phuket mar. biol. cent. res. bull.76: 9–17 (2019)

9
Phuket mar. biol. Cent. Res. Bull.76: 9–17 (2019) DESCRIPTION OF A NEW SPECIES OF Bembradium (SCORPAENIFORMES: BEMBRIDAE) FROM THE ANDAMAN SEA, THAILAND Saki Kishimoto 1 *, Toshio Kawai 2 , Fumihito Tashiro 3 and Charatsee Aungtonya 4 1 Graduate School of Fisheries Sciences, Hokkaido University, Hakodate, Hokkaido 041-8611, Japan 2 Faculty of Fisheries Sciences, Hokkaido University Museum, Hakodate, Hokkaido 041-8611, Japan 3 Fisheries Science Center, Hokkaido University Museum, Hakodate, Hokkaido 041-8611, Japan 4 Phuket Marine Biological Center, Muang District, Phuket 83000, Thailand *Corresponding author: [email protected] ABSTRACT: A new species of Bembradium (Scorpaeniformes: Bembridae) is described on the basis of a single specimen collected in 284–300 m in the Andaman Sea off Phuket, Thailand. The new species, Bembradium magnoculum, differs from all congeners by the following combination of characters: short and wide snout (length 79.9% eye diameter; width at anterior margin of anterior nostril 13.4% SL), large eye (13.2% SL), long upper jaw (20.1% SL), long preanus length (64.8% SL), long predorsal-fin length (41.9% SL), short second dorsal-fin base (57.1% HL), long anal-fin base (23.9% SL), short lower caudal peduncle length (25.8% HL), short pectoral fin (45.9% HL), short pelvic-fin spine (16.0% HL), small spines of similar size on nasal bone and broad dermal flap of anterior nostril. Keywords: Pisces, Bembradium magnoculum sp. nov., Indian Ocean, BIOSHELF INTRODUCTION The genus Bembradium Gilbert, 1905, placed in the family Bembridae, was established for Bembradium roseum Gilbert, 1905, based on a specimen captured off Hawaii. A second species, Bembradium furici Fourmanoir and Rivaton, 1979, was described from a New Caledonian specimen. Since then, this genus has included only the two species, B. roseum recorded from Hawaii, Emperor Seamount Chain, Japan, Taiwan, and New Caledonia, in depths of 138–800 m (Gilbert, 1905; Chave and Mundy, 1994; Rivaton and Bourret, 1999; Yamada et al., 2007; Ho et al., 2009; Fricke et al., 2011; Nakabo and Kai, 2013), and B. furici still known only from New Caledonia in 360–400 m (Fourmanoir and Rivaton, 1979; Poss, 1999; Fricke et al., 2011). In 1999, a single specimen identified as Bembradium was caught in the Andaman Sea off Phuket, Thailand, during the Biodiversity of the Andaman Sea Shelf (BIOSHELF) project operated by the Phuket Marine Biological Center of Thailand. As a result of a comparison of the specimen with types and other specimens of the two congeners, the specimen was found to be a new species. The study provides a formal description of the new species. MATERIALS AND METHODS Specimens examined in this study are in the Kyoto University, Japan (FAKU), Muséum national d’Histoire naturelle, France (MNHN), Phuket Marine Biological Center, Thailand (PMBC), and National Museum of Natural History, Smithsonian Institution, USA (USNM). Counts and proportional measurements gen- erally follow Hubbs and Lagler (1958), and scale counts follow Nakabo (2002). Snout width was measured at the anterior margin of the orbit, and at the anterior margin of the anterior nostril. Standard and head lengths are abbreviated as SL and HL, respectively. Measurements were made to the nearest 0.1 mm with a digital caliper and a divider. Terminology of head spines generally follows Knapp et al. (2000) except for the nasal ridge. Nasal ridge is on the nasal bone.

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Page 1: Phuket mar. biol. Cent. Res. Bull.76: 9–17 (2019)

Phuket mar. biol. Cent. Res. Bull.76: 9–17 (2019)

DESCRIPTION OF A NEW SPECIES OF Bembradium (SCORPAENIFORMES: BEMBRIDAE) FROM THE ANDAMAN SEA, THAILAND

Saki Kishimoto1*, Toshio Kawai2, Fumihito Tashiro3 and Charatsee Aungtonya4

1Graduate School of Fisheries Sciences, Hokkaido University, Hakodate,Hokkaido 041-8611, Japan

2Faculty of Fisheries Sciences, Hokkaido University Museum, Hakodate, Hokkaido 041-8611, Japan

3Fisheries Science Center, Hokkaido University Museum, Hakodate, Hokkaido 041-8611, Japan

4Phuket Marine Biological Center, Muang District, Phuket 83000, Thailand*Corresponding author: [email protected]

ABSTRACT: A new species of Bembradium (Scorpaeniformes: Bembridae) is described on the basis of a single specimen collected in 284–300 m in the Andaman Sea off Phuket, Thailand. The new species, Bembradium magnoculum, differs from all congeners by the following combination of characters: short and wide snout (length 79.9% eye diameter; width at anterior margin of anterior nostril 13.4% SL), large eye (13.2% SL), long upper jaw (20.1% SL), long preanus length (64.8% SL), long predorsal-fin length (41.9% SL), short second dorsal-fin base (57.1% HL), long anal-fin base (23.9% SL), short lower caudal peduncle length (25.8% HL), short pectoral fin (45.9% HL), short pelvic-fin spine (16.0% HL), small spines of similar size on nasal bone and broad dermal flap of anterior nostril.

Keywords: Pisces, Bembradium magnoculum sp. nov., Indian Ocean, BIOSHELF

INTRODUCTION

The genus Bembradium Gilbert, 1905, placed in the family Bembridae, was established for Bembradium roseum Gilbert, 1905, based on a specimen captured off Hawaii. A second species, Bembradium furici Fourmanoir and Rivaton, 1979, was described from a New Caledonian specimen. Since then, this genus has included only the two species, B. roseum recorded from Hawaii, Emperor Seamount Chain, Japan, Taiwan, and New Caledonia, in depths of 138–800 m (Gilbert, 1905; Chave and Mundy, 1994; Rivaton and Bourret, 1999; Yamada et al., 2007; Ho et al., 2009; Fricke et al., 2011; Nakabo and Kai, 2013), and B. furici still known only from New Caledonia in 360–400 m (Fourmanoir and Rivaton, 1979; Poss, 1999; Fricke et al., 2011). In 1999, a single specimen identified as Bembradium was caught in the Andaman Sea off Phuket, Thailand, during the Biodiversity of the Andaman Sea Shelf (BIOSHELF) project operated by the Phuket Marine Biological Center of Thailand. As a result of a comparison of the specimen with types and other specimens of the two congeners,

the specimen was found to be a new species. The study provides a formal description of the new species.

MATERIALS AND METHODS

Specimens examined in this study are in the Kyoto University, Japan (FAKU), Muséum national d’Histoire naturelle, France (MNHN), Phuket Marine Biological Center, Thailand (PMBC), and National Museum of Natural History, Smithsonian Institution, USA (USNM). Counts and proportional measurements gen-erally follow Hubbs and Lagler (1958), and scale counts follow Nakabo (2002). Snout width was measured at the anterior margin of the orbit, and at the anterior margin of the anterior nostril. Standard and head lengths are abbreviated as SL and HL, respectively. Measurements were made to the nearest 0.1 mm with a digital caliper and a divider. Terminology of head spines generally follows Knapp et al. (2000) except for the nasal ridge. Nasal ridge is on the nasal bone.

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TAXONOMYFamily Bembridae

Genus Bembradium Gilbert, 1905

Bembradium Gilbert, 1905: 637 (type species Bembradium roseum Gilbert, 1905); neuter.

Generic diagnosis. Anterior part of head strongly depressed, becoming less so posteriorly; lower jaw short and not protruding; many small spines on suborbital ridge; origin of pectoral fin posterior to vertical through pelvic fin origin; lateral line running near middle of body; body scales large (Gilbert, 1905; Nelson et al., 2016).

Bembradium magnoculum sp. nov.Figs. 1A, 1B, 2, 3A; Tables 1, 2

Holotype. PMBC 30062, 115.9 mm SL, 7°30'N 97°50'E to 7°31'N 97°51'E, 284–300 m depth, 9 Nov. 1999, Agassiz trawl, R/V Chakratong Tongyai, St. I6.

Diagnosis. Snout length 79.9% eye diameter; snout width at anterior margin of anterior nostril 13.4% SL; eye diameter 13.2% SL; upper-jaw length 20.1% SL; preanus length 64.8% SL; predorsal-fin length 41.9% SL; length of second dorsal-fin base 57.1% HL; length of anal-fin base 23.9% SL; length of lower caudal peduncle 25.8% HL; pectoral-fin length 45.9% HL; pelvic-fin spine length 16.0% HL; size of spines on nasal bone small and of similar size; dermal flap of anterior nostril broad.

Description. Counts and measurements are provided in Tables 1 and 2. Body elongate and gradually compressed posteriorly, its profile slightly arched dorsally and nearly straight ventrally, deepest at the first spine of first dorsal fin. Head large, depressed, especially anteriorly. Snout short and wide, strongly depressed, with broad dermal flap on anterior nostril (Fig. 3A). Mouth subterminal; maxilla extending beyond anterior margin of eye, but not reaching past center of eye. Orbit and eye very large; interorbital region very narrow and grooved.

Opercle with two smooth ridges (Fig. 2). Operclular flap well developed. Gill rakers on upper limb three plus 12 (left) or 11 (right) on lower limb, including single raker at angle. Lateral line running nearly parallel to dorsal contour of body. First dorsal fin separated from second fin. First dorsal fin originating above pectoral-fin base. Anal fin origi-nating slightly posterior to vertical through origin of second dorsal fin. Pectoral fin rounded; its tip reaching to vertical through second anal ray base. Posterior tip of depressed pelvic fin reaching vertical through base of seventh dorsal fin spine. Posterior margin of caudal fin slightly rounded. Some small spines of similar size on nasal bone. Lachrymal with (left side) or without (right side) single small spine. Two ridges with small spines on suborbital region; length of spines on anterior ridge [12 (left side) or 11 (right side)] much shorter than on posterior ridge [18 (left side) or 16 (right side)]. Single preocular spine in front of eye; tip of spine forked (left side) or simple (right side). Interorbital region with 17 (left side) or 18 (right side) supraorbital spines. Fourth and fifth infraor-bital bones serrated. Postocular region with 18 (left side) or 14 (right side) small spines. Pterotic with ridge possessing three spines. Parietal with single prominent spine. Two nuchal spines. Single robust spine on preopercle with small supplemental spine just before robust spine; robust spine not reaching to posterior margin of opercle (Fig. 2). Teeth on jaws, vomer, palatine and first basi-branchial conical, uniformly very small; upper-jaw teeth forming separate tooth-bands on each side. Scales ctenoid on body, cheek, occiput, isthmus, jugular, pectoral-fin base and basal 1/6 of pectoral fin; scales on body largest, those on pectoral fin smallest. Coloration when fresh unknown. When pre-served in alcohol, head, body and all fins uniformly yellowish brown.

Distribution. Known from only the sea floor at 284–300 m depth off Phuket, Thailand in the southern Andaman Sea.

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Description of a new species of Bembradium

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Etymology. The specific name magnoculum from the Latin magnus, large, and oculus, eye, refers to the large eyes of this species.

Remarks. In this study, a new species is added to genus Bembradium, bringing the total number of species in this genus to three. They can be distinguished from each other by differences in snout length (79.9% eye diameter in B. magnoculum vs. 87.8–98.6 in B. furici and 86.4–115.7 in B. roseum), snout width at anterior margin of anterior nostril (13.4% SL in B. magnoculum vs. 10.5–12.9 in B. furici and 9.9–11.6 in B. roseum), eye diameter (13.2% SL in B. magnoculum and 12.4–13.4 in B. furici vs. 9.8–11.8 in B. roseum), upper-jaw length (20.1% SL in B. magnoculum vs. 17.9–19.5 in B. furici and 17.7–19.4 in B. roseum), preanus length (64.8% SL in B. magnoculum vs. 58.0–64.3 in B. roseum), predorsal-fin length (41.9% SL in B. magnoculum vs. 38.5–40.5 in B. furici and 35.7–41.0 in B. roseum), length of second dorsal-fin base (57.1% HL in B. magnoculum and 48.9–58.0 in B. furici vs. 58.8–73.6 in B. roseum), length of anal-fin base (23.9% SL in B. magnoculum and 23.3–27.6 in B. roseum vs. 21.7–22.0 in B. furici), length of lower caudal peduncle (25.8% HL in B. magnoculum vs. 30.9–35.0 in B. furici vs. 27.4–38.6 in B. roseum), pectoral-fin length (45.9% HL in B. magnoculum vs. 47.9–53.5 in B. furici and 47.6–60.5 in B. roseum), pelvic-fin spine length (16.0% HL in B. magnoculum vs. 19.7–21.5 in B. furici and 16.6–22.9 in B. roseum), size of spines on nasal bone (small and of similar size in B. magnoculum and B. roseum vs. posteriormost spine much stronger than others in B. furici) and shape of dermal flap of anterior nostril (broad in B. magnoculum and B. furici vs. narrow in B. roseum; the flap broken in holotype of B. roseum) (Fig. 3; Tables 1, 2).

Key to the species of Bembradium1a. Dermal flap of anterior nostril narrow; eye small (9.8–11.8% SL)..…......................... B. roseum1b. Dermal flap of anterior nostril broad; eye large (12.4–13.4% SL)………….......................... 22a. Posteriormost spine on nasal bone much stronger than others; snout long (87.8–98.6% eye diameter) …...………………........................… B. furici2b. Spines on nasal bone small and of similar size; snout short (79.9% eye diameter)….…..…… ……….......…......… B. magnoculum sp. nov.

Comparative materialsBembradium furici: MNHN 1978-0470, holotype of B. furici, 120.2 mm SL, west of the Pines Island, New Caledonia, 360 m depth, trawl; MNHN 2004-2577, 3 specimens, 104.2–117.1 mm SL, 380–389 m depth, 23 Oct. 2003; MNHN 2004-2691, 1 specimen, 99.6 mm SL, 395–400 m depth, 5 Nov. 2003.

Bembradium roseum: USNM 51617, holotype of B. roseum, 71.7 mm SL, Pailolo Channel, Hawaii, 252 m depth; FAKU 1818, 1820, 1821, 3 specimens, 85.3–97.8 mm SL, off Owase, Mie, Japan, 13 May 1936; FAKU 1890, 1 specimen, 70.7 mm SL, off Owase, Mie, Japan, 27 May 1936; FAKU 3103, 1 specimen, 75.3 mm SL, Miya Fish Market, Aichi, Japan, 7 Nov. 1936; FAKU 72512–72514, 3 specimens, 97.4–116.0 mm SL, Emperor Seamount Chain, 23 Nov. 1994; FAKU 72541, 72549, 2 specimens, 104.3–104.9 mm SL, Emperor Seamount Chain, 1994; FAKU 72577, 1 specimen, 110.4 mm SL, Emperor Seamount Chain, 14 Nov. 1994.

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Figure 1. Three species of Bembradium. (A) B. magnoculum sp. nov., PMBC 30062, holotype, 115.9 mm SL; (B) dorsal view of head of B. magnoculum sp. nov., PMBC 30062, holotype; (C) B. furici, MNHN 1978-0470, holotype, 120.2 mm SL; (D) B. roseum, USNM 51617, holotype, 71.7 mm SL. Scale bars 10 mm.

Figure 2. Lateral view of head of Bembradium magnoculum sp. nov., PMBC 30062, holotype, 115.9 mm SL. Spines: LAC, lachrymal; NC, nuchal; NR, nasal ridge; PA, parietal; PO, preopercular; POC, preocular; PSOC, postocular; PT, pterotic; SBO, suborbital; SPO, supraorbital; SS, supplemental preopercular. Scale bar 10 mm.

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Description of a new species of Bembradium

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Figure 3. Dermal flap on anterior nostril: (A) B. magnoculum sp. nov., PMBC 30062, holotype, 115.9 mm SL; (B) B. roseum, FAKU 72541, 104.3 mm SL. Scale bars 1 mm.

ACKNOWLEDGEMENTS

We sincerely thank Yoshiaki Kai (FAKU) for loaning specimens and providing valuable com-ments; William J. Richards (National Marine Fisheries Service, Southeast Fisheries Science Center / NOAA) for his critical reading of an early manuscript and English correction; Somkiat Khokiattiwong (former PMBC Director), Somchai Bussarawit (former Specialist, Marine and Coastal Resources and

Development Institute, Thailand), and staff of the Reference Collection of PMBC for their kind help during research in PMBC; J. Williams, J. Clayton, S. Raredon and K. Murphy (USNM) and P. Pruvost, G. Zora, L. M. Duque-Vélez and A. Laurent (MNHN) for their organization and assistance during FT’s visit. We also thank Hisashi Imamura (Hokkaido University, Japan) and Naohide Nakayama (formerly FAKU) who supported our examination.

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Phuket mar. biol. Cent. Res. Bull.

14

Tabl

e 1.

Mer

istic

cou

nts o

f thr

ee sp

ecie

s of B

embr

adiu

m.

1 *D

ata

for B

. fur

ici p

artly

from

Pos

s (19

99).

2 *D

ata

for B

. ros

eum

par

tly fr

om F

orm

anoi

r and

Riv

aton

(197

9), K

anay

ama

(198

2), Y

atou

(198

5), O

chia

i (19

88),

Imam

ura

(199

7), Y

amad

a et

al.

(200

7), H

o et

al.

(200

9), a

nd N

akab

o an

d K

ai (2

013)

.

B. m

agno

culu

m sp

. nov

.B.

furi

ciB.

rose

umho

loty

peho

loty

peot

hers

1 *ho

loty

peot

hers

2 *St

anda

rd le

ngth

(mm

)11

5.9

120.

299

.6–1

17.1

71.7

70.7

–116

.0C

ount

sD

orsa

l fin

IX-1

2IX

-11

IX-1

1–12

IX-1

2V

III–

XI-

11–1

2A

nal fi

n11

1010

1110

–11

Cau

dal fi

n7+

6=13

No

data

No

data

No

data

6–7+

6–7=

12–1

4Pe

ctor

al fi

n25

24–2

523

–30

23–2

523

–27

Pelv

ic fi

nI,

5I,

5I,

5I,

5I,

5G

ill ra

kers

3+11

–12=

14–1

52+

8=10

2–3+

7=9–

103+

6=9

2–4+

7–12

=10–

15Ve

rtebr

ae26

No

data

No

data

2626

Late

ral-l

ine

scal

es28

2928

–29

2827

–28

Pred

orsa

l sca

les

4N

o da

taN

o da

taN

o da

ta3–

5Sc

ales

abo

ve la

tera

l lin

e2

No

data

No

data

No

data

2–

3.5

Scal

es b

elow

late

ral l

ine

5N

o da

taN

o da

taN

o da

ta5–

8

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Page 7: Phuket mar. biol. Cent. Res. Bull.76: 9–17 (2019)

Description of a new species of Bembradium

15

Tabl

e 2.

Mea

sure

men

ts re

lativ

e to

SL

and

HL

for t

hree

spec

ies o

f Bem

brad

ium

.

B. m

agno

culu

m sp

. nov

.B.

furi

ciB.

rose

umho

loty

peho

loty

peot

hers

holo

type

othe

rs*

Stan

dard

leng

th (m

m)

115.

912

0.2

99.6

–117

.171

.770

.7–1

16.0

Mea

sure

men

ts (%

SL)

Bod

y de

pth

at fi

rst d

orsa

l fin

20.8

20.5

18.3

–20.

615

.914

.7–2

1.9

Bod

y de

pth

at se

cond

dor

sal fi

n16

.518

.315

.9–1

7.5

15.5

12.1

–18.

1B

ody

wid

th3.

2N

o da

taN

o da

taN

o da

ta2.

9–4.

5H

ead

leng

th46

.843

.844

.4–4

6.8

39.6

41.4

–47.

6H

ead

dept

h18

.1N

o da

ta18

.3–2

1.5

15.6

13.5

–18.

8H

ead

wid

th20

.3N

o da

ta17

.6–1

9.5

17.9

16.2

–20.

5Sn

out l

engt

h10

.513

.411

.7–1

2.3

11.3

10.0

–11.

4Sn

out w

idth

at a

nter

ior m

argi

n of

orb

it 15

.4N

o da

ta11

.8–1

4.5

12.6

12.0

–16.

3Sn

out w

idth

at a

nter

ior m

argi

n of

ant

erio

r nos

tril

13.4

No

data

10.5

–12.

910

.59.

9–11

.6Ey

e di

amet

er13

.2N

o da

ta12

.4–1

3.4

9.8

10.0

–11.

8O

rbita

l dia

met

er16

.613

.114

.4–1

5.2

12.6

14.9

–16.

2Po

st o

rbita

l len

gth

19.4

No

data

18.3

–19.

215

.916

.6–1

9.4

Inte

rorb

ital w

idth

2.0

No

data

1.5–

1.8

2.4

1.6–

1.9

Upp

er-ja

w le

ngth

20.1

19.3

17.9

–19.

517

.717

.9–1

9.4

Prea

nus l

engt

h64

.8N

o da

taN

o da

ta58

.058

.7–6

4.3

Pred

orsa

l-fin

leng

th41

.9N

o da

ta38

.5–4

0.5

35.7

37.2

–41.

0Pr

eana

l-fin

leng

th67

.7N

o da

ta63

.1–6

6.9

61.2

62.7

–68.

1Le

ngth

of fi

rst d

orsa

l-fin

base

19.7

No

data

18.1

–21.

120

.415

.3–2

3.0

Leng

th o

f sec

ond

dors

al-fi

n ba

se26

.7N

o da

ta22

.9–2

6.1

29.1

25.3

–29.

1Le

ngth

of a

nal-fi

n ba

se23

.9N

o da

ta21

.7–2

2.0

27.6

23.3

–27.

2

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Page 8: Phuket mar. biol. Cent. Res. Bull.76: 9–17 (2019)

Phuket mar. biol. Cent. Res. Bull.

16

B. m

agno

culu

m sp

. nov

.B.

furi

ciB.

rose

umho

loty

peho

loty

peot

hers

holo

type

othe

rs*

Cau

dal p

edun

cle

dept

h7.

57.

67.

2–7.

57.

55.

8–7.

4Le

ngth

of u

pper

cau

dal p

edun

cle

13.8

17.8

15.0

–17.

516

.312

.8–1

7.7

Leng

th o

f low

er c

auda

l ped

uncl

e12

.114

.014

.0–1

5.8

14.1

11.9

–16.

0Pe

ctor

al-fi

n le

ngth

21.5

23.5

21.6

–22.

8N

o da

ta19

.2–2

5.8

Pelv

ic-fi

n le

ngth

16.7

18.1

18.7

–19.

4N

o da

ta14

.3–1

8.0

Pelv

ic-fi

n sp

ine

leng

th7.

5N

o da

ta8.

9–10

.09.

17.

1–9.

7M

easu

rem

ents

(% H

L)Le

ngth

of s

econ

d do

rsal

-fin

base

57.1

No

data

48.9

–58.

073

.658

.8–6

9.5

Leng

th o

f low

er c

auda

l ped

uncl

e25

.831

.930

.9–3

5.0

35.6

27.4

–38.

6Pe

ctor

al-fi

n le

ngth

45.9

53.5

47.9

–51.

4N

o da

ta47

.6–6

0.5

Pelv

ic-fi

n sp

ine

leng

th16

.0N

o da

ta19

.7–2

1.5

22.9

16.6

–21.

9

Tabl

e 2.

(con

't)

*Dat

a fo

r B. r

oseu

m p

artly

from

Kan

ayam

a (1

982)

, Yat

ou (1

985)

and

Ho

et a

l. (2

009)

.

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Description of a new species of Bembradium

17

REFERENCES

Chave, E.H. and B.C. Mundy. 1994. Deep-sea benthic fish of the Hawaiian Archipelago, Cross Seamount, and Johnston Atoll. Pac. Sci. 48: 367–409.Fourmanoir, P. and J. Rivaton. 1979. Poissons de la pente récifale externe de Nouvelle-Calédonie et des Nouvelles-Hébrides. Cah. Indo Pacif. 1: 405–443.Fricke, R., M. Kulbicki and L. Wantiez. 2011. Checklist of the fishes of New Caledonia, and their distribution in the southwest Pacific Ocean (Pisces). Stuttg. Beitr. Naturk. A Neue Ser. 4: 341–463.Gilbert, C.H. 1905. Section II. The deep-sea fishes of the Hawaiian Islands. In: Jordan, D.S. and B.W. Evermann (eds.). The aquatic resources of the Hawaiian Islands. Part II. Bulletin of the United States Fish Commission. Government Printing Office, Washington. pp. 637–638, 2 pls. Ho, H.C., K.T. Shao and C.W. Chang. 2009. Three new records of the scorpaeniform fishes in Taiwan. Platax 6: 27–34.Hubbs, C.L. and K.F. Lagler. 1958. Fishes of the Great Lakes region. Bull. Cranbrook Inst. Sci. 26: i–xv, 1–213, 1–44 pls.Imamura, H. 1997. Platycephalidae, Parabembridae, Bembridae, Plectrogeniidae and Hoplichthyidae. In: Okamura, O. and K. Amaoka (eds.). Sea fishes of Japan. Yama-Kei Publishers Company Limited, Tokyo. pp. 783. (in Japanese)Kanayama, T. 1982. Bembradium roseum Gilbert. In: Okamura, O., K. Amaoka and F. Mitani (eds.). Fishes of the Kyushu-Palau Ridge and Tosa Bay. Japan Fisheries Resource Conservation Association, Tokyo. pp. 401.Knapp, L.W., H. Imamura and M. Sakashita. 2000. Onigocia bimaculata, a new species of flathead fish (Scorpaeniformes: Platycephalidae) from the Indo-Pacific. Special Publ. J. L. B. Smith Inst. Ichthyol. 64: 1–10.Nakabo, T. 2002. Introduction to Ichthyology. In: Nakabo, T. (ed.). Fishes of Japan with pictorial keys to the species, English edition. Tokai University Press, Tokyo. pp. xxi–xlii.Nakabo, T. and Y. Kai. 2013. Plectrogenidae. In: Nakabo, T. (ed.). Fishes of Japan with pictorial keys to the species. Third edition. Tokai University Press, Hadano. pp. 1950. (in Japanese)Nelson, J.S., T.C. Grande and M.V.H. Wilson. 2016. Fishes of the world. Fifth edition. John Wiley & Sons, New York. xxvii + 707 pp.Ochiai, A. 1988. Platycephalidae. In: Masuda, H., K. Amaoka, C. Araga, T. Uyeno and T. Yoshino (eds.). The fishes of the Japanese Archipelago. Tokai University Press, Tokyo. pp. 307–308. (in Japanese)Poss, S.G. 1999. Bembridae. In: Carpenter, K.E. and V.H. Niem (eds.). The living marine resources of the western central Pacific. Volume 4. Bony fishes part 2 (Mugilidae to Carangidae). FAO Species Identification Guide for Fishery Purposes. FAO, Rome. pp. 2383–2384.Rivaton, J. and P. Bourret. 1999. Les otolithes des poissons de l’Indo-Pacifique. Doc. Sci. Tech. II(2): 1–378.Yamada, U., M. Tokimura, H. Horikawa and T. Nakabo. 2007. Fishes and fisheries of the East China and Yellow Seas. Tokai University Press, Hadano. lxxii + 1262 pp. (in Japanese)Yatou, T. 1985. Bembradium roseum Gilbert. In: Okamura, O. and T. Kitajima (eds.). Fishes of the Okinawa Trough and the adjacent waters II. Japan Fisheries Resource Conservation Association, Tokyo. pp. 728.

Manuscript received: 26 June 2018Accepted: 11 October 2018

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