Bull. Natl. Mus. Nat. Sci., Ser. A, Suppl. 7, pp. 11–18, March 22, 2013

Suttonia coccinea, a New Grammistin Fish from Japan(Acanthopterygii: Serranidae)
Hiromitsu Endo1 and Kenta Kenmotsu1,2
1

Laboratory of Marine Biology, Faculty of Science, Kochi University,
2–5–1 Akebono-cho, Kochi 780–8520, Japan
E-mail: endoh@kochi-u.ac.jp
2
Yasuzuka High School, 129 Kamigata, Yasuzuka, Jouetsu, Niigata 942–0405, Japan

Abstract A new serranid, Suttonia coccinea, is described based on a single specimen of 65.1 mm
SL collected from the rocky shore of Okinoshima Island, Kochi Prefecture, Shikoku Island, southern
Japan. The new species differs from its two congeners, S. lineata and S. suttoni, in the following
characters: shorter lower jaw (its length 60.9% HL vs. 62.5–66.6); lower jaw slightly projecting
anteriorly beyond upper jaw when closed (vs. clearly beyond upper in S. lineata); longest 3rd–5th
dorsal-ﬁn spines shorter (each length 6.8% SL vs. 7.9–9.5); longest 19–20th dorsal-ﬁn soft-rays
longer (each length 18.0% SL vs. 12.6–16.5); longest 16–17th anal-ﬁn soft-ray longer (each longest length 17.2% SL vs. 12.9–15.5); shorter pelvic ﬁn (its length 11.8% SL vs. 13.0–14.8); scaly
inner base of pectoral ﬁns (vs. naked in S. suttoni); fewer pored scales on lateral line (26 vs. 35–41
in S. lineata, 27–35 in S. suttoni); uniform scarlet head and body with bright red ﬁns when fresh(vs. partly pale in S. lineata, uniform vermilion with chrome-yellow margins of vertical ﬁns in S.
suttoni); and a distinct dark spot on opercle (vs. faint in S. lineata). The new species represents thenorthernmost record of the genus.
Key words : Serranidae, Pseudogrammina, Suttonia, new species, Japan.

The serranid genus Suttonia Smith, 1953
belongs to the tribe Grammistini of the subfamily
Epinephelinae, and is composed of two nominal
species, Suttonia suttoni Smith, 1953 and Suttonia lineata Gosline, 1960 (Baldwin and Johnson,
1993; Randall and Baldwin, 1997; Nelson, 2006;
Craig and Hastings, 2007). Members of thissmall genus are less than 10 cm long, and adults
have a reddish body, with a pale medial line on
the head (e.g., Randall and Baldwin, 1997; Randall, 2005). They live in cryptic habitats of coraland rocky reefs (6–30.5 m depth) of tropical andsubtropical waters in the Indo–West Paciﬁc: S.
suttoni is restricted to the eastern Indian Ocean,Pemba Island off Tanzania (type locality) andComoro Islands, and S. lineata is known widelyfrom the eastern Indian Ocean, Cocos-Keeling
Islands and Christmas Island, to the mid-Paciﬁc,Hawaiian Islands (type locality) and Tahiti, andfrom northeastern Australia to Japan (e.g., Ran-dall and Baldwin, 1997; Kuiter, 2004; Randall,
2005, 2007; Hoese et al., 2006; Allen and Erdmann, 2012). Although Allen and Erdmann(2012) included Japan in the range of S. lineata,
they did not refer to the source of their information. To our knowledge, no specimens or underwater photographs of the species taken in Japanare known.
During SCUBA sampling of ﬁshes by the Laboratory of Marine Biology, Faculty of Science,
Kochi University, around Okinoshima Island,
southwest of Kochi, in July of 2010, the ﬁrst
author collected a small serranid of 65 mm SL ata depth of 16 m on a rocky bottom with corals.
The specimen, which was a uniformly bright reddish color with a mesial pale line on the head
when alive, is identiﬁed as Suttonia. Here, we
describe the specimen as a new species, andreport the northernmost record of the genus.

12

H. Endo and K. Kenmotsu

Materials and Methods
Specimens examined are deposited in the following institutions: Bernice P. Bishop Museum,
Honolulu (BPBM); Laboratory of Marine Biology, Faculty of Science, Kochi University(BSKU); National Museum of Nature and Science, Tsukuba (NSMT, formerly National Science Museum, Tokyo); and Royal Ontario
Museum, Toronto (ROM). Counts and measurements follow Randall and Baldwin (1997): e.g.,caudal-peduncle length is measured as the horizontal distance between verticals at the rear baseof the anal ﬁn and the caudal-ﬁn base; the lengthof the preopercular spine is measured along its
ventro-anterior margin to the tip; and pelvic-ﬁnlength is measured from the base of the spine to
the tip of the longest soft ray. Observation ofdentition, cephalic sensory pores, scales, andexternal bony elements were made after cyanine
blue staining. Total length, standard length, andhead length are abbreviated as TL, SL, and HL,
respectively. Fin rays and vertebrae were counted
from radiographs.
Suttonia Smith, 1953
[New Japanese name: Kurenai-toge-megisu-zoku]

Remarks. Members of the genus Suttonia
possess scales with cteni that do not extend posteriorly beyond the membranous portion of thescale, which is a unique character state among
the Pseudogrammina (Fig. 5; Randall and Baldwin, 1997: ﬁg. 11B). Several characters includedin Randall and Baldwinʼs (1997) generic diagnosis need slight modiﬁcation based on dataobtained from the new species: dorsal-ﬁn rays
VII, 22–25; anal-ﬁn rays III, 18–22; pectoral-ﬁnrays 14–17; a single lateral- line with 26−40
scales in adults; longitudinal scale series 50–56;
gill rakers 5–6＋9–12; vertebrae 10＋16–17 (usually 16); body depth 27–33% SL; HL 37–41%
SL; snout length 17–22% HL; caudal-peduncle
length 11–17% HL; caudal-peduncle depth
29–35% HL; maxilla extending posteriorly to avertical at rear edge of orbit; a band of villiform

teeth in jaws with a small canine-like tooth(sometimes a close-set pair) laterally on eachside or a pair on one side of upper-jaw symphysis; a V-shaped patch of villiform teeth on vomer;
palatines with a band of villiform teeth in 1–4
rows; longest dorsal-ﬁn spine 17–28%HL; longest dorsal-ﬁn soft ray 43–50%HL; second analﬁn spine clearly longest, 16–25%HL; longest
anal-ﬁn soft ray 42–49%HL; caudal ﬁn rounded
to slightly rhomboidal, 57–71 %HL; pectoral ﬁns
67–80% HL (26–32% SL); pelvic ﬁn short, its
origin distinctly anterior to base of pectoral ﬁn;absence of cteni on scale edge (Fig. 5); life color
primarily mottled red to scarlet with a mediandorsal pale pink stripe on head from lower lip to
dorsal-ﬁn origin.
Craig and Hastings (2007) indicated that thesubtribe Pseudogrammina (sensu Randall and
Baldwin, 1997), which includes Aporops, Pseudogramma and Suttonia, is monophyletic basedon the genetic analysis of two nuclear and two
mitochondrial genes. This supported an earlier
hypothesis based on morphological data by Baldwin and Johnson (1993). Members of this subtribe lack the dermal toxin “grammistin,” which
is present in other members of the tribe Grammistini (Baldwin and Johnson, 1993).
The new Japanese name of the genus, Kurenai-toge-megisu-zoku, is based on the Japanesename of the new species, Kurenai-toge-megisu.
The new species represents the ﬁrst documented
record of the genus from Japanese waters.
Suttonia coccinea sp. nov.
[New English name: Scarlet Freckle-faced Podge][New Japanese name: Kurenai-toge-megisu](Figs. 1–5, Tables 1–2)

Holotype. NSMT-P 110917 (formerly BSKU
103900) 65.1 mm SL, 79.7 mm TL, Kuboura,
Okinoshima Island, Sukumo City, Kochi Prefecture, Shikoku Island, Japan (32°44′44″N,
132°33′32″E), 16 m depth, hand net by SCUBA,
coll. by H. Endo, 21 July 2010.
Diagnosis. A new species of Suttonia with thefollowing combination of characters: lower jaw

New Serranid Suttonia from Japan

13

Fig. 1. Suttonia coccinea sp. nov., holotype, NSMT-P 110917, 65 mm SL, in fresh condition: A, left lateral view;B, left dorsolateral view of head; C, ventral view of head to anterior of abdomen. Photographed by H. Endo.

Fig. 2. Suttonia coccinea sp. nov., holotype, NSMT-P 110917, 65 mm SL, in preserved condition: A, left lateralview; B, dorsal view of head; C, ventral view of head to anterior of abdomen. Photographed by H. Endo.

moderately short, its length 61% HL); moderately low spinous dorsal ﬁn, its longest spine
length 17% HL; moderately high soft dorsal ﬁn,

its longest soft-ray length 46% HL; moderately
high anal ﬁn, its longest soft-ray length 44% HL;
somewhat short pelvic ﬁn, its length 61% HL);

14

H. Endo and K. Kenmotsu

Table 1. Counts and aspects of squamation of three Suttonia species. G (1960) and R&B (1997) indicate Gosline(1960) and Randall and Baldwin (1997) respectively. One and two asterisks indicate the counts of specimens
larger than 50 and 40 mm SL respectively.
S. coccinea

Standard length (mm)Dorsal-ﬁn rays
Anal-ﬁn rays
Pectoral-ﬁn rays
Pored lateral-line scalesLongitudinal scale series
Vertebrae
Inner base of pectoral ﬁn

S. lineata

S. suttoni

Holotype

this study
n＝12

R&B (1997)
n＝39

G (1960)
n＝6

R&B (1997)
n＝27

Smith (1953)
n＝4

65
VII, 23
III, 19
16
26
51
10＋16
scaly

66–80
VII, 23–25
III, 19–21
15–18
37–41
47–54
10＋16
scaly

17–81
VII, 22–25
III, 19–22
15–17
35–40*
52–56
10＋16–17
scaly

38–78
VII, 23–24
III, 20–22
16
—
58–64
—
—

34–65
VII, 22–24
III, 18–20
14–15
27–35**
50–53
10＋16
naked

63–78
VII, 23–24
III, 19
16
31
—
—
—

pored scales in lateral line 26; scaly inner base ofpectoral ﬁns; head, body, and ﬁns uniform scarlet when fresh; a distinct dark spot on opercle
and dusky to black pigment on dorso-lateral portion of head.
Description. Counts and proportions areshown in Tables 1 and 2. Body somewhat robust,compressed. Spinous dorsal ﬁn much lower thansoft dorsal ﬁn; ﬁrst spine short, its length 71% oflongest spine; second to seventh spines almost
equal in length, third to ﬁfth spines longest; allsoft rays branched. Posterior corners of 19–20th
dorsal- and 16–17th anal-ﬁn soft rays greatly
elongated. First anal-ﬁn spine shortest, slender,
its length 67% of second spine; second spine
robust, longest, its length 16% HL; third spine
slender, slightly longer than ﬁrst; all soft rays
branched. Pectoral ﬁn elongate, just reaching
posteriorly beyond a vertical at anal-ﬁn origin;its base narrow, height 21% HL; upper origin
below dorsal-ﬁn origin; all ﬁn-rays branched.
Pelvic ﬁn short, its origin anterior to lowermost
base of pectoral ﬁn; spine short, its length 13%
HL, 42% of pelvic-ﬁn length; all soft-rays
branched. Anus located immediately anterior toanal-ﬁn origin. Caudal peduncle short, its length
40% of its depth. Caudal ﬁn large, rounded, with
15 principal rays.
Dorsal outline of head nearly straight anteriorly, gently curved posteriorly. Snout short,rounded, moderately wide, its length almost
equal to orbit diameter. Anterior nostril on ante-

Fig. 3. Dorsal view of anterior head of Suttonia
coccinea sp. nov., holotype, NSMT-P 110917,
65 mm SL, showing cephalic sensory pores on
snout and interorbital region: AN, anterior nostril; AIP, anterior interorbital pore; PIP, posterior interorbital pore; PN, posterior nostril; SOP,
suborbital pore. Drawn by N. Nakayama.

rior rim of snout, forming long tube that projects
slightly beyond upper lip when depressed anteriorly, reaching posterior nostril when depressed
posteriorly, its length half of orbit diameter; posterior nostril without tube, oval, close to orbit, its
longitudinal diameter one ﬁfth of orbit diameter.
Eyes somewhat small. Interorbital space narrow,ﬂat, without dermal ﬂap or small tentacle.
Cephalic lateral-line pores (Fig. 3): two pairs ofnasal pores medially near nostrils; pair of anterior interorbital pores near orbital rims; single

New Serranid Suttonia from Japan

15

Fig. 4. Lateral views of head of two Suttonia species: A, Suttonia coccinea sp. nov., holotype, NSMT-P 110917,
65 mm SL; B, Suttonia lineata, BPBM 7974, 67 mm SL. Photographed by H. Endo.

posterior interorbital pore on a mesial plane; suborbital pores 9; preoperculo-mandibular pores 13(4 dentary and 9 preopercular pores); postorbital
pores 6.
Mouth large, somewhat oblique. Maxilla
extending posteriorly beyond a vertical at rearedge of orbit. Tip of lower jaw slightly projecting
anteriorly beyond upper when mouth closed(Figs. 1B, 4A). A band of villiform teeth in jaws
inwardly curved, conical: 4–6 irregular rows nearsymphysis, 2–4 irregular rows posteriorly, teeth
in inner row somewhat enlarged; two large
canine-like teeth on right side of upper-jaw symphysis (presumably two on left side missing);
vomer with a V-shaped patch of villiform teeth;
palatines with a narrow band of villiform teeth in
1–3 rows.
Gill rakers on ﬁrst arch (right side) 5＋9＝14:
upper 6 rakers on lower limb short, digit-likewith ﬁne spinules; other rakers rudimentary,
plate-like with ﬁne spinules. Tongue slender,
without teeth. Left and right branchiostegal
membranes united antero-ventrally near isthmus,
attached to antero-ventral portion of urohyal.
Branchiostegal rays 7.
Opercle with three ﬂat spines, covered by skinFig. 5. Scale of Suttonia coccinea sp. nov., NSMTP 110917, 65 mm SL. Photographed by K. Kenmotsu.

(invisible externally). Opercular ﬂap well developed, extending posteriorly to a vertical at uppermost point of pectoral-ﬁn base. Upper edge of
operculum joined by thin membrane to head.
Spine on posterior edge of preopercle with deep
furrow, slender, projecting ventro-posteriorly,
directed at an angle of about 50 degrees from thehorizontal.
Scales ctenoid, cteni completely absent (Fig.
5), adherent, covering entire head and body
except the following: snout to middle of interorbital space anterior to anterior interorbital pores,
lachrymal, both jaws except postero-dorsal end

16

H. Endo and K. Kenmotsu

of dentary, preopercular spine, posterior rim ofpreopercle, branchiostegal membrane, and gular.
Fins variously covered with smaller scales except
the following: margin of spinous dorsal ﬁn, distal
three-ﬁfths to two-thirds of soft dorsal and analﬁns (naked area widest at the posterior corner ofﬁns), distal half of caudal ﬁn, and distal threefourths of paired ﬁns. Inner base of pectoral ﬁnscaled.
Single lateral line with 26 pored scales, reaching posteriorly to a vertical at space between ﬁrst
and second anal-ﬁn soft rays.
Color when fresh (Fig. 1). Ground color ofhead and body uniformly scarlet; ﬁns bright red;head dusky dorsally, somewhat pale ventrally; areddish to whitish mesial line between lips anddorsal-ﬁn origin; anterior portion of lips dusky
except at mesial line; orbital rim dusky except
ventrally, iris red, pupil black; a wide dusky rectangular blotch behind eye to preopercle and an
elongate blackish spot on opercle; dorsum ofbody slightly dusky; posterior corners of dorsaland anal-ﬁn membranes and distal half of caudal
ﬁn mottled blackish.
Color in alcohol (Fig. 2). Reddish coloration
when fresh completely faded. Head, body, andscaled vertical ﬁn bases yellowish brown,slightly dusky; dorsal portion of body moredusky (Fig. 2A). Dorsal portion of head and anterior portion of both lips, brown (Fig. 2B). Largemarking behind eye and spot on opercle darkbrown. Ventral half of head, mesial line on dorsal
portion of head, unscaled areas of vertical ﬁns,
and paired ﬁns yellowish white. Posterior corners of dorsal- and anal-ﬁn membranes, distal
half of caudal ﬁn mottled pale blackish brown.
Distribution. Known only from the type
locality, Okinoshima Island, southwest of Kochi,
Shikoku Island, Japan, at a depth of 16 m. Taken
from an opening under large rocks.
Etymology. From the Latin coccinea (scarlet)
in reference to its bright reddish coloration. The
new Japanese standard name of the species,“Kurenai-toge-megisu,” is also in reference to its
coloration.
Comparisons. The new species clearly

belongs to the genus Suttonia based on its having
the following combination of characters: dorsalﬁn rays VII, 22; anal-ﬁn rays III, 19; pectoral-ﬁnrays 16; vertebrae 10＋16; a large inclined preopercular spine with furrow; pair of large interorbital pores, one on each side and situated at edgeof orbit; single lateral line; cteni on scales completely absent; and reddish coloration with palemiddorsal stripe from upper jaw to origin of dorsal ﬁn (Table 1; Figs. 1, 3, 5). Suttonia coccinea
differs from S. lineata and S. suttoni in the following proportions: longest dorsal-ﬁn spine
shorter (length 6.8% SL vs. 7.9–9.5 and 7.9
respectively), longest dorsal-ﬁn soft-ray longer(length 18.0 % SL vs. 12.6–16.2 and 16.5), longest anal-ﬁn soft ray longer (length 17.2% SL vs.
12.9–15.5 and 14.1), pelvic ﬁn shorter (length
11.8% SL vs. 13.0–14.6 and 14.8), lower jaw
shorter (length 60.9% HL vs. 62.5–66.6) (Table
2; Fig. 4). Suttonia coccinea differs from S. lineatain having fewer pored scales in the lateral line
(26 vs. 35–41), lower jaw slightly projecting
anteriorly beyond upper jaw when closed (vs.
clearly beyond upper), uniformly red pectoral
ﬁns (vs. pale with reddish margin) and a distinct
dark spot on the opercle (vs. faint spot) (Table 1,
Figs. 1, 2, 4; Randall and Baldwin, 1997: pl.
1–I). Although S. coccinea resembles S. suttoni
in having fewer pored scales in the lateral line
than S. lineata (26 in S. coccinea, 27–35 in S.
suttoni — 31 in the four type specimens of 63–
78 mm SL counted by Smith, 1953), bright reddish coloration, and a distinct spot on the opercle, the new species differs from S. suttoni inhaving scales on the inner base of the pectoral ﬁn(vs. naked) and a uniformly scarlet head andbody with bright red ﬁns (vs. uniform vermilion
with chrome-yellow margins of vertical ﬁns —Smith, 1953). Additionally, Smith (1953: fig. 3)
noted that one of 4 types of S. suttoni has 3 dark
longitudinal lines along the side of the body(scarlet when fresh). However, based on examination of 27 specimens of S. suttoni, including 3
paratypes and ROM 56690, Randall and Baldwin(1997) described 3 grooves along the sides
instead of 3 dark lines. Neither S. coccinea nor S.

New Serranid Suttonia from Japan
Table 2.

17

Proportional measurements of three Suttonia species.

Standard length (mm)As % of standard length
Body depth
Predorsal length
Preanal length
Prepelvic length
Dorsal-ﬁn base length
Anal-ﬁn base length
Longest dorsal-spine length
Longest dorsal soft-ray length
Longest anal-spine length
Longest anal soft-ray length
Caudal-ﬁn length
Pectoral-ﬁn length
Pelvic-ﬁn length
Caudal-peduncle length
Caudal-peduncle depth
Head length
As % of head length
Snout length
Orbit diameter
Interorbital width
Upper-jaw length
Lower-jaw length
Preopercular-spine length

S. suttoni

S. coccinea
NSMT-P
Holotype

S. lineata
BPBM
8 larger spec.

ROM 56690
1 specimen

Smith (1953)
4 types

65.1

65.5–79.8

40.5

63–78

30.4
38.2
69.3
37.5
55.5
33.9
6.8
18.0
6.3
17.2
24.1
26.0
11.8
6.1
12.0
39.3

25.9–30.6
36.6–38.8
62.3–70.8
32.5–38.6
54.6–61.9
31.0–33.2
7.6–9.5
12.6–16.2
5.3–7.6
12.6–15.5
18.3–23.2
24.7–27.8
13.0–14.6
5.6–7.5
10.9–12.7
36.4–37.9

28.9
41.2
67.9
33.8
59.8
32.8
7.9
16.5
6.4
14.1
24.4
29.1
14.8
5.7
11.6
38.5

31–32

20.3
19.9
9.4
50.8
60.9
17.2

16.8–20.4
19.3–21.2
7.6–9.1
49.0–51.9
62.5–66.6
15.1–19.9

16.7
24.4
7.1
51.9
63.0
19.9

lineata has grooves or dark lines laterally on thebody.
The new species represents the northernmost
record of Suttonia based on published specimen
data, and the range of the genus now extends
from Oahu, Hawaiian Islands (type locality of S.
lineata) to Okinoshima Island, southwest of Shikoku Island (32°45′N). Okinoshima Island of
Kochi is located in a subtropical- to temperatewater region off the Paciﬁc coast of southern
Japan (sensu Nakabo, 2002), and is strongly
inﬂuenced by the Kuroshio Current. Hence,rocky reefs in this area are exposed to warm temperatures in the winter and have many developed
corals. This ecosystem undoubtedly provides
cryptic environments suitable for S. coccinea.
Comparative materials. Suttonia suttoni (1specimen): Moheli, Comoro Islands, ROM
56690, 41 mm SL. Suttonia lineata (11): Oahu,
Hawaiian Islands, BPBM 6003 (1 specimen,
72 mm SL), BPBM 6350 (3, 66–80), BPBM

54
8
14.5
8
28
10
12
38–39

7974 (2, 67–79), BPBM 9767 (1, 66), BPBM
22660 (1, 78); Guam, BPBM 7276 (1, 36);
Tahiti, BPBM 8371 (1, 23); Manado, Sulawesi,
Indonesia, BPBM 36744 (1, 29). Pseudogramma
polyacanthum (2): Okinoshima Island, Kochi,
BSKU 71946 (1, 44), BSKU 71948 (1, 30), 19
July 2004. Aporops bilinearis (1): Kikaijima
Island, Amami Islands, BSKU 5425 (holotype of
Aporops japonicus Kamohara, 55).
Acknowledgments
We are most grateful to K. Matsuura (NSMT)and S. Kimura (Mie University) for their efforts
in preparing this supplement. We also thank thefollowing: members of the ﬁeld sampling team at
Okinoshima Island in 2010, Z. Imoto (UsaMarine Biological Institute, Kochi University),
E. Katayama (NSMT), M. Tsuboi and Y. Kimura(Hiroshima University), Y. Takata (Osaka City,
Osaka), M. Yamamura (Tosa Town, Kochi), M.

18

H. Endo and K. Kenmotsu

Hirota (Tosa City, Kochi), K. Tose (Kochi University), K. Matsuno (Aquas Kashiwajima Diving Service, Kochi) and J. Sakaue (Southern
Marine Laboratory, Palau) for their generous
assistance; Boyokan Sawachika (our regular inn
at Okinoshima Island, Kochi) and Sukumo-wan
Japan Fisheries Cooperative (Sukumo City,
Kochi) for assistance and permission for the
sampling respectively; R. Winterbottom (ROM),
A. Y. Suzumoto (BPBM), and G. Shinohara(NSMT) for loaning and cataloging the typespecimen; K. Shibukawa (Nagao Natural Environment Foundation, Tokyo) for providing uswith valuable information about Suttonia; E.
Katayama (NSMT) for obtaining literature; N.
Nakayama (BSKU) for drawing the illustration;
and G. Yearsley (Hobart, Australia) for editing
the English text. We sincerely thank an anonymous reviewer and S. Kimura for their valuablecomments on the manuscript. The study waspartly supported by Grant-in-Aids for ScientiﬁcResearch from the Japan Society for the Promotion of Science, Tokyo, Japan (A19208019 andB24370041), the Grant-in-Aid of “Marine Science Project”of Natural Science Cluster, Sciences
Unit, Kochi University, and the “Kuroshio Sougou Project” of National Museum of Nature andScience, Tsukuba.
Literature Cited
Allen, G. R. and M. V. Erdmann. 2012. Reef ﬁshes of theEast Indies. Vol. I. Tropical Reef Research, Perth.
x＋420 pp.＋end note.
Baldwin, C. C. and G. D. Johnson. 1993. Phylogeny of
the Epinephelinae (Teleostei: Serranidae). Bulletin ofMarine Science, 52(1): 240–283.

Craig, M. T. and P. A. Hastings. 2007. A molecular phylogeny of the groupers of the subfamily Epinephelinae(Serranidae) with a revised classiﬁcation of the Epinephelini. Ichthyological Research, 54: 1–17.
Gosline, W. A. 1960. A new Hawaiian percoid ﬁsh, Suttonia lineata, with a discussion of its relationships anddeﬁnition of the family Grammistidae. Paciﬁc Science,
14(1): 28–38.
Hoese, D. F., D. J. Bray and J. E. Gates. 2006. Grammistini. Podge Soapﬁshes. Pages 1026–1028 in D. F.
Hoese, D. J. Bray, J. R. Paxton and G. R. Allen, eds.
Fishes. In P. L. Beesley and A. Wells, eds. Zoologicalcatalogue of Australia. Volume 35. ABRS & CSIRO
Publishing: Australia Part 2, xxi＋671–1472 pp.
Kuiter, R. H. 2004. Basslets, Hamlets, and their relatives.
A comprehensive guide to selected Serranidae and Plesiopidae. TMC Publishing, Chorleywood. 216 pp.
Nakabo, T. 2002. Characteristics of the ﬁsh fauna ofJapan and Adjacent waters. Pages xliii–lii. T. Nakabo,
ed. Fishes of Japan with pictorial keys to the species,
English edition. Tokai University Press, Tokyo.
Nelson, J. S. 2006. Fishes of the world. Fourth edition.
John Wiley & Sons, Ins., Hoboken, New Jersey.
xx＋601 pp.
Randall, J. E. 2005. Reef and shore ﬁshes of the Southern
Paciﬁc. New Caledonia to Tahiti and the Pitcairn
Islands. University of Hawaiʻi Press, Honolulu.
xii＋707 pp.
Randall, J. E. 2007. Reef and shore ﬁshes of the Hawaiian
Islands. Sea Grant College Program, University of
Hawaiʻi, Honolulu. xiv＋546 pp.
Randall, J. E. and C. C. Baldwin. 1997. Revision of theserranid ﬁshes of the subtribe Pseudogrammina, withdescriptions of ﬁve new species. Indo-Paciﬁc Fishes,(26): 1–56.
Smith, J. L. B. 1953. The ﬁshes of the family Pseudogrammidae from East Africa. Annals and Magazine of
Natural History, Series 12, 6: 548–560.
Manuscript received 3 November 2012; revised 20 January 2013; accepted 31 January 2013.
Associate editor: S. Kimura