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GONOSTOMATIDAE
Bristlemouths
by A.S. Harold
D iagnostic characters: Body moderately elongate (to about 36 cm standard length); head and body
compressed. Relative size of head highly variable. Eye very small (Cyclothone) to moderately large
(Margrethia). Nostrils high on snout, prominent in dorsal view. Mouth large, angle of jaw well posterior to
eye. Premaxillary teeth uniserial (except in Triplophos); dentary teeth biserial near symphysis. Chin
barbel absent. Gill openings very wide. Branchiostegals 12 to 16 (4 to 6 on posterior ceratohyal). Gill
rakers well developed. Pseudobranchiae usually absent (present in Diplophos and Margrethia). Dorsal
fin at or slightly posterior to middle of body (except in Triplophos in which it is anterior). Anal-fin
base moderately to very long. Dorsal fin with 10 to 20 rays; anal fin with 16 to 68 rays; caudal fin forked;
pectoral-fin rays 8 to 16; pelvic-fin rays 5 to 9. Dorsal adipose fin present or absent; ventral adipose fin
absent. Scales deciduous. One or more rows of discrete photophores on body; BR 7-16; OA 0-77; IV
11-51; VAV 3-17; AC 12-51; isthmus photophores (IP) present or absent; posterior orbital
photophore (ORB 2) absent. Parietals well developed; epiocciptals separated by supraoccipital. Four
pectoral-fin radials (except Cyclothone, which has 1). Colour: skin varying from colourless through brown
to black; black and silvery pigmentation associated with photophores.
ORB 2 absent OA
ORB 1 OP
IP Diplophos
PV
VAV AC
IV
Bonapartia
Habitat, biology, and fisheries: Mesopelagic and bathypelagic, oceanic. Development, especially of
photophores, protracted. Diet consists of other fishes and crustaceans.
Stomiiformes: Gonostomatidae 1897
photophores in
Sternoptychidae clusters
photophores in
Astronesthidae, Chauliodontidae, Idiacanthidae, clusters
Malacosteidae, Melanostomiidae, and Stomiidae:
similar arrangement of photophores but body Sternoptychidae
generally more elongate; chin barbel usually present
(reduced or absent in Chauliodontidae, absent in males of Idiacanthidae, and absent in Malacosteus of the
Malacosteidae); jaw teeth greatly enlarged, fang-like; gill rakers absent in adults; usually 3 bony
pectoral-fin radials.
Myctophidae and Neoscopelidae (myctophiform families): also with photophores, but usually with a less
elongate body and the maxilla is completely excluded from the gape by the premaxilla; ventral photophores
usually more widely spaced and not arranged in such regular rows as in the Gonostomatidae.
fang-like
teeth
maxilla
premaxilla
chin barbel
present Astronesthidae Neoscopelidae
2a. VAV photophores 12 to 17 (Fig. 2); ORB photophore below or slightly ahead of anterior
margin of eye; lower gill rakers 7 to 10; a row of small photophores present on posterior
half of lower jaw; pseudobranchiae present; IV photophores 29 to 51 . . . . . . . . . . . . . ® 3
2b. VAV photophores 5 to 7 (Fig. 3); ORB photophore below centre of eye; lower gill rakers
12 to 16; no photophores present on posterior half of lower jaw; pseudobranchiae
absent; IV photophores 24 to 30; dorsal fin well anterior to middle of body . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Triplophos Brauer, 1902
OA
VAV
IV VAV
Fig. 2 Diplophos Fig. 3 Triplophos
4a. Dorsal-fin origin anterior to anal-fin origin (Fig. 5); pseudobranchiae well developed
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Margrethia Jespersen and Tåning, 1919
4b. Dorsal-fin origin above or posterior to anal-fin origin (Fig. 6); pseudobranchiae
inconspicuous or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 5
5a. Anal fin with elongate anterior rays (Fig. 6); OA photophores absent; pectoral-fin rays
14 to 16 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bonapartia Goode and Bean, 1896
5b. Anterior rays of anal fin not elongate; OA photophores present (6 to 16); pectoral-fin
rays 7 to 13 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 6
OA photophores absent
6a. Eyes moderate to small (Fig. 7a); OA photophores 11 to 21; SO photophore usually
present (absent in G. bathyphilum); vertebrae 37 to 40; maxilla with a series of
elongate, subequal teeth separated by a series of shorter, subequal teeth (Fig. 8a);
palatine teeth in a single row; anal-fin rays 20 to 32 . . . . . . . . Gonostoma Rafinesque, 1810
6b. Eyes very small (Fig. 7b); OA photophores 6 to 10; SO photophore absent; vertebrae
29 to 33; maxillary teeth all short and subequal or with occasional slightly longer teeth
distributed at roughly equal intervals (Fig. 8b); palatine teeth in an anterior patch;
anal-fin rays 16 to 21 . . . . . . . . . . . . . . . . . . . . . . Cyclothone Goode and Bean, 1883
eye large
a) Gonostoma
b) Cyclothone
a) Gonostoma b) Cyclothone
Fig. 7 head Fig. 8 teeth on maxilla (upper jaw)
References
Fujii, E. 1984. Family Gonostomatidae. In Fishes of the Japanese Archipelago, edited by H. Masuda, K. Amaoka,
C. Araga, T. Uyeno, and T. Yoshino. Tokyo, Tokai University Press, pp. 48-49.
Paxton, J.R. and J.E. Hanley. 1989. Gonostomatidae. In Zoological catalogue of Australia 7. Pisces: Petromyzontidae
to Carangidae, edited by J.R. Paxton , D.F. Hoese, G.R. Allen, and J.E. Hanley, pp. 183-186.
Schaefer, S., R.K. Johnson, and J. Badcock. 1986. Gonostomatidae. In Smiths’ sea fishes, edited by M.M. Smith and
P.C. Heemstra. Johannesburg, Macmillan South Africa, pp. 247-253.
1900 Bony Fishes
D iagnostic characters: Moderately elongate to deep-bodied (to about 10 cm standard length). Head
moderate to very large, more than 1/3 of standard length in Sternoptyx. Eye large, up to about 1/2 of
head length, directed vertically and telescopic in some Argyropelecus species. Snout short. Mouth small to
moderate in size, terminal to dorsal oblique; jaw teeth generally small, some Argyropelecus species with
well-developed canine teeth. Chin barbel absent. Branchiostegals 6 to 10. Pseudobranch present. Gill
rakers well developed. Dorsal-fin origin usually near middle of body (anterior in Danaphos, posterior in
Araiophos). Anal fin moderate to long-based, interrupted centrally by a group of photophores in some
genera (e.g. Argyripnus, Argyropelecus, Polyipnus). Dorsal fin with 6 to 20 rays; anal fin with 17 to 38 rays;
caudal fin forked; pectoral fin with 11 to 18 rays; pelvic fins with 5 to 7 rays. Dorsal adipose fin usually
present; absent in Araiophos eastropas and Polyipnus latirastrus. Scales present, deciduous. Ventral
photophore series with disjunct clusters of 2 or more photophores. Two ventrolateral rows of
photophores on body; OA 0-10; IV 10-24; VAV 3-32; AC 3-51; paired row of photophores on isthmus (IP);
branchiostgeal photophores 6 (7 in Sonoda); 1 orbital photophore present (ORB 1); posterior orbital
photophore (ORB 2) absent (present in most Phosichthyidae). Three branchiostegal rays originating on
posterior ceratohyal. Four pectoral-fin radials. Photophores Alpha type. Colour: skin light to dark
brown, often with reflective guanine pigment on side of body; silvery and black pigmentation usually
associated with photophores. Some species with dark dorsum pigment and/or with saddle-like markings
and lateral bars or incomplete stripes (Polyipnus).
OP
OA
AC
VAV
IP
Argyropelecus
PV
IV
ORB 1 OP OA
SO
BR Maurolicus
IP PV VAV AC
IV
AC - ventral series posterior to anal-fin origin OP - opercular photophores
BR - on the branchiostegal membranes ORB 1 - anterior to eye
IP - ventral series anterior to pectoral-fin base PV - ventral series between bases of pectoral and pelvic fins
IV - ventral series anterior to pelvic-fin base SO - paired photophores near symphysis of lower jaw
OA - lateral series VAV - ventral series between pelvic-fin base and anal-fin origin
ventral
IP photophores ventral photophores
present or absent photophores not not in clusters
Gonostomatidae in clusters Phosichthyidae
maxilla
photophores
chin barbel not in clusters premaxilla photophores
usually present not in clusters
Astronesthidae Neoscopelidae
References
Ahlstrom, E.H., W.J. Richards, and S.H. Weitzman. 1984. Families Gonostomatidae, Sternoptychidae, and associated
stomiiform groups: development and relationships. In Ontogeny and systematics of fishes, edited by H.G. Moser,
W.J. Richards, D.M. Cohen, M.P. Fahay, A.W. Kendall, Jr., and S.L. Richardson. Amer. Soc. Ichthyol. Herpetol.,
Spec. Publ., (1):184-198.
Baird, R.C. 1971. The systematics, distribution and zoogeography of the marine hatchetfishes (family Sternoptychidae).
Bull. Mus. Comp. Zool., Harvard Univ., 142:1-128.
Fujii, E. 1984. Family Sternoptychidae. In Fishes of the Japanese Archipelago, edited by H. Masuda, K. Amaoka,
C. Araga, T. Uyeno, and T. Yoshino. Tokyo, Tokai University Press, pp. 47-48.
Harold, A.S. 1994. A taxonomic revision fo the sternoptychid genus Polyipnus (Teleostei: Stomiiformes), with an
analysis of phylogenetic relationships. Bull. Mar. Sci., 54:428-534.
Parin, N.V. and S.G. Kobyliansky. 1993. Review of the genus Maurolicus (Sternoptychidae, Stomiiformes), with
re-establishing validity of five species considered junior synonyms of M. muelleri and descriptions of nine new
species. Tr. Inst. Okeanol., 128:69-107.
Paxton, J.R. and J.E. Hanley. 1989. Sternoptychidae. In Zoological catalogue of Australia 7. Pisces: Petromyzontidae
to Carangidae, edited by J.R. Paxton, D.F. Hoese, G.R. Allen, and J.E. Hanley, pp. 190-195.
Stomiiformes: Phosichthyidae 1903
D iagnostic characters: Body generally moderately elongate (to about 30 cm standard length),
somewhat deep-bodied in some Ichthyococcus species. Head and body compressed; head small to
moderately large. Mouth large; premaxillary teeth uni- or biserial, dentary teeth biserial anteriorly, uniserial
posterially; teeth ranging in size from small to large but not highly elongate and fang-like. Chin barbel
absent. Eleven to 22 branchiostegal rays, 4 to 7 on posterior ceratohyal. Gill rakers well developed (true
gill rakers restricted to angle of arch in Woodsia). Pseudobranchiae absent, except Woodsia
meyerwaardeni. Dorsal fin usually near middle of body, its origin well in advance of anal-fin origin
(except Pollichthys which has the dorsal-fin orgin immediately above that of the anal fin). Anal fin
with a moderately long base but terminating posteriorly ahead of the most constricted portion of caudal
peduncle. Dorsal fin with 10 to 16 rays; anal fin with 12 to 33 rays; caudal fin forked; pectoral-fin rays 7 to 11;
pelvic-fin rays 6 to 8. Dorsal adipose fin present (except Yarrella). Scales present, deciduous. Two
ventrolateral rows of well-developed photophores on body, rows of accessory photophores dorsal to main
rows in Yarrella; OA 17-53; IV 19-28; VAV 7-17; AC 12-28; row of photophores on isthmus (IP); 2 orbital
photophores (ORB), except Polymetme and Yarrella which lack the posterior ORB 2. Three
pectoral-fin radials. Usually 2 supramaxillae. Gamma type photophores; serial photophores with a lumen
and a duct. Colour: skin varying from light brown in Pollichthys to dark brown or nearly black in Yarrella,
scales of dorsum often outlined with dark pigment.
ORB OP OA
adipose fin
SO
VAV AC lateral view
IP
BR PV OA
AC
SO
PV ventral view
IP VAV
IV
AC - ventral series posterior to anal-fin origin OP - opercular photophores
BR - on the branchiostegal membranes ORB - anterior (ORB 1) and posterior (ORB 2) to eye
IP - ventral series anterior to pectoral-fin base PV - ventral series between bases of pectoral and pelvic fins
IV - ventral series anterior to pelvic-fin base SO - paired photophores near symphysis of lower jaw
OA - lateral series VAV - ventral series between pelvic-fin base and anal-fin origin
abbreviated terminology of photophores
Habitat, biology, and fisheries: Mesopelagic and bathypelagic adults (Yarrella and Polymetme may be
benthopelagic). Larvae nearer surface than adults. Diet consists mainly of zooplankton, crustaceans in
particular.
ORB
Similar families occurring in the area
Gonostomatidae: photophores Beta type; photophores
present on isthmus (IP) (Diplophos, Manducus and
Triplophos ) or absent ( Bonapartia, Cyclothone,
Gonostoma and Margrethia); posterior orbital (ORB 2) IP
photophore absent; 4 bony pectoral-fin radials. Gonostomatidae
1904 Bony Fishes
photophore
clusters
Sternoptychidae Sternoptychidae
photophore clusters
fang-like teeth
maxilla
References
Fujii, E. 1984. Family Gonostomatidae. In Fishes of the Japanese Archipelago, edited by H. Masuda, K. Amaoka,
C. Araga, T. Uyeno, and T. Yoshino. Tokyo, Tokai University Press, pp. 44-47.
Paxton, J.R. and J.E. Hanley. 1989. Photichthyidae. In Zoological catalogue of Australia 7. Pisces: Petromyzontidae to
Carangidae, edited by J.R. Paxton, D.F. Hoese, G.R. Allen, and J.E. Hanley, pp. 187-189.
Stomiiformes: Astronesthidae 1905
D iagnostic characters: Body fusiform and slightly compressed (to about 21.5 cm standard length,
commonly to 15 cm). Head moderately large, about 25% of standard length. Eye small, less than 1/4 of
head length. Snout short to moderate, its length at most twice eye diameter. Mouth large with a small
number of teeth ranging in size from small to large and fang-like. Vomerine and palatine teeth present.
Elongate chin barbel present; terminal bulb present or absent. Branchiostegal rays 14 to 25. Gill rakers
rudimentary in adults. Dorsal fin usually near middle of body, its origin well in advance of anal-fin
origin. Anal fin terminating posteriorly on constricted portion of caudal peduncle. Dorsal fin with 9 to 21
rays; anal fin with 12 to 28 rays; caudal fin forked; pectoral fins with 6 to 9 rays; pelvic fins with 5 to 9 rays.
Dorsal adipose fin present (except in Rhadinesthes). Scales absent, no hexagonal areas on body. Two
ventrolateral rows of photophores on body; OA 11-56; IV 9-46; VAV 7-28; AC 6-18; anterior portion of
lower row (PV) curving upward at pelvic-fin base and appearing disjunct from posterior portion of
row (VAV); row of photophores on isthmus (IP); many small photophores covering much of body and head,
their greatest concentration ventrally; suborbital photophore small, inconspicuous, located at
anteroventral margin of eye; postorbital photophore prominent, located posteroventrally to eye.
Three pectoral-fin radials. Stomach usually large and black, intestine originating near its anterior end; 1 or
2 pyloric caecae present. Colour: skin usually black, silvery pigmentation occasionally present on flank;
patches of luminous tissue on body in many species.
suborbital postorbital OA
photophore photophore
AC
IP PV VAV
IV
AC - ventral series posterior to anal-fin origin OA - lateral series
IP - ventral series anterior to pectoral-fin base PV - ventral series between bases of pectoral and pelvic fins
IV - ventral series anterior to pelvic-fin base VAV - ventral series between pelvic-fin base and origin of anal fin
abbreviated terminology of photophores
Habitat, biology, and fisheries: Mainly mesopelagic adults (some species benthopelagic), juveniles
have been caught at the surface at night. Diet consists of other mesopelagic fishes and crustaceans.
Similar families occurring in the area
Chauliodontidae: body elongate, maximum depth at back of head; chin barbel short and simple, becoming
reduced or absent in adults; dorsal fin well anterior, its origin between vertical through pectoral and pelvic
fins, first ray prolonged; scale-like areas on body delineated by hexagonal pigmentation pattern.
Idiacanthidae: body highly elongate, eel-like; dorsal fin with very long base (54 to 74 rays), its origin well
anterior to midbody; bases of dorsal- and anal-fin rays with a small sharp spur; dorsal adipose fin absent;
pectoral fins present in larvae, absent in adults; pelvic fins absent in males.
54-74 dorsal-fin rays
hexagonal
pigmentation pattern
pectoral fins
absent in adults
Malacosteidae: no membrane in floor of mouth; chin barbel present or absent; presence of a preorbital light
organ, associated with suborbital organ (absent in Photostomias); dorsal fin located posteriorly, just ahead
of caudal fin; dorsal adipose fin absent; pectoral fins absent or consisting of only free, filament-like rays.
Melanostomiidae: maxillae with erect teeth anteriorly and small, oblique denticles posteriorly; chin barbel
present; dorsal fin located posteriorly, just ahead of caudal fin; dorsal adipose fin absent; pectoral fins
present or absent; suborbital photophore absent. dorsal adipose fin absent
suborbital photophore absent
Stomiidae: body elongate and slender; chin barbel present, with terminal bulb and filaments; scale-like
areas on body delineated by hexagonal pigmentation pattern; dorsal fin located posteriorly, just ahead of
caudal fin; dorsal adipose fin absent.
Gonostomatidae, Phosichthyidae, and Sternoptychidae: these remaining stomiiform families often with
somewhat similar body form and photophore arrangement, but have gill rakers as adults and lack both a
chin barbel and greatly enlarged, fang-like jaw teeth.
small jaw teeth
hexagonal pigmentation no dorsal adipose fin
D iagnostic characters: Body long and slender (to 41 cm standard length), covered by a gelatinous
membrane with luminous inclusions in life. Head small. Mouth large, relative to head, with teeth
ranging in size from small to moderately large and fang-like; posterior teeth of upper jaw (maxilla) not as
small oblique denticles; jaws curved upward slightly. Chin barbel prominent, terminal bulb and filaments
present. Branchiostegals 16 to 18. No well-developed gill rakers in adults. Dorsal fin located well
posteriorly just anterior to caudal fin, directly above anal fin. Anal fin terminating posteriorly just
anterior to constricted portion of caudal peduncle. Dorsal fin with 13 to 23 rays; anal fin with 15 to 25 rays;
caudal fin forked or rounded; pectoral fins with 6 to 9 rays; pelvic fins with 4 or 5 rays. Dorsal adipose fin
absent. Body covered with hexagonal scale-like areas. Two ventrolateral rows of photophores on body;
OA 36-67 or 137-153; IV 41-64 or 89-99; VAV 5-16 or 58-67; AC 14-22; paired row of photophores on
isthmus (IP); 1 or more small photophores associated with hexagonal scale-like areas; suborbital
photophore small, inconspicuous, located at anteroventral margin of eye; postorbital photophore
prominent, located posteroventrally to eye. Colour: body scale rows delineated by a hexagonal
pigmentation pattern.
postorbital photophore OA hexagonal pigmentation pattern present
IP PV
VAV
AC
IV
chin barbel with
bulb and filaments
AC - ventral series posterior to anal-fin origin
suborbital postorbital photophore
IP - ventral series anterior to pectoral-fin base
photophore IV IV - ventral series anterior to pelvic-fin base
OA - lateral series
PV - ventral series between bases of pectoral and pelvic fins
oblique VAV - ventral series between pelvic-fin base and anal-fin origin
denticles
absent
abbreviated terminology of photophores
terminal bulb
filament
Habitat, biology, and fisheries: Mainly mesopelagic (to 1 000 m) as adults, with some species
bathypelagic, to 2 000 m. Migration to near surface at night in some species. Diet consists of other
midwater fishes.
hexagonal pigmentation
Similar families occurring in the area pattern absent dorsal
adipose fin
Astronesthidae: body fusiform and slightly
compressed; chin barbel present, terminal bulb
present or absent; dorsal fin near middle of body, its
origin well ahead of anal-fin origin; dorsal adipose
fin present; no hexagonal pigment areas on body;
anterior portion of ventral photophore row (PV)
curving upward at pelvic-fin base.
Astronesthidae
1908 Bony Fishes
Chauliodontidae: only other stomiiform family with scale-like areas on body delineated by hexagonal
pigmentation pattern; chin barbel short and simple, becoming reduced or absent in adults; dorsal fin well
anterior, near head, first ray prolonged.
Idiacanthidae: body highly elongate, eel-like; dorsal fin with very long base (54 to 74 rays), its origin well
anterior to midbody; bases of dorsal- and anal-fin rays with a small sharp spur; pectoral fins present in
larvae, absent in adults; pelvic fins absent in males; hexagonal pigmentation pattern absent. 54-74
pectoral fins absent hexagonal pigmentation dorsal-fin
st dorsal in adults pattern absent rays
prolonged 1 adipose fin
dorsal-fin ray present
Malacosteidae: no membrane in floor of mouth; chin barbel present or absent; presence of a preorbital
photophore (absent in Photostomias); pectoral fins absent or consisting of only free, filament-like rays;
hexagonal pigmentation pattern absent.
Melanostomiidae: maxillae with erect teeth anteriorly and small, oblique denticles posteriorly; chin barbel
present, variable in length; pectoral fins present or absent; pelvic fins present; hexagonal pigmentation
pattern absent. posterior upper jaw teeth as hexagonal pigmentation
preorbital light hexagonal pigmentation pattern absent small, oblique denticles pattern absent
organ present
no membrane in
floor of mouth Malacosteidae Melanostomiidae
Gonostomatidae, Phosichthyidae, and Sternoptychidae: these remaining stomiiform families often with
somewhat similar body form and photophore arrangement, but have gill rakers as adults and lack the
hexagonal pigmentation pattern, a chin barbel and enlarged, fang-like jaw teeth.
Myctophidae and Neoscopelidae (myctophiform families): also with photophores, but with a less elongate
body, lack a chin barbel at all stages, have gill rakers as adults, and the maxilla is completely excluded from
the gape by the premaxilla; ventral photophores usually more widely spaced and not arranged in such
regular rows as in the Stomiidae.
jaw teeth not enlarged dorsal adipose fin hexagonal pigmentation
and fang-like usually present pattern present dorsal adipose fin present
maxilla
hexagonal pigmentation
chin barbel absent pattern absent
Phosichthyidae premaxilla chin barbel
always absent
List of species occurring in the area Neoscopelidae
Stomias affinis Günther, 1887
Stomias boa (Risso, 1810)
Stomias danae Ege, 1933
Stomias longibarbatus (Brauer, 1902)
Stomias nebulosus Alcock, 1889
Reference
Paxton, J.R. and J.E. Hanley. 1989. Stomiidae. In Zoological catalogue of Australia 7. Pisces: Petromyzontidae to
Carangidae, edited by J.R. Paxton, D.F. Hoese, G.R. Allen, and J.E. Hanley, pp. 206-207.
Stomiiformes: Chauliodontidae 1909
D iagnostic characters: Body long, slender (to about 30 cm standard length), covered by a gelatinous
membrane (usually lost during capture) with luminous inclusions, maximum depth of body at back
of head. Head short and about as deep as long. Eye diameter about four times into head length. Snout
short, its length less than eye diameter. Mouth large with numerous very large teeth on premaxillae and
dentary, some greatly elongated, fang-like and extending over front of head to above eye when
mouth is closed; posterior half of maxillae with numerous minute teeth. Vomerine teeth absent; palatine
teeth present. Chin barbel short and simple, becoming reduced or absent during development.
Branchiostegal rays 12 to 21. Gill rakers rudimentary, represented by tooth plates only. Dorsal fin near
head, its origin about half way between origin of pectoral and anal fins. First dorsal-fin ray greatly
prolonged. Anal fin located posteriorly, near caudal fin. Dorsal fin with 5 to 7 rays; anal fin with 10 to 13
rays; caudal fin forked; pectoral-fin rays 9 to 14; pelvic fin with 6 to 8 rays. Dorsal and ventral adipose fins
present. Five longitudinal rows of scales covering body, delineated by a hexagonal pigmentation pattern.
Two ventrolateral rows of photophores on body; OA 39-50; IV 25-34; VAV 22-30; AC 8-13; row of
photophores on isthmus (IP); 1 or more small photophores associated with each hexagonal area on body;
suborbital and postorbital photophores present. Premaxillae not protractile; epioccipitals well developed
and lateral to supraoccipital; parietals minute. Anterior vertebrae unossified. Three pectoral-fin radials.
Colour: body scale rows delineated by a hexagonal pigmentation pattern.
dorsal fin far forward,
with prolonged 1st ray
hexagonal
suborbital pigmentation pattern
photophore
postorbital dorsal adipose fin
photophore OA present
chin barbel IP PV VA AC
reduced or
absent IV
AC - ventral series posterior to anal-fin origin OA - lateral series
IP - ventral series anterior to pectoral-fin base PV - ventral series between bases of pectoral and pelvic fins
IV - ventral series anterior to pelvic-fin base VAV - ventral series between pelvic-fin base and origin of anal fin
abbreviated terminology of photophores
Habitat, biology, and fisheries: Meso- to bathypelagic, to a maximum depth of 2 800 m, in open ocean;
juveniles undergo vertical migration to near surface at night. Diet consists of other fishes and crustaceans.
Similar families occurring in the area
Stomiidae: only other stomiiform family with hexagonal pigmentation pattern; body more elongate and
slender; chin barbel present, with terminal bulb and filaments; dorsal fin located posteriorly, just ahead of
caudal fin; dorsal adipose fin absent.
Astronesthidae: body fusiform and slightly compressed; chin barbel present, terminal bulb present or
absent; dorsal fin near middle of body, its origin well ahead of anal-fin origin; no hexagonal pigment areas
on body; anterior portion of ventral photophore row (PV) curving upward at pelvic-fin base.
hexagonal pigmentation
no dorsal adipose fin pattern absent
Idiacanthidae: body highly elongate, eel-like; hexagonal pigmentation pattern not present; chin barbel
well-developed; dorsal fin with very long base (54 to 74 rays); bases of dorsal- and anal-fin rays with a
small sharp spur; dorsal adipose fin absent; pectoral fins present in larvae, absent in adults; pelvic fins
absent in males.
Malacosteidae: hexagonal pigmentation pattern not present on body; no membrane in floor of mouth; chin
barbel present or absent; presence of a preorbital light organ, associated with suborbital organ (absent in
Photostomias; dorsal fin located posteriorly, just ahead of caudal fin; dorsal adipose fin absent; pectoral
fins absent or consisting of only free, filament-like rays. dorsal adipose
pectoral fins absent in adults 54-74 dorsal-fin rays preorbital light organ fin absent
dorsal
adipose
fin
absent
no membrane in
floor of mouth
Idiacanthidae Malacosteidae
Melanostomiidae: hexagonal pigmentation pattern not present on body; maxillae with erect teeth
anteriorly and small, oblique denticles posteriorly; chin barbel present, variable in length; pectoral fins
present or absent.
Gonostomatidae, Phosichthyidae, and Sternoptychidae: these remaining stomiiform families often with
somewhat similar body form and photophore arrangement, but have gill rakers as adults and lack the
hexagonal pigmentation pattern, a chin barbel and enlarged, fang-like jaw teeth.
hexagonal pigmentation pattern absent dorsal adipose fin absent 1st dorsal-fin ray
not prolonged
chin barbel
always
Melanostomiidae absent
Phosichthyidae
References
Fujii, E. 1984. Family Chauliodontidae. In Fishes of the Japanese Archipelago, edited by H. Masuda, K. Amaoka,
C. Araga, T. Uyeno, and T. Yoshino. Tokyo, Tokai University Press, pp. 48-49.
Paxton, J.R. and J.E. Hanley. 1989. Chauliodontidae. In Zoological catalogue of Australia 7. Pisces: Petromyzontidae
to Carangidae, edited by J.R. Paxton, D.F. Hoese, G.R. Allen, and J.E. Hanley, p. 205.
Stomiiformes: Melanostomiidae 1911
D iagnostic characters: Body elongate, slender (to about 50 cm standard length), slightly compressed
(except Bathophilus in which the body is short and highly compressed). Head quite small. Jaws large,
about equal to length of head; membrane connecting dentary bones forms floor of mouth. Maxillae with
erect teeth anteriorly and small, oblique denticles posteriorly. Vomerine and palatine teeth present or
absent. Chin barbel present; ranging widely in length, with or without branches or terminal
elaborations; sexually dimorphic in some taxa. Branchiostegal rays 8 to 22. Gill rakers rudimentary,
arches with series of tooth plates only. Dorsal fin located well posteriorly just anterior to caudal fin,
directly above anal fin. Dorsal fin with 9 to 30 rays; anal fin with 9 to 46 rays; caudal fin small and forked;
pectoral fins present or absent, 0 to 47 rays; pelvic fins present, usually with 7 rays, range 4 to 26. Dorsal
adipose fin absent (except in Chirostomias, which does not occur in the area). Scales absent, no
hexagonal areas on body. Two ventrolateral rows of prominent photophores on body (except in some
Bathophilus species); OA 18-72; IV 16-62; VAV 11-24; AC 5-25; paired row of photophores on isthmus
(IP); many small photophores covering much of body and head, often occurring in vertical rows associated
with segments of body musculature; single row of photophores on caudal peduncle; preorbital photophore
absent, except Pachystomias; suborbital photophore absent; postorbital photophore usually
present, located posteroventrally to eye, sometimes reduced or absent in females. Anterior
vertebrae at least slightly modified, allowing greater movement of head; highly reduced in some genera.
Stomach long, moderately distensible, usually pigmented; typically 2 pyloric caecae present. Colour: skin
usually black, sometimes iridescent silver, bronze, or green.
OA dorsal adipose fin
suborbital photophore absent absent
AC
IV
VAV
Habitat, biology, and fisheries: Mainly mesopelagic, occurring in the upper 1 000 m as adults, some
species undergoing vertical migration to near surface at night. Specimens have been caught by open nets
to depths of up to 4 500 m. Some species apparently benthopelagic as adults. Diet consists of other
mesopelagic fishes and some crustaceans. suborbital photophore
dorsal adipose fin
present
present
Similar families occurring in the area
Astronesthidae: dorsal fin near middle of body, its
origin well ahead of anal-fin origin; dorsal adipose
fin present; anterior portion of ventral photophore
row (PV) curving upward at pelvic-fin base;
suborbital photophore present.
Astronesthidae
1912 Bony Fishes
Chauliodontidae: chin barbel short and simple, becoming reduced or absent in adults; dorsal fin well
anterior, near head, anterior ray prolonged; dorsal adipose fin present; pectoral and pelvic fins present;
scale-like areas on body delineated by hexagonal pigmentation pattern.
Idiacanthidae: body highly elongate, eel-like; dorsal fin with very long base (54 to 74 rays), its origin well
anterior to anal-fin origin; bases of dorsal- and anal-fin rays with a small sharp spur; pectoral fins present
in larvae, absent in adults; pelvic fins absent in males.
pectoral fins absent in adults 54-74 dorsal-fin rays
dorsal adipose
fin present
hexagonal
pigmentation pattern
Malacosteidae: no membrane in floor of mouth; chin barbel present or absent; presence of a preorbital
photophore (absent in Photostomias, present in the melanostomiid Pachystomias); pectoral fins absent or
consisting of only free, filament-like rays.
Stomiidae: body elongate and slender; chin barbel present, with terminal bulb and filaments; posterior
maxillary teeth not consisting of a series of small, oblique denticles; scale-like areas on body delineated by
hexagonal pigmentation pattern; pectoral fins present.
preorbital light organ present hexagonal pigmentation
pattern
Gonostomatidae, Phosichthyidae, and Sternoptychidae: these remaining stomiiform families often with
somewhat similar body form and photophore arrangement, but have gill rakers as adults and lack both a
chin barbel and greatly enlarged, fang-like jaw teeth.
Myctophidae and Neoscopelidae (myctophiform families): also with photophores, but with a less elongate
body, lack a chin barbel, have gill rakers as adults, and the maxilla is completely excluded from the gape by
the premaxilla; ventral photophores usually more widely spaced and not arranged in such regular rows as
in the Melanostomiidae.
dorsal adipose fin
usually present
maxilla
Reference
Paxton, J.R. and J.E. Hanley. 1989. Melanostomiidae. In Zoological catalogue of Australia 7. Pisces: Petromyzontidae
to Carangidae, edited by J.R. Paxton, D.F. Hoese, G.R. Allen, and J.E. Hanley, pp. 197-203.
1914 Bony Fishes
D iagnostic characters: Body markedly elongate, eel-like, slightly compressed (females to about
48 cm standard length). Head small. Snout equal to or less than orbit diameter. Mouth large in adult
females, with numerous barbed, hinged, fang-like teeth, variable in size; jaw teeth absent in males; few teeth
present on vomer and palatine. Chin barbel length about twice that of head in females, absent in males.
Gill arches without rakers or teeth. Dorsal fin with very long base, its origin well anterior to
midbody. Anal-fin base length about 1/2 of dorsal-fin base, its origin below middle rays of dorsal fin. Dorsal
and anal fins terminating posteriorly on constricted portion of caudal peduncle. Base of each dorsal and
anal-fin ray with a small, sharp spur. Dorsal fin with 54 to 74 rays; anal fin with 29 to 49 rays; caudal fin
forked; pectoral fins present in larvae, absent in adults; pelvic fins present in females, with 6 rays,
absent in males. Dorsal adipose fin absent. Scales absent. Two main rows of photophores on body
ventrolaterally (see Astronesthidae family figure, page 1905, for definition of abbreviated terms); OA 52-61; IV
31-36; VAV 15-18; AC 13-18; paired row of photophores on isthmus (IP); small light organs scattered over head
and body in patterns on each body segment; suborbital photophore absent; postorbital photophore
prominent, located posteroventrally, equal in size to eye in males, smaller in females. Premaxillae
not protractile. Mesethmoid bilobate anteriorly, with prominent lateral processes proximal to nasal sacs.
Parietals absent. Gasbladder absent. Colour: skin usually black in females, males dark brown; hexagonal
pigment areas lacking.
suborbital photophore postorbital pectoral fin absent in adults 54-74 dorsal-fin rays
absent photophore
chin barbel
female
Habitat, biology, and fisheries: Meso- to bathypelagic, recorded to a depth of 2 000 m. Diet consists
mainly of fishes. Markedly sexually dimorphic; males reaching about 15% of female body size, retaining
some larval features; anterior anal-fin rays modified as intromittent organ. Eyes on the ends of long stalks
in larvae.
Similar families occurring in the area
Stomiidae: body elongate and slender but not eel-like; chin barbel present, with terminal bulb and
filaments; scale-like areas on body delineated by hexagonal pigmentation pattern; dorsal fin short-based
(13 to 23 rays), located posteriorly, just ahead of caudal fin; dorsal and anal-fin rays without basal spur;
pectoral and pelvic fins present.
Astronesthidae: body fusiform and slightly compressed; chin barbel present in males and females;
dorsal-fin short-based (9 to 21 rays), near middle of body, rays without basal spur; dorsal adipose fin
present; anterior portion of ventral photophore row (PV) curving upward at pelvic-fin base.
hexagonal pigmentation 9-12 dorsal-fin rays
pattern 13-23 dorsal-fin rays dorsal adipose
fin present
no membrane in
floor of mouth Malacosteidae Melanostomiidae
Gonostomatidae, Phosichthyidae, and Sternoptychidae: these remaining stomiiform families with
somewhat similar photophore arrangement, but have gill rakers as adults, a relatively short dorsal fin, and
lack both a chin barbel and enlarged, fang-like jaw teeth; only 1 genus, the monotypic Triplophos, has a
highly elongate species.
Myctophidae and Neoscopelidae (myctophiform families): also with photophores, but with a much less
elongate body, lack a chin barbel at all stages, have gill rakers as adults, and the maxilla is completely
excluded from the gape by the premaxilla; ventral photophores are usually more widely spaced and not
arranged in such regular rows as in the Idiacanthidae.
short dorsal-fin base
short dorsal-fin
base
maxilla
premaxilla
chin barbel
chin barbel always absent
absent
Phosichthyidae Neoscopelidae
References
Fujii, E. 1984. Family Idiacanthidae. In Fishes of the Japanese Archipelago, edited by H. Masuda, K. Amaoka, C. Araga,
T. Uyeno, and T. Yoshino. Tokyo, Tokai University Press, pp. 53-54.
Paxton, J.R. and J.E. Hanley. 1989. Idiacanthidae. In Zoological catalogue of Australia 7. Pisces: Petromyzontidae to
Carangidae, edited by J.R. Paxton, D.F. Hoese, G.R. Allen, and J.E. Hanley, p. 208.
1916 Bony Fishes
D iagnostic characters: Body moderately elongate (to about 24 cm standard length), compressed.
Head and eye large relative to body. Snout usually quite short (elongate in Aristomias). Mouth large,
jaws longer than skull, about 15 to 30% standard length; membranes lacking between rami of lower
jaw. Jaw teeth variable in size, some very large and barbed. Vomerine teeth absent; palatine with or
without teeth. Chin barbel present or absent. Branchiostegal rays 9 to 15. Gill arches with teeth reduced
or absent; no well-developed rakers. Dorsal fin located well posteriorly just anterior to caudal fin,
directly above anal fin. Dorsal fin with 14 to 28 rays; anal fin with 17 to 32 rays; caudal fin small, forked;
pectoral fins absent or consisting of 2 to 17 free, filament-like rays; pelvic fins at about midbody, with 5
to 9 rays. Dorsal and ventral adipose fins absent. Scales absent, no hexagonal areas on body. Two
ventrolateral rows of photophores on body (poorly developed and inconspicuous in Malacosteus); OA
7-39; IC 12-22; paired row of photophores on isthmus (IP); many small light organs covering much of body
and head; preorbital light organ present or absent; suborbital photophore present, varying in size from
minute to very large, or absent; postorbital photophore prominent, located posteroventrally to eye.
Premaxillae not protractile. Parietals and posttemporals absent. First few vertebrae unossified. Stomach
distensible, pigmented. Pyloric caecae present or absent. Colour: skin black to dark brown.
postorbital photophore
OA dorsal
adipose fin
absent
preorbital
light VA
organ
AC
IP PV
no membrane in
floor of mouth
IV
AC - ventral series posterior to anal-fin origin
Habitat, biology, and fisheries: Meso- to IP - ventral series anterior to pectoral-fin base
bathypelagic adults, at depths to 4 000 m. Diet IV - ventral series anterior to pelvic-fin base
consists of fishes and crustaceans. Floor of OA - lateral series
lower jaw lacks membranes, which allows the PV - ventral series between bases of pectoral and pelvic fins
jaws to swing widely while feeding, hence the VAV - ventral series between pelvic-fin base and anal-fin origin
common name “loosejaws”.
abbreviated terminology of photophores
Similar families occurring in the area
Astronesthidae: membrane present in floor of mouth; chin barbel present, terminal bulb present or absent;
dorsal-fin near middle of body, its origin well ahead of anal-fin origin; dorsal adipose fin present; anterior
portion of ventral photophore row (PV) curving upward at pelvic-fin base.
Chauliodontidae: membrane present in floor of mouth; body elongate, maximum depth at back of head;
chin barbel short and simple, becoming reduced or absent in adults; dorsal fin well anterior, between
vertical through pectoral and pelvic fins; pectoral and pelvic fins present; scale-like areas on body
delineated by hexagonal pigmentation pattern.
hexagonal dorsal
dorsal adipose pigmentation adipose fin
fin present pattern present
Astronesthidae Chauliodontidae
Stomiiformes: Malacosteidae 1917
Idiacanthidae: body highly elongate, eel-like; membrane present in floor of mouth; dorsal fin with very
long base (54 to 74 rays), its origin well anterior to midbody; bases of dorsal- and anal-fin rays with a small
sharp spur; pectoral fins present in larvae, absent in adults; pelvic fins absent in males.
Melanostomiidae: membrane present in floor of mouth; maxillae with erect teeth anteriorly and small,
oblique denticles posteriorly; chin barbel present, variable in length; pectoral fins present or absent; pelvic fins
present.
preorbital light
pectoral fins absent in adults 54-74 dorsal-fin rays organ absent
membrane
present in floor
Idiacanthidae Melanostomiidae
Stomiidae: body elongate and slender; chin barbel present, with terminal bulb and filaments; membrane
present in floor of mouth; scale-like areas on body delineated by hexagonal pigmentation pattern; pectoral
and pelvic fins present.
Gonostomatidae, Phosichthyidae, and Sternoptychidae: these remaining stomiiform families often with
somewhat similar body form and photophore arrangement, but all have membrane present in the floor of
mouth and gill rakers as adults, and lack both a chin barbel and greatly enlarged, fang-like jaw teeth.
preorbital light
organ absent hexagonal pigmentation pattern preorbital light organ absent dorsal adipose fin
usually present
References
Fujii, E. 1984. Family Malcosteidae. In Fishes of the Japanese Archipelago, edited by H. Masuda, K. Amaoka, C. Araga,
T. Uyeno, and T. Yoshino. Tokyo, Tokai University Press, p. 53.
Paxton, J.R. and J.E. Hanley. 1989. Malacosteidae. In Zoological catalogue of Australia 7. Pisces: Petromyzontidae to
Carangidae, edited by J.R. Paxton, D.F. Hoese, G.R. Allen, and J.E. Hanley, p. 204.
1918 Bony Fishes
Order ATELEOPODIFORMES
ATELEOPODIDAE
Jellynoses (tadpole fishes)
by J.A. Moore
D iagnostic characters: Elongate fishes (to 2 m) with soft, flabby body; trunk short; tail elongate,
compressed and tapering. Head moderate sized; eyes small to moderate; snout pointed or
rounded, very soft and gelatinous; mouth subterminal and protrusible; broad band of small villiform
teeth present or absent on premaxilla and mandible. Branchiostegals 7. No fin spines; short-based
single dorsal fin just behind head with 3 to 12 rays; anal fin elongate and united with caudal fin, 85 to
120 rays total; pectoral-fin rays 12 to 14; pelvic fins jugular, 2 to 4 rays with single long ray in adults.
Scales only present imbedded in lateral-line canal. Colour: light brown to dark purplish brown.
no spines in fins
Habitat, biology, and fisheries: Uncommon benthic or benthopelagic fishes usually taken between 200
and 800 m. Little known of general biology. Some species feed on benthic invertebrates. No commercial
importance.
2 dorsal fins
Similar families occurring in the area
Macrouridae: chin barbel usually present; often
spinous rays in first dorsal fin; long, low second
dorsal fin behind first dorsal fin; scales cover portions
of body and head.
chin
barbel
List of species occurring in the area Macrouridae
Ateleopus indicus Alcock, 1891
Ateleopus japonicus Bleeker,1853
Ateleopus sp.
Ijimaia sp.
Parateleopus microstomus Smith and Radcliffe, 1912
Reference
Smith, M.M. 1986. Family Ateleopodidae. In Smiths’ sea fishes, edited by M.M. Smith and P.C. Heemstra.
Johannesburg, Macmillan South Africa, pp. 404-406.