Caudal peduncle length
Upper jaw lengthPre-dorsal-fin length
Spinous dorsal fin base length
Soft-rayed dorsal fin base length
1st dorsal fin spine length
2nd dorsal fin spine length
3rd dorsal fin spine length
5th dorsal fin spine length
7th dorsal fin spine length
Last dorsal fin spine length
1st dorsal fin soft ray length
Longest dorsal fin soft ray length
Last dorsal fin soft ray lengthPre-anal-fin length
Anal fin base length
1st anal fin spine length
2nd anal fin spine length
1st anal fin soft ray length
Longest anal fin soft ray length
Last anal fin soft ray length
Caudal fin length
Caudal concavity
Pectoral fin length
Pre-pelvic-fin length
Pelvic fin spine length
1st pelvic fin soft ray length
Last pelvic fin soft ray length
Basal width of pelvic fin axillary scale
Basal width of scale between pelvic fin bases

Holotype

Non-types

AMS IB.
4973
Australia

Southwestern Pacific

77.6
XIII, 11
II, 10
20
I, 5
2
8+7
17
8
24
3/9
14
9+23=32

22.6
6.8
45.1
16.9
9.0
10.2
14.3
17.5
9.4
37.4
45.7
14.3
7.9
9.8
11.6
14.8
15.2
11.1
14.9
19.6
7.9
61.9
20.6
4.8
15.9
—
—
7.3
11.1
33.4
31.1
40.1
16.1
29.0
12.9
1.8
2.5

Chromis yamakawai sp. nov.
Holotype

Paratypesn

NSMT-P
111948
Japan

Northwestern Pacificn22.5–93.0

23

83.0

18.2–130.8

107

XIII–XIV (XIII),
11–13 (12)II, 10–12 (11)17–20 (19)
I, 5
2
8+7
15–19 (18)
8–11 (8)25–27 (26)
3/9–10 (9)14
7–9 (8)+21–25 (24)=
29–34 (32)21

XIII, 12

104

21
15
19
21
21
16
15
11
13
17
17

II, 11
19
I, 5
2
8+7
19
8
26
3/9
14
8+23=31

XIII–XIV (XIII),
11–13 (12)II, 10–12 (11)
17–21 (19)
I, 5
2
8+7
16–21 (19)
7–10 (8)21–27 (26)
3/9–10 (9)14
6–9 (7)+20–25 (23)=
27–33 (30)

28.2–37.3 (31.4)
7.0–8.9 (7.5)
39.0–47.7 (43.4)
12.2–20.6 (16.5)
9.0–13.4 (11.0)
9.1–11.3 (10.3)
13.3–15.5 (14.2)
12.9–20.3 (17.5)
9.1–12.2 (10.1)
36.3–41.1 (38.9)
39.3–46.9 (43.6)
11.7–15.8 (13.8)
6.4–10.2 (8.3)
9.5–14.9 (12.1)
11.6–17.3 (14.7)
10.5–17.8 (15.2)
8.8–17.9 (13.6)
6.0–12.7 (10.0)
7.4–17.7 (13.6)
15.0–21.9 (19.1)
5.5–8.8 (7.6)
59.3–72.8 (64.9)
17.9–20.8 (19.0)
4.8–7.4 (6.1)
15.5–19.9 (18.3)
10.5–21.1 (17.4)
14.2–21.1 (18.4)
6.1–9.1 (7.8)
11.4–23.1 (16.7)
24.0–41.5 (36.0)
26.5–33.5 (30.2)
37.0–44.8 (40.3)
15.4–18.7 (16.5)
15.2–29.0 (26.3)
8.8–15.4 (12.3)
0.6–1.8 (1.5)
2.0–2.8 (2.6)

19
19
19
19
19
19
20
19
19
19
16
16
16
16
16
15
16
16
15
16
16
16
19
18
19
19
19
17
14
14
18
16
15
15
16
15
16

23.8
7.1
44.5
18.1
9.4
10.4
14.1
19.2
9.3
38.3
47.6
15.1
6.6
11.0
14.2
15.9
15.7
11.6
12.8
22.0
7.0
68.0
20.0
5.8
18.1
18.8
18.8
7.1
38.4
22.0
29.8
39.3
16.0
23.1
14.6
2.1
3.6

23.7–36.0 (29.7)
5.3–8.9 (7.4)
36.3–50.6 (43.8)
12.0–26.4 (17.5)
7.9–14.3 (9.9)
8.8–11.9 (10.1)
12.7–15.4 (14.0)
10.8–21.2 (18.0)
7.8–11.4 (9.4)
34.8–43.4 (38.2)
34.4–48.2 (44.4)
10.7–17.6 (14.1)
5.3–13.3 (7.8)
9.1–16.1 (11.8)
9.7–18.2 (14.2)
11.4–18.7 (15.2)
11.4–18.7 (14.6)
7.6–14.6 (11.2)
8.3–20.5 (14.8)
13.2–25.3 (20.6)
4.8–11.9 (7.9)
56.3–73.6 (64.6)
14.4–21.2 (19.1)
3.4–8.6 (6.1)
14.3–22.9 (18.1)
13.8–22.9 (18.7)
13.4–22.9 (18.9)
4.3–10.4 (7.8)
27.9–44.8 (36.0)
11.8–25.9 (20.0)
23.5–35.2 (30.2)
32.1–48.6 (41.5)
13.4–18.3 (15.9)
14.6–31.4 (25.7)
9.1–16.2 (12.8)
1.7–2.8 (2.2)
2.2–3.9 (3.0)

97
95
97
95
95
95
101
95
87
95
95
95
93
93
94
95
92
94
81
82
95
94
93
95
94
89
82
94
83
82
93
95
94
80
93
58
59

104
95
101
102
102
70
63
34
90
67
90

200

Table 5.
(1960).

H. Iwatsubo and H. Motomura

Counts and measurements, expressed as percentages in standard length (SL), of Japanese specimens reported by Kamohara
Kamohara (1960) reported as:

Chromis flavomaculatus

Chromis notata

Chromis notata

Re-identified in this study as:

Chromis notata

Chromis notata

Chromis yamakawai

Standard length (mm)Counts
Dorsal fin rays
Anal fin rays
Pectoral fin rays
Pelvic fin rays
Spiniform caudal fin rays
Principal caudal fin rays
Tubed lateral line scales
Pored scales on caudal peduncle
Scale rows in longitudinal series
Scale rows above / below lateral line
Circumpeduncular scales
Gill rakers
Measurements (% SL)
Head length
Snout length
Body depth
Body width
Orbit diameter
Interorbital width
Caudal peduncle depth
Caudal peduncle length
Upper jaw lengthPre-dorsal-fin length
Spinous dorsal fin base length
Soft-rayed dorsal fin base length
1st dorsal fin spine length
2nd dorsal fin spine length
3rd dorsal fin spine length
5th dorsal fin spine length
7th dorsal fin spine length
Last dorsal fin spine length
1st dorsal fin soft ray length
Longest dorsal fin soft ray length
Last dorsal fin soft ray lengthPre-anal-fin length
Anal fin base length
1st anal fin spine length
2nd anal fin spine length
1st anal fin soft ray length
Longest anal fin soft ray length
Last anal fin soft ray length
Caudal fin length
Caudal concavity
Pectoral fin length
Pre-pelvic-fin length
Pelvic fin spine length
1st pelvic fin soft ray length
Last pelvic fin soft ray length
Basal width of pelvic fin axillary scale
Basal width of scale between pelvic fin bases

Holotype

Non-type

Non-type

Non-type

BSKU 8482
Susaki, Kochi

BSKU 4944
Tachibana, Tokushima

BSKU 4946
Tachibana, Tokushima

BSKU 8513
Susaki, Kochi

84.8

80.9

89.6

113.0

XIII, 13II, 11
18
I, 5
2
8+7
17
8
26
4/12
14
—

XIII, 13II, 11
19
I, 5
2
8+7
17
8
26
4/12
14
—

XIII, 13II, 11
19
I, 5
2
8+7
18
8
25
4/10
14
9+21=30

XIII, 12II, 12
19
I, 5
2
8+7
18
—
—
3/9
14
9+21=30

30.5
8.1
50.0
20.3
10.0
10.6
14.5
17.6
10.4
37.4
44.5
16.9
9.2
13.0
15.1
16.5
15.6
14.4
14.7
24.1
8.1
60.0
18.8
6.3
17.2
17.9
19.1
6.1
36.0
17.0
32.7
40.3
16.2
31.6
11.2
—
3.3

31.5
8.4
46.7
17.4
10.3
10.0
14.2
18.5
9.9
38.6
45.5
14.7
9.5
15.6
17.1
17.1
16.1
14.7
16.4
27.1
6.8
61.3
20.4
6.7
—
19.0
—
7.7
38.7
17.1
33.6
43.8
16.4
30.2
13.0
1.6
2.6

34.4
8.3
45.2
39.5
10.5
10.4
14.3
19.7
10.2
36.1
43.6
15.3
11.0
14.0
17.1
16.6
16.7
14.7
16.0
22.6
8.2
63.4
19.8
8.2
18.8
17.8
18.2
8.1
—
—
28.8
45.8
15.4
26.0
12.0
1.9
2.8

28.7
7.0
42.4
15.9
8.7
9.7
12.7
1.2
7.8
39.5
45.2
13.1
7.3
11.0
13.9
15.5
15.1
12.2
—
21.7
7.7
61.2
19.5
6.6
18.8
18.2
18.2
8.3
—
—
27.5
41.7
15.9
24.5
11.9
2.2
2.7

Three species of Western Pacific Chromis201

I. 34910, 113.8 mm SL, KAUM-I. 34911, 102.8 mm SL,
Togashiohama, Oga, 16 October 2010, spear, S. Chiba.
YAMAGATA: KAUM-I. 26534, 97.7 mm SL, Tobi-shimaisland, Sakata, 39°12′N, 139°30′E, 1996, 0–30 m depth, H.
Motomura; KAUM-I. 31026, 34.0 mm SL, KAUM-I. 31027,
38.3 mm SL, KAUM-I. 31032, 39.4 mm SL, KAUM-I. 31033,
34.0 mm SL, KAUM-I. 31037, 37.6 mm SL, KAUM-I. 31038,
36.9 mm SL, KAUM-I. 31039, 38.1 mm SL, KAUM-I. 31040,
35.7 mm SL, 13 October 1996, H. Ida; NSMT-P 73365, 2
specimens, 82.9–84.5 mm SL, Tobi-shima island, 7–8 November 1936; NSMT-P 73464, 103.5 mm SL, Tobi-shima island, August 1936, H. Umemoto. NIIGATA: FAKU 86796,
107.7 mm SL, FAKU 86797, 96.0 mm SL, FAKU 86798,
105.8 mm SL, FAKU 86799, 102.4 mm SL, Tsutsuishi Fishing Port, Itoigawa, 6 June 2003; HUMZ 78484, 94.8 mm SL,
Mano Bay, Sado Island, 25 November 1978. ISHIKAWA:
KAUM-I. 31042, 56.7 mm SL, KAUM-I. 31043, 54.1 mm
SL, KAUM-I. 31044, 61.0 mm SL, KAUM-I. 31045, 60.4 mm
SL, Kanazawa, 14 October 1982, H. Ida. KYOTO: FAKU
60946, 83.4 mm SL, FAKU 60948, 86.2 mm SL, FAKU
60957, 92.3 mm SL, FAKU 90959, 85.6 mm SL, Wakasa Bay,
10 October 1994; FAKU 131446, 3 of 5 specimens, 89.3–
92.1 mm SL, Mio, Kami, 11 July 1997, Hyogo Prefectural
Fisheries Experiment Station; FAKU W 374, 78.5 mm SL,
Nishi-maizuru Fish Market, 25 August 1978, I. Nakamura.
HYOGO: OMNH-P 4332, 77.1 mm SL, Imagoura, Kasumi,
Kinosaki, 22 July 1990, spear, H. Setoguchi; OMNH-P
7933, 20.4 mm SL, Oura, Takano, Kinosaki, 3 September
1994, hand net, T. Suzuki; OMNH-P 8098, 94.8 mm SL,
Moroyose Fishing Port, Moroyose, Hamasaka, Mikata, 7
February 1996, set net, Uwa Fish Store. SHIMANE: FAKU
38334, 50.3 mm SL, FAKU 38349, 77.1 mm SL, FAKU
38350, 77.2 mm SL, FAKU 38351, 70.8 mm SL, FAKU
38484, 84.4 mm SL, FAKU 38485, 86.7 mm SL, FAKU
38487, 81.6 mm SL, Oki Islands, 1950s; KAUM-I. 31041,
97.6 mm SL, Oki-no-shima island, 23 July 1972, H. Ida.
YAMAGUCHI: HMNH-P 5397, 71.9 mm SL, Susa Fishing
Port, Susa, Hagi, 1 August 2002.
SETO INLAND SEA plus EAST CHINA SEA (82 specimens, 33.7–110.5 mm SL). OSAKA: OMNH-P 13704,
57.7 mm SL, Fuke Fishing Port, Fuke, Sennan-misaki, 4 November 1998, H. Ariyama. WAKAYAMA: KAUM-I 28397,
84.7 mm SL, off Osaki Fishing Port, Shimotsu, Kainan,
34°08′01″N, 135°07′55″E, 6 November 2009, line-fishing,
M. Hanano and Y. Yamana; OMNH-P 9601, 81.2 mm SL,
Oi, Miyazaki, Arida, 8 June 1975, T. Kawasaki; OMNH-P
10512, 76.0 mm SL, OMNH-P 10513, 67.2 mm SL, OMNHP 10514, 74.3 mm SL, Hama, Hatsushima, Arida, 29 August
1975, T. Kawasaki; OMNH-P 26833, 83.2 mm SL, Kada, 29
October 1961, K. Hatooka; OMNH-P 26834, 79.0 mm SL,
OMNH-P 26835, 74.2 mm SL, Kada, 4 November 2004,
K. Hatooka. HYOGO: OMNH-P 13800, 93.6 mm SL,
OMNH-P 13801, 92.4 mm SL, Anaga Fishing Port, Anaga,
Minami-awaji, 4 November 1999, line-fishing, K. Hatooka. OKAYAMA: HUMZ 44820, 85.0 mm SL, Seto, Oda.
HIROSHIMA: HMNH–P 3671, 76.2 mm, HMNH-P 3672,
77.1 mm SL, HMNH-P 3673, 49.9 mm SL, Ta-shima island,
Utsumi, Fukuyama, 15 September 2001; HUMZ 70681,

74.6 mm SL, off Onomichi, August 1930. YAMAGUCHI:
HMNH-P 9839, 69.3 mm SL, HMNH-P 9840, 72.7 mm SL,
Murotsu Fishing Port, Murotsu, Kaminoseki, 8 December 2006. KAGAWA: BSKU 65511, 79.5 mm, BSKU65512,
88.9 mm, BSKU 65569, 73.9 mm SL, BSKU 65570, 75.6 mm
SL, BSKU 65571, 76.8 mm SL, BSKU 65572, 84.9 mm SL,
BSKU 65573, 83.7 mm SL, BSKU 65574, 83.2 mm SL,
BSKU 65581, 85.1 mm SL, BSKU 65582, 82.4 mm SL, BSKU
65583, 78.6 mm SL, BSKU 65584, 75.4 mm SL, BSKU 65585,
75.4 mm SL, No Fishing Port, Sakaide, line-fishing, 26 June,
2003; TOKUSHIMA: BSKU 4944, 80.9 mm SL, Tachibana,24 October 1955; BSKU 4946, 89.6 mm SL, Tachibana, 24
October 1955. EHIME: BSKU 38835, 97.3 mm SL, BSKU
38836, 99.7 mm SL, BSKU 38838, 92.7 mm SL, Yahata­
hama, 6 August 1983; BSKU 39015, 92.8 mm SL, Yahatahama, 8 August 1983; OMNH-P 11229, 92.4 mm SL,
OMNH-P 11230, 91.0 mm SL, OMNH-P 11231, 81.6 mm
SL, OMNH-P 11232, 76.8 mm SL, Okuura, Yoshida, Uwajima, 23 December 1997, line-fishing, K. Hatooka; OMNHP 14077, 2 specimens, 82.0–86.9 mm SL, Sakashizu, Uwajima, 11 July 2001, net, Uwajima Fisheries High School;
OMNH-P 14107, 2 specimens, 91.6–107.4 mm Uwajima
Bay, 31 October 2000, line-fishing. OITA: KAUM-I. 32287,
74.3 mm SL, KAUM-I. 32288, 80.5 mm SL, KAUM-I.
32289, 85.6 mm SL, Hiji, Beppu, 33°21′N, 131°32′E, 28
July 2010, 3 m depth, line-fishing, S. Torigoe and H. Iwatsubo. NAGASAKI: BSKU 9863, 82.0 mm SL, Goto Islands,
1955; FAKU S 122, 101.8 mm SL, FAKU S 123, 100.1 mm
SL, Shikoe, Tsushima, 1973; KAUM-I. 28139, 80.3 mm SL,
Tachibana Bay, Ikanoura, 4 April 2010, line-fishing, M.
Okamoto. KUMAMOTO: SNFR 12473, 4 specimens, 85.7–
97.6 mm SL, Minamata, 23 March 1987. KAGOSHIMA:
KAUM-I. 7576, 59.0 mm SL, east of Sakinoyama, Kata­ura, Kasasa, Minami-satsuma, 31°25′44″N, 130°11′49″E,
24 February 2007, 27 m depth, set net, M. Ito; KAUM-I.
10753, 33.2 mm SL, Ohto Port, Kasasa, Minami-satsuma,
31°25′19″N, 130°10′27″E, 6 July 2008, 4–5 m depth, handnet, M. Ito; KAUM-I. 33715, 105.9 mm SL, off Ikara-jimaisland, Nagashima, Izumi, 32°13″N, 130°12′E, 20 October 2010, KAUM Fish Team; KAUM-I. 33770, 110.5 mm
SL, Take-shima island, Nagashima, Izumi, 32°13′26″N,
130°10′51″E, 5–10 m depth, 24 October 2010, line-fishing,
KAUM Fish Team; KAUM-I. 33801, 94.7 mm SL, KAUMI. 33802, 95.3 mm SL, off Ikara-jima island, Nagashima,
Izumi, 32°13′N, 130°12′E, 26 October 2010, KAUM FishTeam. KOREA: BSKU 101353, 89.0 mm SL, BSKU 101354,
108.4 mm SL, BSKU 101355, 110.0 mm SL, BSKU 101364,
108.4 mm SL, BSKU 101365, 110.0 mm SL, BSKU 101366,
104.2 mm SL, BSKU 101367, 97.3 mm SL, BSKU 101368
93.1 mm SL, BSKU 101369, 69.5 mm SL, BSKU 103852,
93.6 mm SL, BSKU 103853, 84.4 mm SL, BSKU 103854,
98.4 mm SL, BSKU 103855, 100.5 mm SL, BSKU 103856,
94.3 mm SL, BSKU 103857, 96.3 mm SL, BSKU 103858,
104.9 mm SL, BSKU 103859, 94.6 mm SL, BSKU 103860,
102.5 mm SL, BSKU 103861, 102.3 mm SL, Jeju Market, Jeju
Island, 5 August 2005, H. Endo.
PACIFIC COAST OF JAPAN plus SOUTHERNCHINA (71 specimens, 26.7–122.3 mm SL). CHIBA:

202

H. Iwatsubo and H. Motomura

KAUM-I. 25873, 68.2 mm SL, off Tokyo University of Marine Science and Technology Field Science Center Tate­yama Station, Banda, Tateyama, 34°58′37″N, 139°46′10″E,
July 2009, 7 m depth, hand net, M. Watai; KPM-NI 21975,
26.7 mm SL, Banda, Tateyama, 4 October 2003, hand net, M.
Honda; KPM-NI 25242, 92.6 mm SL, off Shiramazu, Chikura, Minamiboso, 23 m depth, 9 July 2009, H. Ishikawa.
KANAGAWA: KPM-NI 2443, 103.0 mm, KPM-NI 2444,
105.8 mm SL, Hayakawa Fishing Port, Odawara, set net,
6 June 1996; KPM-NI 2555, 95.2 mm SL, Odawara Fish
Market, Odawara, 6 June 1996, H. Senou; KPM-NI 2571,
100.6 mm SL, 22 June 1996; KPM-NI 2886, 103.8 mm SL,
Odawara Fish Market, Odawara, 3 July 1996, H. Senou;
KPM-NI 15137, 90.5 mm SL, KPM-NI 15769, 86.5 mm
SL, Aburatsubo Bay, Koajiro, Misaki, Miura, 10 March
1996, Mochizuki et al.; KPM-NI 24878, 100.0 mm SL, off
Enoshima island, Sagami Bay, Fujisawa, 20 m depth, 16
August 2009, H. Ishikawa. SHIZUOKA: KAUM-I. 31046,
90.5 mm SL, KAUM-I. 31047, 71.4 mm SL, Numazu, 4 January 1997, H. Ida; KPM-NI 13265, 81.1 mm SL, KPM-NI
13266, 99.4 mm SL, Heda Port, Heda, Tagata, 30 September 2003; NSMT-P 81233, 74.7 mm SL, Masaki breakwater, Shimizu Harbor, Miho Peninsula, 15 July 1982, gill net.
MIE: FAKU 24503, 98.9 mm SL, Washio, 25 October 1954,
Y. Iwai; FRLM 2600, 110.7 mm SL, Wagu, Shima, 22 June
1980, S. Kimura et al.; FRLM 3052, 111.4 mm SL, Wagu,
Shima, 31 May 1981, S. Kimura et al.; FRLM 3223, 95.5 mm
SL, FRLM 3224, 99.5 mm SL, FRLM 3226, 106.8 mm SL,
FRLM 3227, 102.5 mm SL, FRLM 3230, 97.7 mm SL, FRLM
3231, 96.0 mm SL, FRLM 3233, 103.4 mm SL, FRLM 3234,
110.2 mm SL, FRLM 3235, 86.0 mm SL, Wagu, Shima, 23
September 1981, S. Kimura et al.; FRLM 4085, 116.5 mm
SL, Wagu, Shima, 31 July 1983, S. Kimura et al.; FRLM
4512, 98.1 mm SL, FRLM 4514, 83.7 mm SL, Ago Bay, Goza,
Shima, 14 September 1984, S. Kimura et al.; FRLM 6669,
94.5 mm SL, Wagu, Shima, 8 September 1987, S. Kimura
et al.; FRLM 6750, 69.1 mm SL, Wagu, Shima, 19 August
1987, S. Kimura et al.; FRLM 11050, 64.8 mm SL, Ago Bay,Goza, Shima, 29 June 1991, S. Kimura et al.; FRLM 20828,
42.8 mm, Shukutaso, Minami-ise, Watarai, 29 August 1997,
S. Kimura et al. KOCHI: BSKU 56080, 122.3 mm SL, Okino-shima island, Sukumo, line-fishing, 9 July 2001; BSKU
7412, 83.1 mm SL, Kochi Fish Market, 16 August 1957;
BSKU 3416, 74.2 mm SL, BSKU 3417, 72.3 mm SL, Mimase,27 November 1953; BSKU 8559, 77.7 mm SL, BSKU 8560,
86.6 mm SL, Susaki, 7 July 1959; BSKU 52423, 93.4 mm SL,
Kokadai, 16 April 1994; KSHS 8802, 80.6 mm SL, KSHS
8804, 97.9 mm SL, Kochi Central Fish Market, 4 November
1968; KSHS 18585, 96.9 mm SL, Sukumo Bay, Sukumo, 26
July 1979. MIYAZAKI: MUFS 30749, 99.1 mm SL, MUFS
30750, 96.8 mm SL, MUFS 30751, 94.3 mm SL, Iorigawa,
Kadokawa, Higashiusuki, 14 September 2009. CHINA: CAS
55958, 5 specimens, 60.0–81.6 mm SL, CAS 55959, 11 specimens, 54.9–85.2 mm SL, Hong Kong, 27 February 1958, R.
L. Bolin and D. Bromhall; KAUM-I. 31050, 91.9 mm SL,
KAUM-I. 31051, 86.8 mm SL, KAUM-I. 31052, 86.9 mm SL,
KAUM-I. 31053, 85.2 mm SL, KAUM-I. 31054, 84.8 mm SL,
Cheung Chau Island, Hong Kong, 19 November 1995, H.

Ida. TAIWAN: USNM 14159, 87.9 mm SL, Soo Wan Bay,
3–9 m depth, 29 January 1910; USMN 270856, 85.8 mm SL,
southwest of Yeh-liu, 25°12′N, 121°41′E, 8–11 m depth, 19
May 1968, V. Springer et al.
IZU ISLANDS (5 specimens, 65.9–128.8 mm SL). TOSHIMA ISLAND: KPM-NI 25259, 128.8 mm SL, 35 m
depth, 16 May 2009, Y. Ogawa. MIYAKE-JIMA ISLAND:
KAUM-I. 31020, 88.3 mm SL, KAUM-I. 31021, 92.0 mm
SL, 23 June 1972, H. Ida; KAUM-I. 31022, 122.4 mm SL, 25
July 1973; NSMT-P 30735, 65.9 mm SL, Igaya Bay, 30°06′N,
139°30′E, 22 September 1973.
Diagnosis. A species of Chromis with the following
combination of characters: dorsal fin rays XIII–XIV, 11–14(mode XIII, 13); anal fin rays II, 10–12 (11); pectoral fin rays
16–20 (19); principal caudal fin rays 15; spiniform caudal fin
rays 2; scale rows in longitudinal series 23–28 (26); tubed
lateral line scales 16–21 (17); scale rows above lateral line
4–5 (4), scale rows below lateral line 11–12 (11); gill rakers
7–10 (9)+19–24 (21)=27–33 (30); preopercular marginsmooth; caudal fin deeply forked, without prolonged filaments; prominent black blotch covering upper two-thirds ofpectoral fin base throughout life; indistinct white spot at endof dorsal fin base; caudal peduncle without distinct markings; broad dark band along each caudal fin lobe.
Distribution. Chromis notata is distributed only in
East Asia, where it ranges from Akita Prefecture and southwards in the Sea of Japan, including southern Korea, from
Chiba Prefecture to Kyushu along the Pacific coast of Japan,
including the Izu Islands, and in southern China, including Taiwan and Hong Kong (Fig. 4). It does not occur in
the Ryukyu Islands. Senou et al. (2006) reported C. notata
from Ie-jima island, off Okinawa-jima island in the central
Ryukyu Islands, on the basis of a single specimen (KPMNI 4072). This specimen (35.6 mm SL, 15 m depth, 16 June1997) is herein re-identified as Chromis atripectoralis Welander and Schultz, 1951 (H. Senou, pers. comm.).
Synonyms. Bloch (1792) described Perca japonica as a
new species on the basis of a single specimen from Japan.
The dried holotype of P. japonica (ZMB 14630) has apparently been lost (Paepke 1999; this study). Bloch andSchneider (1801) proposed Cichla macrolepidota Bloch andSchneider, 1801 as a new replacement name for P. japonica,
notwithstanding that P. japonica was an available name.
However, C. macrolepidota is regarded here as an available
name since it was described before 1931 (ICZN 1999: article 12.2.3); thus C. macrolepidota is a junior synonym of
P. japonica. Although Paepke (1999) wrote, “P. japonica appears to be a senior synonym of Chromis notatus”, examination of the original description of P. japonica and the drawing of the holotype (reproduced here as Fig. 5) showed thatP. japonica differs from C. notata in having X, 10 dorsal fin
rays, 8 anal fin soft rays, 3 scale rows above and 6 scale rows
below the lateral line, and no black spot on the pectoral finbase or dark brown bands along the caudal fin lobes. Although the taxonomic status of P. japonica is still unknown,
it is regarded here as a nomen oblitum because the name has
never been used since its original description, except for
Bloch and Schneider (1801) and Paepke (1999).

Three species of Western Pacific Chromis203

Fig. 2.
Primary type specimens of nominal species regarded as Chromis notata. A, lectotype of Heliases notatus (RMNH.PISC. 895,
86.8 mm SL); B, holotype of Chromis villadolidi (FMNH 59185, 101.0 mm SL); C, holotype of Chromis miyakeensis (ZUMT 53957, 126.3 mm
SL); D, holotype of Chromis flavomaculatus (BSKU 8482, 84.8 mm SL).

Jordan and Tanaka (1927) described Chromis villadolidi
as a new species on the basis of five specimens from the Sea
of Japan between Tsushima Island and Fukuoka, off northern Kyushu. Aoyagi (1941), Kamohara (1960), and Randall
et al. (1981) regarded C. villadolidi as a junior synonym of
C. notata. Examination of the holotype (FMNH 59185,
101.0 mm SL; Fig. 2B) of C. villadolidi showed it to be identical with C. notata, the former having five scale rows above
the lateral line, 11 scale rows below, and a black blotch restricted to the upper two-thirds of the pectoral fin base (Fig.
2; Table 1). Therefore, the taxonomic status of C. villadolidi
as a synonym of C. notata is confirmed in this study.
Chromis miyakeensis was originally described by Moyer
and Ida (1976) on the basis of eight specimens from Miyake-jima island in the Izu Islands off central Honshu,
Japan. Subsequently, Randall et al. (1981) regarded C. miyakeensis as a subspecies of C. notata, and its subspecies status, i.e., as C. notata miyakeensis, was recognized as well by
Ida (1984) and Aonuma and Yoshino (1993, 2000, 2002).
Chromis notata miyakeensis was supposedly characterized
by a deeper body, compared with C. notata notata (Randall et al. 1981; Aonuma and Yoshino 1993, 2000, 2002),
but our examination of a large series of specimens of C.
notata showed that body depth varies geographically (see‘Geographic variation of Chromis notata’ below). The rangeof the body depths of specimens from Miyake-jima island(45.8–52.3% of SL), including the holotype of C. miyakeensis(50.4%; Fig. 2C), was almost within that of C. notata fromother localities (37.8–51.5%; Tables 1, 2; Figs 6, 7). Thus, C.

miyakeensis is herein regarded as a junior synonym of C. notata. The synonymy of C. flavomaculata is discussed below(see ‘Remarks’ on C. kennensis).
Geographic variation of Chromis notata. Examination of a large series of specimens of C. notata from a wide
area of East Asia showed it to have three morphologicallydistinct populations, viz., in the Sea of Japan, in the Seto Inland Sea plus East China Sea, and along the Pacific coast ofJapan plus southern China (Figs 3, 4, 6, 7; Table 2). The population of the Sea of Japan tends to have a shallower body
depth (mean 44.0% of SL), a narrower body width (17.2%),
and a shallower caudal peduncle depth (14.1%) than the
population of the Pacific coast of Japan plus southern China(48.0%, 19.5%, and 14.8%, respectively; Figs 3, 6; Table 2). In
addition, the dorsal fin spines of the former population tend
to be longer (mean length of first spine 9.9% of SL) thanthose of the latter (9.1%) (Fig. 7; Table 2). The populationof the Seto Inland Sea plus East China Sea has intermediate
values of these morphometrics [e.g., body depth 41.1–50.7%
of SL (mean 45.5%), body width 15.4–23.4% (18.2%), andcaudal peduncle depth 13.3–15.9 (14.4%); Figs 3, 6, 7; Table
2]. Despite these slight morphometric differences among thethree populations, there are no differences in meristics, coloration, or other morphological features among them (Fig.
3; Tables 2, 3). Thus, the three are herein regarded as examples of geographic variation of a single species.
Specimens from the Izu Islands, including the holotype of
C. miyakeensis, tend to have a relatively deep body and caudal peduncle, wide body width, and long dorsal fin spines

204

H. Iwatsubo and H. Motomura

Fig. 3.
Juveniles to adults of Chromis notata from different localities. A–C, specimens from the Sea of Japan (A, KAUM-I. 31033, 34.0 mm
SL; B, KAUM-I. 31045, 60.4 mm SL; C, KAUM-I. 34910, 113.8 mm SL). D–F, specimens from the Seto Inland Sea and East China Sea(D, KAUM-I. 10753, 33.7 mm SL; E, KAUM-I. 7576, 59.0 mm SL; F, KAUM-I. 33715, 104.5 mm SL). G–I, specimens from the Pacific coast ofJapan and southern China (G, KPM-NI 21975, 26.7 mm SL; H, KAUM-I. 25873, 68.2 mm SL; I, KPM-NI 25259, 122.3 mm SL).

Fig. 5.
Reproduction of original painting of Perca japonica(from Bloch 1792: pl. 311, fig. 1).

a deeper body, is identical to C. notata.

Fig. 4.
Distributional records of Chromis notata on the basis of
specimens examined in this study: populations of the Sea of Japan(☐), the Seto Inland Sea plus East China Sea (△), the Pacific coastof Japan plus southern China (○), and the Izu Islands (▽). Somesymbols include more than one specimen.

(Table 2), but these values are almost within those of the
population of the Pacific coast of Japan plus southern China(Figs 6, 7; Table 2). This further supports our contention
that C. miyakeensis, which was characterized only by having

Chromis kennensis Whitley, 1964
[Australian English name: Yellow-spotted Chromis](Figs 8, 9, 11, 12E; Tables 3–4)
Chromis kennensis Whitley, 1964: 182, Fig. 12 (type locality:
Kenn Reef, Coral Sea); Allen 1975: 81, 83, unnumberedfig. (Kenn Reef, Coral Sea; Lord Howe Island; Sydney,
Australia; Ouvea Island, Loyalty Islands, New Caledonia).
Chromis flavomaculata (not of Kamohara): Allen 1991: 68(Ouvea Island, Loyalty Islands, New Caledonia; Lord
Howe Island; Coral Sea; eastern Australia between south-

Three species of Western Pacific Chromis205

Fig. 6.
Relationships of (A) body depth and (B) body width to
standard length in Chromis notata populations of the Sea of Japan(☐), the Seto Inland Sea plus East China Sea (△), the Pacific coastof Japan plus southern China (○), and the Izu Islands (▽).

ern Great Barrier Reef and Sydney); Kuiter 1993: 237,
unnumbered fig. (Coral Sea to Montague Island; Lord
Howe Island); Randall et al. 1997: 258, unnumbered fig.
(Coral Sea; Lord Howe Island; New Caledonia); Randall
2005: 357, unnumbered fig. (southern Great Barrier Reef
to New South Wales; Lord Howe Island; Norfolk Island;
New Caledonia); Allen et al. 2006: 1449, (Lord Howe Island; Great Barrier Reef, Queensland to Montague Island,
New South Wales, Australia); Kuiter and Debelius 2006:
530, unnumbered fig. (Australia); Kato 2011b: 58 (Great
Barrier Reef; New Caledonia; Tonga); Allen et al. 2012:
80 (eastern Australia and Coral Sea); Allen and Erdmann2012: 580 (eastern Australia; Coral Sea; New Caledonia;
northern tip of New Zealand).
Holotype. AMS IB. 4973, 77.6 mm SL, Kenn Reef,
Coral Sea, D. McMichael.

Non-type material examined. 23 specimens, 22.5–
93.0 mm SL—AUSTRALIA: AMS I. 17357-023, 43.0 mm
SL, off Phillip Point, Lord Howe Island, New South Wales,
31°05′S, 159°01′E, G. Allen et al.; AMS I. 17376-008, 4
specimens, 33.6–35.1 mm SL, Lord Howe Island, New South
Wales, 31°50′S, 159°10′E, F. Talbot et al.; AMS I. 17380-006,
2 specimens, 22.5–23.8 mm SL, Lord Howe Island, NewSouth Wales, 31°50′S, 159°10′E, AMS party; AMS I. 17737001, 48.8 mm SL, Long Bay, Sydney, New South Wales,
33°07′S, 151°03′E, R. Kuiter; AMS I. 27139–024, 71.4 mm
SL, Middleton Reef, Coral Sea, 29°48′67″S, 159°06′83″E, A.
Gill et al.; AMS I. 27891-043, 2 specimens, 59.0–62.2 mm
SL, Elizabeth Reef, Coral Sea, 29°09′S, 159°00′E, D. Williams et al.; BPBM 14766, 3 specimens, 30.0–33.0 mm
SL, Lord Howe Island, New South Wales, 20–27 m depth,
6 February 1973, spear, J. Randall and G. Allen; BPBM
33664, 3 specimens, 79.0–82.0 mm SL, Coral Sea, 19°52′S,
158°19′E, 9 m depth, 24 August 1988, J. Randall et al. NEW
CALEDONIA: BPBM 11438, 70.4 mm SL, 1 km northwestof Dumbae Pass, Nouméa, 15–18 m depth, J. Randall et al.;USNM 319941, 31.8 mm SL, Ouvea Island, Loyalty Islands,
20°39′30″S, 166°23′00″E, 18 m depth, 14 November 1991,
J. Williams et al.; USNM 319942, 82.1 mm SL, Bagaat Islet,
Ouvea Atoll, Loyalty Islands, 20°37′18″S, 166°16′08″E,
15–22 m depth, 16 November 1991, J. Williams, et al.;USNM 319943, 74.8 mm SL, Ouvea Island, Loyalty Islands,
20°34′12″S, 166°14′12″E, 5 m depth, 16 November 1991,
J. Williams et al. TONGA: USNM 332432, 28.9 mm SL,
Ha’atafu Beach, Tongatapu, 21°03′58″S, 175°20′03″W, 5.5–
7.9 m depth, 21 October 1993, J. Williams, et al.
Diagnosis. A species of Chromis with the following
combination of characters: dorsal fin rays XIII, 11–13 (mode
XIII, 12); anal fin rays II, 10–12 (11); pectoral fin rays 17–21(19); principal caudal fin rays 15; spiniform caudal fin rays
2; scale rows in longitudinal series 24–27 (26); tubed lateralline scales 15–19 (18); scale rows above lateral line 3, scalerows below lateral line 9–10 (9); gill rakers 7–9 (8)+21–25(24)=29–34 (32); preopercular margin smooth; caudal fin
deeply forked, without prolonged filaments; basal width ofpelvic fin axillary scale 0.6–1.8% of SL; basal width of scale
between pelvic fin bases 2.0–2.8% of SL; prominent black
blotch covering entire pectoral fin base throughout life; nowhite spot at end of dorsal fin base; caudal peduncle without
distinct markings; caudal fin yellowish with no dark band
along each fin lobe.
Distribution. Chromis kennensis is known only fromthe southwestern Pacific Ocean: the Coral Sea (Ken Reef,
Middleton Reef, and Elizabeth Reef), Lord Howe Island,
eastern Australia (Sydney), New Caledonia (Ouvea Island),and Tonga (Tongatapu) (Fig. 9).
Remarks. Kamohara (1960) described Chromis flavomaculata (as flavomaculatus) as a new species on the basisof a single specimen (BSKU 8482; Fig. 2D) from Kochi,
Japan, while also listing an additional non-type specimen(BSKU 4944) of the species from Tokushima, Japan. Examination of the holotype and non-type specimen showed
them to be identical with C. notata in having four scale rows
above the lateral line and 12 scale rows below, a black blotch

206

Fig. 7.

H. Iwatsubo and H. Motomura

Relationships of the dorsal fin spine lengths to standard length in Chromis notata populations. Symbols as in Fig. 6.

Fig. 8.
Photographs of Chromis kennensis. A, AMS IB. 4973, holotype, 77.6 mm SL; B, juvenile, New South Wales, Australia (photo by R.
Kuiter); C, BPBM 14766, juvenile, 31.6 mm SL, Lord Howe Island (photo by J. Randall); D, BPBM 11438, adult, 93.0 mm SL, New Caledonia(photo by J. Randall).

Three species of Western Pacific Chromis207

Fig. 9.
Distributional records of Chromis kennensis (◆) and C.
yamakawai sp. nov. (★) on the basis of specimens examined in thisstudy. Some symbols include more than one specimen.

restricted to the upper two-thirds of the pectoral fin base,and black bands along the caudal fin lobes (Fig. 2; Tables 4,5). Therefore, C. flavomaculata is herein regarded as a juniorsynonym of C. notata. In the same paper, Kamohara (1960)
gave a description of two specimens of C. notata. These two
specimens (BSKU 4946 and 8513) were examined and reidentified by us as C. notata and C. yamakawai sp. nov., respectively. Data for Kamohara’s (1960) specimens are given
in Table 5.
A species of Chromis, previously identified as C. flavomaculata, is herein regarded as comprising two geographically disjunct species: C. yamakawai sp. nov. fromthe northwestern Pacific Ocean and C. kennensis from thesouthwestern Pacific Ocean (Fig. 9). These two species canbe distinguished from C. notata by having three scale rows
above the lateral line and nine or 10 scale rows below (vs 4
or 5 and 11 or 12 in C. notata; Tables 1–5), a black blotch
covering the entire pectoral fin base in adults (vs restricted
to the upper two-thirds of the pectoral fin base), and a yellow caudal fin without black bands (vs a grayish caudal fin
with black bands along the lobes) (Figs 3, 8, 10).
Chromis kennensis, previously regarded as a junior synonym of C. flavomaculata (e.g., Randall et al. 1997; Randall 2005; Allen et al. 2006), differs from C. yamakawai sp.
nov. in coloration, meristics, and morphometrics. In young
specimens, the dark blotch at the base of the pectoral fin isrelatively large in C. kennensis, covering the entire fin base,whereas that of C. yamakawai is small, covering only theFig. 10. Photographs of Chromis yamakawai sp. nov. A, KAUM-I. 40100, juvenile, 31.9 mm SL, Iou-jima island, Kagoshima, Japan; B,
NSMT-P 111948, holotype, 83.0 mm SL, Yoron-jima island, Kagoshima, Japan; C, juvenile, Hachijo-jima island, Izu Islands, Japan (photo by
S. Kato); D, adult, Hachijo-jima island, Izu Islands, Japan (photo by S. Kato).

208

H. Iwatsubo and H. Motomura

Fig. 11. Relationships of (A) basal width of pelvic fin axillary scale and (B) basal width of scale between bases of pelvic fins to standard
length in Chromis kennensis (◆) and C. yamakawai sp. nov. (★).

Fig. 12. Heads of three species of Chromis. A–D, C. notata (A, KAUM-I. 26534, 97.7 mm SL, Tobi-shima island, Yamagata, Japan; B,
KAUM-I. 33715, 104.5 mm SL, Kagoshima, Japan; C, KPM-NI 24878, 100.0 mm SL, Enoshima island, Kanagawa, Japan; D: ZUMT 53957,
126.3 mm SL, Miyake-jima island, Izu Islands, Japan); E, C. kennensis (AMS IB. 4973, holotype, 77.6 mm SL, Kenn Reef, Coral Sea);
F, C. yama­kawai sp. nov. (NSMT-P 111948, holotype, 83.0 mm SL, Yoron-jima island, Kagoshima, Japan). AN, anterior nostril; PN, posterior
nostril.

Three species of Western Pacific Chromis209

upper half of the fin base (the blotch becomes larger withgrowth in C. yamakawai) (Figs 8, 10). In meristics, C. kennensis differs slightly from C. yamakawai in having fewer
tubed lateral line scales (15–19 vs 16–21 in C. yamakawai),and more gill rakers [21–25 (mode 24) lower gill rakers and
29–34 (32) total rakers vs 20–25 (23) and 27–33 (30), respectively] (Table 3). In morphometrics, the basal width ofthe pelvic fin axillary scale is 0.6–1.8% of SL (mean 1.5%)
in C. kennensis, whereas that of C. yamakawai is 1.4–2.8%(2.2%) (Fig. 11; Table 4). The basal width of the scale between the pelvic fins in C. kennensis (2.0–2.8% of SL, mean
2.6%) also tends to be less than that of C. yamakawai(2.2–3.9%, 3.0%) (Fig. 11; Table 4). Moreover, C. yama­kawai grows larger than C. kennensis (largest recorded size
130.8 mm vs 93.0 mm SL).
Chromis kennensis and C. yamakawai are similar to C.
pura Allen and Randall, 2004, which is known only fromthe Alor Islands, Indonesia, in overall body appearance
and general coloration, but C. pura has a yellowish caudal
peduncle, darker yellow fins, and a smaller black blotch at
the base of the pectoral fin (Allen and Randall 2004: figs. 2,3). Chromis pura also differs from the former two species
in having fewer gill rakers (25–27 in C. pura vs 29–34 in C.
kennensis and 27–33 in C. yamakawai).
Chromis yamakawai sp. nov.
[Standard Japanese name: Kihoshi-suzumedai][New English name: Northern Yellow-spotted Chromis](Figs 9–11, 12F; Tables 3–5)
Chromis notatus (not of Temminck and Schlegel): Aoyagi
1941: 181, fig. 11 (southern part of Japan); Kamohara
1960: 4 (Kochi, Japan).
Chromis sp.: Masuda et al. 1975: 96, 285, fig. E on p. 96(southern Japan, including Ryukyu and Ogasawara islands).
Chromis flavomaculatus (not of Kamohara): Masuda et al.
1980: 96, 285, fig. E on p. 96 (southern Japan, including
Ryukyu and Ogasawara Islands); Ida 1984: 192, pl. 185,
fig. F (central Japan to Taiwan).
Chromis flavomaculata (not of Kamohara): Araga et al.
1985: 148, unnumbered fig. (Izu Peninsula and southwards along Pacific coast of Japan; Taiwan); Allen 1991:
68, unnumbered fig. (southern Japan, including Ryukyu
Islands; Taiwan); Aonuma and Yoshino 1993: 809, unnumbered fig. (Ogasawara Islands, Izu Peninsula andsouthwards along Pacific coast of Japan; Taiwan); Masuda
and Kobayashi 1994: 221, fig. 5 on p. 221 (southern
Japan); Randall et al. 1997: 258 (Japan to Taiwan); Ao­numa and Yoshino 2000: 925, unnumbered fig. (Ogasa­
wara Islands, Izu Peninsula and southwards along Pacificcoast of Japan; Taiwan); Aonuma and Yoshino 2002: 925,
unnumbered fig. (Ogasawara Islands, Izu Peninsula andsouthwards along Pacific coast of Japan; Taiwan); Randall
2005: 357 (southern Japan); Kuiter and Debelius 2006:
530, unnumbered fig. (Japan); Konishi 2007: 238, unnumbered fig. (Izu, Ogasawara, and Ryukyu islands, Japan;
Taiwan); Yoshino and Senou 2008: 264, unnumbered figs

(Pacific coast of southern Japan; Hachijo-jima island, Izu
Islands; Ryukyu and Ogasawara islands; Taiwan); Araga
2009: 440 (Ogasawara Islands, Izu Peninsula and southwards along Pacific coast of Japan; Taiwan); Motomura
et al. 2010: 149 (Yaku-shima island, Kagoshima, Japan);
Kato 2011a: 150, unnumbered figs (southern Japan;
Hachijo-jima island; Ryukyu Islands; Ogasawara Islands;
Taiwan); Kato 2011b: 58, unnumbered figs (Hachijo-jimaisland, Izu Islands; Taiwan); Allen et al. 2012: 80, unnumbered fig. (southwestern Japan to Philippines); Allen andErdmann 2012: 580, unnumbered fig. (southern Japan,
including Ryukyu Islands; Taiwan; northern Philippines).
Holotype. NSMT-P 111948 (formerly KAUM-I. 40160),
83.0 mm SL, off west coast of Yoron-jima island, Kagoshima,
Japan, 27°03′21″N, 128°24′39″E, 10 m depth, 15 August
2011, drive fishing, H. Motomura, M. Yamashita, T. Yoshida,
H. Nishiyama, and H. Iwatsubo.
Paratypes. 114 specimens, 18.2–130.8 mm SL—
JAPAN: BPBM 6880, 92.2 mm SL, Ishigaki-jima island, Oki­nawa, May 1968, spear, J. Randall; BPBM 11989, 68.4 mm
SL, Ishigaki-jima island, Okinawa, 22 May 1968, J. Randall and A. Banner; BPBM 35316, 53.5 mm SL, Tatsumi
Bay, Chichi-jima island, Ogasawara Islands, 26 m depth,
7 June 1992, J. Randall and R. Pyle; BPBM 19073, 2 specimens, 68.4–92.2 mm SL, southeast coast of Sesoko Island,
Okinawa, 26 May 1975, spear, J. Randall; BSKU 8513,
11.3 mm SL, Okino-shima island, Sukumo, Kochi, 8 June
1959; BSKU 65128, 114.2 mm SL, BSKU 65129, 120.0 mm
SL, BSKU 65130, 109.3 mm SL, BSKU 69938, 130.8 mm
SL, Okino-shima island, Sukumo, Kochi, line-fishing, 22
August 2002; FAKU 104285, 69.7 mm SL, FAKU 104420,
67.9 mm SL, Ankyaba, Amami-oshima island, 24 December
1958, T. Mashima; FAKU 61749, 90.7 mm SL, FAKU 61753,
88.7 mm SL, Shirahama, Wakayama, 1981–1982; FAKU
121418, 102.6 mm SL, Shirahama, Wakayama, 1984–1985;
FRLM 10757, 77.6 mm SL, Naha, Okinawa, 17 May 1990,
S. Kimura et al.; HMNH-P 3118, 80.9 mm SL, Okinawa,
17 August 2001; HMNH-P 3505, 26.4 mm SL, Seragaki,
Onna, Okinawa, 18 August 2001; HMNH-P 4286, 97.9 mm
SL, Manzamo, Onna, Okinawa, 1 January 2002; HMNH-P
7485, 108.0 mm SL, HMNH-P 7486, 89.6 mm, Okinawa, 27
December 2003, Okinawa Federation of Fisheries Cooperatives; HUMZ 3455, 82.8 mm SL, HUMZ 3456, 73.3 mm
SL, Okinawa; HUMZ 3457, 85.1 mm SL, Okinawa; HUMZ
3461, 79.0 mm SL, Okinawa; HUMZ 39070, 72.9 mm SL,
Chinen, Okinawa, 6 March 1974; HUMZ 46754, 66.3 mm
SL, HUMZ 46758, 66.6 mm SL, Naha Fish Market, Naha,
Okinawa, 29 October 1975; HUMZ 48235, 76.4 mm SL,
Naha, Okinawa, 13 April 1973; HUMZ48567, 85.8 mm SL,
Naha, Okinawa, 13 April 1973; HUMZ 48579, 83.0 mm SL,
Naha, Okinawa, 13 April 1973; KAUM-I. 7211, 96.0 mm SL,
KAUM-I. 7212, 103.0 mm SL, off Kakeroma-jima island,
Setouchi, Amami Islands, Kagoshima, 13 November 2007,
net, FV Uehara-maru; KAUM-I. 29581, 18.2 mm SL, off west
coast of Iou-jima island, Mishima, Kagoshima, 30°15′42″N,
130°15′42″E, 5–20 m depth, 28 May 2010, hand net, KAUM
Fish Team; KAUM-I. 31023, 96.3 mm SL, KAUM-I. 31024,

210

H. Iwatsubo and H. Motomura

106.6 mm SL, KAUM-I. 31025, 104.4 mm SL, Miyake-jimaisland, 25 July 1973, H. Ida; KAUM-I. 40099, 26.2 mm SL,
KAUM-I. 40100, 31.9 mm SL, KAUM-I. 40101, 26.8 mm
SL, off southwest coast of Yoron-jima island, Kagoshima,
27°01′48″N, 128°24′37″E, 2–3 m depth, drive fishing, 14
August 2011, H. Motomura et al.; KAUM-I. 40161, 81.7 mm
SL, KAUM-I. 40162, 86.4 mm SL, KAUM-I. 40163, 79.5 mm
SL, KAUM-I. 40170, 91.8 mm SL, KAUM-I. 40175, 91.0 mm
SL, KAUM-I. 40177, 95.3 mm SL, KAUM-I. 40178, 95.8 mm
SL, KAUM-I. 40181, 50.5 mm SL, KAUM-I. 40183, 91.5 mm
SL, KAUM-I. 40186, 96.1 mm SL, KAUM-I. 40194, 92.8 mm
SL, KAUM-I. 40198, 80.5 mm SL, KAUM-I. 40213, 52.4 mm
SL, KAUM-I. 40214, 55.2 mm SL, KAUM-I. 40216, 54.9 mm
SL, KAUM-I. 40217, 54.6 mm SL, KAUM-I. 40129, 53.9 mm
SL, KAUM-I. 40221, 53.7 mm SL, same data as holotype;
KPM-NI 4686, 95.1 mm SL, KPM-NI 4687, 77.8 mm SL,
Ishigaki-jima island, Okinawa, net, March 1998; KPM-NI
5065, 26.4 mm SL, KPM-NI 5105, 91.2 mm SL, KPM-NI
5106, 89.3 mm SL, Ie-jima island, Okinawa, 4 m depth, handnet, 19 June 1998, T. Nomura; KPM-NI 17973, 121.9 mm SL,
Gunkan Dike, Kaminato Port, Kaminatohigashi, Hachijojima island, Tokyo, line-fishing, 16 August 2000, Y. Miya­zaki; KPM-NI 18542, 112.7 mm SL, Futami Port, Chichijima island, Ogasawara Islands, Tokyo, line-fishing, 12 December 2006, Y. Miyazaki; KSHS 15340, 82.7 mm SL, KSHS
15341, 78.0 mm SL, Okinawa-jima island, March–April
1974; KSHS 16983, 85.3 mm SL, KSHS 16984, 85.6 mm SL,
KSHS 16985, 77.9 mm SL, KSHS 16986, 79.2 mm SL, Naha
Fish Market, Naha, Okinawa, 5 November 1975; NSMT-P
3246–004, 98.0 mm SL, Amami-oshima island, 21 November 1957; NSMT-P 29624, 77.2 mm SL, Miyako-jima island,
Okinawa, 20 m depth, net, 21 July 1988, H. Senou and M.
Aizawa; NSMT-P 30613, 55.4 mm SL, Igaya Bay, Miyakejima island, Izu Islands, 34°06′N, 139°30′E, 17 November
1974; NSMT-P 30742, 67.6 mm SL, Igaya Bay, Miyake-jimaisland, Izu Islands, 34°06′N, 139°30′E, 19 November 1974;
NSMT-P 30811, 79.2 mm SL, NSMT-P 30812, 76.5 mm
SL, Igaya Bay, Miyake-jima island, Izu Islands, 30°06′N,
139°30′E, 10 m depth, 21 December 1976; NSMT-P 32798,
32.9 mm SL, Miyanohama Bay, Chichi-jima island, Ogasa­
wara Islands, 27°06′N, 142°15′E, hand net, 10 June 1990,
J. Okuno; NSMT-P 90966, 103.9 mm SL, NSMT-P 90967,
109.1 mm SL, Futami Bay, Chichi-jima island, Ogasa­wara
Islands, 12 December 2006, Y. Miyazaki; NSMT-P 92979, 2
specimens, 98.6–98.9 mm SL, Keise-jima island, Oki­nawa,
6 July 2003, T. Kon; NSMT-P 32810, 2 specimens, 18.9–
23.4 mm SL, Kominato beach, Chichi-jima Island, Ogasawara Islands, 27°03′N, 142°11′E, 3 m depth, 12 June 1990,
K. Matsuura; NSMT-P 105526, 25.3 mm SL, Sokaru, Amami-oshima island, Kagoshima, 8 m depth, 11 June 1991, K.
Matsuura and M. Aizawa; URM-P 907, 69.1 mm SL, URM-P
908, 67.0 mm SL, URM-P 909, 68.4 mm SL, URM–P 910,
73.3 mm SL, URM-P 911, 70.7 mm SL, Oki­nawa Island;
URM-P 1182, 94.7 mm SL, Tomari Fish Market, Naha,Oki­nawa, 10 December 1981; URM-P 4598, 97.2 mm SL,
Ishigaki Port, Ishigaki-jima island, Okinawa, 6 September
1982, H. Masuda; URM-P 11761, 77.1 mm SL, Oki­nawa,
9 September 1985, Y. Aonuma; URM-P 22905, 89.2 mm

SL, off Tropical Biosphere Research Center Sesoko Station,
University of the Ryukyus, Sesoko-jima island, Oki­nawa, 13
July 1977, T. Yoshino; URM-P 27135, 68.9 mm SL, Amamioshima island, Kagoshima, 27 March 1992, Y. Sakurai et al.;
URM-P 30301, 88.5 mm SL, Okinawa, 17 September 1993,
H. Yoshigou and Y. Sakurai; URM-P 31167, 66.4 mm SL,
Okinawa, 12 February1994, H. Yoshigou; URM-P 33815,
76.0 mm SL, Okinawa, 16 June 1995, H. Yoshigou; URM-P
37846, 100.0 mm SL, Maeda Cape, Okinawa, 11 April 1997,
H. Yoshigou and M. Sekine; URM-P 37901, 92.5 mm SL,
Okinawa, 19 April 1997, N. Takei and H. Yoshigou; URM-P
38096, 71.0 mm SL, Okinawa, 17 May 1997, H. Yoshigou etal.; URM-P 39899, 78.2 mm SL, URM-P 39900, 82.0 mm
SL, Setouchi, Amami-oshima island, Kagoshima, 6 April
1998, Setouchi Fisheries Cooperative; WAM-P 25516–016,
74.7 mm SL, Okinawa. PHILIPPINES: USNM 314196, 2
specimens, 70.8–72.3 mm SL, Y’ami Island, Batan Islands,
21°07′35″N, 121°56′45″E, 15–20 m depth, 26 April 1987, D.
Johnson et al.; USNM 364157, 79.4 mm SL, Sabatan Island, 9November 1908.
Diagnosis. A species of Chromis with the following
combination of characters: dorsal fin rays XIII–XIV, 11–13(mode XIII, 12); anal fin rays II, 10–12 (11); pectoral fin
rays 17–21 (19); principal caudal fin rays 15; spiniform caudal fin rays 2; scale rows in longitudinal series 21–27 (26);
tubed lateral line scales 16–21 (19); scale rows above lateralline 3, scale rows below lateral line 9–10 (9); gill rakers 6–9(7)+20–25 (23)=27–33 (30); preopercular margin smooth;caudal fin deeply forked, without prolonged filaments; basal
width of pelvic fin axillary scale 1.7–2.8% of SL; basal widthof scale between pelvic fin bases 2.2–3.9% of SL; prominent
black blotch covering upper two-thirds of pectoral fin basein juveniles and covering entire fin base in adults; no whitespot at end of dorsal fin base; caudal peduncle without distinct markings; caudal fin yellowish without dark band
along each fin lobe.
Description. Proportional measurements as percentages of SL are given in Table 4. Frequency distributions of
selected meristic characters are presented in Table 3. Features for the holotype are presented first, followed by paratype data in parentheses. Characters given in the diagnosis
are not repeated here.
Body moderately deep, compressed. Dorsal profile ofhead slightly convex. Snout shorter than orbit diameter.
Mouth terminal, small, oblique; posterior margin of maxilla just reaching (or not reaching) to vertical through anterior margin of pupil. Upper jaw with entire row of canine
teeth, 2 or 3 irregular rows of smaller conical teeth anteriorly, 1 such row posteriorly. Lower jaw with row of canine
teeth and anterior inner row of small conical teeth; no small
posterior teeth. No teeth on vomer and palatines. Tongue
obtuse triangular. Gill rakers long and slender, longest on
lower limb near angle, about three-fourths as long as longest
gill filaments. Nostril with fleshy rim, more protrusive on
posterior edge, vertically situated at level of middle of pupil.
Preopercular margin smooth; upper end not reaching levelof upper margin of orbit. Suborbital with free lower margin.
Opercular margin with small, flat spine projecting from be-Three species of Western Pacific Chromis211

neath large scale. Body scales finely ctenoid. Scales covering head except for lips, tip of snout, and narrow zone fromorbit to around nostrils. Anterior lateral line ending below
soft-rayed portion of dorsal fin. Scaly sheath at base of dorsal fin, its width about one-half of pupil diameter at base of
middle of dorsal fin. Scales covering basal parts of dorsal,
pectoral, and anal fins. Small scales covering entire caudal
fin, except for center and posterior margin. Small scales
on basesof pectoral fins. One median scale process extending posteriorly from between bases of pelvic fins, its length
about half that of pelvic spine. Axillary scale above base ofpelvic spine more than one-third as long as that spine. Origin of dorsal fin above third (or fourth) tubed scale. Tip of
pectoral fin not reaching vertical through origin of anal fin.
Depressed pelvic fin reaching to (or just short of) origin of
anal fin. Vertebrae 11+​15=​26.
Color when fresh (based on color photographs of KAUMI. 40100 and NSMT-P 111948; Fig. 10A, B): body olivebrown with dark-brown-margined body scales (more yellowish body in young). Upper head and nape greenish
brown; suborbital and opercular scales sometimes bluish.
Spinous portion of dorsal fin yellowish brown with dark
margin in young, brownish without marginal marking in
adults; posterior half of soft-rayed portion of dorsal fintransparent. Pectoral fin transparent. Pelvic fin yellowish tobrownish anteriorly, transparent posteriorly. Anterior twothirds of anal fin blackish, remaining portion transparent.
Caudal fin lobes yellow in young, dark yellow in adults, andtransparent centrally. Color of preserved specimens similar
to that of fresh specimens, except for dusky brown caudal
fin.
Distribution. Chromis yamakawai is known from the
Izu Islands (recorded from Miyake-jima and Hachijo-jimaislands), the Ogasawara Islands (Chichi-jima island), thePacific coast of southern Japan (Wakayama and Okinoshima island, Kochi), the Ryukyu Islands (Iou-jima, Amami-oshima, Kakeroma-jima, Yoron-jima, Okinawa-jima,Ie-jima, Sesoko-jima, Keise-jima, and Ishigaki-jima islands),
and the Philippines (Batan Islands) (Fig. 9).
Etymology. The species is named for Takeshi Yama­kawa (Kochi, Japan), who first recognized this species as
new and kindly made specimens available to the authors.
Remarks. Chromis yamakawai has previously been regarded as C. flavomaculata, the latter being regarded as a
junior synonym of C. notata in this study (see Remarks forC. kennensis). Comparisons among C. notata, C. kennensis,
C. yamakawai, and C. pura are made in the Remarks forC. kennensis. These four species can be distinguished fromother members of Chromis by having a smooth preopercular margin, a large, black blotch at the base of the pectoralfin, a blackish or brownish caudal peduncle, and a caudal
fin without filamentous rays. Chromis westaustralis Allen,
1976, endemic to Western Australia, is similar to C. notata
in general coloration and morphology, but specimens of C.
westaustralis were not available for this study; detailed comparisons between the two species are required to assess thetaxonomic status of C. westaustralis.
The squamation and sensory pores on the heads of C. no-tata, C. kennensis, and C. yamakawai are shown in Fig. 12.
The distribution of sensory pores on the head varies individually and does not represent a species-specific pattern (Fig.
12).

Acknowledgments
We thank M. McGrouther, A. Hay, and S. Reader (AMS),
A. Suzumoto and L. O’Hara (BPBM), H. Endo (BSKU),
D. Catania (CAS), Y. Kai (FAKU), S. Kimura (FRLM), H.
Yoshi­gou (HMNH), M. Yabe, H. Imamura, and T. Kawai(HUMZ), Y. Iwatsuki (MUFS), K. Matsuura and G. Shino­hara (NSMT), K. Hatooka (OMNH), M. van Oijen and R.
de Ruiter (RMNH), K. Hoshino (SNFR), T. Yoshino, M.
Toda, and K. Miyamoto (URM), J. Williams and S. Raredon (USNM), S. Morrison and G. Moore (WAM), P. Bartsch(ZMB), K. Sakamoto (ZUMT), and H. Ida (formerly Kita­
sato University) for opportunities to examine specimens of
Chromis. We are grateful to T. Yamakawa (formerly Kochi
Senior High School) for sharing his knowledge of the taxonomy of Chromis, M. Ito (Maruse Daikichi Fishery Cooperative), S. Chiba and E. Katayama (NSMT), M. Okamoto(SNFR), S. Dewa (Diving Services Umiannai), K. Furuta(Diving Services Umikobo), Y. Kaji and other staff of the
Wakayama Prefectural Museum of Natural History, K.
Hoshino, S. Torigoe, K. Imai, and other staff of the Oita Marine Palace Aquarium Umitamago, T. Kasedo and T. Ogami(Faculty of Fisheries, Kagoshima University), staff of the
Azuma-cho Fishery Cooperative, H. Sato, H. Sato, K. Kaji­wara, K. Tokuda, S. Hiwatari, and T. Yasunaga (Mishima,
Kagoshima), H. Takahashi (Kagoshima University Activation Center for Yoron Island), K. Tatsuno and other staff
of Yoron Town Hall, T. Takeshita (Yoron Diving Services),
N. Iwamura and other staff of the Yoron Chamber of Commerce and Industry, and Y. Kimura (Graduate School of
Biosphere Science of Hiroshima University) for their kindassistance with collecting specimens of Chromis. We also
thank S. Kato (Regulus Diving) and R. Kuiter (Aquatic Photographics) for kindly providing underwater photographsof C. yamakawai from Hachijo-jima island and C. kennensis
from New South Wales, respectively, J. Randall for providing photographs of C. kennensis, T. Hashimoto (KAUM) for
taking X-ray photographs, M. Matsunuma and G. Ogihara(KAUM) for examining the type specimens of Chromis, H.
Senou (KPM) for information on C. notata from the Ryukyu
Islands, and G. Yearsley (Ellipsis Editing) for checking an
early version of this manuscript. We greatly appreciate Y.
Haraguchi, volunteers, and students of KAUM for theircuratorial assistance. This study was supported in part by
Grants-in-Aid for Scientific Research (B, 24370041 and C,
23580259) from the Japan Society for the Promotion of Science, Tokyo, Japan (JSPS), a Grant-in-Aid for Young Scientists (B, 19770067) from the Ministry of Education, Science,Sports and Culture, Tokyo, Japan, the JSPS Asian Core Program, “Establishment of Research and Education Network
on Coastal Marine Science in Southeast Asia”, the JSPS International Training Program, “Protect Diversity of Biore-

212

H. Iwatsubo and H. Motomura

sources in the Tropical Area”, and the Coastal Area Capability Enhancement in Southeast Asia Project of the ResearchInstitute for Humanity and Nature, Kyoto, Japan.

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