Apogonidae Fishes LRs

CardinalfishesCover_CardinalfishesCover 8/09/19 7:17 PM Page 1
Cardinalfishes of the World – Kuiter & Kozawa

Cardinalfishes of the World
Rudie H Kuiter & Toshikazu Kozawa
This book is an illustrated identification guide to the
Cardinalfishes of the family Apogonidae. Since our earlier
publication (1999) of: Fishes of the Indo-West Pacific,
APOGONIDAE many new species were discovered and much
has changed in their classification. The Atlantic taxa were
added and about 350 species of these ray-finned fishes are
now recognised globally. All are illustrated, except for
some small taxa for which images were not available.
Cardinalfishes vary much in colour and patterns, many are
striped and can look very similar to each other, but in the
Atlantic the majority of species are red in colour, hence
their vernacular name. Large genera comprise a number of
species-complexes in which taxa are difficult to distinguish,
often only by their different colour patterns. The majority
of species are illustrated in living colour and in situ, fish
captured were photographed soonest after, showing fresh
Cardinalfishes
colours, whilst some are presented with original drawings
of the types. Cardinalfishes are in principal marine, but
some are found in brackish water and one genus has fully
adapted to fresh water. They are mostly small percoid fishes
that range in size from about 6 to 20 cm, but a few are
smaller or over 20 cm long. The majority occur on reefs at
of the World
relatively shallow depths, whilst some taxa are known only
from deep trawls. Cardinalfishes are one of a few marinefish
families in which oral brooding of eggs takes place. When
spawning, eggs are released by the female in a gelatinous
mass, which is immediately fertilised by the male and
quickly taken into the mouth. Brooding males are readily
recognised by a deeply expanded mouth and developing
eggs can be seen through the skin.
Anthis – Aquatic Photographics

AQUATIC
PHOTOGRAPHICS
Rudie H
Rudie H Kuiter
Kuiter &
& Toshikazu
Toshikazu Kozawa
Kozawa
CardinalfishesCover_CardinalfishesCover 8/09/19 7:17 PM Page 1
Cardinalfishes of the World – Kuiter & Kozawa

Cardinalfishes of the World
Rudie H Kuiter & Toshikazu Kozawa
This book is an illustrated identification guide to the
Cardinalfishes of the family Apogonidae. Since our earlier
publication (1999) of: Fishes of the Indo-West Pacific,
APOGONIDAE many new species were discovered and much
has changed in their classification. The Atlantic taxa were
added and about 350 species of these ray-finned fishes are
now recognised globally. All are illustrated, except for
some small taxa for which images were not available.
Cardinalfishes vary much in colour and patterns, many are
striped and can look very similar to each other, but in the
Atlantic the majority of species are red in colour, hence
their vernacular name. Large genera comprise a number of
species-complexes in which taxa are difficult to distinguish,
often only by their different colour patterns. The majority
of species are illustrated in living colour and in situ, fish
captured were photographed soonest after, showing fresh
Cardinalfishes
colours, whilst some are presented with original drawings
of the types. Cardinalfishes are in principal marine, but
some are found in brackish water and one genus has fully
adapted to fresh water. They are mostly small percoid fishes
that range in size from about 6 to 20 cm, but a few are
smaller or over 20 cm long. The majority occur on reefs at
of the World
relatively shallow depths, whilst some taxa are known only
from deep trawls. Cardinalfishes are one of a few marinefish
families in which oral brooding of eggs takes place. When
spawning, eggs are released by the female in a gelatinous
mass, which is immediately fertilised by the male and
quickly taken into the mouth. Brooding males are readily
recognised by a deeply expanded mouth and developing
eggs can be seen through the skin.
Anthis – Aquatic Photographics

AQUATIC
PHOTOGRAPHICS
Rudie H
Rudie H Kuiter
Kuiter &
& Toshikazu
Toshikazu Kozawa
Kozawa
ApogonidaeFishes1-21_ApogonidaeFishes1-21 8/09/19 5:18 PM Page I
Cardinalfishes
of the World
Rudie H kuiter & Toshikazu Kozawa

ApogonidaeFishes1-21_ApogonidaeFishes1-21 8/09/19 5:18 PM Page II
Contents
Foreword ..................................................................................1
FAMILY APOGONIDAE ....................................................................1
Key to subfamilies of Apogonidae................................................2
Key to the genera of Apogonidae ................................................2
SUBFAMILY AMIOIDINAE (2 genera) ..................................................4
Amioides Smith & Radcliffe in Radcliffe, 1912......................4
Holapogon Fraser, 1973 .....................................................4
SUBFAMILY APOGONINAE
Tribe Apogonichthyini ..............................................................5
Apogonichthys Bleeker, 1854..............................................6
Foa Jordan & Evermann in Jordan & Seale, 1905 ................8
Fowleria Jordan & Evermann, 1903 ..................................12
Neamia Smith & Radcliffe, 1912 .......................................16
Ozichthys Fraser, 2014 .....................................................17
Vincentia Castelnau, 1872 ................................................18
Tribe Apogonini.....................................................................22
Apogon Lacepède, 1801 ...................................................22
T. Hirata Zapogon Fraser, 1972 .......................................................38
Astrapogon Fowler, 1907..................................................41
Paroncheilus Smith, 1964.................................................53
Phaeoptyx Fraser & Robins, 1970 .....................................43
Tribe Archamiini ....................................................................44
Archamia Gill, 1863 .........................................................44
Cardinalfishes
Taeniamia Fraser, 2013 ....................................................45
of the World Tribe Cheilodipterini...............................................................52
New edition 2019 Cheilodipterus Lacepède, 1801.........................................52
Tribe Glossamiini ...................................................................66
©1999, ©2019 Glossamia Gill, 1863 ........................................................66
Rudie H Kuiter & Toshikazu Kozawa. Yarica Whitley, 1930 ........................................................70
Tribe Gymnapogonini.............................................................71
There is no objection to copying Cercamia Randall & Smith, 1988 ......................................71
parts of this book for personal use. Gymnapogon Regan, 1905 ...............................................74
Please check with authors for other Pseudamiops Smith, 1954 ................................................77
uses – by email: Lachneratus Fraser & Struhsaker, 1991 .............................78
rudiekuiter@optusnet.com.au Tribe Lepidamiini ...................................................................79
nekomimi29q@icloud.com Lepidamia Gill, 1863 ........................................................79
Tribe Ostorhinchini ...............................................................82
Publishers Ostorhinchus Lacepède, 1802...........................................82
Aquatic Photographics Tribe Pristiapogonini ............................................................129
Pristiapogon Klunzinger, 1870 ........................................129
PO Box 124
Pristicon Fraser, 1972 ....................................................135
Seaford, 3198
Tribe Rhabdamiini ...............................................................137
Victoria, Australia
Rhabdamia Weber, 1909 ................................................137
& Tribe Veruluxini ...................................................................140
Anthis (Nexus) Verulux Fraser, 1972 ......................................................140
1-16-1 Yabuta Tribe Siphamiini ..................................................................141
Okazaki, Aichi, 444-2137 Siphamia Weber, 1909 ...................................................141
Japan Tribe Sphaeramiini ..............................................................155
Apogonichthyoides Smith, 1949 .....................................155
Jaydia Smith, 1961 ........................................................165
Nectamia Jordan, 1917 ..................................................170
Xeniamia Jordan, 1917 ..................................................177
Pterapogon Koumans, 1933 ...........................................178
Quinca Mees, 1966 ........................................................180
Sphaeramia Fowler & Bean, 1930 ..................................180
Tribe Zoramiini....................................................................182
Fibramia Fraser & Mabuchi, 2014 ...................................182
AQUATIC Zoramia Jordan, 1917 ....................................................184
SUBFAMILY PAXTONINAE
PHOTOGRAPHICS Paxton Baldwin & Johnson, 1999 ...................................186
Subfamily PSEUDAMIINAE
Pseudamia Bleeker, 1865 ...............................................187
Acknowledgements & list of photographers ............................192
References & Literature Cited or Used ...................................193
Index ..................................................................................195
II Cardinalfishes of the World – Kuiter & Kozawa

ApogonidaeFishes1-21_ApogonidaeFishes1-21 8/09/19 5:18 PM Page 1
Foreword
About the book
A book on this large group of perciform fishes commonly known as cardinalfishes was first published by the
authors in 1999 on CD and in 2001 as a digital on-demand printed hard copy. Since then the classification
of the Apogonidae family has advanced greatly whilst in addition many new discoveries have been made and
this publication is a taxonomically updated and greatly expanded pictorial guide. The scope of illustrating the
species in living colours and in situ remains the same, with images of freshly caught specimens as an option.
We have attempted to illustrate every species with multiple images from the different areas around the world
with the cooperation of the many underwater photographers familiar with these fishes. To ensure the use of
the correct name of the particular species, we made special efforts to consult original descriptions and to
photograph the species from or near the type-localities. For some particular species we have travelled to the
type-localities or the nearest areas that represents the same kind of fauna to secure photographs. The work
is based on detailed studies of the Apogonidae with the advice of recognised experts of various Apogonidae
groups and completion of this new edition was only made possible with photographs generously supplied by
more than 100 divers, ichthyologists and collectors from around the world. Taxonomic problems remain
among several tribes and there is uncertainty of some species in their classification. As taxonomic decisions
in recent works are not always agreed with, names of a some species used by us may differ from the those
of the reviewers and placement to genus needs to be verified for some. Final taxonomic decisions in this
book are made by the authors.
We have followed the most recent classification based on molecular analysis presented by currently recognised
“world experts” on the Apogonidae (Mabuchi et al, 2014), and used associated taxa data presented in the
Catalog of Fishes. Subfamilies, tribes and genera are largely presented in alphabetic order, but species in an
order that accommodates the similar taxa together or in groups, such as species-complexes. Undetermined
and undescribed species are included in the generic sections. Each genus has a brief introduction that gives
the type species, author and date, the name gender, the number of species and provides a basic description,
diagnostics and general information known to date. Details for each species includes present scientific and
vernacular name, original name as described, author and date, their distribution and general information,
including useful comments on similar species or synonymy. Images are captioned with scientific name,
photo-locality, and credited with the photographers initials that are listed in alphabetic order following
acknowledgements. A full index to scientific and common names is provided. References given in the text to
books or papers are listed under References & Literature Cited or Used. For complete taxa citations see:
Catalog of Fishes online (http://research.calacademy.org/research/ichthyology/catalog/fishcatmain.asp).
Family APOGONIDAE
The Apogonidae family with 40 genera comprises 4 subfamilies of which Apogoninae is the largest by far,
divided into many tribes, in 36 genera and has almost 350 described species worldwide (Mabuchi et al,
2014), but many more taxa are known that are undescribed. The members are variously coloured, spotted,
striped or banded in the Indo-West Pacific, whilst in the Atlantic most Apogon taxa are bright red in colour,
hence their common name cardinalfishes. The other subfamilies are small with one or two genera.
Cardinalfishes feature an oval to elongate compressed scaly body, large mouth, have large prominent eyes,
and two dorsal fins, that are short-based often tall that are distinctly separate, with the exception of the
monotypic Paxtoninae, which has a single dorsal-fin. They are a diverse group of perciform, generally
inshore marine reef-dwelling fishes in tropical waters. Most genera members inhabit shallow environments,
live on or near coral and rocky reefs, but some live on muddy substrates, or enter brackish and freshwater.
Only those of the genus Glossamia are fully adapted to freshwater. Vincentia members are endemic to
Australia’s southern temperate waters and a few other genera range to warm-temperate zones or live in deep
water. Most deepwater dwellers are not well known, some taxa from single specimens only.
Many species are secretive, hiding among corals and in caves, crevices, and remain under ledges during the
day and are active at night, but some are opportunistic feeders, school at any time of the day and are active
depending on tidal currents that carry zoo-plankton. Most feeding activities are at night, but some taxa that
are involved in some kind of mimicry with daytime-active fishes are diurnal, swimming openly about over
open substrates. A few of the taxa form large schools during the day. Some species have bioluminescent
organs, producing light either with or without symbiotic bacteria. All known species are reputed to be oral
brooding and their dispersal is limited to the level of development of fry. A few taxa nurse the hatchlings in
the mouth that settle on the substrate and these are locally endemic, whilst species with a minimal hatchling
development have a long pelagic stage and are widespread by oceanic currents.
Cardinalfishes of the World – Kuiter & Kozawa 1
Key to subfamilies of APOGONIDAE (after Mabuchi et al, 2014)

1 Two dorsal fins, I,8–13 in second dorsal fin; first dorsal spines all with uneven lengths ............................................2
– Single dorsal fin, VI,19; spines III–VI similar lengths ...............................................................................Paxtoninae
2 Supramaxilla small or absent, if large, dorsal fin VI–I,9–10 ....................................................................................3
– Supramaxilla large, dorsal fin VII or VIII–I,9–10 .....................................................................................Amioidinae
3 Lateral line single when scales present or absent; if scales absent, lateral line composed of free neuromasts .....Apogoninae
– Lateral lines double, first pored or notched from posttemporal, second abdominal with only notched scales ....Pseudamiinae
Key to the genera of Apogonidae (after Mabuchi et al, 2014)

Note this key uses external characters where possible to afford more use.
1 No pored lateral-line scales or scales absent on body. ............................................................................................2
– Some pored lateral-line scales on body..................................................................................................................3
2 Single dorsal fin, six spines [Paxtoninae] .....................................................................................................Paxton
– Two dorsal fins, seven spines (Gymnapogonini) . ..................................................................................................4
3 One pored lateral line, sometimes partially pored followed by pits or grooves . ........................................................7
– Two inconspicuous lateral-line scales, one dorsal (pored then notched), one ventral (notched) [Pseudamiinae] .......Pseudamia
4 Scales present ....................................................................................................................................................5
– Scales absent ...................................................................................................................................Gymnapogon
5 Second dorsal fin I,8–9 .......................................................................................................................................6
– Second dorsal fin I,12–13 ..................................................................................................................Lachneratus
6 Anal rays 11–13 ....................................................................................................................................Cercamia
– Anal rays 8 or 9 ...............................................................................................................................Pseudamiops
7 Silver or blackish band (bioluminous in life) along ventral side from hyal region extending past abdomen onto caudal
peduncle (Siphamiini). ............................................................................................................................Siphamia
– No such bioluminous bands ................................................................................................................................8
8 Longest procurrent caudal-fin rays segmented ....................................................................................................10
– Longest procurrent caudal-fin rays spinous, not segmented ...................................................................................9
9 Dorsal-fin rays 9; anal-fin rays 9 ..........................................................................................................Sphaeramia
– Dorsal-fin rays 13–15; anal-fin rays 12–14 ...........................................................................................Pterapogon
10 Dorsal-fin rays 8–13 .........................................................................................................................................11
– Dorsal-fin rays 14–15; anal-fin rays 13–14 ..................................................................................................Quinca
11 Lateral-line scales less than 29 ..........................................................................................................................12
– Lateral-line scales greater than 32 (Lepidamiini) ......................................................................................Lepidamia
12 Preopercular ventral edge ossified, serrated, crenulated or smooth .......................................................................13
– Preopercular ventral edge with unossified flap (Apogonini) ...................................................................................15
13 First three infraorbitals with upper edges smooth to crenulated ............................................................................19
– First three infraorbitals with upper edges strongly serrated (Pristiapogonini) ..........................................................14
14 Dorsal fin VI–I,9; Dark spot on body below lateral line under first dorsal fin; dark spot(s) under posterior base of second
dorsal fin; No stripe from snout onto opercle through eye ..........................................................................Pristicon
– Dorsal fin VII–I,9; No spots on body below dorsal fins; Stripe from snout onto opercle through eye, may continue on body
..............................................................................................................................................................Pristiapogon
15 Predorsal scaled ...............................................................................................................................................16
– Predorsal without scales along the center line to origin of first dorsal fin .................................................Astrapogon
16 Anal-fin rays 8 .................................................................................................................................................17
– Anal-fin rays 9 ..................................................................................................................................Paroncheilus
17 Stomach and intestine pale ...............................................................................................................................18
– Stomach and intestine black .....................................................................................................................Zapogon
18 Preopercular flap not extends past vertical edge ..........................................................................................Apogon
– Preopercular flap extends past vertical edge ...........................................................................................Phaeoptyx
19 Supramaxilla large, easy to detect ......................................................................................................................20
– Supramaxilla small difficult to detect or absent ....................................................................................................23
20 First dorsal spines 7 or 8 [Amioidinae] ...............................................................................................................21
– First dorsal spines 6 (Glossamiini) ......................................................................................................................22
21 Anal-fin rays 8; canine teeth; basicaudal bar .............................................................................................Amioides
– Anal-fin rays 7; villiform teeth; spots on body ........................................................................................Holapogon
22 Preopercle edges serrate; only a basicaudal spot ............................................................................................Yarica
– Preopercle edges smooth; body with many markings ...............................................................................Glossamia
23 Anal-fin rays 8–9 ..............................................................................................................................................27
– Anal-fin rays 10–19 .........................................................................................................................................24
24 Edge of preopercle serrated (Archamiini) ............................................................................................................26
2 Cardinalfishes of the World – Kuiter & Kozawa

– Edge of preopercle smooth (Rhabdamiini) ..........................................................................................................25

25 Anal fin rays 10–11; Dorsal fins VII–I,10–11; Some canine teeth .................................Rhabdamia (Bentuviaichthys)
– Anal-fin rays 12–13; Dorsal fins VI–I,9; Villiform teeth ......................................................Rhabdamia (Rhabdamia)
26 Anal-fin rays 15–17; No bars or stripes on head or body; First dorsal-fin spine 1.1–1.4 in second dorsal-fin spine Archamia
– Anal-fin rays 12–19; two yellow bars on head, or 1–23 bars on body, or darkish or yellowish mid-line stripe; First dorsal
spine 1.3–3.4 in second dorsal-fin spine ................................................................................................Taeniamia
27 No canine teeth; some lateral dentary teeth may be slightly enlarged ....................................................................28
– Canine and canoid teeth present (Cheilodipterini) .............................................................................Cheilodipterus
28 First dorsal spines 6–8, if 6 spines then second dorsal-fin rays 8 or anal-fin rays 9 ............................................... 29
– First dorsal spines 6; second dorsal-fin rays 9, anal-fin rays 8 .....................................................................Fibramia
29 First dorsal spines 6–8, if 6 spines then body with one or more stripes extending to caudal fin ...............................32
– First dorsal spines 6; no body stripes .................................................................................................................30
30 Anal-fin rays 9; no cheek mark ..........................................................................................................................31
– Anal-fin rays 8; narrow or broad cheek mark . .........................................................................................Nectamia
31 Preopercle edge serrate; caudal peduncle and/or caudal base with one small dark spot or diffuse large darkish region;
no small dark snout mark .........................................................................................................................Zoramia
– `Preopercle edge smooth; no dark marks on caudal base or caudal peduncle; small dark snout mark ...........................Verulux
32 Stomach and intestine pale; first dorsal spines 7 or 8 ..........................................................................................34
– Stomach and intestine with melanophores to completely blackish; first dorsal spines 7 ..........................................33
33 Fourth dorsal spine longer than third spine; caudal fin emarginate, truncate or rounded ...................................Jaydia
– Third dorsal spine longer than fourth, if fourth longer then caudal fin forked (Ostorhinchini) ..........................Ostorhinchus
34 Edge of preopercle serrated ..............................................................................................................................38
– `Edge of preopercle smooth ...............................................................................................................................35
35 Palatine teeth absent ........................................................................................................................................36
– `Palatine teeth present ......................................................................................................................................Foa
36 Pored lateral-line scales from posttemporal to base of caudal fin ..........................................................................37
– `Pored lateral-line scales usually short, only pits present past dorsal fins ........................................................Fowleria
37 First dorsal fin with 8 spines, or dark mark on opercle if 7 spines ...........................................Neamia and Fowleria*
– `No dark mark on opercle ................................................................................................................Apogonichthys
38 Post-temporal serrate; basisphenoid present ..............................................................................Apogonichthyoides
– `Post-temporal smooth; basisphenoid absent .............................................................................................Vincentia
*Species of Neamia have fused hypurals 2 + 3 and 4 + 5; Species of Fowleria have 5 free hypurals
Not included here, described since the key was published: Ozichthys, which differs from Vincentia in the number of pectoral
fin-rays 17–19 versus 14–16, and Xeniamia atrithorax collected with shrimp trawls from depths of 70–119 m in the China
Sea. It belongs in the Siphamiini and may be closely related to Jaydia.
1st dorsal fin 2nd dorsal fin

spinous 1 spine + soft rays
caudal fin
soft rayed
pectoral fin
soft rayed
anal fin
2 spines + soft rays
ventral fin
(pelvic fin)
1 spine + soft rays
Fin terminology used (Apogon cyanosoma illustration after Bleeker, 1853).

SUBFAMILY AMIOIDINAE
Comprises two monotypic genera: Amioides and Holapogon.
Genus Amioides Smith & Radcliffe, 1912

Fem. Type-species Amia (Amioides) grossidens Smith & Radcliffe, 1912.
The monotypic Amioides is a deep-dwelling (77–267 m) genus known from limited material, but widespread
Indo-Pacific from continental and island locations of East Africa to Japan and Vanuatu (Fraser, 2013).
Diagnosis D. VII–I,9–10; A. II,8; gill rakers 2+9, including rudiments 17–19; one or two pairs of anterior
retrorse canine teeth near symphysis of dentaries on each side, with a row of spaced 2–3 canines and other
slightly enlarged teeth; two large retrorse canine teeth near premaxilla symphysis with band of villiform teeth
on sides; black peritoneum; one or two large prongs on posterior edge of posttemporal with small serrations
along dorsal edge; one supernumerary anal spine; large supramaxilla (after Fraser, 2013).
Deep-dwelling Cardinalfish Amioides polyacanthus

Cheilodipterus polyacanthus Vaillant, 1877. Réunion, western Mascarenes, southwestern Indian Ocean.
Indo-Pacific, known from numerous locations from Indo-West Pacific: Réunion, Mascarenes, Comores, Madagascar and east
to Indonesia and Vanuatu, north to southern Japan, in depths of 77–276 m. Associated with hard bottom, drop-offs, and
caves. Mark Erdmann observed a group of about 10 individuals at 77 meters depth, that were hovering over a big boulder
at Nusa Penida, Bali. Pale brownish silvery, dusky dorsally. Two forward-leaning near-vertical dark lines on head, first below
eye and second over opercle. Axil also dark and a dark band at end of caudal peduncle. A short mid-lateral ragged stripe,
from below first dorsal-fin origin to below end of second dorsal-fin base. Fins yellowish. A large species, length to 25 cm.
A B
Amioides polyacanthus China Seas. ZUK Amioides polyacanthus Taiwan. WCC
C
Amioides polyacanthus Collected from 77 m. Nusa Penida, Bali, Indonesia. MVE

GENUS Holapogon Fraser, 1973

Masc. Type-species Apogon maximus Boulenger, 1888.
The monotypic Holapogon is a deep-dwelling (38–100 m) genus known from limited Indian Ocean material.
Diagnosis D. VIII–I,7, the eighth spine reduced, usually hidden; A. II,7; gill rakers 2+9, including rudiments
17–19; large supramaxilla; villiform teeth on premaxilla, dentary, vomer and palatine, none on ectopterygoid;
preopercular ridge smooth, ventral and posterior edges serrated; infraorbital edges smooth except for one
or two spines on the second bone; one spine on posttemporal; lateral-line complete, scales ctenoid; three
predorsals; two pairs of uroneurals (after Fraser, 1973).
Titan Cardinalfish Holapogon maximus

Apogon maximus Boulenger, 1888. Muscat, Oman, Gulf of Oman, Arabian Sea.
Indian Ocean, known from the coastal regions of Oman and Gulf of Mannar, India, and Andaman Islands, reported from
depths of 38–100 m. Usually trawled in depths of about 80 m or greater. A distinct species by the spotted pattern on the
body, two forward-leaning near-vertical lines over head, first through eye and second from nape over opercle, and adults
with their great body size with large rounded fins. A large species, length to 30 cm.
Holapogon maximus At 38 m. Oman. JER
SUBFAMILY APOGONINAE
SUBFAMILY APOGONINAE comprises 14 tribes:
APOGONICHTHYINI .......5 GLOSSAMIINI .............66 PRISTIAPOGONINI .....129 SPHAERAMIINI .........155
APOGONINI ...............22 GYMNAPOGONINI .......71 RHABDAMIINI ..........137 ZORAMIINI ..............182
ARCHAMIINI ..............44 LEPIDAMIINI ..............79 VERULUXINI............140
CHEILODIPTERINI........52 OSTORHINCHINI .........82 SIPHAMIINI .............141
TRIBE APOGONICHTHYINI
Comprises five genera: Apogonichthys, Foa, Fowleria, Neamia, and Vincentia.
Diagnosis D. VII(I) or VIII+I,7–10; A. II,7–9; head and body with ctenoid scales; pored lateral-line scales,
3–24, scales not pored with groove or pit in scale; preopercle smooth on ridge, serrate or smooth on edges,
where smooth a narrow weakly ossified to unossified flap; three supraneurals; supramaxilla narrow, reduced
or absent; basisphenoid reduced or absent; one pair of uroneurals present; three epurals; five free hypurals
or 1–2 fused and 3–4 fused, one or more fused to terminal centrum; free parhypural; caudal fin emarginate,
truncate or rounded; head and body reddish, brownish or blackish without stripes, often with pale or dark
spots on body (after Mabuchi et al, 2014).

TRIBE APOGONICHTHYINI – GENUS Apogonichthys Bleeker, 1854

Masc. Type-species Apogonichthys perdix Bleeker, 1854.
Five species are recognised: Apogonichthys ahimsa, A. infuscus, A. ocellatus, A. perdix, and A. waikiki,
but some identifications are uncertain. Whitley’s A. ahimsa is an unsolved taxon similar to A. perdix.
Three-spot Cardinalfish
Apogonichthys perdix
Amia perdix Bleeker, 1854. Flores, Indonesia.
Widespread Indo West-Pacific, known from the eastern
Indian Ocean and tropical central West Pacific. Usually
shallow on reefs, secretive in macro-algae and rubble
habitats. D VII, but may have a minute 8th spine, well
hidden under skin. First dorsal fin angular and caudal fin
rounded. Colour tan to reddish with irregular spotting.
Often with 3 small but distinct white spots, one at each
end of dorsal and anal-fin bases, and one basicaudal in
A centre. Length to about 50–60 mm.
Apogonichthys perdix Type, Flores, Indonesia. After Bleeker
B C
Apogonichthys perdix Nha Trang, Vietnam. RWI Apogonichthys perdix Nusa Penida, Bali. GRA
D E
Apogonichthys perdix Taiwan. PCH Apogonichthys waikiki Marquesas Islands. JTW
Gentle Gobbleguts Apogonichthys ahimsa

Apogonichthys ahimsa Whitley, 1959. Heron Island, Qld.
Only known from Heron Island where 10 specimens
were collected, but it is likely to be widespread in
Queensland, and like its close siblings lives secretively in
coral crevices or in shallow macro-algae rubble habitats.
D. VII-I,9; A. II, 8; P. 15; Lat. Line 20; Tr. 1/1/6, and
one predorsal scale (meristics after Whitley). Living
colours are unknown. Largest specimen 32 mm.
Apogonichthys ahimsa appears to be closely related to
A. perdix. The recent Apogonichthyoides chrysurus
synonymy seems incorrect. This is a larger deep-bodied
species (11 cm) with 3 or more predorsal scales, which
occurs at lower latitudes in Queensland and in the Lord
Howe Island region of the Tasman Sea.
Apogonichthys ahimsa Holotype 32 mm TL. GPW

Waikiki Cardinalfish Apogonichthys waikiki

Apogonichthys waikiki Jordan & Evermann, 1903. Oahu Island, Hawaiian Islands.
Possibly restricted to the Hawaiian region. A secretive small species on shallow rocky reefs, and usually seen only at night.
At some stage was treated as a synonym of Apogonichthys perdix a closely related species and was reported as such. Body
dark-mottled and with black spots, showing a whitish band about halfway at night. Small white dots behind end of dorsal fin
and centrally basicaudal. Short dusky to black dashes radiating from posterior of eye. The caudal fin usually reddish. Length
to about 45 mm.
A B
Apogonichthys waikiki Hawaiian Islands. JER Apogonichthys waikiki Hawaiian Islands. JHO
Tan Cardinalfish Apogonichthys infuscus

Apogon (Apogonichthys) infuscus Fourmanoir, 1955.
Abu Latt/Alith Island, Saudi Arabia, Red Sea.
Western Indian Ocean, Red Sea region to South Africa. Tan
with numerous small dark spots. Was treated as a synonym
of A. perdix at some stage. According DNA analysis A.
infuscus is thought to replace A. perdix in the WIO (Sergey
Bogorodsky, pers. comm.). Photo identification uncertain:
pectoral fin unusually long in this small individual shown,
reaching level of below end of dorsal fin; caudal peduncle
deep. Length to 50 mm.
Apogonichthys infuscus Dahab, Egypt, Red Sea. SBO
Ocellated Cardinalfish Apogonichthys ocellatus

Apogon ocellatus Weber, 1913.
Northern Sulawesi, Indonesia.
Widespread Indo-Pacific, known from the east Africa to
Micronesia and Australia, in depths of a few metres on reefs,
secretive in coarse rubble and macro-algae habitats. D. VII,
but may have a tiny 8th spine hidden under skin. Lateral line
distinct with 23 tubed scales. Pale brownish to blackish with
black markings from behind eye and spotting on body. Fins
rounded and dusky. First dorsal fin somewhat angular with a
large blue-black ocellus. Length to 60 mm.
A
Apogonichthys ocellatus Madang, PNG. GRA
B C
Apogonichthys ocellatus Marquesas, French Polynesia. B JTW C JER

TRIBE APOGONICHTHYINI – Genus Foa Jordan & Evermann, 1905

Fem. Type-species Fowleria brachygrammus Jenkins, 1903.
Foa comprises a group of small taxa. Eight described species are recognised:
F. brachygramma .....8 F. hyalina ............11 F. leisi.........................10 F. nivosa ............10
F. fo.........................8 F. landoni ..............9 F. madagascariensis.....11 F. yamba ............10
Several additional undescribed species are known.
Weed Cardinalfish Foa brachygramma

Fowleria brachygrammus Jenkins, 1902. Oahu Island, Hawaiian Islands.
Endemic to Hawaiian Islands. Occurs abundantly in sheltered bays of the region, but cryptic on macro-algae rubble reef and
usually observed at night. Often found in brackish water zones and may enter fresh water (Hoover, 2008). Length to 65 mm.
A B
Foa brachygramma Night. Hawaii. JER Foa brachygramma Night. Kewalo, Hawaii. JHO
Fo Cardinalfish Foa fo
Foa fo Jordan & Seale, 1905. Negros, Cavite, Luzon Island,
Philippines, Manila Bay, South China Sea, western Pacific.
A common widespread Indo-West Pacific species with similar
siblings in many parts of the Indo-Pacific. Very cryptic and
usually occurs on open soft bottom in macro-algae rubble and
soft coral habitats. Brown to near black colour with indistinct
darker banding and white speckles, a white axil spot and like
several congeners with 3 white spots on caudal fin base. First
dorsal and ventral fins similarily coloured to body. Attains
about 40 mm.
A
Foa fo Tulamben, Bali, Indonesia. RHK
B C
Foa fo Males with oral brood. B Philippines. C Ningaloo, Western Australia. B PWO C GJW

E
Foa fo Milne Bay, PNG. RCS
D F
Foa fo Maumere, Flores, Indonesia. RHK
G H
Foa fo Mabul, Borneo, Malaysia. TKO Foa fo Naama Bay, Red Sea. SBO
Landon’s Cardinalfish Foa landoni

Apogonichthys landoni Herre, 1934. Cebu Harbour, Philippines.
West Pacific, Philippines and Indonesia. Rarely seen, presently known only from the type material and a few photographed
individuals. From original description: D. VIII-I, 9; A. II, 7; L.L. 21+2; 2 scale rows above and 6 below L.L.; 2 scale rows
on preopercle; 6 pre-dorsal scales. Mostly red, a pale region between second dorsal and anal fins. Length to 45 mm.
A B
Foa landoni Northern Sulawesi. BLA Foa landoni Night. Philippines. TKO

Harbour Cardinalfish Foa yamba

Foa yamba Fraser, 2014.
Yamba, Iluka Marina, New South Wales, Australia.
One of a complex of similar Indo-Pacific species and it
appears to be restricted to eastern Australia. Fairly common
in Sydney Harbour, very cryptic, usually on open soft bottom
habitats near outcrops or debris. Very similar to Foa fo, in its
dark dusky colour and 3 white spots on caudal-fin base, but
basically a dusky brown with dark small spots and diffused
whitish markings. First dorsal and ventral fins similarily
coloured to body. Previously identified as Foa fo. Attains
about 50 mm.
A
B C
Foa yamba C juvenile. Sydney, Australia. RHK
Snowy Cardinalfish Foa nivosa Leis’ Cardinalfish Foa leisi

Foa nivosa Fraser & Randall, 2011. Majuro Atoll, Marshall Is. Foa leisi Fraser & Randall, 2011. French Polynesia.
Known from Palau, the Marshall Islands and Fiji. Marshall West Pacific, known only from French Polynesia. Rarely
Island specimens were collected in Halimeda beds in 15–27 seen. This species has been collected from near surface to
meters (Fraser & Randall, 2011). Can be expected in various 18 m depth in lagoons, bays and near mouths of small tidal
macro-algae habitats. Colour: head and body reddish to streams. One specimen was collected in a pen shell, Atrina
brownish red with series of tiny whitish spots. A deep-bodied pectinta (Fraser & Randall, 2011). Previously identified as
small taxon, length to 40 mm. Foa fo. Length to 45 mm.
Foa nivosa Ngeruketabel Island, Palau. RWI Foa leisi Paratype. Haurei Bay, Rapa Iti. JTW
White-speckled Cardinalfish Foa sp. 1 Heemstra’s Cardinalfish Foa sp. 2

Undescribed new species (Fraser, pers. comm.) Maybe Foa madagascariensis var. (Fraser, pers. comm.)
Known only from the Maldives, but is probably widespread Known from Mauritius and Mozambique, western Indian
in the western Indian Ocean. Colour reddish brown with Ocean. Colour grey with dusky spots and indistinct barring,
white speckles over head and body. Length about 45 mm. Ventral fins dusky. Length about 45 mm.
Foa sp. 1. Maldives. JER Foa sp. 2. Mauritius. PCH

Snow-flakes Cardinalfish Foa sp. 3

Undetermined taxon, only known from a few photographs.
Based on visible meristics and colour pattern it belongs to the genus Foa and is most similar to F. fo, but the white flecks
colour pattern is different from its congeners. Length about 40 mm.
A B
Foa sp. 3. Okinawa, Japan. AMI Foa sp. 3. Kalimantan Island, Malaysia. KNI
Tide-pool Cardinalfish Foa madagascariensis

Foa madagascariensis Petit, 1931.
Sarodrano, Province of Tuléar, Madagascar, depth 5–6 m.
Red Sea to South African coast, south to Natal, Seychelles
and Madagascar. Body brown or greenish to pale tan with
few faint vertical bars and with numerous dusky-black spots;
ventral fins dusky. Reported as being common among weeds
in tidal pools. Deep-bodied, length to 60 mm.
Foa zuluensis Fowler, 1934, the type of which came from
Natal, is a putative synonym. A
Foa madagascariensis Preserved specimen. Tanzania. SRA
B C
Foa madagascariensis Dahab, Egypt, Red Sea. SBO
Sharp-snout Cardinalfish Foa hyalina

Amia hyalina Smith & Radcliffe, 1912.
Northern Sulawesi, Indonesia.
West Pacific. Philippines to Madang, PNG. Coastal
and lagoons, usually among soft corals in 2–15 m
depth. Distinct by its head profile and the pinkish
colour with red-brown bands. Length to 50 mm.
A B
Foa hyalina Juvenile. Palau, Micronesia. HNA Foa hyalina Madang, PNG. GRA

TRIBE APOGONICHTHYINI – Genus Fowleria Jordan & Evermann, 1903

Fem. Type-species Apogon auritus Valenciennes, 1831.
Genus Fowleria comprises a group of small taxa. Eight described species are recognised:
F. abocellata...........15 F. aurita .................12 F. flammea.............14 F. vaiulae ...............15
F. amblyuroptura....14 F. isostigma ............13 F. marmorata .........12 F. variegata. ...........13
Many Fowleria taxa feature an opercle spot or ocellus, but it is absent in the smaller species, that look more
like members of the genus Apogonichthys, but differ in lateral-line formation with fewer tubed scales.
Crosseye Cardinalfish Fowleria aurita

Apogon auritus Valenciennes, 1831. Mauritius.
Widespread Red Sea and Indo-West Pacific. Occurs in sheltered dense rubble and boulder reefs, usually very shallow, but
reported to 30 m depth. Cryptic, rarely seen during the day and usually comes out at night. Pale to dark orange-brown with
short black lines radiating from posterior of eye; a large black spot on opercle with yellow edge and a short stripe above of
equal width; fins brown in adults. Large species for the genus, length to 10 cm.
A B
Fowleria aurita Red Sea. SBO Fowleria aurita Comores. PCH
C D
Fowleria aurita Kerama, Japan. AON Fowleria aurita Cocos Island, Indian Ocean. GRA
Eared Cardinalfish Fowleria marmorata

Apogonichthys marmoratus Alleyne & Macleay, 1877.
Cape Grenville, Qld, Australia.
Widespread Indo-West Pacific. Clear coastal to outer reefs,
secretive in mixed rubble and algae slopes. To about 20 m
deep, out at night. Pinkish grey with many brown bars. Large
black ear-like opercle spot with a thin margin of yellow. Median
fins mostly red, usually fading on margins. Length to 75 mm.
A
Fowleria marmorata Lizard Island, Queensland. RHK
B C
Fowleria marmorata Red Sea. SBO Fowleria marmorata Madang, PNG. GRA

Dotted Cardinalfish Fowleria isostigma

Apogonichthys isostigma Jordan & Seale, 1906.
Apia, Upolu Island, Samoa.
Widespread Indo-West Pacific and Red Sea. Coastal
mud flats with outcrops and reefs to about 10 m deep.
Very secretive, rarely seen and usually only out at night.
Colour dusky to dark brown with series of small black
spots over sides. Eye yellow. A large opercle spot, black
with yellow sharp edge. Fins plain. Length to 90 mm.
At some stage the Red Sea spotted form was identified

A
as Fowleria punctulata (= F. variegata?).
Fowleria isostigma Al Wajh bank, Red Sea. SBO
B C
Fowleria isostigma Red Sea. SBO Fowleria isostigma Lizard Island, Queensland. RHK
D E
F. isostigma Mururoa Atoll, French Polynesia. JTW Fowleria isostigma Lembeh Strait, Sulawesi, Indonesia. RHK
Variegated Cardinalfish Fowleria variegata

Apogon variegatus Valenciennes, 1832. Mauritius.
Widespread Indo-West Pacific ranging into sub-tropical zones. Coastal bays and harbours, usually dead coral rubble or rocky
habitats to about 10 m deep. Secretive, rarely seen and usually out at night. Colour dusky brown with indistinct banding,
black spots and white speckling over sides and fins or plain. A large yellow-edged black spot on opercle. Length to 80 mm.
A B
Fowleria variegata Night. Amami Island, Japan. HDO Fowleria variegata Okinawa, Japan. TKO
C D
Fowleria variegata (plain form) Red Sea. SBO Fowleria variegata Sydney Harbour, Australia. RHK

Chestnut Cardinalfish Fowleria sp. 1 & 2

Undetermined taxa from Japan and Philippines. West Pacific, unknown distribution. Sp. 2 was caught on a fishing line in
shallow water. Image taken in Philippines shows what seems a different species. First dorsal fin spine minute. Caudal fin is
rounded, they have a prominent eye-size black ear spot, typical for genus, and lateral line ending below second dorsal fin.
Body almost uniformly dark brown and fins brownish or dusky. Length to 90 mm.
A B
Fowleria sp. 1. Anilao, Philippines. ARY Fowleria sp. 2. Okinawa, Japan. TKO
Flame Cardinalfish Fowleria amblyuroptera

Apogon (Apogonichthys) amblyuropterus Bleeker, 1853.
Wahai, Ceram, Indonesia.
West Pacific, only known from southern Indonesia. Coastal
reef slopes and bays, in macro-algae rubble habitats, shallow
to about 20 m depth. Uniformly bright yellow-orange with a
large reddish yellow eye. Red spots in median fins forming
indistinct narrow bands. Lateral line complete with tubed
scales. A small species, length to 40 mm.
B C
Fowleria amblyuroptera A & B females. C male with oral brood. Gilimanuk and Tulamben, Bali, Indonesia. RHK
Stop Light Cardinalfish Fowleria flammea

Fowleria flammea Allen, 1993. Madang, Papua New Guinea
West Pacific, Papua New Guinea, southern Indonesia to Philippines. Coastal, lagoons to outer reefs in 7 to 30 m depth.
Seldom seen during the day. First dorsal fin tall. Deep-red in colour, whitish under eye on cheek and belly, eye yellow to
red, fins clear. Lower lip with fine barring. Length to 50 mm.
A B
Fowleria flammea A holotype B aquarium, Philippines. A GRA B TKO

Dwarf Cardinalfish Fowleria vaiulae complex

Apogon vaiulae Jordan & Seale, 1906. Apia, Upolu Island, Western Samoa.
Widespread Indo-West Pacific but it may comprise a species-complex. Sheltered shallow coastal macro-algae reefs to about
15 m depth. Secretive solitary species, rarely seen and usually out at night. Colour pale pinkish brown with variable, more
or less distinct reddish or dusky banding. Dusky short bands radiating from posterior half of eye. Uncertain genus placement,
lacks the generic opercle-spot, and has a tiny black dash at lateral-line origin. A small species, length to 40 mm.
A B
Fowleria vaiulae Type-form. French Polynesia. JTW Fowleria vaiulae Bali, Indonesia RHK
C D
Fowleria vaiulae Bali, Indonesia. RHK Fowleria vaiulae var. Flores, Indonesia RHK
E F
Fowleria vaiulae var. Maldives. RHK Fowleria vaiulae var. Bali, Indonesia RHK
G H
Fowleria vaiulae var. Kerama, Japan. AON Fowleria vaiulae Comores. 12 mm SL. PCH
I J
Fowleria vaiulae var. abocellata Red Sea. I SBO J JHI

TRIBE APOGONICHTHYINI – Genus Neamia Smith & Radcliffe, 1912

Fem. Type-species Neamia octospina Smith & Radcliffe, 1912.
Genus Neamia comprises four species: N. articycla, N. notula, N. octospina, and N. xenica.
Eightspine Cardinalfish Neamia octospina

Neamia octospina Smith & Radcliffe, 1912.
Palawan, Philippines.
Widespread Indo-West Pacific and Red Seas. Sheltered
shallow coastal to offshore in macro-algae coral reefs,
usually in 3–5 m depth. Secretive solitary species, hiding
deep into crevices. Very pale in colour, slightly pinkish
white with reddish bands radiating from posterior of eye.
It has a high number of pectoral-fin rays, 17–21. A small
species, length to 50 mm.
A
Neamia octospina Rowley Shoals, WA. GRA
B C
Neamia octospina Nusa Penida, Bali, Indonesia. GRA Neamia octospina Farasan Island, Red Sea. SBO
Circular Cardinalfish Neamia articycla

Neamia articycla Fraser & Allen, 2006. Lizard Island, Qld, Australia.
West Pacific known from a few West Pacific locations, southern Japan to northern Australia and Fiji. Reported from 1–40
m depths, living on macro-algae rubble and soft-coral reefs. Pale brownish red with faint banding and few dusky spots on
sides. Fins clear to yellow or red. Dorsal fins with black markings. It has a large round opercle spot with a yellowish edge.
Looks very similar to Fowleria, but it has a complete lateral line with 22–23 scales. Eye orange. Length to 86 mm.
A B
Neamia articycla Indonesia. MVE Neamia articycla Mindanao, Philippines. JTW
C
Neamia articycla Raja Ampat, West Papua, Indonesia. GRA

Gillspot Cardinalfish Neamia notula

Neamia notula Fraser & Allen, 2001. Mauritius.
Indo-Pacific, known from the type-locality at 17 m, from
Bali where photographed at 40 m and one collected at
70 m, also reported from Japan, and the Philippines. It
is a moderately deepwater species which is rarely
observed, but is more active during the day than the very
similar looking Fowleria members, from which Neamia
differs in having a complete lateral-line with tubed scales.
Orange-red in colour and has a reflective black opercle
blotch. Eye red. Length to 45 mm. A
Neamia notula Mauritius. Holotype. PCH
B C
Neamia notula Bali, Indonesia. GRA
Strange Cardinalfish Neamia xenica

Neamia xenica Fraser, 2010. Kiritimati Atoll, Kiribati.
Known from the single specimen shown, 26 mm long,
which was collected from 93 m depth at Kiritimati Atoll,
Kiribati by Richard Pyle, utilizing mixed-gas rebreathers.
When caught it was pinkish red with clear fins (inset) and
eye red. Appears to be juvenile, opercle has a patch of
small dark spots, which shows in preserved specimen,
seemingly in the development or process of forming a
round spot as shown in the congeners Neamia articycla
and N. notula (above).
Neamia xenica Holotype. Kiritimati Atoll, Kiribati. RPY
TRIBE APOGONICHTHYINI Genus Ozichthys Fraser, 2014

Masc. Type-species Apogon albimaculosus Kailola, 1976.
Genus Ozichthys comprises a single species: O. albimaculosus.
Cream-spotted Cardinalfish
Ozichthys albimaculosus
Apogon albimaculosus Kailola, 1976.
Northern Australia.
Known from northern Australia and Papua New Guinea.
Generally reported from 7–37 m, living on soft bottom,
but may be trawled much deeper (62 m). Pale brownish
with cream spots. A short black band from eye to origin
of lateral line and darks spots on median fins. Second
dorsal and anal fin basally with a large yellow-edged
black spot. Length to 86 mm.
Ozichthys albimaculosus From 62 m. Port Hedland, WA. CSIRO

TRIBE APOGONICHTHYINI – Genus Vincentia Smith & Radcliffe, 1912

Fem. Type-species Vincentia waterhousii Castelnau, 1872. (=Vincentia conspersa (Klunzinger, 1872).
Genus Vincentia comprises 5 species: V. badia, V. conspersa, V. macrocauda, V. novaehollandiae, V. punctata.
Endemic to southern Australian waters. Hatchlings well developed, taxa highly localised.
Southern Gobbleguts Vincentia conspersa

Apogon conspersus Klunzinger, 1872.
Hobson Bay, Victoria.
Widespread south coast, including Tasmania. Highly
variable with several localised colour forms or represents
a species-complex. Commonly occurs in shallow coastal
rocky reefs, forming small aggregations in caves and
deep ledge during the day, but reported offshore to
depth of 65 m. Variably greyish brown to dark reddish
brown, speckled with dark spots, and in South Australia
the Vincentia waterhousii form (C) has white spots all
over, including on fins. The Tasmanian form was named
V. lemprieri (B), which has a black preopercle margin
A and black scribble markings over sides. Length to 14 cm,
Vincentia conspersa Flinders, Western Port, Victoria. RHK but usually to about 12 cm.
B C
Vincentia conspersa Bicheno, Tasmania. RHK Vincentia conspersa Edithburg, South Australia. RHK
D
Vincentia conspersa Female. Portsea, Port Phillip Bay. RHK

E F
Vincentia conspersa Portsea, Port Phillip Bay. RHK Vincentia conspersa Juvenile. Portsea. RHK
G H
Vincentia conspersa Flinders, Western Port. RHK Vincentia conspersa Juvenile. Portsea. RHK
I
Vincentia conspersa Male with oral brood. Blairgowrie, Port Phillip Bay. SLK

Scarlet Cardinalfish Vincentia badia

Vincentia badia Allen, 1987.
Point Peron, Western Australia.
Southern Australian coast and nearby islands, common
in South Australia in sheltered coastal bays and seagrass
estuaries, ranging west to Shark Bay, Western Australia.
Variable dark-grey, yellow, greenish or purplish brown to
bright red, including all the fins. Often confused with
Vincentia conspersa, where sympatric, but besides the
usually reddish colour it differs in having larger fins, as
well as more dorsal and anal fin rays. Length to 10 cm.
A
Vincentia badia Night. Venus Bay, SA. RHK
B C
Vincentia badia Port Victoria, SA. RHK Vincentia badia Juvenile. Pearson Island, SA. RHK
Orange Cardinalfish Vincentia punctata

Vincentia punctata Klunzinger, 1880.
King Georges Sound, Western Australia.
From the gulfs and Kangaroo Island, South Australia to
Shark Bay, Western Australia. Coastal rocky reefs,
secretive in caves and ledges in 5–30 m depth, comes
out at night to feed over more open substrates. Brown
to orange, adults with numerous tiny black spots in series
over body-sides along scale-rows. Preopercle-edge black.
Length to about 12 cm.
A
Vincentia punctata Rottnest Island, WA. GJW
B C
Vincentia punctata Castle Rock, WA. RHK Vincentia punctata Juvenile. Lucky Bay, WA. RHK

Smooth Cardinalfish Vincentia macrocauda

Apogon macrocauda Allen, 1987. Cheybe Beach, Western Australia.
Southern Australia from Gulf of St Vincent, SA to Albany, WA. Very secretive in rocky reefs at depths of 8–25 m. Differs
from congeners in having cycloid to weakly ctenoid scales instead of strongly ctenoid scales. Caudal peduncle long and body
more slender when juvenile. Orange-red with whitish blotches, body-colour extending partly onto fins, that are covered with
many half-pupil-size cream spots. Length to 10 cm.
A B
Vincentia macrocauda A juvenile. B large adult. Esperance, Western Australia. NTS
Eastern Cobbleguts
Vincentia novaehollandiae
Apogon novaehollandiae Valenciennes, 1832.
New South Wales, Australia.
East coast of Australia, southern Queensland to central
New South Wales. Commonly occurs in Sydney
Harbour and Botany bay in caves and ledges during the
day, coming out into the open at night to feed, floating
close to the bottom. Body pale to dark coppery brown,
variably with numerous small whitish speckles, colours
may extend onto fins. Dark short dashes radiating from
behind and below the eye. Length to 10 cm.
C D
Vincentia novaehollandiae C juvenile. D male with oral brood. Sydney, NSW. RHK

SUBFAMILY APOGONINAE – TRIBE APOGONINI

The tribe comprises 5 genera: Apogon, Astrapogon, Paroncheilus, Phaeoptyx, and Zapogon.
Diagnosis dorsal fin VI+I,9; anal fin II,8; developed gill rakers 9–19; posttemporal edge smooth or
weakly serrate; anterior nare with low rim and flap; preopercle smooth on ridge, serrate to smooth on
vertical edge, horizontal edge an unossified large flap; head and body with ctenoid or cycloid scales;
pored lateral-line scales 23–25, scales with one pore above center and one below central pore; intestine
and stomach pale (except two species); head and body reddish with or without blackish markings or head
and body brownish without bars or stripes, both color patterns may be translucent in some species.
Other characteristics one supernumerary spine; branched first segmented dorsal and anal ray;
ctenoid or cycloid scales on predorsal, cheek, breast, two pelvic scales, and body; cycloid scale on opercle
and onto base of caudal fin; pored lateral-line scales simple with one pore on upper side and one on
lower side; pectoral fin-rays 11–16; 0–3 supraneurals; pored lateral-line scales from posttemporal to
base of caudal fin; caudal fin forked or rounded; 9+8 segmented principal caudal rays, 15 branched,
upper and lower unbranched; unbranched procurrent rays, longest segmented; teeth on premaxilla,
dentary, vomer, palatine, all villiform (except one species canine-like on premaxilla and dentary) or
absent on palatine; six infraorbitals, bony shelf on third infraorbital; supramaxilla absent; basisphenoid
present, reduced or absent; anterior ceratohyal notched; 10+14 vertebrae; 8 ribs; 8–9 epineurals;
uroneurals absent; three epurals; five free hypurals, 1–2 fused and 3–4 fused, 1–2 fused and 3–4 fused
to terminal centrum; free parhypural; low crest on PU2 (after Mabuchi et al, 2014).
Key to the genera

1 Predorsal scaled...................................................................................................................2
Predorsal without scales along the center line to origin of first dorsal fin...................Astrapogon
2 Anal-fin rays 8 ....................................................................................................................3
Anal-fin rays 9 ...................................................................................................Paroncheilus
3 Stomach and intestine pale ..................................................................................................4
Stomach and intestine black ......................................................................................Zapogon
4 Preopercular flap does not extend past vertical edge.....................................................Apogon
Preopercular flap extends past vertical edge .............................................................Phaeoptyx
TRIBE APOGONINI – Genus Apogon Lacepède, 1801

Masc. Type-species Apogon ruber Lacepède, 1801 (= Mullus imberbis Linnaeus, 1758).
Genus Apogon comprises about 50 species globally, some with uncertain status.
Atlantic East & Central Pacific Indo-West Pacific + Red Sea
A. americanus....................24 A. atradorsatus ................28 A. campbelli ......................36
A. aurolineatus ..................30 A. atricaudus ...................28 A. caudicinctus ..................38
A. axillaris .........................25 A. dammermani...............33 A. coccineus ......................35
A. binotatus.......................25 A. deetsie ........................39 A. crassiceps ......................37
A. gouldi ...........................27 A. dovii ...........................29 A. dianthus........................39
A. imberbis ........................23 A. erythrinus ...................37 A. doryssa .........................35
A. lachneri.........................30 A. guadalupensis .............28 A. erythrosoma ..................36
A. leptocaulus....................27 A. kautamea ....................39 A. hypselonotus .................34
A. maculatus .....................23 A. lativittatus...................34 A. indicus ..........................36
A. mosavi ..........................30 A. marquesensis ..............37 A. kominatoensis ...............38
A. phenax..........................27 A. pacificus......................29 A. seminigracaudus............38
A. pillionatus .....................25 A. posterofasciatus ..........38 A. semiornatus ..................33
A. planifrons......................26 A. retrosella.....................38 A. susanae .........................35
A. pseudomaculatus...........24 A. rubrifuscus ..................39 A. talboti ...........................32
A. quadrisquamatus ...........31 A. tricinctus.......................39
A. robbyi............................31 A. unicolor ........................32
A. robinsi...........................30
A. townsendi .....................26

Mediterranean Cardinalfish
Apogon imberbis
Mullus imberbis Linnaeus, 1758.
Malta, Mediterranean Seas.
Widespread Mediterranean and eastern Atlantic from
Portugal to Gulf of Guinea, Azores, Madeira, Canary
Islands, Cape Verde Islands. Rocky reefs, may form
large schools during the day near or in caves and under
large overhangs. Usually observed in depths of about 10
to 20 m, but reported to 200 m from trawls. Colour
plain red, with or without a black peduncular spot, which
may be lost in adults, and faint dusky band on snout to
eye. Length up to 15 cm, but commonly to 10 cm.
A
Apogon imberbis São Tomé, Africa, east Atlantic. NVR
B C
Apogon imberbis B Canary Islands, east Atlantic. C France, Mediterranean. HDE
Flamefish Apogon maculatus

Apogon maculatus Poey, 1860. Cuba.
West Atlantic. Rocky reefs, secretive in crevices. Usually
seen shallow in 1–20 m depth, but reported to 128 m.
Colour pale to deep red, a black cheek-spot, usually has
a black pupil-size spot just below posterior end of the
second dorsal-fin and a may have a broad black blotch
on caudal-peduncle, but markings highly variable in their
density or presence. Length to about 12 cm.
A
Apogon maculatus Night. San Blas, Panama. GED
B C
Apogon maculatus Juvenile. Caribbean. RMY Apogon maculatus St. Vincent, Caribbean. RCS

Two-spot Cardinalfish Apogon pseudomaculatus

Apogon pseudomaculatus Longley, 1932. Tortugas, Florida, U.S.A.
Western and eastern Atlantic. Rocky reefs, secretive in caves and crevices. Usually observed in the shallows to about 20 m,
but reported from trawls to about 134 m depth. Colour red, with a two black spots, one below end of the second dorsal
and other on upper-half of caudal-fin base. Pearly appearance at night (C). Length to 12 cm.
A B
Apogon pseudomaculatus São Tomé, Africa, east Atlantic. NVR Apogon pseudomaculatus Bahamas. RMY
C D
Apogon pseudomaculatus Ascension Islands, central Atlantic. PWI Apogon pseudomaculatus Brazil, west Atlantic. ACF
Brazilian Flamefish Apogon americanus

Apogon americanum Castelnau, 1855. Bahia, Brazil.
Western central and southwestern Atlantic: a species
endemic to Brazil, including Fernando de Noronha
Archipelago and St. Paul's Rocks. Rocky reefs, secretive
in crevices. Usually shallow in 1–20 m depth. Colour
pale to deep red, often a defused pupil-size spot on the
opercle. All fins red in adults. Usually has a faint dusky
saddle-like marking below second dorsal fin and one on
caudal peduncle, markings are usually more obvious in
juveniles or at night. Length to about 12 cm.
A
Apogon americanus Brazil, west Atlantic. ACF
B C
Apogon americanus B night. C juvenile. Praia do Forte, São João da Mata, Bahia, Brazil, west Atlantic. ACF

Axillary-spot Cardinalfish Apogon axillaris

Apogon axillaris Valenciennes, 1832. Ascension Island.
Central Atlantic: Ascension and St. Helena islands. Rocky reefs, in caves and under large overhangs with rich invertebrate
growth to about 35 m depth. Often seen swimming upside down agains ceiling. Colour plain pinkish red, a distinct black
axilliary spot and faint dusky band from snout through eye over cheek. Length to about 11 cm.
A B
Apogon axillaris A St. Helena Island, central Atlantic. B Ascension Islands, central Atlantic. A RLU B PWI
Broad-saddle Cardinalfish
Apogon pillionatus
Amia pillionatus Böhlke & Randall, 1968.
North side of Cabagua Island, Margarita Island, Venezuela.
Western Atlantic, Florida Keys, Bahamas and Caribbean.
Rocky reefs, secretive in crevices. Usually deeper than
20 m and reported to 122 m depth. Colour pale pink to
orange or red, a short black bar after end of 2nd dorsal
fin, caudal-peduncle usually dusky with 2 narrow white
bars. Eye large and dark. Fins transparent pinkish. A
small species, length to 65 mm.
Apogon pillionatus Jamaica, west Atlantic. PCO
Barred Cardinalfish Apogon binotatus

Amia binotata Poey, 1867. Cuba.
Western Atlantic. Rocky reefs, secretive in crevices.
Usually shallow in 3–30 m depth, reported to 60 m.
Colour pale pinkish orange, pearly appearance at night.
A narrow brownish to black bar under end of the second
dorsal-fin and another on end of caudal-peduncle. Fins
semi-transparent pink. Eye large, without horizontal
lines and very dark. Length to 12 cm.
B
Apogon binotatus Bahamas, west Atlantic. RMY

Pale Cardinalfish Apogon planifrons

Apogon planifrons Longley & Hildebrand, 1940.
Tortugas, Florida, U.S.A.
Usually shallow in 3–30 m depth. Colour pale cream to
pinkish, narrow brownish to black bar under end of the
second dorsal and a second bar posteriorly or patch on
the caudal-peduncle. Fins reddish. Top of eyes yellowish.
Pale pearly appearance at night. Length to about 12 cm.
A
Apogon planifrons Night. Arrraial do Cabo, Brazil. JLG
B C
Apogon planifrons B Exuma, Bahamas. C male with oral brood. Cayman, west Atlantic. B AME C CAB
D E
Apogon planifrons Night. San Andres, Colombia, Brazil, west Atlantic. ACF
Belted Cardinalfish Apogon townsendi

Amia townsendi Breder, 1927. Saddle Rock, Washerwoman Cut, Bahamas.
Western Atlantic. Rocky reefs, secretive in crevices. Usually found very shallow, but reported to 55 m depth. Colour pale
pink to orange with 3 narrow line-like black bars, first under end of the second dorsal, other two at end of caudal-peduncle
filled with brown as a dark blotch. Eye very large and dark. A small species, length to 65 mm.
A B
Apogon townsendi In front, night. San Andres, Brazil. ACF Apogon townsendi Bonaire, west Atlantic. PMA

Slendertail Cardinalfish Apogon leptocaulus

Apogon leptocaulus Gilbert, 1972. Palm Beach County, Florida, U.S.A.
Western Atlantic, Jamaica, Florida and Caribbean in 18–60 m depth, rarely caught or seen. It was photographed to the
west of Montego Bay at about 30 m depth in an area of deep crevices, very much like a deeper reef habitat. Head flattened,
eye large, distinct scalation and the caudal fin is unusually deeply forked with long pointed lobes, unlike any other apogon
in the region. Colour pinkish with broad diffused dusky bands below dorsal fins and posteriorly on caudal peduncle. Fins pale
reddish. Many scales reflective pearly in deep water and at night. Length to 60 mm.
A B
Apogon leptocaulus Montego Bay, Jamaica, west Atlantic. PCO
Deepwater Cardinalfish Apogon gouldi

Apogon gouldi Smith-Vaniz, 1977. Northwestern edge of Bermuda Plateau.
Bermuda to south Caribbean, western Atlantic. Deep water, reported from 55–262 m depth. Body pale to bright red, with
darker red or dusky bands on caudal peduncle, that alternate with whitish bands. The median fins are clear with reddish
areas to the margins and on lobes. Caudal fin shallowly forked with broadly rounded lobes. Length to about 75 mm.
A B
Apogon gouldi Curaçao, west Atlantic. R&A
Mimic Cardinalfish Apogon phenax

Apogon phenax Böhlke & Randall, 1968. Nassau vicinity, Great Bahama Bank, Bahamas.
Western Atlantic, Florida Keys, Bahamas and Caribbean to Brazil. Rocky reefs, secretive in crevices. Usually deeper than 15
m and reported to about 50 m. Colour pinkish brown to deep red with 2 dusky bands on caudal peduncle. Often with dark
reddish along bases of vertical fins. Length to about 10 cm, usually smaller.
A
A B
Apogon phenax Night. San Andres, Brazil. ACF Apogon phenax Jamaica, west Atlantic. PCO

Bar-spot Cardinalfish Apogon retrosella

Amia retrosella Gill, 1863. Baja California, Mexico.
Gulf of California to southern Mexico. Rocky reefs,
secretive in crevices. Usually shallow in 1–20 m depth.
Colour pinkish brown to red, a vertical black stripe from
second dorsal to anal fin just past anus, a pupil-sized
black spot on the caudal fin base. Length to 10 cm.
Apogon retrosella Sea of Cortez. HDE
Plain Cardinalfish Apogon atricaudus

Apogon atricaudus Jordan & McGregor, 1898. Revillagigedos Islands and southern tip of Baja California. Eastern Pacific.
Rocky reefs, secretive in crevices. Usually found shallow in 1–20 m depth. Colour pale to yellow or deep red, broadly dusky
dorsally over nape to caudal peduncle. Variable with or without a dark streak between spines centrally in first dorsal-fin.
Apogon guadalupensis (Osburn & Nichols, 1916) has been placed in synonymy (Fraser, pers. comm.).
A B
Apogon atricaudus A Mapelo Island. B Revillagigedos Islands, Eastern Pacific. RRO
Black-tip Cardinalfish Apogon atradorsatus

Apogon atradorsatus Heller & Snodgrass, 1903.
Santa María Island, Galápagos Islands.
Galápagos and Mapelo Islands, Eastern Pacific. Inhabits
rocky reefs at shallow depths. Caudal fin moderately
emarginate. Head and lower part of body yellowish
brown to red, Upper part of body and caudal peduncle
dusky grey. A distinct black tip on second dorsal-fin and
variabe on lobes of caudal fin, sometimes a black tip on
A anal fin. Length to about 7 cm.
Apogon atradorsatus Revillagigedos Islands, Eastern Pacific. GRA
B
Apogon atradorsatus Cousins, Galápagos, Eastern Pacific. GED

Pink Cardinalfish Apogon pacificus

Mionorus pacificus Herre, 1935. Isabela Island, Galápagos Islands.
Eastern Pacific. Found shallow in coastal waters, usually in less than 15 m depth, living in rocky and coral reefs. Usually
seen in small groups or pairs, Body pale pink to reddish brown. A black stripe on snout through eye and a short narrow
vertical bar below second dorsal fin. Pair of yellow stripes through eyes, edging the black stripe. Length to about 10 cm.
A B
Apogon pacificus Isla Plata, Ecuador. GED Apogon pacificus Night. Coiba Brincanco, Panama. GED
C D
Apogon pacificus Sea of Cortez. HDE Apogon pacificus Sea of Cortez. RLU
Tail-spot Cardinalfish Apogon dovii

Apogon dovii Günther, 1861. Baja California, Mexico.
Gulf of California to Peru. Rock and coral reefs. Usually shallow in 5–20 m depth. Colour pinkish brown to reddish, a black
band running over snout, eye and cheek, a large black spot centrally at end of caudal peduncle. Intermittently a dusky stripe
over anterior half of lateral line. Length to 10 cm.
A B
Apogon dovii Panama, East Pacific. GRA
C D
Apogon dovii Roca Coibita, Panama. GED Apogon dovii Isla Plata, Ecuador. GED

Roughlip Cardinalfish Apogon robinsi

Apogon robinsi Böhlke & Randall, 1968.
Great Bahama Bank, Bahamas.
Usually shallow in 3–30 m depth, reported to 150 m.
Colour pale pink, a narrow dusky bar under end of the
second dorsal to anal fin, and a broadly-square patch on
caudal-peduncle. Fins translucent pink. Has a pearly
appearance dorsally at night. Length to 12 cm.
Apogon robinsi Abrolhos, Mato Verde, Brazil. ABE
Whitestar Cardinalfish Apogon lachneri

Apogon lachneri Böhlke, 1959.
North of the center of Green Cay, Bahamas.
Western Atlantic. Rocky and coral reefs, prefers clear
water habitats, secretive in crevices, usually observed at
night. Often very shallow, from intertidal to about 16 m
deep, but reported to 106 m. Colour pale pinkish
brown, a black saddle spot at end of the second dorsal,
with small white spot just behind. Eyes very large and
dark. A small species, length to 65 mm.
A
Apogon lachneri Night. San Andres, Colombia, Brazil. ACF
B C
Apogon lachneri Bahamas. RMY Apogon lachneri Jamaica. PCO
Dwarf Cardinalfish Apogon mosavi Bridle Cardinalfish Apogon aurolineatus

Apogon mosavi Dale, 1977. Cat Island, Bahamas. Amia aurolineatum Mowbray, 1927.
Western Atlantic, Bahamas, Belize, Haiti, and Jamaica. Siguanea Bay, Isle of Pines, south of western Cuba.
Coral and rocky reefs, and seagrass beds in 12–40 m. Body Western Atlantic. Rocky reefs, secretive in crevices. Usually
plain pale, salmon, with very faint pale brown stripes on the shallow in a few metres depth, but reported to 80 m. Colour
body from opercle edge to the caudal peduncle. Fins with a pale brownish, sometimes with dark short markings radiating
shade of pink. Length to about 35 mm. from behind and below eyes. Length to 65 mm.
Apogon mosavi West Atlantic. CBA Apogon aurolineatus West Atlantic. VTR

Sawcheek Cardinalfish
Apogon quadrisquamatus
Apogon quadrisquamatus Longley, 1934.
Tortugas, Florida.
Western Atlantic, south to Brazil. Usually open bottom
at moderate depths of about 12 to 107 m in seagrasses
or associates with invertebrates, including anemones,
sponges and conch shells. Colour pale dusky bronze to
reddish with series of pale spots forming lines and a faint
pupil-size dusky blotch on caudal peduncle. First dorsal
fin with white streak on membrane following the tallest
spines. Length to about 65 mm.
A
Apogon quadrisquamatus West Atlantic. JGA
B C
Apogon quadrisquamatus Praia do Forte, São João da Mata, Bahia, Brazil. ACF
Striped Cardinalfish Apogon robbyi

Apogon robbyi Gilbert & Tyler, 1997. Twin Cays, Belize.
Western Atlantic, Belize, Jamaica, Colombia, and
Panama south to Brazil. Found shallow in coastal waters,
usually in less than 15 m depth, but reported to 74 m.
Schooling in various habitats including in seagrass beds,
rocky reef crevices, swimming near anemones, and
among long urchin-spines. Body cream, pale salmon to
orange with brownish stripes along scale rows from the
opercle edge to the caudal peduncle. Caudal-peduncle
end has a faint oval brown to black blotch. The fins are
clear or pale yellowish. Length to about 50 mm.
A
Apogon robbyi Bonaire. RMY
B
Apogon robbyi Among long spines of an urchin, like seen with Siphamia members in the Indo-Pacific. Caribbean. HER

Hana Cardinalfish Apogon unicolor

Apogon unicolor Döderlein, 1901. Yokohama, Japan.
Appears to be widespread Indo-west Pacific, but some
records may be based on other species. Reported from
southern Japan in shallow coastal bays, usually in a few
metres depth. Dusky red body and fins, scales on back
with dark margins. Intermittently shows a whitish bar
anteriorly on caudal peduncle. Lateral line 2.5–3 scales
below first dorsal fin. Length to about 12 cm.
Similar to Apogon talboti which has lateral-line running
A higher below first dorsal fin.
Apogon unicolor Okinawa, Japan. TKO
B C
Apogon unicolor Palawan, Philippines. GRA Apogon unicolor Izu, Japan. HKA
Talbot’s Cardinalfish Apogon talboti

Apogon talboti Smith, 1961. Zanzibar.
Widespread Indian Ocean to Indonesia and Red Sea.
Nocturnal, secretive in caves, crevices and coral rubble.
Shallow to about 15 m depth. Pinkish red, scales with
dusky centres. Eyes usually yellowish white and all fins
red. Head profile dorsally straight. Lateral line very high,
1.5 scales below first dorsal fin. Caudal-fin with large
broadly rounded lobes. Length to 10 cm.
A Similar to Apogon unicolor in which the lateral-line runs
Apogon talboti Halmahere, Indonesia. GRA lower on upper sides.
B C
Apogon talboti Tosa Bay, Kochi Prefecture, Japan. HSE Apogon talboti Red Sea. SBO
Comores Cardinalfish Apogon aff. talboti

Undetermined member of the Apogon unicolor group
from the Comores.
Comores off Africa’s east coast, west Indian Ocean.
Sibling of Apogon talboti or a geographical variation
from the Comores with the lateral-line running high on
upper sides and a long caudal peduncle. Plain pinkish
red with a large pale yellowish white eye. Length of
specimen 55 mm.
Apogon aff. talboti Comores. PCH

Dammerman’s Cardinalfish Apogon dammermani

Apogon dammermani Weber & de Beaufort, 1929. Rapa, south-central Pacific.
West to central Pacific, eastern Indonesia, Papua New Guinea to Rapa. A close relative to Apogon talboti which shares the
lateral-line running very high along upper sides. Scales with red margins. Length to 95 mm.
A B
Apogon dammermani Madang, PNG. GRA Apogon dammermani Flores, Indonesia. RHK
Half-banded Cardinalfish
Apogon semiornatus
Apogon semiornatus Peters, 1876. Zanzibar.
Widespread Indo-west Pacific, ranging into sub-tropical
zones on the east coast of Australia. Rocky and rubble
reefs in caves and crevices. Shallow to about 30 m
depth. Transparent grey and red, Red on snout, a short
black band from behind eye to anus and another from
above eye to caudal fin base. Length to about 75 mm.
Below juveniles sharing a crevice with a juvenile Pterois
volitans, a tropical expatrate, and a pair of pufferfish
Canthigaster calisterna, a local temperate species.
A
Apogon semiornatus Iriomote Island, Japan. KYA
C
Apogon semiornatus Southern Oman. JHO
B D
Apogon semiornatus B small juveniles. Montague Island, southern NSW. D Tulamben, Bali, Indonesia. RHK

Elusive Cardinalfish Apogon sp. 1

Known from the southern Red Sea and Myanmar,
trawled from sand bottom in depths of 60–80 m. This
taxon will be described by Gon et al. (in prep.).
D. VI–I,9; A. II,8; P. 12; LL. complete, 24; predorsal
scales 7; scales between first dorsal fin and lateral line
1.5; body depth 3.1 in SL; head large, its length 2.2 in
SL; first dorsal-fin spine moderately long, 1.7 in length
of the second spine; preopercle ridge smooth, posterior
margin serrate; caudal fin forked with rounded lobes.
Colour including fins reddish orange, scale with dark
edges; faint dark dots of various sizes across caudal
peduncle close to caudal-fin base. Length to 7 cm.
Apogon sp. 1. Saudi Arabia, Red Sea. SBO
Black-margin Cardinalfish Apogon aff. unicolor Long-tail Cardinalfish Apogon sp. 2

Undescribed species of the Apogon unicolor group. Undetermined species.
Collected from New Caledonia. Lateral line runs high on West Pacific, but only known from the east coast of Australia.
upper sides similar to Apogon dammermani. A more Cryptic, solitary in caves and crevices of rocky reefs. Similar to
slender species in the group. Caudal peduncle long, fin Apogon crassiceps, but a larger taxon, much more elongated
large and forked with somewhat pointed rounded lobes. with long caudal peduncle, colour deeper pinkish red dorsally
Red, including fins, body scales with dark-dusky margins. and ventrally, and eyes black. Black along spine and abdomen.
Caudal fin with a broad blackish posterior margin, proba- A long second spine in first dorsal-fin, head profile flat dorsally.
bly black in live. Length to about 60 mm. Intestine black. Length to 80 mm.
Apogon aff. unicolor New Caledonia. RWI Apogon sp. 2. Sydney, NSW. RHK
Long-spine Cardinalfish Apogon hypselonotus Broadband Cardinalfish Apogon lativittatus

Apogon hypselonotus Bleeker, 1855. Apogon lativittatus Randall, 2001. Marquesas Islands.
Batu Islands, Indonesia. Marquesas Islands, Eastern Central Pacific. Rocky reefs,
Little known species from Pacific. Like the similar species secretive in crevices and caves, shallow to moderate depths.
probably secretive in crevices in 20–40 m depth. Eye very Colour pale to deep red, a broad dusky band from lateral-line
large suggesting being nocturnal and that it lives at moderate origin to caudal fin base. A second broad band from behind
depths or in the back of dark caves. Body colour pinkish to eye over pectoral-fin base to about anal fin and a narrower
reddish transparent. Dorsal-fin spines acceptionally long, band high on back along dorsal-fin bases. The banded dark
longest subequal to body depth. Length to about 45 mm. pattern is very similar to Apogon semiornatus, but bands
look faded anteriorly. Length to about 8 cm.
Apogon hypselonotus American Samoa. RCW Apogon lativittatus Marquesas Islands. SMI

Night Cardinalfish Apogon doryssa

Apogon doryssa Jordan & Seale, 1906. Japan.
Indo-West Pacific. Usualy occurs long drop-offs and
steep slopes in caves and ledges during the day, but
rarely seen, even where common. Out at night on the
bottom of caves or near reef on open substrate. Head
pinkish red with a pearly spot on cheek, and pearly spots
scattered dorsally on body outline, body mostly pinkish
semi-transparent with red along outline. Length to 50 mm.
B C
Apogon doryssa A/B Bali. C Flores, Indonesia. RHK
Red-Sea Ruby Cardinalfish

Apogon coccineus
Apogon coccineus Rüppell, 1838. Red Sea.
Red Sea and western Indian Ocean. Found in various
reef habitats, hiding during the day, but swimming over
open substrate close to reefs at night. Semi-transparent
red at night with iridescent blue spots scattered dorsally
on head and along the back. Length to about 45 mm.
A
Apogon coccineus Safaga, Egypt, Red Sea. RHK
B C
Apogon coccineus B Al Lith, Saudi Arabia, Red Sea. C Oman. B SBO C PWO
Susan’s Cardinalfish Apogon susanae

Apogon susanae Greenfield, 2001. Ifaluk Atoll, Caroline Islands, Micronesia.
Western Pacific. Body transparent pale pinkish, reddish long dorsal fin and caudal fin bases. A pale transparent area on
snout in front of eye. Length to about 40 mm.
A B
Apogon susanae A French Polynesia. B Ie-jima Island, Okinawa Prefecture, Japan. A JTW B HSE

Campbell’s Cardinalfish Apogon campbelli

Apogon campbelli Garman, 1903. Delagoa Bay.
Western Indian Ocean and Red Sea. Clear coastal and
island rocky reefs. Secretive in coral and rubble reefs and
out in the open at night. Usually in less than 25 m depth.
Caudal fin forked with broad round lobes. Very similar to
Apogon coccineus, but is stockier, deeper bodied with a
blunter snout and longer second spine in first dorsal-fin.
Colour pinkish red, slightly translucent with reflective
scales dorsally. Length to about 45 mm.
A
Apogon campbelli Egypt, Red Sea. RHK
B C
Apogon campbelli Farasan Island, Red Sea. SBO Apogon campbelli Aliwal Shoals, South Africa. RHK
Red Sea Night-cardinalfish Apogon erythrosoma

Apogon erythrosoma Randall, 2003. Marsa Ktana, Egypt, Gulf of Aqaba, Red Sea.
Red Sea and northwestern Indian Ocean. Rocky reefs, secretive. Usually seen at night, shallow in 1–20 m depth. Glass-like
transparent, colour very pale pinkish to reddish, usually small pearly spots dorsally and on opercle. Length to 45 mm.
A B
Apogon erythrosoma Dahab, Red Sea. SBO Apogon erythrosoma Egypt, Red Sea. RHK
Indian Night-cardinalfish Apogon indicus

Apogon indicus Greenfield, 2001. Mauritius, Mascarenes, southwestern Indian Ocean.
Widespread Indian Ocean to West Pacific. Rocky reefs, secretive. Usually seen at night, shallow in 1–20 m depth. Colour
almost transparent pale cream or pinkish to reddish, usually small pearly spots dorsally and on opercle. Length to 45 mm.
Looks identical to Apogon erythrosoma, and is only marginally different in pectoral-fin and gill raker counts.
A B
Apogon indicus Ryukyu Islands, Japan. HDO Apogon indicus Maldives. RHK

Ruby Cardinalfish Apogon crassiceps

Apogon crassiceps Garman, 1903. Fiji.
Reported as widespread central to southwest Pacific, but
may comprise a species-complex. Clear coastal and
island rocky reefs. Secretive in caves along drop-offs and
out in the open at night. Usually in less than 25 m depth.
Caudal fin forked with rounded lobe tips. Snout short
and rounded. Colour pinkish red, slightly translucent.
Length to about 45 mm. A
Apogon crassiceps Sydney, NSW. RHK
B C
Apogon crassiceps Surin Island, Thailand. USA Apogon crassiceps Byron Bay, NSW. BHU
Hawaiian Ruby-cardinalfish Apogon erythrinus

Apogon erythrinus Snyder, 1904. Puako Bay, Hawaii Island, Hawaiian Islands.
Endemic to the Hawaiian Islands with several similar species throughout the Indo-Pacific. Rocky reefs, secretive in crevices.
Usually shallow in 1–20 m depth. Colour transparent pale to deep red, a dusky cheek spot. Usually faint dusky markings
below dorsal fin origins and on caudal peduncle. Length to about 50 mm.
A B
Apogon erythrinus Hawaiian Islands. JHO
Marquesas Ruby-cardinalfish
Apogon marquesensis
Apogon marquesensis Greenfield, 2001.
Ua Pou, Marquesas Islands.
Endemic to the Marquesas Islands. Originally collected
from cave in less than 10 m depth. Compared to similar
species the dorsal fin spines are short. Colour pale to
deep red, somewhat transparent. Length to 45 mm.
B C
Apogon marquesensis A drawing of the type-specimen. After Greenfield. B/C specimens from the Smithsonian. SMI

ApogonidaeFishes22-43_ApogonidaeFishes22-43 11/09/19 9:20 AM Page 38
Dark-tail Cardinalfish
Apogon seminigracaudus
Apogon seminigracaudus Greenfield, 2007.
Vanua Levu, Fiji.
Widespread western Pacific: from southern Japan and
Philippines to Fiji, Tonga Islands and Samoa. Cryptic in
often silty coral and rubble reefs, shallow to 30 m depth,
usually solitary. Colour orange red, slightly translucent,
with dusky band on caudal peduncle fading into body
A and lower lobe of caudal fin, which is more prominent
Apogon seminigracaudus Palawan, Philippines. GRA in juveniles. Length to about 50 mm.
B C
Apogon seminigracaudus Iriomote Island, Japan. TKO Apogon seminigracaudus Kerama Island, Japan. ANO
Kominato Cardinalfish
Apogon kominatoensis
Apogon kominatoensis Ebina, 1935.
Tidepools at Kominato, Chiba.
Maybe restricted to southern Japan. Was recorded and
confused with Apogon coccineus restricted to the west
Indian Ocean region. Shallow, reported from sheltered
bays at depths of less than 5 m. Colour deep red, yellow
eye. Caudal fin with broad rounded lobes. A small red
species, length to 45 mm.
Apogon kominatoensis Izu Oshima, Japan. HSE
Rapa Cardinalfish Apogon caudicinctus

Apogon caudicinctus Randall & Smith, 1988.
Rapa, south-central Pacific.
West Pacific, Fiji and Solomon Islands to Japan.
Paratypes listed from Japan seem different and may be
based on different sibling species. Secretive and most
specimens were collected with the use of chemicals from
caves and crevice to 40 m depth. Colour brownish with
dusky banding below end of second dorsal fin and on
end of caudal peduncle. Length to 12 cm.
Apogon caudicinctus Florida Island, Solomons. JER
Rear-bar Cardinalfish
Apogon posterofasciatus
Apogon posterofasciatus Allen & Randall, 2002.
Florida Island, Solomon Islands.
Central west Pacific, Philippines to Solomon Islands and
Fiji. At moderate depths in 18 to 37 m depth in caves
and ledges. Body and fin colour yellowish or reddish,
slightly translucent, caudal peduncle with two faint dusky
broad bands, first partly into body, second basicaudal.
Length to about 60 mm.
Apogon posterofasciatus Lesser Sunda Islands, Indonesia. GRA

Rapanui Cardinalfish Apogon kautamea

Apogon kautamea Greenfield & Randall, 2004.
Tahai, Easter Island, southeastern Pacific.
Only known from Easter Island, southeastern Pacific.
Rocky reefs, secretive in crevices in 20–40 m depth.
Colour somewhat transparent pinkish to reddish. Eye
yellowish. Length to about 50 mm.
Apogon kautamea Easter Island, southeastern Pacific. JER
Deetsie’s Cardinalfish Apogon deetsie

Apogon deetsie Randall, 1998.
Oahu Island, Hawaiian Islands.
Hawaiian Islands, maybe endemic to the region. Rocky
reefs, secretive in the back of dark caves and crevices.
Usually occurs shallow in 1–20 m depth. Colour pale to
deep red, a dusky cheek spot, a defused pupil-size spot
high on the opercle. Usually faint broad dusky banding
below second dorsal fin and on the caudal peduncle.
Apogon deetsie Hawaiian Islands. JHO
Three-bar Cardinalfish Apogon tricinctus

Ostorhinchus tricinctus Allen & Erdmann, 2012.
Palawan, Philippines.
Widespread Indo-West Pacific: Comores; Indonesia east
to Palau and Fiji, south to Papua New Guinea. Clear
coastal and island rocky reefs. Cryptic in the back of
caves and crevices in coral and rubble reef slopes in
about 20 m depth and probably deeper. Colour plain
pinkish red to orange, slightly translucent when juvenile,
with some dusky banding. Length to about 50 mm.
Apogon tricinctus Palawan, Philippines. GRA
Flat-head Cardinalfish Apogon rubrifuscus

Apogon rubrifuscus Greenfield & Randall, 2004.
Motu Tautara, Easter Island.
Easter Island, southeastern Pacific. Type was collected
from a cave, depth 39 meters. Colour brownish red,
faintly dusky over upper sides, posteriorly and on caudal
peduncle. Fins large, transparent reddish, caudal fin long
and forked with large rounded lobes. First dorsal-fin tall,
sharply angular, second spine longest. Hear profile flat
dorsally. Eye creamish white. Length to about 10 cm.
Apogon rubrifuscus Easter Island, southeastern Pacific. JER
Pink-flower Cardinalfish Apogon dianthus

Apogon dianthus Fraser & Randall, 2002. Palau Islands.
Widespread Indo-West Pacific: Comores; Indonesia east
to Palau and Fiji, south to Papua New Guinea. Clear
coastal, off shore and island rocky reefs. Cryptic in caves
and crevices in coral and rubble reef slopes in about 20
m depth and probably deeper. Colour plain pinkish red
to orange, slightly translucent when juvenile. Length to
about 50 mm.
Apogon dianthus Brunei. GRA

TRIBE APOGONINI – Genus Zapogon Fraser, 1972

Masc. Type-species Apogon evermanni Jordan & Snyder, 1904.
Genus comprises two species: Zapogon evermanni and Z. isus. The two species were placed in Zapogon
Fraser, 1972, as a subgenus of Apogon. There are only minimal differences from Apogon as a genus that
cannot be determined easily, including genetically (Mabuchi et al, 2014).
Cave Cardinalfish Zapogon evermanni

Apogon evermanni Jordan & Snyder, 1904. Hawaii.
Circumglobal in tropical and subtropical seas, common in Indonesia at depths of about 20 m. Reported from the Atlantic
in 3–250 m depth. Rocky reefs, secretive in deep crevices, and usually swimming upside down on the ceiling of large caves
or overhangs. Brown to reddish yellow. A black stripe from snout through eye over cheek, a black spot followed by a white
spot, together at end of second dorsal fin base. This species has tiny body-scales compared to the nearly identical looking
sibling Apogon isus (following species). Length to about 12 cm.
A B
C D
Zapogon evermanni A–C Tulamben, Bali. Indonesia. D Maldives. RHK
Red Sea Cave-cardinalfish Zapogon isus

Apogon isus Randall & Bohlke, 1981. Red Sea.
Red Sea and Indian Ocean. Rocky reefs, secretive in crevices. Usually shallow in 1–20 m depth. Colour pale brown to red,
a black stripe from snout through eye over cheek, a black spot and white spot together at end of second dorsal fin base.
Looks identical in colour and morphology to the widespread Apogon evermanni above, but has much larger body-scales,
clearly visible in the images. Length to about 10 cm.
A B
Zapogon isus Night. Sharm El Naga, Red Sea. ARY Zapogon isus Angarosh Reef, Red Sea. SBO

TRIBE APOGONINI – Genus Astrapogon Fowler, 1907

Masc. Type-species Apogonichthys stellatus Cope, 1867.
Genus comprises three species: Astrapogon alutus, A. puncticulatus and A. stellatus.
‘Distinguished from subgenus Apogonichthys Bleeker, Nat. Tijds, Ned. Ind., VII, 1854, pp. 312, 321 (type
Apogonichthys perdix Bleeker), by the long ventrals, which reach well beyond the origin of the anal.’ Small
dark-looking fishes with long ventral fins, reaching far on base of anal-fin in two species; a near rounded tail,
slightly indented centrally. Eyes large with dark streaks radiating posteriorly. D. VI–I,9. A. II,8. P. 14–16.
Scales cycloid. Lateral line complete with 23–25 scales. No median predorsal scales (after Randall, 1968).
Conch Cardinalfish Astrapogon stellatus

Apogonichthys stellatus Cope, 1867.
Nassau, New Providence Island, Bahamas.
Western Central Atlantic: Bermuda, Florida, and
Bahamas to the Antilles and Rio de Janeiro, Brazil.
Prefers clear waters of oceanic islands in depths to
about 25 m. Commensal with the Queen Conch-shell
Lobatus gigas, in which found in the mantle cavity, and
with the bivalve Atrina rigida. Up to 5 individuals have
been found living in the same conch. Feeds at night on
small crustaceans near the bottom, away from the shell.
Greyish bronze to near black with numerous black
speckles. Several dark streaks radiating from eye. A
small species, length to about 50 mm.
Astrapogon stellatus In living conch-shell. Belize. PHU
Black-fin Cardinalfish
Astrapogon puncticulatus
Apogonichthys puncticulatus Poey, 1867. Cuba.
Western Central Atlantic: Bermuda, south Florida, and
Bahamas to the Antilles, and south to Rio de Janeiro,
Brazil. Prefers clear water seagrass habitats and along
reef-faces. Drifts near to the bottom in close vicinity to
their safety retreat. Often hides in empty shells and it
has been reported from mantle cavity of the conch shell
Strombus pugilis. Colour pale greyish bronze to dusky
with black and white speckles. Fins usually yellowish
with small dusky spots. Ventral fins very large, reaching
far over anal fin and colour is often more blackish.
Dorsal fin with black anterior margin. Both dorsal fins
and caudal fin may have whitish margin on lobes.
A
Astrapogon puncticulatus Bahia, Brazil. ACF
B C
Astrapogon puncticulatus Key Largo, Florida. NDL Astrapogon puncticulatus Brazil, west Atlantic. JLG

Bronze Cardinalfish
Astrapogon alutus
Apogon alutus Jordan & Gilbert, 1882.
Snapper banks off Pensacola, Florida, U.S.A. (from a
fish-stomach content).
Tropical western Central Atlantic. Prefers clear waters of
oceanic islands in seagrasses to moderate depths. Body
colour, underlaying bronze, but mostly covered with
dusky and black spots, some forming short dashes.
Somewhat similar to Astrapogon puncticulatus, but
readily distinguished by the much shorter ventral fins,
only just reaching anal fin. Fins mostly dusky with fine
A black spots and shading. Length to about 65 mm.
Astrapogon alutus Bahamas, west Atlantic. LJO
B C
Astrapogon alutus West Atlantic. VTR Astrapogon alutus West Atlantic. VTR
TRIBE APOGONINI – Genus Paroncheilus Smith, 1964

Masc. Type-species Paroncheilus stauchi Smith, 1964, =Paroncheilus affinis (Poey, 1875).
Genus Parocheilus is monotypic.
Diagnosis VI dorsal spines, fins deeply divided as VI–I,9; anal fin II,9; scales cycloid to weakly ctenoid;
lateral-line complete, 23–24 scales. Jaws with single row of teeth on dentary and premaxilla, some
enlarged as prominent canines. Preopercular ridge smooth; serrated posterior-edge, ventral edge not well
ossified (after Fraser, 1972).
Big-tooth Cardinalfish Paroncheilus affinis

Chilodipterus affinis Poey, 1875. Havana, Cuba.
Western and eastern Atlantic. Rocky reefs, schooling in
caves and under large overhangs with rich invertebrate in
about 20 to 100 m depth. Colour translucent salmon, to
pink, an indistinct dusky or yellowish band from snout
through eye over cheek. Fins clear. Length to 75 mm.
A B
Paroncheilus affinis Cape Verde Islands, east Atlantic. PWI Paroncheilus affinis Caribbean, west Atlantic. PHU

TRIBE APOGONINI – Genus Phaeoptyx Fraser & Robins, 1970

Fem. Type-species Amia conklini Silvester, 1915.
Genus Phaeoptyx comprises three Atlantic species: P. conklini, P. pigmentaria and P. xenus.
Diagnosis VI dorsal spines, fins divided as VI–I,9; anal fin II,8; scales cycloid; lateral-line complete,
24–25 scales. Both jaws with narrow bands of fine teeth (slightly enlarged in P. pigmentaria). Preopercular
ridge smooth; serrated posterior-edge, ventral edge not well ossified (after Fraser, 1972).
Freckled Cardinalfish Phaeoptyx conklini

Amia conklini Silveste, 1915. Coral reef off Guanica Harbor, Puerto Rico, Caribbean Sea.
Western Central Atlantic. Commonly found on shallow rocky and coral habitats, in empty conch shells or rubble. Abundant
where Acropora corals grow. Scales with dark margins. Median fins with black bands. Length to about 65 mm.
A B
Phaeoptyx conklini West Atlantic. VTR Phaeoptyx conklini West Atlantic. VTR
Sponge Cardinalfish Phaeoptyx xenus

Apogon xenus Böhlke & Randall, 1968. Off Nassau, Great Bahama Bank, Bahamas.
Western Central Atlantic. Occurs on sponge reef, hiding deep in tubular sponges. Comes out to feed at night. Body pale
brownish with small dusky spots, a black tail-base spot. Fins mostly clear, unmarked. Length to about 60 mm.
A B
Phaeoptyx xenus West Atlantic. VTR Phaeoptyx xenus West Atlantic. JER
Dusky Cardinalfish Phaeoptyx pigmentaria

Monoprion pigmentarius Poey, 1860. Cuba.
Widespread tropical western and eastern Atlantic (Gulf of
Guinea islands), Ascension Island. Secretive, hiding deep
in cave during the day. Drifts above reefs at night to
feed. Somewhat transparent, shiny silvery brown with
numerous small dusky spots A dusky tail-base spot. Fins
clear, unmarked. Length to about 65 mm.
A
Phaeoptyx pigmentaria Bahia, Brazil. ACF
B C
Phaeoptyx pigmentaria Bahia, Brazil. ACF Phaeoptyx pigmentaria São Tomé, Africa, east Atlantic. NVR

SUBFAMILY APOGONINAE – TRIBE ARCHAMIINI

Tribe Archamiini comprises 2 genera: Archamia and Taeniamia.
Diagnosis. Hypurals 1 and 2 and hypurals 3 and 4 fused together, fused to terminal centrum without
urostylar sheath; 2nd through 7th ribs with expanded flanges; soft anal-rays 15–17; basicaudal dark
spot subequal to pupil; body translucent in life, without stripes or bars (after Fraser, 2013b). Similar to
Apogon, but anal-fin has long base and many more rays.
TRIBE ARCHAMIINI – Genus Archamia Gill, 1863

Masc. Type-species Apogon bleekeri Günther, 1859. Genus Archamia is monotypic.
Bleeker’s Cardinalfish Archamia bleekeri

Apogon bleekeri Günther, 1859. Ambon, Indonesia.
Widespread Indo-West Pacific, if represented by a single
species. Inshore, often silty rocky reefs and mangroves.
Juveniles form schools, large adults usually pair. Transparent
yellowish grey, sides reflective silvery, a pupil-sized black spot
centrally on caudal-fin base. Yellowish on mouth. Fins clear,
transparent. Length to about 85 mm.
E–G a deep-bodied form as adults, from Durban Harbour,

South Africa has a more rounded overall shape with a highly
elevated back and appears to represent a different taxon.
A
B C
Archamia bleekeri C juvenile. A Maumere, Flores. B/C Singaraja, Bali. Indonesia. RHK
D E
Archamia bleekeri Singapore. GRA Archamia cf bleekeri Natal, South Africa. DPO
F G
Archamia cf bleekeri Undetermined deep-bodied high-back form. Durban harbour, South Africa. RHK

TRIBE ARCHAMIINI – Genus Taeniamia Fraser, 2013

Fem. Type-species Archamia leai Waite, 1916.
Genus Taeniamia comprises 15 named taxa:
T. ataenia .................50 T. flavofasciata..........45 T. lineolata...............47 T. pallida..................51
T. biguttata ..............46 T. fucata ...................48 T. macroptera ..........48 T. sansibarica ...........49
T. bilineata ...............51 T. kagoshimana........47 T. melasma ..............46 T. zosterophora. .......50
T. buruensis..............49 T. leai.......................46 T. mozambiquensis ..45
As Archamia, but with bars on head and body or darkish yellowish mid-body stripes.
Yellow-barred Cardinalfish
Taeniamia flavofasciata
Archamia flavofasciata Gon & Randall, 2003.
Durban, KwaZulu-Natal, South Africa.
Western Indian Ocean: Eastern South Africa to
Madagascar. In sheltered bay at moderate depth on open
substrate around outcrops, In aggregations hovering
above in about 20 m depth or more. Deep-bodied and
identified by the many yellow bars over the lower sides.
A distinct black pupil-sized spot centrally on caudal-fin
base. Dorsals, anal and ventral fins with white leading
margin. Length to 85 mm.
A
B C
Taeniamia flavofasciata At type-locality. Durban, South Africa. RHK
Mozambique Cardinalfish
Taeniamia mozambiquensis
Archamia mozambiquensis Smith, 1961. Mozambique.
Western Indian Ocean, Zanzibar to Sodwana Bay.
Coastal coral reef habitats, usually in loose aggregations
with branching corals to about 15 m depth. Pale whitish
grey, bluish dorsally with 2 brown to orange longitudinal
thin lines that may break up in to a series of spots with
growth or age. A distinct pupil-sized black spot centrally
on caudal-fin base. Length to 85 mm.
A
B C
Taeniamia mozambiquensis B male with oral brood. Sodwana Bay, South Africa. A DPO B/C GRA

Lea’s Cardinalfish Taeniamia leai

Archamia leai Waite, 1916. Norfolk Island.
South Pacific Oceanic locations off southern Queensland
and Tasman Sea, northern New South Wales, including
New Caledonia and Norfolk Island. Protected habitats on
coral reefs and in lagoons to about 15 m depth. Pale
brown to yellow posteriorly, dusky dorsally. Broad dusky
to black band from below 2nd dorsal to vent. A double
short bar above pectoral-fin base and a dark stripe on
snout through eye with bluish white margins. Fins pale
A pink to reddish. Length to 9 cm.
Taeniamia leai Norfolk Island, South Pacific Ocean. NCO
B C
Taeniamia leai Cook Island, northern New South Wales, Australia. ISH
Ear-blotch Cardinalfish Taeniamia biguttata

Archamia biguttata Lachner, 1951.
East Indies; Philippines; Samoa.
Western Pacific: Indonesia, north to the Philippines and
Taiwan, south to Papua New Guinea and east to Samoa.
Shallow coastal reef crests among corals and in caves,
usually in less than 10 m depth. Deep-bodied with a
large ‘ear’ spot and numerous thin orange, near vertical
lines. With or without a pupil-sized black spot centrally
on caudal-fin base A black band slightly angled backward
A from below eye. A short yellow line across snout. Length
to about 10 cm.
Taeniamia biguttata Milne Bay, Papua New Guinea. PWO
B C
Taeniamia biguttata With and without basicaudal spot. Maumere, Flores, Indonesia. RHK
Tear-spot Cardinalfish Taeniamia melasma

Archamia melasma Lachner & Taylor, 1960.
Northern Territory, Australia.
Northern Australia. Uncertain distribution due to the
similarity with Taeniamia biguttata. It differs from that
species in the black blotching that is faint below the eye
and the blotch above opercle that runs down ‘tear-like’.
It has only few thin yellow lines anteriorly on the body
close to pectoral-fin base. Apparently this species always
lacks a basicaudal spot and the numerous orange thin
lines along entire sides in similar taxa, unless a third
taxon is involved. Length to 85 mm.
Taeniamia melasma Cassini Island, WA. GRA

Shimmering Cardinalfish
Taeniamia lineolata
Apogon lineolatus Cuvier, 1828. Red Sea.
Red Sea, northwestern Indian Ocean: Gulf of Aden.
Coastal coral reef habitats, usually in loose aggregations
in coarse rubble reefs to about 10 m depth. Out at night
over open substrate, swimming close to the bottom.
Yellow on head and over abdomen, greyish dorsally and
many reddish, thin vertical lines. Pale semitransparent
whitish grey at night, bluish dorsally with many vertical
lines formed by close-set minute black or red spots. A
large, up to eye-size basicaudal spot. Length to 85 mm. A
B
Taeniamia lineolata A night colour. Egypt, Red Sea. RHK
Kagoshima Cardinalfish
Taeniamia kagoshimana
Apogon kagoshimanus Döderlein, 1883.
Kagoshima, Japan.
North Western Pacific, Japan. Shallow coastal reef crests
among corals and in caves, usually in less than 10 m
depth. A member of the Taeniamia fucata-complex that
has the numerous orange thin vertical lines along entire
sides. Distinguished in this complex by the brownish
blotch at eye level over the first few vertical lines closely
above and behind the pectoral-fin base. Length to about
85 mm. It was previously treated as Archamia dispilus
a putative junior synonym. A
B C
Taeniamia kagoshimana A/B Iriomote Island. C Kerama Island, Japan. A KYA B AON C TKO

Faint-lined Cardinalfish
Taeniamia macroptera
Apogon macropterus Cuvier & Valenciennes, 1828.
Java, Indonesia.
Widespread tropical West Pacific. Occurs in estuaries
and sheltered bays, often silted habitats. Congregates in
large schools, usually floating just above large branching
coral beds. Distinguished from the similar many-lines
species by the caudal peduncle being largely dusky to
black and without stripe across snout. Length to 10 cm.
B C
Taeniamia macroptera A Tulamben, Bali. B Maumere, Flores. C Gilimanuk, Bali, Indonesia. RHK
Painted Cardinalfish Taeniamia fucata

Apogon fucatus Cantor, 1850. Pinang, Malaysia.
Widespread tropical Indo-West Pacific and Red Sea.
Some geographical variations and several very similar
species. Typical form with numerous yellow to orange
thin lines along entire sides, vertical on upper and
angling forward on lower half. A yellow stripe across
snout. An up to eye-size black blotch on caudal-fin base,
variable from small to large, occasionally absent. Length
to about 10 cm.
A
B C
Taeniamia fucata Gilimanuk, Bali, Indonesia. RHK Taeniamia fucata Egypt, Red Sea. RHK
D E
Taeniamia fucata Queensland, Australia. RHK Taeniamia fucata Manado, Indonesia. RHK

Zanzibar Cardinalfish
Taeniamia sansibarica
Apogon (Archamia) sansibaricus Pfeffer, 1893.
Zanzibar, Tanzania, western Indian Ocean.
Western Indian Ocean, Zanzibar, Seychelles and Maldives.
Sheltered often silty coral habitats, manily in lagoons
with branching corals and large soft corals. Usually in
aggregations in shallow depths to about 6 m depth.
Similar to Taeniamia fucata, but somewhat transparent,
pale yellowish grey with numerous indistinct thin yellow
lines, Spot on centre of caudal-fin base small or absent.
Snout with yellow line across. Length to 85 mm.
A
B C
Taeniamia sansibarica Maldives. RHK
Yellow-barred Cardinalfish
Taeniamia buruensis
Apogon buruensis Bleeker, 1856.
Buru Island, Molucca Islands, Indonesia.
West Pacific, Throughout Indonesia, Thailand and
Philippines, east to Fiji. Coastal bays and usually found
in brackish water, entering tidal portions of freshwater
streams. Forms aggregations in shallow depths to about
3 m depth. Pale, somewhat transparent creamish grey
with two black lines and a pupil-size spot centrally on end
of caudal peduncle. Length to 85 mm. A
Taeniamia buruensis Buru Island, Indonesia. After Bleeker
B
Taeniamia buruensis Milne Bay, Papua New Guinea. GRA

Girdled Cardinalfish
Taeniamia zosterophora
Apogon zosterophorus Bleeker, 1856.
Ambon, Indonesia.
Tropical West Pacific with geographical variations. In
corals to 15 m depth. A pair of narrow orange to red
bars over gill-cover is typical. Also considered typical is a
broad black near-vertical band below second dorsal fin,
at slight angle to about in front of anal-fin base, but it can
be faint or completely absent. A small black spot on mid
A caudal-fin base. Length to 75 mm.
B C
Taeniamia zosterophora Flores, Indonesia. RHK
D E
Taeniamia zosterophora Pulau Putri, Java, Indonesia. RHK Taeniamia zosterophora Iriomote Island, Japan. RHK
Painted Cardinalfish Taeniamia ataenia

Archamia ataenia Randall & Satapoomin, 1999.
Mentawai Islands, West Sumatra, Indonesia.
Eastern Indian Ocean: Andaman Sea and Sumatra.
Usually forming schools in staghorn corals om reef crests
of protected reefs. Very similar and looks identical to the
Taeniamia zosterophora form without the black band
under the second dorsal fin, but apparently it always
lacks this black band. Length to about 10 cm.
A
Taeniamia ataenia Pulau Weh, Sumatra, Indonesia. GRA
B C
Taeniamia ataenia Surin Island, Thailand. USA

Black-spot Glass-cardinalfish
Taeniamia bilineata
Archamia bilineata Gon & Randall, 1995. Red Sea.
Red Sea endemic. Usually found schooling in caves and
swimming above coral head, shallow to about 30 m
depth. Glass-like transparent with several longitudinal
black thin lines and a pupil-sized or larger spot centrally
on caudal-fin base. A small species, length to 35 mm.
C D
Taeniamia bilineata C at night. Egypt, Red Sea. RHK
Pallid Cardinalfish Taeniamia pallida

Archamia pallida Gon & Randall, 1995. Masirah Island, Oman.
Northwestern Indian Ocean: Oman and Kenya, usually in shallow in 3–12 m depth. Schooling in weedy habitats in Kenya.
Pale, somewhat transparent grey with traces of two brown to blackish lines or series of small spots and pupil-sized spot on
centre of caudal-fin base, but spot absent in Kenya populations (B), possibly a different species. Length to 50 mm. Differs
from the similar Taeniamia bilineata in having more pectoral-fin rays, deeper body and a small or no tail spot.
A B
Taeniamia pallida Masirah Island, Oman. JER Taeniamia pallida Kenya. DGO

SUBFAMILY APOGONINAE – TRIBE CHEILODIPTERINI

Comprises the single genus Cheilodipterus Lacepède, 1801. Masc. Type-species C. lineatus Lacepède,
1801. Genus Cheilodipterus presently has 16 recognised species:
C. alleni.................53 C. isostigma ...........63 C. nigrotaeniatus ..65 C. pygmaios............64
C. arabicus.............61 C. heptazona .........54 C. novemstriatus ....64 C. quinquelineatus ..62
C. artus .................58 C. lachneri .............60 C. parazonatus.......65 C. singapurensis......57
C, intermedius .......56 C. lineatus .............55 C. persicus .............61 C. zonatus ..............65
Gon, 1994 revised the genus, but some taxa remain uncertain, representing species-complexes.
Diagnosis Members of the Apogoninae: VI+I,9; anal fin II,8; head and body with ctenoid scales;
pored lateral-line scales 24–26; canine teeth on premaxilla and dentary; preopercle ridge smooth,
edges serrate; three supraneurals; reduced supramaxilla; one pair of uroneurals; three epurals; five free
hypurals; free parhypural; caudal fin forked or emarginate; dark stripes on head and body, no stripes
in second dorsal or anal fin, basicaudal dark spot, band or no spot. Other characteristics one super-
numerary dorsal spine; branched first segmented dorsal and anal ray; ctenoid scales on predorsal,
cheek, breast, two pelvic scales, and body; ctenoid scale on opercle and onto base of caudal fin; pored
lateral-line scales with multiple pores; pectoral fin-rays 11–14; three supraneurals; 9+8 segmented
principal caudal rays, 15 branched, upper and lower unbranched; unbranched procurrent rays, longest
segmented; villiform teeth on vomer and palatine; six infraorbitals, bony shelf on third infraorbital;
basisphenoid present; anterior ceratohyal smooth or notched; 10+14 vertebrae; 8 ribs; 7–8 epineurals;
stomach and intestine pale with pale peritoneum; low crest on PU2 (after Mabuchi et al, 2014).
A favoured habitat of cardinal fishes is formed by the shading of a village jetty platform, built in protected location, the pylons
supported by rubble on the bottom. An amazing numbers of different species comprising several genera of the Apogonidae
family were mixed, filling the water column high above. Most taxa stayed near the substrate, but the dominating species
Cheilodipterus artus (Pacific form) was swimming in the open. Maumere Bay, Flores, Indonesia. RHK
Allen’s Cardinalfish Cheilodipterus alleni

Cheilodipterus alleni Gon, 1994. Madang, PNG.
West Pacific. Philippines, Indonesia and PNG. Usually
solitair along drop-offs in caves. Has 8–9 longitudinal
stripes, 5 of which ending on caudal-fin base. Dorsal fin
with black tip. Caudal-fin base dusky and outer rays black.
Eye lacks double lines, iris yellow. length to about 22 cm.
At some stage all Cheilodipterus taxa with 8 lines were
synonymised in C. macrodon, but they clearly comprise a
species-complex. This taxon was recognised as valid,
named C. alleni Gon, 1994, but it is identical Paramia
octolineatus (A) in Bleeker’s ATLAS. However, as
Bleeker’s description was presented under the C.
macrodon name, and he seemed to have assumed the A
species to be synonymus at the time after drawing it.
Paramia octolineatus Indonesia. Bleeker’s illustration.
B C
D E
Cheilodipterus alleni B Milne Bay, Papua New Guinea. C–E Maumere, Flores, Indonesia. RHK
F
Cheilodipterus alleni Typical species individual. Maumere, Flores, Indonesia. RHK

Pacific Tiger-cardinalfish
Cheilodipterus heptazona
Cheilodipterus heptazona Bleeker, 1849.
Jakarta, Indonesia.
Widespread West Pacific. Usually solitairy or in pairs
along reefs margins. Has 8–9 longitudinal tripes that end
abruptly to a white section of caudal peduncle. Caudal fin
base with a large black spot in centre when juvenile, base
and outer rays dusky in adults. Dorsal fin without black tip.
Head yellow in juveniles. length to at least 16 cm.
A The Pacific form of Cheilodipterus lineatus, commonly
referred to as C. macrodon, is not synonymus.
Cheilodipterus heptazona Milne Bay, PNG. RHK
B C
Cheilodipterus heptazona B Lizard Island, Queensland. C post-larval juvenile, Sydney Harbour, Australia. RHK
D E
Cheilodipterus heptazona Rowley Shoals, WA. RHK Cheilodipterus heptazona Tulamben, Bali. RHK
F
Cheilodipterus heptazona Typical Pacific form at type-locality. Pulau Putri, Java, Indonesia. RHK

Indian Tiger-cardinalfish
Cheilodipterus lineatus
Cheilodipterus lineatus Lacepède, 1801. Indian Ocean.
Widespread Indian Ocean and Red Sea. Usually solitairy
along drop-offs in caves. Has 8–9 longitudinal tripes that
end abruptly to a white section of caudal peduncle. Caudal
fin base with a central black spot when juvenile, often all
black in adults and outer rays black. Dorsal fin with black
tip in the Red Sea. Eye with double yellow stripe and
snout yellow in juveniles. length to 25 cm.
This taxon is generally referred to as Cheilodipterus A
macrodon, type-locality also from the Indian Ocean.
Cheilodipterus lineatus KwaZulu-Natal, South Africa. DKI
B C
Cheilodipterus lineatus B at night. Typical Red Sea form. Safaga, Egypt, Red Sea. RHK
D E
Cheilodipterus lineatus D post-larval juvenile. E subadult. Egypt. Red Sea. D RHK E SBO
F
Cheilodipterus lineatus Typical Indian Ocean form. Maldives. RHK

Intermediate Tiger-cardinalfish
Cheilodipterus intermedius
Cheilodipterus intermedius Gon, 1994.
Sesoko Island, Ryukyu Islands, Okinawa, Japan.
West Pacific, mainly Japan and China Seas, south to
Papua New Guinea. A solitary species in rubble reefs in
sheltered habitats, deep lagoons and caves to 20 m depth.
Similar to Cheilodipterus heptazona, but caudal fin base
lacks dusky bar and instead is yellow with a small black
central spot, becoming whitish in large individuals. Outer
caudal fin rays black. Length to about 20 cm.
A
Cheilodipterus intermedius Milne Bay, PNG. RHK
B
Cheilodipterus intermedius Kochi Prefecture, Japan. THI
C
Cheilodipterus intermedius Kochi Prefecture, Japan. THI D
E
Cheilodipterus intermedius Male with oral brood. Typical species individual. Kashiwajima, Japan. RHK

A B
Cheilodipterus males with their oral brood. A C. lineatus in the Maldives, B C. heptazona, Tulamben, Bali, Indonesia. and D
C. intermedius, Japan, on opposite page. The two above species are generally lumped under the name C. macrodon as a
very widespread Indo-Pacific taxon, but they are Indian and Pacific siblings that show differences in colour and it seems in the
canines features as well. They are treated as valid taxa and both have geographical forms as well. The brood comprises a great
number of tiny eggs and fry are pelagic, dispersing of vaste distances determined by ocean currents. RHK
Singapore Tiger-cardinalfish
Cheilodipterus singaporensis
Cheilodipterus singaporensis Bleeker, 1859. Singapore.
Widespread tropical West Pacific. A coastal species in silty
habitats, among boulders in seagrass areas. Juveniles
among sea-urchin spines. Shallow to about 10 m depth.
Dusky-brown in colour with 4 longitudinal black lines and
a small black caudal-fin base spot. Juveniles with white
longitudinal line from upper of eye ending as a white spot
above the black basicaudal spot, which has some yellow
above and below. Fins plain. Length to about 20 cm. A
Cheilodipterus singaporensis Pulau Putri, Java. RHK
B
C
Cheilodipterus singaporensis B inset, small juvenile. Maumere, Flores, Indonesia. RHK

Arrow-tooth Cardinalfish
Cheilodipterus artus
Cheilodipterus artus Smith, 1961. Seychelles.
Widespread Indo-West Pacific, but two forms: Indian (this
page) and Pacific (next page). Schooling in the Pacific,
often among staghorn corals. Forms small aggregations
along reefs in the Indian Ocean. Basicaudal spot large in
the Indian Ocean and small in the Pacific. Adults with
black band on caudal-fin base. Body lines black to orange,
alternating thin and thick in Indian Ocean and of even
A width in the Pacific. Length to about 12–20 cm.
B C
E
Cheilodipterus artus Typical Indian Ocean form. Maldives. RHK

A B
C
Cheilodipterus artus B male with oral brood. Bali, Indonesia. RHK
E
D Cheilodipterus artus Bali, Indonesia. RHK
F
Cheilodipterus artus Typical Pacific Ocean form. Flores, Indonesia. RHK

Lachner’s Cardinalfish
Cheilodipterus lachneri
Cheilodipterus lachneri Klausewitz, 1959.
Al Ghardaqa, Red Sea.
A Red Sea endemic, occurring south to Sudan, but rare.
Records from elsewhere appear to be based on the
misidentification of other species. Sheltered, often silty
reef habitats. Solitaire or loose small aggregations. Has
numerous thin lines, numbers increasing with growth, up
to 13 in adults, 3 or 4 of which thickened. A large black
spot centrally on caudal-fin base surrounded by yellow in
juveniles. Length to 14 cm.
Adults have 2 or 3 over the gills versus 4 or 5 lines in the
A almost identical looking Cheilodipterus arabicus.
Cheilodipterus lachneri Safaga, Egypt, Red Sea. HDE
B C
Cheilodipterus lachneri Red Sea. SBO Cheilodipterus lachneri Egypt, Red Sea. RHK
D E
Cheilodipterus lachneri Red Sea. SBO Cheilodipterus lachneri Egypt, Red Sea. RHK
F
Cheilodipterus lachneri Typical adult. Red Sea. SBO

Arabian Cardinalfish Cheilodipterus arabicus

Perca arabicus Gmelin, 1788. Red Sea. (Cheilodipterus caninus Smith, 1949, from Mozambique is a junior synonym).
Red Sea and tropical west Indian Ocean. Sheltered, often silty reef habitats. Solitaire or loose small aggregations. Has
numerous thin lines, numbers increasing with growth, up to 16 in adults, none of which obviously thickened. A small black
horizontally-elongate spot centrally on caudal-fin base, spot with yellow surround and a white area on caudal peduncle.
Adults have 4 or 5 thin black lines over gill-cover that are fewer and much thicker in the otherwise very similar
Cheilodipterus lachneri. Juveniles with only the principal body lines almost identical to C. quinquelineatus, but in latter
the basicaudal black spot is round and not horizontally elongate. Length to 18 cm.
A B
Cheilodipterus arabicus Juvenile. Red Sea. RHK Cheilodipterus arabicus Typical adult. Jeddah, Red Sea. SBO
Persian Cardinalfish
Cheilodipterus persicus
Cheilodipterus persicus Gon, 1994. Persian Gulf.
Persian Gulf and Arabian Sea to west of the Gulf of
Oman. Schools along reef walls in front of caves. A
whitish species with many thin black longitudinal lines,
alternating thin and thick, increasing in numbers with
growth. A large black spot centrally on caudal-fin base
with yellow surrounding blotch. Length to 15 cm.
Rarely sympatric, but similar to Cheilodipterus arabicus
and C. lachneri, adults best distinguished by their whitish
body colour. Juveniles with few lines could be mistaken for
C. quinquelineatus which is also whitish in body colour,
but basicaudal black spot is horizontally elongate.
A
Cheilodipterus persicus Male with oral brood. Oman. RFI
B C
D E
Cheilodipterus persicus B/C Di Hamri, Socotra, Arabian Seas. SBO. D/E Oman. RFI

Five-line Cardinalfish
Cheilodipterus quinquelineatus
Cheilodipterus quinquelineatus Cuvier, 1828.
Society Islands.
Widespread tropical Indo-Pacific, ranging into cooler
sub-tropical zones as expatriates. Common in various
habitats from coastal, silty to outer reef walls to about
35 m depth. Single or in small aggregations in corals or
caves. Colour white with 4 distinct black longitudinal
lines along side, a fifth ventrally. A large bright yellow
blotch at end of caudal peduncle and onto fin base with
A a smaller than pupil-size round black spot centrally at
level just above mid-lateral line. Length to 10 cm.
Cheilodipterus quinquelineatus Egypt, Red Sea. RHK
B C
Cheilodipterus quinquelineatus Rowley Shoals, WA. RHK Cheilodipterus quinquelineatus Sydney. RHK
D
Cheilodipterus quinquelineatus Flores, Indonesia. RHK
Thin-lined Cardinalfish Cheilodipterus sp. 1

Undetermined species, apparently undescribed (Thomas Fraser, pers. comm.).
Known from islands and Coron Lake of the Philippines. Colour whitish grey, sides with 3 black and 1 white longitudinal
thin lines, an indistinct short 4th line along dorsal-fin bases. Stripes thickest on head. Estimated length about 8 cm.
A B
Cheilodipterus sp. 1. Calamain Island, Philippines. NBA Cheilodipterus sp. 1. Coron Lake, Philippines. GRA

Toothy Cardinalfish
Cheilodipterus isostigma
Cheilodipterus isostigma Schultz, 1940.
New Guinea.
West Pacific, Philippines to Australia and Fiji in the
South Pacific. Clear coastal to outer reef lagoon
slopes with rich coral growth, usually among staghorn
corals. Similar to Cheilodipterus quinquelineatus in
which mid-lateral line lines up with lower edge of the
tail-spot, whilst in this species it lines up with centre.
This species has large canines in the inner part of the
lower jaw. Length to 11 cm.
A
Cheilodipterus isostigma Maldives. RHK
B C
Cheilodipterus isostigma Maldives. RHK Cheilodipterus isostigma Rowley Shoals, WA. RHK
D
Cheilodipterus isostigma Flores, Indonesia. RHK
Western Toothy-cardinalfish Cheilodipterus cf. isostigma

Undetermined species known from a few photographs, ranging from Java, Indonesia to the Red Sea.
Indian Ocean and Red Sea. Appears to replace Cheilodipterus isostigma in some areas. Differs in line pattern with lowest
ventral line rising at vent towards the pectoral-fin base and the basicaudal black spot is larger. Length to about 11 cm.
A B
Cheilodipterus cf. isostigma Egypt, Red Sea. FLI Cheilodipterus cf. isostigma Pulau Putri, Java, Indonesia. RHK

Dwarf Toothy-cardinalfish
Cheilodipterus pygmaios
Cheilodipterus pygmaios Gon, 1994. Red Sea.
A Red Sea endemic. Various habitats from coastal, silty to
coral outcrops, seen to about 40 m depth. Solitary or in
small aggregations. Colour white with 4 relatively thick
distinct black longitudinal lines along side, a fifth ventrally.
A bright yellow blotch at end of caudal peduncle with a
small black spot centrally and dorsally. Length to 7 cm.
Very similar to Cheilodipterus novemstriatus, but a
A much smaller species and 4th line is of even thickness.
Cheilodipterus pygmaios Egypt, Red Sea. RHK
B C
Cheilodipterus pygmaios Red Sea. SBO Cheilodipterus pygmaios Egypt, Red Sea. RHK
Five-line Cardinalfish
Cheilodipterus novemstriatus
Cheilodipterus novemstriatus Rüppell, 1838. Red Sea.
Known only from the Red Sea and Arabian seas. Found
on protected shallow reefs to about 10 m depth. Usually
seen seeking shelter among long-spined sea-urchins in
small aggregations. Colour whitish with 4 distinct black
longitudinal lines along side, a fifth ventrally. Fourth line
with thickened section over abdomen. Caudal-fin base
with a small black spot centrally, a yellow one above and
A topped by a second small black spot. Length to 10 cm.
Cheilodipterus novemstriatus Red Sea. SBO
B
Cheilodipterus novemstriatus Safaga, Egypt, Red Sea. RHK

Mimic Cardinalfishes – Cheilodipterus spp.

Some fishes mimic other species that are avoided by predators because of defensive properties, such as venomous bites,
which is called Batesian mimicry, Three forms of a Cheilodipterus are known to mimic the venomous sabre-toothed
blennies of the Meiacanthus genus. Besides colour the forms are identical and if they are different species with mimicry
involved is uncertain. In other fishes, often only juveniles mimic different sympatric species, but grow into identical adults.
Mimic Cardinalfish 1
Cheilodipterus nigrotaeniatus
Cheilodipterus nigrotaeniatus Smith & Radcliffe, 1912.
Zamboanga, Philippines.
West Pacific, Philippines to southern Indonesia. Swims
openly about during the day over open substrate. Mimics
the saber-toothed blenny Meiacanthus grammistes in
looks as well as swimming using the tail, rather than the
pectoral fins. Yellowish with 3 black longitudinal thick
stripes and few small spots on caudal fin. Length 10 cm.
A
Cheilodipterus nigrotaeniatus Lembeh Strait, Indonesia. RHK
B C
Model Meiacanthus grammistes Indonesia. RHK Cheilodipterus nigrotaeniatus Lembeh Strait, Indonesia. RHK
Cheilodipterus zonatus
Cheilodipterus zonatus Smith & Radcliffe, 1912.
Rita Island, Philippines.
West Pacific, Philippines, Malaysia and Solomon Islands.
Swims openly about during the day over open substrate.
This form mimics the saber-toothed blenny Meiacanthus
geminatus. Ventrally yellow from below black mid-lateral
stripe, white above and dusky dorsally. Length 10 cm.
Cheilodipterus zonatus Sabah, Borneo. GRA
Cheilodipterus parazonatus
Cheilodipterus parazonatus Gon, 1994.
Zamboanga, Philippines.
West Pacific, Philippines to southern Indonesia. Swims
openly about during the day over open substrate. Mimics
the saber-toothed blenny Meiacanthus vittatus. Similar to
Cheilodipterus zonatus, but ventrally white, midlateral
line is of variable width with a white narrow border or line A
dorsally and back uniformly grey above. Length 10 cm. Cheilodipterus parazonatus Milne Bay, PNG. RHK
B C
Model Meiacanthus vittatus Milne Bay, PNG. RHK Cheilodipterus parazonatus Solomon Islands. SPO

SUBFAMILY APOGONINAE – TRIBE GLOSSAMIINI

Comprises 2 genera: Glossamia and Yarica.
Diagnosis D. VI+I,8–13; A. II,8–16; head and body naked or with large weakly ctenoid or cycloid
scales; LL scales 23–24 or a series of free neuromasts to caudal-fin base, interrupted mid-line lateral
line present or absent; preopercle ridge smooth, edge with one or more spines at angle; 0–2 ossified
supraneurals; supramaxilla, paired uroneurals and basisphenoid absent; 2–3 epurals; hypurals 1–2
fused, 3–4 separate or fused to urostyle; parhypural separate or fused to hypurals 1–2; caudal fin
forked or rounded; pterosphenoid and parasphenoid joined excluding prootic from internal orbit rim.
Other characteristics one supernumerary dorsal-fin spine; unbranched or branched first segmented
dorsal-fin ray, branched first anal-fin ray; cycloid or weakly ctenoid scales on predorsal, cheek, breast,
two pelvic scales, and body; cycloid scale on opercle and onto base of caudal fin; pectoral-fin rays
10–14; 9+8 segmented principal caudal rays, 13–15 branched, upper one or two and lower one or
two unbranched; unbranched procurrent rays, longest segmented; some enlarged teeth on premaxilla
and dentary, canine or villiform on vomer, villiform on palatine; six infraorbitals, bony shelf on third
infraorbital present or absent; 1–2 postcleithra; anterior ceratohyal smooth or notched; 10+14 or
9+15 (Cercamia) vertebrae; 7–8 ribs; 0–6 epipleurals; low crest on PU2 (after Mabuchi et al, 2014).
Genus Glossamia Gill, 1863 – SPECIES ACCOUNTS after Gerald R. Allen.
Fem. Type-species Apogon aprion Richardson, 1842.
Genus Glossamia comprises presently 10 recognised species:
G. aprion.............66 G. gjellerupi.........68 G. sandei .............68 G. wichmanni ......68
G. arguni .............69 G. heurni.............69 G. timika .............67
G. beauforti .........69 G. narindica ........68 G. trifasciata........67
Australian Mouth-almighty
Glossamia aprion
Apogon aprion Richardson, 1842. Port Essington, NT.
Freshwater of tropical Australia and southern-central New
Guinea. A solitary species favouring still environments
such as lake margins and swamps, but it also occurs in
flowing streams. Males orally incubate eggs, numbering
several hundreds, about 3 mm in diameter. Colour greyish
brown with distinct mottled pattern, often with 3 irregular
dark bars on body, one below each dorsal fin and one on
base of caudal fin. Also an oblique bar from snout through
A eye and a conspicuous small black spot in whitish patch
Glossamia aprion Kakadu, Northern Territory. NLI below eye. Length to 20 cm, usually to 10 cm.
B C
Glossamia aprion Daintree River, Queensland. RCS Glossamia aprion Timika, Irian Jaya. GRA
D E
Glossamia aprion Lake Echam, Queensland. PWO Glossamia aprion Juvenile. Lake Bariene, Qld. GRA

Timika Mouth-almighty
Glossamia timika
Glossamia timika Allen, Hortle & Renyaan, 2000.
Minajerwi River system, Papua, Indonesia.
Freshwaters of the Timika area, Irian Jaya. Favours small
streams flowing through primary rainforest. It sometimes
shares this habitat with Glossamia sandei. Similar to G.
trifasciata, but has extra, narrower bars between the
three primary body bars. The pattern is consistant in both
juveniles and adults. Length to 12 cm.
B C
Glossamia timika B juvenile. Timika, Irian Jaya. GRA
Three-bar Mouth-almighty
Glossamia trifasciata
Apogon trifasciatus Weber, 1913.
Lorentz River, Irian Jaya.
Freshwaters of southern New Guinea and Irian Jaya. A
solitary species found in rivers, small creeks and swamps.
Often in streamside vegetation or log jams. A stocky
species, pale brownish with 3 blackish bars, frequently
with bronze, golden yellowish hue on breast and
abdomen. Eggs about 2.5 mm in diameter, hatchlings 10
mm long are well developed, exhibiting the characteristic
body bars of adults. Length to 13 cm. A
B
Glossamia trifasciata B Hatchlings, 10 mm long. Ningurum, PNG. GRA

Slender Mouth-almighty
Glossamia narindica
Apogon narindica Roberts, 1978.
Middle Fly River, Papua New Guinea.
Freshwaters of southern-central New Guinea. A poorly
known fish, less than 20 specimens have been collected,
all from the Bensback and Middle Fly Rivers. Has a more
slender body than its congeners. Occurs in swamps and
along edges of streams among aquatic vegetation and log
debris. Colour brown with silvery reflections on head and
lower sides. Length to 13 cm.
Glossamia narindica Fly River, PNG. GRA
Barred Mouth-almighty Glossamia sandei

Apogon sandei Weber, 1908. Wagani River, Urana basin, southern New Guinea.
Central southern New Guinea between the Puran River, PNG and lake Yamur, Papua. Lowland rivers and small tributaries
to about 100 m elevation, often in rainforest. Congregates among log snags, frequently in roots of large fallen trees. This
species keeps tiny young in mouth for an undetermined period. Juveniles distinctly barred, the pattern fading with growth.
A large species, length to 23 cm.
A B
Glossamia sandei Timika, Irian Jaya. GRA
Wichmann’s Mouth-almighty Glossamia wichmanni

Apogon wichmanni Weber, 1908. New Guinea.
Fresh waters of northern New Guinea. A poorly known species that has been collected in several streams of central-northern
New Guinea between the Bewani Mountains, Papua New Guinea, and the Tawarin River, Irian Jaya. Similar to G. gjellerupi
but generally has fewer bands and are vertical on caudal peduncle. Found solitary among log debris, streamside vegetation
and marsh area on edges of lakes. A large species, growing to about 20 cm.
A B
Glossamia wichmanni A male. B female. PNG. GRA
Gjellerup’s Mouth-almighty
Glossamia gjellerupi
Apogon gjellerupi Weber & de Beaufort, 1929.
Sepik & Mamberamo Rivers, New Guinea.
Freshwaters of northern New Guinea. Most common
Glossamia from the Markham River, Papua New Guinea
to the Mamberamo, Irian Jay, in elevation up to 600 m.
River A solitary species in most habitats. Hatchlings well
developed with a prominent orange yolk-sac. Similar to
Glossamia wichmanni, but usually has more oblique
bands on the body. Length to 18 cm.
Apogon abo Herre, 1936 (described from the Sepik River)
Glossamia gjellerupi Pagwi, PNG. GRA is a junior synonym.

Heurn’s Mouth-almighty Glossamia heurni

Apogon heurni Weber & de Beaufort, 1929. Idenburg & Otker rivers, Mamberamo system, Papua Province, Indonesia.
Mamberamo River to Wapoga River systems of Papua Province. Lowland floodplains in streams, lakes, and swamps. Light
brown to silvery grey with thin oblique black lines on body. Length to 15 cm.
A B
Glossamia heurni A juvenile. Siewa, Wapoga River system. GRA
Beaufort’s Mouth-almighty Glossamia beauforti

Apogon beauforti Weber, 1908. New Guinea.
Fresh waters of northern New Guinea. A poorly known species that has been collected in several streams of central-northern
New Guinea between the Bewani Mountains, Papua New Guinea, and the Tawarin River, Irian Jaya. Similar to G. heurni
but body-lines run longitudinally straight. Found solitary among log debris, streamside vegetation and marsh area on edges
of lakes. A large species, growing to about 20 cm long.
A B
Glossamia beauforti A Lake Sentani. B Dabra Mamberamo. GRA
Arguni Mouth-almighty
Glossamia arguni
Glossamia arguni Hadiaty & Allen, 2011.
Wahisewar River, Arguni Bay region.
West Papua Province, Indonesia. Known only from the
Arguni Bay region’s freshwater streams, which represents
the farthest west record for the genus in the region of
New Guinea and lays approximately 130 km north-west
of Lake Yamur, previous limit of distribution. Colour grey
with a tinge of brown and a pattern of dark-dusky blotching
along the back and forming broad bars below lateral line.
Length to 12 cm.
A
B C
Glossamia arguni A/C adults. B juvenile. Arguni Bay region. GRA

TRIBE GLOSSAMIINI – Genus Yarica Whitley, 1930

Fem. Type-species Apogon torresiensis Castelnau, 1875 (= Apogon hyalosoma Bleeker, 1852).
Genus Yarica is thought to be monotypic.
Yarica is included in this tribe based on morphological data only. Support for combining Glossamia
and Yarica are similar body shapes, freshwater habitat preferences, a large supramaxilla of similar
shape, a single supernumerary dorsal spine and nine epineurals, but it needs confirmation.
The widespread distribution of the single representative Yarica hyalosoma indicates that this genus has
hyaline characteristics imbedded in its life history, able to adapt to a wide range of salinities.
Humpbacked Cardinalfish
Yarica hyalosoma
Apogon hyalosoma Bleeker, 1852. Singapore.
Widespread tropical Indo-West Pacific, Japan to Australia
and west to Africa. Lives in mangrove habitats, entering
freshwater streams. Colour greyish, dusky dorsally silvery
below. Reflective silvery on side of head. Membrane after
2nd spine on dorsal fin black. A peduncular black spot
variable from pupil to eye diameter size, which may relate
to habitat if a single taxon. Distinguished from Glossamia
by the much larger scales, a basicaudal spot and opercle
edges are serrate versus smooth. Length to 17 cm.
Very similar to Fibramia amboinensis of the tribe
A
Zoramiini that also lives in mangrove habitats.
Yarica hyalosoma Aquarium, Townsville, Qld. RHK
B C
Yarica hyalosoma Small basicaudal spot form. Phuket, Thailand. USA
D
Yarica hyalosoma Large basicaudal spot form (Y. torresiensis). Darwin, from small tidal creek. NLI

SUBFAMILY APOGONINAE – TRIBE GYMNAPOGONINI

Comprises 4 genera (key below) and an estimated 25 species.
Diagnosis Dorsal fin VI+I,8–13; anal fin II,8–16; head and body naked or with large weakly ctenoid
or cycloid scales; lateral-line scales 23–24 or lateral line a series of free neuromasts to base of caudal
fin, interrupted mid-line lateral line present or absent; preopercle ridge smooth, edge with one or more
spines at angle; 0–2 ossified supraneurals; supramaxilla, paired uroneurals and basisphenoid absent;
2–3 epurals; hypurals 1–2 fused, 3–4 separate or fused to urostyle; parhypural separate or fused to
hypurals 1–2; caudal fin forked or rounded; pterosphenoid and parasphenoid joined excluding prootic
from internal orbit rim. Other characteristics one supernumerary dorsal-fin spine; unbranched or
branched first segmented dorsal-fin ray, branched first anal-fin ray; cycloid or weakly ctenoid scales on
predorsal, cheek, breast, two pelvic scales, and body; cycloid scale on opercle and onto base of caudal
fin; pectoral-fin rays 10–14; 9+8 segmented principal caudal rays, 13–15 branched, upper one or two
and lower one or two unbranched; unbranched procurrent rays, longest segmented; some enlarged
teeth on premaxilla and dentary, canine or villiform on vomer, villiform on palatine; six infraorbitals,
bony shelf on third infraorbital present or absent; 1–2 postcleithra; anterior ceratohyal smooth or
notched; 10+14 or 9+15 (Cercamia) vertebrae; 7–8 ribs; 0–6 epipleurals; low crest on PU2 (after
Mabuchi et al, 2014).
Key to Genera
1 Scales present ............................................................2
Scales absent.............................................Gymnapogon
2 Second dorsal fin I,8–9 ...............................................3
Second dorsal fin I,12–13 ...........................Lachneratus
3 Anal rays 11–13. ............................................Cercamia
Anal rays 8 or 9 ........................................Pseudamiops
TRIBE GYMNAPOGONINI – Genus Cercamia Randall & Smith, 1988

Fem. Type-species Cercamia cladara Randall & Smith, 1988.
Genus Cercamia has about 7 species: C. cladara, C. eremia, C. melanogaster, and C. spp. nov, some
descriptions are in process. Little known and rarely observed small transparent nocturnal fishes.
Frail Cardinalfish Cercamia cladara
Cercamia cladara Randall & Smith, 1988. Rapa, reef at entrance to Haurei.
West and Central Pacific. Solitary, lives on reef margins in about 10 to 20 m depth. D. VI–I,9; A. II,12–13; P. 10. Body
depth 4.4 (4.2–4.5) in SL (after Randall & Smith, 1988); Scales deciduous. Glassy-transparent, sometimes with tiny reddish
spots, most colour from internal parts, gills red and abdominal area brown and black with silvery, bluish white reflective line,
bending down close to anus. Caudal fin forked with rounded tips. Length to 45 cm.
A B
Cercamia cladara Iriomote Island, Japan. KYA Cercamia cladara. Bali, Indonesia. RHK
C
Cercamia cladara Ishigaki Island, Japan. TKO

Black-belly Cardinalfish
Cercamia melanogaster
Cercamia melanogaster Allen, Erdman & Mahardini, 2015.
West Papua Province, Indonesia.
West Pacific, Papua New Guinea. Solitary, drifting close
to substrate near reefs at night in depths of about 10 to
40 m. D, VI–I,9; A. II,11–13; P. 9–11. Body glass-like
transparent, red band on snout and behind eye, stomach
A region black with a thin white line from top of eye towards
anus. Eyes dusky yellow. Length to 45 mm.
Cercamia melanogaster Raja Ampat Islands, West Papua. GRA
B
Cercamia melanogaster West Papua Province, Indonesia. GRA
Red-face Cardinalfish Cercamia sp. 1

Undescribed Cercamia from Palau and Indonesia.
A description is in preparation (Fraser, pers. comm.).
West Pacific, from Palua and Indonesia to Papua New
Guinea. Solitary, drifting close to substrate near reefs at
night in about 10 to 30 m depths. Glass-like transparent,
head bright red, stomach region black with a white line
from upper of opercle towards anus. Eyes black. Length
to about 45 mm.
A
Cercamia sp. 1. Manado, Indonesia. RHK
B
Cercamia sp. 1. Milne Bay, PNG. PWO

Glassy Cardinalfish Cercamia eremia

Cercamia eremia Allen, 1987. South Murion Island, WA.
Appears to be widespread in the West Pacific, records
elsewhere based on similar taxa. Drifts close to substrate
near reefs at night in 2 to 40 m depths. D. VI–I,9; A.
II,12–13; P. 9–11. Semi-transparent, showing very little
pigmentation with orange chromophores concentrated on
head and scale-margins on body. Silvery reflective on
abdomen. Length to 45 mm. A
Cercamia eremia Holotype. GMO
B C
Cercamia eremia Raja Ampat Islands, West Papua. GRA Cercamia eremia Flores, Indonesia. RHK
Delicate Cardinalfish Cercamia sp. 2

Undetermined species, apparently new. A description is
in preparation (Bogorodsky, pers. comm.).
Red Sea, very similar to Cercamia melanogaster, having
the same black abdomen and bluish white line, but with
the presence of numerous tiny orange chromophores as
shown in C. eremia. An orange broad band over head, on
snout and behind eye. Length to about 45 mm.
B C
Cercamia sp. 2. Dahab, Red Sea. SBO Cercamia sp. 2. Al Wajh, Red Sea. SBO
D
Cercamia sp. 2. A & D Obhur, Red Sea. SBO

TRIBE GYMNAPOGONINI – Genus Gymnapogon Regan, 1905

Masc. Type-species Gymnapogon japonicus Regan, 1905.
Genus contains ten nominal species: Gymnapogon annona, G. foraminosus, G. janus, G. japonicus, G.
urospilotus and G. vanderbilti have 9 or 10 soft dorsal, 9 or 10 anal rays and 12 to 14 pectoral rays, the
lower two and upper two unbranched. G. africanus, G. melanogaster, G. philippinus and G. sagittarius
have 9 soft dorsal rays, the first ray unbranched and 8 soft anal rays, the first ray branched. G. japonicus
and G. velum have a rounded caudal fin with 13, 14 or 15 branched principal caudal rays. G. africanus,
G. annona, G. melanogaster, G. philippinus, G. sagittarius, G. vanderbilti have a forked caudal fin with 13
branched principal caudal rays and 2 unbranched upper and lower principal caudal rays. A single bifid-like
preopercle spine is shared by Gymnapogon annona, G. melanogaster and undescribed forms. The presumed
wide-spread B-marked species complex made up of G. urospilotus, G. vanderbilti and other undescribed
forms have at least one small upper preopercle spine in addition to the larger single spine near the angle of
the preopercle (after Fraser, 2016). Six named species are included, but some identifications are tentative.
G. urospilotus may be synonymus with G. vanderbilti or is restricted to the Marshall islands region in the Pacific.
Several species have been photographed in their larval or post-larval stage, about 15–20 mm long, showing
greatly elongated ventral fins and the body with reddish pigmentation forming diffused blotching. It seems
likely that all members of the B-spot complex have these features that change to the reduced fins and plain
transparent whitish appearance once settling on the substrate. Pelagic larval fishes often have long filaments
that mimic stinging cells of other planktonic creatures such as jellyfish or look like some other potentially
harmful creature. Little is known about the geographical distribution of each species.
It is apparent that a number of taxa are new to science. The genus is under review by Thomas Fraser.
Japanese Naked-cardinalfish
Gymnapogon japonicus
Gymnapogon japonicus Regan, 1905.
Inland Sea, Japan.
West Pacific, Southern Japan and Philippines. Secretive,
usually only collected with chemicals. Semi-transparent
whitish. The adult looks superficially somewhat like a
goby. Caudal fin rounded. Length to 9 cm.
Gymnapogon japonicus Shimane Prefecture, Japan. TSU
Philippines Naked-cardinalfish
Gymnapogon philippinus
Henicichthys philippinus Herre, 1939.
Luzon Island, Philippines.
Western Pacific: Indonesia and Philippines north to
Taiwan and Ryukyu Islands, south to New Caledonia.
Shallow water, including tidepools. Plain semi-transparent
whitish. Rounded snout. Caudal fin forked with rounded
lobes. Length to 9 cm.
Gymnapogon philippinus Ambon, Indonesia. ARY
Australian Naked-cardinalfish
Gymnapogon annona
Gymnapogon annona Whitley, 1936.
Lindeman Island, Queensland, Australia.
West Pacific, reported from Queensland and probably
occurs in Indonesia and Papua New Guinea. Secretive,
usually only collected with chemicals from shallow coastal
reefs. Length to about 50 mm.
Gymnapogon annona Holotype. after Whitley
African Naked-cardinalfish
Gymnapogon africanus
Gymnapogon africanus Smith, 1954. Mozambique.
Western Indian Ocean species. Smith's description of
Gymnapogon africanus in part: Transparent, colour
being visible: only the pupil, the aliments, a red arterial
axial streak and melanophores chiefly on the occiput with
a group of some on the snout. Shallowly forked caudal
Gymnapogon africanus Oman. JER fin. Length to about 50 mm.

Slender Naked-cardinalfish Gymnapogon sagittarius

Gymnapogon sagittarius Yoshida, Kawai & Motomura, 2019. Ha Long Bay, Vietnam.
Only known from a few trawled specimens off the Andaman Sea, Malaysia, Vietnam and east to Papua New Guinea. The
species is named sagittarius in reference to its slendernesss. Caudal fin is forked. Length to about 70 mm.
A B
Gymnapogon sagittarius A Milne Bay. Papua New Guinea. B Raja Ampat, West Papua. GRA
Highfin Naked-cardinalfish
Gymnapogon velum
Gymnapogon velum Fraser, 2019. Western Australia
Only known from two trawled specimens off northern
west and east Australia in 50–60 m depth. The species is
named velum (Latin: sail) in reference to the long and
deep dorsal and anal fins. Caudal fin is rounded. Length
to about 60 mm.
Gymnapogon velum Holotype. Fins partly reconstructed. CSIRO
B-spot Naked-cardinalfish
Gymnapogon vanderbilti-complex
Gymnapogon vanderbilti Fowler, 1938. Kiritimati.
Widespread Indo-West Pacific, but comprises a species
complex. Its members sharing the B-shaped basicaudal
spot. Secretive, usually only collected with chemicals from
shallow coastal reefs. Pelagic larval stages have very long
ventral fins and some of these may have long extended A
rays. Length to about 50 mm.
Gymnapogon vanderbilti Adult. Flores, Indonesia. RHK
B C
Gymnapogon sp. Marquesas Isands. SMI Gymnapogon sp. Flores, Indonesia. RHK
D E
Gymnapogon sp. Philippines. LWI Gymnapogon sp. Comores Islands. RWI
F G
Gymnapogon sp. Mauritius. PCH Gymnapogon sp. Post-larval. Comoros Islands. RWI

Gymnapogon spp. – pelagic stages

Undetermined taxa photographed at night in open water. The long fan-like ventral fins have 5 principal rays that are
branched close to the base, headed by a feeble spine, giving the false impression of having 11 fin rays.
A B
Gymnapogon sp. 1. Post-larval. Japan. HKO Gymnapogon sp. 1. Pelagic stage. Japan. RMI
C D
E
Gymnapogon sp. 2. Pelagic stage. Japan. RMI

F
Gymnapogon sp. 3. Pelagic stage. Solomon Islands. MSC
G
Gymnapogon sp. 4. Pelagic stage. Coral Sea. GED
TRIBE GYMNAPOGONINI – Genus Pseudamiops Smith, 1954

Masc. Pseudamiops pellucidus Smith, 1954.
Genus comprises 5 species: P. diaphanes, P. gracilicauda, P. pellucidus, P. phasma, and P. springeri.
Transparent, small larval-like fishes sheltering in caves and ledges during the day and rarely noticed.
Specimens are usually collected when using chemicals. Caudal fins rounded.
Limpid Cardinalfish
Pseudamiops pellucidus
Pseudamiops pellucidus Smith, 1954.
Malindi, Kenya, western Indian Ocean.
Western Indian Ocean: South and eastern Africa to
Madagascar. A slender species, long caudal-peduncle and
caudal-fin strongly rounded. Length to about 50 mm.
Pseudamiops pellucidus After Smith

Slender-tail Cardinalfish
Pseudamiops gracilicauda
Gymnapogon gracilicauda Lachner, 1953.
Bikini Atoll, Marshall Islands.
Indo-West Pacific: Cocos-Keeling and Christmas islands
(eastern Indian Ocean) east to Marshall Islands and Rapa,
north to the Ryukyu Islands, south to northern Australia
and New Caledonia. Transparent, with slight pinkish hue.
This species has a very long caudal peduncle and a large
Pseudamiops gracilicauda Guam, Micronesia. DBU rounded caudal fin. Length to about 50 mm.
Transparent Cardinalfish
Pseudamiops diaphanes
Pseudamiops diaphanes Randall, 1998.
Makaha, Oahu Island, Hawaiian Islands.
Central Pacific: Johnston Atoll and Hawaiian Islands.
Glass-like transparent except for internal organs and the
blackish vertebral column. Rarely seen during the day.
Drifts high above the bottom at night in the pursuit of
zooplankton. Length to about 40 mm.
Pseudamiops diaphanes Hawaiian Islands. JER
Marquesas Naked-cardinalfish Pseudamiops phasma

Pseudamiops phasma Randall, 2001. Nuku Hiva, Taiohae Bay, Marquesas Islands.
Distribution: Marquesas Islands. Length to about 40 mm.
Pseudamiops phasma Marquesas Islands. SMI
Springer’s Cardinalfish
Pseudamiops springeri
Pseudamiops springeri Gon & Bogorodsky, 2013.
Gulf of Aqaba, Egypt.
Red Sea endemic, collected from Gulf of Aqaba, Egypt
and Al Wajh, Saudi Arabia. All specimens were collected
as solitary from deep crevices in the reef in depths of
5–18 m. Length to 40 cm.
Pseudamiops springeri Al Wajh, Saudi Arabia. SBO
TRIBE GYMNAPOGONINI – Genus Lachneratus Fraser & Struhsaker, 1991

Masc. Type-species Lachneratus phasmaticus Fraser & Struhsaker, 1991. Monotypic.
Phantom Cardinalfish
Lachneratus phasmaticus
Lachneratus phasmaticus Fraser & Struhsaker, 1991.
Hawaiian Islands.
Indo-West Pacific: Comores, Mauritius (Mascarenes), Fiji,
Marquesas Islands, Hawaiian Islands; probably more wide-
spread. A rarely seen species, typically lives in the back of
long tube-like caves. One specimen is known from about
100 m depth. Semi-transparent, pinkish red. Stomach
black with a white line. Length to about 50 mm.
Lachneratus phasmaticus Hawaii. JHO

SUBFAMILY APOGONINAE – TRIBE LEPIDAMIINI

Comprises the single genus. Lepidamia Gill, 1863.
Diagnosis D. VII(I)-I,9 or VIII-I,9; A II,8; head and body with ctenoid scales; pored lateral-line scales
35–48; preopercle ridge smooth, edges serrate; three supraneurals; supramaxilla absent; basisphenoid
present; one pair of reduced uroneurals present; three epurals; five free hypurals; parhypural separate;
two autogenous haemal spines; two supernumerary dorsal spines, caudal fin forked.
Other characteristics first fin ray in second dorsal fin branched and segmented; first anal-fin ray
branched and segmented; ctenoid scales on predorsal, cheek, breast, four pelvic scales, and body;
ctenoid scales on opercle and onto base of caudal fin; pored lateral-line scales with multiple pores; 15
branched, upper and lower unbranched; developed gill rakers 8–17; unbranched procurrent rays,
longest segmented; 10+14 vertebrae; 8 ribs; 9 epipleurals; teeth in multiple rows on premaxilla,
dentary, vomer, palatine, all villiform; six infraorbitals, bony shelf on third infraorbital; stomach and
intestine pale, peritoneum pale; low crest on PU2 (after Mabuchi et al, 2014). Lepidamia has been
collected only from continental marine waters from South Africa to China (Gon, 1995).
TRIBE LEPIDAMIINI – Genus Lepidamia Gill, 1863

Fem. Type-species Apogon kalosoma Bleeker, 1852.
Genus Lepidamia comprises 4 nominal species: L. kalosoma, L. multitaeniata, L. natalensis and L.
omanensis. As species juveniles have distinct colour patterns, but adults are similar.
Pin-striped Cardinalfish Lepidamia kalosoma

Apogon kalosoma Bleeker, 1852. Bangka, Indonesia.
Known from Indonesia, ranging from Sumatra’s northeast
coast to Borneo and Taiwan. Lives in caves and ledges on
shallow coastal reef in 3–15 m depth. Pre-dorsal scales
6–8. LL 36. P. 15. Pale grey to pinkish brown with many
dusky to brown narrow lines long scale-rows, in juveniles
some lines are thickened mid-laterally and along upper
sides, with a black spot on centre of caudal-fin base. Fins
pink to red in adults. Length to 14 cm.
Adults have short median fins, similar in size to
Lepidamia omanensis and L. natalensis. Bleeker’s L.
noordzieki is more juvenile and a putative synonym, but
A
the two taxa may comprise Indian and Pacific siblings.
Lepidamia kalosoma Brunei. GRA
B C
Lepidamia kalosoma Brunei. GRA Lepidamia kalosoma Taiwan. WCC
D E
D Lepidamia kalosoma Banka, NE Sumatra. E L. noordzieki Java, south coast. After Bleeker

Natal Cardinalfish Lepidamia natalensis

Apogon natalensis Gilchrist & Thompson, 1908.
KwaZulu-Natal, South Africa. Western Indian Ocean. The
juveniles may have a distinctive pattern, but adult is much
like Lepidamia omanensis and L. kalosoma. Pre-dorsal
scales 2–5. LL. 40–47. P. 15. Has four scale-rows or
more above lateral line and up to 12 below the highest
point. Adult colour red with brownish dusky thin lines
along scale-rows. Fins deep red, smallish and rounded.
Length to 18 cm.
Lepidamia natalensis KwaZulu-Natal, South Africa. DKI
Oman Cardinalfish Lepidamia omanensis

Apogon (Lepidamia) omanensis Gon & Mee, 1995.
Masirah Island, Oman.
Northwestern Indian Ocean: Oman and Red Sea. Shallow
coastal reefs, tidepools to about 10 m depth, staying close
to shelter during the day. Red with dusky thin lines along
scale rows. Pre-dorsal scales 3–5. LL 43–48. P 16–17.
Up to 12 scale-rows below lateral line. Juveniles have
three black stripes along sides, upper continuing along
caudal peduncle. Pupil-size black spots on dorsal-fin
bases, anal-fin base and another smallish on caudal-fin
A base. Fins red in adults. Caudal fin with dusky margin in
Lepidamia omanensis Oman. PWO adults. Length to 18 cm.
B C
Lepidamia omanensis Juvenile. Shuway'ir, Oman. JER Lepidamia omanensis Juvenile. Oman. PWO
Small-scale Cardinalfish
Lepidamia multitaeniata
Apogon multitaeniatus Cuvier, 1828. Red Sea.
Endemic to the Red Sea and Gulf of Aden. Shallow in few
metres depth, living in dark holes and reef-crevices. Red
with this dusky lines along scale rows. Pre-dorsal scales
3–6. LL 36–40. P. 14. Up to 9 scale-rows below lateral
line. Juveniles have three black stripes along sides, a black
spot below each dorsal-fin base, and a large blotch on the
centre of the peduncle and caudal-fin base. Fins deep red
A in adults. Length to 16 cm.
B C
Lepidamia multitaeniata C juvenile. Farasan Island. Red Sea. SBO

Malan Cardinalfish Lepidamia cf. omanensis

Northern Indian Ocean along continental margin from
Djibouti, India, Pakistan to south coast of Java. Mainly
known from relatively shallow coastal trawls or line-fishing
in bays. Pinkish with reddish brown thin lines along scale
rows. Subadults have a black mid-lateral stripe and a
pupil-size black spot on centre of caudal-fin base. Upper
stripe on 2nd dorsal fin base into spot on posterior end.
Fins red in adults, and first dorsal-fin develops a black
blotch. Pectoral axil dusky to black. Length to 18 cm.
Undetermined taxon, possibly is a Lepidamia omanensis A
variation or it is Bleeker’s L. noordzieki. Lepidamia cf. omanensis Maskali Island, Djibouti. JER
B C
Lepidamia cf. omanensis Trawled, large adults. Pakistan. MEH
D E
Lepidamia cf. omanensis Oman. JER Lepidamia cf. omanensis Oman. JER
F G
Lepidamia cf. omanensis F juvenile. G subadult. Hurghada, Red Sea. DKI
H I
Lepidamia cf. omanensis Shuway'ir, Oman. JER Lepidamia cf. omanensis India. JER

SUBFAMILY APOGONINAE – TRIBE OSTORHINCHINI

Comprises the single genus Ostorhinchus.
Diagnosis D. VI+I,9 or VII+I,9; A II,8–9; head and body with ctenoid scales; pored lateral-line scales
6–26; preopercle ridge smooth, edges serrate; 2–3 supraneurals; supramaxilla absent; basisphenoid
present; one pair of reduced uroneurals present or absent; three epurals; five free hypurals or 1–2
fused; parhypural separate; caudal fin forked; head and/or body with one or more light or dark stripes,
bars rarely present, basicaudal spot or broad mark present or absent. Other characteristics one or
two supernumerary dorsal spines; branched first segmented fin ray in second dorsal-fin; first anal-fin
ray branched and segmented; ctenoid scales on predorsal, cheek, breast, two pelvic scales, and body;
ctenoid scale on opercle and onto base of caudal fin; pored lateral-line scales with multiple pores;
pectoral fin-rays 11–16; three supraneurals; 9+8 segmented principal caudal rays, 15 branched, upper
and lower unbranched; unbranched procurrent rays, longest segmented; teeth in one or multiple rows
on premaxilla, dentary, vomer, palatine, all villiform or absent on palatine, sometimes enlarged, but not
canine on side of dentary; six infraorbitals, bony shelf on third infraorbital; anterior ceratohyal smooth
or notched; developed gill rakers 10–26; 10+14 vertebrae; 8 ribs; 8 epipleurals; blackish stomachs and
intestines; low crest on PU2 (after Mabuchi et al, 2014).
TRIBE OSTORHINCHINI – Genus Ostorhinchus Lacepède, 1802

Fem. Type-species Ostorhinchus fleurieu Lacepède, 1802.
Presently more than 95 species are recognised in the genus:
O. angustatus............120 O. hoevenii ...............127 O. properuptus..........106
O. aphanes................103 O. holotaenia ............109 O. pselion .................103
O. apogonoides ...........83 O. ishigakiensis .........127 O. quadrifasciatus .......92
O. aroubiensis ...........118 O. jenkinsi...................96 O. regula...................118
O. aterrimus .............117 O. kiensis ....................94 O. relativus ...............122
O. atrogaster ...............99 O. komodoensis ..........83 O. rueppellii..............103
O. aureus ....................86 O. leptofasciatus .........97 O. schlegeli ...............112
O. brevispinis ............118 O. leslie ....................109 O. sealei....................123
O. bryx........................95 O. limenus ................115 O. selas.....................103
O. capricornis..............86 O. lineomaculatus .....102 O. semilineatus ...........96
O. cavitensis..............109 O. luteus...................107 O. septemstriatus ........93
O. chalcius ................148 O. maculiferus ..........126 O. sinus ....................122
O. cheni ....................113 O. margaritophorus...122 O. taeniophorus ........115
O. chrysopomus ........122 O. melanoproctus......101 O. unitaeniatus............97
O. chrysotaenia .........111 O. microspilos .............85 O. victoriae ...............116
O. cladophilos .............99 O. moluccensis..........125 O. victoriae aff. .........116
O. compressus...........121 O. monospilus...........125 O. wassinki ...............107
O. cookii ...................116 O. multilineatus ........121 O. yamato ...................99
O. cyanosoma ...........104 O. nanus .....................99 O. sp. 1.....................102
O. cyanotaenia ..........111 O. neotes ..................100 O. sp. 2.....................102
O. dispar...................101 O. nigripes ................128 O. sp. 3.....................102
O. doederleini ...........114 O. nigrocinctus ............87 O. sp. 4.....................105
O. elizabethae .............90 O. nigrofasciatus .......119 O. sp. 5.....................105
O. endekataenia........111 O. norfolcensis ............87 O. sp. 6.....................106
O. fasciatus .................89 O. notatus ...................97 O. sp. 7.....................107
O. flagelliferus ..........126 O. novemfasciatus.....120 O. sp. 8.....................107
O. flavus .....................86 O. ocellicaudus............98 O. sp. 9.....................108
O. fleurieu...................85 O. oxina....................124 O. sp. 10...................110
O. franssedai.............101 O. oxygrammus.........123 O. sp. 11...................113
O. fukuii ...................112 O. parvulus ...............100 O. sp. 12...................118
O. griffini ..................126 O. pallidofasciatus ....117 O. sp. 13...................118
O. gularis ....................99 O. pallidus ..................98
O. hartzfeldii.............110 O. pleuron...................90
The genus Ostorhinchus comprises many clades that may be assigned to subgenera or other genera in the future.
The status of many species remain uncertain, often forming complexes comprising similar members. Many of the
treated synonyms may need to be revisited as some names, such as Apogon fasciatus, or A. cyanosoma, were
commonly applied to similar looking, but different species as a ‘catch all’ and as widespread Indo-Pacific taxa, but
many of the complex-members appear to have a limited geographical range and due to erroneous synonymy some
of the recently described species were possibly named a long time ago.

Short-tooth Cardinalfish
Ostorhinchus apogonoides
Cheilodipterus apogonoides Bleeker, 1856.
Manado, Sulawesi, Indonesia.
Widespread Indo-West Pacific. Inshore, harbours and
lagoons to outer reef habitats along drop-offs in caves to
about 50 m depth, usually in pairs. Juveniles expatriate
to subtropical water, commonly found in the Sydney
region among boulders on rocky reefs. Plain yellow to
orange, greyish dorsally. Double blue line from snout
though eye over head followed by blue spots scattered
centrally on body, black or brown on snout between the
lines. Length to 10 cm. A
Ostorhinchus apogonoides Sulawesi, Indonesia. RHK
B C
Ostorhinchus apogonoides Sydney, Australia. RHK Ostorhinchus apogonoides Maldives. RHK
D E
Ostorhinchus apogonoides Bali, Indonesia. RHK Ostorhinchus apogonoides Flores, Indonesia. RHK
Komodo Cardinalfish Ostorhinchus komodoensis

Apogon komodoensis Allen, 1998. Komodo Islands, Indonesia.
West Pacific, known from Indonesia to the Philippines. Occurs in small groups on sheltered reefs with branching corals in
5–12 m depth. Pale whitish yellow to coppery brown, greyish dorsally ending in black on caudal peduncle where abruptly
changing to white on caudal-fin base. A pair of blue-white lines on snout through eyes, black on snout in between, and a
narrow red vertical streak on opercle margin. Length to 8 cm.
A B
Ostorhinchus komodoensis Komodo, Indonesia. A GRA B TTO

Ring-tail Cardinalfish
Ostorhinchus aureus
Ostorhinchus aureus Lacepède, 1802.
Mascarenes, southwestern Indian Ocean.
lagoons to outer reef habitats to about 30 m depth.
Small juveniles solitary. Adults in pairs or form large
schools in caves and in the water column above outcrops
in tidal zones to feed on zooplankton when the currents
are running. Colour orange-yellow, black nose stripe
running over eye with blue edges, grey dorsally and large
adults have a broad black basicaudal bar usually with
concave sides. Length to about 13 cm.
Brood comprises numerous tiny eggs. Hatchlings poorly
developed and have a long pelagic stage, expatriating
A over vast geographical regions by surface currents.
C
B Ostorhinchus aureus Juvenile. Sydney, Australia. RHK
D E F
Ostorhinchus aureus A pair. B/D male with fresh oral brood. E/F brood near hatching stage. Bali, Indonesia. RHK
G H
Ostorhinchus aureus G in cave. Milne Bay, PNG. H feeding high up in water column. Gilimanuk, Bali, Indonesia. RHK

Flower Cardinalfish
Ostorhinchus fleurieu
Ostorhinchus fleurieu Lacepède, 1802. Pacific Ocean.
lagoons to outer reef habitats to about 35 m depth. Both
juvenile and adults may form large schools. Adults often
high above outcrops during the day in tidal currents
when feeding on zooplankton. Pinkish to yellow with a
black stripe on nose through eye with bluish white lines
as borders, and a black basicaudal spot, variable from
pupil to eye-size, sometimes forming a band. This
species is virtually identical to Ostorhinchus aureus, but
in O. aureus the black peduncular band is larger with
concave sides. When sympatric in areas the variations in
the band can make it difficult to distinguish between
them (E). Length to about 13 cm.
A
Ostorhinchus fleurieu Durban, South Africa. RHK
B C
Ostorhinchus fleurieu Durban, South Africa. RHK Ostorhinchus fleurieu Aliwal Shoal, South Africa. RHK
D E
Ostorhinchus fleurieu Oman. RFI Ostorhinchus fleurieu or O. aureus Flores, Indonesia. RHK
Micro-spot Cardinalfish Ostorhinchus microspilus

Apogon microspilus Allen & Randall, 2002. Lombok, Indonesia.
Indonesia, known from a few specimens in the Lombok area. On reef margins in about 20 m depth. Overall pale orange,
reddish on cheek, a miniscule spot on centre of caudal-fin base and a pair of blue-silver lines on snout and through eye.
A B
Ostorhinchus microspilus Lombok, Indonesia. A TTO B JER

Capricorn Cardinalfish
Ostorhinchus capricornis
Apogon capricornis Allen & Randall, 1993.
One Tree Island, Queensland, Australia.
East coast of Australia from tropical southern
Queensland to Montague Island, in southern NSW.
Sheltered rocky or coral reef habitats, shallow coastal to
deep offshore. Adults usually in pairs or small groups.
Small juveniles form aggregation in caves. Similar to
Ostorhinchus apogonoides, but the sides have narrow
yellow bars and also has a pupil-sized black basicaudal
spot. Length to about 10 cm.
A
Ostorhinchus capricornis Keppel Island, Queensland. RHK
B C
Ostorhinchus capricornis From type-locality. RHK Ostorhinchus capricornis Juvenile. Sydney. RHK
D E
Ostorhinchus capricornis D adult. E small juveniles in cave. Montague Island, southern NSW, Australia. RHK
Lemon-belly Cardinalfish Ostorhinchus flavus

Apogon flavus Allen & Randall, 1993. Norfolk Island.
Southwestern Pacific, Norfolk and Lord Howe Islands.
Schools on shallow rocky and coral reefs to about 25 m
depth. Plain brownish to orange-yellow, a dusky band
on snout and a small black basicaudal spot. Similar to
Ostorhinchus capricornis, but has more black on lips
and is less ornamental on head. Length to 10 cm.
A B
Ostorhinchus flavus Lord Howe Island. A ESC B MPF

Black-rim Cardinalfish Ostorhinchus nigrocinctus

Apogon nigrocinctus Smith & Radcliffe, 1912. Philippines.
West Pacific, northern Australia to Philippines. Mainly
known from trawls, but observed schooling near large
rocky outcrops at 30 m in a current prone channel. Grey
with pale pinkish sides, a narrow black bar on caudal-fin
base with small spot in centre. Length to about 85 mm.
A B
Ostorhinchus nigrocinctus Lembeh Strait, Sulawesi, Indonesia. RHK
Norfolk Cardinalfish Ostorhinchus norfolcensis

Apogon norfolcensis Ogilby, 1888. Norfolk Island.
Southwestern Pacific, Norfolk Island, Lord Howe Island and New Caledonia. Locally abundant in lagoons and on reefs, may
form large schools. Plain brownish, scales above lateral line with dusky spots, along belly with pale spots. Whitish banding
on head, below dorsal-fins gap, and on caudal peduncle. A pupil-size black basicaudal spot. Length to 10 cm.
A B
C
Ostorhinchus norfolcensis A Lord Howe Island. B adult. C schooling. Norfolk Island. A/C MPF B RCS

Striped Cardinalfishes of the Ostorhinchus fasciatus/kiensis groups

There is much confusion to the identity among the basically grey species with a black mid-lateral stripe from tip of snout to
end of tail and a black stripe above that runs from nape above eye along upper sides ending close or on caudal peduncle.
There are many taxa that fit in this pattern. The first was named Mullus fasciatus by Shaw in White 1790 from the Sydney
subtropical region and the name Apogon fasciatus has long been used as a ‘catch-all’ name for many species with similar
patterns occurring throughout the tropical Indo-Pacific. Currently recognised “world experts” considered A. fasciatus as
widely distributed. Fraser, 2005 listed nine nominal taxa in the group: Apogon fasciatus (Shaw in White, 1790), Apogon
quadrifasciatus Cuvier in Cuvier & Valenciennes, 1828, Apogon monogramma Günther, 1880, Apogon septemstriatus
Günther, 1880, Apogon evanidus Fowler 1904, Apogon elizabethae (Jordan & Seale, 1905), Apogon quinquestriatus
Regan, 1908, Apogon kiensis Jordan & Snyder, 1901 and Apogon bryx Fraser, 1998. The two latter species, of the O.
kiensis complex have VI first-dorsal spines, whilst all others have VII. Two of the listed taxa, A. monogramma and A.
evanides do not belong in the O. fasciatus-group, whilst the others are treated valid species here. Now placed in
Ostorhinchus, many additional species in the group, that have clearly different striped-patterns remain undescribed. Like
the sympatric O. limenus, O. fasciatus is part of a fauna which has a limited sub-tropical geographical distribution in New
South Wales and southern Queensland. O. fasciatus is not a putative widespread tropical species as suggested by Fraser,
2005. He examined many specimens in the fasciatus group from numerous Indo-West Pacific locations, but worked
with preserved material that lost much of the essential diagnostic colour patterns, resulted in lumped them as fasciatus based
on a criteria of meristics, the counts of fin-ray and gill-rakers, but none of these taxa have matching patterns with the Sydney
O. fasciatus (shown below). Striped species show little variation, but some can be expected. However, when the stripes run
in different positions on the head and body or have features that are never shown in others, they are not just variations. Of
the ‘O. fasciatus’-group, only the taxon O. quadrifasciatus appears to be a widespread species in the Indian Ocean and
Red Sea, which is commonly misidentified and illustrated as O. fasciatus, due to the assumed synonymy.
Many cardinal fishes are highly localised, which is related to oral brooding and the level of parental care. The species that
produce few and highly developed eggs, some may even take care of their offspring for a short period, are highly localised
and endemic to very small areas. Minimal differences on a genetical level between populations of the same species may
occur because of oral brooding and their limited larval dispersal. In contrast, species such as Ostorhinchus aureus produce
numerous tiny eggs that hatch less developed fry may have a long planktonic pelagic stage and consequently are dispersed
over vast geographical regions. Whilst based on distinctive differences in the striped pattern it is clear that the O. fasciatus
group comprises many different species, that may need to be backed up with a DNA study to satisfy the skeptics. Even
among the species shown here with multiple images it is likely that more cryptic taxa are present. Unpublished molecular
studies revealed that in the China Seas there are at least 4 potential ‘fasciatus’ species present (Bogorodsky, pers. comm.).
Typical adult Ostorhinchus fasciatus Freshly preserved specimen. Vaucluse Bay, Sydney Harbour. RHK
A grey fish, whitish ventrally. The black midlateral-line has a white line edging ventrally which the lacks bars or spots below,
seen in several tropical taxa, but may have a faint dark stripe there instead. It has a thin black line from the top of the eye
over the straight section lateral-line which is not present in most other ‘fasciatus’ species. A black line from nape runs curved
along contour of back above the lateral-line on upper sides, ending dorsally on caudal peduncle to fin-base. It has a thin
white line above, but lacks the black line above along dorsal-fin bases present in some similarly striped species.

Australian Striped Cardinalfish

Ostorhinchus fasciatus
Mullus fasciatus Shaw, 1790. Sydney, Australia.
East coast of Australia only. Inshore, harbours and muddy
bays, shallow to 50 m. Rocky reefs, schooling or in
pairs. Pale grey with black lines. Mid-lateral line straight,
from snout, through eye, bordered below by a white line,
to caudal-fin margin. Juveniles have black basicaudal
spot, which with growth is absorbed by mid-lateral line.
A long line from over nape along upper sides to caudal
peduncle. Usually a thin short line from above eye on
the lateral line to below start of 2nd dorsal fin. No black
line along dorsal-fin bases. Length to about 15 cm. A
B C
E F
Ostorhinchus fasciatus E juvenile. Sydney Harbour. F male with oral brood. Port Stephens. A–E RHK F DHA

Elizabeth’s Cardinalfish
Ostorhinchus elizabethae
Apogon elizabethae Jordan & Seale, 1905. Hong Kong.
Coastal China Seas. Grey dorsally with black and white
lines. Mid-lateral line from tip of snout, through eye to
caudal-fin margin, usually slightly curved up centrally,
ventrally with white and dark bars. No short line from
above eye on to the lateral line. A black line from over
nape to caudal peduncle. No black line along dorsal-fin
bases. A white thin line closely above each black line.
Fins marked as in type, ventral-tip dusky. Length to 9 cm.
Not a synonym of Apogon fasciatus (Shaw, 1790), but
A Ostorhinchus pleuron (Fraser, 2005) looks identical and
Ostorhinchus elizabethae Type-locality, Hong Kong. RYU appears to be synonymus.
B C
Ostorhinchus elizabethae Type illustration. After J&S Ostorhinchus elizabethae Hong Kong. RCH
D E
Ostorhinchus elizabethae (O. pleuron paratype) Vietnam. RWI Ostorhinchus elizabethae Hong Kong. KUC
Indonesian Striped Cardinalfish

Ostorhinchus cf. fasciatus-1
Undetermined species.
Komodo to West Papua, Indonesia, and Philippines. On
open muddy substrates in small groups, pairs or single.
Grey with thick black lines. Mid-lateral line straight, from
snout, through eye, to caudal-fin margin, bordered by
white lines, lower with sparse barring below. Dusky line
from upper of eye to descend of lateral line and a line
from over nape to caudal peduncle. A black line above,
with white thin line in between black along dorsal-fin
A bases. Blue basally in anal fin. Length to about 12 cm.
B C
Ostorhinchus cf. fasciatus-1. Maumere, Flores, Indonesia. RHK Ostorhinchus cf. fasciatus-1. Philippines. ARY

Mud-Bay Striped Cardinalfish

Only known from Tomini Bay, Sulawesi, Indonesia. On
open muddy substrates among forests of seawhips at 25
m depth, in small to large groups. Grey with black and
white lines. Mid-lateral line straight, from snout, through
eye, to caudal-fin margin. A thinner straight line low on
upper side, from nape sloping downwards to upper of
caudal peduncle. No line above along dorsal-fin bases.
Female has short white barring below black mid-lateral
line. Length to about 10 cm.
A
B C
D
Ostorhinchus cf. fasciatus-2. A males. B juvenile. C male. D pair, female on right. Tomini Bay, Sulawesi, Indonesia. RHK
Lembeh Striped Cardinalfish

Only observed in Lembeh Strait at 30 m near large
rocks, in strong current zone. Grey with three black
lines, white lines and white barring. Mid-lateral line
straight, thick on snout and through eye, to posterior
margin of caudal-fin and white barring below. A black
line from snout over above eye to upper of caudal
peduncle with thin white line above. A black line from
mid-nape along the dorsal-fin bases. Length to 12 cm.
A
B C
Ostorhinchus cf. fasciatus-3. Lembeh Strait, Sulawesi, Indonesia. RHK

Pearly-striped Cardinalfish
Ostorhinchus quadrifasciatus
Apogon quadrifasciatus Cuvier, 1828. India.
Indian Ocean, Africa and east to Bali, Indonesia, Red
Sea and Mediterranean immigrant. Coastal, shallow to
deep water on muddy bottom. Grey with two black lines.
Mid-lateral line straight, from tip of snout, through eye
to caudal-fin margin, bordered by white line below, lines
may faint on posterior part of body in adults. Pearly lines
bordering lateral-line, offset on crossing descend. A
black line over nape reaching upper of caudal peduncle,
A pearly line closely above, no black line along dorsal-fin
bases. Fins reddish in adults. Ventrals with a distinct
A white leading edge. Length to about 12 cm.
B C
Ostorhinchus quadrifasciatus Gilimanuk, Bali, Indonesia. RHK O. quadrifasciatus Trawled, 12 cm. Pakistan. HBO
D E
Ostorhinchus quadrifasciatus Lengths 7 & 9 cm. Red Sea. SBO
F
Ostorhinchus quadrifasciatus Subadults with reddish fins. Gilimanuk, Bali, Indonesia. RHK

Japanese Striped-cardinalfish
Ostorhinchus cf. quadrifasciatus
West Pacific, southern tropical Japan and China Sea.
Coastal sheltered bays on muddy substrates. Grey with
two black narrow lines. Mid-lateral line straight, from
snout, through eye, to caudal-fin margin, with short
white bars (female) and spots below. A black thin line
from nape to end of caudal peduncle, an indistinct dusky
thin line above, following the dorsal profile closely to
below dorsal-fins gap. Length to about 12 cm. A
B C
Ostorhinchus cf. quadrifasciatus B juvenile. Iriomote Island A/C Okinawa, Japan. A TKO B KIM C KYO
Short-stripe Cardinalfish
West Pacific, ranging west to Singapore and north to
southern Japan. Grey with three black lines. Mid-lateral
line straight, thick on snout and through eye, to margin
of caudal-fin and white barring below. A black line from
nape with thin white line above along upper side, ending
below end of second dorsal fin. Vertical droplet-like
white dashes below mid-lateral line. No black line along
the dorsal-fin bases. Length to about 12 cm. Ostorhinchus cf. fasciatus-4. Singapore. TKO
Seven-stripe Cardinalfish
Ostorhinchus septemstriatus
Apogon septemstriatus Günther, 1880. Arafura Sea.
Widespread western Pacific, ranging from tropical
Australia to the Philippines. Lives at moderate depths to
52 m. Pale grey and two distinct black lines which have
white thin lines bordering closely above. Mid-lateral line
from tip of snout through eye, to posterior margin of
caudal fin. A black line from top of snout high along side
and along upper of caudal peduncle. A black thin line
from mid-nape divides and continues closely along each
side of the dorsal-fin bases. Length to about 9 cm.
A
Ostorhinchus septemstriatus Port Hedland, WA. CSIRO
B C
Ostorhinchus septemstriatus Philippines. B JGR C ARY

Short-line Rifle-cardinalfish
Ostorhinchus kiensis
Apogon kiensis Jordan & Snyder, 1901.
Wakonoura, Kii, Japan.
Appears to be restricted to sub-tropical southern Japan.
Sheltered coastal muddy bays to deep water, usually seen
solitary, but may form large schools on open substrates
near remote rocky or coral outcrops and large sand
anemones. Grey with two black and three white lines.
Mid-lateral line thick and straight, from tip of snout,
through eye, to end of caudal-fin. Upper black and white
lines short, from top of snout reaching to below or past
end of the second dorsal fin. Length to 8 cm.
A
Ostorhinchus kiensis Osezaki, Japan. TKO
B C
Ostorhinchus kiensis C juvenile. Kochi Prefecture, Japan. THI
D
Ostorhinchus kiensis Osezaki, Japan. RHK

Long-line Rifle-cardinalfish
Ostorhinchus cf. kiensis
Known only from Sydney Harbour, but based on
other sub-tropical Sydney fishes, it probably ranges
north to Capricornia, Queensland. Rocky reefs in
5 to 25 m depth, often seen swimming among
schools of Ostorhinchus fasciatus. Mid-lateral line
broad from tip of snout to end of caudal fin. A
black stripe with a white upper-edge from top of
snout over eye, running high along upper sides to
end of caudal peduncle. Length to about 8 cm.
Appears to be a close relative of the Japanese
Ostorhinchus kiensis.
A
B C
Ostorhinchus cf. kiensis Watson’s Bay, Sydney Harbour. RHK
Bryx Cardinalfish Ostorhinchus bryx

Apogon bryx Fraser, 1998. Luzon Island, Philippines, South China Sea.
Widespread indo-west Pacific. Shallow coastal protected bays with muddy substrates, but has been reported to 155 m depth.
Mid-lateral black line broad from tip of snout to end of caudal fin, bordered by white lines. A thin black stripe from top of
snout over eye, running along upper sides to about midway of caudal peduncle. Paralleled by a white line closely above to
end of 2nd dorsal-fin base. A small pale-blue spot low on gill-cover below white line. Length to about 8 cm.
A
Ostorhinchus bryx Flores, Indonesia. RHK B
C D
Ostorhinchus bryx Philippines. ARY Ostorhinchus bryx Gilimanuk, Bali, Indonesia. RHK

Half-lined Cardinalfish
Ostorhinchus semilineatus
Apogon semilineatus Schlegel, 1843. Japan.
Subtropical Japanese waters. Sheltered bays, rocky reef
habitats, often forming massive schools under jetties and
offshore to 45 m deep. Adults may pair off, but regroup
after spawning. Yellow on lower sides, dusky dorsally.
Head with 3 black lines. One from tip of snout through
eye to opercle margin. Another from top of snout along
upper side to below dorsal-fins gap and third from nape
dorsally along fin-bases. First dorsal fin with black tip and
caudal-fin base with a pupil-sized black spot centrally.
Length to 12 cm.
A
B D
Ostorhinchus semilineatus B school in which many males with oral brood. Izu Peninsula, Japan. RHK
False Ear-spot Cardinalfish

Ostorhinchus jenkinsi
Apogon jenkinsi Evermann & Seale, 1907. Philippines.
Tropical West Pacific, southern Japan to Australia. Clear
water lagoons, forming schools in branching corals. Pale
brownish grey, a small ‘ear’ spot, dark nose stripe an
pupil-sized peduncular spot or narrow bar. Was confused
with Ostorhinchus notatus which is very similar, but has
short black lines on nape. Length to 12 cm.
A B
Ostorhinchus jenkinsi A schooling. Lizard Island, Queensland, Australia. B pair. Komodo, Indonesia. A RHK, B TTO

Ear-spot Cardinalfish
Ostorhinchus notatus
Sparus notatus Houttuyn, 1782. Japan.
Subtropical Japanese waters. Replaced by the
similar tropical Ostorhinchus jenkinsi further
south, which caused many erroneous records.
Common on rocky reefs in the regions of the Izu
Peninsula, where in pairs, but usually forming
schools. Pale brownish grey, a small black ‘ear’
spot, a pupil-sized peduncular spot, and has the
lines on snout to inter-orbitals that distinguishes it
from O. jenkinsi. Length to about 11 cm.
B
Ostorhinchus notatus Kochi, Japan. THI
C D
Ostorhinchus notatus Kagoshima, Japan. SDE Ostorhinchus notatus Kashiwajima, Japan. RHK
Pale-cheek Cardinalfish Mid-stripe Cardinalfish

Ostorhinchus leptofasciatus Ostorhinchus unitaeniatus
Ostorhinchus leptofasciatus Allen, 2001. Apogon unitaeniatus Allen, 1995. Western Australia.
Raja Ampat, West Papua, Indonesia. Coastal muddy habitat and shallow bays in 1 to at least 22 m
West Pacific, known from New Guinea and northeastern depth. Most specimens known from shallow prawn trawls in
Australia on inner reefs in 10–20 depth. Colour pale-brown, about 10 m depth. Reflective silvery, dusky dorsally. A short
cheek whitish, Black striped on snout on dorsal fins close to black dash on opercle behind eye followed by a pale yellow
base, on caudal-fin base and blackish area above anal-fin mid-lateral stripe to caudal-fin base. Length to 8 cm.
base. Length to 12 cm.
Member of the Ostorhinchus notatus group, lacking ‘ear-spot’.
Ostorhinchus leptofasciatus Raja Ampat, West Papua. GRA Ostorhinchus unitaeniatus NT, Australia. GRA

Tail-eye Cardinalfish Ostorhinchus ocellicaudus

Apogon ocellicaudus Allen, Kuiter & Randall, 1994.
Ashmore Reef, Timor Sea.
West Pacific, northwestern Australia, southern Indonesia
north to Sulawesi, Papua New Guinea and the Solomon
Islands. Coastal coral reefs, pairing, in small groups,
occasionally forming schools on remote coral outcrops
in 15–30 m depth. Pale greyish, snout with brown-black
stripes, a yellowish mid-lateral stripe and a large black
basicaudal spot with a white halo. Length to 8 cm.
A B
Ostorhinchus ocellicaudus Lembeh Strait, Sulawesi, Indonesia. RHK
Pale Cardinalfish Ostorhinchus pallidus

Apogon pallidus Allen & Erdmann, 2017.
Star Harbour, Makira Island, Solomon Islands.
Known only from two locations in the Solomon Islands. It
occurs in aggregations hovering about 1–2 m above the
bottom at depths of 18–30 m, usually mixed with other
species. Pinkish, semi-translucent on back, grading to
pale yellowish on lower head, breast, and abdomen;
dusky brown stripe from snout to front of eye, continued
faintly behind eye to rear edge of upper opercle; lower
A sides with series of about 14–16 brown short bars. A
Ostorhinchus pallidus Freshly preserved holotype. GRA black pupil-size basicaudal spot. Length to 8 cm.
B C
Ostorhinchus pallidus Guadalcanal, Solomon Islands. GRA
Mosaic-line Cardinalfish Ostorhinchus cladophilos

Apogon cladophilos Allen & Randall, 2002. Lizard Island, Queensland, Australia.
West Pacific, Flores, Indonesia to northeastern Australia. Occurs on inner reefs in about 20–30 depth. Often found in silty
habitat swimming among soft corals on reef slopes. Semi-transparent, a single black stripe from snout to caudal-peduncle,
thickest on opercle behind eyes, tapering out thin, reaching well short of the black near eye-size spot on caudal-fin base.
Stripe made up of spots in a mosaic-pattern. Length to 45 mm.
A B
Ostorhinchus cladophilos Maumere, Flores, Indonesia. RHK

Gular Cardinalfish Ostorhinchus gularis

Apogon gularis Fraser & Lachner, 1984.
Southern Red Sea, Yemen.
Known from southern Red Sea, Arabian Seas and Gulf
of Oman ranging south to South Africa. Usually trawled
in 30 to 40 m depth. Pale greyish with pale yellowish
hue, a dusky stripe on snout through eye. First dorsal fin
with a black tip. Length to about 65 mm.
Apogon smithvanizi Allen & Randall, 1994, is a junior
synonym. Ostorhinchus gularis KwaZulu-Natal, South Africa. DKI
Black-stomach Cardinalfish
Ostorhinchus atrogaster
Amia atrogaster Smith & Radcliffe 1912.
Luzon Island, Philippines.
Known from trawls to 80 m depth in the Philippines and
northern Australia. Colour in alcohol given: cream-buff,
traces of silvery band from scapula to caudal base; tip of
chin and snout to middle of eye dusky black; tip of 1st
dorsal blackish; caudal dusky distally; other fins hyalin;
peritoneum white; stomach black. Length to 60 mm. Ostorhinchus atrogaster Holotype. After S&R
Slender Cardinalfish Ostorhinchus yamato

Ostorhinchus yamato Yoshida, Hayashi & Motomura, 2018.
Shizuoka, Japan.
Only known from Southern Japan. Occurs at depths of
about 20 m. It may co-occurs with the similar looking
Rhabdamia gracilis, which is also pale, somewhat
transparent whitish, but not as slender and has a small
low-set black basicaudal spot. Length to 60 mm. A
Uncertain genus assignment, exceptionally slender.
Ostorhinchus yamato Otzuki-cho, Shizuoka, Japan. THI
B C
Ostorhinchus yamato B holotype. Kagoshima, Japan. C paratype. Shizuoka, Japan. B HMO C MHA
Pygmy Cardinalfish Ostorhinchus nanus

Apogon nanus Allen, Kuiter & Randall, 1994.
Maumere Bay, Flores, Indonesia.
West Pacific, Indonesia from Bali & Borneo to PNG.
Occurs in small to large aggregations on silty inner reefs
in about 3–20 depth. Greenish grey, semi-transparent,
an ochre mid-lateral stripe from eye to caudal-peduncle
with a white line bordering below. Length to 60 mm,
usually about 45 mm.
A
B C
Ostorhinchus nanus A Tulamben, Bali. B/C Maumere, Flores, Indonesia. RHK

Mini Cardinalfish Ostorhinchus neotes

Apogon neotes Allen, Kuiter & Randall, 1994.
Flores, Indonesia.
Widespread tropical West Pacific. Forms schools in
15–20 m depth along reef wall and on slopes rich with
soft corals. Transparent and small, looking larval-like and
easily overlooked. Black line mid-laterally ending with a
pupil-sized peduncular spot. A white line from below eye
over abdomen that are internally black. Black along
bases of dorsals and anal-fin. Length to 30 mm.
Smallest known member of the genus, but belong to a
group that apparently is subgeneric.
A
B
Ostorhinchus neotes Maumere Bay, Flores, Indonesia. RHK
Red-spot Cardinalfish Ostorhinchus parvulus

Apogon parvulus Smith & Radcliffe, 1912. Tataan Pass, Philippines.
Widespread tropical West Pacific from western, eastern and southern Indonesia to southern Japan. Typically schooling and
congregating around large coral heads in coastal and inner reefs to depth of about 10 m. This species is readily distinguished
from the other small transparent species by the distinct red spot on the base of the caudal fin. Length to 40 mm.
A B
Ostorhinchus parvulus Maumere, Flores, Indonesia. RHK Ostorhinchus parvulus Borneo, Malaysia. RHK
C
Ostorhinchus parvulus Pair, male with oral brood on the right. Amami, Japan. RHK

Whitespot Cardinalfish Ostorhinchus dispar

Apogon dispar Fraser & Randall, 1976. Palau.
Widespread tropical West Pacific. Clear coastal waters
and outer reef walls. Lived in caves along steep walls.
Solitary or in pairs, usually with fan corals in 15–50 m
depth. Semi-transparent, black stripe from snout
through eye, red following eye and as a thin reddish
black line to end of caudal peduncle. A pupil-size black
spot with a small white one directly above on caudal-fin
base. Length to 75 mm. A
Ostorhinchus dispar Tulamben, Bali, Indonesia. RHK
B C
Ostorhinchus dispar Maumere, Flores, Indonesia. RHK
Black-vent Cardinalfish Ostorhinchus melanoproctus

Apogon melanoproctus Fraser & Randall, 1976. Solomon Islands.
West Pacific, Philippines to Indonesia, Papua New Guinea and Solomon Islands. Clear coastal water and outer reef wall.
Lived in caves along steep walls. Types were collected in 18–37 m depth. Looks very similar to Ostorhinchus dispar by the
spotting on the tail, but has a black spot on vent, reddish ventrals, and is smaller when fully grown. Length to 45 mm.
A B
Ostorhinchus melanoproctus Milne Bay, Papua New Guinea. PWO
Faint-striped Cardinalfish Ostorhinchus franssedai

Apogon franssedai Allen, Kuiter & Randall, 1994. Flores, Indonesia.
West Pacific, southern Indonesia to northern Australia and New Guinea. Lived in caves along steep walls, mainly outer reef
habitat. Depth range 15–40 m, but rarely seen in less than 25 m. Semi-transparent, with dusky-yellow stripes from snout
to end of caudal peduncle. An eye-size black spot on caudal-fin base. Length to 75 mm.
A B
Ostorhinchus franssedai Maumere, Flores, Indonesia. RHK

Belly-spot Cardinalfish Ostorhinchus sp. 1

Undetermined species, appears to be undescribed.
Indian Ocean, Maldives and Sumatra, western Indonesia. Lived in caves along steep walls, mainly outer reef habitat. Depth
range 15–40 m, but rarely seen in less than 25 m. Semi-transparent, with 3 dusky stripes from snout over head, fading into
body. A larger-size black spot on caudal-fin base and a white blotch on gular. Length to 75 mm.
A B
Ostorhinchus sp. 1. A Maldives. B Pulau Weh, Sumatra, Indonesia. A RHK B TTO
Sharp-eye Cardinalfish Ostorhinchus sp. 2

West Pacific, southern Japan to Philippines and Palau,
but may involve several very similar transparent looking
taxa with a large black basicaudal spot. Pale greyish
transparent and a large black spot on caudal-fin base. A
very thin white line from lateral-line origin to the large
black caudal spot, encircling much of it over the top.
Ostorhinchus sp. 2. Palau, Micronesia. HNA
Trim Cardinalfish Ostorhinchus sp. 3

West Pacific, known only from photograph, but lives in
caves and probably widespread. Found swimming with
Ostorhinchus melanoproctus at 18 m depth during a
night dive. Pale greyish, transparent and a large black
spot on caudal-fin base. A very thin red mid-lateral line
from behind eye to a large black spot on caudal-fin base.
Ostorhinchus sp. 3. Milne Bay, Papua New Guinea. PWO
Half-barred Cardinalfish Ostorhinchus lineomaculatus

Apogon lineomaculatus Allen & Randall, 2002. Tulamben, Bali, Indonesia.
West Pacific, southern Indonesia and northern Australia to northern Indonesia. Coastal slopes, usually deep, swimming in
pairs or small groups above coral bommies. Out at night to hunt over open substrate, venturing well away from reefs. A
black mid-lateral line with barring below. A pupil-size black spot on caudal-fin base. Length to 10 cm.
A B
Ostorhinchus lineomaculatus A Tulamben, Bali, type-locality. B Flores, Indonesia. RHK

Invisible Cardinalfish
Ostorhinchus aphanes
Ostorhinchus aphanes Fraser, 2012.
Uchelbeluu Reef, Palau.
Little known, only from Palau, where collected from 73 m,
probably deep-water dwelling species. Semi-transparent
with faint dusky bands and yellow line from above eye to
below base of second dorsal-fin. Length to 60 mm.
Ostorhinchus aphanes Holotype. Uchelbeluu Reef, Palau. RWI
Anklet Cardinalfish Ostorhinchus pselion

Apogon pselion Randall, Fraser & Lachner, 1990. Himeira, Sinai Peninsula, Egypt, Gulf of Aqaba, Red Sea.
A Red Sea endemic. Shallow coral reefs to about 35 m depth. Semi-transparent, with dusky-yellow stripes from snout to
caudal peduncle. A near eye-size black spot on caudal-fin base with a white halo. Length to 50 mm.
A B
Ostorhinchus pselion Farasan Island, Saudi Arabia, Red Sea. SBO
Shooting-star Cardinalfish Ostorhinchus selas

Apogon selas Randall & Hayashi, 1989. Madang, Papua New Guinea.
West Pacific, southern Japan to Indonesia and Solomon Islands. Shallow coral reefs to about 45 m depth. A broad heavy
dusky-yellow mid-lateral stripe and a large black spot on caudal-fin base. Upper line in eye yellow. Length to 50 mm.
A B
Ostorhinchus selas Amami, Japan. RHK Ostorhinchus selas Milne Bay, Papua New Guinea. RHK
Western Gobbleguts Ostorhinchus rueppellii

Apogon rueppellii Günther, 1859. Australian seas. Western Pacific and southeastern Indian Ocean: Western Australia,
Northern Territory (Australia) and Papua New Guinea. Mainly Australia’s west coast ranging south into subtropical zone.
Pale grey with black dots along lateral line and few scattered elsewhere. Length to 12 cm.
A B
Ostorhinchus rueppellii A Fremantle, Geograph Bay. B Monte Bellow Islands, Western Australia. RHK

Orange-lined Cardinalfish
Ostorhinchus cyanosoma
Apogon cyanosoma Bleeker, 1853.
Lawajong, Solor Island, Timur Province, Indonesia.
West Pacific: Philippines to southern Indonesia, north-
eastern Australia and New Caledonia. Records from
elsewhere are probably based on other similar species.
Coastal and lagoons to about 20 m depth, often in
schools. Pale grey, dark on snout, with narrow orange
lines from head to caudal-fin base. Mid-lateral line may
end in a spot, which is orange-red at night. Lower two
lines on body thinner than interspaces. Top and bottom
of eye yellow. Length to about 65 mm.
Ostorhinchus rubrimaculata Randall & Kulbicki, 1998,
A is a putative junior synonym, its description based on the
Ostorhinchus cyanosoma Tulamben, Bali, Indonesia. RHK red-spot night-form (C).
B C
Ostorhinchus cyanosoma Day and night colours. Gilimanuk, Bali, Indonesia. RHK
D
Ostorhinchus cyanosoma Near type-locality. Northeastern Flores, Indonesia. RHK

Pacific Orange-lined Cardinalfish

Ostorhinchus sp. 4
Undetermined species, apparently undescribed.
Widespread tropical West Pacific, southern Japan to
Bali, Indonesia and New Guinea. Coastal and inner reef,
in pairs or small aggregations in caves. Similar to
Ostorhinchus cyanosoma, but lower 3 orange lines
much thicker than interspaces. Eye large and nearly all
black with a hint of orange in the iris. No peduncular
spot. Length to 70 mm. A
Ostorhinchus sp. 4. Iriomote Island, Japan. RHK
B C
Ostorhinchus sp. 4. Amami, Japan. RHK Ostorhinchus sp. 4. Bali, Indonesia. RHK
D E
Ostorhinchus sp. 4. Iriomote Island, Japan. D TKO E RHK
Indian Orange-lined Cardinalfish

Ostorhinchus sp. 5
Widespread west Indian Ocean and Red Sea. Coastal
and inner reef, in pairs or aggregations. Similar to
Ostorhinchus sp. 4, but lines on upper sides very thin
and interspaces wider, ending on caudal-fin base. Head
reddish below and behind eye, snout and nape black
with white lines. Eye large and nearly all black. Lacks a
basicaudal spot. Length to 70 mm.
B C
Ostorhinchus sp. 5. B night colour. Safaga, Egypt, Red Sea. RHK

Southern Orange-lined Cardinalfish

Ostorhinchus properuptus
Lovamia properupta Whitley, 1964.
Frederick Reef, Coral Sea.
Eastern Australia, south to sub-tropical central NSW,
New Caledonia, PNG and west to Sumatra, Indonesia,
but Ostorhinchus properuptus appears to comprise a
species-complex. Coastal and offshore on rocky and
coral reefs to about 30 m depth, solitary or in pairs.
Lower half of body solid orange with a single narrow
pale blue line at lower eye-level. A short line over cheek.
In true species orange lines above eye-level narrow with
A wider interspaces. Length to about 85 mm.
Ostorhinchus properuptus Solitary Islands, NSW. RHK Often identified as Ostorhinchus cyanosoma.
B C
Ostorhinchus aff. properuptus Milne Bay, PNG. RHK O. properuptus Juvenile. Sydney, NSW. RHK
D E
Ostorhinchus aff. properuptus D Pulau Putri, Java. E Pulau Weh, Sumatra, Indonesia. D RHK E RCS
Maldivian Cardinalfish Ostorhinchus sp. 6

Indian Ocean, Maldives. Possibly widespread in the region. On outer reef margins and deep lagoons. May form large
schools. Was reported as Ostorhinchus cyanosoma, but is related to the O. properuptus group. Very pale yellowish,
showing very faint yellow lines. Snout black with bluish white lines, ending just past eyes. Length to 55 mm.
A B
Ostorhinchus sp. 6. Maldives. RHK

Hibernia Cardinalfish
Ostorhinchus sp. 7
Timor Sea, geographical distribution is not known. This
taxon looks very similar to Ostorhinchus cyonosoma in
colour and having the pattern of narrow orange lines
along the body and caudal peduncle, but interspaces
more bluish and lines continue over caudal fin and there
is no suggestion of a basicaudal spot. Length 50 mm.
Ostorhinchus sp. 7. Timor Sea, Western Australia. GRA
Kupang Cardinalfish Ostorhinchus wassinki

Apogon wassinki Bleeker, 1861. Kupang, Timor.
Timor Sea, distribution unclear. Name often applied to members of the Ostorhinchus cyonosoma complex, but is possibly
of the O. properuptus complex. The type 60 mm long, was described as yellow with 2 or 3 reddish lines and pinkish fins.
A B
Ostorhinchus wassinki A type, after Bleeker. B ? Cassini Island, Kimberley Coast, Western Australia. GRA
Dusky Orange-lined Cardinalfish

Ostorhinchus sp. 8
Monte Bello Archipelago, Western Australia. Sandstone
reefs in tidal channels. A member of the Ostorhinchus
cyonosoma complex with black stripes on head that
fade to dusky yellow stripes on body. Interspaces
whitish. Length 70 mm.
Ostorhinchus sp. 8. Monte Bello Islands, Western Australia. RHK
Yellow Cardinalfish Ostorhinchus luteus

Apogon luteus Randall & Kulbicki, 1998. Marshall Islands.
Micronesia to Fiji Islands. A member of the Ostorhinchus
properuptus complex. Shallow coastal to 50 m depth.
Canary yellow, two white lines from snout through eye,
with brown to black in between. Length to 50 mm.
B C
Ostorhinchus luteus Guam, Micronesia. A/B RHK C RMY

Izu Cardinalfish Ostorhinchus sp. 9

Undescribed species known only from southern Japan,
occurring from the Izu Peninsula to the Kagoshima
region. Locally common on shallow rocky reefs, in pairs
or schooling near holes and caves. Lower half of sides
orange with white lines, lower as a series of dots. Upper
half of sides pale grey with narrow orange lines. Snout
with black and white line. Length to about 85 mm.
Usually referred to Ostorhinchus properuptus, but this

species is solid orange over the abdominal area. Instead
it shows similarities with O, holotaenia, especially in the
lower line as spots and the striped pattern on the tail.
A
Ostorhinchus sp. 9. Pair. Kashiwajima, Japan. RHK
B C
D E
Ostorhinchus sp. 9. B–E spawning sequence. Kochi Prefecture, Japan. THI
F G
Ostorhinchus sp. 9. F schooling on rubble-rock reef. G juvenile. Osezaki, Japan. F RHK G TKO

Copper-striped Cardinalfish Ostorhinchus holotaenia

Apogon holotaenia Regan, 1861. Muscat, Oman, Gulf of Oman, Arabian Sea.
Widespread Indian Ocean and Red Sea, east to Indonesia. Coastal reefs in about 15–35 m depth. Variable with yellow to
brown stripes. Mid-lateral line continues on caudal fin to its margin. Lower pale line break up into spots. Length to 8 cm.
A B
Ostorhinchus holotaenia Socotra. SBO Ostorhinchus holotaenia Maldives. RHK
C D
Ostorhinchus holotaenia Sodwana, South Africa. RHK Ostorhinchus holotaenia Seychelles. GRA
Red-tail Cardinalfish Ostorhinchus leslie

Ostorhinchus leslie Schultz & Randall, 2006.
Rose Atoll, American Samoa.
American Samoa and Vanuatu regions. It is usually
found at moderate depths around reef margins in 20–35
m. A distinctive species, bright yellow changing to red
on caudal peduncle, colour intensifying on tail-base.
Ostorhinchus leslie Rotuma, Fiji. JTW
Cavite Cardinalfish Ostorhinchus cavitensis

Amia cavitensis Jordan & Seale, 1907.
Cavite, Luzon Island, Philippines.
Known from Singapore and Bali to Philippines. Coastal,
often silty habitat on muddy substrates with rubble or
rocky outcrops. Pale grey above, white below. A broad
mid-lateral yellow to green stripe with white borders.
Black on snout and a pupil-size black spot on caudal fin,
just beyond base. Length to 65 mm.
A
B C
Ostorhinchus cavitensis Bali, Indonesia. RHK Ostorhinchus cavitensis Singapore. RHK

Silver-lined Cardinalfish
Ostorhinchus hartzfeldii
Apogon hartzfeldii Bleeker, 1852.
Ambon Island, Molucca Islands, Indonesia.
West Pacific from Philippines to Indonesia. Records
from elsewhere probably based on sibling species. In
coastal bays on open muddy substrates, usually close to
rubble outcrops or corals. Juveniles with black broad
stripes and white lines on head and a white line from
nape high along sides to caudal-fin base. Adult mostly
brown to near black, retaining white lines on head and
A high on along side. A pupil-size black spot slightly above
Ostorhinchus hartzfeldii Flores, Indonesia. RHK centre on caudal-fin base. Length to about 12 cm.
B C
Ostorhinchus hartzfeldii B Mabul, Borneo, Malaysia. C Derawan, Kalimantan, Sulawesi Sea. B TKO C RHK
D E
Ostorhinchus hartzfeldii D night. E juvenile. Mabul, Borneo, Malaysia. RHK
Golden-lined Cardinalfish
Ostorhinchus sp. 10
Undescribed species from tropical eastern Australia,
ranging south into subtropical zones. Solitary or in pairs
on shallow rocky or rubble reefs, near holes and caves.
Head with black stripes, fading to greenish yellow on
body. A pair of white lines from snout through eye and
edging the mid-lateral stripe. A thin yellow line from
nape, running high along back following dorsal profile to
end of caudal peduncle. A pupil-size black spot centered
on caudal fin close to base. Length to about 85 mm.
A
B C
Ostorhinchus sp. 10. C juvenile. B Moreton Bay. A/C Keppel Island. Queensland, Australia. RHK

High-fin Cardinalfish
Ostorhinchus chrysotaenia
Apogon chrysotaenia Bleeker, 1852.
Jakarta, Java, Indonesia.
Indonesia, northwestern Australia, north to Japan, east
to Solomon Islands. Shallow coastal reef flats with rocky
boulder substrates and coral bommies to about 10 m
depth. Juveniles pale whitish with dark stripes. Adults
orange, blue lines on head and faint brown body-stripes,
darkest dorsally. An intermittent black basicaudal spot on
lateral-line or above. Dorsal-fins sharply pointed, 2nd fin
very tall and sickle shaped. Length to about 12 cm.
Amia cyanotaenia Bleeker, 1853 is a putative synonym.
A
B C
Ostorhinchus chrysotaenia B juvenile. Mabul, Borneo, Malaysia. A/C Maumere, Flores, Indonesia. RHK
D E
Ostorhinchus chrysotaenia D Maumere, Flores, Indonesia. E Raja Ampat, West Papua. D RHK E GRA
Bangka Cardinalfish
Ostorhinchus endekataenia
Apogon endekataenia Bleeker, 1852.
Bangka or Lepar Island, Indonesia.
West Pacific, Indonesia, Malaysia, Brunei and also
Singapore. Sheltered silty habitats, forming small groups
with rubble or rocky outcrops on open bottom. Juveniles
with 3 black stripes and a large eye-size black spot on
caudal-fin base. Stripes change in colour with growth to
a dusky brown. Length to about 14 cm.
B C
Ostorhinchus endekataenia B juvenile. Singapore. RHK

Schlegel’s Cardinalfish
Ostorhinchus schlegeli
Apogon schlegeli Bleeker, 1855. Japan.
Subtropical Japan, possibly ranging into northern China
Sea. Records from elsewhere doubtful. In coastal waters
on rocky slopes and walls in caves and ledges, shallow to
60 m depth. Pinkish overall, adult with 7 long blackish
brown lines along body, fewer and black in juvenile, and
a larger than pupil-size black spot centrally on caudal-fin
base. Length to about 12 cm.
A Was confused with Ostorhinchus endekataenia due to
Ostorhinchus schlegeli Osezaki, Japan. RHK a mix-up of Bleekers drawings of the types.
B
Ostorhinchus schlegeli Izu, Japan. RHK
C D
Ostorhinchus schlegeli C juvenile. Kochi Prefecture, Japan. D male with oral brood. Kagoshima, Japan. C THI D SDE
Futasuji Cardinalfish Ostorhinchus fukuii

Apogon fukuii Hayashi, 1990.
Coastal water off Ukui, Wakayama Prefecture, Japan.
Indo-Pacific, subtropical waters of Japan, New Zealand
and eastern South Africa. Usually solitary along rocky
walls at moderated depth, 35–60 m. A large eye-size
black spot centrally on caudal peduncle close to base. A
mid-lateral line from snout through eye, reaching just
short of peduncular spot and a slightly shorter line from
nape along upper side reaching well past below end of
second dorsal-fin base. Length to about 85 mm.
B C
Ostorhinchus fukuii Izu Peninsula, Japan. RHK Ostorhinchus fukuii KwaZulu-Natal, South Africa. DKI

Munahosi Cardinalfish Ostorhinchus cheni

Apogon cheni Hayashi, 1990. Lejima Island, Okinawa
Prefecture, Japan, depth 70-100 meters.
Known from captures in southern Japan, Guam, Taiwan,
west to Sri Lanka, and south to Bali, Indonesia, where
photographed (B). Lives on deep rocky reefs to 100 m
depth. Dark thin body lines, upper just reaching past
dorsal fins, irridescent blue lines on head. A smaller than
pupil-size black basicaudal spot and a tiny dusky spot below
1st dorsal spine at eye-level. Length to 16 cm.
A
Ostorhinchus cheni Guam, Micronesia. RMY
B C
Ostorhinchus cheni At 70–80 m. Tulamben, Bali. TSH Ostorhinchus cheni Philippines. HMO
Thin-lined Cardinalfish Ostorhinchus sp. 11

Tropical Western Australia, south to Rottnest Island.
Shallow rocky reefs in caves and ledges, to about 20 m
depth. Body pale greyish pink to yellowish pink and a
pupil-size black basicaudal spot. Three to 5 black to
brown thin longitudinal lines, distinct and blackish in
juveniles, fading in adults to two long pale dusky brown
thin lines reaching caudal peduncle. Length to 14 cm.
B C
Ostorhinchus sp. 11. B large juvenile. C pair, male with oral brood. Monte Bello Islands, Western Australia. RHK
D E
Ostorhinchus sp. 11. D juvenile. Northwest Cape. E Port Hedland. Western Australia. D GRA E NCO.

Four-line Cardinalfish
Ostorhinchus doederleini
Apogon doederleini Jordan & Snyder, 1901.
Nagasaki, Japan.
Widespread West Pacific, but comprises a complex of
geographical forms of undetermined status. Pale grey to
pinkish with four long black body lines thin, thickest up
to pupil-size width on cheek. Lines narrower in some
populations. An about pupil-sized black basicaudal spot,
proportionally larger in juveniles. Fins pinkish, turning
red at night. Length to 16 cm.
A
Ostorhinchus doederleini Kashiwajima, Japan. RHK
B
Ostorhinchus doederleini Night colour. Izu, Japan. RHK C
D E
Ostorhinchus doederleini C juvenile. Iriomote Island, Japan. KYA
F G
Ostorhinchus doederleini Ehime, Japan. THI Ostorhinchus doederleini Nuptial. Kochi, Japan. THI
H I
Ostorhinchus doederleini Moreton Bay, Qld. RHK Ostorhinchus doederleini Juvenile. Sydney. RHK

Sydney Cardinalfish
Ostorhinchus limenus
Apogon limenus Randall & Hoese, 1988.
Sydney Harbour, New South Wales, Australia.
Eastern Australia, southern Qld, to southern NSW. On
rocky and coral reefs below overhangs in pairs or small
groups, shallow to about 20 m depth, but trawled in
deep water as well. Pale greyish pink with black lines and
pink to red fins. Stripes thick along lower sides and thin
and high on upper sides, with a wide space between 2nd
and 3rd with short stripe on head. A pupil-size or larger
black spot on caudal-fin base. Length to 14 cm. A
Ostorhinchus limenus Keppel Island, Queensland. RHK
B C
D E
Ostorhinchus limenus B small juvenile. C male. D subadult. E night colours. Sydney Harbour, NSW. RHK
Reef-flat Cardinalfish
Ostorhinchus taeniophorus
Apogon taeniophorus Regan, 1908. Maldives.
Widespread Indo-Pacific. Shallow reef flats, usually in a
few metres depth under coral plates. Pale pinkish dusky
with four main black stripes, lower two very heavy, close
together on lower half, upper two much thinner on
upper half. Mid-lateral line thickens on the base of the
caudal fin as a spot, lines continuing a little onto the fin.
Fins reddish. Length to 14 cm. A
Ostorhinchus taeniophorus Maumere, Flores, Indonesia. RHK
B C
Ostorhinchus taeniophorus Mauritius. PCH Ostorhinchus taeniophorus Tulamben, Bali, Indonesia. RHK

Northern Striped-cardinalfish
Ostorhinchus victoriae
Apogon victoriae Günther, 1859. Victoria River, NT.
Northwestern Australia and Northern Territory. Coastal
reef in shallow water where it was observed to about 15
m depth. Pale with reddish brown soft-edged stripes that
are narrower than interspaces. A half-length stripe from
top of eye to below 2nd dorsal fin. Pectoral-fin base
dark, a pupil-sized black spot on caudal-fin base, and has
a very large anal fin. Length to about 14 cm.
A rarely observed species due to the remoteness of the
region. The type-locality was originally not stated, but
Ostorhinchus victoriae Northern Western Australia. RCS revealed by Macleay, 1881.
Western Striped-cardinalfish
Ostorhinchus aff. victoriae
Western Australia from Geograph Bay to Shark Bay,
subtropical. Clear coastal waters and offshore rocky
reefs to about 20 m depth. Stripes thick, reddish brown
with narrower pale interspaces. Three long sharp-edged
stripes from snout to base of the caudal-fin. Interspaces
narrower, pale grey to whitish. A half-length stripe from
upper of eye, ending about below dorsal fins gap. Fins
pinkish to red. A larger than pupil-size black basicaudal
A spot with a whitish halo. Length to 14 cm.
B C
Ostorhinchus aff. victoriae A Carnac Island. B Busselton. C Fremantle. Southern Western Australia. RHK
Cook’s Cardinalfish Ostorhinchus cookii

Apogon cookii Macleay, 1881. Endeavour River and Erub,
Torress Strait Islands, Queensland, Australia.
Widespread Indo-West Pacific, and Red Sea. Sheltered
coastal bays and lagoons to about 10 m depth, but
occurs also offshore in moderate depths. Black stripes
very heavy along lower sides. Mid-lateral line usually
ends in a rounded blotch on the caudal-fin base. Fins
pinkish. Length to about 10 cm.
A
Ostorhinchus cookii Bali, Indonesia. RHK
B C
Ostorhinchus cookii Northeastern Sulawesi, Indonesia. RHK Ostorhinchus cookii Socotra. SBO

Pale-striped Cardinalfish Ostorhinchus pallidofasciatus

Apogon pallidofasciatus Allen, 1987. North West Cape, Western Australia.
Northwestern Australia and Northern Territory. In tidal channels and silty rubble reef in shallow water, observed to about
15 m depth. Adults may look completely black with indistinct pale lines breaking the otherwise uniform dark colour into
broad stripes. An pupil-sized black spot on caudal-fin base which may be indistinct. Length to about 13 cm.
B C
Ostorhinchus pallidofasciatus Female with pale stripes, male plain. Monte Bello Islands, Western Australia. RHK
Black Cardinalfish
Ostorhinchus aterrimus
Apogon aterrimus Günther, 1867.
Cape York, Queensland, Australia.
West Pacific, eastern Australian coast from Cape York to
Moreton Bay. Sheltered coastal bays on silty rubble reef
in shallow water, observed to about 10 m depth. A deep
bodied species with large fins. Adults may look almost
black with indistinct darker stripes, one mid-laterally and
one on upper sides. Juveniles dark greenish and are
more distinctly striped, 4 of which reaching caudal-fin
base, have reddish dusky fins, and an indistinct black
pupil-sized basicaudal spot. Length to about 16 cm.
B C
Ostorhinchus aterrimus B subadult. North Stradbroke Island, Moreton Bay, Queensland, Australia. RHK

Aroub Cardinalfish
Ostorhinchus aroubiensis
Apogon aroubiensis Hombron & Jacquinot, 1853.
Aroub, Malaysia.
Only known from the single type, which was just under
6 cm in total length and may be subadult. Colour was
given as silvery green with four black stripes; fins green
with broad brown border. The Apogon aroubiensis
name was often used for other species, but of the known
striped species, only juvenile Ostorhinchus aterrimus
has four lines along the caudal peduncle and is perhaps
its closest relative. It may also live in similar estuarine
Ostorhinchus aroubiensis Holotype. after H&J silty coastal habitats.
Tail-stripe Cardinalfish
Ostorhinchus sp. 12
Tropical Western Australia. Only known from the Monte
Bello Islands in stong current prone tidal channels under
low overhangs. Heavy black lines on head and body with
whitish narrow interspaces. Mid-lateral stripe over the
entire length from tip of snout to caudal-fin’s posterior
margin. Line from above eye very short, on head only.
Ostorhinchus sp. 12. Monte Bello Islands, WA. RHK
Heavy-stripe Cardinalfish Ostorhinchus sp. 13

Known from coastal central Queensland. A distinct species among the multistriped taxa by the heavy mid-lateral stripe, a
narrow black stripe from nape high along upper sides and a pupil-size black basicaudal spot with pale halo. Length 10 cm.
A B
Ostorhinchus sp. 13. Magnetic Island, Queensland. RHK
Ruler Cardinalfish Ostorhinchus regula Tiny-spine Cardinalfish Ostorhinchus brevispinis

Apogon regula Fraser & Randall, 2003. Apogon brevispinis Fraser & Randall, 2003.
Condor Reef, Caroline Islands. Rurutu, Austral Islands, South Pacific.
Deeper waters of Guam and Condor Reef in the Carolina Rurutu Island in the Austral Islands and Rangiroa, Tuamoto
Islands, 30–60 m. Type was collected from 55 meters. Archipelago. Deep water coral slopes at 50–60 m depth.
Silvery with pale-brown broad stripes, 3 of which reaching Pale creamy white with three tan-brown broad stripes that
caudal-fin base. Length to 45 mm. reach caudal-fin base. Length to 45 mm.
Ostorhinchus regula Paratype. Guam, Micronesia. JER Ostorhinchus brevispinis Rangiroa. JER

Black-striped Cardinalfish
Ostorhinchus nigrofasciatus
Apogon nigrofasciatus Lachner, 1953.
Yuro Island, Bikini Atoll, Marshall Islands.
Widespread Indo-Pacific, but a taxa-complex with at
least 3 forms. Coastal to outer reef slopes and walls in
narrow ledges and caves to at least 50 m depth. Mostly
black with thin white or yellow lines between the very
broad black stripes running from snout to caudal-fin
base. Fins pink in some forms. Length to 14 cm.
A
B C
Ostorhinchus nigrofasciatus B juvenile. A Montague Island. B/C Sydney Harbour. NSW, Australia. RHK
D E
Ostorhinchus nigrofasciatus Iriomote Island, Japan. KYA Ostorhinchus nigrofasciatus Sipadan, Borneo. RHK
F
Ostorhinchus nigrofasciatus Maldives. RHK G
H
Ostorhinchus nigrofasciatus Maumere, Flores, Indonesia. RHK

Narrow-striped Cardinalfish
Ostorhinchus angustatus
Amia angustata Smith & Radcliffe, 1911.
Malanipa Island, east of Zamboanga, Philippines.
Widespread Indo-West Pacific, and Red Sea. Clear
coastal reef crests and slopes, where usually shallow, and
drop-offs to about 25 m depth. Black stripes and whitish
interspaces close to subequal width. Mid-lateral stripe
ends into a spot on the base of the caudal fin. Fins pink
translucent to reddish. Length to about 11 cm.
A
Ostorhinchus angustatus Tulamben, Bali, Indonesia. RHK
B C
Ostorhinchus angustatus Maumere, Flores, Indonesia. RHK Ostorhinchus angustatus Maldives. RHK
Three-line Cardinalfish
Ostorhinchus novemfasciatus
Apogon novemfasciatus Cuvier, 1828.
Timor; Guam, Mariana Islands, western Pacific.
Tropical Indo-West Pacific. Shallow protected coastal
reefs, often silty habitat. usually in pairs, swimming close
to shelter. Three main black body stripes that reach to
centre of caudal fin, the outer two with ends converging.
Mid-lateral line continues thinly over caudal fin, and may
reach posterior margin. Length to 10 cm.
B C
Ostorhinchus novemfasciatus A male with oral brood. Singapore. B Flores, Indonesia. C night colour. Kalimantan. RHK
D
Ostorhinchus novemfasciatus Pair, male with oral brood on the right. Maumere, Flores, Indonesia. RHK

Multi-lined Cardinalfish
Ostorhinchus multilineatus
Amia multilineata Bleeker, 1874.
Ceram Island, Molucca Islands, Indonesia.
Tropical West Pacific, Widespread Indonesia west to
Java, north to Philippines and also Solomon Islands.
Coastal reefs and lagoons, usually with large coral
bommy outcrops in 2–25 m depth. Head with broad
black stripes over snout and cheeks; body with numerous
thin lines, the numbers increasing with growth, few of
which slightly thickened. Fins pink to reddish. Length to
about 11 cm. A
Ostorhinchus multilineatus Maumere, Flores, Indonesia. RHK
B C
Ostorhinchus multilineatus Mabul, Borneo. RHK Ostorhinchus multilineatus Night. Manado, Indonesia. RHK
Blue-eyed Cardinalfish
Ostorhinchus compressus
Amia compressa Smith & Radcliffe, 1911. Philippines.
Widespread tropical West Pacific. Coastal reefs and
outer reef lagoon. Usually schooling among staghorn
corals. Juveniles whitish with black narrow lines and a
yellow peduncle blotch with a black spot centrally, just
like in young Cheilodipterus spp. (D). Stripes in adults
are thicker and reddish black; eyes look blue in natural
light and fins reddish. Length to about 12 cm.
A
Ostorhinchus compressus Milne Bay, PNG. RHK
B C
Ostorhinchus compressus Flores, Indonesia. RHK Ostorhinchus compressus Lizard Island, Qld, Australia. RHK
Ostorhinchus
D Cheilodipterus E
Ostorhinchus compressus Juv. Flores, Indonesia. RHK Ostorhinchus compressus Pulau Putri, Java, Indonesia. RHK

Sister Cardinalfish Ostorhinchus relativus Recess Cardinalfish Ostorhinchus sinus

Apogon relativus Randall, 2001.Marquesas Islands. Apogon sinus Randall, 2001.
Known only from the Marquesas Islands were common, Rurutu, Austral Islands, South Pacific.
apparently replacing the similar Ostorhinchus angustatus Occurs in very shallow water (one from a tide pool) at the
on shallow reef-flats. Pale creamy white with three black to heads of deep bays. Pale greyish brown with seven narrow
brown stripes that reach caudal-fin base. Mid-lateral line dark stripes on the body, five of which reaching caudal-fin
ends in black spot. Length to 11 cm. base. Length to 16 cm.
Ostorhinchus relativus Holotype. Marquesas Islands. JER Ostorhinchus sinus Holotype. Marquesas Islands. JER
Cheek-spots Cardinalfish
Ostorhinchus chrysopomus
Apogon chrysopomus Bleeker, 1854.
Makassar, Sulawesi, Indonesia.
Western Pacific: Indonesia to Philippines. Sheltered
coral reefs, usually in about 5–10 m depth, but ventures
down to about 25 m. Yellowish white with two orange to
blackish long lines. Very similar to Ostorhinchus sealei,
but cheek with 5 or 6 orange spots instead of two bars.
Pupil-size black spot slightly above centre on caudal-fin
A base. Length to 65 mm.
C
B Ostorhinchus chrysopomus. Singapore. TKO
D
Ostorhinchus chrysopomus B night colour. C juvenile. D pair. Sulawesi, Indonesia. RHK

Short-band Cardinalfish
Ostorhinchus oxygrammus
Apogon oxygrammus Allen, 2001.
Raja Ampat Islands, Irian Jaya Province, Indonesia.
Known only from the type-locality area. Occurs on
macro-algal reef rubble or coral-rock outcrops in about
35–50 m depth. A whitish species with a single broad
black stripe mid-laterally, extending onto caudal fin. No
line on snout. A small species, length to 50 mm.
Ostorhinchus oxygrammus Raja Ampat, West Papua. GRA
Cheek-bar Cardinalfish Ostorhinchus sealei

Apogon sealei Fowler, 1918. Philippines.
West Pacific, Indonesia to the Philippines and also in
north-western Australia. Along reef margins in small to
large schools. Usually in less than 10 m depth, but found
to about 25 m depth. Pale yellowish white, juveniles with
three black thin lines, one mid-laterally, followed by a
small caudal-fin base spot and two lines along upper
side. Lines fade in large adults and posteriorly break up
into orange spots. Cheek with two orange bars. Length
to about 10 cm. A
B C
Ostorhinchus sealei Maumere, Flores, Indonesia. RHK Ostorhinchus sealei Mabul, Borneo, Malaysia. RHK
D
Ostorhinchus sealei An amazing number of individuals forming a school under a village jetty at Pulau Besar, with several
other Apogonidae members and a blenny joining the dense school. Maumere, Flores, Indonesia. RHK

Pearly Cardinalfish
Ostorhinchus margaritophorus
Apogon margaritophorus Bleeker, 1854.
Batjan, Indonesia.
Western Pacific: Malaysia and Indonesia to Philippines,
and Papua New Guinea to Solomon Islands. Shallow to
about 5 m, in small groups in weedy reef and seagrass
habitats. A white mid-lateral stripe and red to brown
stripes above and below. Lower two side stripes have
short connecting bars, forming a series of white spots in
between. First dorsal fin with the longest spine white,
A
followed by red-brown membrane. Length to 65 mm.
Ostorhinchus margaritophorus Mabul, Borneo, Malaysia. RHK
B C
D E
Ostorhinchus margaritophorus Sulawesi, Indonesia. RHK Ostorhinchus margaritophorus Singapore. RHK
Harrow Cardinalfish Ostorhinchus oxina

Apogon oxina Fraser, 1999. Madras, India.
Indian Ocean, Arabian region to India and Sri Lanka. Coastal rubble reef, usually forming pairs or small schools. Closely
related to Ostorhinchus moluccensis. Pale silver grey with thin brownish black lines, four of which over head, mid-lateral
line to base of caudal fin. A line from above eye to end of second dorsal-fin base to a white spot. Mid-lateral line crossed by
four or five thin bars over abdominal section. Length to about 8 cm.
A B
Ostorhinchus oxina Aquarium, from Sri Lanka. FZE

Moluccen Cardinalfish
Ostorhinchus moluccensis
Apogon moluccensis Valenciennes, 1832.
Widespread eastern Indian Ocean and West Pacific.
Locally abundant in estuaries and coastal bays in silty
habitats to depths of 25 m. Pale to dark brown with
white and brown to black stripes on head and a distinct
white spot below the end of second dorsal-fin base. It
has barring on abdominal area that can be turned on and
off at will. Length to about 10 cm.
Apogon ventrifasciatus Allen, Kuiter & Randall, 1994 A
is a putative synonym. Ostorhinchus moluccensis Gilimanuk, Bali, Indonesia. RHK
B C
Ostorhinchus moluccensis Manado, Indonesia. RHK
D E
Ostorhinchus moluccensis Tulamben, Bali, Indonesia. RHK
Yellow-eye Cardinalfish
Ostorhinchus monospilus
Apogon monospilus Fraser, Randall & Allen, 2002.
Philippines.
Indonesia, Australia to Philippines. Pale to dark brown
with white and dusky stripes on head and a distinct white
spot below the end of second dorsal-fin base.
Intermittent broad mid-lateral brown band. Similar to
Ostorhinchus moluccensis, but eye yellow, no banding
and white spot less distinct. Length to about 10 cm.
A
Ostorhinchus monospilus Gilimanuk, Bali, Indonesia. RHK
B C
Ostorhinchus monospilus Qld, Australia. RHK Ostorhinchus monospilus & moluccensis Philippines. TKO

Spotted Cardinalfish Ostorhinchus maculiferus

Apogon maculiferus Garrett, 1864. Hawaiian Islands.
Hawaiian Islands endemic. Common in the cooler northwestern chain of the islands where is may form large schools in the
vicinity of caves or below overhangs. Dusky to pale brown with series of small dark spot along scale rows, some forming
more or less distinct lines. Fins semi-transparent pale brown or yellowish. Length to 14 cm.
A B
Ostorhinchus maculiferus Hawaiian Islands. JHO
Easter Island Cardinalfish

Ostorhinchus chalcius
Apogon chalcius Fraser. & Randall, 1986.
Easter Island.
Only known from Easter Island. Plain brownish on head
and upper sides, lower sides with reflective silvery thinly
barred pattern. Some dusky blotching over snout and
dorsally on head. Length to about 16 cm.
Ostorhinchus chalcius Easter Island. JER
Hook-fin Cardinalfish Coach-whip Cardinalfish

Ostorhinchus griffini Ostorhinchus flagelliferus
Amia griffini Seale, 1910. Bantayan Island, Philippines. Amia flagelliferus Seale, 1910. Madras, India.
Indonesia, Borneo and Philippines. Rock and weed bottom Western Indian Ocean: South and East Africa to
with scattered corals in 2–10 m depth. Lower part of head Madagascar. Rocky and rubble habitat. Colour uniformly
and sides reddish brown. Sides below lateral line with dusky pinkish red, intensifying on head and may show a barred
bars. Pale over the back and caudal peduncle. Juveniles with pattern on sides below lateral line. Short bluish white lines
longitudinal stripes, a yellowish caudal peduncle and a small over snout and eye, filled in between with black. A small
black spot centrally on caudal-fin base. Length to 14 cm. black spot on caudal-fin base. Length to about 13 cm.
Ostorhinchus griffini Sabah, Borneo. GRA Ostorhinchus flagelliferus Sodwana, South Africa. DKI

Flag-fin Cardinalfish Ostorhinchus hoevenii

Apogon hoevenii Bleeker, 1854.
Widespread West Pacific, ranging from southern Japan
to northern Australia, west to Java and east to the
Solomon Islands. Sheltered bays and reefs, shallow to at
least 30 depth. Often seek protection in long-spined
urchins or other invertebrates, including anemones, and
forming dense schools. Pale brown or grey to dusky.
First dorsal-fin is distinct by its white posterior margin. A
tiny white spot in front of first dorsal and one below the
end of second dorsal-fin base. Eye yellow. Length to
about 10 cm, but usually much smaller.
Identified in Japan as Ostorhinchus ishigakiensis, which A
is a putative junior synonym. Ostorhinchus hoevenii Male with oral brood. Bali, Indonesia. RHK
B C
Ostorhinchus hoevenii Flores, Indonesia. RHK Ostorhinchus hoevenii (ishigakiensis) Kerama Island, Japan. RHK
D E
Ostorhinchus hoevenii Lizard Island, Qld, Australia. RHK Ostorhinchus hoevenii Gilimanuk, Bali, Indonesia. RHK
F
Ostorhinchus hoevenii & O. monospilus Some swimming among anemone tentacles. Sulawesi, Indonesia. FSN

Black-foot Cardinalfish
Ostorhinchus nigripes
Apogon nigripes Playfair, 1857. Zanzibar, Tanzania.
Western Indian Ocean, Kenya, Mozambique and
Zanzibar. Always among weeds in shallow water, usually
forming large dense schools. Brooding males mix in with
the schools. Deep-bodied for the genus with a tall first
dorsal fin and long ventrals. Body and fins brownish to
yellow with some silver striped barring on cheek and
body, mostly below lateral line. Length to about 8 cm.
Placement of this unusual taxon in Ostorhinchus is

uncertain and needs to be verified. In shape this taxon
looks like it may belong in the Tribe Sphaeramiini.
A
Ostorhinchus nigripes Zanzibar. GRA
B C
D
Ostorhinchus nigripes Several males with oral brood in the school. Kenya. DGO

SUBFAMILY APOGONINAE – TRIBE PRISTIAPOGONINI

Contains two genera: Pristiapogon, and Pristicon.
Diagnosis D. VI+I,9 or VII+I,9; A. II,8; head and body with ctenoid scales; pored lateral-line scales
23–25; preopercle ridge serrate, edges serrate; infraorbitals serrate; three supraneurals; supramaxilla
absent; basisphenoid present; one pair of uroneurals or absent; three epurals; five free hypurals;
parhypural separate; one or two supernumerary dorsal spines; caudal fin forked.
Other characteristics first segmented fin-ray in second dorsal-fin branched, first anal-fin ray
branched and segmented; ctenoid scales on predorsal, cheek, breast, two pelvic scales, and body;
ctenoid scales on opercle and onto base of caudal fin; pored lateral-line scales with multiple pores;
P 12–16; 9+8 segmented principal caudal rays, 15 branched, upper and lower unbranched;
unbranched procurrent rays, longest segmented; teeth in one or multiple rows on premaxilla, dentary,
vomer, palatine, all villiform or absent on palatine; six infraorbitals, bony shelf on third infraorbital;
anterior ceratohyal smooth or notched; developed gill rakers 10–19; 10+14 vertebrae; 8 ribs; 8
epipleurals; low crest on PU22 (after Mabuchi et al, 2014).
Pristiapogon has 6 and Pristicon 7 spines in the first dorsal fin.
TRIBE PRISTIAPOGONINI – Genus Pristiapogon Klunzinger, 1870

Masc. Type-species Apogon fraenatus Valenciennes, 1832.
According to Fraser & Lachner, 1985, Pristiapogon comprises 6 described species: P. abrogramma, P. exostigma,
P. fraenatus, P. kallopterus, P. menesemus, and P. taeniopterus, but the first two are uncertain, whilst P. urostigma
was not included, as it is not typical for the genus. There is much confusion about Bleeker’s type-specimens, but
his description and illustrations are accurate, e.g. Apogon kallopterus is applied to the species shown on the next
two pages. Application of Bleeker’s names here are based on the Bleeker descriptions and his figures.
Spiny-eye Cardinalfish
Pristiapogon fraenatus
Apogon fraenatus Valenciennes, 1832.
New Guinea and Guam.
Widespread Indo-Pacific and Red Sea, with expatriates
ranging to sub-tropical waters. Various shallow coral and
rocky reef habitats, but observed to 47 m depth (D).
Black mid-lateral stripe, pupil width on head, tapering
evenly to end of caudal peduncle, greenish grey above
and pinkish grey below. Normally has a centrally placed
black pupil-size basicaudal spot, but it may be absent at
night or in deep water. Length to about 10 cm. A
Pristiapogon fraenatus Moreton Bay, Queensland. RHK
B C
Pristiapogon fraenatus Night. Maldives. RHK Pristiapogon fraenatus Iriomote Island, Japan. KYA.
D E
Pristiapogon fraenatus D Menjangan Island, Bali. E typically schooling under a ledge. Maumere, Flores, Indonesia. RHK

Tapered-line Cardinalfish Pristiapogon kallopterus

Apogon kallopterus Bleeker, 1856. Manado, Sulawesi.
Widespread Indo-Pacific and Red Sea. Various shallow coral and rocky reef habitats, ranging to 60 m depth on walls.
Brownish grey with black mid-lateral stripe, pupil width on head, tapering evenly to near end of the caudal peduncle.
Posterior edges of dorsal fins slightly rounded. Black streaks on membranes of first dorsal fin distinguishes this taxon from
its siblings (A/B). Usually a black basicaudal spot above line, variable in size up to pupil-size, but occasionally absent. Outer
margins of caudal-fin black. Body depth about 30% in SL (approx 3.6 in length by Bleeker). Length usually to 10 cm.
P. exostigma (Jordan & Starks, 1906), was applied to this taxon in the genus revision (Fraser & Lachner, 1985), whilst P.
kallopterus was applied to a larger species and deeper bodied species, presented here as P. urostigma on following pages,
which up to subadult size has a non-tapering and more ribbon-like black mid-lateral line, usually ending above or in line with
a basicaudal spot and lacks the first dorsal-fin’s black streaks on membranes and dark outer margins of caudal-fin. Fraser
examined the types and is of the opinion that the species shown here as O. kallopterus is O. exostigma and this name was
applied to this taxon in the genus revision (Fraser & Lachner, 1985). We think Bleeker’s illustration and description applies
to the species featured here. The basicaudal spot is clearly above the level of the mid-lateral line.
A B
Pristiapogon kallopterus 10 cm. After Bleeker Pristiapogon kallopterus Type-form. Flores, Indonesia. RHK
C D
Pristiapogon kallopterus Night colours. Japan. TKO Pristiapogon kallopterus Bali, Indonesia. RHK
E
Pristiapogon kallopterus Night colours, with basicaudal spot. Maldives. RHK

F G
Pristiapogon kallopterus Spotless and basicaudal-spot forms on the same coral head. Maldives. RHK
H I
Pristiapogon kallopterus Day and night. Safaga, Egypt, Red Sea. RHK
J K
Pristiapogon kallopterus Type-form. Iriomote I., Japan. KYA Pristiapogon kallopterus Lizard Island, Queensland. RHK
L
Pristiapogon kallopterus Night colours, without a basicaudal spot. Maldives. RHK

Spiny-head Cardinalfish
Pristiapogon urostigma
Amia urostigma Bleeker, 1874. Singapore.
Widespread Indo-Pacific and Red Sea. Various shallow
coral and rocky reef habitats, ranging to 30 m depth.
Large adults are greyish brown, with or without a black
caudal spot, Young with broad, ribbon-like mid-lateral
black stripe and pupil-size black caudal spots in line with
stripe. Posterior margins of dorsal fins slightly concave,
last ray elongated. Opercle and preopercle margins with
finely serrated edges. Body depth 35% or more in SL.
(approx 3.25 in TL by Bleeker). Length to 15 cm.
A Pristiapogon kallopterus, was applied to this taxon in
the genus revision (Fraser & Lachner, 1985).
Pristiapogon urostigma Bali, Indonesia. RHK
B C
Pristiapogon urostigma Spotless form. Sipadan, Borneo. RHK Pristiapogon urostigma Mabul, Borneo. RHK
D E
Pristiapogon urostigma Singapore. After Bleeker Pristiapogon urostigma Sipadan, Borneo. RHK
F
Pristiapogon urostigma Type-form. Note typical posterior shapes of dorsal and anal fins. Bali, Indonesia. RHK

G H
Pristiapogon urostigma Night. Kakeroma, Japan. TKO Pristiapogon urostigma Iriomote Island, Japan. KYA
I J
Pristiapogon urostigma Night. Tulamben, Bali. RHK Pristiapogon urostigma Tulamben, Bali. RHK
K L
Pristiapogon urostigma Juvenile and adult. Aliwal shoals, South Africa. RHK
M N
Pristiapogon urostigma Double basicaudal spot variation. Egypt, Red Sea. RHK

Insular Cardinalfish Pristiapogon taeniopterus

Apogon taeniopterus Bennett, 1836. Mauritius.
Islands of the Pacific and Indian Ocean. Reported from Mauritius, Cocos-Keeling Islands and in the eastern I.O. from
Christmas Island. In the Pacific most oceanic islands, including Micronesia and Polynesia. Dorsally grey with dusky spots on
scales, to pinkish below, Fins with black streak and caudal-fin base mostly black as a large blotch or band and outer rays
black. Length to about 18 cm. A & B are juveniles and C is adult.
A B
Pristiapogon taeniopterus Christmas Island. I.O. GRA Pristiapogon taeniopterus Mauritius. PCH
C
Pristiapogon taeniopterus Mururoa Atoll, French Polynesia. JTW
Band-fin Cardinalfish Pristiapogon menesemus

Apogon menesemus Jenkins, 1903. Oahu Island, Hawaiian Islands.
Reported as Indo-West Pacific, but it has been confused with Pristiapogon taeniopterus and may be more restricted as an
endemic to the Hawaiian region. Reflective brownish. Black steaks in fins are more defined than in P. taeniopterus and it
lacks the black blotch or band on the caudal-fin base. Length to about 18 cm.
A B
Pristiapogon menesemus A night colour. Hawaiian Islands. JHO

TRIBE PRISTIAPOGONINI – Genus Pristicon Fraser, 1972

Masc. Type-species Apogon trimaculatus Cuvier, 1828.
Presently genus Pristicon has 3 described species: P. rhodopterus, P. rufus and P. trimaculatus.
Species of Pristiapogon usually have a darkish single mid-lateral stripe and a basicaudal spot, whilst the Pristicon
taxa have bars or saddles below the dorsal fins and a spot at the base of the caudal fin and/or opercle.
Three-spot Cardinalfish
Pristicon trimaculatus
Apogon trimaculatus Cuvier, 1828.
Moluccas, Indonesia.
Widespread tropical Indo-Pacific. Coastal reef slopes to
outer reef walls to about 20 m depth. Secretive in caves,
usually out at night. Pale grey to brown with vertical and
slightly angled black bars below dorsal fins. Second bar
Y-shaped. Juveniles with a distinct first dorsal-fin, black
above first bar and white posterior margin. Usually a
black round spot on cheek and caudal peduncle, but
sometimes faded. Length to 14 cm.
A
Pristicon trimaculatus Anilao, Philippines. TKO
B C
Pristicon trimaculatus Juvenile. Flores, Indonesia. RHK Pristicon trimaculatus Night colour. Mine Bay, PNG. RHK
D E
Pristicon trimaculatus Maumere, Flores, Indonesia. RHK
F G
Pristicon trimaculatus Mine Bay, PNG. RHK Pristicon trimaculatus Iriomote Island, Japan. KYA

False Three-spot Cardinalfish

Pristicon rhodopterus
Apogon rhodopterus Bleeker, 1852. Singapore.
Western Pacific: from Singapore, Indonesia to southern
Japan, and east to Solomon Islands. Clear coastal reef
slopes to outer reef walls to about 20 m depth. Secretive
in caves, usually out at night. Pale grey to brownish with
a black bar below each dorsal fin and a small peduncular
spot. Scales with a black thin margin. Very similar to
Pristicon trimaculatus, but bar below second dorsal fin
simple and lacks cheek spot. Length to about 15 cm.
A
Pristicon rhodopterus Pulau Putri, Java, Indonesia. RHK
B
Pristicon rhodopterus Palau, Micronesia. HNA
Rosy Cardinalfish Pristicon rufus

Apogon rufus Randall & Fraser, 1999. Malakal Pass, Palau Island.
Western Pacific: Indonesia, Papua New Guinea east to Fiji, north to Ryukyu Islands. Solitary, usually seen on outer reef
slopes in 15–80 m depth. Reddish brown to deep red. Diffused broad dark bars below dorsal fins, small black cheek and
basicaudal spots. Length to about 12 cm.
A B
Pristicon rufus Philippines. ARY Pristicon rufus Kerama Island, Japan. AON

SUBFAMILY APOGONINAE – TRIBE RHABDAMIINI

Contains the genus, Rhabdamia Weber, 1909, but may have subgenera.
Diagnosis D VI+I,9 or VII+I,10–11; A II,11–13; head and body with weakly ctenoid or cycloid; pored
lateral-line scales 23–24; preopercle ridge smooth, edges smooth; two supraneurals; supramaxilla
absent; basisphenoid present or absent; anterior pair of uroneurals reduced or absent; three epurals;
hypurals 1+2+3+4 fused, the plate fused to terminal centrum; parhypural separate; no autogenous
haemal spines; 1–2 supernumerary dorsal spines; caudal fin forked.
Other characteristics first segmented fin-ray in second dorsal-fin branched, first anal-fin ray branched and
segmented; P 13–17; 9+8 segmented principal caudal rays, 15 branched, upper and lower unbranched;
developed gill rakers 17–31; unbranched procurrent rays, longest segmented; villiform teeth in one or two
rows on premaxilla, or with a few canines (Bentuviaichthys), one villiform row on dentary and vomer, one
villiform row or absent on palatine; six infraorbitals, bony shelf reduce or absent on third infraorbital;
stomach and intestine pale with melanophores and silvery peritoneum with melanophores; 10+14 vertebrae;
low crest on PU2 (after Mabuchi et al, 2014).
This tribe remains unresolved at this stage. Some species are yet to be described.
TRIBE RHABDAMIINI – Genus Rhabdamia Weber, 1909

Fem. Type-species Rhabdamia clupeiformis Weber, 1909 (=R. gracilis).
Presently genus Rhabdamia has 4 described species: R. gracilis, R. nigrimentum, R. novaluna, and R. spilota.
The previously included R. cypselura Weber, 1909 is now placed in the genus Verulux Fraser, 1972.
Luminous Cardinalfish Rhabdamia gracilis

Apogonichthys gracilis Bleeker, 1856.
Manado, Sulawesi, Indonesia.
Widespread tropical Indo-West Pacific, but a taxon
which may comprise more than one. Various shallow
coral and rocky reef habitats, ranging to 60 m depth on
walls and rocky outcrops, where usually in schools.
Transparent pale grey and has a tiny black spot on lower
end of the caudal peduncle. Only upper tip of caudal fin
black. Blackish line along anal-fin base, continuing on
caudal peduncle. Length to about 60 mm. A
B C
Rhabdamia gracilis Maumere, Flores, Indonesia. RHK
D E
Rhabdamia gracilis Rowley Shoals, WA. RHK Rhabdamia gracilis Iriomote Island, Japan. TKO

Indonesian Doubles-cardinalfish
Rhabdamia spilota
Rhabdamia spilota Allen & Kuiter, 1994.
Bali, Indonesia.
Southern Indonesia from Bali to West Papua, Indian
Ocean and west to the Red Sea. Coastal reef sand slopes
with outcrops and along reef margins. Typically swims in
tight pairs, occasionally in small groups formed by multiple
pairs. Usually in about 40 m depth and occasionally seen
as shallow as 25 m. Transparent pale grey with a thin
A whitish mid-lateral line and small black spots just below
Rhabdamia spilota Tulamben, Bali, Indonesia. RHK on head and abdominal. Length to about 60 mm.
B C
Rhabdamia spilota Komodo, Indonesia. TTO Rhabdamia spilota Bali, Indonesia. RHK
African Doubles-cardinalfish Rhabdamia sp. 1

Undetermined species, apparently new.
Western Indian Ocean, known only from the upper KwaZulu-Natal coast, reported as Rhabdamia spilota. Coastal reefs,
and may form massive schools numbering thousands. Swims in tight pairs during spawning periods. Transparent pale grey
with a thin whitish mid-lateral line and both caudal fin tips black. Black spots on cranium. Length to about 60 mm.
A B
C
Rhabdamia sp. 1. Sodwana Bay, South Africa. DKI

New-moon Cardinalfish
Rhabdamia novaluna
Rhabdamia novaluna Yoshida, Mabuchi & Motomura,
2018. Kerama Islands, Okinawa, Japan.
Japan, Marshall Islands, Indonesia, Papua New Guinea,
Solomon Islands, Australia and Fiji. Typically swims in
tight pairs or in large schools. Body transparent pale
greyish, reflective silvery on head and abdomen, with
bluish white and blackish mid-lateral lines. Similar to
Rhabdamia gracilis, but the low-set basicaudal spot is
larger, usually more distinct. Length to about 50 mm.
A
B C
Rhabdamia novaluna Triton Bay, West Papua. GRA Rhabdamia novaluna Raja Ampat, West Papua. GRA
Japanese Doubles-cardinalfish Rhabdamia sp. 2

Undetermined species known only from tropical Japan.
Caudal fin with both tips black and no basicaudal spot. It has been confused with the similar Rhabdamia spilota, but lacks
the small black spots just below the mid-lateral line on the head and abdominal. Length to 50 mm.
A B
Rhabdamia sp. 2. Kagoshima aquarium. SDE Rhabdamia sp. 2. Kochi Prefecture, Japan. THI
Black-chin Cardinalfish Rhabdamia nigrimentum

Bentuviaichthys nigrimentum Smith, 1961. Red Sea.
A Red Sea endemic. Reef habitats along margins, types were trawled in 25 m depth. Transparent pale cream with internal
blackish lines, blue on abdomen and an orange-dusky ventral line along anal-fin base and caudal peduncle. Tip of lower jaw
black. Length to 65 mm.
A B
Rhabdamia nigrimentum A Farasan Island. B Dumsuk Island, Red Sea. SBO

SUBFAMILY APOGONINAE – TRIBE VERULUXINI

Contains the single genus Verulux.
Diagnosis D VI–I,9; A II,9; head and body with weakly ctenoid or cycloid scales; pored lateral-line scales 24;
preopercle ridge smooth, edges smooth; one supraneural; supramaxilla absent; basisphenoid present;
uroneurals absent; two epurals; fused hypurals 1+2+3+4, fused to terminal centrum; no autogenous
haemal spines; one supernumerary dorsal spine; bioluminescent organ under cleithrum; caudal fin forked.
Other characteristics first segmented fin-ray in second dorsal-fin branched, first anal fin-ray branched
and segmented; 9+8 segmented principal caudal rays, 15 branched, upper and lower not branched;
developed gill rakers 12–15; unbranched procurrent rays, longest segmented; P 14–16; 10+14
vertebrae; 8 ribs; 5–6 epipleurals; teeth in one row on premaxilla, dentary, vomer, palatine, all villiform,
or absent on palatine; six infraorbitals, bony shelf absent on third infraorbital; stomach and intestine
black with silvery peritoneum; low crest on PU2 (after Mabuchi et al, 2014).
TRIBE VERULUXINI – Genus Verulux Fraser, 1972

Fem. Type-species Rhabdamia cypselurus Weber, 1909.
Genus Verulux has two named species: V. cypselurus, V. solmaculata and one undescribed.
Snout-spot Cardinalfish Verulux cypselurus

Rhabdamia cypselurus Weber, 1909. Kawa, western Ceram, Maluku Province, Indonesia.
Widespread tropical Indo-West Pacific. Coastal reef slopes to outer reef drop-offs to depths of about 50 m, usually adjacent
to deep water with large coral heads. In caves during the day, forming dense schools. spreading out at night, floating well
above the substrate in pursuit of zooplankton. Transparent, reflective internal parts. A small black spot or dash on snout,
and caudal fin with dark streaks on outer rays. Dusky ventral stripe along anal fin and caudal peduncle. Length to 60 mm.
A
Verulux cypselurus Manado, Indonesia. RHK
B C
Verulux cypselurus Tulamben, Bali, Indonesia. RHK Verulux cypselurus Iriomote Island, Japan. KYA
Sun-spot Cardinalfish
Verulux solmaculata
Verulux solmaculata Yoshida & Motomura, 2016.
D'Entrcasteaux Islands, Papua New Guinea.
Eastern Indonesia, Papua New Guinea and western
Australia Schooling on protected coral reefs in about
10–20 m depth. Semi-transparent pale brownish, and
intermittently with a pupil-size black basicaudal blotch; a
dusky longitudinal band on the caudal-fin lobes, and
short black stripe or dash from tip of snout to the eye.
Verulux solmaculata Raja Ampat, West Papua. GRA

Yellow-nose Cardinalfish Verulux sp. 1

A likely Red Sea endemic, known south to the Gulf of
Aden, apparently replacing the widespread Verulux
cypselurus in the region. Body transparent pinkish or
yellowish, reflective internal parts. Blackish along lateral
line. A few minute black spots on snout, and behind eye.
Snout yellow. Vent orange at the start of a series of
dusky spots forming a ventral stripe along anal fin and
caudal peduncle. Length to 60 mm.
B C
Verulux sp. 1. Lahami Bay, Red Sea. SBO
SUBFAMILY APOGONINAE – TRIBE SIPHAMIINI

Contains the single genus Siphamia.
Diagnosis D VI–VII + I,7–11; A II,7–11; pored lateral-line scales 0–24; preopercular ridge smooth,
edges smooth to serrate; supraneurals 1–2; supramaxilla absent; basisphenoid absent; uroneurals absent;
two epurals; hypurals 1+2 and 3+4 fused into two plates, upper plate fused to terminal centrum;
parhypural separate; one or two supernumerary dorsal spines; caudal fin emarginate or forked; bacteria
bioluminescent system from hyal region to along body above or past anal-fin base.
Other characteristics first segmented fin-ray in second dorsal-fin branched, first anal-fin ray branched
and segmented; ctenoid, cycloid or spinoid scales; median predorsal scales 0–6; P 11–16; segmented
principal caudal rays 9+8, 15 branched, upper and lower unbranched; unbranched procurrent rays,
longest segmented; villiform teeth in one or multiple rows on premaxilla, dentary, vomer, palatine, or absent
on palatine; developed gill rakers 6–18; lateral-line scales usually with a vertical row of free neuromasts;
six infraorbitals, bony shelf on third infraorbital; anterior ceratohyal smooth or notched; developed gill
rakers 10–26; 10+14 vertebrae; 8 ribs; 8 epipleurals; stomach, intestine and peritoneum generally pale
with variously sized melanophores; low creston PU2 (after Mabuchi et al, 2014).
TRIBE SIPHAMIINI – Genus Siphamia Weber, 1909

Fem. Type-species Rhabdamia tubifer Weber, 1909.
Mostly tropical taxa, commonly known as siphonfishes; species have strong association with spiny invertebrates
such as long-spined Diadema urchins and Crown-of-thorn Sea-star. Eggs are usually relatively large and a brood
contains about 200 eggs. The recent synonymy (Gon & Allen, 2012) or identity is questionable for a few species
and some are treated as valid here. Genus Siphamia comprises 28 nominal described species:
S. arabica ...................148 S. fraseri.....................154 S. roseigaster ..............143
S. argentea .................151 S. fuscolineata ............146 S. senoui ....................150
S. arnazae...................151 S. goreni.....................148 S. spinicola .................153
S. brevilux ..................153 S. guttulata.................154 S. stenotes ..................154
S. cephalotes ..............142 S. jebbi .......................150 S. tubifer ....................144
S. corallicola...............143 S. majimai ..................147 S. tubulata..................149
S. cuneiceps................142 S. mossambica ............148 S. versicolor ..................145
S. cyanophthalma .......150 S. papuensis ...............152 S. zaribae ......................145
S. elongata .................151 S. permutata...............153
S. fistulosa..................150 S. randalli...................151
Little Siphonfish Siphamia cephalotes

Scopelus cephalotes Castelnau, 1875.
Adelaide, South Australia.
Coastal waters of southern Australia from central New
South Wales and Tasmania to southern Australia and
Western Australia. Usually in small loose groups on low
weedy reefs with Ecklonia kelp or seagrasses to depths
of about 10 m. Greenish grey to a coppery brown in
relation to habitat. Snout short and rounded, whilst jaws
are sub-equal. Fins clear. Brood with gel-like base.
A Hatching after about 14 days. Smallest cardinalfish in
the cool south coast region, length to 50 mm.
In some southern populations the lower jaw is clearly
protruding, looking much like Siphamia cuneiceps.
B C D
Siphamia cephalotes A male with oral brood. B female, typical form. C/D brood removed. Flinders, Victoria. RHK
E F
Siphamia cephalotes Whyalla, South Australia. Type-locality. RHK
G H
Siphamia cephalotes Frankston, Victoria. RHK Siphamia cephalotes Broughton Island, NSW. RHK
Wedgehead Siphonfish Siphamia cuneiceps

Siphamia cuneiceps Whitley, 1941. Fraser Island, Queensland, Australia.
Known from central Queensland to central NSW and Western Australia from the Perth area to Shark Bay, but may also
occur on the south coast, east to South Australia. Shallow to 30 m depth in seagrasses and kelp-reef. ?Synonym of
Siphamia cephalotes, but has fewer gill-rakers, a more pointed snout and protruding lower jaw. Silvery brown, often with
reddish broad stripe mid-laterally and white below on abdomen. A blackish basicaudal blotch. Length to about 50 mm.
A B
Siphamia cuneiceps A male with oral brood. B female, typical form. Sydney, NSW. RHK

Silver Siphonfish Siphamia roseigaster

Apogon roseigaster Ramsay & Ogilby, 1887.
Parramatta River, Sydney, Australia.
Eastern Australia, Queensland and New South Wales.
Reported from the northern territory, but appears to
involve other species in tropical waters. Estuarine, often
in the upper reaches near freshwater. Schooling in rocky
habitats. Semi-transparent pale grey with pinkish hue,
reflective silvery, dusky dorsally. Queensland fish more
transparent and have a small black spot following second
dorsal-fin base. Length to 75 mm in Sydney, NSW.
Smaller, to about 60 mm in tropical Queensland. A
B C
Siphamia roseigaster B night colour. Upper Sydney Harbour, Australia (near type-locality). RHK
D E
Siphamia aff. roseigaster Tropical form. Moreton Bay, southern Queensland. RHK
Coral Siphonfish Siphamia corallicola

Siphamia corallicola Allen, 1993. Madang, PNG.
Indo-Malay Archipelago, Indonesia, Papua New Guinea
to the Philippines. Usually in small groups among
branching corals or sponges, in protected shallow bays
and estuaries. Mostly brownish with silvery white spots
on head, a dash angling up from behind eye axil, some
barring on body and grey blotching dorsally over the
back. Length to 40 mm.
A
Siphamia corallicola Madang, Papua New Guinea. GRA
B C
Siphamia corallicola Bali, Indonesia. RHK

Two-spot Siphonfish Siphamia tubifer

Siphamia tubifer Weber, 1909. Northeastern point of
Timor, Timor Island, southern Malay Archipelago.
Eastern Indian Ocean, Philippines, Indonesia to Papua
New Guinea. Usually in small to large groups in among
long-spined urchins. Either all dark brown with two white
spots, one behind each base of the dorsal fins, often with
series of small brownish dark spots, or white with three
longitudinal black narrow lines, lowest line very narrow.
Similar to Siphamia versicolor, but instead of one it has
2 distinct white spots, behind each dorsal-fin bases and
the axil is dark. Length to 50 mm.
A Name often used throughout the Indo-Pacific for several
Siphamia tubifer Philippines. ARY other similar species that have striped forms.
B C
Siphamia tubifer Milne Bay, PNG. RHK Siphamia tubifer Madang, PNG. GRA
D
Siphamia tubifer Urchin: Asthenosoma varium, host to plain and striped colour forms. Northeast Bali, Indonesia. TTO

One-spot Siphonfish Siphamia versicolor*

Amia versicolor Smith & Radcliffe, 1911.
Cataingan Bay, Masbate, Philippines.
West Pacific, Caroline Islands and north to Japan.
Usually in small loose groups in among spines of
Diadema urchins. Usually completely black, but with a
single white spot behind base of second dorsal-fin and a
white axil. May also shows as a white form with three
narrow black longitudinal lines, lowest line thickest.
Length to 45 mm.
Often treated as a synonym of Siphamia tubifer which
has a white spot following each of the dorsal-fin bases.
*Was not recognised as valid by Gon & Allen, 2012. A
Siphamia versicolor Kashiwajima, Japan. RHK
B C
Siphamia versicolor Kashiwajima, Japan. TKO Siphamia versicolor Kashiwajima, Japan. RHK
Zarib’s Siphonfish Siphamia zaribae*

Siphamia zaribae Whitley, 1959.
Heron Island, Queensland, Australia.
Lizard and Heron Islands. Queensland, Australia. Forms
small groups in association with urchins, Diadema spp,
Crown-of-thorns Seastar and soft corals. Type described
with 6 spines in the first dorsal fin and was illustrated as
such. Eye very large. Juveniles white with three heavy
black longitudinal lines, lower interspace wider than
stripes, upper line thickened to a blotch above posterior
edge of eye. Adults plain shiny greenish dark brown.
Length to 36 mm. A
*Was not recognised as valid by Gon & Allen, 2012. Siphamia zaribae Queensland, Australia. PWO
B
Siphamia zaribae With Crown-of-thorns Seastar. Lizard Island, Queensland, Australia. RHK

Philippines Siphonfish
Siphamia fuscolineata
Siphamia fuscolineata Lachner, 1953. Philippines.
Philippines to southern Japan. Forms small groups in
association with Diadema spp. It clearly shows 7 spines
in the first dorsal fin, first very small. Eye relatively small.
Juveniles white with three heavy black longitudinal lines,
lines wider than interspaces, and reaching caudal-fin
base. Adults plain coppery brown and fins pale reddish
brown, Length to about 35 mm.
A
Siphamia fuscolineata Okinawa, Japan. TKO
B C
Siphamia fuscolineata Mabul, Borneo, Malaysia. RHK Siphamia fuscolineata Okinawa, Japan. TKO
D E
Siphamia fuscolineata Okinawa, Japan. TKO

Tokyo Siphonfish Siphamia majimai

Siphamia majimai Matsubara & Iwai, 1958.
Urasokari, Amami-oshima Island, Japan.
West Pacific, Philippines and north to Japan. Usually in
small loose groups in among spines of Diadema urchins.
Usually blackish brown all over. Has 6 spines in the first
dorsal fin. Fins with tiny red spots. Length to 45 mm.
Whitley’s Siphamia zaribai shares the 6 spines in the
first dorsal fin, but synonymy seems incorrect.
A
Siphamia majimai Taiwan. PCH
B C
Siphamia majimai Hachijo Island, Japan. KTA Siphamia majimai Juvenile. Iriomote Island, Japan. KYA
D
Siphamia majimai Sokodo Bay, Hachijo Island, Japan. SKA

Goren’s Siphonfish Siphamia goreni

Siphamia goreni Gon & Allen, 2012.
Dahlak Archipelago, Dahlak Kebir, ship wreck, Eritrea.
Southern Red Sea and Arabian Gulf, Oman. Various
reef habitats, among spines of sea-urchins in small
groups. Grey to brownish, densely covered with black
spots over head and body. Light-organs end at end of
caudal-peduncle. Length to 35 mm.
A
Siphamia goreni Farasan, Red Sea. SBO
B C
Siphamia goreni Oman. PWO Siphamia goreni Amaq, Red Sea. SBO
Arabian Siphonfish Siphamia arabica

Siphamia arabica Gon & Allen, 2012. United Arab Emirates.
Known from Arabian Peninsula coast of the Gulf of Oman at depths of 3–16 m. Shelters among the spines of sea urchins.
Body usually covered with black speckles and spot all over, giving it an overall brown look with orange fins. Has 7 spines in
first dorsal fin. Length to 40 mm.
A B
Siphamia arabica Gulf of Oman. RFI Siphamia arabica holotype. EHE
Mozambique Siphonfish Siphamia mossambica

Siphamia mossambica Smith, 1955. Mozambique, Africa.
Western Indian Ocean: South and East Africa and Oman to Seychelles, Comoros, Madagascar, Réunion and Mauritius.
Coastal shallow reefs and large tide pools in small loose groups in among spines of Diadema urchins. Usually covered with
black speckles and spot all over, fins peppered with minute red spots. Has 7 spines in first dorsal fin. Length to 40 mm.
A B
Siphamia mossambica Natal, South Africa. PCH Siphamia mossambica Mauritius. JER

Black-coral Siphonfish Siphamia tubulata

Apogon tubulatus Weber, 1909.
Sumbawa Island, Indonesia and Papua New Guinea.
Western Pacific: reported widespread from southern
Japan to northern Australia, but name may involve other
species. Forms small groups in soft corals and among
spines of sea-urchins on reef flats to about 10 m depth.
Whitish silvery with pale greenish brown to blackish
banding deviating from a mid-lateral stripe, forming
various patterns. Length to 50 mm. A
B
Siphamia tubulata Kashiwajima, Japan. RHK
C D
Siphamia tubulata Kochi Prefecture, Japan. THI
E F
Siphamia tubulata Japan. ANA Siphamia tubulata Kashiwajima, Japan. TKO

Red-eye Siphonfish Siphamia fistulosa

Siphamia fistulosa Weber, 1909.
Bima Bay, Sumbawa, Indonesia.
Western Pacific: Indonesia, Brunei to Southern Japan
Lives at moderate depths to about 30 m in branching and
soft corals. Head mostly orange with large orange to red
eye. Body semi-transparent grey, scales on abdominal
region with orange margins. Light-organ ends just past
anal-fin base. Length to 30 mm.
Siphamia senoui Gon & Allen, 2012 was named from
Iriomote Island, but it looks identical to S. fistulosa.
A Counts suggest modally having fewer lateral-line scales,
but it seems to be a junior synonym.
Siphamia fistulosa Komodo, Indonesia. JER
B C
Siphamia fistulosa (S. senoui). B Iriomote Island. C Amami-Oshima, Japan. B KYA C MEN
Jebbs’s Siphonfish Siphamia jebbi

Siphamia jebbi Allen, 1993. Madang, Papua New Guinea.
Indo-West Pacific: Philippines and Malaysia south to Micronesia and to Tonga, Fiji and Australia. Sheltered bays and lagoons.
in 15–30 m depth in small groups among certain corals. Some dark spots, mostly covered with orange dots and dashes over
head and body, fading posteriorly. Length to 30 mm.
A B
Siphamia jebbi Cenderawasih Bay, West Papua. GRA
Blue-eye Siphonfish
Siphamia cyanophthalma
Siphamia cyanophthalmus Gon & Allen, 2012.
Maumere, Flores, Indonesia.
Western Pacific: Indonesia to Papua New Guinea. Found
on sheltered reefs among corals in 15–30 m depth.
Pinkish, body semi-transparent, silvery reflections on
side of head, a pair of blue lines through eye and many
orange spots over head and anterior part of the body.
Siphamia cyanophthalma Raja Ampat, West Papua. GRA

Arnaz’s Siphonfish Siphamia arnazae

Siphamia arnazae Allen & Erdmann, 2019.
Nuakata Island, Milne Bay, Papua New Guinea.
Currently known only from two locations in Milne Bay
Province of eastern Papua New Guinea. In addition to
the type locality it was found at Sideia Island, which lies
approximately 43 km southwest of Nuakata. It occurs on
sheltered reef slopes at depths of 8–20 m. In groups of
about 10–30 fish, mainly in Seriotopora hystrix coral
colonies, but also small Acropora heads with close-set
branches. Translucent pale pinkish to reddish orange
dorsally on head, with dense covering of variable-sized
orange spots on head and anteriorly on body. Eye black
with two vertical bluish white zones on outer of iris.
Siphamia arnazae Milne Bay, Papua New Guinea. MVE
Randall’s Siphonfish Siphamia randalli

Siphamia randalli Gon & Allen, 2012.
Moorea, Society Islands.
Southern Pacific: Society Islands and Cook Islands.
Greenish grey with small black dots dorsally, sides with
large black area posteriorly divided by a white stripe and
reflective silvery over abdominal region and lower part of
head. Length to 30 mm.
Siphamia randalli French Polynesia. RWI
Deepwater Siphonfish Siphamia argentea

Siphamia argentea Lachner, 1953. Northern Balabac Strait, Philippines.
Western Pacific: Philippines and Indonesia. A deepwater species, photographed to 70 m depth and reported from just over
100 m. A moderately deep-bodied species compared to the similar congeners and has a relatively small eye. Pale silvery
brownish with several dark saddle-like blotches and bar on caudal-fin base. Red striations on organ. Length to 40 mm.
A B
Siphamia argentea Bali, Indonesia. GRA Lembeh Strait, Sulawesi, Indonesia. MVE
C D
Siphamia argentea Lembeh Strait, Sulawesi, Indonesia. GRA

Papuan Siphonfish Siphamia papuensis

Siphamia papuensis Gon, Allen, Erdmann & Gouws, 2014. Raja Ampat Islands, West Papua Province, Indonesia.
Only known from the Raja Ampat region. A deepwater species, photographed at 66 m to 70 m depth. Pale brownish and
semi-transparent, with faint dusky banding, anterior ones shallow, forward pointing chevron marks, that tend to partially
merge anteriorly. Length to 40 mm.
A B
Siphamia papuensis holotype. Pulau Balbal, Raja Ampat Islands, West Papua. MVE
C D
Siphamia papuensis Fiabacet, Indonesia. C MVE D GRA
Sinai Siphonfish Siphamia permutata

Siphamia permutata Klausewitz, 1966.
Hurghada, Red Sea Governorate, Egypt, Red Sea.
Red Sea, probably an endemic. Usually in small loose
groups in among spines of Diadema urchins. Usually
plain blackish brown all over, peppered with tiny red
spots, has no white spots following bases of dorsal fins
as in S. tubifer it is confused with. Fins pale brownish.
Pale form with three longitudinal black stripes. Has 6
spines in first dorsal fin. Length to 45 mm.
The name of its senior synonym Beanea trivittata
Steindachner, 1902, was suppressed.
A
Siphamia permutata Sharm el Moya Bay, Red Sea. SBO
B C
Siphamia permutata Obhur, Red Sea. SBO Siphamia permutata Sharm el Moya, Red Sea. SBO

Reef Siphonfish Siphamia spinicola

Siphamia spinicola Gon & Allen, 2012. Kinde Reef, northeast coast, New Caledonia.
Western Pacific: Indonesia south to New Caledonia, Chesterfield Islands and Yap. Coral reefs in 5–20 m depth. Silvery dark
yellow-brown with series of small pale spots forming indistinct vertical lines. Eye as yellow-white. Length to 40 mm.
A B
Siphamia spinicola Sulawesi, Indonesia. SWM Siphamia spinicola New Caledonia.. JER
Slender Siphonfish Siphamia elongata

Siphamia elongata Lachner, 1953. Philippines.
Western Pacific: Philippines, Brunei, Indonesia and
Papua New Guinea. Coastal, often silty habitat with
remote outcrops seeking shelter in branching corals,
crinoids and among sea-urchin spines. A slender species
with dense brown to greenish grey blotching with whitish
interspaces. Length to about 45 mm.
A
Siphamia elongata Mabul, Borneo, Malaysia. RHK
B
Siphamia elongata Milne Bay, Papua New Guinea. PWO
Short-light Siphonfish Siphamia brevilux

Siphamia brevilux Gon & Allen, 2012.
Madang Province, Papua New Guinea.
Western Pacific: Papua New Guinea. Among spines of
Diadema sea-urchins in small groups at 8–30 m depth.
Light-organ not extending beyond anal-fin base. Overall
silvery to reddish brown. Fins transparent pale brown.
Length to 35 mm.
Siphamia brevilux Madang, Papua New Guinea. GRA

Darnley Siphonfish Siphamia guttulata Fraser’s Siphonfish Siphamia fraseri

Apogon guttulatus Alleyne & Macleay, 1877. Siphamia fraseri Gon & Allen, 2012.
Darnley Island, Torres Strait, Queensland, Australia. Kadavu Island, Kavala Bay, Fiji.
Only known from types. Was said to be very numerous at Southwestern Pacific: New Caledonia, Tonga and Fiji. Coral
Darnley Island in holes in the rocks at low water. D, VII, I.8, reefs in 5–20 m depth. Silvery sides, dorsally dark brown with
A. II.8. Silvery, three longitudinal dark lines on sides, lower 2 numerous variable sized dark dots. Length to 40 mm.
reaching tail-base. Length to 30 mm.
Siphamia guttulata after A&M Siphamia fraseri Tonga Mala Island, Vava'u Group. JTW
Lined Siphonfish Siphamia stenotus

Siphamia stenotus Gon & Allen, 2012.
Triton Bay, West Papua, Indonesia.
Known from the type-locality and ranging along
southern Indonesia west to at least Bali. Forms
aggregations among crinoids and sponges on
coastal reefs in 10–15 m depth. Semi-transparent
white and silvery with brownish dark stripe from tip
of snout through eye to base of caudal-fin and a
short line from nape over eye to below gap
between dorsal fins. Length to 30 mm.
A
Siphamia stenotus Triton Bay, West Papua, Indonesia. GRA
B D
Siphamia stenotus North-west Bali, Indonesia. GRA

SUBFAMILY APOGONINAE – TRIBE SPHAERAMIINI

Contains six genera: Apogonichthyoides, Jaydia, Nectamia, Pterapogon, Quinca and Sphaeramia.
Diagnosis D. VII+I,9–10, VII(I)+I,9–14 or VIII+I,9–10 with eighth spine reduced to a tiny visible spine
or a nubbin hidden under skin supported by a free sixth distal radial; A. II,8–13; head and body with
ctenoid scales; pored lateral-line scales 23–27; preopercle ridge smooth, edges serrate; 2–3 supraneurals;
supramaxilla absent; basisphenoid present; one pair of uroneurals; three epurals; five free hypurals or
1+2 fused and 3+4 fused with 3+4 fused to terminal centrum; parhypural separate; two autogenous
haemal spines; two supernumerary dorsal spines; caudal fin forked, emarginate or truncate.
Other characteristics first segmented fin-ray in second dorsal-fin branched, first anal-fin ray branched
and segmented; ctenoid scales on predorsal, cheek, breast, two pelvic scales, and body; ctenoid scales
on opercle and onto base of caudal fin; 1–3 predorsal scales cycloid or ctenoid; pored lateral-line scales
with multiple pores; pectoral-fin rays 13–17; 9+8 segmented principal caudal rays, 15 branched, upper
and lower unbranched; developed gill rakers 7–38; unbranched procurrent rays, longest segmented or
spinous; third or fourth dorsal spine longest; 10+14 or 10+15 vertebrae; 8 ribs; 8–9 epipleurals; teeth
in one or multiple rows on premaxilla, dentary, vomer, palatine, all villiform; six infraorbitals, bony shelf
on third infraorbital; stomach and intestine pale or black with pale peritoneum; low crest on PU (after
Mabuchi et al, 2014).
TRIBE SPHAERAMIINI – Genus Apogonichthyoides Smith, 1949

Masc. Type-species Amia uninotata Smith & Radcliffe, 1912.
Genus Apogonichthyoides comprises 23 described species:
A. atripes....................157 A. gardineri ................160 A. pharaonis ...............159
A. brevicaudatus .........161 A. heptastygma...........157 A. regani.....................163
A. cantoris ..................164 A. maculipinnis...........163 A. sialis ......................161
A. cathetogramma ......161 A. melas .....................158 A. taeniatus ................159
A. chrysurus ...............161 A. miniatus.................158 A. timorensis ..............163
A. enigmaticus ............160 A. niger ......................156 A. uninotatus..............155
A. erdmanni ...............158 A. nigripinnis..............164 A. umbratilis...............162
A. euspilotus...............159 A. opercularis .............164
Centre-spot Cardinalfish
Apogonichthyoides uninotatus
Amia uninotata Smith & Radcliffe, 1912.
Bisucay Island, Cuyos Islands, Philippines.
Known only from the Philippines. Mangrove fringes of estuaries
and brackish lakes. Shallow to about 10 m depth, single or in
pairs with rocky or coral outcrops. Pale brownish with a black
eye-size round spot mid-body, which has a thin pale-yellow halo.
B
Apogonichthyoides uninotatus Philippines. GRA

Black-body Cardinalfish
Apogonichthyoides niger
Apogon niger Döderlein, 1883. Japan.
Northwestern Pacific: northern Vietnam east to
Philippines, north to Japan. Forms loose
aggregations comprising pairs, occasionally in
large groups. Shallow sheltered bays to deep
water on muddy substrates near outcrops of
rock or sponges. Variable pale to dark brown
with blackish blotching or many narrow bars.
Dorsal, ventral and anal fins brown to black.
A
Apogonichthyoides niger Osezaki, Japan. KIM
B D
Apogonichthyoides niger B oral brood. D small juvenile. Japan. C Hong Kong variation. B & D THI C SWO
E F
Apogonichthyoides niger Pair during courtship. Ehime, Japan. RHK

Two-eyed Cardinalfish Apogonichthyoides atripes

Amia atripes Ogilby, 1916. Sandgate, Queensland, Australia.
Tropical and subtropical waters of Australia, south to central New South Wales. Deep in tropical water where usually trawled
with prawns. In New South Wales in coastal bays and estuaries on muddy substrates with rocky out crops or debris that have
hiding spaces. Dark blackish brown with a large yellow-edged black ocellus mid-body below first dorsal fin. Dorsals, ventral
and anal fins with body colour. Caudal fin transparent, whitish to yellow. Length to 9 cm.
A B
Apogonichthyoides atripes Large adults. Sydney Harbour, New South Wales, Australia. RHK
C D E
Apogonichthyoides atripes C 10 mm. D 30 mm. E 45 mm. Sydney Harbour, New South Wales, Australia. RHK
Seven-spot Cardinalfish
Apogonichthyoides heptastygma
Apogon heptastygma Cuvier, 1828. Red Sea.
Red Sea endemic. Sheltered inshore rubble reef habitat,
Secretive in crevices in small groups, coming out at night
on open sandy bottom. Pale brownish grey, with a small
ocellus mid-body below first dorsal fin. Small black spots
along dorsal-fin bases and up to four along mid-line of
caudal peduncle. First dorsal-fin with first long spine and
membrane black, second white. Leading edge of ventrals
white. A small species, largest seen about 45 mm long.
B C
Apogonichthyoides heptastygma Safaga, Egypt, Red Sea. RHK

Black Cardinalfish
Apogonichthyoides melas
Apogon melas Bleeker, 1848. Sumbawa, Indonesia.
Western Pacific and western Australia: Cocos-Keeling
Islands, Western Australia to New Caledonia, north to
Ryukyu Islands. Solitary in coral reefs, hiding in dark
places during the day, coming out at night, but stays
close to reefs. Completely dark brown with a black round
spot in second dorsal fin during the day. Pale with black
banding at night and black spot in second dorsal fin with
white edge. Length to 10 cm.
Its junior synonym Apogon noumeae was named by
Whitley in 1959, based on a fish with night pattern (B)
A which is clearly shown in his illustration of the holotype.
Apogonichthyoides melas Sulawesi, Indonesia. RCS
B C
Apogonichthyoides melas Night colour. Java, Indonesia. RHK Apogonichthyoides melas Iriomote Island, Japan. KYA
Little Red Cardinalfish Apogonichthyoides miniatus

Apogonichthyoides miniatus Fraser, 2010. Cartier Island, Timor Sea.
Southern Indonesia and Timor Sea. A deep water species, known only from trawls. A distinctly coloured species by its red
colour and a large eye-size black spot below first dorsal fin, slightly above eye-level. Length to 50 mm.
A B
Apogonichthyoides miniatus Alor, Indonesia. GRA Apogonichthyoides miniatus Timor Sea. DBR
Erdmann’s Cardinalfish
Apogonichthyoides erdmanni
Apogon erdmanni Fraser & Allen, 2011.
Misool, West Papua, Indonesia.
Known only from a single specimen, collected at 72 m
depth. Pale brown with a dot and two stripes behind and
below eye, a pupil-size black round spot below the first
dorsal fin just above eye-level, followed by some faint
dusky barring. Caudal fin yellow. Leading dorsal spine
darkish. Length to 52 mm.
Apogonichthyoides erdmanni Raja Ampat, West Papua. MVE

Egyption Cardinalfish Apogonichthyoides pharaonis

Apogon pharaonis Bellotti, 1874. Suez, Gulf of Suez, Egypt, Red Sea.
Red Sea, western Indian Ocean: East Africa east to India; Mediterranean Sea (Red Sea immigrant). Solitary or small groups
in rocky reefs and seagrass beds. Pale with black banding below dorsal fins and on caudal peduncle. An ocellus mid-body
below first dorsal fin. Ventrals brown to black. Length to 10 cm.
A B
Apogonichthyoides pharaonis Mediterranean. PWI Apogonichthyoides pharaonis Red Sea. ARY
Two-belt Cardinalfish
Apogonichthyoides taeniatus
Apogon taeniatus Cuvier, 1828. Red Sea.
Red Sea, western Indian Ocean: East Africa, Gulf of
Oman and Persian Gulf to Madagascar; Mediterranean
Sea (Red Sea immigrant). Coastal coral or rocky reefs
and mangroves, forming aggregations. Pale grey with
black bands situated below dorsal fins, an ocellus in the
first band, mid-body below the first dorsal fin. Indistinct
dusky lines along scale rows. Ventrals with white leading
edge and a small black mark on tip. Length to 10 cm.
A
Apogonichthyoides taeniatus Arabian Gulf. GRA
B C
Apogonichthyoides taeniatus Oman. RFI Apogonichthyoides taeniatus Farasan Island, Red Sea. SBO
Sparsely-spotted Cardinalfish
Apogonichthyoides euspilotus
Apogon euspilotus Fraser, 2006.
Ile aux Canards, Noumeb´a, New Caledonia.
Known only from New Caledonia. Life colours
unknown. Preserved brownish with dark spots on head
and body, few in fins. Dark stripes radiating from behind
eye. Caudal fin weakly emarginate with shallow central
indent. Length 7 cm.
Apogonichthyoides euspilotus Holotype. after Fraser

Double-band Cardinalfish
Apogonichthyoides enigmaticus
Apogon enigmaticus Smith, 1961.
Durban, KwaZulu-Natal, South Africa.
Reported as Indo-West Pacific, but maybe restricted to
the west Indian Ocean region. Shallow coastal rocky
reefs in about 10–15 m depth, but also in tide-pools.
Juveniles in small groups. Pale brown, scales with dusky
edges. Dorsal fins with black leading edges. Ventrals with
white leading edges. Juveniles and subadults with narrow
black bars, one below front of each dorsal fin that fade
in large adults. Length to 14 cm.
Recently used Apogonichthyoides dhofar is a junior
synonym. A. pseudotaeniatus Gon, 1986, is putative
synonym as well, as the bars and spot fade in large adults.
A
B C
Apogonichthyoides enigmaticus Oman. PWO Apogonichthyoides enigmaticus Juvenile. Oman. RFI
D E
Apogonichthyoides enigmaticus Subadult. Red Sea. SBO Apogonichthyoides enigmaticus Aqaba. TPA
Gardiner’s Cardinalfish
Apogonichthyoides gardineri
Apogon gardineri Regan, 1908.
Cargados Carajos, Indian Ocean.
Western Indian Ocean, type was collected at 55 m
depth, a Mascarene Plateau endemic. Lives at moderate
depths offshore. Pale brown with three diffused black
bands, saddle-like below each dorsal fin and ring-like on
caudal peduncle. First dorsal-fin’s anterior half blackish
extended from body-band. Eye yellow. Length to 10 cm.
Apogonichthyoides gardineri Mauritius. JER

Two-belt Cardinalfish
Apogonichthyoides sialis
Amia sialis Jordan & Thompson, 1914.
Suruga Bay, Japan.
Amia cathetogramma Tanaka, 1917.
Tanabe, Wakayama Prefecture, Japan.
Japan to China Seas, Brunei and Philippines. Coastal
sheltered bays, forming aggregations on rocky reefs and
open bottom outcrops. Body pale grey with black bands
situated below dorsal fins, and a small basi-caudal spot,
head dusky. The description of Apogonichthoides sialis
states body without markings except basi-caudal spot.
Bands may fade in adults, and it is a presumed synonym. A
Apogonichthyoides sialis Ehime, Japan. RHK
B C
Apogonichthyoides sialis Ehime, Japan. THI Apogonichthyoides sialis With oral brood. Ehime, Japan. RHK
High-fin Cardinalfish
Apogonichthyoides chrysurus
Apogon chrysurus Ogilby, 1908. Lord Howe Island.
Southwestern Pacific, southern Queensland, and Lord
Howe Island region. Protected coastal bays, harbours
and lagoons. Usually only seen at night, drifting in a
floating way above the substrate. Deep-bodied, plain
pale yellowish to brownish, may show indistinct barring
on deepest part of body. Dorsal fins tall, joined at bases,
first fin tallest. Length to 10 cm.
Apogonichthyoides chrysurus was placed in Vincentia
at some stage. A record from Heron Island, north of its
geographical range, is based on Apogonichthys ahimsa
Whitley, 1959, misplaced in synonymy, a different small
more slender taxon with 1 predorsal scale, which is close
to or may be synonymus with A. perdix. Apogonichthyoides chrysurus Lord Howe Island. NCO
Many-band Cardinalfish Apogonichthyoides brevicaudatus

Apogon brevicaudatus Weber, 1909. Jedan Island, Aru Islands, Indonesia.
Northern half of Australia and Indonesia. Usually trawled offshore. Pale brown to yellowish with longitudinal brown stripes
along scale-rows. Fins brown to dusky, a black spot basally on second dorsal and anal fins. Length to 11 cm.
A B
Apogonichthyoides brevicaudatus A northern WA. B Montcomery Reef. Western Australia. GRA

Cryptic Cardinalfish
Apogonichthyoides umbratilis
Apogon umbratilis Fraser & Allen, 2010.
Northwest Shelf, Western Australia.
West Pacific from northern Australia and Papua New
Guinea and north to Brunei. Type was trawled in 46 m
depth. Secretive and solitary. Pale brown, with faint dark
banding below dorsal fins and on caudal peduncle. Short
dark dashes radiating from behind eye. First dorsal fin
whitish with a dark centre or black blotch. Ventral and
A anal fins with white markings. Length to 6 cm.
Apogonichthyoides umbratilis Milne Bay, PNG. RCS
B C
Apogonichthyoides umbratilis West Papua, Indonesia. GRA Apogonichthyoides umbratilis Brunei. GRA
Japanese Cryptic Cardinalfish Apogonichthyoides aff. umbratilis

Undetermined taxon, only known from southern Japan. Occurs in shallow protected coastal bays to about 10 m depth on
rocky reef crests and outcrops of rocks on silty substrates. Secretive and solitary. Pale brown, with faint dark banding show-
ing more distinct at night, one each below dorsal fins and on caudal peduncle. First dorsal-fin whitish, sometimes with a dark
or black blotch centrally. Length to 6 cm.
Lacks strong body markings, but night-pattern very similar to Apogonichthyoides umbratilis.
B C
Apogonichthyoides aff. umbratilis A male. B female. C night. Ehime, Japan. A/B RHK C THI

Timor Cardinalfish
Apogonichthyoides timorensis
Apogon timorensis Bleeker, 1854.
Kupang, Timur, Timor Sea, eastern Indian Ocean.
Red Sea, Indo-West Pacific: East Africa and Madagascar
east to Solomon Islands, north to Taiwan, south to
northern Australia. Secretive shallow coastal reefs and
lagoons, but also deep offshore on muddy substrates.
Pale yellowish or brownish to dusky with diffused broad
dark banding. Whitish blotch with black edges angling
backwards from behind lower part of eye. Fins yellowish.
A
Apogonichthyoides timorensis Lembeh Strait, Sulawesi. RHK
B C
Apogonichthyoides timorensis Bali, Indonesia. RHK Apogonichthyoides timorensis Timor, Indonesia. CBA
Speckle-head Cardinalfish
Apogonichthyoides maculipinnis
Apogon maculipinnis Regan, 1908.
Haddhunmathi Atoll, Maldives, Indian Ocean.
Known only from the Maldives, the type-specimen and
the individual in the photograph, which as taken at 40
m depth. Secretive under objects near rubble reef during
the day. Colour brownish grey with dusky banding and
small dark spots. Short dark dashes radiating from
behind eye. Fins with tiny spots and first dorsal-fin with
black blotch. Length to about 10 cm.
Apogonichthyoides maculipinnis Maldives. MVE
Regan’s Cardinalfish
Apogonichthyoides regani
Apogon regani Whitley, 1951. Replacement name for
Apogon punctatus Regan, 1908. Indian Ocean.
Western Indian Ocean. Lives at about 50 m depth.
Brownish grey with small black spot in longitudinal series
on body scales. Fins dusky. Length to about 75 mm.
Apogonichthyoides regani Seychelles. JER

Pearly-cheek Cardinalfish
Apogonichthyoides opercularis
Apogon opercularis Macleay, 1878.
Port Darwin, Northern Territory, Australia.
Australia: Queensland to Western Australia. Mainly
known from trawls. Colour dull vinous red, including fins
except the pectorals more or less black; a black mark
extending from beneath the eye to angle of preopercle,
and opercle there is a margaritaceous patch with a black
patch above it (according to original description). Length
to about 10 cm.
Apogonichthyoides opercularis Darwin, Australia. JER
Riouw Cardinalfish
Apogonichthyoides cantoris
Apogon cantoris Bleeker, 1851. Riouw, Indonesia.
A taxon only known from Bleeker’s description. He had
obtained 3 specimens from Bintang and Ambon, nearly
12 cm long. Colour given by Bleeker as illustrated.
Undetermined and uncertain genus placement.
Apogonichthyoides cantoris Indonesia. after Bleeker
Black-finned Cardinalfish
Apogonichthyoides nigripinnis
Apogon nigripinnis Cuvier, 1828.
Java, Indonesia; Puducherry, India.
Widespread Indian Ocean: South Africa to Java. Brown
with three broad black bands. Fins except pectorals
mostly black. Caudal fin slightly rounded. An eye-size
ocellus mid-body below the first dorsal fin, but usually is
largely obscured in the black band. Length to 14 cm.
Name is commonly applied in error to similar species
that show an ocellus below the first dorsal fin at about
eye-level, including Apogonichthyoides atripes, A.
pharaonis, and A. taeniatus
B C
Apogonichthyoides nigripinnis Durban, South Africa. DKI Apogonichthyoides nigripinnis India. after Day

TRIBE SPHAERAMIINI – Genus Jaydia Smith, 1961

Fem. Type-species Apogon ellioti Day, 1875.
Genus Jaydia comprises 17 described species:
J. albomarginata .........169 J. hungi...................167 J. quartus .................170
J. argyrogaster ............170 J. lineata.................168 J. queketti ................167
J. carinata ..................166 J. melanopus...........166 J. striata...................170
J. catalai.....................170 J. novaeguineae.......169 J. striatodes..............170
J. ellioti ......................165 J. photogaster .........168 J. truncata................169
J. erythrophthalma .....170 J. poeciloptera ........165
Usually moderately-deep water (40–100 m), mud dwelling species utilising holes in the bottom for shelter.
Synonymy treatments of certain species in the genus revision (Gon, 1995) could not be supported. Such taxa
presented here as valid are commented on in the species-accounts.
Elliot’s Cardinalfish Jaydia ellioti

Jaydia ellioti Day, 1875. India.
Indo-West Pacific and Red Sea: Gulf of Oman east to
Philippines and Marshall Islands, north to Taiwan; and
Mediterranean Sea (Red Sea immigrant). Inshore in
coastal bays and harbours, relatively shallow to 80 m
depth. Pale tan whitish grey, dusky dorsally. Dorsal fins
white, first with tip broadly black, second and anal fin
with a black submarginal stripe. Caudal fin rounded with
a black posterior margin. A black line angling backwards
from below eye. Length to 15 cm.
Synonymy of this rounded-tail taxon with the square-tailed
Jaydia truncata by Gon, 1995 seems incorrect, whilst J.
smithi Kotthaus, 1970, is a putative synonym.
Jaydia ellioti From trawl, Red Sea. SBO
Yellow-fin Cardinalfish Jaydia poeciloptera

Apogon poecilopterus Cuvier, 1828. Java, Indonesia.
Indo-West Pacific: India east to Philippines, north to
southern Japan, south to northern Australia and New
Caledonia. Usually trawled inshore on muddy bottoms
near river mouths. Line in holes in the bottom during the
day, observed at 25 m depth (below). Grey, head dark,
scales with dusky edged bars, that form as longitudinal
lines in a zig-zag style along scale-rows on upper sides.
Ventrals, anal and caudal fins yellow, of which the anal
fin is brightest in colour. First dorsal with dark tip.
Length to about 14 cm. A
Jaydia poeciloptera From trawl, northern Australia. CSIRO
B C
Jaydia poeciloptera In a typical mud-habitat at 25 m depth. North-eastern Sulawesi, Indonesia. RHK

Monster Cardinalfish Jaydia melanopus

Apogon melanopus Weber, 1911.
Wardakau, Aru Island, Indonesia.
West Pacific, Indonesia, Western Australia and Northern
Territory (Australia). Trawled in coastal waters. Reported
from 2–62 m depth. Yellowish brown to silvery grey, a
dusky band below first dorsal fin and dusky blackish spot
at each end of dorsal and anal fins bases or forming a
second band. Fins dusky. Caudal fin emarginate. Length
to about 15 cm.
Jaydia melanopus From trawl, northern WA. CSIRO
Blue-finned Cardinalfish Jaydia carinata

Apogon carinatus Cuvier, 1828. Japan.
Western Pacific and southeastern Indian Ocean. Trawled in depths of about 50–60 m depth, and is rarely seen shallower.
Underwater photographs were taken at 23 and 40 m depth on a muddy substrate. Body grey with faint broad banding.
Dorsal, anal, ventral and caudal fins mostly grey to yellow. Second dorsal fin posteriorly with a basal eye-size white-edged
black spot. Ventral and anal fins grey to yellow, but may turn deep blue at night. Anal fin with black margin. Caudal fin
rounded. A dusky wedge-shaped short bar angling backwards from below eye. Length to 12 cm.
A B
Jaydia carinata Japan. ANA Jaydia carinata Kanagawa Prefecture, Japan. HSE
C
Jaydia carinata Night colours. Suruga Bay, Japan. SNA

Hungi Cardinalfish Jaydia hungi

Jaydia hungi Fourmanoir & Nhu-Nhung, 1965. Nossi Be, Madagascar.
Indian Ocean: East Africa, Comoros and Madagascar to Indonesia, at depth range of 18-62 m. Inclusion of Indonesia is
based on the photograph. Whitish grey below, dusky dorsally, extended with long saddle-like marking over sides. Caudal fin
rounded. First dorsal-fin white with the tip broadly black. Second dorsal fin with dusky bands, along margin and up from
base. Anal and ventral fins plain white or bluish. A short black line angling backwards from below eye. Length to 10 cm.
Jaydia hungi Bali, Indonesia. MYA
Quekett’s Cardinalfish Jaydia queketti

Apogon queketti Gilchrist, 1903. KwaZulu-Natal, South Africa.
Red Sea, western Indian Ocean: South Africa to Persian Gulf, and to east India; Mediterranean Sea immigrant from Red
Sea. Mainly trawled on sandy bottom to about 75 m depth. Silvery whitish grey, body with series of brown spots. Fins
whitish, first dorsal fin with a large black blotch. Caudal and anal fins with a black margin. Length to 10 cm.
Jaydia queketti Madras, India. JER

Multi-bar Cardinalfish Jaydia striata Grey-banded Cardinalfish Jaydia striatodes

Amia striata Smith, 1912. Luzon, Philippines. Apogon (Jaydia) striatodes Gon, 1997. Andaman Sea.
Indo-West Pacific: Pakistan to Philippines to Papua New Eastern Indian Ocean, western Pacific: Andaman Sea east to
Guinea, Malaysia, and Thailand. Trawled at moderate Philippines, north to northern Vietnam and southern China.
depths. Pale yellowish grey to silvery with 9–11 black narrow Trawled in depths of about 100 m. Body pale grey to silvery
bars on sides. Fins yellowish, often with faint dusky broad with 6–9 dusky bars on sides. Eye yellowish. Caudal fin
margins. Caudal fin slightly rounded. Length to 10 cm. rounded. Length to 10 cm.
Jaydia striata Ryoung, Thailand. HMO Jaydia striatodes Beibu Gulf, China. ZYU
Ten-bar Cardinalfish Jaydia lineata

Apogon lineatus Temminck & Schlegel, 1843.
Nagasaki, Japan
Indo-West Pacific, but mainly known from the northern
West Pacific. Primarily deepwater, usually trawled on
sandy bottom to about 100 m depth, but adults have
been observed at night by divers in Japan as shallow as
3 and 4 m depth. Silvery grey, body with about 8–10
black thin bars along sides from behind head, to last one
basicaudal. Caudal fin usually slightly rounded, but shape
variable to truncate or emarginate with shallow central
A indent. Length to about 10 cm.
Jaydia lineata Akita Prefecture, Japan. HSU
B C
Jaydia lineata Male with oral brood. Fukui Prefecture, Japan. TTA Jaydia lineata Japan. MHA
D
Jaydia lineata Subadult. Nagasaki, Japan. TKO

New Guinea Cardinalfish

Jaydia novaeguineae
Apogon novaeguineae Valenciennes, 1832. New Guinea.
Known from Java, southwestern Indonesia to Papua
New Guinea Usually trawled in shallow depths. Body
with faint chevron bars, interspaced with iridescent pale
blue which extends over side of head. Dorsal profile of
nape convex. Eyes yellowish white. Fins all clear. Caudal
fin distinctly emarginate. Length to about 85 mm.
The inclusion of other species as synonyms may cause
erroneous reporting of geographical distributions.
Jaydia novaeguineae Madang, Papua New Guinea. GRA
White-edged Cardinalfish Jaydia albomarginata

Amia albomarginata Smith & Radcliffe, 1912. Fish market, Cavite, Luzon Island, Philippines.
Red Sea and Indo-West Pacific. Trawled on sandy of muddy substrates in 40–100 m depth. Plain silvery grey, with whitish
reflective sides of head and body. Eyes yellow. First dorsal-fin with dusky tips over five spines. A faint sub-marginal dusky
line in second dorsal fin. Ventral and anal fins with white leading margins and caudal fin with a distinct white lower edge.
Dorsal fins with leading black margins and upper edge of caudal fin with a dusky margin. Caudal fin truncate to emarginate.
Length to about 10 cm. Synonymy by Gon, 1995, with the above shown Jaydia novaeguinae, seems incorrect.
A B
Jaydia albomarginata Red Sea. SBO Jaydia albomarginata Southern Japan. HSE
Flat-tail Cardinalfish Jaydia truncata

Apogon truncatus Bleeker, 1854.
Padoz Reef, Madang Lagoon, Papua New Guinea.
Reported from the Persian Gulf, India, Burma, Thailand,
Indonesia and New Guinea, but some records may be
based on other species. Depth range 15–110 m. Plain
pale silvery whitish brown with faint dusky bars in life,
reported to show only after preservation. Head with
dark area above eye and dorsal profile of nape flat or
slightly concave. Eyes pale yellow. Caudal fin typically
truncate to slightly indented centrally. Top of first dorsal
fin broadly blackish. Length to 6 cm.
Several similar species, but with a rounded caudal-fin,
A
that were were incorrectly put in synonymy.
Jaydia truncata Adult. Oman. JER
B C
Jaydia truncata after Bleeker Jaydia truncata Subadult. Mindoro, Philippines. JTW

Silver-cheek Cardinalfish Jaydia photogaster

Apogon photogaster Gon & Allen, 1998. Padoz Reef, Madang Lagoon, Papua New Guinea.
Known only from Madang, Papua New Guinea and Waigeo. Solitary or small groups in lagoon patch-reefs in 18–52 m
depth. Pale yellowish grey to silvery with 9–10 dusky bars on sides. Silvery light-organ on belly. Length to 6 cm.
A B
Jaydia photogaster Raja Ampat, West Papua, Indonesia. A GRA B MVE
Brown-spotted Cardinalfish Jaydia argyrogaster Quart Cardinalfish Jaydia quartus

Apogon argyrogaster Weber, 1909. West New Guinea. Apogon quartus Fraser, 2000.
Northwestern Australia to West Papua. Lives in sand rubble Saya de Malha Bank, Indian Ocean.
habitats in gorgonians and sponges in 15–86 m depth. Pale Know only from the type-location, taken in depths of about
greyish brown with many pupil-size brown spots on body. 60 m. Life colours unknown. Has an ocellus like marking on
Short black lines radiating from behind eyes. Length to 6 cm. side below first dorsal-fin at eye-level. Length to 65 mm.
Jaydia argyrogaster West Papua, Indonesia. GRA Jaydia quartus Holotype. after Fraser
Catala’s Cardinalfish Jaydia catalai Orange-eye Cardinalfish

Apogon (Nectamia) catalai Fourmanoir, 1973. Jaydia erythrophthalma
New Caledonia. Jaydia erythrophthalma Gon, Liao & Shao, 2015.
Known only from New Caledonia from about 15 m depth. West Pacific: Philippines. Reef-associated, caught in depth
Little known species, probably reef associated. Deep-bodied range of 115–469 m. Body whitish with a hint of orange.
for the genus, grey in colour with some dark mottling and Eye orange, upper sides of body with 2 series of dark-brown
has a yellow eye. Dorsals, anal and ventral fins dusky. Caudal spots (suffused with orange in life) of which largest spots of
fin slightly rounded. Length to about 7 cm. top series with a pale centre. Length to 75 mm.
Jaydia catalai New Caledonia. JER Jaydia erythrophthalma Philippines. after authors

TRIBE SPHAERAMIINI – Genus Nectamia Jordan, 1917

Fem. Type-species Apogon fuscus Quoy & Gaimard, 1825.
Genus Nectamia comprises 9 described species:
N. annularis................174 N. ignitops..................176 N. similis ....................177
N. bandanensis ...........175 N. luxuria ...................176 N. viria .......................175
N. fusca ......................172 N. savayensis ..............173 N. zebrina...................174
Dusky looking species, collectively known as the ”bandanensis” group. Members are variously distributed in
the tropical Indo-Pacific and Red Sea. Reef dwellers, secretive during the day sheltering in caves, ledges and
often in small groups among the bases of large branching coral-heads. Ventures out solitary at night to feed
on open substrates, usually staying close to their shelter.
Key to the adult species of the Nectamia (after Fraser, 2008)

1. Oblique cheek mark expanded near eye; basicaudal spot, saddle or bar; body depth generally greater than 42% of
standard length....................................................................................................................................................2
Oblique cheek mark narrow throughout length; basicaudal spot central and diffuse in adults; body depth generally less
than 42% of standard length. Indo-Central Pacific .............................................................................Nectamia fusca
2. Lobes of the caudal fin with dark bands near upper and lower edges ........................................................................3
` Caudal fin dusky, no dark bands in lobes near upper and lower edges ......................................................................6
3. Well developed light bars on side of body, may have narrow or broad dark areas between the light bars .....................4
No well developed light bars, occasionally a few faint light bars. Western Indian - Central Pacific..................................
............................................................................................................................................. Nectamia savayensis
4. Light bar present on the opercle flap, a few narrow light bars on body, dorsal saddles under the first and second dorsal
fins. West Pacific ..........................................................................................................................Nectamia similis
No bar on the opercle flap, no dorsal saddle under the first and second dorsal fins ....................................................5
5. Pale bars with wide darker area on body, black caudal-peduncle saddle extends below lateral line but not as a complete
uniform band. Red Sea-Gulf of Aden ............................................................................................Nectamia zebrina
Caudal-peduncle mark above lateral-line; pale bars with narrow dark bars on body. Indian-Central Pacific .....................
...................................................................................................................................................Nectamia luxuria
6. Dark caudal-peduncle band extend below lateral-line scales; dorsal saddles present or faint .........................................7
No pale bars; no dorsal saddles; broad cheek mark. Red Sea........................................................Nectamia annularis
7. Two dorsal saddles, no pale bars on body...............................................................................................................8
Dorsal saddle present under second dorsal fin; a few faint pale bars sometimes present. West Pacific......Nectamia viria
8. Total gill rakers and rudiments usually 26–29. West Pacific ......................................................Nectamia bandanensis
Total gill rakers and rudiments usually 24–27. South China Sea ....................................................Nectamia ignitops
Some individuals could not be identified based on colourations that were seemingly derived from different taxa, suggesting
hybridisation between sibbling species may be taking place.
Nectamia sp. Maldives. RHK Nectamia sp. Iriomote Island, Japan. KYA
This fish looks very much like Nectamia annularis from the This fish is possibly Nectamia savayensis. but fins are all
Red Sea, but this species is not known from the Maldives. clear, lacking the normal or white and black margins. It
The ring-like band on the caudal peduncle is grey instead of shows no dark or light barring on the body to suggest any
black. Possibly a variation of N. savayensis. other known species.

Ghost Cardinalfish Nectamia fusca

Apogon fuscus Quoy & Gaimard, 1825.
Guam, Mariana Islands, western Pacific.
Widespread Indo-West Pacific and Red Sea. Solitary or
small groups on coral or rocky reefs in about 10–20 m
depth. Plain brownish to yellowish grey, and body may
show indistinct barring on deepest part and a diffused
dark blotch on caudal peduncle, which shows distinct at
night only. A black thin line angling backwards from
below eye. Length to about 85 mm.
Name Ostorhinchus guamensis Valenciennes, 1852,

often used for this species, is a junior synonym.
A
B C
Nectamia fusca C night. Safaga, Egypt, Red Sea. RHK
D
Nectamia fusca A typical adult. Maumere, Flores, Indonesia. RHK
E F
Nectamia fusca ? Sizes 45 and 50 mm, possibly subadult colouration. Sydney, Australia. RHK

Savay Cardinalfish Nectamia savayensis

Apogon savayensis Günther, 1872. Savay, Samoa.
Widespread Indo-West Pacific. Usually solitary on coral or rocky reefs in about 5–25 m depth. Plain brownish grey, dusky
on back, a grey to black short saddle on caudal peduncle, usually all above lateral line. A black wedge-shaped bar angling
backwards from below eye. White horizontal short line(s) in eye. Dorsal fins with dusky to black anterior margins, caudal fin
with blackish and white margins on outer rays. Length to 11 cm.
A
Nectamia savayensis Maumere, Flores, Indonesia. RHK
B C
Nectamia savayensis B night. Manado, Indonesia. RHK
D E
Nectamia savayensis Night. Maldives. RHK

Zebra Cardinalfish Nectamia zebrina

Apogon zebrinus Fraser, Randall and Lachner, 1999.
Egypt, Gulf of Aqaba, Red Sea.
Solitary or small groups on coral or rocky reefs in about
15 m depth. Solid dark grey dorsally, extending down
into 7–8 bars on sides with narrow whitish interspaces.
A black band on caudal peduncle, broad dorsally, and a
wedge-shaped black line angling backwards from below
eye. Length to about 85 mm.
B C
Nectamia zebrina Safaga, Egypt, Red Sea. RHK Nectamia zebrina Red Sea. JNE
Black-ring Cardinalfish Nectamia annularis

Apogon annularis Rüppell, 1829.
Sinai Coast, Egypt, Gulf of Suez.
Solitary and secretive on rubble reefs to about 10 m
depth, usually seen drifting over open substrates at night.
Head and body plain grey, caudal peduncle whitish with
a black complete band. A thin wedge-shaped black line
angling backwards from below eye. Fins yellowish with
dark and white leading edges. Length to 70 mm.
B C
Nectamia annularis C night. Safaga, Egypt, Red Sea. RHK

Banda Cardinalfish Nectamia bandanensis

Apogon bandanensis Bleeker, 1854. Banda, Indonesia.
Indo-West Pacific: Cocos-Keeling and Christmas Island,
Indonesia east to Marshall Islands and Samoa, Australia
north to southern Japan. Solitary or small groups on
coral or rocky reefs in about 15 m depth. Pale brownish,
caudal peduncle whitish, posteriorly with a broad dark
dusky band, black above lateral line. Shows three broad
bands at night. A well defined wedge-shaped black line
angling backwards from below eye. Length to 10 cm.
A
Nectamia bandanensis Gilimanuk, Bali, Indonesia. RHK
B C
Nectamia bandanensis Night. Sulawesi, Indonesia. RHK Nectamia bandanensis Night. Philippines. HFR
Bracelet Cardinalfish Nectamia viria

Apogon viria Fraser, 2008. Sangay Siapo Islands, Philippine Islands.
Western Pacific: Malaysia, Indonesia and Philippines to Australia and to Fiji. In small aggregation on shallow coral rich reefs
or coarse rubble accumulations. Dusky brown, 3–4 narrow pale bars on deepest part of the body, caudal peduncle largely
white with a broad black and brown band, of which upper half black. Caudal-fin outer margins yellow. A sharply defined
wedge-shaped black line angling backwards from below eye. Length to about 10 cm.
A B
Nectamia viria Gilimanuk, Bali, Indonesia. RHK Nectamia viria Maumere, Flores, Indonesia. RHK
C D
Nectamia viria Palau, Micronesia. HNA Nectamia viria Bali, Indonesia. RHK

Yellow-edged Cardinalfish Nectamia luxuria

Apogon luxuria Fraser, 2008. Ceram, Molucca Islands, Indonesia.
Indo-West Pacific: Maldive Islands (Indian Ocean) to central Pacific, Australia to Philippines. Coastal to outer reef lagoons.
Low in corals during the day, usually in small groups. Distinct by the bright-yellow leading margins on median fins. Body with
many narrow grey bars on deepest part and a black saddle mark on caudal peduncle. Length to 70 mm.
A
Nectamia luxuria Andaman Sea. USA
B C
Nectamia luxuria Flores, Indonesia. RHK
Fire-eye Cardinalfish Nectamia ignitops

Apogon ignitops Fraser, 2008. Tioman Island, Malaysia.
Northern West-Pacific: known from Malaysia, Gulf of Thailand, Vietnam, and north to Japan. Secretive in crevices and on
sheltered coral-rich reefs forming aggregations low in staghorn corals. Distinct by the bright red eye. Silvery, brownish, dusky
dorsally and head, caudal peduncle pale with a dusky band. A short wedge-shaped dark line angling backward from below
eye and a narrow yellow to yellow-brown stripe along mouth below the eye. Length to about 10 cm.
A B
Nectamia ignitops Kerama Island, Japan. MAT Nectamia ignitops Iriomote Island, Japan. TKO

Look-alike Cardinalfish Nectamia similis

Apogon similis Fraser, 2008. Kabaena Island, Banda Sea, Indonesia.
Western Pacific: Malaysia, Indonesia, Papua New Guinea and Queensland (Australia) to Philippines and to Palau. Sheltered
coral reefs, in dense branching corals during the day to about 10 m depth. Dusky grey with few narrow pale bars on deep
part of body. Peduncular band dusky with black blotch above lateral line. Length to 70 mm.
A B
Nectamia similis A night. Maldives. RHK
TRIBE SPHAERAMIINI – Genus Xeniamia Fraser & Prokofiev, 2016

Fem. Type-species Xeniamia atrithorax Fraser & Prokofiev, 2016.
Genus Xeniamia comprises a single species: P. atrithorax.
Black-chest Cardinalfish
Xeniamia atrithorax
Xeniamia atrithorax Fraser & Prokofiev, 2016. South
China Sea off Khanh Hoa Province, Vietnam.
South China Sea, Vietnam and southern Taiwan. Most
specimens were collected in shrimp trawls on muddy
substrates from depths of 70–119 m. Usually this
species appeared in groups of about 10–60 specimens
per trawl. Pale semi-transparent-cream with a hint of
orange and a black region from ventrals to anus. Length
to 35 mm. Appears to be closely related to Siphamia,
but lacks the luminous organ.
A
B
Xeniamia atrithorax Freshly caught off Dong-gang, Pingtung, Taiwan. A HMO B KKO

TRIBE SPHAERAMIINI – Genus Pterapogon Koumans, 1933

Fem. Type-species Pterapogon kauderni Koumans, 1833.
Genus Pterapogon comprises a single species: P. kauderni.
Banggai Cardinalfish Pterapogon kauderni

Pterapogon kauderni Koumans, 1833. Banggai Islands.
An attractive cardinalfish, unique among marine fishes in not
only incubating eggs like other marine cardinalfishes, but
also accommodating hatchings for several days in the
mouth. Of all apogons it probably has the largest eggs,
measuring 2.5 mm in diameter, and fewest in number, with
up to about 20. Juveniles stay together and seek protection
among the long spines of Diadema urchins in seagrass
habitat in shallow depths, just below low tide mark, to about
2 m depth and also use stinging anemones for protection.
Being highly developed at birth is the reason for the limited
dispersal of their offspring, in contrast to species such as
Ostorhinchus aureus, that has thousands of tiny eggs and
pelagic fry, which consequently is very widespread in the
Indo-Pacific. Length to 7 cm.
Originally it was only known from the Banggai Islands, off
central-east Sulawesi. The recent re-discovery of this species
has led to great interest from the aquarium trade and this
A attractive species was also introduced to other several other
Pterapogon kauderni Banggai Islands, Indonesia. FSN islands in the region by collectors.
B C
Pterapogon kauderni Gravid female and courtship. Hong Kong aquarium. RHK
D
Pterapogon kauderni Semi-adults and juveniles, typical behaviour. Banggai Islands. GRA

E F
Pterapogon kauderni Hatchling, aquarium. TKO Pterapogon kauderni Captured with hatchlings. GRA
G H
Pterapogon kauderni Juveniles with anemone. GRA Pterapogon kauderni Juvenile, aquarium. TKO
I
Pterapogon kauderni Schooling above refuge-providing branching coral. Banggai Islands. GRA

TRIBE SPHAERAMIINI – Genus Quinca Mees, 1966

Fem. Type-species Quinca mirifica Mees, 1966.
Genus Quinca comprises a single species: Q. mirifica.
Sail-fin Cardinalfish Quinca mirifica

Quinca mirifica Mees, 1966. Cockatoo Island, Yampi Sound, Kimberley Division, Western Australia.
It appears to be restricted to a small region off north Western Australia from about Exmouth to Derby, the Buccaneer Archipelago.
It lives in shallow coastal protected reefs in caves and ledges. A moderately deep-bodied species, but looks extra high with the large
and tall dorsal, anal and ventral fins. Completely dark brown to nearly black, but may show pale saddle-like markings over the back
or white areas during courtship. Length to 14 cm.
A B
Quinca mirifica Kimberley and Exmouth regions. A GRA B NCO
TRIBE SPHAERAMIINI –
Genus Sphaeramia Fowler & Bean, 1930
Fem. Type-species Apogon nematopterus Bleeker, 1856.
Genus Sphaeramia comprises two species: S. orbicularis
& S. nematoptera.
Polka-dot Cardinalfish Sphaeramia orbicularis
Apogon orbicularis Cuvier, 1828. Java, Indonesia.
Red Sea, East Africa east to Caroline Islands, Gilbert Islands, Fiji and
Tonga, north to southern Japan, south to southern Indonesia and
New Caledonia to northern Australia. In mangroves and harbours, often
in surface waters. Deep-bodied, pale grey, a black pupil-width line from
front of first dorsal-fin over abdomen, continuing as a black margin on
A ventral fin. Black spots, small on head and near pupil-size posteriorly
Sphaeramia orbicularis Flores, Indonesia. RHK on body. Length to about 75 mm.
B C
Sphaeramia orbicularis Derawan, Borneo. RHK Sphaeramia orbicularis Pulau Putri, Java, Indonesia. RHK

Pyjama Cardinalfish Sphaeramia nematoptera

Apogon nematopterus Bleeker, 1856. Ambon, Indonesia.
Western Pacific: Indonesia east to Caroline Islands and Fiji, north to
Yaeyama Islands, south to northern Queensland (Australia) and New
Caledonia. Sheltered coastal reef slopes and lagoon with rich coral
growth. Forms large schools in branching corals during the day. In
depths to about 10 m. Deep bodied with tall fins, second dorsal-fin
with whip-like filament reaching anal-fin tip in adults. Head yellow,
eye bright red with two vertical white lines on sides. A broad black
band over first dorsal-fin, extended over body below. Rest of body
white with many half pupil-size red spots. Length to 85 mm.
A
Sphaeramia nematoptera Flores, Indonesia. RHK
C
B Sphaeramia nematoptera Iriomote Island, Japan. KYA
D
Sphaeramia nematoptera Schooling in among branching corals. Flores, Indonesia. RHK

SUBFAMILY APOGONINAE – TRIBE ZORAMIINI

Contains two genera: Fibramia and Zoramia.
Diagnosis D VI–I,9; A II,8–9; head and body with ctenoid scales; pored lateral-line scales 24;
preopercle ridge smooth, edges serrate; three supraneurals; supramaxilla absent; basisphenoid present;
anterior pair of uroneurals; three epurals; five free hypurals; parhypural separate; two autogenous
haemal spines; one supernumerary dorsal spine; caudal fin forked. Other characteristics first segmented
fin-ray in second dorsal-fin branched, first anal-fin ray branched and segmented; second dorsal spine
longest, very long and filamentous in some species of both genera; ctenoid scales on predorsal, cheek,
breast, two pelvic scales, and body; ctenoid scales on opercle and onto base of caudal fin; cycloid or
ctenoid predorsal scales1–3; pored lateral-line scales simple with one pore above and one below
midline; pectoral-fin rays 13–17; 9+8 segmented principal caudal rays, 15 branched, upper and lower
unbranched; developed gill rakers 7–38; unbranched procurrent rays, longest segmented; 10+14
vertebrae; 8 ribs; 7–8 epipleurals; teeth in one, two or multiple rows on premaxilla, dentary, vomer,
palatine, all villiform; six infraorbitals, bony shelf absent or on third infraorbital; stomach and intestine
black with silvery peritoneum; low crest on PU2 (after Mabuchi et al, 2014).
TRIBE ZORAMIINI – Genus Fibramia Fraser & Mabuchi 2014

Fem. Type-species Fibramia thermalis Cuvier, 1829.
Genus Fibramia comprises three described species: F. amboinensis, F. lateralis and F. thermalis. Coastal
marine fishes, some may enter freshwater.
Sangi Cardinalfish Fibramia thermalis

Apogon thermalis Cuvier, 1829. Ceylon.
Indo-West Pacific: East Africa and Madagascar to Australia and
Vanuatu, north to southern Japan. Coastal, protected clear waters in
small aggregation with outcrops on reefs and sand to about 20 m depth.
Semi-transparent to dusky above, a dark stripe over snout through eye,
fading on to body. Anterior margin of dorsal fin black. A small black spot
following bases of both dorsal fins and a basicaudal spot about pupil-size.
For a long time this taxon was known as Apogon sangiensis Bleeker,
1857, which is a putative junior synonym.
A
Fibramia thermalis Bali, Indonesia. RHK
B C
Fibramia thermalis Derawan, Borneo. RHK Fibramia thermalis Pulau Putri, Java, Indonesia. RHK
D E
Fibramia thermalis Iriomote Island, Japan. TKO Fibramia thermalis Iriomote Island, Japan. KYA

Coastal Cardinalfish Fibramia amboinensis

Apogon amboinensis Bleeker, 1853. Ceylon.
Western Pacific: Japan to Indonesia and Papua New Guinea. Coastal, brackish waters, entering freshwater. Semi-transparent, whitish
and silvery reflective. A black downward curving thin line from top of operculum to caudal-peduncle, followed by a small basicaudal spot.
Similar to Fibramia lateralis, the next species, but head profile concave versus convex above eyes. Length to about 11 cm.
A B
Fibramia amboinensis Northern Borneo, Malaysia. GRA Fibramia amboinensis Flores, Indonesia. RHK
Pin-stripe Cardinalfish Fibramia lateralis

Apogon lateralis Valenciennes, 1832.
Vanikoro Island, Santa Cruz Islands, Western Pacific.
Indo-West Pacific: East Africa, Seychelles, Madagascar and Mauritius
east to Caroline Islands (Micronesia), Tonga and Samoa, north to
Japan, south to Australia and New Caledonia. Coastal, protected clear
waters near rivers and may enter freshwater. Semi-transparent to dusky
above, a dark stripe over snout through eye, fading on to body. A thin
black downward curving line from top of operculum to caudal-peduncle,
followed by a small basicaudal spot. A small black spot following eye often
present. First dorsal-fin with leading black margin. Length to 85 mm.
B C
D E
Fibramia lateralis Maumere, Flores, Indonesia. RHK

TRIBE ZORAMIINI – Genus Zoramia Jordan, 1917

Fem. Type-species Apogon graeffii Günther, 1873. {=Zoramia leptacantha (Bleeker, 1856)}.
Genus Zoramia comprises six described species: Z. flebila, Z. fragilis, Z. gilberti, Z. leptacantha. Z. perlita, and
Z. viridiventer. Apparently there are more species, yet to be described.
Long-spine Cardinalfish Zoramia leptacantha

Apogon leptacanthus Bleeker, 1856. Molucca Islands, Indonesia.
Red Sea, Indo-West Pacific: East Africa, Madagascar and Seychelles
east to Marshall Islands, Tonga and Samoa, south to Australia and
New Caledonia. Clear coastal reefs to outer reef lagoons, protected
waters in small to large aggregation with staghorn-coral outcrops on reefs
and sand to about 15 m depth. Disperses at night over sandy and rubble
substrates to feed on small swimming crustaceans. Semi-transparent pale
greenish to bluish on head and abdomen with yellow and blue narrow bars
on deepest part of the body. Length to about 8 cm.
A
Zoramia leptacantha Flores, Indonesia. RHK
B C
Zoramia leptacantha Iriomote Island, Japan. KYA Zoramia leptacantha Ashmore Reef, WA. BHU
D
Zoramia leptacantha Maumere, Flores, Indonesia. RHK

Pearly Cardinalfish, Zoramia perlita

Apogon perlitus Fraser & Lachner, 1985. Palau.
Indo-West Pacific: Andaman Sea to Palau and Japan.
Coastal, often silty reefs in sheltered bays and estuaries,
forming schools closely above branching corals. Slender
for genus, semi-transparent grey. A small white saddle spot
immediately behind second dorsal-fin base, a dusky streak
on the anal-fin base and a basicaudal black small spot. No
black caudal-fin tips. Length to 50 mm.
A
Zoramia perlita Gilimanuk, Bali, Indonesia. RHK
B C
Zoramia perlita Pulau Putri, Java, Indonesia. RHK Zoramia perlita Iriomote Island, Japan. TKO
Gilbert’s Cardinalfish, Zoramia gilberti

Amia gilberti Jordan & Seale, 1905. Philippines.
Malaysia, Indonesia, Papua New Guinea to the Philippines,
east to Palau and Yap. Sheltered reef lagoons in rich coral
crests, in pairs or small aggregations. Semi-transparent,
abdominal region reflective silvery blue. Dorsally and upper
sides sometimes brownish grey and densily peppered with
yellow miniscule-sized spots from snout to a dusky caudal
peduncle. Ventrals reddish. A prominent dark spot usually
present on opercle flap, and has a small black basicaudal
spot in pale fish, which maybe obscured when peduncle is
dusky. Caudal-fin tips black. Length to 50 mm.
A
Zoramia gilberti Ashmore Reef, WA. BHU
B
Zoramia gilberti Palau, Micronesia. HNA

Sparsely-spotted Cardinalfish Zoramia viridiventer

Apogon viridiventer Greenfield, Langston & Randall, 2005.
Madang Province, Papua New Guinea.
Southern Japan, Malaysia, Indonesia, Australia to the Marshall
Islands. In small aggregation in branching corals on sheltered reefs to
about 15 m depth. Semi-transparent, reflective white or silvery blue on
belly and gill-cover. Few blue spots scattered on head and anteriorly on
body. A basicaudal black spot up to about pupil-size and caudal fin with
black tips. Eyes yellow. Length to about 6 cm.
A
Zoramia viridiventer Madang, PNG. GRA
B C
Zoramia viridiventer Iriomote Island, Japan. KYA Zoramia viridiventer Flores, Indonesia. RHK
Tearful Cardinalfish Zoramia flebila

Apogon flebila Greenfield, Langston & Randall, 2005.
Vanua Balavul Island, Northern Lau Group, Fiji.
West Pacific, Papua New Guinea to Fiji. Sheltered reefs in coral at about
10–15 m depth. Semi-transparent greenish brown dorsally and upper
sides. reflective white or silvery blue on belly and gill-cover. Anal-fin base
with dusky and pale blue line. Few blue spots scattered on head below eye
and in series anteriorly on body. A small black basicaudal spot up to about
pupil-size, but may be obscured by a dusky caudal-peduncle. Eyes yellow
dorsally, blue ventrally. Caudal fin with black tips. Length to 55 mm.
Zoramia flebila Madang, PNG. GRA
Fragile Cardinalfish Zoramia fragilis

Apogon fragilis Smith, 1961. Pinda, Mozambique, western Indian Ocean.
West Indian Ocean from African coast to the Maldives. Sheltered lagoons in corals, moving out over sand at night. Bluish transparent,
dusky dorsally from interorbital to caudal-fin and ventrally along caudal peduncle. A small basicaudal spot, first-dorsal and caudal fin with
small black tips. Length to 50 mm.
Zoramia fragilis Night. Maldives. RHK

Blue-barred Cardinalfish Zoramia sp. 1

Known only from Maumere Bay Flores, Indonesia, Coastal,
usually very shallow near freshwater outlets in lagoons.
Schooling among mangrove roots or staghorn corals on
inner reefs at hight tides. Semi-transparent, reflective white
or silvery blue on belly and gill-cover. Five blue narrow bars on
deepest section of the body. A basicaudal black spot about
pupil-size. Caudal fin with black tips. Ventral fins reddish
with blue tips. First dorsal-fin blue in large individuals. Eyes
yellow dorsally and blue ventrally. Length to about 50 mm.
B C
D
Zoramia sp. 1. Maumere, Flores, Indonesia. RHK

SUBFAMILY PAXTONINAE Fraser & Mabuchi, 2014

Type-genus Paxton Baldwin & Johnson, 1999. Represented by a single genus.
Diagnosis D. (continues), VI,19; A. I,15–16; internal support of spines by serial proximal-middle
radials in near articulation with gaps between each spine, similar distance between 6th and 7th
elements; sixth proximal-middle radial without serial spine or ray; fifth and seventh proximal-middle
radials with serial spine and ray respectively; one supernumerary dorsal spine; supraneurals
absent; first anal proximal-middle radial curved; 9+8 branched principal caudal fin-rays; caudal fin
truncate or slightly rounded; vertebrae 10+14; epineurals on first two vertebrae; rod-like ribs on
3rd to 10th vertebrae; epineurals present on ribs of 3rd to 9th vertebrae; supramaxilla and
basisphenoid absent; six infraorbitals, without shelf on third, only first and second in contact, third
sixth not in contact and all small; medial and lateral extrascapular absent; preopercle ridge smooth
and edge with single large spine at angle, preopercle, including spine, covered by skin; prootic
excluded along internal orbit ring by pterosphenoids and parasphenoid; parietal separated by
supraoccipital; a unique postfrontal bone; uroneurals absent; two epurals; haemal spines for PU3
and PU4 each fused to centra; parhypural fused to hypurals 1+2; terminal centrum fused with
hypurals 3+4; hypural 5 absent; second epibranchial articulating with third pharyngobranchial;
anterior and posterior ceratohyals sutured together by a few interdigitating struts medially;
anterior ceratohyal not per forated; seven branchiostegals, anterior three ventrally followed
by two on distal side of anterior ceratohyal, two on distal side of posterior ceratohyal; single
postcleithrum (after Mabuchi et al, 2014).
Genus Paxton Baldwin & Johnson, 1999

Fem. Type-species Paxton concilians Baldwin & Johnson, 1999.
Genus Paxton is monotypic. Four specimens of this percoid fish were collected by trawl off northwestern
Australia and prior to determining it to be an apogonid were believed to represent an undescribed genus of
grammistin serranid. As is the case with some of the closely related Pseudamia, juveniles may secretively live
in shallow habitats of the Monte Bellow Islands or Kimberley coast, regions rarily visited by divers.
Paxton's Cardinalfish Paxton concilians

Paxton concilians Baldwin & Johnson, 1999. Northeast off the Monte Bello Islands, Western Australia.
Known only from Western Australia. Occurring on the continental shelf to about 80 m depth, types came from about 65 m depth.
Juveniles may occur in shallow water, expected to live secretively in rocky crevices. Paxton concilians is unique among the Apogonidae
taxa in having a single continuous dorsal fin, VI,19; anal fin I,15–16. Also the number of soft rays is unusually high in both fins.
Generically it shares many characters with Pseudamia. Body mostly pale creamish to yellow, gill cover and abdominal region
silvery white. Median fins whitish, dorsal with a distinct black outer margin. Length to 85 mm.
Paxton concilians Freshly preserved holotype 63.6 mm SL. Near Monte Bello Islands, Western Australia. WWH

SUBFAMILY PSEUDAMIINAE Smith, 1954

Type genus Pseudamia Bleeker, 1865. Represented by a single genus.
Diagnosis D. VII, as VI+I,8–9; A. II,8–10; wide separation between dorsal fins; internal support
of dorsal spines by serial proximal-middle radials not in close articulation, far apart at 6th between
and 7th elements; one supernumerary dorsal spine; 1–2 supraneurals; first anal proximal-middle
radial straight; 9+8 principal caudal fin-rays, 15 branched, upper and lower unbranched, caudal
fin rounded or rhomboid, mid-line fin-ray longest; supramaxilla and basisphenoid absent;
preopercle ridge smooth, edge serrated at angle or smooth; prootic narrowed along internal orbit
rim, not quit excluded by pterosphenoid and parasphenoid; anterior pair of uroneurals absent,
posterior pair of uroneurals reduced or absent; two epurals; hypurals 1–2 fused, 3–4 fused,
separate from urostyle; hypural 5 free; parhypural free; free medial extrascapular; neuromasts
reduced to a simple cross-hatch on head, linear and vertical rows on body, present on caudal fin;
cephalic pore system simple without canal flutes; lateral-line scales 23–43, first few scales pored,
followed by notched scales, a lower lateral line of notched scales begins on abdomen; cycloid
scales on body (after Mabuchi et al, 2014).
Genus Pseudamia Bleeker, 1865

Fem. Type-species Cheilodipterus polystigma Bleeker, 1860 (= Pseudamia amblyuroptera).
Genus Pseudamia comprises seven species: P. amblyuroptera, P. gelatinosa, P. hayashii, P. nigra, P. rubra,
P. tarri, and P. zonata.
White-jaw Cardinalfish Pseudamia amblyuroptera

Cheilodipterus amblyuropterus Bleeker, 1856. Buru Island, Molucca Islands, Indonesia.
Western Pacific: Singapore, Indonesia, Papua New Guinea to the Philippines, east to Palau and Yap islands. Shallow estuaries near
freshwater run-offs, usually found in depths less than 2 m. Body with black spots and streaks, a white area below eye with one or
two small black spots. Fins blackish and an indistinct pupil-size basicaudal spot above centre. Length to 85 mm.
Pseudamia amblyuroptera Manado, Indonesia. RHK
Hayashi’s Cardinalfish Pseudamia hayashii

Apogon hayashii Randall, Lachner & Fraser, 1985.
Ngargol Island, Palau Islands.
Red Sea, Indo-West Pacific: Gulf of Aden; Indonesia, Japan,
Australia, east to Micronesia and Samoa. Sheltered reefs in
coral, shallow to 65 m depth. Semi-transparent black when
juvenile; brown to black when adult. Caudal fin black, large
and rounded with transparent corners. Length to 10 cm.
A
Pseudamia hayashii Tulamben, Bali, Indonesia. RHK
B C
Pseudamia hayashii Maldives. RHK Pseudamia hayashii Socrota. SBO

Charcoal-banded Cardinalfish Pseudamia zonata

Pseudamia zonata Randall, Lachner & Fraser, 1985. Mactan Island, Cebu, Philippines.
Western Pacific: Indonesia to Fiji, north to Japan. Solitary, far in the back of small caves during the day, hovering in front at night in
depths of 10 to 30 m. Slender-bodied species, whitish grey with large fins that are mostly black in adults, black broad bands below
dorsal fins and one basicaudal as an elongate blotch. A large species in the genus, length to about 21 cm.
A B
Pseudamia zonata Milne Bay, PNG. PWO Pseudamia zonata Iriomote Island, Japan. KYA
C
Pseudamia zonata Juvenile. Tulamben, Bali, Indonesia. RHK
D
Pseudamia zonata Rabaul, PNG. RCS

Gelatinous Cardinalfish
Pseudamia gelatinosa
Cheilodipterus gelatinosa Smith, 1956.
Aldabra Islands, western Indian Ocean.
Red Sea, Indo-West Pacific: East Africa and Madagascar
east to Society Islands, Japan, and Australia. Shallow to 40
m depth. Body with small black square spots that have a
pale centre and tiny spots forming numerous fine lines
along the body. Median fins with black bands and whitish
corners. Length to 11 cm.
A
Pseudamia gelatinosa Tulamben, Bali, Indonesia. RHK
B C
Pseudamia gelatinosa Kerama Island, Japan. AON Pseudamia gelatinosa Lizard Island, Qld, Australia. RHK
D
Pseudamia gelatinosa Yemen, Red Sea. PCH
Tarr’s Cardinalfish Pseudamia tarri

Apogon tarri Randall, Lachner & Fraser, 1985.
Jana Island, Saudi Arabia, Persian Gulf.
Western Indian Ocean, known only from the Persian Gulf.
Secretive on sheltered reefs in coral rubble, shallow to about
15 m depth. Pale pinkish grey with numerous black tiny
spots. Fins mostly dusky, pale marginally along bases.
Caudal fin dusky and strongly rounded. Length to 50 mm.
Pseudamia tarri Jana Island, Saudi Arabia, Persian Gulf. JER
Ogasawara Cardinalfish Pseudamia rubra

Apogon rubra Randall, Lachner & Fraser, 1985.
Chichi-jima, Ogasawara Islands, Japan.
Western North Pacific. Sheltered reefs in coral-caves or
among rubble in a depth of few metres. Pale greyish, scales
with reddish margins. A dusky spot following eye and a
pupil-size dusky spot posteriorly on caudal peduncle.
Pseudamia rubra Holotype. Chichi-jima, Japan. JER

Acknowledgements
We are most grateful to Gerry Allen, Sergey Bogorodsky, Thomas Fraser, Martin Gomon, Ross Robertson for checking/reviewing
various sections of the manuscript, sharing information and their comments. A special thank you to the numerous photographers,
artists and institutions listed below for their generous contributions, making this worldwide project possible.
List of photographers initials

ACF – Alfredo Carvalho Filho, Brazil MAT – Moriyama Atsushi. Japan
AMI – Akihiko Mishiko, Japan. Via FishPix MEH – Mohammad Eusuf Hasan, Pakistan. Via Fishbase
ANA – Atsushi Nagano, Japan. Via FishPix MEN – Masako Enomoto, Japan
AON – Atsushi Ono, Japan MHA – Masayoshi Hayashi, Japan, Yokosuka City Museum
ARY – Andrey Ryanskiy MPF – Malcolm Francis, NIWA, New Zealand
BHU – Barry Hutchins, WA Museum MSC – Mike Scotland, www.mikescotland.com.au
BNA – Benjamin Naden MVE – Mark V. Erdmann, Conservation International
CAB – Cindy Abgarian, cgillsphotos.com, USA MYA – Michihiko Yamada, Japan. Via FishPix
CBA – Christhoph Bausch, Germany NBA – Nicolas Bailly, USA
CSIRO – Commonwealth Scientific and Industrial Research NCO – Neville Coleman, Australia
Organisation NDL – Ned De Loach, USA
DBR – Dianne Bray, Museums Victoria NLI – Nathan Litjens, Australia
DBU – David Burdick, Guam NTS – Nobuaki Tsutaki, https://tsutaki.net/
DGO – Dawn Goebbels, South Africa NVR – Nuno Rodrigues Vasco, www.nunovascorodrigues.com
DHA – David Harasti, Australia PCH – Phil C. Heemstra, SAIAB, South Africa
DKI – Dennis King, South Africa PHU – Paul Human, USA
DPO – Dennis Polack, FishwisePro, South Africa PWI – Peter Wirtz, https://www.researchgate.net/
EHE – Elaine Heemstra, SAIAB, South Africa profile/Peter_Wirtz
ESC – Erik Schlogl, Australia PWO – Phil Woodhead, Wet Image, Cairns
FLI – Francois Libert, France R&B – Robertson & Baldwin, STRI
FSN – Frank Schneidewind, Germany RCH – Raymond Chan, Hong Kong
FTE – Frank Teigler, Germany, www.luxfotodesign.de RFI – Richard Field, Oman
GED – Graham Edgar, University of Tasmania RHK – Rudie H. Kuiter, Aquatic Photographics, Australia
GPW – Gilbert P. Whitley. Australian Museum RLU – Roger Lubbock, England
GLM – Glen Moore, Western Australian Museum RMI – Ryo Minemizu, Japan, Phenomenons.
GRA – Gerry R. Allen, Conservation International RMY – Rob Myers, Coral Graphics
GTA – Go Tabinuki, Japan RPY – Richard Pyle, Hawaii
GJW – Glen J. Whisson, Aqua Research and Monitoring RRO – Ross Robertson, Smithsonian Tropical Research
Services, Western Australia Institute, Panamá
HBO – Hamid Badar Osmany, Pakistan. Via Fishbase RWI – Richard Winterbottom, Canada
HDE – Helmut Debelius, IKAN, Germany RYU – Rowena Yue, Hong Kong
HDO – Hideo Dohra, Japan. Via FishPix SBO – Sergey Bogorodsky, Station of Naturalists, Omsk, Russia
HER – Harry Erhardt, Germany SDE – Shinichi Dewa, Japan
HFR – Herbert Frei, Germany SKA – Shoichi Kato, Japan
HKA – Hiroaki Kaneko, Japan. Via FishPix SLK – Steven L. Kuiter, Wildlife Photographics, Australia
HMO – Hiroyuki Motomura, Kagoshima University Museum, SMI – Smithsonian Library, USA
Japan SNA – Seishi Nakano, Japan. Via FishPix
HSE – Hiroshi Senou, Kanagawa Prefectural Museum SPO – Sally Polack, FishwisePro, South Africa
of Natural History. Japan SRA – Sandra Raredon, USA
ISH – Ian Shaw, Australia STRI – Smithsonian Tropical Research Institute, Panamá
JER – John E. Randall, Bishop Museum, Hawaii SWM – Scott W. Michael, USA
JGA – Jim Garin, USA SWO – Stephen Wong, Hong Kong
JGR – Jim Greenfield, www.oceaneyephoto.com THI – Tomonori Hirata, Japan
JHI – Johan Hinterkircher, Germany TKO – Toshikazu Kozawa, Anthis (Nexus)
JHO – John Hoover, Hawaii TPA – Thomas Paulus, Germany
JLG – João Luiz Gasparini, Brazil TSH – Takumi Shiraishi, Aquarius Divers
JNE – John Neuschwander, Netherlands TSU – T. Susuki, Japan
JTW – Jeff T. Williams, Div. of Fishes, Museum Support Center TTA – Taizo Taguchi, Japan. Via FishPix
KIM – Keisuke Imai. Japan TTO – Takamasa Tonozuka, Dive & Dive, Bali
KNI – Kenji Nin. Japan. Via FishPix USA – Ukkrit Satapoomin, Phukett
KKO – Keita Koeda. National Museum of Marine Biology & VTR – Van Tassell & Robertson, STRI
Aquarium, Taiwan WCC – Wei-Chusn Chiang. Via Fishbase
KTA – Kotaro Tanaka, Japan WWH – William White, CSIRO
KUC – Keido Uchino, Japan. Via FishPix ZUK – Via zukan.com
KYA – Korechika Yano, Japan ZYU – Zhengsen Yu. Via Fishbase
LJO – Louis Johnston, STRI

REFERENCES & LITERATURE CITED

Allen, G.R. 1985. A new species of cardinalfish (Apogonidae) from Northern Australia and the Aru Islands.
=description of Apogon fusovatus.
Allen, G.R. 1987. New Australian Fishes, Part 2, Four new species of Apogonidae.
=descriptions of Apogon pallidofasciatus, Rhabdamia eremia, Vincentia badia & V. macrocauda
Allen, G.R. 1992. A new species of cardinalfish (Apogonidae) from Northern Australia. =description of Pseudamia nigra.
Allen, G.R. 1993. Cardinalfishes (Apogonidae) of Madang Province, Papua New Guinea, with
Descriptions of Three New Species. =descriptions of Fowleria flammea, Siphamia corallicola & S. jebbi.
Allen, G.R. 1995. A new species of cardinalfish (Apogon: Apogonidae) from Northern Australia.
=description of Apogon unitaeniatus.
Allen, G.R. 1998. A new species of cardinalfish (Apogonidae) from the Komodo Islands, Indonesia.
=description of Apogon komodoensis.
Allen, G.R. & Erdmann, M.V. 2012 Reef fishes of the East Indies. Apogonidae 366–418, Volumes I–III.
Tropical Reef Research, Perth, Australia, i–xiv + 1292 pp
Allen, G.R., & R.H. Kuiter. 1994. Descriptions of two new species of cardinalfishes (Apogonidae) from Malaysia and Indonesia.
=descriptions of Apogon sabahensis & Rhabdamia spilota.
Allen, G.R., & R.H. Kuiter, J.E. Randall. 1994. Descriptions of five new species of cardinalfishes (Apogonidae: Apogon)
from Maumere Bay, Flores, Indonesia and surrounding regions.
=descriptions of Apogon ocellicaudus, A. nanus, A. neotus, A. franssedai & A. ventrifasciatus.
Allen, G.R., & S.M. Morrison. 1996. A new species of Cardinalfish (Apogonidae) from northern Australia.
=description of Apogon fuscomaculatus.
Allen, G.R., & J.E. Randall. 1993. Three new species of Cardinalfishes (Apogonidae) from Australia and Adjacent Seas.
=descriptions of Apogon virgulatus, A. flavus & A. capricornis.
Allen, G.R., & J.E. Randall. 1993. A new species of cardinalfish (Apogon: Apogonidae) from Arabian Seas.
=descriptions of Apogon smithvanizi.
Allen, G.R., & J.E. Randall. 1995. Apogon virgulatus Allen et Randall, a junior synonym of
Apogon cavitiensis Jordan et Seale. (cavitensis is correct spelling)
Allen, G.R., & R.C. Steene. 1995. Notes on the ecology and behaviour of the Indonesian
cardinalfish (Apogonidae) Pterapogon kauderni Koumans.
Baldwin, C.C. & Johnson, G.D. 1999. Paxton concilians: a new genus and species of Pseudamine apogonid
(Teleostei: Percoidei) from northwestern Australia: the sister group of the enigmatic Gymnapogon. Copeia
1999 (4), 1050–1073.
Bleeker, P. 1852-1879. Various publications, Nat. Tijds. Ned. Ind., Verh. Akad., Amsterdam.
Bleeker, P. 1873. Atlas Ichthyology. Volume 7.
Bleeker, P. 1876. Atlas Ichthyology. Volume 8.
Castelnau, F.L. 1878. Australian fishes. New or little known species. Proceedings of the Linnean Society of New South Wales,
2 (3), 225–248.
Day, F. 1875. The fishes of India; being a natural history of the fishes known to inhabit the seas and Fresh waters of India,
Burma, and Ceylon. London. 1, 1–168.
Fraser, T.H. 1972. Comparative osteology of the shallow water cardinal fishes (Perciformes: Apogonidae) with references to the
systematics and evolution of the family. Ichthyological Bulletin of the J. L. B. Smith Institute of Ichthyology, 34, i–v +1–105.
Fraser, T.H. 1973. Evolutionary significance of Holapogon, a new genus of cardinal fishes (Apogonidae), with a redescription of its
type-species, Apogon maximus. The J.L.B Smith Institute of Ichthyology Special Publication, 10, 1–7
Fraser, T.H. 1998. A new species of cardinalfish (Apogonidae) from the Philippines with
comments on species of Apogon with six first dorsal spines. =description of Apogon bryx.
Fraser, T.H. 2005. A review of the species in the Apogon fasciatus group with a description of a new species of cardinalfish
from the Indo-West Pacific (Perciformes: Apogonidae). Zootaxa, 924, 1–30.
Fraser, T.H. 2008. Cardinalfishes of the genus Nectamia (Apogonidae, Perciformes) from the Indo-Pacific region with descriptions
of four new species. Zootaxa, 1691, 1–52.
Fraser, T.H. 2013a. Family status of and the senior synonym of Coranthus (Percomorpha: Apogonidae). Copeia, 2013, 23–30.
Fraser, T.H. 2013b. A new genus of cardinalfish (Apogonidae: Percomorpha), redescription of Archamia and resemblances and
relationships with Kurtus (Kurtidae: Percomorpha). Zootaxa, 3714 (1), 1–63.
Fraser, T.H. & Allen, G.R. 2010. Cardinalfishes of the genus Apogonichthyoides Smith, 1949 (Apogonidae) with a
description of a new species from the West-Pacific region. Zootaxa, 2348, 40–56.
Fraser, T.H. & E.A. Lachner. 1985. A Revision of the Cardinalfish Subgenera Pristiapogon
and Zoramia (Genus Apogon) of the Indo-Pacific Region.
Fraser, T.H. & J.E. Randall. 1976. Two new Indo-West Pacific Cardinalfishes of the genus Apogon.
=descriptions of Apogon dispar & melanoproctus.
Gilbert, Carter R. & James C. Tyler. 1997. Apogon robbyi, a new cardinalfish (Perciformes: Apogonidae) from the
Caribbean Sea. Bulletin of Marine Scence. 60(3): 764–781.
Grant, E.M. 1987. Fishes of Australia.
Gomon, M.F. J.C.M. Glover & R.H. Kuiter. 1994. Fishes of Australia’s South Coast.
Gon, O. 1994. Revision of the Cardinalfish Genus Cheilodipterus (Perciformes: Apogonidae),
with Description of Five New Species.
Gon, O. 1995. Revision of the cardinalfish subgenus Lepidamia (Perciformes, Apogonidae, Apogon). Israel Journal of Zoology,
41, 1–22.
Gon, O. 1996. Revision of the cardinalfish subgenus Jaydia (Perciformes, Apogonidae, Apogon). Transactions of the Royal
Society of South Africa, 51, 147–194.
Gon, O. & Allen, G.R. 2012. Revision of the Indo-Pacific cardinalfish genus Siphamia (Perciformes: Apogonidae). Zootaxa,
3294, 1–84.
Gon, O., Liao, Y.-C. & Shao, K-T, 2015. A new species of the cardinalfish genus Jaydia (Teleostei: Apogonidae) from the
Philippines. Zootaxa, 3980 (2), 286–292.
Gon, O. & Randall, J.E. 2003. Revision of the Indo-Pacific cardinalfish genus Archamia (Perciformes: Apogonidae, with

descriptions of a new species. Indo-Pacific Fishes, 35, 1–49.

Greenfield, D. W. 2001. Revision of the Apogon erythrinus complex (Teleostei: Apogonidae). Copeia 2001 (no. 2): 459-472.
Greenfield, D.W., Langston, R.C. & Randall, J.E. 2005. Two new cardinalfish species of the Indo-Pacific fish genus Zoramia.
Proceedings of the California Academy of Science s, Fourth Series, 56, 625–637
Hayashi, M. 1990. Two new cardinalfish (Apogonidae: genus Apogon) from the Indo-West
Pacific. =includes descriptions of Apogon fukuii & A. sheni.
Hayashi, M., et al. 1997. First record of apogonid fish, Apogon franssedai (Pisces:
Apogonidae) from Ryukyu Islands, southwestern Japan.
Hoover, John P. 1994. Hawaii’s Fishes. Mutual Publishing. Honolulu, Hawaii.
Humann, Paul & Ned DeLoach. 2002. Reef fish identificationm Florida Caribbean Bahamas.
General, Apogonidae: photographs: 17 species.
Ida, H., & J.T. Moyer. 1974. Apogonid Fishes of Miyake-Jima and Ishigaki-Jima, Japan, with
descriptions of a New Species. =including description of Apogon ishigakiensis.
Jordan, D.S., & J.O. Snyder. 1901. List of fishes collected in 1883 and 1885 by Pierre Louis
Jouy and preserved in the United States National Museum, with descriptions of six new species.
Kawanabe, H., & N. Mizuno. Freshwater Fishes of Japan.
Includes brackisch waters. Apogonidae: photographs: Apogon amboinensis, Sphaeramia orbicularis.
Kamohara, T. 1967. Coloured Illustrations of The Fishes of Japan. General. Apogonidae: drawings:
Apogonichthys carinatus, Apogon lineatus, A. doederleini, A. semilineatus, A. notatus
Kottelat, M., et al. 1993. Freshwater Fishes of Western Indonesia and Sulawesi. Includes brackisch waters.
Apogonidae: photographs: Apogon amboinensis, A. hyalosoma, A. sangiensis, A. lateralis.
Kuiter, R.H. 1992. Tropical Reef-Fishes of the Western Pacific, Indonesia and Adjacent Waters.
Kuiter, R.H. 1993. Coastal Fishes of South-Eastern Australia. General, Apogonidae: photographs: 25 species.
Kuiter, R.H. 1994. Southeast Asia Tropical Fish Guide. General, Apogonidae: photographs: 21 species.
Kuiter, R.H. 1997. Guide to Sea Fishes of Australia. General, Apogonidae: photographs: 42 species.
Kuiter, R.H. 1998. Photo Guide to Fishes of the Maldives. General, Apogonidae: photographs: 34 species.
Kuiter, R.H. & S.L. Kuiter 2018. Coastal Sea-Fishes of South-Eastern Australia. Aquatic Photographis, Seaford.
Lachner, E.A. 1953. Family Apogonidae: cardinalfishes. In Schultz et al., Fishes of the Marshall and Marianas Islands.
Leis, J.M., Meyer, O., Hay, A.C. & Gaither, M.R. 2015. A coral-reef fish with large, fast conspicuous larvae and small,
cryptic adults. Copeia, 103 (1), 78–86.
Mabuchi, K., Fraser, T.H., Song, H., Azuma, Y., & Nishida, M. 2014. Revision of the systematics of the cardinalfishes
(Percomorpha: Apogonidae) based on molecular analyses and comparative reevaluation of morphological characters.
Zootaxa, 3846 (2), 151–203
Masuda, H., & G.R. Allen. 1987. Sea Fishes of the World. General, Apogonidae: photographs: 55 species.
Masuda, H., et al. 1984. The Fishes of the Japanese Archipelago. General, Apogonidae: photographs: 70 species.
Masuda, H., & Y. Kobayashi. 1994. Grant Atlas of Fish Life Modes. General, Apogonidae: photographs: 60 species.
McCulloch, I.S.R. 1929-1930. A checklist of the fishes recorded from Australia.
Mees, G.F. 1966. A new fish of the family Apogonidae from tropical Western Australia. Journal of the Royal Society of
Western Australia, 49 (3), 83–84.
Munro, A.R. 1960. Handbook of Australian Fishes.
Myers, R.F. 1989. Micronesian Reef Fishes.
Nakabo, T. 1993. Fishes of Japan with Pictorial Keys to the Species.
Okamura, O., & K. Amaoka. 1997. Sea Fishes of Japan. General, Apogonidae: photographs: 67 species.
Paxton, J.R., et al. 1989. Zoological Catalogue of Australia. Vol. 7. PISCES.
Radcliffe, L. 1912. Descriptions of fifteen new fishes of the family Cheilodipteridae,
from the Philippine Islands and contiguous waters.
Randall, J.E. 1983. Red Sea Reef Fishes. General, Apogonidae: photographs: 14 species.
Randall, J.E. 1995. Coastal Fishes of Oman. General, Apogonidae: photographs: 34 species.
Randall, J.E., G.R. Allen, & R.C. Steene. 1990. Fishes of the Great Barrier Reef and Coral Sea.
Randall, J.E., & R.C. Anderson. 1993. Annotated Checklist of the Epipelagic and Shore Fishes of the Maldive Islands.
Randall, J.E., & J.E. Bohlke. 1981. The Status of the Cardinalfishes Apogon evermanni and A. anisolepis
(Perciformes: Apogonidae), with Description of Related New Species from the Red Sea.
Randall, J.E., & M. Hayashi. 1990. Apogon selas, a New Cardinalfish from the Western Pacific.
Randall, J.E., & D.F. Hoese. 1988. Apogon limenus, a new species of cardinalfish
(Perciformes: Apogonidae) from New South Wales.
Randall, J.E., & H. Ida. 1993. Pseudamia rubra, a New Cardinalfish (Perciformes: Apogonidae) from the Ogasawara Islands.
Randall, J.E., & M. Kulbicki. 1998. Two new Cardinalfishes (Perciformes: Apogonidae) of the
Apogon cyanosoma complex from the western Pacific, with notes on the status
of A. wassinki Bleeker. =including descriptions of Apogon luteus & A. rubrimacula.
Randall, J.E., & E.A. Lachner. 1986. The status of the Indo-West Pacific cardinalfishes.
Apogon aroubiensis and A. nigrofasciatus.
Randall, J.E., E.A. Lachner, & T.H. Fraser. 1985. A Revision of the Indo-Pacific Apogonid
Fish Genus Pseudamia, with Descriptions of Three New Species.
Randall, J.E., E.A. Lachner, & T.H. Fraser. 1990. On the validity of the Indo-Pacific cardinalfishes Apogon aureus
Lacepede),
and A. fleurieu (Lacepede), with description of a related new species from the Red Sea.
Randall, J.E., & C. Lavett Smith. 1988. Two New Species and a New Genus of Cardinalfishes
(Perciformes: Apogonidae) from Rapa, South Pacific Ocean. =descriptions of Apogon caudicinctus & Cercamia
cladara.
Randall, J.E., Lachner, E.A. & Fraser, T.H. 1985. A revision of the Indo-Pacific apogonid fish genus Pseudamia, with
descriptions of three new species. Indo-Pacific Fishes, 6, 1–23.
Sainsbury, K.J. et al. 1984. Continental Shelf Fishes of Northern and North-Western Australia.
Smith, J.L.B. 1961. Fishes of the family Apogonidae of the western Indian Ocean and the Red Sea. Ichthyological Bulletin,
Department of Ichthyology, Rhodes University, 22, 373–418.
Weber, M. & deBeaufort, F.L. 1929. The fishes of the Indo-Australian Archipelago. V. Anacanthini, Allotriognathi,
Heterostomata, Berycomorphi, Percomorphi, families Kuhliidae, Apogonidae, Plesiopidae, Priacanthidae,

Centropomidae. Brill, E.J. Ltd., Leiden, 5, pp. i‒xiv + 458.

Whitley, G.P. 1951. New Fish Names and Records.
Whitley, G.P. 1959 (September, 18). More Ichthyological Snippets.
Whitley, G.P. 1964. Fishes from the Coral Sea and the Swain Reefs. =including description of Apogonichthys coggeri.
Yoshida, T., M. Hayashi and H. Motomura. 2018. Ostorhinchus yamato, a new species of cardinalfish (Perciformes:
Apogonidae) from Japan. Ichthyological Research v. 66 (no. 2): [1-9] 230-238.
Yoshida, T., M. Hayashi and H. Motomura. 2018. Rhabdamia novaluna, a new species of cardinalfish (Perciformes:
Apogonidae) from the western Pacific Ocean, with comments on the synonymy of Rhabdamia gracilis. Ichthyological
Research v. 66 (no. 1): [1-11] 129-139.
Yoshida, T. and H. Motomura. 2017. Distributional Range Extension and Live Coloration of the Indo-Pacific Deepwater
Cardinalfish Ostorhinchus cheni (Perciformes: Apogonidae). The Japanese Society of Systematic Zoology. Species
Diversity 22: 225–230.
Yoshida, T., T. Kawai and H. Motomura. 2019. Gymnapogon sagittarius, a new species of cardinalfish (Perciformes:
Apogonidae) from the South China Sea and Andaman Sea. The Ichthyological Society of Japan.
Yoshida, T., K. Koeda and H. Motomura. 2018. Distributional Range Extension of Xeniamia atrithorax (Perciformes:
Apogonidae) in the northern South China Sea. The Japanese Society of Systematic Zoology. Species
Diversity 23: 239–241.
Websites consulted
http://fishesofaustralia.net.au/
http://researcharchive.calacademy.org/research/ichthyology/catalog/fishcatmain.asp
http://fishpix.kahaku.go.jp/fishimage-e/search.html
https://www.fishbase.se/search.php
https://www.fishwisepro.com/
https://stri.si.edu/
Index scientific names

affinis, Paroncheilus ..............42 maculatus ........................23 umbratilis ..............................162
africanus, Gymnapogon .........74 marquesensis ....................37 aff. umbratilis ........................162
ahimsa, Apogonichthys ............6 mosavi .............................30 uninotatus .............................155
albimaculosus, Ozichthys .......17 pacificus ...........................29 Apogonichthys ............................6
albomarginata, Jaydia .........169 phenax .............................27 ahimsa ......................................6
alleni, Cheilodipterus ............53 pillionatus ........................25 infuscus .....................................7
alutus, Astrapogon ................42 planifrons ........................26 ocellatus ....................................7
amblyuroptera, Fowleria ........14 posterofasciatus ................38 perdix ........................................6
amblyuroptera, Pseudamia ..189 pseudomaculatus ..............24 waikiki ......................................7
amboinensis, Fibramia .........183 quadrisquamatus ..............31 APOGONINAE ....................................5
americanus, Apogon ..............24 retrosella ..........................28 APOGONICHTHYINI ........................5
Amioides ...............................4 robbyi ..............................31 APOGONINI ................................22
polyacanthus ......................4 robinsi ..............................30 ARCHAMIINI ...............................44
AMIOIDINAE ...............................4 rubrifuscus .......................39 CHEILODIPTERINI .........................52
angustatus, Ostorhinchus ....120 seminigracaudus ...............38 GLOSSAMIINI ..............................66
annona, Gymnapogon ...........74 semiornatus .....................33 GYMNAPOGONINI .........................71
annularis, Nectamia ............174 sp. 1 ................................34 LEPIDAMIINI ................................79
aphanes, Ostorhinchus ........103 sp. 2 ................................34 OSTORHINCHINI ..........................82
Apogon ...............................22 susanae ............................35 PRISTIAPOGONINI .......................129
americanus .......................24 talboti ..............................32 RHABDAMIINI ............................137
atradorsatus .....................28 aff. talboti ........................32 SIPHAMIINI ...............................141
atricaudus ........................28 townsendi ........................26 SPHAERAMIINI ...........................155
aurolineatus .....................30 tricinctus ..........................39 VERULUXINI .............................140
axillaris ............................25 unicolor ...........................32 ZORAMIINI ................................182
binotatus ..........................25 aff. unicolor ......................34 apogonoides, Ostorhinchus ...........83
campbelli ..........................36 Apogonichthyoides .........155 aprion, Glossamia ........................66
caudicinctus .....................38 atripes ............................157 arabica, Siphamia .......................148
coccineus ..........................35 brevicaudatus .................161 arabicus, Cheilodipterus ...............61
crassiceps .........................37 cantoris ..........................164 Archamia ...................................44
dammermani ....................33 chrysurus .......................161 bleekeri ...................................44
deetsie ..............................39 enigmaticus ....................160 argentea, Siphamia ....................151
dianthus ...........................39 erdmanni .......................158 arguni, Glossamia .........................69
doryssa .............................35 euspilotus .......................159 argyrogaster, Jaydia ....................170
dovii .................................29 gardineri ........................160 arnazae, Siphamia ......................151
erythrinus .........................37 heptastygma ...................157 aroubiensis, Ostorhinchus ...........118
erythrosoma .....................36 maculipinnis ...................163 articycla, Neamia .........................16
gouldi ...............................27 melas .............................158 artus, Cheilodipterus ....................59
guadalupensis ...................28 miniatus .........................158 Astrapogon ................................41
hypselonotus ....................34 niger ..............................156 alutus ......................................42
imberbis ...........................23 nigripinnis ......................164 puncticulatus ...........................41
indicus .............................36 opercularis .....................164 stellatus ...................................41
kautamea .........................39 pharaonis .......................159 ataenia, Taeniamia .......................50
kominatoensis ..................38 regani ............................163 aterrimus, Ostorhinchus .............117
lachneri ............................30 sialis ..............................161 atradorsatus, Apogon ...................28
lativittatus ........................34 taeniatus ........................159 atricaudus, Apogon ......................28
leptocaulus .......................27 timorensis ......................163 atripes, Apogonichthyoides .........157

atrithorax, Xeniamia ..................177 ellioti, Jaydia ..........................165 africanus ..............................74

aureus, Ostorhinchus ....................84 elongata, Siphamia .................153 annona ................................74
aurita, Fowleria ............................12 endekataenia, Ostorhinchus ....111 japonicus .............................74
aurolineatus, Apogon ...................30 enigmaticus, Apogonichthyoides 160 philippinus ...........................74
axillaris, Apogon ..........................25 erdmanni, Apogonichthyoides .158 sagittarius ............................75
badia, Vincentia ...........................20 eremia, Cercamia ......................73 sp. .......................................76
bandanensis, Nectamia ...............175 erythrinus, Apogon ...................37 vanderbilti ...........................75
beauforti, Glossamia .....................69 erythrophthalma, Jaydia .........170 velum ..................................75
biguttata, Taeniamia .....................46 erythrosoma, Apogon ...............36 hartzfeldii, Ostorhinchus .........110
bilineata, Taeniamia .....................51 euspilotus, Apogonichthyoides ...159 hayashii, Pseudamia ...............189
binotatus, Apogon ........................25 evermanni, Zapogon .................40 heptastygma, Apogonichthyoides ..157
bleekeri, Archamia .......................44 fasciatus, Ostorhinchus .............89 heptazona, Cheilodipterus ........54
brachygramma, Foa ........................8 Fibramia ..............................182 heurni, Glossamia .....................69
brevicaudatus, Apogonichthyoides 161 amboinensis .......................183 hoevenii, Ostorhinchus ...........127
brevilux, Siphamia ......................153 lateralis .............................183 Holapogon ...............................5
brevispinis, Ostorhinchus ............118 thermalis ...........................182 maximus ................................5
bryx, Ostorhinchus .......................95 fistulosa, Siphamia .................150 holotaenia, Ostorhinchus ........109
buruensis, Taeniamia ....................49 flagelliferus, Ostorhinchus ......126 hungi, Jaydia ..........................167
campbelli, Apogon .......................36 flammea, Fowleria ....................14 hyalina, Foa ..............................11
cantoris, Apogonichthyoides .......164 flavofasciata, Taeniamia ............45 hyalosoma, Yarica .....................70
capricornis, Ostorhinchus .............86 flavus, Ostorhinchus .................86 hypselonotus, Apogon ..............34
carinata, Jaydia ..........................166 flebila, Zoramia ......................186 ignitops, Nectamia ..................176
catalai, Jaydia ............................170 fleurieu, Ostorhinchus ..............85 imberbis, Apogon .....................23
caudicinctus, Apogon ...................38 fo, Foa .......................................9 indicus, Apogon ........................36
cavitensis, Ostorhinchus .............109 Foa ............................................8 infuscus, Apogonichthys .............7
cephalotes, Siphamia ..................142 brachygramma .......................8 intermedius, Cheilodipterus ......56
Cercamia ...................................71 fo ..........................................8 isostigma, Cheilodipterus ..........63
cladara ....................................71 hyalina ................................11 isostigma, Fowleria ...................13
eremia .....................................73 landoni ..................................9 isus, Zapogon ...........................40
melanogaster ...........................72 leisi .....................................10 japonicus, Gymnapogon ............74
sp. 1 ........................................72 madagascariensis .................11 Jaydia ...................................165
sp. 2 ........................................73 nivosa ..................................10 argyrogaster .......................170
chalcius, Ostorhinchus ................126 sp. 1 ....................................10 carinata .............................166
Cheilodipterus ..........................52 sp. 2 ....................................10 catalai ................................170
alleni .......................................53 sp. 3 ....................................11 ellioti .................................165
arabicus ...................................61 yamba .................................10 erythrophthalma ................170
artus ........................................58 zuluensis ..............................11 hungi .................................167
heptazona ................................54 Fowleria .................................12 lineata ...............................168
intermedius .............................56 abocellata ............................15 melanopus .........................166
isostigma .................................63 amblyuroptera .....................14 novaeguineae .....................169
cf. isostigma ............................63 aurita ..................................12 photogaster ........................170
lachneri ...................................60 flammea ..............................14 poeciloptera .......................165
lineatus ...................................55 isostigma .............................13 quartus ..............................170
nigrotaeniatus ..........................65 marmorata ...........................12 queketti .............................167
novemstriatus ..........................64 sp. 1 & 2 .............................14 smithi ................................165
octolineatus .............................53 vaiulae .................................15 striata ................................168
parazonatus .............................65 variegata ..............................13 striatodes ...........................168
persicus ...................................61 fraenatus, Pristiapogon ...........129 truncata .............................169
pygmaios .................................64 fragilis, Zoramia .....................186 jebbi, Siphamia .......................150
quinquelineatus .......................62 franssedai, Ostorhinchus .........101 jenkinsi, Ostorhinchus ..............96
singaporensis ...........................57 fraseri, Siphamia ....................154 kagoshimana, Taeniamia ...........47
sp. 1 ........................................62 fucata, Taeniamia .....................48 kallopterus, Pristiapogon ........130
zonatus ....................................65 fukuii, Ostorhinchus ...............112 kalosoma, Lepidamia ................79
cheni, Ostorhinchus ....................113 fusca, Nectamia ......................172 kauderni, Pterapogon .............178
chrysopomus, Ostorhinchus ........122 fuscolineata, Siphamia ............146 kautamea, Apogon ...................39
chrysotaenia, Ostorhinchus .........111 gardineri, Apogonichthyoides ..160 kiensis, Ostorhinchus ................94
chrysurus, Apogonichthyoides ....161 gelatinosa, Pseudamia ............191 kominatoensis, Apogon ............38
cladara, Cercamia .........................71 gilberti, Zoramia .....................185 komodoensis, Ostorhinchus ......83
cladophilos, Ostorhinchus .............98 gjellerupi, Glossamia ................68 lachneri, Apogon ......................30
compressus, Ostorhinchus ..........121 Glossamia ..............................66 lachneri, Cheilodipterus ............60
concilians, Paxton ......................188 abo ......................................68 landoni, Foa ...............................9
conklini, Phaeoptyx ......................43 aprion ..................................66 lateralis, Fibramia ...................183
conspersa, Vincentia .....................18 arguni ..................................69 lativittatus, Apogon ..................34
cookii, Ostorhinchus ...................116 beauforti ..............................69 leai, Taeniamia .........................46
corallicola, Siphamia ..................143 gjellerupi .............................68 leisi, Foa ...................................10
crassiceps, Apogon .......................37 heurni ..................................69 Lepidamia ..............................79
cuneiceps, Siphamia ...................142 narindica .............................68 kalosoma .............................79
cyanophthalma, Siphamia ..........150 sandei ..................................68 multitaeniata .......................80
cyanosoma, Ostorhinchus ...........104 timika ..................................67 natalensis ............................80
cypselurus, Verulux ....................140 trifasciata ............................67 noordzieki ............................79
dammermani, Apogon ..................33 wichmanni ...........................68 omanensis ...........................80
deetsie, Apogon ............................39 goreni, Siphamia ....................148 cf. omanensis .......................81
dianthus, Apogon .........................39 gouldi, Apogon .........................27 leptacantha, Zoramia ..............184
diaphanes, Pseudamiops ...............78 gracilicauda, Pseudamiops ........78 leptocaulus, Apogon .................27
dispar, Ostorhinchus ...................101 gracilis, Rhabdamia ................137 leptofasciatus, Ostorhinchus .....97
doederleini, Ostorhinchus ...........114 griffini, Ostorhinchus ..............126 leslie, Ostorhinchus ................109
doryssa, Apogon ...........................35 gularis, Ostorhinchus ................99 limenus, Ostorhinchus ............115
dovii, Apogon ...............................29 guttulata, Siphamia ................154 lineata, Jaydia ........................168
elizabethae, Ostorhinchus .............90 Gymnapogon .........................74 lineatus, Cheilodipterus ............55

lineolata, Taeniamia ....................47 aterrimus .............................117 sp. 2 ..................................102

lineomaculatus, Ostorhinchus ...102 atrogaster ...............................99 sp. 3 ..................................102
luteus, Ostorhinchus .................107 aureus ....................................84 sp. 4 ..................................105
luxuria, Nectamia .....................176 brevispinis ............................118 sp. 5 ..................................105
macrocauda, Vincentia ................21 bryx .......................................95 sp. 6 ..................................106
macroptera, Taeniamia ................48 capricornis .............................86 sp. 7 ..................................107
maculatus, Apogon .....................23 cavitensis .............................109 sp. 8 ..................................107
maculiferus, Ostorhinchus .........126 chalcius ................................126 sp. 9 ..................................108
maculipinnis, Apogonichthyoides ....163 cheni ....................................113 sp. 10 ................................110
madagascariensis, Foa .................11 chrysopomus ........................122 sp. 11 ................................113
majimai, Siphamia ....................147 chrysotaenia .........................111 sp. 12 ................................118
margaritophorus, Ostorhinchus .....124 cladophilos .............................98 sp. 13 ................................118
marmorata, Fowleria ...................12 compressus ..........................121 taeniophorus ......................115
marquesensis, Apogon ................37 cookii ...................................116 unitaeniatus .........................97
maximus, Holapogon ....................5 cyanosoma ...........................104 victoriae .............................116
melanogaster, Cercamia ..............72 dispar ..................................101 yamato ................................99
melanoproctus, Ostorhinchus ....101 doederleini ...........................114 oxina, Ostorhinchus ................124
melanopus, Jaydia ....................166 elizabethae .............................90 oxygrammus, Ostorhinchus .....123
melas, Apogonichthyoides .........158 endekataenia ........................111 Ozichthys ...............................17
melasma, Taeniamia ...................46 fasciatus .................................89 albimaculosus ......................17
menesemus, Pristiapogon ..........134 cf. fasciatus-1 .........................90 pacificus, Apogon .....................29
microspilus, Ostorhinchus ...........85 cf. fasciatus-2 .........................91 pallida, Taeniamia ....................51
miniatus, Apogonichthyoides ....158 cf. fasciatus-3 .........................91 pallidofasciatus, Ostorhinchus .117
mirifica, Quinca ........................180 cf. fasciatus-4 .........................93 pallidus, Ostorhinchus ..............98
moluccensis, Ostorhinchus ........125 flagelliferus ..........................126 papuensis, Siphamia ...............152
monospilus, Ostorhinchus .........125 flavus .....................................86 parazonatus, Cheilodipterus ......65
mosavi, Apogon ..........................30 fleurieu ..................................85 Paroncheilus .........................42
mossambica, Siphamia ..............148 franssedai ............................101 affinis ..................................42
mozambiquensis, Taeniamia ........45 fukuii ...................................112 parvulus, Ostorhinchus ...........100
multilineatus, Ostorhinchus ......121 griffini ..................................126 Paxton ..................................188
multitaeniata, Lepidamia ............80 gularis ....................................99 concilians ...........................188
nanus, Ostorhinchus ...................99 hartzfeldii ............................110 PAXTONINAE ..............................188
narindica, Glossamia ...................68 hoevenii ...............................127 pellucidus, Pseudamiops ...........77
natalensis, Lepidamia .................80 holotaenia ............................109 perdix, Apogonichthys ................6
Neamia .....................................16 jenkinsi ..................................96 perlita, Zoramia ......................185
articycla .................................16 kiensis ...................................94 permutata, Siphamia ..............152
notula ....................................17 cf. kiensis ...............................95 persicus, Cheilodipterus ............61
octospina ...............................16 komodoensis ..........................83 Phaeoptyx ..............................43
xenica ....................................17 leptofasciatus .........................97 conklini ...............................43
Nectamia ................................171 leslie ....................................109 pigmentaria .........................43
annularis ..............................174 limenus ................................115 xenus ...................................43
bandanensis .........................175 lineomaculatus .....................102 pharaonis, Apogonichthyoides 159
fusca ....................................172 luteus ...................................107 phasma, Pseudamiops ...............78
ignitops ................................176 maculiferus ..........................126 phenax, Apogon .......................27
luxuria .................................176 margaritophorus ...................124 philippinus, Gymnapogon .........74
savayensis ............................173 melanoproctus .....................101 photogaster, Jaydia .................170
similis ..................................177 microspilus .............................85 pigmentaria, Phaeoptyx ............43
viria .....................................175 moluccensis .........................125 pillionatus, Apogon ..................25
zebrina .................................174 monospilus ..........................125 planifrons, Apogon ...................26
nematoptera, Sphaeramia .........181 multilineatus ........................121 pleuron, Ostorhinchus ..............90
neotes, Ostorhinchus .................100 nanus .....................................99 poeciloptera, Jaydia ................165
niger, Apogonichthyoides ..........156 neotes ..................................100 polyacanthus, Amioides ..............4
nigrimentum, Rhabdamia ..........139 nigripes ................................128 posterofasciatus, Apogon ..........38
nigripes, Ostorhinchus ..............128 nigrocinctus ...........................87 Pristiapogon ........................129
nigripinnis, Apogonichthyoides .164 nigrofasciatus .......................119 fraenatus ...........................129
nigrocinctus, Ostorhinchus ..........87 norfolcensis ............................87 kallopterus .........................130
nigrofasciatus, Ostorhinchus .....119 notatus ..................................97 menesemus ........................134
nigrotaeniatus, Cheilodipterus .....65 novemfasciatus .....................120 taeniopterus .......................134
nivosa, Foa .................................10 ocellicaudus ...........................98 urostigma ...........................132
norfolcensis, Ostorhinchus ..........87 oxina ...................................124 Pristicon ...............................135
notatus, Ostorhinchus .................97 oxygrammus ........................123 rhodopterus .......................136
notula, Neamia ...........................17 pallidofasciatus ....................117 rufus ..................................136
novaeguineae, Jaydia ................169 pallidus ..................................98 trimaculatus .......................135
novaehollandiae, Vincentia .........21 parvulus ...............................100 properuptus, Ostorhinchus ......106
novaluna, Rhabdamia ...............139 pleuron ..................................90 pselion, Ostorhinchus .............103
novemfasciatus, Ostorhinchus ...120 properuptus .........................106 Pseudamia ...........................189
novemstriatus, Cheilodipterus .....64 pselion .................................103 amblyuroptera ...................189
ocellatus, Apogonichthys ...............7 quadrifasciatus .......................92 gelatinosa ..........................191
ocellicaudus, Ostorhinchus ..........98 cf. quadrifasciatus ..................93 hayashii .............................189
octospina, Neamia ......................16 regula ..................................118 rubra .................................191
omanensis, Lepidamia .................80 relativus ...............................122 tarri ...................................191
opercularis, Apogonichthyoides ....164 rueppellii .............................103 zonata ...............................190
orbicularis, Sphaeramia .............180 schlegeli ...............................112 PSEUDAMIINAE ...........................189
Ostorhinchus ...........................82 sealei ...................................123 Pseudamiops .........................77
aff. victoriae .........................116 selas ....................................103 diaphanes ............................78
angustatus ...........................120 semilineatus ...........................96 gracilicauda .........................78
aphanes ...............................103 septemstriatus ........................93 pellucidus ............................77
apogonoides ...........................83 sinus ....................................122 phasma ................................78
aroubiensis ...........................118 sp. 1 ....................................102 springeri ..............................78

pseudomaculatus, Apogon .........24 fistulosa ..............................150 tubulata, Siphamia ...................149

Pterapogon ...........................178 fraseri ..................................154 umbratilis, Apogonichthyoides .162
kauderni .............................178 goreni ..................................148 unicolor, Apogon ........................32
punctata, Vincentia ....................20 guttulata .............................154 uninotatus, Apogonichthyoides 155
puncticulatus, Astrapogon ..........41 jebbi ....................................150 unitaeniatus, Ostorhinchus .........97
pygmaios, Cheilodipterus ...........64 majimai ...............................147 urostigma, Pristiapogon ...........132
quadrifasciatus, Ostorhinchus .....92 mossambica .........................148 vaiulae, Fowleria ........................15
quadrisquamatus, Apogon ..........31 papuensis ............................152 vanderbilti, Gymnapogon ...........75
quartus, Jaydia ........................170 permutata ............................152 variegata, Fowleria .....................13
queketti, Jaydia .......................167 randalli ...............................151 velum, Gymnapogon ..................75
Quinca ...................................180 roseigaster ...........................143 versicolor, Siphamia .................145
mirifica ...............................180 senoui .................................150 Verulux ..................................140
quinquelineatus, Cheilodipterus ....62 spinicola ..............................153 cypselurus ...........................140
randalli, Siphamia ....................151 stenotus ..............................154 solmaculata .........................140
regani, Apogonichthyoides .......163 tubifer .................................144 sp. 1 ...................................141
regula, Ostorhinchus ................118 tubulata ...............................149 victoriae, Ostorhinchus .............116
relativus, Ostorhinchus .............122 versicolor .............................145 Vincentia .................................18
retrosella, Apogon .....................28 zaribae ................................145 badia .....................................20
Rhabdamia ...........................137 solmaculata, Verulux ................140 conspersa ..............................18
gracilis ................................137 Sphaeramia ...........................180 macrocauda ...........................21
nigrimentum .......................139 nematoptera ........................181 novaehollandiae ....................21
novaluna .............................139 orbicularis ...........................180 punctata ................................20
sp. 1 ...................................138 spilota, Rhabdamia ...................138 viria, Nectamia .........................175
sp. 2 ...................................139 spinicola, Siphamia ..................153 viridiventer, Zoramia ................186
spilota .................................138 springeri, Pseudamiops ...............78 waikiki, Apogonichthys ................7
rhodopterus, Pristicon ..............136 stellatus, Astrapogon ..................41 wassinki, Ostorhinchus .............107
robbyi, Apogon ..........................31 stenotus, Siphamia ...................154 wichmanni, Glossamia ...............68
robinsi, Apogon .........................30 striata, Jaydia ...........................168 Xeniamia ...............................177
roseigaster, Siphamia ...............143 striatodes, Jaydia .....................168 atrithorax ............................177
rubra, Pseudamia .....................191 susanae, Apogon ........................35 xenica, Neamia ..........................17
rubrifuscus, Apogon ...................39 Taeniamia ................................45 xenus, Phaeoptyx .......................43
rueppellii, Ostorhinchus ...........103 ataenia ..................................50 yamato, Ostorhinchus ................99
rufus, Pristicon ........................136 biguttata ................................46 yamba, Foa ................................10
sagittarius, Gymnapogon ............75 bilineata ................................51 Yarica .......................................70
sandei, Glossamia ......................68 buruensis ...............................49 hyalosoma .............................70
sansibarica, Taeniamia ...............49 flavofasciata ..........................45 Zapogon ..................................40
savayensis, Nectamia ................173 fucata ....................................48 evermanni .............................40
schlegeli, Ostorhinchus .............112 kagoshimana .........................47 isus .......................................40
sealei, Ostorhinchus .................123 leai ........................................46 zaribae, Siphamia ....................145
selas, Ostorhinchus ..................103 lineolata ................................47 zebrina, Nectamia ....................174
semilineatus, Ostorhinchus .........96 macroptera ............................48 zonata, Pseudamia ...................190
seminigracaudus, Apogon ..........38 melasma ................................46 zonatus, Cheilodipterus ..............65
semiornatus, Apogon .................33 mozambiquensis ....................45 Zoramia .................................184
senoui, Siphamia .....................150 pallida ...................................51 flebila .................................186
septemstriatus, Ostorhinchus .....93 sansibarica .............................49 fragilis .................................186
sialis, Apogonichthyoides .........161 zosterophora ..........................50 gilberti ................................185
similis, Nectamia ......................177 taeniatus, Apogonichthyoides ...159 leptacantha .........................184
singaporensis, Cheilodipterus .....57 taeniophorus, Ostorhinchus ......115 perlita .................................185
sinus, Ostorhinchus ..................122 taeniopterus, Pristiapogon ........134 sp. 1 ...................................187
Siphamia ...............................141 talboti, Apogon ..........................32 viridiventer ..........................186
arabica ................................148 tarri, Pseudamia .......................191 zosterophora, Taeniamia ............50
argentea ..............................151 thermalis, Fibramia ..................182
arnazae ...............................151 timorensis, Apogonichthyoides .163
brevilux ...............................153 townsendi, Apogon .....................26
cephalotes ...........................142 tricinctus, Apogon ......................39
corallicola ...........................143 trifasciata, Glossamia .................67
cuneiceps ............................142 trimaculatus, Pristicon ..............135
cyanophthalma ....................150 truncata, Jaydia .......................169
elongata ..............................153 tubifer, Siphamia ......................144

Apogonidae Fishes LRs

Uploaded by

Document Information

Original Title

Copyright

Available Formats

Share this document

Share or Embed Document

Sharing Options

Did you find this document useful?

Is this content inappropriate?

Copyright:

Available Formats

Apogonidae Fishes LRs

Uploaded by

Copyright:

Available Formats

CardinalfishesCover_CardinalfishesCover 8/09/19 7:17 PM Page 1

Cardinalfishes of the World – Kuiter & Kozawa

Anthis – Aquatic Photographics

Cardinalfishes of the World – Kuiter & Kozawa

Anthis – Aquatic Photographics

Rudie H kuiter & Toshikazu Kozawa

II Cardinalfishes of the World – Kuiter & Kozawa

Key to subfamilies of APOGONIDAE (after Mabuchi et al, 2014)

Key to the genera of Apogonidae (after Mabuchi et al, 2014)

2 Cardinalfishes of the World – Kuiter & Kozawa

– Edge of preopercle smooth (Rhabdamiini) ..........................................................................................................25

1st dorsal fin 2nd dorsal fin

Fin terminology used (Apogon cyanosoma illustration after Bleeker, 1853).

Genus Amioides Smith & Radcliffe, 1912

Deep-dwelling Cardinalfish Amioides polyacanthus

4 Cardinalfishes of the World – Kuiter & Kozawa

GENUS Holapogon Fraser, 1973

Titan Cardinalfish Holapogon maximus

Holapogon maximus At 38 m. Oman. JER

Cardinalfishes of the World – Kuiter & Kozawa 5

TRIBE APOGONICHTHYINI – GENUS Apogonichthys Bleeker, 1854

Gentle Gobbleguts Apogonichthys ahimsa

6 Cardinalfishes of the World – Kuiter & Kozawa

Waikiki Cardinalfish Apogonichthys waikiki

Tan Cardinalfish Apogonichthys infuscus

Apogonichthys infuscus Dahab, Egypt, Red Sea. SBO

Ocellated Cardinalfish Apogonichthys ocellatus

Cardinalfishes of the World – Kuiter & Kozawa 7

TRIBE APOGONICHTHYINI – Genus Foa Jordan & Evermann, 1905

Weed Cardinalfish Foa brachygramma

8 Cardinalfishes of the World – Kuiter & Kozawa

Landon’s Cardinalfish Foa landoni

Cardinalfishes of the World – Kuiter & Kozawa 9

Harbour Cardinalfish Foa yamba

Snowy Cardinalfish Foa nivosa Leis’ Cardinalfish Foa leisi

White-speckled Cardinalfish Foa sp. 1 Heemstra’s Cardinalfish Foa sp. 2

Foa sp. 1. Maldives. JER Foa sp. 2. Mauritius. PCH

10 Cardinalfishes of the World – Kuiter & Kozawa

Snow-flakes Cardinalfish Foa sp. 3

Tide-pool Cardinalfish Foa madagascariensis

Sharp-snout Cardinalfish Foa hyalina

Cardinalfishes of the World – Kuiter & Kozawa 11

TRIBE APOGONICHTHYINI – Genus Fowleria Jordan & Evermann, 1903

Crosseye Cardinalfish Fowleria aurita

Eared Cardinalfish Fowleria marmorata

12 Cardinalfishes of the World – Kuiter & Kozawa

Dotted Cardinalfish Fowleria isostigma

At some stage the Red Sea spotted form was identified

Variegated Cardinalfish Fowleria variegata

Cardinalfishes of the World – Kuiter & Kozawa 13

Chestnut Cardinalfish Fowleria sp. 1 & 2

Flame Cardinalfish Fowleria amblyuroptera

Stop Light Cardinalfish Fowleria flammea

14 Cardinalfishes of the World – Kuiter & Kozawa

Dwarf Cardinalfish Fowleria vaiulae complex

Cardinalfishes of the World – Kuiter & Kozawa 15

TRIBE APOGONICHTHYINI – Genus Neamia Smith & Radcliffe, 1912

Eightspine Cardinalfish Neamia octospina

Circular Cardinalfish Neamia articycla

16 Cardinalfishes of the World – Kuiter & Kozawa

Gillspot Cardinalfish Neamia notula

Strange Cardinalfish Neamia xenica

Neamia xenica Holotype. Kiritimati Atoll, Kiribati. RPY