Phuket mar. biol. Cent. Res. Bull. 80: 21–51 (2023)
DOI: 10.14456/pmbcrb.2023.2

DEEP-SEA FISHES FROM THE ANDAMAN SEA BY R/V CHAKRATONG TONGYAIDURING 1996–2000. PART 7: ORDER STOMIIFORMES
Kota Obata1*, Kohei Mita1, Tetsuro Senda1, Toshio Kawai1, Fumihito Tashiro2,
Hisashi Imamura1, 2, Charatsee Aungtonya3 and Surapong Banchongmanee3
Faculty, Graduate School / School of Fisheries Sciences, Hokkaido University, Hakodate,
Hokkaido 041-8611, Japan
2
Fisheries Science Center, The Hokkaido University Museum, Hakodate, Hokkaido 041-8611, Japan
3
Phuket Marine Biological Center, Mueang Phuket District, Phuket 83000, Thailand*Corresponding author: obata.kota.z1@elms.hokudai.ac.jp
1

ABSTRACT: Stomiiform fishes collected by the project team of the Biodiversity of the Andaman Sea
Shelf (BIOSHELF) during 1996–2000 were identified as 28 species belonging to 5 families in 2 suborders.
This study represents the first records of the following 11 species from the Andaman Sea: i.e., 1 speciesof Diplophidae: Manducus greyae (Johnson, 1970); 1 species of Phosichthyidae: Pollichthys mauli (Poll,
1953); 9 species of Stomiidae: Astronesthes formosana Liao, Chen and Shao, 2006, Astronesthes indopacifica
Parin and Borodulina, 1997, Borostomias mononema (Regan and Trewavas, 1929), Bathophilus pawneei
Parr, 1927, Eustomias (Haploclonus) bifilis Gibbs, 1960, Leptostomias multifilis Imai, 1941, Melanostomias
melanops Brauer, 1902, Photonectes (Photonectes) albipennis (Döderlein, 1882) and Idiacanthus fasciolaPeters, 1877.
Key words: Diplophidae, Gonostomatidae, Sternoptychidae, Phosichthyidae, Stomiidae, Thailand, BIOSHELF

INTRODUCTION
Deep-sea fishes collected by the project teamof the Biodiversity of the Andaman Sea Shelf(BIOSHELF) (see details in Aungtonya et al. 2000)
have been studied taxonomically as following
orders (sensu Nelson 2006): Scorpaeniformes by
Kawai et al. (2017) and Kishimoto et al. (2019),
Beryciformes and Stephanoberyciformes by Kimura
et al. (2019a; b), Albuliformes, Ateleopodiformes andLampriformes by Kawai et al. (2020a), Argentiniformes
by Senda et al. (2020), Perciformes by Kawai et al.
(2020b) and Pleuronectiformes and Tetraodontiformesby Kawai et al. (2020c). Following the aforementioned
studies, species of Stomiiformes (sensu Nelson2006) are newly reported in this study.

MATERIALS AND METHODSAll specimens, which were caught from deep-sea
waters of the Andaman Sea by BIOSHELF(Aungtonya et al. 2000), have been kept in 70%
ethyl alcohol after fixation by 10% formalin. These
specimens have been deposited at the ReferenceCollection of Phuket Marine Biological Center,
Phuket, Thailand (PMBC). Several specimens weretransferred to the Hokkaido University Museum,
Hakodate, Japan (HUMZ). Comparative materials
are deposited at PMBC and HUMZ.
Counts and measurements follow Hubbs andLagler (1958) for Diplophidae, Gonostomatidae
and Phosichthyidae, Baird (1971) for Argyropelecus(Sternoptychidae), Harold (1994) for Polyipnus(Sternoptychidae), Haruta and Kawaguchi (1976)
for Sternoptyx (Sternoptychidae) and Gibbs et al.
(1983) for Stomiidae. Photophore terminology
follows Grey (1964) and Johnson (1970) for
Diplophidae, Gonostomatidae and Phosichthyidae,
Baird (1971) for Argyropelecus (Sternoptychidae),
Harold (1994) for Polyipnus (Sternoptychidae),Haruta and Kawaguchi (1976) for Sternoptyx(Sternoptychidae) and Morrow (1964a) andWeitzman (1986) for Stomiidae. Barbel
terminology of Stomiidae follows Morrow andGibbs (1964) and Gibbs et al. (1983). Standard
and head lengths are abbreviated as SL and HL,
respectively. Measurements were made to the nearest
0.1 mm with digital caliper. Vertebrae were counted
from radiographs. Specimens without detailed
examination, meaning only species identificationand measurement of SL, are tagged with asterisk after
catalog number.

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Phuket mar. biol. Cent. Res. Bull.

Differences of some characters of each species
between the present and previous studies are treatedas intraspecific variations. This is because thesedifferences are too small for interspecific differences
and other characters are congruent with diagnosesof each species.
SPECIES LIST
Suborder Gonostomatoidei
Family Diplophidae
Diplophos cf. taenia
Fig. 1
Diagnosis. Anal-fin rays 65; IC photophores 104;
LLP photophores 104; dorsal-fin base situated at
middle of body; two photopuncts present ahead of
SO photophore; last two AC photophores grouped,
not separated (present study).
Materials. PMBC 30562, 1 specimen, 72.4 mm SL,
St. J8, 7°15′N 97°33′E to 7°15′N 97°30′E, otter trawl,
473–494 m depth, 18 Feb. 2000; PMBC 30563*, 1
specimen, SL unmeasured due to damage, St. E7,8°30′N 97°01′E to 8°29′N 97°03′E, Agassiz trawl,
449–446 m depth, 8 Feb. 2000.
Distribution. Andaman Sea (present study).
Remarks. These specimens are close to Diplophos
australis Ozawa, Oda and Ida, 1990 and Diplophos
taenia Günther, 1873 in having 65 anal-fin rays
(61–67 in the former species vs. 59–72 in the latter:
Grey 1964; Mukhacheva 1978; Ozawa et al. 1990;
Koeda and Ho 2019). Although these two species
can be almost distinguished by the number ofvertebrae (33–37 + 50–54 = 84–91 vs. 37–41 +52–60 = 90–100: Ozawa et al. 1990), those of
the present specimens could not be examined
due to their poor condition. Diplophos taenia isreported from tropical to temperate waters in world
oceans including the Andaman Sea, while D.
australis only from South Atlantic and South Pacificsouth from 40°S (e.g., Ozawa et al. 1990; Rajan etal. 2013; Kenaley and Stewart 2015a; Villarins etal. 2022). Therefore, this study tentatively identified
these specimens as Diplophos cf. taenia.

Manducus greyae (Johnson, 1970)
Fig. 2
Diagnosis. Anal-fin rays 55–59; total vertebrae
76; IC photophores 83–89; dorsal-fin origin nearer
to snout than caudal peduncle; 4–5 photopunctspresent ahead of SO photophore, arranged inirregular posteriorly directed series; last two AC
photophores not grouped, distance between last two AC
photophores equal to or wider than that of proceeding AC
photophores (Johnson 1970; Mukhacheva 1978).
Material. PMBC 30564, 1 specimen, 104.8 mm
SL, St. E7, 8°30′N 97°07′E to 8°29′N 97°40′E, otter
trawl, 435–444 m depth, 9 Feb. 2000.
Distribution. Indian Ocean off Sumatra and Javaislands, and western Pacific from South China Sea
to Papua New Guinea (Johnson 1970; Mukhacheva
1978; Ozawa and Oda 1986; Kailola 1987; Harold
1999; Gloerfelt-Tarp and Kailola 2022) including
Andaman Sea (present study).
Remarks. In the Indian Ocean, this species has
been known only from off Sumatra and Java islands(Mukhacheva 1978; Gloerfelt-Tarp and Kailola2022). Therefore, this study is the first record of
the species from the Andaman Sea.
Triplophos hemingi (McArdle, 1901)
Fig. 3
Diagnosis. Dorsal-fin rays 9–11; anal-fin rays
54–63; gill rakers on first gill arch 8–9 + 11–16 =23–25; branchiostegal rays 11–17; total vertebrae
58–62; BR photophores 8–14; IV photophores 24–30;
VAV photophores 5–7; IC photophores 68–76;
premaxilla well elongated more than toothed
portion of maxilla and forming most of lower
margin of upper jaw; mouth and opercular bones
oblique; dorsal-fin base situated well ahead of
middle of body; anal-fin base longer than half of
SL; SO photophore present; small photophores
arranged in a line along upper jaw; photophorespresent on isthmus; four or five lines of photophorespresent on lateral body (McArdle 1901; Brauer 1902;
Poll 1953; Grey 1964; present study).

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Deep-sea fishes from the Andaman Sea

Materials. PMBC 30502*, 7 specimens, 136.2–
180.3 mm SL, St. Z4, 7°34'N 97°03'E to 7°35'N
97°04'E, 660–633 m depth, otter trawl, 25 Jan.
1999; PMBC 30565, 1 specimen, 138.9 mm SL,
St. G8, 8°00'N 97°06'E to 8°00'N 97°04'E, 508–
518 m depth, otter trawl, 20 Nov. 1999; PMBC
30566*, 1 specimen, 94.2 mm SL, PMBC 30567,
1 specimen, 72.9 mm SL, St. J8, 7°15'N 97°33'E to
7°15'N 97°30'E, 473–494 m depth, otter trawl, 18
Feb. 2000; PMBC 30568, 6 specimens, 81.7–114.8mm SL, St. L8, 6°46'N 97°33'E to 6°44'N 97°35'E,

513–501 m depth, otter trawl, 22 Feb. 2000; PMBC
30569, 4 specimens, 152.5–171.0 mm SL, St. J10,
7°20'N 97°14'E to 7°22'N 97°13'E, 655–651 m
depth, otter trawl, 28 Jan. 1999; PMBC 30570,
6 specimens, 70.6–96.6 mm SL, PMBC 30571,
2 specimens, 73.7–92.7 mm SL, PMBC 30572,
12 specimens, 62.3–94.9 mm SL, St. Z2, 7°42'N
97°28'E to 7°42'N 97°18'E, 464 m depth, otter
trawl, 23 Jan. 1999; PMBC 30573*, 9 specimens,
94.4–161.7 mm SL, St. Z3, 7°42'N 97°20'E to
7°42'N 97°18'E, 493–322 m depth, otter trawl,
24 Jan. 1999.

Figure 1. Diplophos cf. taenia, PMBC 30562, 72.4 mm SL. Scale bar 10 mm.

Figure 2. Manducus greyae, PMBC 30564, 104.8 mm SL. Scale bar 10 mm.

Figure 3. Triplophos hemingi, PMBC 30565, 138.9 mm SL. Scale bar 20 mm.

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Phuket mar. biol. Cent. Res. Bull.

Distribution. Tropical to subtropical waters in
Atlantic, Indian and western Pacific Oceans (e.g.,
Grey 1964; Schaefer et al. 1986a; Quéro et al.
1990a; Harold 1999; 2002; Randall and Lim 2000;
Psomadakis et al. 2019; Koeda 2020a; Sutton et al.
2020; Villarins et al. 2022).
Remarks. The characters of present specimens
are different from those of previous descriptions in
the followings: 9–11 dorsal-fin rays in the present
specimens (vs. 10–11 in previous descriptions),
20–23 total gill rakers on the first arch (vs. 23–25),
14–17 branchiostegal rays (11–14) and 11–14 BR
photophores (8–11) (McArdle 1901; Brauer 1902;
Poll 1953; Grey 1964). However, other characters ofthem are congruent with diagnosis of T. hemingi:
e.g., premaxilla well elongated more than toothed
portion of maxilla; dorsal-fin base situated wellahead of middle of body; 4–5 lines of photophorespresent on lateral body (McArdle 1901; Brauer
1902; Poll 1953; Grey 1964). Therefore, we tentatively
identified the present specimens as T. hemingi.
Grey (1964) indicated intraspecific variationsin this species between the Indian and Atlantic
Oceans: IV photophores 29–30 and VAV photophores
5–6 in Indian Ocean vs. 24–25 and 7 in Atlantic.
However, the present study rejects Grey’s (1964)
hypothesis because of findings of specimens having 24
IV photophores (one individual of PMBC 30568)
and 7 VAV photophores (one individual of PMBC
30572). On the other hand, present study regards
the morphological differences of the following
characters as the geographical variations between
both oceans: i.e., total gill rakers on the first gillarch 20–23, branchiostegal rays 14–17 and BR
photophores 11–14 in the Indian Ocean vs. 23–25,
11–14 and 8–11 in the Atlantic (McArdle 1901;
Brauer 1902; Grey 1964; present study).
Family Gonostomatidae
Cyclothone acclinidens Garman, 1899
Fig. 4
Diagnosis. Maxillary teeth on posterior series
oblique at acute angle, enlarged posteriorly, notalternated with shorter teeth; vomerine teeth present;
gill rakers between upper and lower limbs of firstgill arch 2; gill rakers on hypobranchial of first gillarch 5–6; gill filaments on hypobranchial not fused;anus situated at just middle between pelvic-fin baseand anal-fin origin, or slightly nearer to pelvic-fin base;supracaudal luminous gland well developed,
reaching dorsal-fin base; area just anterior toanal-fin origin transparent or sparsely pigmented(Mukhacheva 1964; 1974; 1980; Kawaguchi 1971;
Badcock 1982; Miya 1994a).
Materials. PMBC 30574, 2 specimens, 24.1–25.6mm SL, St. B10, 9°11′N 96°12′E to 9°10′N 96°14′E,
689–549 m depth, otter trawl, 11 Feb. 1997.
Distribution. Tropical to temperate waters ofworld oceans (e.g., Mukhacheva 1964; 1974; 1980;
Kawaguchi 1971; Badcock 1984a; Quéro et al.
1990a; Harold 2002; Paxton et al. 2006a; Sutton
et al. 2020).
Remarks. Some of the diagnostic characters of
C. acclinidens could not be checked in the present
specimen because of its damage: e.g., extension of
supracaudal luminous grand to dorsal-fin base, andtransparent or sparsely pigmented region anterior
to anal-fin origin. However, the present study
identified the specimen as C. acclinidens basedon the morphology of teeth on upper jaw, thenumber of gill rakers, the morphology of gill
filament, and the situation of anus (see diagnosis).
Cyclothone alba Brauer, 1906
Fig. 5
Diagnosis. BR photophores 8; OA photophores 6;
vomerine teeth absent; gill raker between upperand lower limbs of first gill arch 1; gill filamentson hypobranchial fused; series of OA photophores
not reaching behind pelvic-fin base; photophores
on anal-fin base 9; body mostly transparent except
dorsal head with V-shaped pigmented area
(Mukhacheva 1964; 1974; Bond and Tighe 1974;
Miya 1994a).
Materials. PMBC 30501*, 1 specimen, SL unmeasured
due to damage, PMBC 30575, 1 specimen, 22.5mm SL, PMBC 30576, 1 specimen, 21.8 mm SL,
St. J8, 7°15′N 97°30′E to 7°15′N 97°33′E, 473–494
m depth, otter trawl, 18 Feb. 2000.
Distribution. Tropical to temperate and subarctic
waters of Atlantic, Indian Ocean, western and central
Pacific, and eastern Pacific off Chile (e.g., Kawaguchi
1971; Mukhacheva 1974; Badcock 1984a; Quéro

25
Deep-sea fishes from the Andaman Sea
et al. 1990a; Harold 2002; Paxton et al. 2006a;
Fricke et al. 2009; Kenaley and Stewart 2015b;
Sutton et al. 2020).
Cyclothone pallida Brauer, 1902
Fig. 6
Diagnosis. Maxillary teeth on posterior series

moderately oblique, moderately enlarged posteriorly;
enlarged teeth on posterior series of maxillaryalternated with small teeth (male) or without them(female); vomerine teeth present; gill rakers between
upper and lower limbs of first gill arch 2; gill rakers
on hypobranchial of first gill arch 4; gill filamentson hypobranchial not fused; anus closer to pelvic-fin
base than anal-fin origin; supracaudal luminous

Figure 4. Cyclothone acclinidens, PMBC 30574, 24.1 mm SL, Scale bar 5 mm.

Figure 5. Cyclothone alba, PMBC 30576, 21.8 mm SL. Scale bar 5 mm.

Figure 6. Cyclothone pallida, PMBC 30083, 29.2 mm SL. Scale bar 5 mm.

26
Phuket mar. biol. Cent. Res. Bull.
gland not well developed; pigmentation on snout
region in dorsal head, and on dorsal- and analfin bases; area anterior to anal-fin origin usuallytransparent and sometimes sparsely pigmented(Kawaguchi 1971; Mukhacheva 1974; Badcock
1982; Miya 1994a; b).

Diagnosis. VAV photophores 4–5; supracaudal andinfracaudal luminous glands 1 and 2, respectively;
anus closer to anal-fin origin than middle between
pelvic-fin insertion and anal-fin origin; dorsal-fin
origin situated above anal-fin origin; pectoral-fin
tip not reaching behind pelvic-fin base; pelvic-fin
tip not reaching anus; adipose fin present; numerous
tiny photophores present on body and head; circle
luminous glands present under each OA photophore
except first OA photophore (Grey 1964; Mukhacheva1972).

PMBC 30578, 4 specimens, 134.7–164.7 mm
SL, St. B10, 9°11′N 96°12′E to 9°10′N 96°14′E,
689–549 m depth, otter trawl, 11 Feb. 1999; PMBC
30579, 1 specimen, 86.2 mm SL, PMBC 30580*, 2
specimens, 153.5–164.1 mm SL, St. E9, 8°28′N
95°59′E to 8°30′N 95°58′E, 649–550 m depth,
otter trawl, 5 Feb. 1999; PMBC 30581, 1 specimen,
115.7 mm SL, St. G8, 8°00′N 97°04′E to 8°00′N
97°06′E, 508–518 m depth, otter trawl, 20 Nov.
1999; PMBC 30582, 1 specimen, 63.7 mm SL,
station unknown, 25 Jan. 1999; PMBC 30583, 10
specimens, 60.0–94.4 mm SL, PMBC 30584*, 1
specimen, 53.2 mm SL, PMBC 30585, 8 specimens,
60.0–102.9 mm SL, PMBC 30586*, 1 specimen,
57.6 mm SL, PMBC 30587, 1 specimen,
71.0 mm SL, PMBC 30588*, 3 specimens, 56.8–
65.5 mm SL, PMBC 30589*, 4 specimens, SL
unmeasured due to damages, PMBC 30590*, 1
specimen, 60.5 mm SL, PMBC 30591*, 9
specimens, 62.0–92.3 mm SL, St. J8, 7°15′N
97°30′E to 7°15′N 97°33′E, 473–494 m depth,
otter trawl, 18 Feb. 2000; PMBC 30592, 1 specimen,
66.6 mm SL, PMBC 30593*, 2 specimens, 143.3–
152.6 mm SL, St. J10, 7°20′N 97°14′E to 7°22′N
97°13′E, 655–651 m depth, otter trawl, 28 Jan.
1999; PMBC 30594*, 1 specimen, 157.4 mm SL,
St. J10, 7°14′N 97°15′E to 7°15′N 97°15′E, 689–
687 m depth, Agassiz trawl, 19 Feb. 2000; PMBC
30595, 4 specimens, 109.1–189.5 mm SL, St. J10,7°15′N 97°14′E to 7°15′N 97°16′E, 662–696 m
depth, otter trawl, 19 Feb. 2000; PMBC 30596*, 2
specimens, 51.7–164.6 mm SL, PMBC 30597, 1
specimen, 72.9 mm SL, PMBC 30598, 5 specimens,
68.0–89.4 mm SL, PMBC 30599*, 1 specimen,
58.1 mm SL, St. L8, 6°46′N 97°33′E to 6°44′N
97°35′E, 513–501 m depth, otter trawl, 22 Feb.
2000; PMBC 30600*, 3 specimens, 55.4–67.5 mm
SL, PMBC 30601*, 6 specimens, 46.2–71.1 mm
SL, St. Z2, 7°42′N 97°28′E to 7°42′N 97°31′E, 464
m depth, otter trawl, 23 Jan. 1999; PMBC 30602*,
1 specimen, 89.1 mm SL, St. Z3, 7°42′N 97°18′E
to 7°42′N 97°20′E, 493–322 m depth, otter trawl,
24 Jan. 1999; PMBC 30603, 1 specimen, 155.0
mm SL, station and date unknown.

Materials. PMBC 30500*, 9 specimens, 147.1–
204.6 mm SL, St. Z4, 7°34'N 97°03'E to 7°35'N
97°04'E, 660–633 m depth, otter trawl, 25 Jan.
1999; PMBC 30577*, 3 specimens, 73.7–153.2mm SL, St. B8, 9°09′N 96°16′E to 9°10′N 96°18′E,
489–504 m depth, otter trawl, 11 Feb. 1999;

Distribution. Tropical to temperate waters inworld oceans (e.g., Grey 1964; Kawaguchi 1971;
Mukhacheva 1972; Badcock 1984a; Quéro et al.
1990a; Harold 1999; 2002; Paxton et al. 2006a;
Kenaley and Stewart 2015b; Psomadakis et al.
2019; Sutton et al. 2020).

Material. PMBC 30083, 1 specimen, 29.2 mm SL,
sex unknown due to damage of maxilla, St. J10,
7°20′N 97°15′E to 7°22′N 97°13′E, 655–651 m
depth, otter trawl, 28 Jan. 1999.
Distribution. Tropical to temperate and subarctic
waters of world oceans (e.g., Kawaguchi 1971;
Mukhacheva 1974; Badcock 1982; 1984a; Quéro
et al. 1990a; Miya 1994b; Harold 1999; 2002; Paxton
et al. 2006a; Kenaley and Stewart 2015b; Sutton
et al. 2020).
Remarks. Morphology of the teeth on upper jaw
and the supracaudal luminous gland could not bechecked in the present specimen due to its damage,which is one of the diagnostic characters of C.
pallida. However, the present study identified thespecimen as C. pallida based on the number of gillrakers, the morphology of gill filament, the situation ofanus, presence of pigmentation on snout and dorsaland anal-fin bases, and presence of transparent area
anterior to anal-fin origin (see diagnosis).
Sigmops elongatus (Günther, 1878)
Fig. 7

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Deep-sea fishes from the Andaman Sea
Family Sternoptychidae
Argyropelecus affinis Garman, 1899
Fig. 8Diagnosis. Dorsal-fin rays 9; dorsal profile of bodynot significantly raised posterior to dorsal blade;dorsal blade low, its height less than one-third ofits length; sphenotic spine absent on postorbital
region; postabdominal spines 2; transparent region
below abdominal photophores relatively narrowed;
supra-abdominal, preanal, anal, and subcaudal
photophores arranged regularly and straightly;
photophores of anal and subcaudal series notgrouped, separated from each other by small gaps
(Schultz 1961; 1964; Baird 1971; Borodulina 1978).
Materials. PMBC 17705, 1 specimen, 45.0 mm
SL, St. Z4, 7°34'N 97°03'E to 7°35'N 97°04'E,
660–633 m depth, otter trawl, 25 Jan. 1999; PMBC
17706, 1 specimen, 47.5 mm SL, St. B8, 9°10'N
96°18'E to 9°09'N 96°16'E, 489–504 m depth, otter

trawl, 11 Feb. 1999; PMBC 17707, 1 specimen,
38.2 mm SL, St. J10, 7°15'N 97°16'E to 7°15'N
97°14'E, 662–696 m depth, otter trawl, 19 Feb.
2000; PMBC 30604*, 1 specimen, 39.0 mm SL,
St. J8, 7°15'N 97°33'E to 7°15'N 97°30'E, 473–494
m depth, otter trawl, 18 Feb. 2000.
Distribution. Tropical to subtropical waters in world
oceans (e.g., Schultz 1961; 1964; Baird 1971;
1986; Borodulina 1978; Badcock 1984b; Quéro etal. 1990b; Aizawa and Doiuchi 2013a; Satapoomin
2011; Sutton et al. 2020; Villarins et al. 2022).
Argyropelecus sladeni Regan, 1908
Fig. 9Diagnosis. Dorsal-fin rays 9; total gill rakers on
first gill arch 17–22; enlarged canine teeth onlower jaw absent; upper preopercular spine curved
upward extending well beyond posterior borderof preopercle; postabdominal spines 2, anterior

Figure 7. Sigmops elongatus. (A): PMBC 30579, 86.2 mm SL; (B): PMBC 30595, 189.5 mm SL. Scalebars 20 mm.

28
Phuket mar. biol. Cent. Res. Bull.
one squared and blunt; no spines on scales below
subcaudal photophores; supra-abdominal, preanal,
anal, and subcaudal photophores arranged irregularly,
not straightly; subcaudal photophores well separated
from anal photophores; no pigmentation on outermost
caudal-fin rays (Schultz 1961; 1964; Baird 1971;
Borodulina 1978).
Materials. PMBC 17708, 3 specimens, 31.8–45.0mm SL, PMBC 17709, 6 specimens, 34.1–55.4mm SL, St. J8, 7°15'N 97°33'E to 7°15'N 97°30'E,
473–494 m depth, otter trawl, 18 Feb. 2000;
PMBC 17710, 1 specimen, 32.1 mm SL, St. J10,
7°15'N 97°16'E to 7°15'N 97°14'E, 662–696 m
depth, otter trawl, 19 Feb. 2000; PMBC 17711, 1
specimen, 18.2 mm SL, St. L8, 6°46'N 97°33'E to
6°44'N 97°35'E, 513–501 m depth, otter trawl, 22
Feb. 2000.
Distribution. Tropical to temperate waters inworld oceans (e.g., Schultz 1961; 1964; Baird
1971; 1986; Borodulina 1978; Quéro et al. 1990b;
Paxton et al. 2006b; Satapoomin 2011; Aizawa andDoiuchi 2013a; Harold et al. 2015; Sutton et al.
2020; Villarins et al. 2022).
Polyipnus asper Harold, 1994
Fig. 10
Diagnosis. Anal-fin rays 15–17; gill rakers on firstgill arch 5–6 + 12–14 = 17–20; ACB photophores
8–10; palatine teeth absent; posttemporal bonewith serrate external keels; posttemporal spines 2;
longitudinal parietal keel discontinuous; opposed
median spines on posterior region of parietal 2;
dentate scales present below PV, VAV and ACC
photophores, respectively; dentate scales present
or absent below ACB photophores; lateral dark
pigment bar broad and long, not extending ventrally
further than level of center of orbit (Harold 1994;
Harold et al. 1998; present study).
Materials. PMBC 17712, 1 specimen, 48.8 mm
SL, St. Z2, 7°42'N 97°28'E to 7°42'N 97°31'E,
464 m depth, otter trawl, 23 Jan. 1999; PMBC
17713, 2 specimens, 26.5–47.8 mm SL, St. C8,
9°00'N 96°15'E to 9°00'N 96°13'E, 478–480 m
depth, Agassiz trawl, 3 Feb. 2000; PMBC 17714, 1

specimen, 55.5 mm SL, St. E7, 8°30'N 97°01'E to
8°29'N 97°03'E, 449–446 m depth, Agassiz trawl, 8Feb. 2000; PMBC 17715, 4 specimens, 52.3–59.3mm SL, St. E7, 8°30'N 97°07'E to 8°29'N 97°04'E,
435–444 m depth, otter trawl, 9 Feb. 2000; PMBC
21474, 3 specimens, 42.1–43.1 mm SL, PMBC
30605*, 1 specimen, 31.5 mm SL, station and dateunknown.
Distribution. Andaman
Satapoomin 2011).

Sea(Harold

1994;

Remarks. Polyipnus asper has conspicuous dentate
scales below ACB photophores based on previous
descriptions (Harold 1994; Harold et al. 1998)
and judged from the figure of the holotype (fig.
42 in Harold 1994). However, present specimens
have very weak or invisible denticles on scales
below ACB photophores. This suggests P. asper has
intraspecific morphological variation of denticles onscales below ACB photophores.
Sternoptyx obscura Garman, 1899
Fig. 11
Diagnosis. Maximum body depth from base of
first dorsal-fin ray to base of postabdominal spineless than 82% SL; posterior body depth from endof dorsal-fin base to ventral margin of anal photophores
less than 33% SL; area between transparent regionof anal-fin base and anal photophores not sphenoid
in shape (Schultz 1961; 1964; Baird 1971; Haruta
and Kawaguchi 1976; Borodulina 1978; Aizawa
and Doiuchi 2013a).
Materials. PMBC 17716, 1 specimen, 28.5 mm
SL, St. Z4, 7°34'N 97°03'E to 7°35'N 97°04'E,
660–633 m depth, otter trawl, 25 Jan. 1999; PMBC
17717, 1 specimen, 24.0 mm SL, PMBC 30692, 1
specimen, 21.1 mm SL, St. J10, 7°15'N 97°16'E to
7°15'N 97°14'E, 662–696 m depth, otter trawl, 19
Feb. 2000.
Distribution. Tropical to subtropical waters inIndian and Pacific Oceans (e.g., Schultz 1961;
1964; Baird 1971; 1986; Haruta and Kawaguchi
1976; Borodulina 1978; Satapoomin 2011; Aizawa
and Doiuchi 2013a; Harold et al. 2015).

29
Deep-sea fishes from the Andaman Sea
Suborder Phosichthyoidei
Family Phosichthyidae
Pollichthys mauli (Poll, 1953)
Fig. 12
Diagnosis. Anal-fin rays 22–30; gill rakers on firstgill arch 5 + 11–12 = 16–17; ORB photophores 2;
OP photophores 3; photophores from isthmus topectoral fin 9; AC photophores 18–21, with 13–15

on anal-fin base; anal-fin origin just below orslightly posterior to dorsal-fin origin; anal-fin basemuch longer than dorsal-fin base; SO photophorepresent; eighth and/or ninth IV photophores slightly
elevated (Poll 1953; Grey 1964; Kawaguchi 1971).
Material. PMBC 30606, 1 specimen, 21.8 mm SL,
St. L6, 6°45'N 98°06'E to 6°44'N 98°05'E, 303–313
m depth, Agassiz trawl, 23 Feb. 2000.

Figure 8. Argyropelecus affinis, PMBC 17706, 47.5 mm SL. Scale bar 10 mm.

Figure 9. Argyropelecus sladeni, PMBC 17709, 55.4 mm SL. Scale bar 10 mm.

30
Phuket mar. biol. Cent. Res. Bull.
Distribution. Tropical to temperate waters in northernand eastern Atlantic, Indian Ocean off Réunion andIndonesia, tropical to subtropical waters in western
Pacific (e.g., Grey 1964; Kawaguchi 1971; Badcock
1984c; Schaefer et al. 1986b; Quéro et al. 1990c;
Harold 1999; 2002; Liao et al. 2008; Fricke et al.
2009) including Andaman Sea (present study).
Remarks. Situations of the dorsal- and anal-fin
origins and pigmentation of the dorsal head of thepresent specimen well resembled those of juvenileof this species (Ozawa 1976: fig. 3F).
In the Indian Ocean, this species has been
known only from off Réunion, Madagascar andIndonesia (Shcherbachev et al. 1986; Fricke et al.
2009; Gloerfelt-Tarp and Kailola 2022). Therefore,
this study is the first record of the species from theAndaman Sea.
Vinciguerria nimbaria(Jordan and Williams, 1895)
Fig. 13
Diagnosis. Gill rakers on first gill arch 17–26; totalvertebrae 39–44; premaxilla with 6–10 small
subequal teeth; ORB photophores 2; IV photophores
21–24; VAV photophores 8–11; summation of ventral
two series of photophores 64–73; SO photophorepresent (Grey 1964; Kawaguchi 1971; Gorbunova
1972; Johnson and Barnett 1975; Johnson and Feltes
1984).
Materials. PMBC 30503*, 4 specimens, 18.8–22.5mm SL, St. Z2, 7°42'N 97°28'E to 7°42'N 97°31'E,
464 m depth, otter trawl, 23 Jan. 1999; PMBC
30607, 1 specimen, 23.9 mm SL, St. J8, 7°15'N
97°33'E to 7°15'N 97°30'E, 473–494 m depth, otter
trawl, 18 Feb. 2000.
Distribution. Tropical to temperate waters in northern Atlantic, western central Atlantic and Indian
Oceans except Red Sea, and Pacific (e.g., Kawaguchi
1971; Gorbunova 1972; Badcock 1984c; Schaefer
et al. 1986b; Quéro et al. 1990c; Harold 1999;
Paxton et al. 2006c; Kenaley and Stewart 2015c).

Family Stomiidae
Subfamily Astronesthinae
Astronesthes formosana Liao, Chen and Shao, 2006
Fig. 14
Diagnosis. Dorsal-fin rays 17–20; anal-fin rays
13–15; gill filaments on first gill arch short, their
number more than 40 pairs; total vertebrae 45–48;
IC photophores 49–54; OA photophores 31–34;
chin barbel well developed, length increasing withbody growth, reaching 50–80% HL (specimens
over 40–50 mm SL); tip of barbel slightly swollen
or rounded; anal fin originating anterior to verticalthrough end of dorsal fin; anal-fin base shorter thandorsal-fin base; IV photophores arched outward on
pelvic-fin base; dense aggregations of luminous
tissue present on head and/or body; single luminous
tissue present on opercle; luminous tissue absent
on upper jaw and present on lower jaw; luminous
tissue not developed on head and body; smear-likeluminous patches or more than 1 pair of prominent
luminous patches present on dorsal head behind
upper nostril; no caudal light glands (Goodyear
and Gibbs 1970; Parin and Borodulina 1997; 2003;
Liao et al. 2006; Koizumi et al. 2021; Rajeev et al.
2022; present study).
Materials. PMBC 30366, 1 specimen, 57.3 mm
SL, St. Z2, 7°42′N 97°28′E to 7°42′N 97°31′E, 464
m depth, otter trawl, 23 Jan. 1999; PMBC 30369,
1 specimen, 90.3 mm SL, St. J8, 7°15′N 97°33′E
to 7°15′N 97°30′E, 473–494 m depth, otter trawl,
18 Feb. 2000; PMBC 30382, 1 specimen, 48.2 mm
SL, HUMZ 230163, 1 specimen, 81.0 mm SL, St.
E7, 8°30′N 97°07′E to 8°29′N 97°04′E, 435–444 m
depth, otter trawl, 9 Feb. 2000.
Distribution. Arabian Sea, western Pacific aroundTaiwan and Japan (Liao et al. 2006; Koizumi et al.
2021; Rajeev et al. 2022) including Andaman Sea(present study).
Remarks. The number of total vertebrae of present
specimens (45–46) are slightly different from thatof previous descriptions (46–48: Liao et al. 2006;
Koizumi et al. 2021; Rajeev et al. 2022).

31
Deep-sea fishes from the Andaman Sea
In the Indian Ocean, this species has been
known only from the Arabian Sea (Rajeev et al.
2022). Therefore, this study is the first record ofthis species from the Andaman Sea.

Astronesthes indopacifica Parinand Borodulina, 1997
Fig. 15
Diagnosis. Dorsal-fin rays 17–20; anal-fin rays
13–16; gill filament on first gill arch short, their

Figure 10. Polyipnus asper, PMBC 17715, 58.9 mm SL. Scale bar 10 mm.

Figure 11. Sternoptyx obscura, PMBC 17716, 28.5 mm SL. Scale bar 10 mm.

32
Phuket mar. biol. Cent. Res. Bull.
number more than 40 pairs; total vertebrae 45–49;
IC photophores 50–54; OA photophores 31–34;
chin barbel well developed, length increasing withgrowth, reaching 60–90% HL (specimens over
40–50 mm SL); tip of barbel simple, usually not
swollen or rounded, or rarely swollen or rounded;
anal fin originating anterior to vertical through endof dorsal fin; anal-fin base shorter than dorsal-finbase; IV photophores arched outward on pelvic-finbase; dense aggregations of luminous tissue present
on head and/or body; single luminous tissue present
on opercle; no luminous tissue on both jaws; luminous
tissue not developed on head and body; smear-likeluminous patches or more than 1 pair of prominent
luminous patches present on dorsal head behind
upper nostril; no caudal light glands (Goodyear
and Gibbs 1970; Parin and Borodulina 1997;
2003; Liao et al. 2006; Koizumi et al. 2021; present
study).
Materials. PMBC 30368, 1 specimen, 53.8 mm SL,
St. J8, 7°15′N 97°33′E to 7°15′N 97°30′E, 473–494

m depth, otter trawl, 18 Feb. 2000; PMBC 30378, 1
specimen, 46.1 mm SL, St. G8, 8°00′N 97°06′E to8°00′N 97°04′E, 508–518 m depth, otter trawl, 20 Nov.
1999.
Distribution. Tropical to temperate waters inIndian, and western and central Pacific Oceans(e.g., Parin and Borodulina 1997; Liao et al. 2006;
Paxton et al. 2006d; Stewart and Kenaley 2015a)
including Andaman Sea (present study).
Remarks. The number of anal-fin rays of present
specimens (12–15) are slightly different from thatof previous descriptions (13–16: Liao et al. 2006;
Koizumi et al. 2021).
In the Indian Ocean, A. indopacifica has been
known only from the western and central regions(e.g., Parin and Borodulina 1997; 2003). Therefore,
this study is the first record of this species from theAndaman Sea.

Figure 12. Pollichthys mauli, PMBC 30606, 21.8 mm SL. Scale bar 5 mm.

Figure 13. Vinciguerria nimbaria, PMBC 30607, 23.9 mm SL. Scale bar 5 mm.

33
Deep-sea fishes from the Andaman Sea
Astronesthes martensii Klunzinger, 1871
Fig. 16
Diagnosis. Dorsal-fin rays 11–13; anal-fin rays 17–
21; total vertebrae 50–52; IC photophores 58–64;
OA photophores 35–38; anal-fin base longer thandorsal-fin base; length of chin barbel usually notless than HL; terminal bulb of barbel without
appendages; depth of caudal peduncle 1.2–1.7 times
in its length; 2–3 middle photophores in AC series

elevated posterior to anal fin; no aggregations ofluminous tissue on head; one black spot present on
caudal peduncle behind anal fin; transverse pigment
bands around accessory photophores present (Parinand Borodulina 1995; 2003).
Materials. PMBC 30693, 2 specimens, 70.2–73.2mm SL, St. J7, 7°15′S 7°53′E to 7°16′S 7°52′E,
356–360 m depth, Agassiz trawl, 17 Feb. 2000.

Figure 14. Astronesthes formosana, PMBC 30369, 90.3 mm SL. Scale bar 10 mm.

Figure 15. Astronesthes indopacifica, PMBC 30368, 53.8 mm SL. Scale bar 10 mm.

Figure 16. Astronesthes martensii, PMBC 30693, 73.2 mm SL. Scale bar 10 mm.

34
Phuket mar. biol. Cent. Res. Bull.
Comparative materials. PMBC 30370, 1 specimen,
81.5 mm SL, HUMZ 230161, 1 specimen, 76.2 mm
SL, eastern Indian Ocean, 300–400 m depth, otter
trawl, 8 Sep. 1980.
Distribution. Tropical waters in Indian and western
Pacific Oceans (e.g., Gibbs 1964a; 1986a; Parinand Borodulina 1995; 2003; Harold 1999;
Psomadakis et al. 2019; Gloerfelt-Tarp and Kailola2022).
Borostomias mononema(Regan and Trewavas, 1929)
Fig. 17
Diagnosis. Anal-fin rays 14–19; IC photophores
65–74 (usually 70 or more); VAL photophores
21–25; OA photophores 46–49; postorbital organs
2; anterior part of both jaws with fangs not muchlarger than other teeth; teeth on both jaws curved;
row of AC photophores straight, not curved upwardbehind anal-fin base (Gibbs 1964a; 1984a; 1986a;
Borodulina 2008; present study).
Materials. PMBC 30367, 1 specimen, 76.8 mm
SL, St. J8, 7°15′N 97°30′E to 7°15′N 97°32′E,
490–479 m depth, Agassiz trawl, 18 Feb. 2000;
PMBC 30372, 1 specimen, 192.2 mm SL, St. J8,7°21′N 97°26′E to 7°20′N 97°25′E, 520–531 m
depth, otter trawl, 27 Jan. 1999; PMBC 30390, 1
specimen, 193.8 mm SL, St. J8, 7°15′N 97°33′E
to 7°15′N 97°30′E, 473–494 m depth, otter trawl,
18 Feb. 2000; PMBC 30381, 2 specimens, HUMZ
230162, 1 specimen, SL unmeasured due todamage, station and date unknown.
Distribution. Tropical to temperate waters in eastern
Atlantic, Indian and western Pacific Oceans (Gibbs
1964a; 1984a; 1986a; 1990a; Fujii 1983; Paxton etal. 2006d; Stewart and Kenaley 2015a; Sutton etal. 2020) including Andaman Sea (present study).
Remarks. The number of anal-fin rays and IC
photophores of present specimens (14–18 and 70–
74, respectively) are slightly different from thoseof previous descriptions (15–19 and 65–72: Gibbs
1984a; 1986a).
Borostomias mononema has been known fromthe eastern Indian Ocean (e.g., Gibbs 1964a; Paxton
et al. 2006d), but has not been recorded from theAndaman Sea (e.g., Monkolprasit et al. 1997;

Satapoomin 2011; Rajan et al. 2013; Psomadakis
et al. 2019). Therefore, this study is the first recordof this species from the Andaman Sea.
Subfamily Stomiinae
Stomias affinis Günther, 1887
Fig. 18
Diagnosis. Pectoral-fin rays 6–7; total vertebrae
66–72; premaxillary teeth 5–6; dentary teeth 9–12;
small teeth on each side of dentary around symphysis
3–4; teeth on palatine 2; PV photophores 41–46;
VAV photophores 5–9; IC photophores 73–82;
OV photophores 40–46; VAL photophores 4–8;
OA photophores 46–53; rows of hexagonal areas
dorsal to lateral series of photophores 6; lengthfrom snout to pelvic-fin insertion 73.1–77.0% SL;
distance between pelvic-fin insertion and anal-fin
origin shorter than length of anal-fin base; barbel
length usually about equal to HL; longest mandibular
tooth shorter than longest premaxillary tooth (Ege
1934; Morrow 1964b; Gibbs 1969; Shcherbachev
and Novikova 1976; Fink and Fink 1986; present
study).
Materials. PMBC 30361, 1 specimen, 63.3 mm
SL, St. J8, 7°15′N 97°33′E to 7°15′N 97°30′E,
549–689 m depth, otter trawl, 11 Feb. 1999; PMBC
30362, 3 specimens, 78.2–107.6 mm SL, station
unknown, 25 Jan. 1999; PMBC 30373, 1 specimen,
82.2 mm SL, PMBC 30510*, 2 specimens, 69.6–
82.2 mm SL, St. Z2, 7°42′N 97°28′E to 7°42′N
97°31′E, 464 m depth, otter trawl, 23 Jan. 1999;
PMBC 30506*, 6 specimens, 41.4–118.4 mm SL,
St. E7, 8°30'N 97°07'E to 8°29'N 97°04'E, 435–444
m depth, otter trawl, 9 Feb. 2000; PMBC 30507*,
1 specimen, 121.2 mm SL, St. B8, 9°10′N 96°18′E
to 9°09′N 96°16′E, 489–504 m depth, otter trawl,
11 Feb. 1999; PMBC 30508*, 4 specimens, 78.9–
143.2 mm SL, St. B10, 9°11′N 96°12′E to 9°10′N
96°14′E, 689–549 m depth, otter trawl, 11 Feb.
1999; PMBC 30509*, 2 specimens, 89.2–119.2mm SL, PMBC 30513*, 4 specimens, 99.8–60.1
mm SL, PMBC 30515*, 1 specimen, 45.3 mm SL,
St. J8, 7°15′N 97°33′E to 7°15′N97°30′E, 473–
494 m depth, otter trawl, 18 Feb. 2000; PMBC
30511*, 1 specimen, 141.4 mm SL, St. G8, 8°00′N
97°06′E to 8°00′N 97°04′E, 508–518 m depth, otter
trawl, 20 Nov. 1999; PMBC 30512*, 2 specimens,
96.0–120.2 mm SL, St. J10, 7°20′N 97°14′E to7°22′N 97°13′E, 655–651 m depth, otter trawl, 28

35
Deep-sea fishes from the Andaman Sea
Jan. 1999; PMBC 30514*, 3 specimens, 102.3–
139.8 mm SL, St. Z3, 7°42′N 97°20′E to 7°42′N
97°18′E, 493–322 m depth, otter trawl, 24 Jan.
1999; HUMZ 230157, 1 specimen, 119.1 mm SL,
HUMZ 230158, 1 specimen, 145.0 mm SL, station
unknown, 25 Jan. 1999.
Distribution. Tropical to temperate waters in world
oceans (e.g., Ege 1934; Morrow 1964b; Gibbs 1969;1990b; Shcherbachev and Novikova 1976; Paxton etal. 2006e; Rajan et al. 2013; Kenaley and Stewart
2015d; Koeda 2020b; Sutton et al. 2020).
Remarks. The number of OA photophores ofpresent specimens (46–48) is slightly different
from that of previous descriptions (48–53: Gibbs
1969; Shcherbachev and Novikova 1976).
Stomias nebulosus Alcock, 1889
Fig. 19
Diagnosis. Pectoral-fin rays 6–7; total vertebrae
59–65; premaxillary teeth 16–25; dentary teeth
9–16; teeth on palatine 2–3; PV photophores
33–38; VAV photophores 5–9; IC photophores
66–73; OV photophores 32–38; VAL photophores
5–9; OA photophores 40–45; rows of hexagonal

areas dorsal to lateral series of photophores 6;
length from snout to pelvic-fin origin 66.7–70.6%
SL; distance between pelvic- and anal-fin origins
shorter than length of anal-fin base; barbel length
usually about equal to HL; longest mandibular
tooth longer than longest premaxillary tooth (Ege
1934; Morrow 1964b; Gibbs 1969; Shcherbachev
and Novikova 1976; Fink and Fink 1986; present
study).
Materials. PMBC 30360, 5 specimens, 64.1–91.9mm SL, PMBC 30526*, 1 specimen, 99.1 mm SL,
St. B10, 9°11'N 96°12'E to 9°10'N 96°14'E, 689–
549 m depth, otter trawl, 11 Feb. 1999; PMBC
30364, 2 specimens, 51.1–59.9 mm SL, PMBC
30517*, 44 specimens, 42.7–98.0 mm SL, PMBC
30518*, 3 specimens, SL unmeasured due to damage,
PMBC 30520*, 3 specimens, 72.0–103.6 mm SL,
PMBC 30525*, 1 specimen, 93.8 mm SL, St. E7,
8°30'N 97°07'E to 8°29'N 97°04'E, 435–444 m
depth, otter trawl, 9 Feb. 2000; PMBC 30516*, 44
specimens, 50.6–101.5 mm SL, PMBC 30521*,
3 specimens, 36.0–107.3 mm SL, PMBC 30522,
6 specimens, 48.2–82.0 mm SL, PMBC 30527*,
6 specimens, 40.7–75.3 mm SL, St. J8, 7°15′N
97°33′E to 7°15′N97°30′E, 473–494 m depth, otter
trawl, 18 Feb. 2000; PMBC 30519*, 1 specimen,

Figure 17. Borostomias mononema, PMBC 30367, 76.8 mm SL. Scale bar 10 mm.

36
Phuket mar. biol. Cent. Res. Bull.
88.8 mm SL, PMBC 30528*, 1 specimen, 106.1mm SL, St. G8, 8°00′N 97°06′E to 8°00′N
97°04′E, 508–518 m depth, otter trawl, 20 Nov.
1999; PMBC 30523*, 1 specimen, 70.5 mm SL,
St. Z2, 7°42′N 97°28′E to 7°42′N 97°31′E, 464 m
depth, otter trawl, 23 Jan. 1999; PMBC 30524*, 1
specimen, 77.2 mm SL, St. L8, 6°46′N 97°33′E to6°44′N 97°35′E, 513–501 m depth, otter trawl, 22
Feb. 2000; HUMZ 230155, 1 specimen, 66.3 mm
SL, HUMZ 230156, 1 specimen, 63.5 mm SL, St.
J8, 7°15′N 97°33′E to 7°15′N 97°30′E, 549–689 m
depth, otter trawl, 11 Feb. 1999.
Distribution. Tropical to temperate waters in Indian
and western and central Pacific Oceans (e.g.,
Ege 1934; Morrow 1964b; Gibbs 1969; 1986b;
Shcherbachev and Novikova 1976; Rajan et al.
2013; Koeda 2020b).
Remarks. The number of teeth on the anterior endof the palatine of the present specimens (2–3) isslightly different from that of previous descriptions (2:Gibbs 1969; Shcherbachev and Novikova 1976).
Chauliodus sloani Bloch and Schneider, 1801
Fig. 20
Diagnosis. Total vertebrae 53–62; VAVphotophores 22–28; IC photophores 64–72 (rarely 62
or 63); OA photophores 43–48 (rarely 42 or 49); rowsof hexagonal areas dorsal to lateral series ofphotophores 5; fourth premaxillary tooth longer than
third; dorsal-fin origin over fifth to nineth (rarely
fourth or tenth) OV photophore; postorbital
photophore round or nearly so; postorbital
photophore present below or anterior to verticalthrough posterior edge of eye (Morrow 1961;
1964a; Gibbs and Hurwitz 1967; Parin andNovikova 1974).
Materials. PMBC 30359, 3 specimens, 118.0–144.8mm SL, PMBC 30534*, 17 specimens, 74.9–167.8mm SL, PMBC 30553*, 1 specimen, ca. 55 mm
SL, PMBC 30560*, 100 specimens, 69.5–170.7mm SL, PMBC 30561*, 87 specimens, 65.3–173.9mm SL, St. J8, 7°15'N 97°33'E to 7°15'N 97°30'E,
473–494 m depth, otter trawl, 18 Feb. 2000; PMBC
30535*, 17 specimens, 61.3–137.9 mm SL, St. L8,
6°46'N 97°33'E to 6°45'N 97°35'E, 513–501 m
depth, otter trawl, 22 Feb. 2000; PMBC 30536*,

13 specimens, 102.9–158.2 mm SL, St. Z3, 7°42'N
97°20'E to 7°42'N 97°18'E, 493–322 m depth, otter
trawl, 24 Jan. 1999; PMBC 30537*, 3 specimens,
120.7–146.3 mm SL, PMBC 30554*, 1 specimen,
111.2 mm SL, PMBC 30556*, 1 specimen, 108.1mm SL, PMBC 30558*, 1 specimen, 95.7 mm SL,
St. G8, 8°00'N 97°06'E to 8°00'N 97°04'E, 508–
518 m depth, otter trawl, 20 Nov. 1999; PMBC
30538*, 132 specimens, 62.3–158.2 mm SL, PMBC
30539*, 34 specimens, 108.9–152.5 mm SL, St.
E7, 8°30'N 97°07'E to 8°29'N 97°04'E, 435–444
m depth, otter trawl, 9 Feb. 2000; PMBC 30540*,
30 specimens, 52.9–168.0 mm SL, St. B10, 9°11'N
96°12'E to 9°10'N 96°14'E, 689–549 m depth, otter
trawl, 11 Feb. 1999; PMBC 30541*, 1 specimen,
126.1 mm SL, St. C10, 9°00'N 96°15'E to 9°00'N
96°13'E, 478–480 m depth, Agassiz trawl, 3 Feb.
2000; PMBC 30542*, 1 specimen, 124.7 mm SL,
St. J8, 7°15'N 97°30'E to 7°15'N 97°32'E, 490–479
m depth, Agassiz trawl, 18 Feb. 2000; PMBC
30543*, 2 specimens, 105.4–149.0 mm SL, PMBC
30546*, 1 specimen, 70.1 mm SL, PMBC 30548*,
1 specimen, 131.8 mm SL, St. J10, 7°20'N 97°14'E
to 7°22'N 97°13'E, 655–651 m depth, otter trawl,
28 Jan. 1999; PMBC 30544*, 4 specimens, 71.3–
120.9 mm SL, station unknown, 25 Jan. 1999;
PMBC 30545*, 10 specimens, 45.7–85.3 mm SL,
PMBC 30557*, 1 specimen, 60.1 mm SL, PMBC
30559*, 21 specimens, 76.2–157.0 mm SL, St. Z2,
7°42'N 97°'28E to 7°42'N 97°31'E, 464 m depth,
otter trawl, 23 Jan. 1999; PMBC 30547*, 2 specimens,
99.4–ca. 105 mm SL, St. K11, 7°00'N 97°18'E to
7°00'N 97°21'E, 828–684 m depth, Agassiz trawl,
16 Nov. 1999; PMBC 30549*, 1 specimen, 111.5mm SL, station and date unknown; PMBC 30550*,
1 specimen, 134.3 mm SL, St. L8, 6°45'N 97°36'E
to 6°44'N 97°34'E, 482–507 m depth, Agassiz
trawl, 22 Feb. 2000; PMBC 30551*, 4 specimens,
113.9–143.5 mm SL, St. C8, 9°00'N 96°15'E to
9°00'N 96°13'E, 478–480 m depth, Agassiz trawl,
3 Feb. 2000; PMBC 30552*, 1 specimen, 150.4mm SL, St. E7, 8°30'N 97°01'E to 8°29'N 97°03'E,
449–446 m depth, Agassiz trawl, 8 Feb. 2000; PMBC
30555*, 2 specimens, ca. 125–158.5 mm SL, St. E9,8°30'N 95°58'E to 8°28'N 95°58'E, 649–650 m depth,
otter trawl, 5 Feb. 1999; HUMZ 230153, 1 specimen,
165.0 mm SL, HUMZ 230154, 1 specimen, 169.6mm SL, St. J10, 7°15′N 97°16′E to 7°15′N 97°14′E,
662–696 m depth, otter trawl, 19 Feb. 2000.

37
Deep-sea fishes from the Andaman Sea

Figure 18. Stomias affinis, PMBC 30362, 107.6 mm SL. Scale bar 10 mm.

Figure 19. Stomias nebulosus, PMBC 30360, 91.9 mm SL. Scale bar 10 mm.

Figure 20. Chauliodus sloani, PMBC 30359, 144.8 mm SL. Scale bar 10 mm.

38
Phuket mar. biol. Cent. Res. Bull.
Distribution. Tropical to temperate waters inworld oceans (e.g., Morrow 1961; 1964a; Gibbs
and Hurwitz 1967; Parin and Novikova 1974; Gibbs
1984b; 1986c; Parin 1990; Paxton et al. 2006e;
Rajan et al. 2013; Stewart 2015; Psomadakis et al.
2019; Sutton et al. 2020).
Subfamily Melanostomiinae
Bathophilus pawneei Parr, 1927
Fig. 21
Diagnosis. Pectoral-fin rays 2 (rarely 1); total vertebrae
45; body depth 9.6–17.8% SL; pelvic-fin basesituated equidistant between dorsal and ventral
profiles of body (Morrow and Gibbs 1964; Barnett
and Gibbs 1968).
Material. PMBC 30375, 1 specimen, 32.6 mm SL,
St. Z2, 7°42′N 97°28′E to 7°42′N 97°31′E, 464 m
depth, otter trawl, 23 Jan. 1999.
Distribution. Tropical to temperate waters in
Atlantic, Indian and western Pacific Oceans (e.g.,Morrow and Gibbs 1964; Barnett and Gibbs 1968;
Gibbs and Barnett 1990; Paxton et al. 2006f;
Aizawa and Doiuchi 2013b; Koeda 2020b; Sutton
et al. 2020; Villarins et al. 2022) including Andaman
Sea (present study).
Remarks. Barnett and Gibbs (1968) included theIndian Ocean in the distribution of B. pawneei.
However, this species has never been known fromthe Andaman Sea (e.g., Monkolprasit et al. 1997;
Satapoomin 2011; Rajan et al. 2013; Psomadakis
et al. 2019). Therefore, this study is the first recordof this species from the Andaman Sea.
Eustomias (Haploclonus) bifilis Gibbs, 1960
Fig. 22
Diagnosis. Pectoral-fin rays 3; pelvic-fin rays 7;
total vertebrae 56–59; premaxillary teeth 6–10;
mandibular teeth 6–10; IC photophores 65–68; OC
photophores 59–62; main stem of barbel ending insingle bulb; barbel having long single branch with
one distal bulb; slender barbel stem without external
pigmentation; single row of black spots present
along part of barbel stem; deep groove on ventral
body absent (Gibbs 1960; Morrow and Gibbs
1964; Parin and Pokhilskaya 1974; Gibbs et al.
1983; Prokofiev 2018; present study).

Material. PMBC 30388, 1 specimen, 46.2 mm SL,
St. J8, 7°15′N 97°33′E to 7°15′N 97°30′E, 473–494
m depth, otter trawl, 18 Feb. 2000.
Distribution. Tropical to temperate waters in Indian
and western and central Pacific Oceans (e.g., Gibbs
1960; Parin and Pokhilskaya 1974; Fujii 1982;
Paxton et al. 2006f; Aizawa and Doiuchi 2013b;
Koeda 2020b; Gloerfelt-Tarp and Kailola 2022)
including Andaman Sea (present study).
Remarks. Although E. (H.) bifilis is widelydistributed in the Indian Ocean (Parin andPokhilskaya 1974), this species has never beenrecorded from the Andaman Sea (e.g., Monkolprasit
et al. 1997; Satapoomin 2011; Rajan et al. 2013;
Psomadakis et al. 2019). Therefore, this study isthe first record of this species from the Andaman
Sea.
Leptostomias multifilis Imai, 1941
Fig. 23
Diagnosis. Dorsal-fin rays 19–21; anal-fin rays
24–27; base of barbel stem with filaments; bulb ofbarbel with one pair of filaments near its base; bulb
of barbel not split; distal part of bulb of barbel withbrush of many short filaments (Imai 1941; Morrow
and Gibbs 1964; Fujii 1984).
Materials. PMBC 30374, 1 specimen, 159.3 mm
SL, St. J8, 7°15′N 97°33′E to 7°15′N 97°30′E,
473–494 m depth, otter trawl, 18 Feb. 2000; PMBC
21475, 1 specimen, 66.8 mm SL, PMBC 30504*, 1
specimen, 184.6 mm SL, station and date unknown.
Distribution. Around Taiwanese and Japanesewaters and southern Kuril Islands (e.g., Imai 1941;
Parin and Sokolovsky 1976; Fujii 1984; Savinykh
et al. 2004; Liao et al. 2008; Aizawa and Doiuchi
2013b) and Andaman Sea (present study).
Remarks. Leptostomias multifilis has been known
only from Taiwan, Japan and Kuril Islands (e.g.,
Imai 1941; Parin and Sokolovsky 1976; Savinykh
et al. 2004; Liao et al. 2008). Therefore, this study
is the first record of this species from the Indian
Ocean.

39
Deep-sea fishes from the Andaman Sea

Figure 21. Bathophilus pawneei, PMBC 30375, 32.6 mm SL. Scale bar 10 mm.

Figure 22. Eustomias (Haploclonus) bifilis, PMBC 30388, 46.2 mm SL. Scale bar 10 mm.

40
Phuket mar. biol. Cent. Res. Bull.
Melanostomias melanops Brauer, 1902
Fig. 24
Diagnosis. IC photophores 60–64; OA photophores
39–41; no luminous tissue on dorsal head; distal
part of barbel expanded; core of barbel stem
continuing in tip of barbel as an axis; tip of barbel
with terminal filament and with/without single
bulb beside axis; barbel stem heavily pigmented
only on its base, sparsely pigmented beyond base
(Regan and Trewavas 1930; Morrow and Gibbs
1964; Parin and Pokhilskaya 1978).

97°30′E, 444–473 m depth, otter trawl, 18 Feb.
2000; PMBC 30385, 1 specimen, 148.8 mm SL, St.
J10, 7°21′N 97°33′E to 7°20′N 97°32′E, 520–532
m depth, otter trawl, 27 Jan. 1999; PMBC 30505*,
1 specimen, 143.5 mm SL, station and date unknown.
Distribution. Tropical to subtropical waters inworld oceans (e.g., Gibbs 1960; 1984c; 1986d;
Morrow and Gibbs 1964; Parin and Pokhilskaya
1978; Gibbs and Barnett 1990; Paxton et al. 2006f;
Aizawa and Doiuchi 2013b; Kenaley and Stewart
2015e; Koeda 2020b; Sutton et al. 2020).

Material. PMBC 30387, 1 specimen, 51.6 mm SL,
St. J8, 7°15′N 97°33′E to 7°15′N 97°30′E, 473–
494 m depth, otter trawl, 18 Feb. 2000.

Photonectes (Photonectes) albipennis(Döderlein, 1882)
Fig. 26

Distribution. Tropical to subtropical waters inworld oceans (e.g., Morrow and Gibbs 1964; Parinand Pokhilskaya 1978; Gibbs 1984c; Gibbs andBarnett 1990; Aizawa and Doiuchi 2013b; Kenaley
and Stewart 2015e; Sutton et al. 2020) including
Andaman Sea (present study).

Diagnosis. BR photophores 5–7; IV photophores
37–38; premaxillary teeth biserial; premaxillary
and dentary teeth heterogenous in juvenile andhomogenous in adult; vertical fins not covered withskin; pectoral fin absent; elongate barbel with aminute terminal bulb and a single terminal filament(sometimes with secondary bulblets); dorsal branchof barbel absent; barbel stem entirely pigmented; 1–2
white luminous shoulder spots present above gill
opening; blue luminous tissue absent on head andbody; IP photophore origin at middle of isthmus; all
VAV photophores equally spaced (Klepadlo 2011;
Prokofiev 2019; Prokofiev and Frable 2021).

Remarks. Melanostomias melanops is widelydistributed in the Indian Ocean except the Andaman
Sea (Morrow and Gibbs 1964; Parin and Pokhilskaya
1978; Monkolprasit et al. 1997; Satapoomin 2011;
Rajan et al. 2013; Psomadakis et al. 2019). Therefore,
this study is the first record of this species from theAndaman Sea.
Melanostomias valdiviae Brauer, 1902
Fig. 25
Diagnosis. IC photophores 56–59; OA photophores
36–39; no luminous tissue on dorsal head; distal
part of barbel not expanded except tip; core of barbel
stem continuing in tip of barbel as an axis; tip ofbarbel without terminal filament, with 3–4 bulbs
beside axis (1–2 bulbs on each side of axis); barbel
stem entirely pigmented except tip (Regan and
Trewavas 1930; Morrow and Gibbs 1964; Parin andPokhilskaya 1978; Aizawa and Doiuchi 2013b).
Materials. PMBC 30389, 1 specimen, 57.1 mm
SL, St. B8, 9°10′N 96°18′E to 9°09′N 96°16′E,
489–504 m depth, otter trawl, 11 Feb. 1999; PMBC
30376, 1 specimen, 139.0 mm SL, St. Z3, 7°42′N
97°20′E to 7°42′N 97°18′E, 322–493 m depth, otter
trawl, 24 Jan. 1999; PMBC 30377, 1 specimen,
111.1 mm SL, St. J10, 7°15′N 97°33′E to 7°15′N

Material. PMBC 30371, 1 specimen, 193.1 mm
SL, St. J10, 7°20′N 97°14′E to 7°22′N 97°13′E,
585–655 m depth, otter trawl, 27 Jan. 1999.
Distribution. Tropical to temperate waters inIndian Ocean and western and central Pacific (e.g.,Morrow and Gibbs 1964; Fujii 1982; Paxton etal. 2006f; Aizawa and Doiuchi 2013b; Prokofiev
2019; Gloerfelt-Tarp and Kailola 2022) including
Andaman Sea (present study).
Remarks. Photonectes (Photonectes) albipennis
has been known from the eastern and western
Indian Ocean (Paxton et al. 2006f; Prokofiev 2019;
Gloerfelt-Tarp and Kailola 2022). However, thisspecies has never been known from the Andaman
Sea (e.g., Monkolprasit et al. 1997; Satapoomin
2011; Rajan et al. 2013; Psomadakis et al. 2019).
Therefore, this study is the first record of this species
from the Andaman Sea.

41
Deep-sea fishes from the Andaman Sea
Photonectes (Photonectes) coffea Klepadlo, 2011
Fig. 27
Diagnosis. BR photophores 5–7; IV photophores
37–39; premaxillary teeth biserial; premaxillary
and dentary teeth heterogenous in juvenile andhomogenous in adult; vertical fins not covered
with skin; pectoral fin absent; elongate barbel withtwo bulbs and a single terminal filament; dorsal
branch of barbel absent; barbel stem entirely
pigmented; 1–2 white luminous shoulder spots
present above gill opening; blue luminous tissue

absent on head and body; IP photophore origin at
middle of isthmus; all VAV photophores equally
spaced (Klepadlo 2011; Flynn and Klepadlo 2012;
Prokofiev 2019; Prokofiev and Frable 2021).
Materials. PMBC 30379, 1 specimen, 71.7 mm
SL, St. E7, 8°30′N 97°07′E to 8°29′N 97°04′E,
435–444 m depth, otter trawl, 9 Feb. 2000; PMBC
30380, 1 specimen, 89.5 mm SL, St. G8, 8°00′N
97°06′E to 8°00′N 97°04′E, 508–518 m depth,
otter trawl, 20 Nov. 1999; PMBC 30383, 1 specimen,
91.5 mm SL, St. Z4, 7°34′N 97°03′E to 7°35′N

Figure 23. Leptostomias multifilis, PMBC 30374, 159.3 mm SL. Scale bar 10 mm.

Figure 24. Melanostomias melanops, PMBC 30387, 51.6 mm SL. Scale bar 10 mm.

42
Phuket mar. biol. Cent. Res. Bull.
97°04′E, 660–633 m depth, otter trawl, 25 Jan.
1999; PMBC 30384, 2 specimens, 89.3–134.0 mm
SL, St. E8, 8°32′N 96°04′E to 8°31′N 96°07′E,
478–488 m depth, otter trawl, 6 Feb. 1999; PMBC
30386, 3 specimens, 109.5–182.5 mm SL, St. J8,7°15′N 97°33′E to 7°15′N 97°30′E, 473–494 m
depth, otter trawl, 18 Feb. 2000; HUMZ 230164, 1
pecimen, 135.3 mm SL, HUMZ 230165, 1 specimen,
125.8 mm SL, St. J8, 7°15′N 97°33′E to 7°15′N
97°30′E, 473–494 m depth, otter trawl, 18 Feb.
2000.
Distribution. Andaman Sea, and tropical to subtropicalwaters in western and central Pacific (Klepadlo 2011;
Prokofiev 2019; Koeda 2020b).
Subfamily Idiacanthinae
Idiacanthus fasciola Peters, 1877
Figs. 28, 29
Diagnosis. VAV photophores 13–18; dorsal-fin
origin anterior to pelvic-fin base; dorsal-fin origin at 22–25th vertebrae; anal-fin origin at 41–
44th vertebrae; pelvic-fin insertion at 25–27thvertebrae; length from pelvic-fin insertion to anal-fin
origin less than length of anal-fin base (Gibbs
1964b; 1984d; Hulley 1986; Gon et al. 2022).
Materials. PMBC 30363, 5 specimens, 73.1–
122.2 mm SL, PMBC 30529*, 1 specimen, 90.2
mm SL, St. E7, 8°30′N 97°07′E to 8°29′N 97°04′E,
435–444 m depth, otter trawl, 9 Feb. 2000; PMBC
30365, 2 specimens, 244.8–267.9 mm SL, PMBC
30531*, 1 specimen, 209.5 mm SL, St. J10, 7°20′N
97°14′E to 7°22′N 97°13′E, 655–651 m depth, otter
trawl, 28 Jan. 1999; PMBC 30530*, 1 specimen,
153.9 mm SL, St. J10, 7°15'N 97°16'E to 7°15'N
97°14'E, 662–696 m depth, otter trawl, 19 Feb.
2000; PMBC 30532*, 2 specimens, 234.1–257.3mm SL, St. Z3, 7°42′N 97°20′E to 7°42′N 97°18′E,
493–422 m depth, otter trawl, 24 Jan. 1999; PMBC
30533*, 1 specimen, 156.1 mm SL, St. L8, 6°46′N
97°33′E to 6°44′N 97°35′E, 513–501 m depth, otter
trawl, 22 Feb. 2000; HUMZ 230159, 1 specimen,
157.2 mm SL, HUMZ 230160, 1 specimen, 73.6mm SL, St. E7, 8°30′N 97°07′E to 8°29′N 97°04′E,
435–444 m depth, otter trawl, 9 Feb. 2000.

Distribution. Tropical and subtropical waters in
Atlantic, Indian and Pacific Oceans (e.g., Gibbs
1964b; 1984d; Hulley 1986; Krueger 1990; Harold,
1999; Randall and Lim 2000; Paxton et al. 2006g;
Aizawa and Doiuchi 2013c; Stewart and Kenaley
2015b; Koeda 2020b; Sutton et al. 2020; Gon etal. 2022) including Andaman Sea (present study).
Remarks. Most of the present specimens have apointed appendage and a terminal filament in the
tip of the barbel (Fig. 29A), which is congruent
with the morphology of the tip of the barbel ofthe previous description (Gibbs 1964b). On theother hand, one individual of PMBC 30363 has
no appendage and terminal filament in the tip ofthe barbel (Fig. 29B). This study identified thisindividual as I. fasciola because the other charactersof this individual are congruent with the diagnosisof I. fasciola: e.g., dorsal-fin origin at 22–25thvertebrae; anal-fin origin at 41–44th vertebrae (Gonet al. 2022). The aberrant morphology of the tip ofthe barbel of this individual is possibly caused bydamage or regeneration.
Although I. fasciola has been known from thewestern and eastern Indian Ocean (e.g., Hulley
1986; Paxton et al. 2006g), this species has neverbeen known from the Andaman Sea (e.g.,
Monkolprasit et al. 1997; Satapoomin 2011; Rajan
et al. 2013; Psomadakis et al. 2019). Therefore,
this study is the new record of this species from theAndaman Sea.
ACKNOWLEDGEMENT
We are grateful to Ms. Jirapath Pattharakowit(PMBC) and staff of the Reference Collection of
PMBC for their help of our research in PMBC. We
also deeply thank A. Prokofiev (Russian Academyof Sciences, Russia) and O. Gon (South African
Institute for Aquatic Biodiversity, South Africa)
for providing literature. We also express gratitude
to two referees for their helpful and valuablesuggestions and comments. This study was partially
supported by JSPS KAKENHI Grant Number
JP23K05364 and JST SPRING Grant Number
JPMJSP2119.

43
Deep-sea fishes from the Andaman Sea

Figure 25. Melanostomias valdiviae, PMBC 30385, 148.8 mm SL. Scale bar 10 mm.

Figure 26. Photonectes (Photonectes) albipennis, PMBC 30371, 193.1 mm SL. Scale bar 10 mm.

Figure 27. Photonectes (Photonectes) coffea, PMBC 30386, 137.0 mm SL. Scale bar 10 mm.

44
Phuket mar. biol. Cent. Res. Bull.

Figure 28. Idiacanthus fasciola, PMBC 30365, 244.8 mm SL. Scale bar 10 mm.

Figure 29. Tip of chin barbel in Idiacanthus fasciola. (A): PMBC 30363, 82.0 mm SL; (B): PMBC 30363,
148.7 mm SL. Scale bars 1 mm.

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Manuscript received: 25 January 2023
Accepted: 3 June 2023

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