Bull. Natl. Mus. Nat. Sci., Ser. A, Suppl. 4, pp. 21–29, March 29, 2010

A New Species of the Flathead Genus Inegocia(Teleostei: Platycephalidae) from East Asia
Hisashi Imamura1,2
1

Fisheries Science Center, the Hokkaido University Museum,
3–1–1 Minato-cho, Hakodate, Hokkaido 041–8611, Japan
2
Present address: Laboratory of Marine Biology and Biodiversity (Systematic Ichthyology),
Faculty of Fisheries Sciences, Hokkaido University,
3–1–1 Minato-cho, Hakodate, Hokkaido 041–8611, Japan
E-mail: imamura@fish.hokudai.ac.jp

Abstract A new species of platycephalid, Inegocia ochiaii, is described based on specimens,
ranging from 47.2–444.0 mm SL, collected from southern Japan and Taiwan. Inegocia ochiaii differs from other platycephalids in having the following combination of characters: IVIII first dorsal-fin rays, usually 11 second dorsal and anal-fin rays, 63–76 oblique body scale rows slanting
downward and backward above the lateral line, a long snout (31.7–34.9% HL), a long andbranched iris lappet, the posterior margin of the orbit lacking distinct pit, the suborbital ridge with
2 spines (preorbital spine absent), a single large interopercular flap present, sensory tubes from thesuborbitals and preopercle not covering cheek region, and scales anteriorly on the lateral line witheither single posterior opening or with 2 openings and scales posteriorly on the lateral line with 2
openings.
Key words : Platycephalidae, New species, Inegocia, East Asia.

Cuvier in Cuvier and Valenciennes (1829)
originally described a platycephalid, Platycephalus guttatus, based on a single specimen
collected from Nagasaki, Japan. After the original description, Temminck and Schlegel (1843)
also described the species and illustrated thespecies, indicating many small dark dots on thedorsal surface of the head and body. Matsubara
and Ochiai (1955) gave detailed description thespecies based on Japanese specimens and placed
it into the genus Inegocia Jordan and Thompson,
1913, owing to the long and branched iris lappet
and well developed interopercular flap. Many authors (e.g., Anonymous, 1962; Ochiai, 1984;
Nakabo, 1993; Imamura, 1997; Kim et al., 2005)
subsequently reported the species from southern
Japan to the East and South China seas under thename I. guttata. However, others considered thatthe species is a junior synonym of Cociella crocodila (Cuvier, 1829) (e.g., Jordan and Richardson, 1908; de Beaufort and Briggs, 1962). Re-

cently, Imamura and Yoshino (2009) demonstrated the conspecificity of C. crocodila and P. guttatus, and that the species from southern Japan to
the East and South China seas commonly recognized as I. guttata was undescribed. The species
previously misidentified as I. guttata is herein described as new.
Materials and Methods
Counts and measurements were made according to Hubbs and Lagler (1958) and were routinely taken from the left side, while gill rakers
were counted on the right side. A small first dorsal spine was expressed by using plus sign. Measurements were made with calipers to the nearest
0.1 mm accuracy. Terminology of head spines
follows Knapp et al. (2000). Institutional
acronyms are from Eschmeyer (1998), except
for Hokkaido University Museum, Hakodate(HUMZ) and National Museum of Nature and22Hisashi Imamura

Science, Tokyo (NSMT). Standard and headlengths are abbreviated as SL and HL, respectively.
Inegocia ochiaii sp. nov.
[Japanese name: Wani-gochi]

Wakasa Bay southward to East China Sea); Nakabo,
2002: 617, unnumbered fig. (Tosa Bay and Wakasa
Bay southward to East China Sea); Kim et al., 2005:
235, fig. (Cheju Island, Korea).
Platycephalus guttatus (not Cuvier in Cuvier and Valenciennes, 1829): Temminck and Schlegel, 1843: 39, pl.
15, fig. 2 (bay of Nagasaki, Japan).

(Figs. 1–4)
Inegocia guttata (not Cuvier in Cuvier and Valenciennes,
1829): Matsubara and Ochiai, 1955: 82, pl. 1 [southern
Japan, including Maizuru (Kyoto Prefecture), Nobeoka(Miyazaki Prefecture), Urado (Kochi Prefecture) andEast China Sea]; Matsubara, 1955: 1121 (southern
Japan, including Maizuru, Nobeoka, Urado, Nagasaki
and East China Sea); Anonymous, 1962: 256, fig. 64-7
(Haimen, China, South China Sea); Masuda et al.,
1975: 342, pl. 146-E (central Honshu southward to
China Sea); Ochiai, 1984: 321, pl. 288-K (Tosa andWakasa Bays, Japan, to the South China Sea); Yamada,
1986: 331 (East China Sea); Lindberg and Krasyukova,
1987: 153, fig. 90 (Sea of Japan); Shao and Chen,
1987: 88, fig. 27 (Taiwan); Shao and Chen, 1993: 256,
fig. 64-7 (Taiwan); Nakabo, 1993: 537, unnumberedfig. (Tosa Bay and Wakasa Bay, Japan, southward to
East China Sea); Masuda and Kobayashi, 1994: 91, fig.
5 (Sagami and Wakasa Bays, Japan, to South China
Sea); Lee and Joo, 1995: 115, figs. 1–2 (Cheju Island,
Korea); Yamada et al., 1995: 119, fig. (southern Japan,
East and South China seas); Imamura, 1997: 219(southern Japan to South China Sea); Lee and Joo,
1998: 222, fig. 8 (Pusan and Cheju Island, Korea);
Nakabo, 2000: 617, unnumbered fig. (Tosa Bay andHolotype. BSKU 48578, 272.4 mm SL, Kashiwajima Island, Kochi Prefecture, Japan (35.2°N, 132.7°E),
hand net, 28 Nov. 1990.
Paratypes. 8 specimens. BSKU 4176, 138.1 mm SL,
Usa, Tosa, Kochi Prefecture, Japan, Aug. 1954; BSKU
37647, 258.8 mm SL, Mogi Bay, Nagasaki Prefecture,
Japan, 16 Oct. 1982; BSKU 60856, 94.5 mm SL, Uranouchi Bay, Tosa, Kochi Prefecture, Japan, 4 m depth, 8
Dec. 2002; BSKU 94659, 423.1 mm SL, fish market, Mimase, Kochi Prefecture, Japan, 25 Apr. 2009; BSKU
95332, 444.0 mm SL, fish market, Mimase, Kochi Prefecture, Japan, 7 March 2008; HUMZ 193076, 329.3 mm SL,
fish market, Taipei, Taiwan, 19 May 2005; NSMT-P 5772,
236.2 mm SL, Aso Bay, Tsushima Islands, Nagasaki Prefecture, Japan, 12–14 July 1968; NSMT-P 54160, 261.8mm SL, Suno-saki Point, southern part of Boso Peninsula,
Chiba Prefecture, Japan, 3 m depth, hand net, 15 July
1992.
Non type. NSMT-P 35729, 47.2 mm SL, Sokodo,
northeast coast of Hachijo-jima Island, Izu Islands, Japan(33°7N, 139°49E), 3 m depth, hand net, 15 July 1992.

Diagnosis. A new species of Inegocia withIVIII first dorsal-fin rays, usually 11 second

Fig. 1. Dorsal (upper) and lateral (lower) views of Inegocia ochiaii sp. nov., BSKU 48578, 272.4 mm SL, Kashiwa-jima Island, Kochi Prefecture, Japan (ethanol preserved condition) (photos by H. Imamura).

New Inegocia from East Asia

dorsal-fin rays, 11 anal-fin rays, 63–76 oblique
body scale rows slanting downward and backward above lateral line, a long snout (31.7–
34.9% HL), a long and branched iris lappet, posterior portion of orbit lacking distinct pit, suborbital ridge with 2 spines (preorbital spine absent),a single large interopercular flap present, sensory
tubes from suborbitals and preopercle not covering cheek region, and scales anteriorly on lateralline with either single posterior opening or with 2
openings, scales posteriorly on lateral line with 2
openings.
Description. Data for the holotype are presented first, followed by paratype data in parentheses or brackets (occasionally non type dataalso shown when necessary): first dorsal-fin rays
IVIII (IVIII); second dorsal-fin rays 11 (11);anal-fin rays 11 (10 in 1, 11 in 7); pectoralfin rays 2 (upper, unbranched)13 (middle,
branched)8 (lower, unbranched)23 [1–3 (usually 2)10–147–921–22 (1 paratype, BSKU
4176, abnormally with 37616; 1 non typewith 20 unbranched rays)]; pelvic-fin rays I, 5
(I, 5); branched caudal-fin rays 6 (upper)6(lower)12 (5 or 64–610–12); scales in lateral line 53, anterior 2 scales with spine [52–54,
2 or 3 with spine]; oblique body scale rows slanting downward and backward above lateral line
74 (66–76); oblique body scale rows slanting
downward and forward above lateral line 105(95–107); gill rakers 156 (145 in 1,
156 in 7). Proportions as % SL: HL 35.8
(35.4–38.0); predorsal length 36.9 (36.8–39.1);
length of first dorsal-fin base 19.0 (18.5–20.6);
length of second dorsal-fin base 26.2 (25.8–
27.7); length of anal-fin base 28.2 (27.2–29.7);
caudal peduncle length 10.2 (9.1–10.8); caudal
peduncle depth 4.7 (4.6–5.1); snout length 11.5
(11.7–12.6); orbital diameter 6.6 (6.3–8.3); upper
jaw length 13.1 (12.8–13.9); lower jaw length20.4 (20.0–21.9); interorbital width 3.0 (2.2–4.0); pectoral-fin length 14.2 (13.4–16.3); pelvicfin length 25.1 (23.5–28.0); caudal-fin length18.9 (17.2–22.1); length of first spine of first dorsal fin 2.9 (2.1–3.5); length of second spine offirst dorsal fin 13.2 (13.9–15.3); length of first23

ray of second dorsal fin 12.7 (11.7–13.4); lengthof first anal-fin ray 6.9 (6.7–8.1). Proportions as% HL: snout length 32.1 (31.7–34.1); orbital diameter 18.6 (16.5–22.2); upper jaw length 36.7
(36.1–37.0); lower jaw length 57.0 (54.9–59.3);
interorbital width 8.3 (6.0–10.8).
Body depressed, mostly covered with ctenoid
scales, some cycloid scales on undersurface.
Head flattened, length 2.8 (2.6–2.8) in SL; postorbital region, opercle and nape scaled. Snout
slender, much longer than orbital diameter, snout
length 3.1 (2.9–3.2) in HL. Upper surface of eye
without papillae. Iris lappet long and wellbranched, and not attaining center of eye dorsally,broad and simple ventrally (Fig. 3A) (eachbranch short and simple in a non type in 47.2 mm
SL, Fig. 3B). Interorbit moderately broad andshallowly concave, width 12.0 (9.2–14.8) in HL.
Posterior margin of orbit lacking distinct pit. Top
and side of head possessing spines. Nasal withsingle small spine. Anteroventral margin oflachrymal without spines. Single sharp preocularspine present, lacking small spines on its anteriorbase (Fig. 2). Preorbital spines absent. Suborbital
ridge with 2 spines below eye. Supraorbital ridge
roughly serrated posteriorly (Fig. 2). Postorbital
region, pterotic, parietal, supratemporal, supracleithrum and left side of posttemporal with single spine, respectively; right side of posttemporalwith 2 spines. Preopercle with 2 spines [or 3 (single smallest lowermost spine added) in right sideof BSKU 4176 and both sides of BSKU 60856];
upper longest, not reaching posterior margin of
opercle, bearing single supplementary spine.
Ridge of upper and lower opercular spines without serrations. Single large interopercular flap
present. Maxilla reaching anterior margin of eye,length 2.7 (2.7–2.8) in HL. Villiform teeth inbands on palatine and jaws, except for anteromedial portion of upper jaw with small and slenderconical teeth; tooth band on upper jaw with distinct notch medially. Small conical teeth in 2 separate patches on vomer. Lip margins without
papillae. Sensory tubes from suborbitals and preopercle undeveloped, not covering cheek region.
Scales anteriorly on lateral line with either single

24

Hisashi Imamura

Fig. 3. Lateral view of iris lappet of Inegocia
ochiaii sp. nov. A, BSKU 37647, paratype,258.8 mm SL, Mogi Bay, Nagasaki Prefecture,
Japan; B, NSMT-P 35729, non type, 47.2 mm
SL, Sokodo, northeast coast of Hachijyo-jimaIsland, Izu Islands, Japan.
Fig. 2. Dorsal view of head of Inegocia ochiaii
sp. nov., BSKU 37647, paratype, 258.8 mm
SL, Mogi Bay, Nagasaki Prefecture, Japan.

posterior opening or with 2 openings, scales posteriorly on lateral line with 2 openings. First dorsal fin originating slightly posterior to opercularmargin. First and second dorsal fin narrowly separated. Pectoral fin rounded posteriorly, length
2.5 (2.3–2.8) in HL. Posterior tip of pelvic fin attaining anal-fin origin (position of posterior tip ofpelvic fin ranging from anterior to anal fin tobase of third ray), pelvic-fin length 1.4 (1.3–1.6)
in HL. Caudal fin slightly rounded posteriorly,
length 1.9 (1.7–2.2) in HL.
Color in alcohol. In holotype (Fig. 1), dorsal
and lateral portions of head and body brown,densely scattered with small blackish dots; lower
side of head and body pale brown. Head with 2

dark brown bands dorsally, crossing eyes and opercular region, respectively; posterior band indistinct. Body with 7 dark brown bands dorsally; 2indistinct and 1 distinct bands below first dorsal
fin, 1 indistinct and 2 distinct bands below second dorsal fin; and 1 distinct band on caudal peduncle. First dorsal fin with broad marginal darkbrown band; anteroventral region of first dorsal
fin with several dark brown spots. Second dorsal
fin with many dark brown spots; several spots
somewhat continuous and tending to form
oblique bands. Pectoral fin with small brown todark brown spots. Pelvic fin with large brown todark brown spots; several spots with blackish
center. Anal fin pale basally and dusky distally.
Caudal fin with brown to dark brown spots ofvarious sizes; several spots with blackish center;narrow and broad dark brown bands on basal andNew Inegocia from East Asia

25

Fig. 4. Underwater photograph of Inegocia ochiaii sp. nov., showing whole body (A) and close up of head (B)(specimen not preserved, photos by H. Endo).

distal portions, respectively; posterior margin
narrowly pale.
Color when alive (based on underwater photographs, Fig. 4). Dorsal and lateral portions ofhead and body grayish, mottled by small brownish and blackish spots, and densely scattered withsmall blackish dots. Head with 2 indistinct bands

formed by pale brownish to brownish spots dorsally, crossing eyes and opercular region, respectively; distinct dark brownish marking present
below eye. Body with 6 bands dorsally; 2 indistinct pale brownish bands below first dorsal fin;
1 indistinct pale brownish and 2 distinct darkbrownish bands below second dorsal fin; and26

Hisashi Imamura

1distinct blackish band on caudal peduncle. First
dorsal fin with 1 blackish marginal marking andseveral dark brown and lighter brown spots. Second dorsal fin with many brownish spots. Pectoral fin with small brownish to blackish brownish spots. Caudal fin with pale brownish to darkbrownish spots of various sizes.
Distribution. Known only from East Asia,
ranging from southern Japan to the East andSouth China seas, including Sagami and Wakasa
Bays, Hachijo-jima Island, southern Korean
Peninsula, Taiwan and China (e.g., Matsubara
and Ochiai, 1955; Anonymous, 1962; Nakabo,
2002; Kim et al., 2005; this study).
Etymology. Named in honor of Dr. Akira
Ochiai, for his many contributions to fish taxonomy, including those for the Platycephalidae.
Remarks. Inegocia ochiaii has been mistakenly identified as Platycephalus guttatus, ajunior synonym of Cociella crocodila (see Imamura and Yoshino, 2009) known from southern
Japan to the South China Sea. However, I. ochiaii
is easily separable from C. crocodila in havingsuch characters as a single large interopercular
flap (vs. interopercular flap absent in C. crocodila), and scales anteriorly on the lateral line witheither single posterior opening or with 2 openings and scales posteriorly on the lateral line with
2 openings (all lateral-line scales with single
opening), and preorbital spine absent (present).
In addition, I. ochiaii has a longer snout thanC. crocodila (31.7–34.9% HL in I. ochiaii vs.
26.6–29.3% HL in C. crocodila, Fig. 5) (data forC. crocodila from Knapp, 1996, Imamura andYoshino, 2009 and this study).
Inegocia ochiaii resembles 2 congeners, I. harrisii (McCulloch, 1914) and I. japonica (Cuvier,1829) (see Imamura and Yoshino, 2009 forauthorship of I. japonica) in having a long andbranched iris lappet dorsally, lacking a distinct
pit on the posterior margin of the orbit, and lacking sensory tubes from the suborbitals and preopercle on the cheek region. However, I. ochiaii iseasily distinguishable from these 2 species inhaving a greater number of oblique body scalerows slanting downward and backward above theFig. 5. Relationship of snout length (%HL) andstandard length (mm) in Inegocia ochiaii sp.
nov. (solid circle) and Cociella crocodila (opencircle). Asterisks on right shoulder of symbol
indicate holotype.

lateral line (63–76 in I. ochiaii vs. 52–55 in I.
harrisii and 53–55 in I. japonica). Inegocia ochiaii has a large interopercular flap, whereas I. harrisii lacks the flap and I. japonica possesses a finger-like flap. Inegocia ochiaii has scales anteriorly on the lateral line with either a single posterior
opening or with 2 openings, and scales posteriorly on the lateral line with 2 openings. In contrast,
all lateral-line scales have 2 openings in I. harrisii and I. japonica. In addition, Inegocia ochiaii
differs from I. japonica in having a fewer numberof second dorsal and anal-fin rays (usually both
11 in I. ochiaii vs. usually both 12 in I. japonica)(data for I. harrisii and I. japonica from Knapp,
1999 and this study).
Differences between Inegocia ochiaii andCymbacephalus bosschei (Bleeker, 1860), known
from the Philippines to northern Australia, andLeviprora inops (Jenyns, 1840), known fromsouthern Australia, are also discussed here because the latter 2 species possess 3 characters incommon with the 3 species of Inegocia. Cymbacephalus bosschei had been placed into the genus
Inegocia by several authors (e.g., Gloerfelt-Tarp
and Kailola, 1984; Imamura, 1996) until Knapp(1999) put it into the genus Cymbacephalus
Fowler, 1938. Inegocia ochiaii is easily separablefrom C. bosschei and L. inops in having 11 second dorsal and anal-fin rays, and the suborbital

New Inegocia from East Asia

ridge with 2 spines (vs. 12 second dorsal andanal-fin rays in C. bosschei and L. inops, andsuborbital ridge with single spine in C. bosschei
and with no spines in L. inops). In addition, theinteropercular flap and lateral-line scales are also
helpful in separating I. ochiaii from C. bosschei(broad interopercular flap with shallow incisions,and all lateral-line scales with single opening
posteriorly in C. bosschei). Inegocia ochiaii isalso distinguishable from L. inops by having
IVIII first dorsal-fin rays (IVII in L. inops)(data for C. bosschei from Knapp, 1999 and thisstudy, and those for L. inops from Kuiter, 1994
and this study).
Comparative materials. Cociella crocodila(13 specimens, all from Japan, including holotype of P. guttatus, ZMB 726, 355.9 mm SL, see
Imamura and Yoshino, 2009 for data of examined
specimens, except for newly added NMW 74721,
397.8 mm SL). Cymbacephalus bosschei (7specimens, all from northern Australia): AMS
I.33461-021, 191.2 mm SL; AMS IB.4093, 133.7mm SL; NTM S.12444-022, 232.3 mm SL; NTM
S.12892-005, 104.6 mm SL; NTM S.12962-005,
205.6 mm SL; NTM S.14472-017, 139.6 mm SL;
NTM S.14665-005, 152.4 mm SL; NTM
S.15532-005, 132.7 mm SL. Inegocia japonica(20 specimens, whereabouts of types unknown,see Imamura and Yoshino, 2009 for data of examined specimens); Inegocia harrisii (8 specimens, all from northern Australia): AMS E.2844(lectotype of Insidiator harrisii), 169.7 mm SL;
AMS I.20473-009, 107.4 mm SL; AMS I.21830006, 168.9 mm SL; AMS I.34398-038, 102.5 mm
SL; NTM S.14354-008, 124.7 mm SL; NTM
S.14356-004, 175.7mm SL; NTM S.14362-011,
145.8 mm SL. Leviprora inops (5 specimens, allfrom southern Australia): BMNH 1917.7.14.76(holotype of Platycephalus inops), 340.1 mm SL;
CSIRO C2365, 292.0 mm SL; WAM P.27590027, 260.8 mm SL; WAM P.27881-002, 148.6mm SL; WAM P.28016-001, 240.8 mm SL.
Acknowledgments
I express my sincere thanks to T. Iwamoto

27(CAS) for critically reading a draft manuscript
and providing important comments. I am gratefulto P. Bartsch (ZMB), H. Endo (BSKU), A. Graham (CSIRO), H. Larson, G. Dally and R.
Williams (NTM), J. Maclaine (BMNH), K. Matsuura and G. Shinohara (NSMT), M. McGrouther (AMS), S. Morrison (WAM) and H.
Wellendorf (NMW) for providing the opportunity
to examine specimens. H. Endo kindly provided
color photographs of I. ochiaii. Finally, I sincerely thank K. Matsuura for the opportunity to publish this description of the new species. This
study was partly supported by an Ito Grant for
Ichthyology, Fujiwara Natural History Foundation to the author.
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Manuscript received 16 June 2009; revised 29 July 2009;
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Associate editor: K. Matsuura.