Species Diversity 21: 187–196
25 November 2016
DOI: 10.12782/sd.21.2.187

Distribution Range Extensions of Parapercis bicoloripes and P.
diplospilus (Perciformes: Pinguipedidae) in the South China
Sea and the Adjacent Waters, with Notes on Ontogenetic
Changes in P. bicoloripes
Mizuki Matsunuma1,8, Abd Ghaffar Mazlan2, Aziz Arshad3, Ying Giat Seah2,4,Siti Tafzilmeriam S. A. K.4, Azwarina Azmi Ramasamy4,
Ricard P. Babaran5, Yoshino Fukui6, and Hiroyuki Motomura7
1 Laboratory of Marine Biology, Faculty of Science, Kochi University, 2-5-1 Akebono, Kochi 780-8520, Japan

E-mail: k1139853@kadai.jp

2 School of Fisheries and Aquaculture Sciences, Universiti Malaysia Terengganu, 21030 Kuala Terengganu, Terengganu, Malaysia
3 Department of Aquaculture, Faculty of Agriculture, Universiti Putra Malaysia, 43400 UPM Serdang, Selangor, Malaysia

4 South China Sea Repository and Reference Center, Institute of Oceanography and Environment, Universiti Malaysia Terengganu,

21030 Kuala Terengganu, Terengganu, Malaysia

5 University of the Philippines Visayas, 5023 Miag-ao, Iloilo, Philippines

6 The United Graduate School of Agricultural Sciences, Kagoshima University, 1-21-24 Korimoto, Kagoshima 890-0065, Japan
7 The Kagoshima University Museum, 1-21-30 Korimoto, Kagoshima 890-0065, Japan
8 Corresponding author(Received 10 May 2016; Accepted 7 September 2016)
Parapercis bicoloripes Prokofiev, 2010, previously known only from waters off Vietnam, is recorded from the northern
Gulf of Thailand, the east coast of the Malay Peninsula, northern Borneo, and Panay, Philippines. Detailed examination of
27specimens (66.1–136.0 mm standard length) revealed significant growth-related changes in several body proportions andcoloration. In addition, 11 specimens (44.2–74.1 mm standard length) of Parapercis diplospilus Gomon, 1981, previously recorded from Vietnam, the Philippines, Indonesia, Papua New Guinea, and Australia, are reported for the first time from theGulf of Thailand and off Terengganu State, Malaysia, east coast of the Malay Peninsula.
Key Words: sandperches, distribution, morphology, Thailand, Malaysia, Philippines, Pacific Ocean.

Introduction
The South China Sea overlaps the Coral Triangle, a regionof exceptional marine diversity (Hoeksema 2007). Randalland Lim (2000) listed at least 3365 species of marine fishes
from the South China Sea, including the Gulf of Thailand.
Although a number of regional studies on marine fish faunas in this region have been published recently (e.g., Matsu­
numa et al. 2011; Yoshida et al. 2013; Kimura et al. 2015),
studies on the diversity of marine fishes in this region are
still ongoing.
Eleven specimens of a small sandperch collected from theGulf of Thailand and off Kuala Terengganu, Malaysia (eastcoast of the Malay Peninsula), were identified as Parapercis
diplospilus Gomon 1981, a species previously recorded frommany localities in the southeastern Indian and western Pacific oceans, including Vietnam, the Philippines, Indonesia,
Papua New Guinea, and Australia (Allen et al. 2006; Prokofiev 2008; Allen and Erdmann 2012). In addition, 27 specimens of Parapercis bicoloripes Prokofiev, 2010, previously
known only from the original description of Vietnamese
specimens, were collected from many localities in the South

China Sea and the adjacent waters, including the Gulf ofThailand, Malaysia and the Philippines. These new recordsof two sandperch species are reported here as a further stepin understanding marine fish diversity in this region.

Materials and Methods
Counts and measurements generally follow Randall et al.
(2008) and were generally made on the left side. Standard
and head lengths are abbreviated as SL and HL, respectively.
Gill rakers, including all rudiments, were counted on theouter side of the first right side arch. The last ray of boththe soft dorsal fin and anal fin was split to the base, but wascounted as a single ray, each being associated with a single
pterygiophore. Scale row counts above the lateral line include the dorsalmost longitudinal row of small scales beneath the dorsal-fin base, expressed as 0.5. The distribution
map was prepared using Quantum GIS 2.2 (Quantum GIS
Development Team 2014), with data from ETOPO1 (Amante and Eakins 2009). Specimens examined in this study aredeposited at the following institutions: Laboratory of Marine Biology, Faculty of Science, Kochi University (BSKU),© 2016 The Japanese Society of Systematic Zoology

Mizuki Matsunuma et al.

188

Kochi; the Kagoshima University Museum (KAUM), Kagoshima; National Museum of Nature and Science (NSMT),
Tsukuba; South China Sea Repository and Reference Center,
Institute of Oceanography and Environment, Universiti Malaysia Terengganu (UMTF), Kuala Terengganu; and Ichthyology Collection, Museum of Natural Sciences, University of
the Philippines Visayas (UPVMI), Miagao.
Parapercis bicoloripes Prokofiev, 2010
(Figs 1–3; Tables 1–2)
Material examined. Twenty-seven specimens, 66.1–
136.0 mm SL. Thailand: KAUM–I. 23099, 100.2 mm SL,
KAUM–I. 23124, 110.9 mm SL, KAUM–I. 23128, 100.5 mm
SL, KAUM–I. 23158, 100.9 mm SL, KAUM–I. 24153,
128.4 mm SL, northern Gulf of Thailand, bottom trawl (ob-tained at fish market in Mahachai). Malaysia (Terengganu
State, east coast of Malay Peninsula): KAUM–I. 16935,
120.4 mm SL, off Kuala Terengganu, bottom trawl, 11 December 2008 (obtained at Chendering fishing port, Kuala
Terengganu); KAUM–I. 41715, 119.1 mm SL, KAUM–I.
41716, 102.7 mm SL, 48 km off Chendering, Kuala Terengganu, 05°16′N, 103°11′E, 70–90 m depth, bottom trawl, 10
September 2011; KAUM–I. 79750, 123.3 mm SL, KAUM–I.
79751, 108.1 mm SL, KAUM–I. 79752, 104.9 mm SL,
KAUM–I. 79753, 126.2 mm SL, KAUM–I. 79754, 136.0 mm
SL, KAUM–I. 79755, 123.1 mm SL, KAUM–I. 79756,
126.0 mm SL, KAUM–I. 79757, 118.4 mm SL, KAUM–I.
79758, 115.3 mm SL, KAUM–I. 79759, 119.8 mm SL,
KAUM–I. 79760, 107.9 mm SL, KAUM–I. 79761, 132.7 mm
SL, UMTF 6569, 129.1 mm SL, UMTF 6570, 109.9 mm
SL, UMTF 6571, 79.0 mm SL, UMTF 6572, 102.4 mm SL,

Fig. 1. Fresh specimens of Parapercis bicoloripes from Malaysia (A–B) and the Philippines (C). A, KAUM–I. 79754, 136.0 mm SL, off Kuala
Terengganu; B, KAUM–I. 16935, 120.4 mm SL, off Kuala Terengganu; C, KAUM–I. 69435, 66.1 mm SL, off Miagao, Iloilo, Panay Island.

Two Parapercis species from the South China Sea

189

Fig. 2. Distribution of Parapercis bicoloripes (triangles) and P. diplospilus (circles). Closed and open symbols indicate previously known andnew records, respectively.

Fig. 3. Ontogenetic changes in relationship of snout length (circles) and fleshy orbit diameter (triangles) as percentage of standard
length to standard length (mm) in Parapercis bicoloripes.

UMTF 6573, 123.9 mm SL, off Kuala Terengganu, bottom
trawl. Malaysia (Sabah State, Borneo): KAUM–I. 12491,
96.2 mm SL, off Kota Kinabalu, bottom trawl (obtained
at fish market in Kota Kinabalu). Philippines: KAUM–I.
69435, 66.1 mm SL, off Miagao, Iloilo, Panay Island, 24 February 2015.
Description. Morphometrics, expressed as percentagesof SL, and frequency distribution of selected meristics areshown in Tables 1 and 2. Dorsal-fin rays V, 21; anal-fin rays
I, 17; pectoral-fin rays all branched except for uppermost
ray; pelvic-fin rays I, 5; branched caudal-fin rays 15 (8 and
7 in dorsal and ventral series, respectively); pseudobranchialfilaments 14–20 (18); branchiostegal rays 6.
Body subcylindrical, moderately elongate; moderately
deep, depth at anal-fin origin 4.8–5.8 in SL; width at pectoral-fin base 5.7–7.3 in SL. Caudal peduncle short, deep,
its depth 0.9–1.1 in length. Head moderately large, length
3.5–3.9 in SL; snout blunt, length slightly longer than orbit
diameter; eye relatively large, fleshy orbit diameter 3.5–4.3
in HL; interorbital space moderately wide, least bony width

10.2–15.6 in HL.
Mouth small, slightly oblique, forming angle of about 20°
to horizontal axis of head; lower jaw slightly protruding beyond tip of upper jaw; posterior margin of upper jaw reaching vertical drawn through anterior margin of pupil. Upper
jaw with outer row of 22–29 incurved caniniform teeth oneach side, anterior pair greatly enlarged, about 3 times larger
than adjacent teeth in size, other teeth increasingly smaller
posteriorly; inner band of villiform conical teeth behindouter caniniform row of upper jaw, broadest anteriorly with
4 or 5 rows maximum, gradually narrowing posteriorly to 1
row at rear of jaw. Front of lower jaw with usually 3 pairsof incurved, enlarged caniniform teeth in distinctly separated outer row (irregular row of 5 pairs of variously sized
caniniform teeth in KAUM–I. 79753 regarded as abnormal
condition), outermost tooth largest, strongly canted posteriorly, about 3 times size of innermost tooth; inner band ofvilliform conical teeth medial to caniniform teeth at frontof lower jaw, about 4–6 tooth rows in broadest portion, followed by row of small, incurved conical teeth, 2–4 teeth inmiddle of tooth row moderately enlarged; vomer with single
crescentic row of 6–8 moderately large conical teeth, all subequal in size; palatine teeth absent.
Gill membranes free from isthmus, with broad free fold;developed gill rakers short, spinous, longest raker about
one-third length of longest gill filament. Nostrils small, anterior nostril with short membranous tube, its tip not reaching posterior nostril when laid back; posterior nostril poreovate with low fleshy rim.
Pores of cephalic sensory system connected by canals
under skin. Three pores in row above maxilla; 3 pores near
nostrils, including 1 pore above and 1 below posterior nostril, 1 between nostrils; 2 pores on anterior margin of snout;row of 7–11 pores in infraorbital series, with several branches in posterior portion; 6–8 pores in row on posterior margin of preopercle; interorbital space with 2 pairs of pores inanterior half, irregular medial row of several small pores inposterior half; occiput with numerous small pores; ventral
surface of lower jaw with 5 pores on each side, anteriormost
pore behind chin.
Opercle bearing short, posteriorly-directed, flattened
spine approximately at level of ventral edge of orbit in lateralview; posterior margin of subopercle slightly expanded pos-Mizuki Matsunuma et al.

190

Table 1. Morphometrics, expressed as percentages of SL, of Parapercis bicoloripes and P. diplospilus from the South China Sea and the adjacent waters.
P. bicoloripes

P. diplospilus

27

11

Number of specimens
Standard length (mm)66.1–136.0

Mean

54.1–74.1

Mean

Body depth at anal-fin origin
Body width
Head length
Snout length
Fleshy orbit diameter
Fleshy interorbital width
Bony interorbital width
Upper-jaw lengthPre-dorsal-fin lengthPre-anal-fin length
Pre-pelvic-fin length
Dorsal-fin base length
1st dorsal-fin spine length
2nd dorsal-fin spine length
3rd dorsal-fin spine length
4th dorsal-fin spine length
5th dorsal-fin spine length
Longest dorsal-fin soft ray length
Anal-fin base length
Anal-fin spine length
Longest anal-fin soft ray length
Pectoral-fin length
Pelvic-fin length
Caudal-fin length
Caudal-peduncle depth
Caudal-peduncle length

17.2–20.6
13.6–17.6
25.7–28.7
6.8–9.1
6.2–7.9
2.9–4.1
1.7–2.6
10.1–12.5
26.2–30.6
43.4–49.9
23.1–25.7
61.7–66.2
2.5–3.5
4.1–5.8
5.7–7.4
6.0–9.6
4.4–7.2
13.8–17.5
41.3–48.6
3.3–5.4
12.2–14.6
19.2–22.0
20.6–25.7
18.1–21.4
8.5–10.2
9.0–11.1

19.2
15.8
26.9
8.3
6.9
3.5
2.2
11.5
28.2
45.5
24.2
64.3
3.0
4.8
6.4
7.4
5.9
15.9
45.4
4.5
13.4
20.2
22.6
19.8
9.6
9.9

14.7–16.8
14.3–17.4
25.7–27.9
6.5–7.7
6.8–8.4
2.4–3.3
1.0–1.6
8.4–9.4
26.5–29.5
43.0–47.7
21.2–23.1
61.2–68.7
1.6–2.7
3.5–5.0
5.3–7.1
7.1–8.2
4.5–5.4
12.9–14.3
46.2–50.1
5.3–6.6
9.3–10.6
18.4–20.6
20.1–23.2
18.4–20.6
7.7–8.5
6.3–8.2

15.7
16.1
27.2
7.0
7.6
2.8
1.4
8.7
28.4
45.4
22.4
65.5
2.2
4.4
6.2
7.7
4.9
13.8
47.8
6.2
9.8
19.5
22.0
19.5
8.1
7.5

Table 2.
waters.

Frequency distribution of selected meristics of Parapercis bicoloripes and P. diplospilus from the South China Sea and the adjacentPectoral-fin rays

P. bicoloripes
P. diplospilus

14

15

2

9

Pored lateral line scales

Scale rows above lateral line1

Scale rows above lateral line2

4.5

3.5

13

10

16

17

18

52

53

54

55

3.5

2

23

2

1

14
1

8
2

1

11

Scale rows below lateral line
10
P. bicoloripes
P. diplospilus

11

3

3

4

4.5

8

16
2

Circumpeduncular scale rows

12

13

14

8

9

10

11

23

24

25

26

2

17

4

5
1

10
4

6
2

1

1
6

4

10

9

1

2

7

Pre-dorsal-fin scale rows

Upper gill rakers

P. bicoloripes
P. diplospilus

4

Lower gill rakers

Total gill rakers

4

5

6

7

8

9

10

11

12

3

4

7

2

1
1

4
2

6

4

1

10
1

11

12

13

14

15

16

17

18

19

1

2

2

2

3

5

—

1

2

1to first dorsal-fin spine base; 2to first dorsal-fin soft ray base.

teriorly, with 8–10 large serrae; posterior edge of interopercle with 2 or 3 weakly developed serrae; preopercle roundedwith 11–20 serrae in all; posttemporal area with small exposed bony cleft, its posterior margin smooth.
Lateral line continuous, 3–5 pored lateral-line scales extending onto caudal fin; body entirely covered with devel-

oped ctenoid scales, scales on dorsolateral portion with ca.
35–50 short cteni; opercle, nape, and dorsal-postorbital region with developed ctenoid scales; cheek covered entirely
with weakly developed ctenoid scales, possessing ca. 25–30
short cteni; subopercle, interopercle, snout, interorbital
space, occipital area and both jaws naked; small, elongate

Two Parapercis species from the South China Sea

ctenoid basal scales extending onto pectoral and caudal fins;dorsal, anal, and pelvic fins without basal scales.
Origin of dorsal fin approximately above upper pectoral fin base; dorsal-fin spines relatively short; 4th dorsal-fin
spine longest; membrane between 5th spine and 1st soft ray
attached near tip of 5th spine, moderately incised; all softrays subequal in length, 19th ray (sometimes 18th or 20th,
rarely 10th or 11th) longest. Origin of anal fin below base of
5th or 6th dorsal-fin soft ray; anal-fin spine weak, slender;
15th or 16th (rarely 14th) soft ray longest. Caudal fin rounded, upper edge slightly pointed posteriorly. Pectoral fin elongate, with rounded posterior contour, 9th or 10th ray longest, its tip almost reaching vertical drawn through anal-fin
origin. Pelvic fin origin slightly anterior to level of pectoralfin base, pelvic-fin spine under skin closely attached to 1st
soft ray; 4th soft ray longest, its tip not reaching anus when
depressed in specimens of >79 mm SL, just reaching anus in
66.1 mm SL specimen (KAUM–I. 69435).
Fresh coloration. Body pale red, whitish ventrally (Fig.
1); scales on reddish portion of body tinged centrally withyellow; ca. 7 large, indistinct, dark Y- or V-shaped markings laterally on body, from behind pectoral fin to caudal
peduncle; indistinct rounded red blotch (blackish in smallest specimen, KAUM–I. 69435) on upper part of caudal-finbase, its size subequal to pupil. Head pale red; cheek with 6
or 7 radial rows of reddish or yellow spots (each cheek scale
tinged with reddish or yellow); straight, narrow yellow linefrom anteroventral edge of orbit to dorsal margin of upper
jaw; eye yellow or pink, tinged with dark red dorsally. Dorsal fin semi-translucent, membrane of soft-rayed portion
with ca. 5–8 narrow, wavy, yellow longitudinal stripes, these
becoming irregular and conjoined posteriorly. Anal fin pale
white, tinged with yellow basally; distal part broadly tinged
with red. Pectoral fin pale yellow. Pelvic fin pale red (white
in KAUM–I. 69435), basal part broadly blackish, ray tips
whitish. Caudal fin semi-translucent, with 5–9 radial yellow bands (or radial rows of small yellow spots), number
increasing and width narrowing with growth; ventral edge
tinged with red.
Distribution. Parapercis bicoloripes has heretofore beenrecorded only from the Gulf of Thailand, the South China
Sea, and the Sulu Sea, including Nha Trang and Phan Thiet
bays, Vietnam (Prokofiev 2008, 2010); the northern Gulf ofThailand; off the east coast of the Malay Peninsula and off
northern Borneo; and at Panay Island, the Philippines (thisstudy, Fig. 2). Although Mohsin and Ambak (1996: 537, 737,
figs 413, 860) recorded Ryukyupercis gushikeni (Yoshino,1975) as “Parapercis gushikeni (Yoshino)” from Malaysia
and adjacent waters, a photograph of their R. gushikeni had
the diagnostic coloration of P. bicoloripes (see below). Similarly, P. diplospilus and R. gushikeni recorded from Malaysia
by Ambak et al. (2010: 230, 231, unnumbered figs) can be
identified as P. bicoloripes.
Remarks. The original diagnosis of Parapercis bicoloripes by Prokofiev (2010) included: dorsal-fin rays V, 20 or
21; anal-fin rays I, 17 [18 soft rays in Prokofiev (2010)]; pectoral-fin rays 17; pored lateral-line scales 55–57; scale rows
above lateral line to first dorsal-fin spine base 3.5; frontal ca-

191

nines in lower jaw 6; total gill rakers 12–14; head and bodyentirely covered with ctenoid scales; fifth dorsal-fin spine
connected with first soft ray by membrane along most of its
length; upper lobe of caudal fin slightly elongate, pointed;
no dark markings on head, body, and median fins; pelvic
fin bicolored. The present specimens are largely consistent
with this diagnosis, with the following exceptions: pectoralfin rays [16–18 vs. 17 in Prokofiev (2010)], scale rows above
the lateral line (3.5 or 4.5 vs. 3.5), pored lateral-line scales(52–55 vs. 55–57), and total gill rakers (12–19 vs. 12–14).
The pored lateral-line scale count differences may have arisen from differing counting methods between the two studies. Prokofiev (2010) probably including several pored scales
extending onto the caudal fin. The other minor differencesin counts most likely represent intraspecific variation.
More notably, the present specimens differ from Prokofiev (2010) in most proportional measurements, including
head length (25.7–28.7% SL in the former vs. 25.2–26.4%
SL in the latter); pre-dorsal-fin length (26.2–30.6% SL vs.
23.9–28.4% SL); pre-anal-fin length (43.4–49.9% SL vs.
42.5–45.5% SL); pre-pelvic-fin length (23.1–25.7% SL vs.
20.9–23.2% SL); caudal-fin length (18.1–21.4% SL vs. 15.8–
18.1% SL); pectoral-fin length (19.2–22.0% SL vs. 16.9–
18.4% SL); pelvic-fin length (20.6–25.7% SL vs. 19.4–23.9%
SL); caudal-peduncle length (9.0–11.1% SL vs. 11.4–14.6%
SL); and caudal-peduncle depth (8.5–10.2% SL vs. 8.2–9.7%
SL). Of these, the minor differences in lengths of the headand pelvic fin, and in caudal-peduncle depth, very likely
represent individual variation within the species. Differences in pre-dorsal- and pre-anal-fin lengths may be explained by ontogenetic morphological changes (see below)
in as much as the material used in the two studies ranged
from 66.1 to 136.0 mm SL (present study) and from 88 to
155 mm SL (Prokofiev 2010). The remaining differences between the present specimens and Prokofiev (2010) (lengthsof the caudal peduncle, pre-pelvic-fin portion, and caudal
and pectoral fins) possibly resulted from different measuring techniques, details of which were not provided by Prokofiev (2010).
Notwithstanding the above-mentioned discrepancies,
the present specimens are identified here as P. bicoloripes,
based on their color pattern. Although 13 additional speciesof Parapercis were newly described subsequent to Prokofiev(2010), P. bicoloripes can be distinguished easily from all thecongeners by its unique coloration (Ho and Shao 2010; Liaoet al. 2011; Allen and Erdmann 2012; Ho et al. 2012, 2014;
Johnson and White 2012; Sparks and Baldwin 2012; Hoand Johnson 2013; Chen et al. 2014; Johnson et al. 2014; Ho2015).
Parapercis bicoloripes, having been previously reported
from Vietnam as Parapercis sp. 1 by Prokofiev (2008), wasdescribed by Prokofiev (2010) on the basis of 20 specimensfrom Nha Trang and Phan Thiet bays, Vietnam. The species
had also been recorded by Mohsin and Ambak (1996) andAmbak et al. (2010) from Malaysia and adjacent waters, but
was misidentified by them as P. diplospilus or R. gushikeni.
The present specimens, therefore, represent the first confirmed records of P. bicoloripes from Thailand, Malaysia, andMizuki Matsunuma et al.

192

the Philippines, indicating a widespread distribution of thespecies in the Gulf of Thailand, the South China Sea, andthe Sulu Sea (Fig. 2).
Morphological changes with growth. Analysis of the
26 examined specimens (66.1–136.0 mm SL) indicated thatseveral body proportions change with growth in P. bicoloripes. Relative lengths of the snout, bony interorbital space,and upper jaw tend to become greater with growth; in contrast, relative lengths of the fleshy orbit diameter, pre-dorsalfin portion, pre-anal-fin portion, second dorsal-fin spine,
anal-fin spine, and pelvic fin become shorter with growth(Fig. 3). No other significant growth-related proportionalchanges were apparent.
The blotch on the upper part of the caudal-fin base appeared to become paler and more indistinct with growth.
The smallest examined specimen (KAUM–I. 69435,
66.1 mm SL) had a distinct blackish-red blotch on the base(Fig. 1C), in contrast to larger specimens, which had apoorly defined red blotch (Fig. 1A–B). Moreover, the number and width of the yellow bands on the caudal fin become
greater and narrower, respectively, with growth. Specimensof <80 mm SL (KAUM–I. 69435 and 79764) possessed
about five caudal-fin bands, whereas six to nine bands were
present in specimens of >96 mm SL. The bands were relatively wide in the smallest examined specimen, about twice
as wide as the spaces between them (Fig. 1C). In contrast,
the bands of large specimens were relatively narrow, with amaximum width subequal to or clearly narrower than thespaces between them (Fig. 1A–B).
Although gill raker numbers varied considerably in P.
bicoloripes (12–19; Table 2), no significant relationshipwas recognized between their number and either standard
length or capture locality, the range in number apparently
being as aspect of individual variation.
Parapercis diplospilus Gomon, 1981
(Figs 2, 4A–B, 5; Tables 1–2)
Material examined. Eleven specimens, 44.2–74.1 mm
SL. Thailand: KAUM–I. 22880, 67.5 mm SL, northern
Gulf of Thailand, bottom trawl (obtained at Angsila fishingport, Chonburi Province); NSMT-P 55320, 3 specimens,
44.2–54.9 mm SL, off Songkla; NSMT-P 70071, 3 specimens,
66.5–70.4 mm SL, possibly Thailand (no further locality
data); NSMT-P 70944, 61.3 mm SL, Thailand (no further locality data); NSMT-P 115466, 55.8 mm SL, Thailand (no further locality data). Malaysia: KAUM–I. 16930, 54.1 mm SL,
off Kuala Terengganu, Terengganu, bottom trawl (obtainedat Cendering fishing port, Kuala Terengganu). Philippines:
KAUM–I. 80706, 74.1 mm SL, off Iloilo, Panay Island (obtained at Tigbauan Market, Iloilo), 11 November 2015.
Description. Morphometrics, expressed as percentagesof SL, and frequency distribution of selected meristics areshown in Tables 1–2. Dorsal-fin rays V, 22; anal-fin rays I,
18; pectoral-fin rays all branched except for uppermost ray;pelvic-fin rays I, 5; branched caudal-fin rays 15 (8 and 7 indorsal and ventral series, respectively); pseudobranchial filaments 10 (1) or 11 (1); branchiostegal rays 6.

Body subcylindrical, moderately elongate, deep, depth
at anal-fin origin 3.1–4.2 in SL; greatest width at pectoralfin base 3.4–5.0 in SL. Caudal peduncle short, deep, depth
0.8–1.0 in length. Head small, its length 1.6–2.7 in SL; snout
short, pointed, its length subequal to orbit diameter; eye relatively large, fleshy orbit diameter 3.3–4.0 in HL; interorbital
space narrow, least bony width 17.2–26.7 in HL.
Mouth small, slightly oblique, forming angle of about 20°
to horizontal axis of head; lower jaw slightly protruding beyond upper jaw tip; posterior margin of upper jaw reachingvertical drawn through anterior margin of pupil. Upper jaw
with outer row of 18–30 incurved caniniform teeth on eachside (number increasing with growth), anterior 2–4 caniniform teeth slightly larger, other teeth increasingly smaller
posteriorly; inner band of villiform conical teeth behindouter canine row of upper jaw, broadest anteriorly with 3–5
rows maximum, gradually narrowing posteriorly to 1 row
at rear of jaw. Front of lower jaw with usually 4 pairs of incurved enlarged, caniniform teeth (3 and 4 on left and rightsides, respectively, in KAUM–I. 16930; 7 or 8 in total) in distinctly separate outer row, innermost tooth usually smallest,
its size about half that of other teeth; inner band of villiform
conical teeth medial to caniniform teeth at front of lower
jaw, broadest portion with about 5 tooth rows, followed byrow of small, incurved conical teeth, 3–10 teeth in middle oftooth row slightly enlarged; vomer with crescentic patch of
2 or 3 rows of small conical teeth, subequal in size; palatine
teeth absent.
Gill membranes free from isthmus, with broad free fold;developed gill rakers short and spinous, longest raker about
one-fourth as long as longest gill filament. Nostrils small,anterior nostril with short membranous tube, its tip notreaching posterior nostril when laid back; posterior nostril
pore ovate or elliptical with slightly raised rim.
Pores of cephalic sensory system connected by canals
under skin. Row of 3–4 pores above maxilla; 3 pores near
nostrils, including 1 pore above and 1 below posterior
nostril, 1 between nostrils; 2 pores on anterior margin of
snout; row of 11–14 pores in infraorbital series, with several
branches in posterior portion; 10 or 11 pores in row on posterior margin of preopercle; interorbital space with 2–5 pairsof pores on anterior half, small pores scattered medially on
posterior half; occiput with numerous small pores; ventral
surface of lower jaw with 5 pores on each side, anteriormost
pore behind chin.
Opercle bearing strong, posteriorly-directed, flattened
spine, approximately at level of ventral edge of orbit in lateral view; posterior margin of subopercle well expanded
posteriorly, with 13–19 serrae, upper serrae relatively large,
number of serrae increasing with growth; posterior edge ofinteropercle with 2 or 3 weak serrae; preopercle rounded,
posteroventral edge with about 6 serrae; posttemporal areawith small exposed bony cleft with several indistinct bonyprotuberances on its posterior margin.
Lateral line continuous, about 3 pored lateral-line scales
extending onto caudal fin; body entirely covered with developed ctenoid scales, scales on dorsolateral portion withca. 40 cteni; head, including cheek, preopercle, interopercle,

Two Parapercis species from the South China Sea

193

Fig. 4. Fresh specimens of Parapercis diplospilus from Thailand (A) and the Philippines (B), and P. ommatura from Japan (C). A, KAUM–I.
22880, 67.5 mm SL, northern Gulf of Thailand; B, KAUM–I. 80706, 74.1 mm SL, off Iloilo, Panay Island; C, BSKU 81924, 84.6 mm SL, Tosa
Bay, Kochi Prefecture.

Fig. 5. Smallest examined specimen of Parapercis diplospilus(preserved). NSMT-P 55320, one of three specimens, 44.2 mm SL,
off Songkla, Thailand.

subopercle, and opercle, with ctenoid scales; no scales on
snout, interorbital space, occipital area, or both jaws; basesof pectoral and caudal fins covered with small, elongate ctenoid scales; dorsal, anal, and pelvic fins without basal scales.
Origin of dorsal fin posterior to vertical drawn throughupper origin of pectoral fin; dorsal-fin spines relatively
short; 4th dorsal-fin spine longest; membrane between 5th
spine and 1st soft ray attached near tip of 5th spine, deeply incised; anterior soft rays subequal in length, 3rd or 4th
ray longest. Origin of anal fin below base of 5th dorsal-fin

soft ray; anal-fin spine weak, slender; 4th soft ray longest.
Caudal fin truncate to slightly rounded, upper edge (andlower edge in KAUM–I. 80766) slightly pointed. Pectoral fin
slightly elongate, with rounded posterior contour, 7th or 8th
ray longest, its tip reaching vertical drawn through anal-fin
origin. Pelvic fin origin anterior to vertical drawn throughpectoral-fin base, spine covered with skin closely attachedto first soft ray; 4th soft ray longest, depressed tip almost
reaching anus.
Fresh coloration. Body creamy-white, whiter ventrally,
darkening dorsally and laterally. (Fig. 4A–B); lower half oflateral body surface above anal fin with transverse row ofabout 6 large, brown blotches. Head creamy-white, postorbital region and nape brownish. Snout with indistinct narrow brown band (width about half pupil diameter) extending from anterior margin of orbit to tip of snout. Cheek
creamy-white, with 2 brown blotches below eye; anterior
blotch small, situated between ventral margin of orbit andposterior tip of upper jaw; posterior blotch below postero-

194

Mizuki Matsunuma et al.

ventral margin of orbit, with slight yellowish tinge. Eye yellow to dull orange, tinged with red posterodorsally; pupil
black. Opercle generally brownish. Membrane of spinous
portion of dorsal fin pale yellow; membrane between 1st
and 3rd spines brownish. Soft-rayed portion of dorsal fin
with pale yellow membrane and about 3 poorly definedbrownish stripes. Anal-fin semi-translucent, membranes
between each ray with brownish tinge. Pectoral-fin membrane semi-translucent, rays with brownish tinge, base withoblique narrow brown band. Pelvic fin pale yellow, with numerous minute melanophores throughout; spine and tips
of soft rays white. Two small dark brown ocellated spots on
caudal-fin base, subequal to pupil in size. Caudal-fin semitranslucent, with ca. 3 narrow, dull orange bands; posterior
margin with narrow zone of dark orange tinge.
Distribution. Parapercis diplospilus is distributed in
the southeastern Indian and western Pacific oceans (Fig.
2): Nha Trang Bay, Vietnam (Prokofiev 2008); Gulf of Thailand (this study); off Terengganu, Malaysia (east coast of
Malay Peninsula) (this study); Visayan Islands, the Philippines (Gomon 1981; this study); Kaimana, West Papua, Indonesia (Allen and Erdmann 2009); Milne Bay, Papua New
Guinea (Allen and Erdmann 2012); and Australia, including
Queensland (Johnson 1999; Robins and McGilvray 1999),
Northern Territory (Larson et al. 2013), and Western Australia (Hutchins 2001, 2003; Allen et al. 2006). Although
Ambak et al. (2010) recorded the species from Malaysia,
their photograph is re-identified here as P. bicoloripes (seeabove).
Remarks. The present specimens, from Thailand, Malaysia, and the Philippines, agree closely with the originaldescription and figures of P. diplospilus given by Gomon(1981), including the diagnostic characters of the species[dorsal-fin rays V, 22, with last spine connected by deeply
incised membrane to first soft ray; anal-fin rays I, 18; pectoral-fin rays 15; scale rows below lateral line 10; circumpeduncular scale rows 23; 7 or 8 (usually 4 pairs) of caniniform teeth in front part of lower jaw; and 2 prominent
dark spots on caudal-fin base]. They differ slightly from
Gomon (1981) in having a slightly longer fourth dorsal-fin
spine, 7.1–8.2% SL (vs. 6.4–7.8% SL in the latter), regarded
here as intraspecific variation. Gomon (1981) included a detailed comparison of P. diplospilus with P. ommatura Jordan
and Snyder, 1902 (Fig. 4C), which is similar to the former inoverall body appearance. Parapercis diplospilus can be readily distinguished from P. ommatura by the two dark spots on
the caudal-fin base [vs. a single (upper) spot in the latter]and blotches on the cheek (vs. longitudinal stripes) (Gomon
1981; Fig. 4).
Parapercis diplospilus, originally described by Gomon(1981) on the basis of 21 specimens from the Philippine
islands, has subsequently been recorded from many localities in the southeastern Indian and western Pacific oceans,
although not from the Gulf of Thailand or east coast of the
Malay Peninsula. The present specimens, therefore, represent the first records of the species from these two localities.
The record of P. diplospilus from Nha Trang Bay (Prokofiev 2008) may need to be reviewed, as the five Vietnamese

specimens described differed significantly from Gomon’s(1981) description in having 17 or 18 pectoral-fin rays (vs.
14–16 in the latter), six caniniform teeth on the front partof the lower jaw (vs. seven or eight), and a relatively greater pelvic-fin length of 25.6–27.5% SL (vs. 19.9–23.6% SL).
Of these characters, dentition has been treated as an important diagnostic feature of Parapercis species (Cantwell1964). In addition, Prokofiev (2008) considered the difference in pelvic-fin length of the Vietnamese specimens as
likely size-related intraspecific variation [the latter specimens being smaller, 37–45 mm SL vs. 48.7–78.0 mm SL in
Gomon (1981)]. However, the smallest specimen examined
here (44.2 mm SL, one of NSMT-P 55320; Fig. 5) clearly had
relatively shorter pelvic fins than the Vietnamese specimens(23.0% SL in the former vs. 25.6–27.5% SL in the latter).
Moreover, Prokofiev (2008) stated that the posterior tip ofthe depressed pelvic fin extended well beyond the anal-fin
origin in the Vietnamese specimens, such not being so inthe present smallest specimen. However, the relationship between pelvic-fin length and standard length can not be considered further here due to the limited material available.
Comments on records of Parapercis ommatura. Shima­da (2013) considered Parapercis ommatura Jordan and Snyder, 1902 to be distributed in the northwestern
Pacific Ocean, including China, Taiwan, and Japan. Although Prokofiev (2008) recorded P. ommatura from Nha
Trang Bay, Vietnam, Liao et al. (2011) included P. ommatura sensu Prokofiev (2008) in the synonymy of their new
species, Parapercis lutevittata Liao, Cheng, and Shao, 2011.
Recently, Ray et al. (2015) recorded P. ommatura, based on
21 specimens (71.9–100.0 mm SL), from the northern Bay
of Bengal, off West Bengal, India. These specimens differedslightly from P. ommatura [as redescribed by Imamura
and Yoshino (2007) on the basis of 16 specimens (64.5–
101.9 mm SL), including the holotype of the species, fromJapan and China] in having 13–14 gill rakers (vs. 8–13 in thelatter) and 61 or 62 lateral-line scales [vs. 56–58; 56–60 in
Shimada (2013)]. Moreover, the Indian specimens also differed from P. ommatura from the northwestern Pacific (Shinohara 1997; this study, Fig. 1C) in coloration, including
bright orange stripes on the cheek (Ray et al. 2015: fig. 1)(vs. brownish in the latter) and a small, black spot on eachmembrane between the third and fifth dorsal-fin spines (vs.
absent). Accordingly, the Indian specimens recorded by Ray
et al. (2015) may represent a species distinct from P. ommatura.
Comparative material. Parapercis diplospilus: UPVMI
316, 57.5 mm SL, off Iloilo, Panay Island, Philippines (obtained at Central Market, Iloilo City) (photograph only).
Parapercis ommatura: BSKU 81924, 84.6 mm SL, Tosa Bay,
Kochi Prefecture, Japan, 30 m depth, RV Toyohata-maru.

Acknowledgments
We are deeply grateful to P. Musikasinthorn, S. Ratmuangkhwang, P. Sreesamran, S. Arbsuwan, S. Ekajit, andN. Ngamtampong (Kasetsart University, Bangkok); N. Muto

Two Parapercis species from the South China Sea

(formerly Research Institute for Humanity and Nature,
Kyoto); K. Koeda, T. Yoshida, H. Hata, K. Eguchi, and A.
Yoshiura (KAUM); staff of the South China Sea Repository
and Reference Center, Institute of Oceanography and Environment, and the School of Fisheries and Aquaculture Sciences, Universiti Malaysia Terengganu; and U. Alama andother staff of UPVM for their generous assistance. We also
gratefully thank K. Sasaki, H. Endo, T. Naito and M. Sato
(BSKU); N. Nakayama and R. Asaoka (formerly BSKU);
K. Matsuura, G. Shinohara, M. Nakae, E. Katayama, andK. Kuriiwa (NSMT); and S. Chiba (formerly NSMT) for
their kind hospitality during the first author’s visit to their
institutions; H. Endo for providing a photograph of P.
ommatura; and students and volunteers of KAUM for curatorial assistance. G. Hardy (Ngunguru, New Zealand) readthe manuscript and provided help with English. This study
was conducted under a Memorandum of Agreement forjoint research made by and among the Department of Agriculture of the Republic of the Philippines (DA), the University of the Philippines-Visayas (UPV), the Kagoshima
University Museum, the Research Institute for Humanity
and Nature, and Tokai University, facilitated by S. L. Sanchez [Bureau of Fisheries and Aquatic Resources (BFAR),
DA]. P. J. Alcala (DA) providing a Prior Informed Consent
Certificate and I. P. Cabacaba and S. M. S. Nolasco (BFAR,DA) providing a fish specimen Export Certificate (No. 201639812). We thank the staff of Office of the Vice-Chancellor
for Research and Extension, UPV, UPV Museum of Natural
Sciences, College of Fisheries, UPV, including S. S. Garibay,
U. B. Alama, V. G. Urbina, L. H. Mooc, C. J. N. Rubido, E.
P. Abunal, A. M. T. Guzman, R. S. Cruz, A. C. Gaje, and R.
F. M. Traifalgar, and graduate students of College of Fisheries, UPV for their support to this research collaboration. This study was supported in part by JSPS KAKENHI
Grant Numbers JP26241027, JP24370041, JP23580259, andJP26450265; a JSPS Fellow grant (PD: 16J00047); the JSPS
Core-to-Core Program, “Research and Education Network
on Southeast Asian Coastal Ecosystems”; the “Coastal Area
Capability Enhancement in Southeast Asia Project” of the
Research Institute for Humanity and Nature, Kyoto, Japan;
the “Biological Properties of Biodiversity Hotspots in Japan”
project of the National Museum of Nature and Science, Tsukuba, Japan; and the “Establishment of Research and Education Network on Biodiversity and Its Conservation in theSatsunan Islands” project of Kagoshima University adopted
by the Ministry of Education, Culture, Sports, Science andTechnology, Japan.

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