COASTAL FISHES OF THE
WESTERN INDIAN OCEAN:

A Longstanding Vision Realized

South African Institute for Aquatic Biodiversity
Somerset Street, Private Bag 1015, Grahamstown,
Eastern Province, South Africa, www.saiab.ac.za
email: WIO@saiab.ac.za
Tel: +27 (0) 46 603 5800
Fax: +27 (0) 46 622 2403

COASTAL FISHES OF THE WESTERN INDIAN OCEAN:

A Longstanding Vision Realized
Origins

Volume 2

In 1949, not long after the publication of The Sea Fishes of Southern Africa, JLB Smith
was offered the directorship of the Natal Museum in Pietermaritzburg. He turned theoffer down, as he wanted to focus on a book on the fishes of the Western Indian Ocean,which he said would take him a year or two. The vision was never fulfilled.
In 1986, the final successor to The Sea Fishes ofSouthern Africa, Margaret Smith and Phil Heemstra’s
Smiths’ Sea Fishes, was published. Margaret died in
1987. Some 10 years later, Phil resurrected JLB’s ideaof a book on the fishes of the Western Indian Ocean.
And now, 20 years after the re-birth of JLB Smith’sidea, this is about to become a reality.

Who put the book together?
Coastal Fishes of the Western Indian Ocean, with main
editors Philip Heemstra, Elaine Heemstra, David
Ebert, Wouter Holleman and John Randall, is to be
published early in 2018. It is the culmination of thework of more than 100 authors, photographersand illustrators, of editors, proofreaders, reviewers
and others, over a period of more than 20 years.
Amongst the major contributors are Phil Heemstra,
Dave Ebert, Dave Smith, Bruce Collette, Stuart Poss,Jack Randall, Gerry Allen, Helen Larson, Danny
Hensley and Kunio Amaoka, Eric Anderson andKeiichi Matsuura.

The Western Indian Ocean (WIO) is home to oneof the richest biotas of marine fishes on the planet,comprising some 3600 species, or about 20% of theworld’s marine fish fauna. Although the WIO regionis the site for most of the earliest scientifically valid
descriptions of Indo-Pacific fishes, the extent of its
fish fauna is still poorly known. Despite considerable
effort by ichthyologists over the past two centuries,
many new species of fishes are described each year:of the 329 new marine species described between
Plate 20
2002 and 2012, 140 were from the WIO.

Coverage and contents

The resulting publication, far more substantial
than JLB Smith could have imagined, arranged in
MyLIOBAtIDAE, Family
eaglerays;
Family RhInOptERIDAE,
CownoseRays
MyLIOBAtIDAE, eaglerays; Family
RhInOptERIDAE,
CownoseRays
six volumes,
covers species that occur in waters
generally shallower than 200 m. In this work, theFamily MyLIOBAtIDAE, eaglerays; Family RhInOptERIDAE, Cownose Rays Western Indian Ocean includes the Red Sea andPersian Gulf and extends to Cape Point, South Africa,
and Kanyakumari, the southernmost tip of India.
22
1
1
Some authors have included fishes from the Gulfof Mannar and wider Sri Lanka.
3

1

2

3

3
4

5a

Volume 15a66Volume 3
Volume 4
Volume 5
Volume 6

Introductory chapters which include the oceanography of the WIO, the origins of coral reefs,
an account of the people who laid the foundations of our knowledge of the WIO’s fishes, andmore. The remainder of the volume covers the agnaths and condrichthyans of the area.
This volume includes a brief overview of the evolution of bony fishes, their anatomy and a key
to the orders. Then follows accounts of families of the Orders Elopiformes to Mugiliformes.
The remainder of the non-perciforms teleosts, from the Atheriniformes to the Scorpaeniformes.
A key to the families of Perciformes, and the families Latidae to Chaetodontidae
The key to the families, and the families Oplegnathidae to Draconettidae.
The key to the families, and the families Gobiidae to Molidae, and the Latimeriidae.

Each taxon has its appropriate keys and, for several of the families, a glossary of terms used in the accounthas been included. The volumes are well illustrated, with black-and-white illustration in keys and for species,
where appropriate, and colour plates of the species. The inside back and front covers of each volume will
have various maps of the Western Indian Ocean.

Format and cost
The book will measure about 220mm wide x 280mm deep. Consideration is being given to both hard-coverand soft-cover binding. The cost for a hard-cover will be in the region of R3500 per set ($270, €240)and soft-cover R2000 ($150, €130).

Expression of interest
If you are interested in this publication, please 1) access the reply form on our website
http://www.saiab.ac.za/coastal-fishes-of-the-western-indian-ocean.htm and complete the online form,
OR 2) complete the form below, scan and mail it to WIO@saiab.ac.za, OR 3) post it to:
WIO Fishes – Expression of Interest, SAIAB, Private Bag 1015, Grahamstown, 6140, South Africa
This will help assess the number of copies to have printed.
Your name:Institution:
Address:

Email Address:
Binding preference:

Hard cover

Soft coverNr. of sets

Purpose:Institution

Library

PersonalOrder Beryciformes

es)Class Osteichthyes (Bony Fishby Phillip C� Heemstra, bony, platecompressed, with enlarged
Body oval but somewhat
to form acarinate spine and fused
like scales , bearing a sharp,
stout, ridged
dorsal fins, first with 4–7
solid, bony armour. Two
dorsal fin
necting membranes; second
spines without intercon
a transparentted) rays connected bywith 11–12 soft (segmen
fin forked,
to soft dorsal fin; caudal
membrane; anal fin similarfinstips, branched rays 17; pelvic
the lobes short, with rounded
fin spines
tiny soft rays; the pelvic
with a large spine and 3–4
of belly
fit into a groove along sides
and
and
erect
lockedbecanlight organ on
short, bluntly rounded;
when depressed. Snoutl.
lower jaw. Solitary, demersa
family3–400 m. An Indo-PacificPineapple fishes live in
pic
species in WIO. The monoty
1
species;4
andwith 2 genera
has the jaws
endemic to Australia. It
Cleidopus gloriamaris is
al space, and the light organ
suborbitnarrowstrongly curved,
when mouth is closed).
(hidden
jaw
lower
the
of
is on the sideby Phillip C� Heemstra, bony, plateSL
Monocentris japonica: 11 cm
compressed, with enlargedview(O. Alvheim, IMR); ventral
Body oval but somewhat
near
e and fused to form a
spinof lower jaw with light organs
ate
carin
p,
shar
(both
a
symphysis shown by arrows
like scales , bear1aing
ed
ridg
t,
stou
Mozambique).
with 4–7
first
fins,
al
dors
Two
solid, bony armour.
nd dorsal fin
ecting membranes; secospines without interconn 2ad by a transparent
ecte
Family Anomalopidae
conn
rays
ted)
men
with 11–12 soft (segFlashlight fishes
fin; caudal fin forked,
al
dors
soft
to
lar
simi
by John E� McCosker
membrane; anal fin
pelvics fins
Monocentris Bloch & Schneider, 1801eye. Body
cent bacteria at
ded tips, branched rays 17; Genu
organ located beneath eachorgan containing lumines
A prominent luminous
Jaws straight; lightcompressed. Dorsal fin
the lobes short, with rounspines chin. Three species, 1 in WIO.
SL at fusiform, laterally
pelvic fin
or somewh
cm
deep
the
11
;
front of lower jaw undered fins.
ica:
rays
japon
soft
ris
tiny
ocent
separate spinous and soft-ray
3–4
Mon
3a
e and
continuous or divided into
strongly ctenoid.
with a large spin
minute,belly
Scalesofsview
blunt.
al
Mouth oblique; snout
s; none on vomer.
(O. Alvheim, IMR); ventr
fit into a groove along sideMonocentris japonica
in bands on jaws and palatine
Teeth small,nearscan be locked erect and and
on
n
organ
orga
light
ed.
light
Gill-rakers well develop
of lower jaw withs organ is continuously
short, bluntly rounded;
Pineapplefish
from the subocular luminoui, Japan). by arrowLights (bothd using
when depressed. Snout
1782: 329, Pl. 2 (Nagasak
n
4a.
show
Key to Families
bacteria and can be occlude
Gasterosteus japonicus Houttuyn symphysis
1986*;
produced by luminous
is usedSFSA*; Smith 1986*; Kotlyarl.
isms to block the light. Light
1a. Pelvic fins with 1 huge spine (which can be locked erect)
Monocentris japonicus Smith
rotational or lid-like mechan
Fricke 1999.
1996*;ue).
lower jaw. Solitary, demersa
Michelly
mbiq
1996*;
predators and to attractfami
and 3 or 4 rudimentary rays; scales enlarged,
KotlyarMoza
confusetoRandal 1995*;-PacificIndoAn
to see by, to communicate,m.
~100 m)0
with sharp ridges and spines, forming a bony armour(below
3–40
2004*
water
in
deep
Heemstra&in
live
Heemstraprey. Adults of most species
ssresembling a pineapple � � � � � � � � � � � � � � � � � � � � MONOCENTRIDAE
Pineapple fishes live
moonleic
during
feed
otyp
to
face
the reef
1b. Pelvic fins with 1 spine and 6B12 soft rays; body not encased
in WIO. The mon
during the day, and ascend
biology.)
LL 12–17; GR 5–7 / 11–14.
1977, for a review of theirin bony armour � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � �2
ra4a. and 4 species; 1 species
9–12; A 10–11; P 13–15;/jaws
�2
nights. (See McCosker,
with � � � genee behaviour,theVertebrae 11+15. Jaws straight
their nocturnal secretiv. It hasDBDIV–VII
ralia
Aust
to
Rarely seen because of7
1.3–1.9, HL 1.8–2.4 in SL.
with
mic
Head,
genera2a. Prominent luminous organ immediately
Sixendevomer.
onis
all tropical oceans.
and palatines, none
dopus gloriamaris
flashlight fishes occur inwith villiform
below eye � � � � � � � � � � � � � � � � � � � � � � � � � � � Clein on jawsscales
katoptron,
orgateeth� � ANOMALOPIDAE
likely (such as Anomalops
outlined in black; head naked.
e, and the light
2b. No light organ beneath eye � � � � � � � � � � � � � � � � � � � � � � � � � � � �
species, 1 in WIO, others
body and fins yellow, the�3ia to the south central Pacific)
ed, narrow suborbital spac
curv
gly
distributed from Indones
widely
stron
coral
ed).
on
(2007).
al.
et
th is Aclos
3a. Jaws with large, fang-like teeth; dorsal-fin basein caves and under ledges
be found. Recently reviewetoOrd er Berynocturnal species, founds inr jaw (hidden when mou
2 to 4 times as long as anal-fin base � � � � � � � � � � � ANOPLOGASTERIDAE
ciformes d by Baldwin
near wrecks and reefs. Juvenile
is on the side of the lowe
and rocky reefs, also on sand3b.

T

his is a poorly understood group that currently is
considered to include seven families: Berycidae,
Holocentridae, Anoplogasteridae, Trachichthyidae,
Anomalopidae, Diretmidae, and Monocentridae. They areprobably an artificial assemblage of unrelated taxa.
Most species live in deep marine waters and avoid bright
light, although may come closer to the surface at night. The
Holocentridae, which have their greatest diversity on coral
reefs, are largely or entirely nocturnal. The have relatively largeeyes. During the day they tend to remain hidden in crevaces, incase or under ledges.
Diagnosis: fin spines usually well developed (rudimentary
or absent in anoplogasterids, and in the dorsal and anal fins ofdiretmids); pelvic fins with a strong spine and 6–12 soft rays
(only 3 or 4 rudimentary rays in Monocentridae): caudal fin
with true (unpaired) small procurrent spines at base of caudal
fin and 18 or 19 principal caudal-fin rays. Six families that canbe recognised as coastal fishes occur in the WIO.

Family Anomalopidae

Jaws with minute teeth; dorsal and anal-fin bases subequal � � � � � � � � � 4

4a.
4b.

PLATE xx

Genus Monocentris

60

Order Beryciformes

1609 WIO Minion + Myriad2013.indd 60

1

Beryx mollis: 21 cm SL (offSomalia; from Kotlyar 1993).

Family Anoplogasteridae

Trachichthodes spinosus Gilchrist, 1903:
204, pl. 13 (type locality: off Kei
River mouth [Cape Morgan], South
Africa); Barnard, 1925a: 363*;
Fowler, 1935: 374; Smith, 1949: 152*.
Centroberyx spinosus: Busakhin, 1982:
14*; Heemstra, 1986: 410*; Smale etal., 1995: 78.

Gill, 1862

(total). Anterior edgeeryx
Gilchrist, 1903 and Austrob
2 in WIO. Trachichthodess of Centroberyx.
McCulloch, 1911 are synonym

Beryx splendens

Slender alfonsino

B. decadactylus)
142 (text only, figure is of
Beryx splendens Lowe, 1834:(type locality: Madeira).
h, 1929:
49; Günther, 1887: 33; McCulloc
Beryx splendens: Lowe, 1843a:
24; Abe, 1972:
1949: 150*; Krefft, 1961:
Smith,
542*;
1936a:
130; Fowler,
1976: 74*;
& Sonoda, 1973: 286* Krefft,
525; Karrer, 1973: 221; Woods
410*;
1984: 108* ; Heemstra, 1986:
al.,etMasuda6*;
1982:
Busakhin,Dubochkin &1987: 73; Ivanin, 989: 159;
Maul, 1986: 741*; Kotlyar,
1995: 78*; Adam et al.,1990: 257*; Smale et al.,
Kotlyar, 1989: 1; Mundy,

D V–VI, 14–15; A IV, 14–16; P 13;
V I, 7; LL continuous, notextending onto caudal-fin rays, 40–47;
GR 10–11/ 20–22. Depth
47–53% SL; head 35–40; length dorsal-finbase 38–43; length
anal-fin base 32–34; predorsal 38–44;
preanal 58–64; caudalfin 28–33; snout 15–20% HL; eye 31–47.
Posterior margin ofscales with denticles of varying lengths,
no ctenii. Cheek scales
5. Fulcral scale in pelvic axil well developed.
No head spines.
Dorsal-fin origin on vertical throughpelvic-fin origin. Anal fin
origin on vertical through middleof dorsal fin. Pectoral finslonger than pelvic fins.

cie s
Wio specie
tosWio LLspe
Key toKey
anal-fin origin 15; total GR
scale rows between andn origin 15; total GR
rows between LL and anal-fi
LL 40–47; transverse scale� � � � � � C. spinosus�����������������
30–34 � � � � � � � � � � � �anal-fin origin 18; total GR
scale rows between LL and
1b. LL 53–62; transverse� � � � C. druzhinini�����������������
�
�
�
�
�
�
�
�
�
�
�
�
9
Top of head, iris, back and caudal fin
druzhinini
bright red, body
Centroberyx26–2
de
1a.

1b.

LL 40–47; transverse� � � � � � C. spinosus�����������������
30–34 � � � � � � � � � � � �
origin 18; total GR
rows between LL and anal-fin
LL 53–62; transverse scale� � � � C. druzhinini�����������������
26–29 � � � � � � � � � � � �

1a.

e is of B. decadactylus)
834: 142 (text only, figur
golden.
(type locality: Saya
Caudal fin with white margin andi Busakhin, 1981: 1728*).
white base, the latter area
Trachichthodes druzhinin
wider in juveniles than adults. Dorsal,
Malha Bank).
anal and paired finsMasuda et al., 1984:: 33; McCulloch, 1929:
unpigmented in young, paired fins becoming
Busakhin, 1982: 9, 14-15*;
Centroberyx druzhinini:
1843a: 49; Günther,1998:1887
yellowor60*.
golden
1994:
i,
11*.
Masuda & Kobayashin adults.
109* ; Kotlyar, 1993: 190;
, 1961: 24; Abe, 1972:
Known only from west of Mossel
Bay (Cape Barracouta),
2*; Smith, 1949: 150*; Krefft
LL continuous,
V I, 7; LL continuous, not
V I, 9–13;
South Africa, north to off Durban. Busakhin’s
P 14–19;
74*;
A IV, 25–32;
12–15; A IV, 15–17; P 13;:
V–VII,
1976
D,(1982)D III–V, 12–15;
Depthstatement,
KrefftDepth
16–23.
1973: 286* 5–7/16–21; pyloric caeca 23–34.
mistakenly citing Fowler (1935 [notn rays, 53–62; GR 5–9/
1936]) as the source, that
extending onto caudal-fi; Woods & Sonoda,
65–70 (69–82 total); GR
lengthbase 15–22; length
the species occurs north of Durban
length dorsal-fin base 34–35;;
29–36; length 1986
:n 410*is probably correct, but
39–42% SL; head 31-36;
29–37% SL;; headcaudalstra, dorsal-fipreanal 50–67; distanceal 42–47; preanal 58–64;
currently unverified by specimens.
: 108*base Heemde24–366 m.
Sayarange:
26–34; predorsal 37–46;
ity: Depth
anal-fin base 28–29; predors
Masuda et al., 1984anal-fin
20-26% HL;
Posterior margin of , 1981: 1728* (type local
base 36-44; snout
23–28% HL; eye 31–34.
khin
n to caudal-fin ochk
snout
Busa
i
&
39–42;
in
fin
hinin
rows
dorsal-fin insertio
s druzar ctenii
Dub
Cheek scale
hode
159;
hicht
989:
Trac
in,
-sized denticles, no ctenii.
wedge-shaped rows of triangul
Ivan
with
uniform73;:
with
Scales1987
scales29-40.
rough
eye
lyar,
but
surface; scales
ped pad on inneretin pelvic axil. No head spines,
disc-sha
scale
ridges;al.,fulcral
into
Small
5.
formednotAdampresent.
). Dorsal-fin origin on vertical
Malha Bankcheeks.
fulcral scale in pelvic axil
le et al., 1995:; 78*;
areas around eyes and on
da et al., 1984:
on D and A bases enlargeddy, 1990: 257*; Sma
set lateralvertical throughfins. Anal-fin origin onBusa
Single pair of head spines
khin, 1982: 9, 14-15*; Masu
through origin of pelvic druzhinin
i: pelvic fins.
Cheek scales in 5–6 rows.
soft
rober
origin on vertical with dorsal
Pectoral fins longer thanCent
fin.yxof dorsal
middleto anterior nostril. Anal-fin
1994: 60*.
i,yashanterior to vertical throughslightly
Koba&
originndaPelvic-fi
Masu
rays 10–11.
: 190;
rest of body silveryar,fin1993
than pelvic finsbright red,
109* ; Kotly
and caudal
n. Pectoral fins longer

Centroberyx druzhinini

pectoral-fin insertio
in adults.

3; LL continuous,
V, 25–32; P 14–19; V I, 9–1
a 23–34. Depth
caec
ric
pylo
21;
/16–
5–7
); GR
15–22; length
base
n
al-fi
dors
th
leng
29–36;
67; distance
50–
nal
prea
46;
37–
34; predorsal 64
4; snout 20-26% HL;n to caudal-fin base 36-4
s of triangular ctenii
with wedge-shaped row
r surface; scales; disc-shaped pad on inne

red, sides
and gill chambers brightHead, iris, back, fins, mouth
70 cm TL.
of body silvery-pink. Attains
Depthof all three major oceans.
Tropical to subpolar waters
range: 25–1240 m.

Order Beryciformes

1609 WIO

64
Minion + Myriad2013.indd

Top of head, backAttains 25 cm.
pink, other fins pink or red.
nes,
recorded from Japan, Philippi
Tropical Indo-West Pacific,
Saya de Malha
western Indian Ocean at
the South China Sea andrange: 100–200 m.
Depth
scar.
Bank and Madaga

ous, notfrom SSF).
17; P 13; V I, 7; LL continu
D V–VII, 12–15; A IV, 15–th62; GR 5–9/ 16–23. Dep
53–
,
rays
n
al-fi
caud
extending onto
base 34–35; length
n
al-fi
dors
th
leng
6;
39–42% SL; head 31-3
64; caudalorsal 42–47; preanal 58–
anal-fin base 28–29; predr margin of
erio
Post
34.
31–
eye
HL;
fin 39–42; snout 23–28%
ek scale rows
d denticles, no ctenii. Che
scales with uniform-size
spines, but roughcale in pelvic axil. No head

PLATE xxby M� Eric Andeson

Body oval, thick anteriorly, compressed
posteriorly with aslender peduncle; head deeper thanlong. Mouth huge, oblique,upper jaw protrusile, Juveniles withsmall depressible teeth;
adult with long well-separated fangs
on jaws; the longest pair at
lower jaw symphysis received into socketsof orbital cavity whenmouth is closed; tongue fills oral cavity
when mouth is closed;
palatines with or without teeth. maxilla
extending well past theeye; eye diameter subequal to greatest
depth of maxilla. Gillrakers long and slender in juveniles, but
short, with 1–3 spinules
in adults. Branchiostegal rays 8 ornine. Dorsal fin single, set
midway between head and tail, its basemore than three times
that of anal fin; caudal fin emarginate,with 17 branched rays;
pectoral fins paddle-shaped, shorter
than head; pelvic finsorigin below rear end of pectoral-finbase; no fin spines except
for procurrent caudal-fin rays at baseof fin. Scales with a spiny
saucer-like plate fused to a short pedicel
connecting to a thinplate embedded in the skin. Lateral line
an open groove crossed
by well-separated bridges of scaly
skin. Vertebrae without
lateral projections; ribs not attachedto centra; neural spinesof abdominal vertebrae depressed. Adults
bathypelagic in 700
to 5000 m; juveniles epipelagic andmesopelagic. One genus,
Anoplogaster Gill with 2 species, one
in WIO.

1

3

Anoplogaster cornuta

5

Fangtooth2017/07/14 11:36 AM7

D 16–20; A 7–9 ; P1 13–16 ; P2 7; BD
1.8–2.2, HL 2.5–2.9 in SL;
eye diameter much smaller than interorbital
width. Juveniles <20 mm SL with a pair of horn-likespines on top of head, andanother pair (one on each preopercle)
extending ventrally.

eye. Body
organ located beneath each
A prominent luminous
pressed. Dorsal fin
com
ally
later
,
orm
fusif
deep or somewhat
and soft-rayed fins.
separate spinous
61
Family Monocentridae
continuous or divided into
5b.
ute, strongly ctenoid.
min
es
Scal
t.
blun
t
Mouth oblique; snou
; none on vomer.
tines
pala
and
jaws
on
ca
s
Teeth small, in band
Monocentris japonid.
Gill-rakers well develope
organ is continuously
Pineapplefish
2a asaki, Japan).
the subocular luminous
from
t
Ligh
(Nag
2
Pl.
1782: 329,
be occluded using
tuyn
can
Hout
and
icus
eria
japon
s
bact
s
Gasterosteu
ar 1986*;
produced by luminoulight. Light is usedtheh SFSA*; Smith 1986*; Kotly
k
Smit
bloc
icus
to
japon
sms
is
hani
centr
Mono
.
rotational or lid-like mecs and to attract*; Michel 1996*; Fricke 1999
ator
1996
pred
ar
use
Kotly
*;
conf
1995
to
al
,
Rand
to see by, to communicate
w ~100 m)*r (belo
wate
deep
in
PLATE xx
liveies
Heemstra & Heemstra 2004
t speces
mos
Order
Beryciform
prey. Adults ofng moonless
duri
feed
to
face
reef
the
during the day, and ascend
biology.)
5–7 / 11–14.
1977, for a review of their
1; P 13–15; LL 12–17; GRnights. (See McCosker,
D IV–VII / 9–12; A 10–1 2a2b 5. Jaws straight
etive behaviour,
secr
l
11+1
urna
e
noct
ebra
Vert
their
HL 1.8–2.4 in SL.
Rarely seen because of
BD 1.3–1.9,
er. Head,ns. Six genera with 7
1
and palatines, none on vom
s occur in all tropical ocea
flashlight fishewith villiform teeth on jaws
naked.
2
malops katoptron,
head
k;
Ano
as
blac
h
in
(suc
y
ined
likel
outl
scalesspecies, 1 in WIO, others
body and fins yellow, the 2b
h central Pacific)
sout
the
to
a
nesi
Indo
widely distributed fromon coral(2007).
es
al.
et
ledg
er
win
und
Bald
by
and
s
wed
d in cave
to be found. Recently revie
A nocturnal species, founniles in3
near wrecks and reefs. Juve
sand
on
also
,
reefs
y
rock
and
r (157 m).
1
been trawled in deeper wate
2
4
shallow water, adults haver
Key to Gen era
h seas. Light organs on lowe
luminous organ
re in roug
asho
wn
4
thro
nally
entral scutes discontinuous;
asio
Occ
17 cm.
ins
D single, continuous; mid-v
Atta
.
1a.
ans)
raisedtace
crus(benthic
preyeyelid-like flap that can be
te
black
a
loca
to
with
eye,
5
used
than
r
are
longe
jaw
blepharon
Mossel� � � � � � � � � � � � � � � Photo
to South Africa (south tothe organ � � � � � � � � � �
covertoIndo-West Pacific: Red Seaa,te spinous and soft-rayed fins;de Malha Bank, Sri Lank
6 s Oman, Saya
1b. D divided into separa
ritiu
Mau
a,
r
Indi
shorte,Bay)light organnes, Taiwan,
mid-ventral scutes continuous;
alops Kner, 1868
nesia, Australia, Philippi
to cover it � � � � � � � Anom
Somalia, Yemen to Indothan eye, without a black flap
and.
Zeal
New
and
n
Japa
a,
Chin
3

4

8

Adult body, head, fins and inside of mouthdark brown to black.
Attains 16 cm SL. Worldwide in tropicaland temperate seas. No
commercial importance.

6

9

3

Family Monocentridae

4

10

7
11

Family Holocentridae
Anoplogaster cornuta: 16 cm SL (off New
Jersey;Woods & Sonoda 1973).

juvenile (Comoros).

5

6

2a Sargocentron diadema: 7 cm SL
18 cm SL (Red Sea; from CFSA).
cm SL (South
1 Sargocentron caudimaculatum:
Myriad2013.indd 61
SL (Chagos). 4 Sargocentron ittodai: 13
Minion
CFSA).+3 Sargocentron inaequalis: 7 cm
Africa; from
(South WIO
SL160912 cm SL (Red Sea).
2b Sargocentron diadema: 15 cm
7 cm SL (Comoros). 6 Sargocentron marisrubri:
cm SL (Solomon
Africa). 5 Sargocentron macrosquamis:
SL (Comoros). 9 Sargocentron praslin: 14
8 Sargocentron microstoma: 11 cm(Seychelles).
SLcm
1111 cm SL (Red Sea).
7 Sargocentron melanospilos:
cm SL (Mauritius). 11 Sargocentron rubrum:Ids). 10 Sargocentron punctatissimum: 10

Centroberyx spinosus: 6cm (South Africa;

1609 WIO Minion + Myriad2013.indd

organ
l scutes discontinuous; luminous
D single, continuous; mid-ventra
raisedeyelid-like flap that can belonger than eye, with a black� � � Photoblepharon�����������������
to cover the organ � � � � �and soft-rayed fins;D divided into separate spinous
shorterorgan; light
mid-ventral scutes continuous
Kner, 1868
to cover it � � � � � � � Anomalops
than eye, without a black flap

2017/07/14 11:36 AM

Fangtooths

anal-finCentroberyx druzhinini: ~14 cm SL (Mauritius;
O. Alvheim, IMR).
Dorsal-fin base longer thanes. Dorsalute spinules. No head spin
min
with
gin
mar
r
erio
post
62
36–
LL
7;
Centrob
rays
eryx
ic
spinosus
pelv
SSF).
soft rays 12–16; Short alfinsino
Africa; from
(South5–7; anal: 24 cmes
splendens
Beryx
spin
fin
ies,
e with spinules. Five spec
bon
l
nasa
of
edge
r
erio
Ant
(totsal).
Centroberyx
Austroberyx
Genu
on scales;base. No ctenii
es Gilchrist, 1903 andlonger thanhichthod
Trac
Dorsal. anal-fin
Dorsal2 fininbaseWIO
spinules. No head spines.
posterior margin with minute
7; LL 36–62 s of Centroberyx.
pelvic rays
12–16;
rays
nym
soft
anal
syno
fin spines 5–7;
1911 are . Five species,
McCulloch,of nasal bone with spinules

1b.

2017/07/14 11:36 AM.indd 58

Genus Centroberyx Gill, 1862base. No ctenii on scales;

1a.

.indd 611609 WIO Minion + Myriad2013

Vietnamy reported from Japan,
museum specimens. Presentl
and the
Ocean around Socotra
and the northwestern Indianrange: 200-900 m.
Error Seamounts. Depth

Key to Gener aby John E� McCosker

1609 WIO Minion + Myriad2013.indd 59

1609 WIO Minion + Myriad2013

th Africa; from SSF).
Beryx splendens: 24 cm (SouBloch & Schneider, 1801

5a.
bacteria at
n containing luminescent
Jaws straight; light orga
WIO.
in
1
ies,
spec
e
Thre
.
chin
front of lower jaw under59

2017/07/14 11:36 AM

2017/07/14 11:36 AM

Anal fin with 4 spines and 12–30 soft rays, each maxilla with2 supramaxillae � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � BERYCIDAE
Anal fin with 2 or 3 spines, 8–12 rays;
one supramaxilla � � � � � � � � � � � � � � � � � � � � � � � � TRACHICHTHYIDAE

5b.

Fishes)Flashlight fishes(157 m).
been trawled in deeper watershallow water, adults haveon lowerin rough seas. Light organs
Occasionally thrown ashore
17 cm.
(benthic crustaceans). Attains
jaw are used to locate prey
Mosselto South Africa (south to
Indo-West Pacific: Red Sea
Lanka,
Saya de Malha Bank, Sri
Bay), India, Mauritius Oman,
Taiwan,ia, Australia, Philippines,
Somalia, Yemen to Indones.
China, Japan and New Zealand5a.

Dorsal fin with 10–13 strong spines � � � � � � � � � � � � � � HOLOCENTRIDAE
Dorsal fin with 3–8 spines � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 5

5a.

Order Osteichthyes (Bony).
off Somalia; from Kotlyar 1993

ns

Pineapple fishes

Pineapple fishesn, Vietnam
esently reported from Japa
tra and the
Soco
nd
arou
an
Oce
ndian
m.
-900
range: 200

noFamily Monocentridae

Family Monocentridae

1 Monocentris japonica: 13 cm (South

Africa; from CFSA). 2 Photoblepharon
steinitzi: 7 cm SL (Comoros; PCH.).
PCH). 4 Beryx splendens: 19 cm SL (Mozambique; PCH).
5 Centroberyx druzhinini: 21 cm

3 Beryx decadactylus: 25 cm FL (Madeira;

8

Family Monocentridae

SL (South Africa; L. Scott).
6 Centroberyx spinosus: 20 cm adult;
4 cm juvenile (South Africa; from CFSA).

65
66

65

67

2017/07/14 11:36 AM

91609 WIO Minion + Myriad2013.indd

2017/07/14 11:36 AM
66

1609 WIO Minion + Myriad2013.indd

67
2017/07/14 11:36 AM

6beneath pectoral fin inse
young.
Top of head, jaws, irisand sides of body silveryInner surface of scales without pad nearposterior margin;
Found in coastal and op
anal fin origin on vertical with insertionof D;
pyloric caeca 12–20 � � Key
the23–26%Atlan� � to� �Wiobase 30–37; snout� � species� � � � � � � � � � � � � � � � � � � � � caudal-fin
southern I
HL; eyetic,
37–46. Scales
B�
withmollis
irregular vertical rows of ctenii formed
into low, parallelridges; disc-shaped pad onoceans.ofDepth
inner surface
7–10
scales; scales range
on D
and A bases enlarged; fulcral
Four pairs of spines
1a Inner surface of scale from Beryx
splendens with ovoid pad near posterior
margin (redrawn from Busakhin 1982).

1b.

cEtORhInIDAE, Basking shark

rtilagenous Fishes

22 Megachasma pelagios, megamouth shark,(Eastern Indian Ocean).

_TEMPLATE_Op

eners_EM1.indd

1.9 m TL juvenile

Family LAMnIDAE, Mackerel sharks

2-3

1 anal spine; pelvic spine
Body deep. D single, continuous;us; LL scales enlarged;
absent; mid-ventral scutes discontinuo
longer than eye, with a
25-30 GR on first arch. Light organ large,
to cover the organ. Two
black eyelid-like flap that can be raised(spelt palpebratus
species, 1 in WIO; the other, P. palpebratum
from central Indonesia
by most authors), is widely distributed
to the central Pacific.

Family Berycidae

Key to species
1a.
1b.

1a.

2a.

223

Inner surface of scales with ovoid, cartilaginous
pad nearposterior margin; anal fin origin before avertical with insertion of D;
pyloric caeca 23–100 � � � � � � � � � � �����������������������

scale present in pelvic axil; cheekon head: 1) above eye; 2) in front of; and 2
scales in 4–5 rows. Head spine lengths
decrease with age. Anal3) below nostrils; 4) at base of lower jaw
by M� Eric Anderson
fin origin on vertical with dorsal softon vertical with spine above
8–9. Pelvic fin origin
eye;pectoral fin insertion. Pelvicrays
beneatheye large, rounded,
SL;
in
3
about
fins may be elongate indorsal
Head
.
soft
rays
compressed
1a Inner surface of scale from Beryx
16–20; LL 56–61 (not countin
Body ovoid,
young.
gscaleslarge, oblique;
on
Mouth
C);
large.
usuallysplendens
Nostrils
steep.
with ovoid pad near posterior
snout short andTop of head, jaws, iris, back and fins
teeth
depth
bands;
in
40–50%
small,
teethSL;
marginJawpyloric
ae.
bright(redrawn
red. Gill cover
caeca
from
supramaxill
74–100 � � � � � � Busakhin
maxilla naked, 2� � � � 1982).
and sides� � B. decadacof body silvery-pink. Attains 60 cm
1b. Inner surface of scales without
tylus
Dorsal fin triangular, often with

Alfonsinos

by a large white spot on dorsal
Anteriormost lateral-line pore preceded� � � � � � � � � � � � � P. palpebratum
corner of opercle; pelvic rays typically 7
pelvic rays 6 � � � � P. steinitzi
White spot on dorsal corner of opercle absent;

Photoblepharon steinitzi
Flashlight fish(notes on the ecology of a “newPhotoblepharon sp. Fridman 1972: 1
species” from the Gulf of Eilat).
Abe & Haned (sic) 1973: 57, Figs. 1-4
Photoblepharon palpebratus steinitzi
1986: 412*.
(type locality, Ras Burka, Red Sea). McCosker,
McCosker & Lagios 1975:
Photoblepharon palpebratus: non Boddaert:
2 (biology of Comoro Island specimens).
Rosenblatt 1987: 161
Photoblepharon steinitzi: McCosker &(steinitzi elevated to specific rank).

V I, 5–6; GR (7–8) +D II–III, 16–18; A I–II, 13–14; P 15–17;
eye 3.4 in head. Body and
21–22). Depth 2.3–2.6; head 1.9–2.1;
10 cm TL.
fins dark brown to grey-black. Attains
Comore, occupies
Known from Red Sea and Isle Grande
and transits to shallow
crevices and caves of deep reef diurnallyduring new moon. Most
water nocturnally to feed, particularly
undiscovered elsewhere
probably more widely distributed but
in western Indian Ocean.

present on vomer and palatines.
with 3–7 spines and 12–
first to third soft ray greatly prolonged,
12–30 soft rays. Pectoral fin
20 rays; anal fin with 3–4 spines andwith 1 spine and 7–13 soft
wedge-shaped, rays 13–19. Pelvic fin
rays. Scales large,
branched
16–17
rays. Caudal fin forked, withon inner surface in Beryx.
thin, ctenoid; fleshy disc often present
single, complete, pored LL
Branchiostegal rays 7–8. Lateral line
(Beryx), or
spinesfew
withscales 36–82. Anterior part of headwell developed, 22–34
spines absent (Centroberyx). Gill rakers
Pyloric caeca 20–100. Gas
on first arch. Vertebrae 10 + 14–15.
bladder present.
break and upper
Medium-sized fishes of the shelf-slope
up in shallower depths,
slope (about 200–600 m), may shoal
eight species. Two species
especially at night. Two genera withtaken commercially in theof Beryx are locally abundant andto fishmeal and oil or
WIO in moderate quantities and reduced
fish in the orient.
sold fresh. More highly prized as food

TL.
Found in coastal and open ocean seamountareas throughout
the Atlantic, southern Indian, westernand central Pacificoceans. Depth range 7–1000 m.

pad near posterior margin;
anal fin origin on vertical with insertionof D;
pyloric caeca 12–20 � � � � � � � � � � � �� � � � � � � � � � � � � � � � � B� mollis

2a.

Four pairs of spines on head: 1) above eye;2) in front of; and3) below nostrils; 4) at base of lower jaw
on vertical with spine above eye;dorsal soft rays 16–20; LL 56–61 (not counting
scales on C);
depth 40–50% SL; pyloric caeca 74–100
� � � � � � � � � � � � B. decadactylus

2a Beryx decadactylus with four
pairs of spines on head.

2b.

2a Beryx decadactylus with four
pairs of spines on head.

2b.

One pair of spines on head in front of eye;dorsal soft rays 13–15; LL 65–70;
depth 29–40% SL; pyloric caeca 23–34
� � � � � � � � � � � � � � � B. splendens

One pair of spines on head in front of eye;dorsal soft rays 13–15; LLBeryx
decadactylus: 26 cm (Madeira; from Woods
65–70;& Sonoda 1973).
depth 29–40% SL; pyloric
Beryxcaeca
23–34
decadactylus� � � � � � � � � � � � � � � B. splendens
Alfonsino
Beryx mollis

Key to Genera

ctenii cover exposed
Dorsal-fin base shorter than anal-fin base;scutes) on belly
surface of scale; no V-shaped scales (ventral
1–4 pairs of spines;
between pelvic fins and anus; head with(usually 3), soft rays 25–30;
dorsal-fin spines 3–5; anal fin spines 3–4
LL 61–82 � � � � � � � � � � Beryx
pectoral-fin rays 14–19; pelvic-fin rays 9–13;
scale ctenii,
nobase;anal-fin
1b. Dorsal-fin base longer thanrow of midventral scutes
posterior margin of scale with spinules only;
without spines;
on belly between pelvic fins and anus; headrays 12–16;
dorsal-fin spines 5–7; anal-fin spines 4, soft
LL 36–62� � � � � � � � � Centroberyx
pectoral-fin rays 13–14; pelvic fin rays 7;

1a.

Beryx decadactylus Cuvier, 1829: 151(type locality: Lisbon, Portugal?);Cuvier, in Cuvier & Valenciennes, 1829:
222.
Beryx decadactylus: Lowe, 1843a: 49;
Günther, 1887: 33; Fowler, 1911: 173;
McCulloch, 1914: 96; Fowler, 1928: 96*;
Maul, 1954: 12*; Krefft, 1961:
24*; Penrith, 1967: 528; Abe and Maruyama,
1971: 1* ; Woods & Sonoda,
1973: 282* Krefft, 1976: 73; Busakhin,
1982: 3* ; Masuda et al., 1984: 109*
Heemstra, 1986: 409*; Maul, 1986: 740*;
Mundy, 1990: 257.
Beryx delphini Valenciennes, in Cuvier& Valenciennes, 1833: 454 (typelocality: SE of Madagascar); Krefft, 1961:
26.
Beryx borealis Düben & Koren, 1846:
33 (type locality: near Bergen,
Norway); Krefft, 1961: 26.
Actinoberyx jugeati Roule, 1923: 1027
(type locality: Morocco?); Roule,
1924: 68*.
Beryx longipinnis Barnard, 1925: 504(type locality: South Africa); Barnard,
1927: 363*; Fowler, 1936: 1268.
Actinoberyx longipinnis: Smith, 1949:
151.

Beryx mollis Abe, 1959: 157* (type locality:

Sagami, Bay, Japan);

Zama &Beryx
Kotlyar,1993:
decada187–190*.
ctylus: 26 cm (MadYasuda, 1979: 149; Busakhin, 1982: 8*;

Beryx decadactylus

Alfonsino

D IV, 12–14; A IV–V, 24–31; P 15–19;
V I, 10–11; LL continuous,
64–77 (total); GR 6–7/ 16–20; pyloric
caeca 12–20. Depth 35–
45% SL; head 31–38; length dorsal-finbase 16–22; length analfin base 28–37; predorsal 45–52; preanal
56–64; distance dorsalfin insertion to caudal-fin base ca. 40-45;
snout 16-25% HL; eye33-47. Scales with rows of triangularctenii and no disc-shaped
pad on inner surface; scales on D and
A bases enlarged; fulcral
scales in pelvic axil well developed.
Cheek scales in 6–7 rows.
Single pair of head spines set lateral
to anterior nostril. Anal
fin origin on vertical with dorsal fin
insertion. Pelvic fin origin
slightly anterior to vertical through pectoral
fin insertion. Pelvic
fin about as long as pectoral fin in adults.

Beryx mollis

Beryx decadactylus Cuvier, 1829: 151

Beryx mollis Abe, 1959: 157* (typ(type locality : Lisbon, Portugal?);surface of scales;
exposed
CuvierYasuda, 1979: 149; Busakhin, 19, ainfleshy,
Head spines present; ctenii coverCuvier & Valenciennes, 1829: 222.
usually
abdomen;onridgeswithno enlarged scales
Plate 8scales; dorsal base shorter
disc-shaped pad on inner surface of Beryx decada
ctylus: Lowe, 1843a: 49; Günther, 1887:
prolonged dorsal-fin ray;33; Fowler, 1911: 173;
than anal base; some specimens with
12-20;
III–V,
D
spinules;withoutbone
McCulloch, 1914: 96; Fowler
anterior edge of nasal
Body and fins pinkish redD
12–14
, 1928:; Ageneral
IV–V, 24–31; P
D III–V,
withIV,
Three species, all in WIO.
silvery
16–20; A96*;
tones on side,
III–IV,Maul,
25–30; P1954:
Krefft,14–18; V12*;
A III, 25–30; V I, 9–13; total LL 61–82.
1961:
I, 9–10;
LL
colourlighter than the other two species ofcontinuous, 56–61 (60–73 total); GR
Beryx; caudal fin
24*; Penrith, 1967: 528; Abe
5–7/16–19; pyloric caeca
lobes darker than body. Attains
64–77
and
74–100.
; GR 6–7/ 16–2
41 cm TL.(total)
DepthMaruyama,
40–50% SL;
Probably widespread
1971:
head 30–38;
1*dorsal-fin; Woods
lengthbase &
in Sonodtropical anda,
temperate Indo-West Pacific. This species
19–24; length anal-fin base 35–36;
was lost
1973: 282* Krefft, 1976: 73;
predorsal length 44–59;
in synonymy with B. splendens
45%20SL;Busakhpreanalhead
length ca.in,1982:
years (Zama
31–38; length d55–60;and Yasuda,; Masud
distance3*
froma et al.,dorsal fin insertion
to 1984:
109* 1982), so thereformay
1979; Busakhin,
be many misidentified
Heemstra, 1986: 409*; Maul, 1986: 740*;
area below the eye isfin base
28–37; predorsal 45–
Photoblepharon steinitzi: ~10cm; the white
Mundy, 1990: 257.
SSF).
from
(Comoros;luminous
Beryx delphini Valenciennes, in Cuvier
fin insertion to caudal-fin bas& Valenciennes, 1833: 454 (typeFamily Monocentridae 63
locality : SE of Madagascar); Krefft, 1961:
33-47. Scales with26.
rows of tri
62 Order Beryciformes
Beryx borealis Düben & Koren, 1846:
33 (type locality : near Bergen,pad on inner surface; scales o
Norway); Krefft, 1961: 26.
scales in pelvic axil well deve
Actinoberyx jugeati Roule, 1923: 1027
(type locality : Morocco?); Roule,
Single pair of head spines se
1924: 68*.
fin origin on vertical with do
Beryx longipinnis Barnard, 1925: 504(type locality : South Africa); Barnard,slightly anteriPLATE
Plate 8orxx to vertical thr
1927: 363*; Fowler, 1936: 1268.
Family OphIchthIDAE, Snake-eels, sand-eels
fin about as long as pectoral fi
Order Beryciformes
Actinoberyx longipinnis: Smith, 1949:
151.
1
Genus Beryx Cuvier, 1829

Family OphIchthIDAE, Snake-eels, sand-eels

1

mus, basking shark (no size or locality).

Class CHON DRC
HTH Y ES – Ca

shark
Family MEGAchASMIDAE, Megamouth

Genus Photoblepharon Weber, 1902

Ord er Beryciformes

1609 WIO Minion + Myriad2013.indd

63

2017/07/14 11:36 AM

2017/07/14 11:36 AM

621609 WIO Minion + Myriad2013.indd

2017/07/14 11:03
AM

Plate 7

Family MEGAchASMIDAE, Megamouth

Family cEtORhInIDAE, Basking shark

shark

3

4

2
1

1 Cetorhinus maximus, basking shark (no2 Megachasma pelagios, megamouth shark,(Eastern Indian Ocean).

size or locality).

1.9 m TL juvenile

Apterichtus flavicaudus, yellowtail sand-eel,
80 cm TL (Hawaii; JER).

Dcrocodilinus,
III–V, 16–20; A III–IV
Brachysomophis
crocodilesnake-, 25–30; P 14–18; 2V I, 9–10; LL
continuous,
56–61 (60–73 total); GR 5–7/16–19;
eel, 44 cm TL (Rodrigues;
PCH).
pyloric caeca
74–100. Depth 40–50% SL; head 30–38; length dorsal-fin base
19–24; length anal-fin base 35–36; predorsal length 44–59;
Brachysomophis cirrocheilos, smudged snakeBrachysomophis crocodilinus,
snakepreancrocodile
al length
ca. 55–60; distance from dorsal fin
eel, 125 cm TL (Indonesia; JER).
eel, 44 cm TL (Rodrigues; PCH).
insertion to

Brachysomophis cirrocheilos, smudged snakeeel, 125 cm TL (Indonesia; JER).

Family LAMnIDAE, Mackerel sharks

1

5

Apterichtus flavicaudus, yellowtail sand-eel,
80 cm TL (Hawaii; JER).

6
3

2

4

Body and fins pinkish red wi
colour lighter than the otherlobes darker than body. Attainin tropical and temperate Indoin synonymy with B. splendens
1979; Busakhin, 1982), so th

5
6

mako,
tra 2004). 4 Isurus oxyrinchus, shortfin
male (South Africa; from Heemstra & Heems
6 Lamna nasus, porbeagle,).
archarias, white shark, ~4 m TL mature(Japan
TL
m
2.3
mako,
n
mako,
n
shortfilongfioxyrinchus,,
Isurus4
Heemstra 2004).
IsurusHeemstra &paucus
fromtramale2004).
(South Africa;5& Heems
porbeagle,~4 m TL mature
white shark,
Heems
carcharias, tra
3 Carcharodonn mako, 2.3 m TL (Japan). 6 Lamna nasus,le (South Africa; from& Heemstra 2004). 5 Isurus paucus, longfi~2.5 m TL female (South Africa; from Heemstra
immature male (NE Atlantic).
ic).
94 cm TL
ture male (NE Atlant
Family SphyRnIDAE, Hammerhead sharkssBrachysomophis henshawi, reptilian snake-eel,
78 cm TL (Oman; JER).

Family SphyRnIDAE, Hammerhead shark

4

3

Brachysomophis henshawi, reptilian snake-eel,
78 cm TL (Oman; JER).

Callechelys catostoma, striped snake-eel,
54 cm TL (Marshall Ids; JER).

Callechelys marmorata, marbled snake-eel(Red Sea; SV Bogorodsky).

4

3 1609 WIO Minion + Myriad2013.indd 63

Callechelys catostoma, striped snake-eel,
54 cm TL (Marshall Ids; JER).

Callechelys marmorata, marbled snake-eel(Red Sea; SV Bogorodsky).

6

5

8

Callechelys marmorata,marbled snake-eel,
32 cm TL (Maldives; JER).

Cirrhimuraena playfairii, fringelip snake-eel,
24 cm TL (South Africa; JER).

Lamnostoma orientalis, oriental snake-eel,
21 cm TL (Indonesia; JER).

Leiuranus semicinctus, halfbanded snake-eel,
42 cm TL (Maldives; JER).

7

Ichthyapus vulturis, vulture snake-eel,
21 cm TL (Hawaiian Ids; JER).

10

9

8

(Mozambique).
hammerhead, 47 cm TL immature female
mokarran,m TL (N Australia). 8 Sphyrna lewini, scalloped
7 Eusphyra blochii, winghead shark, 1.8
from Heemstra & Heemstra 2004). 10 Sphyrna, ~70 cm TL immature female (South Africa;
9 Sphyrna zygaena, smooth hammerhead
great hammerhead, 2.4 m TL (Red Sea).

58

8

7

7

Muraenichthys schultzei, bluntnose worm-eel,
9 cm TL (Eritrea; SV Bogorodsky).

7

5

10

Holocentridae

Familyacanthorhinus: 18 cm (Gulf of Aqaba;
1-12-09 (Mozambique; R. Bills). 2 Ostorhinchus
1 Neoniphon sammara: M43 mangrove
seychellense: 15 cm SL (Mauritius; O. Alvheim,n: 21 cm (Red Sea; JER). 4 Sargocentron
D. Golani). 3 Ostorhynchus hypsipterygio
tiere: 22 cm (Maldives; JER).
30 cm (Marshall Ids; JER). 6 Sargocentron
spiniferum:
Sargocentron
5E).
IMR/ASCLM(Maldives; JER).
JER). 8 Sargocentron violaceum: 34 cm
Sargocentron tiereoides: 15 cm (Maldives;

Callechelys marmorata,marbled snake-eel,
32 cm TL (Maldives; JER).

Cirrhimuraena playfairii, fringelip snake-eel,
24 cm TL (South Africa; JER).

Ichthyapus vulturis, vulture snake-eel,
21 cm TL (Hawaiian Ids; JER).

6

69

57

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69

Plate 5

Family SynGnAthIDAE, Seahorses, pipefishes
brown spots andHead and body brownish with darker
X-shaped (some doubled)
mottling; mid-side of body with fiverows of dark spots
dark brown blotches; 1st dorsal fin with
fin with short verticalforming indistinct bands; base of caudal
with posteriormost
dark bar or crescent, which may coalesce
mm SL.
mid-side X-shaped blotch. Attains 95part)*; Smith & Smith 1963
Glossogobius giuris: Smith 1959b (inHoda 1980*; SSF No. 240.44*;(in part); Hoese & Winterbottom 1979;
Maugé 1986a; Keith et al. 1999*.

7–9 rays; pectoral fins
Second dorsal fin 8 or 9 rays; anal fin
scales 14–25,
17–23 rays; LS 29–35; TRB 8½–11. Predorsal
Opercle with patch ofreaching forward to or just behind eyes.
or with small patch of
cycloid scales dorsally; preopercle naked
1st dorsal fin may
adults,In
adults.
large
in
eye
behindscalesnot greatly so.
have at least 2nd spine filamentous, but

species (only type
Distribution WIO: a poorly known
from Mauritius
specimens examined), recorded only
(20 syntypes).
a different species.
are
Remarks Two syntypes from India

brown, with 4–7
Head and body light brown to yellowish
along side androunded to elongate dark brown blotches
blotch on caudal-fin
large dark brown triangular to elongate
fin with distinct
dorsal
1st
fin;
onto
base, partly extending
spine, specimens
1st
of
base
near
spot
dark brown to blackish
band across tips of 1st
about 55 mm SL or smaller with blackof fin, black pigment
top
over
dorsal-fin spines or black blotch
cm SL.
35
Attainsremains on spines only in adults.

Glossogobius tenuiformis (Fowler 1934)

496–497, Fig. 49 (St Lucia Lake, 32
Glossogobius tenuiformis Fowler 1934b:al, South Africa).
km from mouth, northern KwaZulu-Natpart).
Glossogobius giuris: Hoese 1986b (in(in part)*.
Glossogobius callidus: Whitfield 1998

pectoral fins 18–21
Second dorsal fin 9 rays; anal fin 8 rays;
scales cycloid, 11–16,
rays; LS 28–30; TRB 9–10. Predorsal
preopercular margin.
reaching above, or just in front of, rear
fin with 2nd spine
Cheek and opercle naked. First dorsal
filamentous.

Family MOnAcAnthIDAE

Glossogobius kokius (Valenciennes 1837)
Mauritius flathead goby

Val. 1837: 68 (Mauritius, Malabar
Gobius kokius Valenciennes in Cuv. &and Pondicherry, India).
Val., 1837: 78 (Mauritius);
Gobius filosus Valenciennes in Cuv. &Smith 1959b.

Glossogobius kokius: Akihito & Meguro

PLATE 13Natal flathead goby

estuaries and freshwater
Remarks Widely distributed, enters
Several species are likely
river systems, extending well inland.
confused under this name.

from SSF).

Hippocampus cameleopardalis,
giraffe seahorse, 52 mm SL(S Mozambique; PCH).

Hippocampus capensis, Knysna
seahorse (South Africa;
J Lockyear).

Hippocampus debelius, softcoralseahorse, female (Red Sea;
S Kahlbrock).

Hippocampus debelius, softcoralseahorse, male (Red Sea;
S Kahlbrock).

Hippocampus fuscus, drab
seahorse, 55 mm TL (Red Sea;
SV Bogorodsky).

Hippocampus jayakari, spiny
seahorse, 73 mm TL (Gulf of
Aqaba; JER).

Hippocampus kuda, yellow
seahorse, 137 mm TL(Mozambique; O Alvheim, IMR).

Hippocampus natalensis, cryptic
seahorse (S Mozambique;
V Fraser).

Hippocampus suezensis,
Egyptian seahorse, 35 mm SL(Persian Gulf; JER).

Hippocampus tyro, 61 mm TL
female holotype (Seychelles;
JER).

Hippocampus sp. 1, African
prickly seahorse (South Africa;
G de Valle).

Hippocampus sp. 3, Sodwana
puff seahorse (South Africa;
N Ayliffe).

Hippocampus sp. 3, Sodwana puff
seahorse (South Africa;
C van Jaarsveld).

Kyonemichthys rumengani,pygmy pipedragon (Red Sea;
SV Bogorodsky).

Lissocampus bannwarthi, Suez
pipefish, 134 mm SL (Egypt;
JER).

Glossogobius kokius, syntype (Mauritius).

African coast to South
Distribution Indo-West Pacific: East
Madagascar, Pakistan,
Africa (Umtata River mouth), Aldabra,
to northern Australia.
Red Sea, Réunion and Seychelles to Japan

Glossogobius giuris, 23 cm SL (South Africa;

Hippocampus borboniensis,
Réunion seahorse, 60 mm TL
male (Seychelles; JER).

1975*; Maugé 1986a; Fricke 1999.

8 or 9 rays; pectoral fins
Second dorsal fin 9–10 rays; anal fin
scales cycloid,
17–20 rays; LS 24–33; TRB 9–10. Predorsaledge of preopercle.
14–17, reaching to just in front of rear
very small patch of
with
or
naked
Cheek naked; opercle
spine longest,dorsal-fin
Secondlly.
cycloid scales anterodorsa
filamentous in large males.

brown, with 4–8
Head and body yellowish brown to lightvertically elongate(usually 4) pairs of small dark brownspots interspersed,
blotches along side, may be smaller brownbase and two similarand small dark brown blotch at caudal
form Y-shaped blotch; 1stspots on base of fin, may coalesce todistally; irregular dusky
dorsal fin without black blotch or bandspot at base of 1st spine.
band along base which may form dark
and body covered in fine
When alive, dorsolateral part of headSL.
orange to red speckles. Attains 74 mm, Filefishes

Africa (Zotsha River
Distribution WIO: endemic to Southmouth to Richards Bay, KwaZulu-Natal).
Has been
Remarks Found in estuaries and freshwaters.
in South African riverconfused with G. callidus and G. giuris
species has been collected
systems. A specimen resembling thisfrom Réunion.

2

1

100
FAMILY Gobiidae

Plate 31

Order Perciformes

61

3

Family MOnAcAnthIDA

4

Plate 15

Plate 31

E, Filefishes

Family GOBIIDAE, Gobies

5
2

2

1
3
3

1

6a
4
4

6

6b

5
5
7

6a

9
8

6b

7

10

7

12
11a

8

11b

9

8

9

15
13

10

14
11

16

18

12

10

17

14
20

13
19

11

1 Paramonocanthus arabicus,
Gulf filefish, 7 cm SL (Oman).
2 Paramonacanthus frenatus,
3 Paramonacanthus nematophwedgetail filefish, 6 cm SL
orus, seagrass filefish, 9 cm
(S Mozambique)
SL (Red Sea). 4 Paramonocanthus
5 Pervagor aspricaudus, orangeta
pusillus, blackstriped filefish
il filefish, 9 cm SL male (Mauritiu(Mozambique).
(b) 5 cm SL (S Mozambique).
s). 6 Pervagor janthinosoma,
7 Pervagor randalli, Randall’s
redtail filefish, (a) 8 cm SL(Mauritius),
filefish, 4 cm SL (Red Sea).
9 Stephanolepis diaspros, reticulate
8 Stephanolepis auratus, porky,d filefish, ~11 cm SL (Oman).
19 cm SL (South Africa).
11 Thamnaconus erythraee
10 Thamnaconus arenaceu
nsis, Aqaba filefish, 19 cm
s,
sandy
fi
lefi
sh, 31 cm TL (South Africa).
SL (Red Sea). 12 Thamnacon
13 Thamnaconus melanopr
us fajardoi, spotted filefish,
octes, Oman filefish, ~19 cm
16 cm TL (Mauritius).
TL (Oman). 14 Thamnacon
us modestoides, drab filefish,
32 cm SL (Kenya).

12

307

13

(Vietnam).
goby, 23 mm SL male paratype
pygmy
annosum, greybeard pygmy
Trimma bisella, twosaddle
goby (Maldives). 2 Trimma
pygmy goby (Red Sea). 5
1 Trimma anaima, pallid pygmy
4 Trimma barralli, Barrall’s
ila, hotlips pygmy goby,
Africa). 7 Trimma daleroche
pygmy goby, 19 mm SL (Yemen).
3 Trimma avidori, Avidor’s
pygmy goby, 20 mm SL (Southsus, longspine pygmy goby
polkadotm,filamento
corallinu
Trimma9s). 6 Trimma
Caledonia).
goby, 24 mm SL (Mauritiu
goby (a) 24 mm SL,
pygmy goby, 17 mm SL (NewpygmylEmery’s
yellowtai
emeryi,atum,
Trimma8flavicaud.
17 mm SL paratype (Chagos)
mm SL (Red Sea). 11 Trimma
Griffith’s pygmy goby,, Fishelson’s pygmy goby, 27
Africa). 13 Trimma griffithsi,
goby(Red Sea). 10 Trimma fishelsoni
pygmy goby, 20 mm SL (South
halonevum, pimple pygmy
12 Trimma flammeum, flame
SL (Rodrigues). 15 Trimma(b) 24 mm SL (both Red Sea).
cutface pygmy goby, 21 mm
Mendelssohn’s pygmy goby,(Chagos). 14 Trimma haima,. 17 Trimma mendelssohni,
17 mm SL female paratype
SL female holotype (Chagos)
crescentwing pygmy goby
Hoese’s pygmy goby, 21 mms). 19 Trimma omanensis,
hoesei,(ComoroTrimma
male
16
SL
s).
mm
27
(Maldivegoby,
Trimma naudei, Naude’s pygmy
22 mm SL (Rodrigues). 18
Sea).
(Red
i, Sheppard’s pygmy goby
(Oman). 20 Trimma sheppard

59

14