FAO SPECIES CATALOGUE

FIR/S125 Vol. 7, Part 1
FAO Fisheries S yn o p s is No. 125, Volum e 7, Part 1
FAO SPECIES CATALOGUE
VOL. 7. CLUPEOID FISHES OF THE WORLD
(suborder CLUPEOIDEI)
AN ANNOTATED AND ILLUSTRATED CATALOGUE
OF THE HERRINGS, SARDINES, PILCHARDS, SPRATS, SHADS
ANCHOVIES AND WOLF-HERRINGS
Part I - Chirocentridae, Clupeidae and Pristigasteridae
í*
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winpii»»hiiaiitfiÿ’nffy
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UNITED NATIONS DEVELOPMENT PROGRAMME
FOOD AND AGRICULTURE ORGANIZATION OF THE UNITED NATIONS
FAQ F is h e r ie s S y n o p s is No. 1 2 5 , Volume 7, Part 1
FAO
VOL. 7
SPECIES
CLUPEOID
(Suborder
F IR /S125 V ol. 7 , P art 1
CATALOGUE
FISHES OF THE WORLD
CLUPEOIDEI)
An A n n o ta te d an d Illustrated C a ta lo g u e o f th e
H errin gs, S a r d in e s , P ilc h a r d s, S p r a ts , S h a d s , A n c h o v ie s an d W olf-h errings
P art 1 - C h iro c en tr id a e, C lu p e id a e a n d P r is tig a s te r id a e
prepared by
P e te r J .P . W h iteh ea d
B ritish M u seu m (N atural H isto ry )
Crom well R oad, L ondon SW7 5BD, UK
UNITED NATIONS DEVELOPMENT PROGRAMME
FOO D AND AGRICULTURE ORGANIZATION OF THE UNITED NATIONS
R om e, 1 9 8 5
The d e sig n a tio n s em p lo y ed an d the p resen tatio n
of m a terial in th is p u b lic a tio n do n o t im ply the
e x p ressio n o f any o p in io n w h atso ev er on the
p art o f the F ood and A gricu ltu re O rganization
o f th e U n ite d N a tio n s c o n c e rn in g th e le g a l
status o f any country, territo ry , city or area or
o f its au th o rities, or co n cern in g the d elim itatio n
o f its frontiers or boundaries.
M -4 3
IS B N 9 2 -5 -1 0 2 3 4 0 -9
All
rights
reserved.
No
p a rt o f
this
p u b licatio n
m ay
be
stored in a retriev al system , or tran sm itted in any form or by
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A p p licatio n s
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w ith a statem e n t o f the p u rp o se and ex ten t o f the rep ro d u ctio n , sh o u ld
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O rganization o f the U nited N ations, V ia delle Term e di C aracalla, 00100
R om e
Italy.
Ill
P R E P A R A T IO N
OF THIS DOCUM ENT
The present publication w as prepared under the FA O R egular Program m e, w ith the support o f the United
N atio n s D evelopm en t P ro g ram m e, as p art o f the U N D P /F A O
G lobal P ro je c t onSurvey and Identification o f the
W o rld ’s M arine Fishery R esources (GLO/82/001).
It is the seventh w orldw ide species catalogue in the FA O
Fisheries Synopses series.
The author is the forem ost specialist in clupeoid taxonom y and has also had considerable experience in the
field, especially w ith regard to clupeoid fisheries in tropical and subtropical areas. He has provided the clupeoid
fam ilies for the FA O identification sheets for all the areas so far published (except the M editerranean, Fishing
A rea 37).
A ll illustrations signed TW w ere kindly provided by D r Thosaporn W ongratana (Chulalongkorn U niversity,
Bangkok)! the rem ainder were redraw n (and m odified) at FA O from the literature or from the author’s sketches,
under the supervision o f the author and the editor.
The index o f scientific and vernacular names was prepared in collaboration w ith FA O ’s Fishery Inform ation,
Data and Statistics Service.
In producing these catalogues and other taxonom ic works w ithin the Global Project, FA O is very much aware
that the participation o f specialists depends not only on their willing cooperation, but also on the moral and other
support that they them selves receive for such w ork from their own institutions. In the present case the support of
the B ritish M useum (N atural H istory) is g ratefu lly acknow ledged.
T echnical Editor:
D r W alter Fischer, Fishery R esources and Environm ent D ivision, FAO
Illu strato rs:
0. Lidonnici and P. L astrico, FAO, Rome. A ll drawings initialled “T.W .” were
draw n from specimens by D r Thosaporn W ongratana, Chulalongkorn U niversity,
Bangkok, Thailand
A BSTRACT
This is the seventh in the FA O series o f w orldw ide illustrated catalogues
(synopses) o f m ajo r groups o f o rg an ism s th a t en ter m arine fish eries.
The p resen t
c a ta lo g u e includes all clu p e o id fishes (suborder C L U P E O I D E I - h e rrin g s , sa rd in e s,
pilchards, sprats, shads, anchovies and w olf-herrings), com prising 4 fam ilies, 80 genera
and over 300 species, contributing to more than a quarter o f the w orld fish catch. K eys
are given to the fam ilies, subfamilies and genera, w ith a glossary o f technical term s and
m easurem ents, and a review o f m ajor literature sources w ithin each FAO fishing area.
G enera and species are d iag n o sed, w ith d raw in g s, scien tific and v ern acu lar nam es,
inform ation on habitat, biology, fisheries, and distribution (w ith map): reference is also
m ade to subspecies and to any tax o n om ic or oth er problem s. The o ccu rren ce ot the
sp e c ie s w ith in th e F A O fish in g areas is ta b u la te d .
A ll scien tific nam es a p p lie d to
clupeoid species are given in the synonym ies, and these as well as the vernacular names
are indexed: there is also a complete bibliography o f a ll literature cited.
D istribution:
A uthor
FA O F isheries D ep artm en t
FAO Regional Fisheries Officers
Regional Fisheries Councils
and Com m issions
Selector SM
F or b ib lio g rap h ic purposes th is docum ent
be cited as follows:
W h iteh ead ,
1985
should
P .J.P .,
F A O s p e c i e s c a ta lo g u e .
Vo 1.7. Clupeoid fishes o f the world.
An a n n o ta te d and illu s tra te d c a ta ­
logue o f the h errin g s, sard in es, p il­
ch ard s, sp rats, an ch o v ies and w olfh errin g s. P a rt 1 - C h ir o c e n t r i d a e ,
C lupeidae and P ristig asterid ae. FA O
Fish.Synop.. (1 2 5 )V o l .7, P t.1:303 p.
click fo r n e x t p a g e
click fo r p re v io u s p a g e
iv
TABLE
OF C O N T E N T S
Code
INTRODUCTION
1.1
................................................................................
C lu p e o id s in World F is h e r i e s
.
1
■ ■
3
c a t c h e s ...............................................................
■ ■
3
.
.
9
.
.
10
Clupeoid
1.1.2
Fishing
1. 1. 3
U tiliza tion o f c l u p e o i d s ..............................................
the
.
..............................................
1.1.1
of
Page
g e a r ....................................................................
1 .2
Plan
1 .3
P ro b lem s
1 .4
T e c h n ic a l T e r m s, M e a s u r e m e n t s a n d C o u n ts .......................
.
1 .5
L i t e r a t u r e .................................................................................................
■ ■
14
1 .6
A c r o n y m s .................................................................................................
.
.
18
.
.
19
of
C a t a l o g u e .........................................................
Id en tifica tio n
ORDER CLUPEIFORMES
2 .1
SUBORDER CLUPEOIDEI
.
.
■ ■ 11
.
C h i r o c e n t r i d a e ...............................................................
.
.
. CHIROC
C h i r o c e n t r u s .....................................................................................
.
.
. CHIROC Chiroc
................................................................................
................................................................................
.
.
. CHIROC
...........................................................................
.
F a m il y
C.
C.
2 .2
....................................................
F a m il y
dorab
nudus
C lu p eid a e
. . .
.
.
.
.
.
,.
.
................................................................
.
.
D . acu ta
........................................................................
D. e l o p s o i d e s ........................................................................
.
.
E t r u m e u s ......................................................................................
.
12
.
22
.
.
.
.
23
CHIROC Chiroc 2 .
.
.
.
.
.
23
24
. C L U P .............................
.
.
25
.
.
26
Chiroc
1
2.2.1 Subfam ily D u s s u m i e r i i n a e ....................................................
Dussumieria
.
.
.
.
.
.
.
28
.
.
. CLUP Duss 1 .
. CLUP Duss 2 .
.
.
.
.
.
.
.
.
.
.
28
29
.
.
. CLUP Etm
.
.
...
E. teres
...............................................................................
E. w h i t e h e a d i ........................................................................
.
.
.
.
. CLUP Etru 1 . . . .,
. CLUP Etru 2 ■ ■ ■ .,
.
.
. 30
. 32
S p r a t e l l o i d e s ...............................................................................
.
.
. C L U P Spratel .
.
.
.
.
.
.
.
.
. CLUP
.C L U P
.C L U P
.CLUP
...............................................................................
.
.
.CLUP Jenk
l a m p r o t a e n i a .................................................................
m a j u a ...............................................................................
p a r v u l a ...............................................................................
s t o l i f e r a ...............................................................................
.
.
.
.
.
.
.
.
.CLUP
.CLUP
.CLUP
.CLUP
S.
S.
S.
S.
d e l i c a t u l u s ........................................................................
g r a c i l i s ...............................................................................
l e w i s i ...............................................................................
r o b u s t u s ........................................................................
Jenkinsia
J.
J.
J.
J.
2 .2 .2
Subfamily
C l u p e i n a e ..........................................................
. CLUP Duss
.
.
.
.
.
.
33
1
2
3
4
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
33
34
35
36
.
.
.
.
.
37
. . .
. . .
. . .
. . .
.
.
.
.
.
.
.
.
.
.
.
.
37
39
40
41
Spratel
Spratel
Spratel
Spratel
Jenk
Jenk
Jenk
Jenk
.
1
2
3
4
30
.
. 42
V
Page
C ode
S p rattu s
S*
£L
£L
S*
ÍL
C LU P
an tipodum .
fu eg en sis
m uelleri
n o v aeh o llan d iae
sprattus .
CLU P S pratt 4 .
C LU P S p ratt 2 .
C LU P S p ratt 5 .
C LU P S p ratt 3 .
C LU P S p ratt 1 ■
C lu p eo n ella
CLUP
C.
Ç
C LU P
C LU P
C LU P
C LU P
abrau
c u ltriv en tris
IL e n g rau lifo rm is
IL grim m i .
S a r d i n a .............................
£L p ilch ard u s .
S ard in op s
¿L
IL
IL
IL
S.
H aren g u la
IL
IL
IL
IL
.
caeru leu s
m ela n o stic tu s
n eo p ilch ard u s
o c e lla tu s .. ■
sagax
O.
IL
O.
IL
O.
clu p eo la
h u m eralis
ja g u a n a
th rissin a
.
.
A - c lu p e o id e s .
A - le io g a s te r .
A . sirm .
.
.
£L alb ella .
£L a tricau d a
S . au rita
.
.
45
.
.
.
.
■
.
.
.
.
■
45
46
47
48
49
.
. 51
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
51
52
53
54
.
.
.
.
.
.
55
C LU P Sardi 1 .
.
.
.
.
.
55
C LU P S ardop
.
.
. . .
57
.
.
.
.
.
57
58
59
60
61
C LU P S ardi
C LU P
C LU P
C LU P
C LU P
C LU P
C LU P
C LU P
C LU P
C LU P
C LU P
.
.
.
.
.
.
S ardop
S ardop
S ardop
S ardop
S ardop
H ar
H ar
H ar
H ar
O pi
O pi
O pi
O pi
O pi
.
4
5
1
2
3
. .
. .
. .
. .
. .
1
3
2
4
.
.
.
.
.
.
.
.
.
.
.
.
C LU P O pi
b lackburni
casteln au i
d isp ilo n o tu s .
gotoi .
k o n in g sb erg eri
lossei ■
p u n ctatu s .
q u ad rim acu latu s
sp ilu ru s
.
Species A .
S pecies. B .
S pecies. C.
S ard in ella
C lu p n la
C lu p n la
C lu p n la
C lu p n la
■
.
1
2
3
4
C LU P
C LU P
C LU P
CLUP
.
b erlangai
b ulleri .
lib e rta te .
m ed irastre .
og lin u m .
A m b ly g aster
C lu p n l a
C LU P H ar
H e rk lo tsic h th y s
H.
IL
IL
H.
IL
IL
IL
IL
IL
IL
H.
IL
.
■
.
O p isth o n em a
S prat
.
.
.
.
.
.
.
.
.
.
.
.
. . .
. . .
. . .
. . .
63
64
65
66
. . .
67
.
.
.
.
.
68
69
70
71
72
. . . .
2
3
4
5
1
.
.
.
.
.
.
.
.
.
.
. 63
.
.
.
.
.
. .
. .
. .
. .
. .
C LU P H erk . . . .
. . .
73
C LU P
C LU P
C LU P
CLUP
C LU P
C LU P
C LU P
C LU P
C LU P
C LU P
C LU P
C LU P
H erk
H erk
H erk
H e rk
H erk
H erk
H erk
H erk
H erk
H erk
H erk
H erk
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
. .
.
.
.
.
.
.
.
.
.
.
.
.
74
15
76
11
78
79
80
81
82
83
84
85
CLUP
Ambl
6 . . .
7 . . .
5 . . .
8 . . .
9 . . .
3 . . .
1 . . .
2 . . .
4 . . .
10. . .
11. . .
12. . .
.
.
.
.
.
.
.
.
.
.
.
86
C LU P A m b l 1 .
C LU P A m bl 2 .
C LU P A m bl 3 .
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
86
87
88
CLUP Sardi . . . .
C LU P Sardi 6 .
C L U P Sardi 16
C LU P Sardi 1 .
.
.
.
.
.
.
. . .
90
. . .
. . .
. . .
91
92
93
C ode
S.
S.
S.
S.
S.
S.
S.
S.
S.
S.
S.
S.
S.
S.
S.
S.
S.
S.
b r a c h y s o m a ...........................................
.................................... CLUP
b r a s i l i e n s i s ...........................................
f i j i e n s e ..................................................
f i m b r i a t a ..................................................
.................................... CLUP
g i b b o s a ..................................................
h u a l i e n s i s ..................................................
j u s s i e u i ..................................................
lem u ru
...........................................
l o n g i c e p s ..................................................
m a d e r e n s i s ...........................................
m a r q u e s e n s i s ........................................... ■ .................................... CLUP
m e l a n u r a .................................................. ■ .................................... CLUP
n e g l e c t a ...........................................
........................................... CLUP
r ic h a rd s o n i...........................................
........................................... CLUP
r o u x i ..................................................
........................................... CLUP
s i n d e n s i s .................................................. ■ .................................... CLUP
t a w i l i s .................................................................................................... CLUP
z u n a s i ..................................................
........................................... CLUP
C lupea ...................................................................................................................CLUP
P age
Sardi
Sardi
Sardi
Sardi
Sardi
Sardi
Sardi
Sardi
Sardi
Sardi
Sardi
Sardi
Sardi
Sardi
Sardi
Sardi
Sardi
Sardi
■
5 ■
11■ ■.
17.
.
..
7 ■ ■ .
8 ■ ■ .
18
■ .
19
■ .
.
14
.
3 ■
.
2 ■
20
■ .
4■ .
15
■ .
22
■ .
12
■ .
13 ■ .
23
■ .
.
24
Clup
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
. 95
. 96
. 97
. 98
. 100
. 101
. 102
.103
. 104
. 106
. 107
. 108
. 109
.1 1 0
-Ill
. 112
.1 1 3
.113
.
.
.1 1 5
.
.
.
.
.1 1 5
.1 1 7
C. h a r e n g u s ...........................................
C. p a l l a s i i ...........................................
........................................... CLUP
........................................... CLUP
C lup
Clup
E s c u a l o s a ..........................................................
........................................... CLUP
Esc ■
.
.
.1 1 8
Esc 2 .
Esc 1 .
.
.
.
.
. 118
.1 1 9
F.. e l o n g a t a .................................................. ........................................... CLUP
E. t h o r a c a t a .................................................. ........................................... CLUP
P la t a n ic h t h y s ......................................................... ........................................... CLUP
P. p la t a n a ......................................................... ........................................... CLUP
1
2
.
.
.
.
. 121
1 .
.
.
.
. 121
P lat .
P la t
.
.
R a m n o g a s t e r ......................................................... ........................................... CLU P
R am no .
.
.
. 123
R. a r c u a t a ...................................................
........................................... C LU P
R. m e l a n o s t o m a ........................................... ........................................... CLU P
Ranino 1
R am no 2
■
.
■
.
■123
. 124
Rhino
.
.
. 125
Rhino 1.
Rhino 2
.
■
.
■
. 126
■ 127
L i l e ....................................................................... ........................................... CLUP
Lile ■
■
■
■ 128
T,. p i q u i t i n g a .................................................. ........................................... CLUP
L. s t o l i f e r a ...........................................
........................................... CLUP
Lile 1
L ile 2
■
.
.
.
.
.
. 128
.1 2 9
S t r a n g o m e r a ......................................................... ........................................... CLUP
Strang
■
.
.
. 130
Strang 1
.
.
.1 3 0
R h in o s a r d in ia ......................................................... ........................................... CLUP
R. a m a z o n ic a .................................................. ........................................... CLUP
R. b a h i e n s i s .................................................. ........................................... CLUP
S. b e n t i n c k i .................................................. ........................................... CLUP
.
2.2.3 S ubfam ily P e l l o n u l i n a e .......................................................................................................................................... 132
C y n o t h r i s s a ...................................................
........................................... CLUP
Cyno
.
. 136
C. a n s o r g i i ...........................................
C. m ento
...........................................
C. S p e c i e s ...........................................
........................................... CLUP
........................................... CLUP
........................................... CLUP
Cyno 1
Cyno 2
C y n o sp. .
.
.
.
. 136
. 137
. 138
O d a x o t h r i s s a ...................................................
........................................... C LU P
O daxo .
.
. 139
O. l o s e r a ..................................................
O. v i t t a t a ...................................................
........................................... C LU P
........................................... C LU P
O daxo 1
O daxo 2
.
.
. 139
. 140
P e l l o n u l a ..........................................................
........................................... CLUP
P ellón ■
.
. 142
P. l e o n e n s i s ...........................................
P. v o rax
...........................................
........................................... CLUP
........................................... CLUP
P ellón 1
P ellón 2
.
.
. 142
. 144
.
C ode
P age
N a n n o t h r i s s a ..........................................................
............................. C LU P N an n ■
■
■
.
.
145
N. parva
..........................................................
N . s t e w a r t i ..........................................................
............................. C LU P N an n 1
............................. C LU P N an n 2
■
■
■
■
.
.
.
.
145
146
P o e c i l o t h r i s s a ..........................................................
.............................C LU P P o e c ■
■
■
.
.
147
P. c e n tr a li s ..........................................................
P. c o n s i c a ..........................................................
P. m o e r u e n s i s ..................................................
............................. C LU P P o e c 1
............................. C LU P P o e c 2
............................. C LU P P o e c 5
.
■
■
.
■
■
.
.
.
.
.
.
148
149
150
M ic r o th r is s a .................................................................
............................. C LU P M icro ■
■
■
.
.
151
M. m i n u t a ..........................................................
M . r o v a u x i ..........................................................
M. S pecies A ..................................................
............................. C LU P M icro 1
............................. C LU P M icro 2
............................. C LU P M icro 3
■
■
■
■
■
■
.
.
.
.
.
.
151
152
153
.
.
154
.
.
.
155
156
157
.
158
P o t a m o t h r i s s a ..........................................................
P. a c u t i r o s t r i s ..................................................
P. o b tu s ir o s tr is ..................................................
P. w h i t e h e a d i ..................................................
S to lo th r is s a .................................................................
S. t a n g a n i c a e ...................................................
............................. C LU P P o tam o 1 ■
■
............................. C LU P P o tam o 3 ■
■
.
.
.
.............................C LU P S tolo ■
■
.
............................. C LU P Stolo 1
■
■
.
■
.
158
■
■
.
.
159
............................. C LU P L im no 1 ■
............................. C LU P L im no 2 ■
■
■
.
.
.
.
160
161
.............................C LU P S ierr
■
■
.
.
162
............................. C LU P S ierr 1
■
■
.
.
162
T h r a t t i d i o n .................................................................
............................. C LU P T h rat ■
■
■
.
.
163
T. n o c t i v a g u s ..................................................
............................. C LU P T h rat 1
.
.
.
.
163
............................. C LU P L aev
■
■
.
.
164
............................. C LU P L aev 1
.
.
.
.
165
.............................C LU P C ongo
■
■
.
.
166
.
.
.
166
.
.
167
L im n o th rissa.................................................................
L. m iodon ..........................................................
L. s t a p p e r s i i ..................................................
S ie r r a th r is s a .................................................................
S. le o n e n s is ..........................................................
L a e v i s c u t e l l a ..........................................................
L. d e k im p e i..........................................................
C o n g o t h r i s s a ..........................................................
C. g o s s e i ..........................................................
G i l c h r i s t e l l a ..........................................................
G. a e s t u a r i u s ..................................................
S a u v a g e l l a .................................................................
.............................C LU P L im no
■
■
.............................C LU P C ongo 1 .
............................. C LU P G il
.
.
.
............................. C LU P G il 1 .
.
.
.
.
168
............................. C LU P Sauv
■
■
.
.
169
■
S. m a d a g a s c a r i e n s i s ....................................
............................. C LU P Sauv 1
.
.
.
.
169
S p r a t e l l o m o r p h a ..................................................
............................. C LU P S p ratm
.
.
.
.
170
............................. C LU P S p ratm 1 .
.
.
.
170
............................. C LU P E h ir
S. b i a n a l i s ..........................................................
F.hi r a v a ........................................................................
F.. f l u v i a t i l i s ..................................................
N a v e l l a ........................................................................
D. m a l a b a r i c a ..................................................
.
.
.
.
.
171
............................. C LU P E h ir 1 .
.
.
.
.
172
............................. C LU P D ay
.
.
.
.
.
173
............................. C LU P D ay 1 .
.
.
.
.
173
v in
C ode
Page
C l u p e o i d e s ............................................................................................................C L U P C lu p eo i
CL b o r n e e n s i s
CLUP
CL hy p se lo so m a
CLUP
CL p a p u e n si s
CLUP
CL v e n u lo su s
..............................................................................................C L U P
.
.
C lu p eo i 1C lu p eo i 2 .
C lu p eo i 3 .
C lu p eo i 4 .
.
■
■
■
.
.
174
■
■
■
.
-174
.175
.176
.177
C o r i c a ................................................................................................................... C L U P C o r i c ...............179
CL l a c i n i a t a
CL s o b o r n a
C lu p e ic h th y s
CL a c sa rn c n si s
CL bleekeri
CL g o n i o g n a t h u s
CL p e r a k e n s i s
P o ta m a lo s a
IL r i c h m o n d i a
C L U P C oric 1
C LU P C oric 2
.
C LU P C lupei
C LU P
C LU P
C LU P
CLUP
2 .2 .4
.
■
.
179
■ ■ 180
.
.
.
.181
■
.
■
■
■
.
■
■
.182
.183
.183
-184
C lupei 1 ■
C lupei 2 .
C lupei 3 ■
C lupei 4 ■
.
C L U P P o tam l
■
■
■ .186
C L U P P o tam l
1 ■
■
■ .186
■
■
■ .187
1 ■
2 ■
■
■
■ .188
■ ■ 188
H y p e rlo p h u s ............................................................................................................ C LU P H y p er
IL tr a n s lu c id u s
IL v itta tu s
■
C LU P H y p er
C L U P H y p er
S ubfam ily A l o s i n a e ............................................................................................................................................190
A lo s a .......................................................................................................................... CLUP A l o s .............. 191
A . a e s t i v a l i s . ............................................................................................ CLUP Alos 1 ................................... 192
A* a l a b a m a e . .............................................................................................CLUP Alos 2 ...................................193
Ax a lo s a ........................................................................................................... CLUP Alos 5 ................................... 194
A* b ra sh n ik o v i ............................................................................................ CLUP Alos 8 ................................... 195
A
c a s p i a ................................................................................................... CLUP Alos 7 ...................................197
A* c h r y s o c h l o r i s ..................................................................................... CLUP Alos 3 ...................................198
A* f a l l a x ....................................................................................................CLUP Alos 6 ...................................199
— k e ssl e r i ................................................................................................... CLUP Alos 15....................................201
A - m aeo tica
.............................................................................................CLUP Alos 9 ...................................202
— m e d i o c r i s . ..............................................................................................CLUP A los 13 . . . .
203
A p o n t i c a .....................................................................................................CLUP A los 10 . . . .
204
— p se u d o h a re n g u s .......................................................................................CLUP A los 14 . . . .
205
A
s a p i d is s im a .............................................................................................CLUP Alos 4 .................................. 206
A* s a p o s h n i k o v i . ..................................................................................... CLUP Alos 1 ...................................208
A^ s p h a e r o c e p h a la ..................................................................................... CLUP A los 12
. . . .
209
B re v o o rtia
EL
EL
EL
EL
EL
EL
C L U P B r e v o .............. 210
a u r e a ............................................................................................................ C LU P
g u n t e r i .....................................................................................................C L U P
p a t r o n u s .....................................................................................................C L U P
p e c t i n a t a .............................................................................................. C L U P
s m i t h i ..................................................................................................... C L U P
ty ra n n u s
..............................................................................................C L U P
E th m id iu m
B revo 4
■
B revo 5
.
B revo 1 .
B revo 6
.
B revo 2
■
B revo 3 ■
■ ■
.
. .211
212
.
. .213
■ ■ ■ 214
215
C L U P E t h m ...............217
IL m a c u l a t u m ..............................................................................................C L U P E thm l
.
.
. .
217
E t h m a l o s a ............................................................................................................ C L U P E t h ...............218
IL f i m b r i a t a ..............................................................................................C L U P E th 1 ...............219
H i l s a . ....................................................................................................................C L U P H i l s ...............220
IE k e l e e ........................................................................................................... C L U P H ils 1
.
.
.
.
220
IX
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Page
T e n u a l o s a ............................................................................................................. C L U P T e n u ...............222
U
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U
H
U
ilis h a
m a c ru ra
re e v e s ii
th i b a u d e a u i
to li
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CLUP
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. . . .
. . . .
■
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222
223
224
225
226
G u d u s ia .................................................................................................................... C L U P G u d ...............228
£L c h a p r a
(L v a r i e g a t a
2 .2.5
C L U P G ud 1
C L U P G ud 2
.
.
.
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228
229
S ubfam ily D o r o s o m a t i n a e .................................................................................................................................... 230
D o ro so m a
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a n a le
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p re te n e n s e
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C L U P D oros
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C lu p a n o d o n ............................................................................................................. C L U P C lu p an
£L t h r i s s a ...................................................................................................... C L U P C lu p an
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232
. 232
233
234
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236
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237
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1 . . . .
239
.
240
C L U P K ono
.
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CLUP Kono 1 .
240
N e m a ta lo s a ............................................................................................................. C L U P N e i l i ............... 241
LL a r a b i c a .......................................................................................................C L U P
ÍL c o m e .......................................................................................................C L U P
LL e r e b i .......................................................................................................C L U P
N. ilv e n s is ....................................................................................................... C L U P
LE g a l a t h e a e ............................................................................................... C L U P
N. j a p o n i c a ...............................................................................................C L U P
ÍL n a s u s .......................................................................................................C L U P
N . p a p u e n s i ...............................................................................................C L U P
LE v l a m i n g h i ...............................................................................................C L U P
N eili
N eili
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N eili
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242
243
244
245
246
247
248
250
251
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A n o d o n to sto m a ...................................................................................................... C L U P ....................... 252
A, c h a c u n d a
A s e la n g k a t
A. t h a i l a n d i a e
G o n ia lo s a
G. m a n m i n a
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C L U P A nod 2
C L U P A nod 3
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254
255
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C L U P G o n ..................................... 256
C L U P Gon 1
C L U P Gon 2
C L U P Gon 3
.
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256
257
258
.
.
F a m i l y P r i s t i g a s t e r i d a e ..............................................................P R I S T ................................................ 259
I l i s h a ..................................................................................................................... P R IS T I l i s h ....................................261
L
L
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I.
L
L
L
L
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a f r i c a n a ....................................................................................................... P R IS T
a m a z o n i c a ................................................................................................P R IS T
e l o n g a t a ....................................................................................................... P R IS T
f i l i g e r a ...................................................................................................P R IS T
f u r t h i i ....................................................................................................... P R IS T
k a m p e n i ....................................................................................................... P R IS T
m a c r o g a s t e r ................................................................................................ P R IS T
m e g a l o p t e r a ................................................................................................ P R IS T
m e l a s t o m a ................................................................................................ P R IS T
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263
264
265
266
261
268
269
270
272
X
C ode
T.
T.
T.
T.
T.
novacula
o b fu scata
p ristisa stro id e s
sirish a i
striatu la
P ello n a
P. casteln aean a
P.
davi
P. d itch ela
P. flavipinnis
P. harrow eri
lu tip in n is ..................................................
C h iro cen tro d o n
C. b le e k e r ia n u s ...........................................
N e o o p i s t h o p t e r u s ...........................................
N. cu b an u s
N. tro p icu s
...........................................
...........................................
O p i s t h o p t e r u s ..................................................
O.
O.
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12
13
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273
274
275
276
277
.................................... P R IS T P e li
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280
281
282
283
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.................................... PR IS T P lio 1 .
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.................................... P R IS T C hiro
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286
.................................... P R IS T C hiro 1
.
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286
.................................... P R IS T N eop
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287
.................................... P R IS T N eop 1.
.................................... P R IS T N eop 2 .
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288
288
.................................... P R IS T O pis
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290
3.
4.
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6.
1.
2.
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290
291
292
293
294
296
.................................... P R IS T O dont .
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297
.................................... P R IS T O dont 1
.................................... P R IS T O dont 2
.................................... P R IS T O dont 3
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298
300
.................................... P R IS T P rist
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.................................... P R IS T
.................................... P R IS T
.................................... P R IS T
.................................... P R IS T
.................................... P R IS T
P l i o s t e o s t o m a ...................................................
P.
.................................... P R IS T
.................................... P R IS T
.................................... P R IS T
.................................... P R IS T
.................................... P R IS T
dovii
..................................................
e f f u l e e n s ..................................................
e q u i t o r i a l i s ...........................................
m acrops
t a r d o o r e ..................................................
v a l e n c i e n n e s i ....................................
O d o n to en ath u s
O. c o m p r e s s u s ...........................................
O. m u c r o n a t u s ...........................................
O. p a n a m e n s i s ...........................................
P ristig a ste r
Page
P eli
P e li
P e li
P e li
P e li
3
4
1
5
2
.................................... P R IS T P lio
.................................... P R IS T
.................................... P R IS T
.................................... P R IS T
.................................... P R IS T
.................................... P R IS T
.................................... P R IS T
O pis
O pis
O pis
O pis
O pis
O pis
.
P. c a y a n a ............................................................
R aco n d a
..................................................................
R. r u s s e lia n a ...................................................
.............................
P R IS T R aco n 1
.
click for n e x t p a g e
click fo r p re v io u s p a g e
1.
INTRODUCTION
This catalogu e includes tax o n o m ic re f e re n c e s to all the known species o f clupeoid fish es (four fam ilies, 80
genera, more than 300 species), regardless o f their im portance or not to fisheries. Their classification is shown in
Fig. 1. The exact totals are not yet certain, but it is m ore than a century since G ünther attem pted in his
Catalogue o f fishes in the B ritish M useum (Vol. 7, 1868) to list a ll the valid species; F ow ler’s Catalog o f world
fishes (clupeoid part 1973), also tried to list all known clupeoid fishes, but the nam es and synonym ies are nowadays
m erely confusing. The p resen t a ttem p t is more an expression o f the ‘state o f the a rt’ than a definitive statem ent,
but it may serve to p in -p o in t th o se areas re q u irin g m o re ta x o n o m ic tre a tm e n t and it su m m a riz e s th e many
tax o n o m ic and n o m en clatu ra 1 changes that have been made in rec en t years.
In a group such as the clupeoids, w ith num erous and often closely sim ilar species, it is inevitable that some
true species have long been c o n c e a le d in th e sy n o n y m ie s o f o th e rs , w h i l e very many m o r e nam es have been
proposed than are actu a lly needed. A s a result, much o f our biolo g ical know ledge, e x cep t in the case o f a few
w ell-know n species o f m ainly the c o o le r w aters, is tied to identifications and nam es w hich may not be correct,
thus seriously lim iting the cred ib ility o f many b io lo g ic a l stu d ies.
T h e r e is o ften little th a t can be done in
retro sp ect; specim ens w ere n o t kept and m odern diag n o stic fe a tu re s w ere not m entioned in the publicatio n . For
this reason, many apparently useful references have been omitted here and the synonym ies are often confined to
the original proposal o f a synonym.
The clupeoid fishes are o f prim e im portance to fisheries. In fact, they represent the largest suborder o f
non-d o m esticated v erteb rates h arv ested by m an. H alf the w orld ca tc h o f fishes comes from about sixty species o f
various groups, but a third o f those prime species are clupeoids. U nfortunately, the com piling o f fishery data for
clupeoids has been ham pered, as w ith biological studies, by poor identifications and incorrect nom enclature.
U n d er such conditions, rational exploitation o f clupeoid resources, especially in tropical and subtropical seas, has
o ften been im possible. N e ith e r is there any g u aran tee th a t m odels d eriv ed from th e b etter know n co o l-w ater
species are ap p lica b le to the low latitu d e fish eries.
B oth b iological and fish eries stu d ies, th e refo re , m ust be u n derp in n ed
the present catalogue will contribute to this end.
by sound taxonom y. It is hoped that
The c lu p e o id fishes are united in the su b o rd er C lu p eo id ei o f the o rd er C lu p eifo rm es. They include the
herrings, sardines, sprats, shads, etc., the w olf-herrings and the anchovies. In general, the herring-like clupeoids
(fam ilies Clupeidae, Pristigasteridae) are easily recognized by their keel o f scutes along the belly, small and often
poorly toothed m ouths, and silvery appearance. S im ilarly, the anch o v y -lik e clupeoids (E ngraulididae) are usually
d istinctive b ec a u se o f their projecting, pig-like snout, larg e mouth and ‘underslung’ low er jaw . The fang-like teeth
o f the w olf-herrings (Chirocentridae) are also distinctive. On the w hole, fishery w orkers have little difficulty in
picking clupeoids out o f the net.
In spite o f this clup eo id Took’ h o w ev er, th ere are so many ex cep tio n s to any o f the m ore obv io u s and
e x te rn a l fe a tu re s used in the past to diagnose the clupeoids that m odern diagnoses rely mainly on sm all in te rn a l
ch aracters, such as details o f the caudal fin skeleton and the characteristic coupling o f the sw im bladder, inner ear
and head canal system. One result o f this difficulty is that in the ea rlier literatu re the clu p eo id s w ere freq u ently
allied w ith the tenpounders (E lo p id ae), ladyfishes (A lb u lid ae), milkfishes (C h an id ae ) and certain other Tower’ (i.e.
primitive) bony fishes and relegated to the order Isospondyli. In fact, there was no common ancestry to a ll these
groups and the clu p eo id s are rath er d istan tly related to the others.
For exam ple, the tenpounders and tarpons
(E lops, M egalops) and the ladyfishes (A lbula) have a rib b o n -lik e, lep to cep h alu s larv a, w hich allies them w ith the
eels. The clupeoids are now placed in the distinct suborder C lupeoidei and, w ith the suborder D enticipitoidei (sole
m odern genus D e n tic e p s), c o m p rise the o rd er C lu p eifo rm es.
Their relationship to fossil form s and to all o ther
bony fishes is shown in Fig. 2.
T hrou gho ut the C atalo g u e the term clup eoid refers to m em bers o f the su b o rd er C lu p eo id ei; clupeid refers
to the fam ily C lu p eid ae, p ristig asterid to the fam ily P ristig a ste rid a e , and engraulid to the fam ily E n g rau lid idae;
the subfam ily nam es are sim ilarly trea ted (c lu p ein e , p e llo n u lin e, a lo sin e , etc.).
p e l l o n u l in a e
C L U P E O ID E I
CL U PE ID A E
ALOSINAE
Fig. 1
C lassification o f clupeoid fishes
click fo r n e x t p a g e
click for p re v io u s p a g e
4 -Arm iq a tu s
EUTELEOSTEI
+ O rnate g ulum
IU. C re t. )
-t-EUimichthvs
(L. C re t. I -[ -P ala eo d e n tic e p s
D enticeps
IU. C re t. )
CLUPEOID
FISHES
lOlig.
IRec. )
S u b o rd e r'
S ubo rd e r
CLUPEOIDEI
DENTICIPITOIDEI
O rder
CLUPEIFORMES
DIVISION
DIVISION 2
S u p e ro rd e r
CLUPEOMORPHA
F ig . 2 R e la tio n s h ip s o f c lu p e o id -lik e fish e s (fo s s il fo rm s, fro m
L ow er and U pper C retaceous, E o c e n e and Oligocene, show n w ith + )
1.1 C lupeoids in W orld F is h e r ie s
1.1.1 Clupeoid catches
M ore clupeoid fishes are caught (by w eight, but presum ably also by num ber) than m em bers o f any other
single system atic group o f fishes. The follow ing figures, from the FAO Y earbook o f fishery statistics (1982) bear
th is o u t.
A ll m a rin e and fre s h w a te r fishes
C lu p e o id fishes
C o d - lik e fishes
T u n as
F la t f i s h e s
67 769 371 to n s
18
8 9 7 7 3 1 to n s
10
9 6 9 0 4 2 to n s
2
59 3 2 1 2 to n s
1
126 8 7 9 to n s
= 1 7 .9 %
= 1 6 .2 %
= 3 .8 %
= 1 .7 %
This huge contribution made by clupeoid fish e s to w orld fisheries can be expressed in other ways. For
example, the top 56 fish species ( i.e . those harvested at o v e r 100 000 tons in 1982) accounted for 40 004 027 tons
or about 60% o f the w orld fish c a tc h . O f these 56 top sp ecies, no less than 18 w ere clupeoids, w hich contributed
16 693 748 tons or about 24% o f the world fish c a tc h . A gain, in 1982 there were 24 fish sp ecies harvested at o v e r
h alf a m illio n tons. O f these
24 s p e c ie s, 10 w ere c l u ­
peoids,
w hich
contributed
ju s t o v e r 20% to the total
w orld fish c a tc h . In that
year, the second m ost e x ­
ploited fish in the w orld was
a clupeoid (Sardinops m elano­
stictus o f Japan) , as also the
third (R sagax o f P eru and
Chile) and the seventh, ninth
and tenth (E ngraulis ringens
o f Peru and C hile, form erly
the m ost exploited fish in the
w orld; C lupea harengus o f
the N orth A tlantic; Sardina
pilchardus o f the northeast
A tlantic and M editerranean).
In fact, the m ost h eavily
exploited fish in the w hole
history o f w orld fishing was
the P eruvian anchoveta E.
tin g en s o f w hich o v e r 13
m illio n to n s w a s la n d e d in
1970.
F ig .3 C lu p eo id s as a p ercen tag e o f th e to tal fish c a tc h
in each fishing area (1 9 8 2 fig u re s)
In four o f the FAO fishing a re a s (see Fig. 3), c lu p e o id s
fish in g a re a s th e y a c c o u n t fo r o v e r a q u a r te r o f th e
a c c o u n t fo r over h a if the fish catch and in an o th er six
f is h catch.
4
The size o f the clupeoid fisheries is the result o f tw o m ain factors. F irst, the m ajority o f clupeoids feed
close to the base o f the food chain and can thus benefit more directly from nutrient-rich areas w here there is a
seasonal or more c o n tin u o u s bloom o f plankton. Second, clupeoids are alm ost alw ays schooling fish es and thus
extremely vulnerable to nets (especially purse sein es), w hich can catch a la rg e volume o f fish in a short time. Since
it is the cooler high latitude seas and the areas o f upw elling that are richest in plankton, it is here that the m ajor
clupeoid fisheries exist, as the following figures show (d iv ision s not based on FAO fishing a re a s ; 1982 to ta ls ).
A l l c lu p e o id sp ecies
C ool or upw elling areas
S u b tro p ic a l a r e a s
T ro p ic a l a r e a s
18
15
1
1
897
603
777
516
731
531
278
922
to n s
tons
to n s
to n s
= 82.6%
= 9.4%
= 8.0%
N ot only do the cooler-w ater clupeoids d o m in a te the total clupeoid catch, but they r e p r e s e n t nearly a
quarter o f the world fish production. The 1982 catch o f the Japanese pilchard (Sardinops m elanostictus) w as alone
more than the c o m b in e d catch o f all tropical and subtropical clupeoids, while the catch o f the South A m erican
pilchard (Sk sag ax ) was alm ost that amount. In tropical w aters, clupeoids are re p r e s e n te d by more species, none o f
w hich can rival in abundance the cool-water species. The only w arm -w ater clupeoids whose stocks approach those
o f the c o o le r w aters are m em bers o f the Sardinella aurita com plex (Sk aurita in the eastern and w estern Central
A tlantic, plus Sk brasiliensis; Sk longiceps in the Indian Ocean; ík lem uru in Indonesia, w estern A ustralia and
Japan), also the unrelated Sk m aderensis o ff W est A frica.
H ow ever, in each area these fishes are associated w ith
at least som e upwelling during the year and in at le a s t som e part o f their natural range.
The predom inance o f the cool-w ater clupeoids does not m ean that the tropical and subtropical clupeoid
fisheries are unim portant. First, they may in som e cases be c a p a b le o f further developm ent by more m echanized
exploitation and better processing facilities, w hereas the cool-w ater clupeoid fisheries are usually fully or ev en
over-exploited and are often prone to collapse. Second, the w arm -w ater clupeoid fisheries are frequently a
subsistence rather than a cash resource in exactly those a re a s w here other sources o f anim al protein are
inadequate or unused. Certainly, their c a sh value is low, but so also is their local p u rc h ase price, so that their
contribution to nutrition in such a re as is not to be ju dged m erely by contrasting w arm -w ater w ith cool-w ater
catches.
g . •}>• a ,
The la r g e s t
clupeoid
catches (see Fig. 4) com e
from the southeast Pacific
(Fishing A rea 87; 27.4% o f
all
clupeoid catches), based
primarily on the Peruvian pil­
chard
and
the
P eru v ian
24.6
anchoveta, and also from the
northw est P a c i f i c ( F is h in g
A rea 61; 24.5% o f all clu ­
peoid catches), based prim a­
2.4
rily on the Japanese pilchard.
N ext in im portance are th e
northeast
A tlantic
(Fishing
A rea 27), the eastern Central
Pacific (Fishing A rea 77), the
M editerranean and B lack Sea
(Fishing A rea 37) and the
w estern
C entral
A tlan tic
(Fishing A rea 31), each o f
Clupeoid catches by
w hich c a tc h e s o v e r a m illio n
catch
tons o f clupeoids annually.
The rem aining a re a s , not all o f them tropical, e a ch c a tc h e s
term s, the entire Pacific yields ahout 60% o f all clupeoids,
a b o u t 3% .
nr -ae «
jr . y -g
V«
(X)
fishing area as a percentage o f the total world
o f clupeoids (1 9 8 2 fig u re s)
three quarters o f a m illion tons or less. In general
the A tlantic ahout 30% and the Indian O cean only
As in other groups o f fish e s, the cool-w ater and upwelling a re a s support few clupeoid species and the main
com ponent o f the catch may come from a single species, or perhaps two. Often these are a pilchard ( Sardinops)
and an anchovy (E n g rau lis), w hose relative abundance may oscillate as a result o f a com plex ecological
relationship betw een the two (further m odified by bird or fish predation and hum an exploitation). The subtropical
and especially the tropical a re a s , on the other hand, support a great diversity o f clupeoids and the fisheries are
m ultispecies. A purse seine o r a heach seine may bring in ten
or more clupeoid species w hich
h a v e schooled more
or less together, but little is know n yet o f the factors that determ ine the relative abundance o f each species and
the effect than one species may have on another. In term s o f diversity, the richest area is the Indo-W est Pacific
( F is h in g A re a s 5 1 , 5 7 , 71), w ith about 160 clupeoid species or h a lf the total clupeoid species know n, yet it
produces only 7% o f the world clupeoid catch.
5
C atches o f clupeoids have increased,especially since the
second w orld
w ar, som etim es very much in
advance o f the general rate o f increase in fish production (especially during the boom years o f the Peruvian
anchoveta fishery in the late sixties and early seventies) (Fig. 5). D om inating the clupeoid increase have been the
pilchard (Sardinops) and anchovy
(E ngraulis) fish eries and no t the trad itio n a l n o rth ern fish eries for h errings
(C lu p ea), nor the tropical clupeoid fisheries. Characteristic, how ever, o f those clupeoid species w hich dominate
the fisheries o f particular areas is
a tendency to oscillate rather drastically in their abundance (Fig. 6). A lthough
the production o f good or bad year-classes (w hich may vary by a factor o f ten or m ore) can be related to
ecological factors, the precise role played by fisheries is still not clear. The considerable fluctuations in catches
o f A tlanto-S candian herrings (C lupea harengus), both before the second w orld w ar (27 to 108 thousand tons) and
after (33 to 78 thousand tons) may merely have reflected fishing effort itself, not interm ittent over-exploitation.
N evertheless, w ith the collapse o f this fishery in 1970 follow ing heavy exploitation o f the outstanding 1959 and
1960 year-classes, a period o f strict regulation has seen an apparent recovery. W hether low stocks can produce
good recruitm ent even w hen ecological factors are favourable is not yet certain. The dram atic collapse o f the
Peruvian anchoveta fishery after the m id-seventies poses the same problem , as also that o f the C alifornian
pilchard collapse in the early fifties.
60 -
-
SO - -
40 -
-
-
60%
CLUPEOIDS
30 -
20 -
-
-
TOTAL
MARINE
FISH
CATCH
-
-
20 %
TOTAL
MARINE
CLUPEOID
CATCH
1938
F ig.5
19S0
1960
1970
1980
W orld catches o f marine fishes and o f clupeoid fishes, 1938 and 1949-1983 (in millions o f tons, on left).
A lso, the clupeoid catch as a percentage o f world catch (% on right).
6
tl
|- \ .y s /
m ton*
\
Brevoortia tyrannus
Brevoortia spp.
V
./
~ \/—
/
Sardinops meianosticus
Engraulis encrasicolus__
Sardinops eaeruicus
1955
1975
Fia. 6
195S
1980
1970
Trends in catches o f m ajor clupeoid species,
(divisions on left are 500 000 tons)
1955-1983
Since mem bers o f the same clupeoid genus often have a fairly sim ilar biology, and since sym patric mem bers
o f a genus may not always have been separated in the catch statistics, it is o f interest to know w hich genera play
a m ajor role in fisheries. In fact, 10 clupeoid genera, com prising about 36 species, contribute over 90% o f the
total world clupeoid catch (Fig. 7). The follow ing figures show the catches for 1982:
S pecies
C atch in tons
% o f total clupeoid catch
Sardinops (4 species)
7 820 021
41.4
Engraulis (6 species)
3 570 522
18.9
B rev o o rtia (3 species)
1 257 906
6.6
C lupea (2 species)
1 207 478
6.4
Sardinella (6 species)—^
1 056 324
5.6
Sardina (1 species)
896 975
4.7
Opisthonema (2 species)
576 555
3.1
Sprattus (1 species)
525 135
2.8
C lu p eo n ella (1 species)
434 655
2.3
Stolephorus (about 10 species)
231 344
1.2
93.0
1/
Only S. aurita.
brasiliensis.
lem uru.
longiceps.
m ad e re n sis.
neglecta
7
The order o f these genera varies a little from year to year, depending on the state o f the stocks. Other
genera w hich have been o f im portance in previous years (i.e. a catch o f over 100 000 tons) are the round herring
Etrum eus (1 species), the w o lf herrings (C hirocentrus, 2 species) and the A frican freshw ater Stolothrissa and
Lim nothrissa (1 species each).
C learly these are the genera that m erit further fishery, biological and taxonom ic
study.
7.82
1.06
ce «t
Sardinella
Sardinops
— 1*0
3.57
Sardina
ijf
0.57
Engraulis
1.26
Opisthonema
- 3 0 ----0.53
Brevoortia
m—
Sprattus
1.21
0.43
— 20
Clupea
Clupeonella
0.23
Stolephorus
O t/j
z
_
UJ
.
2
CL
— 10—
S'*
O
CL
UJ M
a. _
I / I __
F ig . 7
01 XI
o
to ™
R e la tiv e im p o rtan ce o f the te n m ajo r clu p eo id gen era (as % o f to tal clu p eo id catch;
f is h e s ta g g e d w ith th e i r c a tc h in m illio n s o f to n s ; 1982 f ig u r e s )
1.1.2 F ishing gear
Since clupeoids are mainly schooling fishes and thus most easily caught by nets, quite a wide variety o f nets
and netting techniques has been developed to suit particular environm ental and economic restraints. A lthough the
m ajor clupeoid catches come from fully mechanized large-scale netting operations, it is w orth recording also the
sm all-scale artisanal gear w hich in hum an and nutritional term s is nevertheless im portant in many tropical and
subtropical areas. Even when clupeoids are merely a bycatch in such artisanal gear, earful sorting can result in
large baskets containing one or two species o f small Stolephorus w hich are as much appreciated as the large ‘tab le’
f is h e s .
S u rrou n d in g nets : Used to encircle schools at or near the surface. There are two kinds, both usually set
w ith a single boat (or w ith an auxiliary craft).
Purse seine - a running line at the bottom o f the wall o f netting that has been set around the school closes
the net like a purse. U sed w ith or w ith o u t a lig h t to a ttrac t the fish es. E m ployed for the S ard in ella
fisheries o ff W est A frica and the Sardinops and Engraulis fisheries o ff Peru, Chile and elsewhere.
Lam para net - no purse lines, but w ith a spoon-shaped sm all-m eshed central bunt and slender w ide-m eshed
lateral wings. U sed w ith or w ithout a light. Em ployed generally for the same clupeoids as the purse seine.
Seine nets : Used betw een surface and bottom, or at the bottom, to encircle schools, w ith or w ithout a bag
at the cen tre and u su ally h au led by a ro p e at each e n d .T h ere are tw o m eth o d s o f o p eratio n .
B each seine - set from the land, either by w ading or from a small boat, then hauled to the beach. Very
comm only used in most tropical or subtropical areas for small inshore clupeoids (Sardinella, Elerklotsichthys,
T hryssa, Stolephorus, etc.).
B oat seine - operated from a boat, w ith long ropes to herd the fishes, usually hauled along the bottom; the
m ost representative is the D anish seine (used for flatfishes, but less frequently for clupeoid fishes).
T raw ln ets :
B oth b ottom traw ls and m id w ater traw ls w ill catch clupeoid fish es, but ap art from a few
species subject to aimed fisheries they generally form a b y c atc h and are not economically exploited in this way.
L iftn ets : Elorizontal nets that are raised w hen fish are suspected to be over the net or have been attracted
there by light. There four tw o main kinds.
P o rtab le - small hand nets used from river banks, jetties, etc.
B o a t-o p e ra te d - bag nets and blanket nets w orked from one or more boats.
clupeoid fishes, especially if used w ith a light.
E x trem ely effectiv e for sm all
S h o re -o p e ra te d - portable or fixed; a h o rizo n tal net is lifted by an o ften in g en io u s co u n ter-w eig h t or is
w inched by hand or m achine. E ffectiv e for sm all clu p eo id s, esp ecially i f used w ith a light. E m ployed in
Southeast Asia.
R ig -o p e ra te d - a large bamboo rig or tow er in w ater o f 5 to 15 m encloses a square net w hich can be
w inched up; from the platform on top a light is low ered to attract the fish. Com m on along the Java Sea
coast o f In d o n esia, w here they are called bagan. E ffectiv e for sm all clupeoids (esp ecially S ard in ella,
Thryssa and Stolephorus).
F a llin g gear : C ast nets produce rather few clupeoids.
G illnets : V ertically-hung nets, at the surface, m idw ater or bottom, w ith a mesh size that usually ‘g ills’ a
restricted size range o f fish es; often set in long lines or ‘fleets’ o f nets.
D riftin g gillnets (d riftn e ts) w ere the
trad itio n al g ear for the n o rth ern h errin g fish eries (C lupea).
S hort gillnets are also used in rivers, being drifted
downstream to catch runs o f anadrom ous fishes. They are used in this way for shads (Tenualosa species) in India.
T raps : Either w ith a non-return funnel or relying on the inability o f the fish to turn and escape (aided by
the pressure o f the w ater
current). Three m ain types catch clupeoids.
Fvke nets - conical n o n -retu rn bag w ith w in g (s) held ap art by stakes; used in
som etimes caught in fyke nets.
sh allo w w ater. Shads
are
Stow nets - conical or w ed g e-sh ap ed nets freq u en tly used in estu arie s or lo w er parts o f riv ers w here the
cu rren t is strong, the net being held by stakes or anchors.
C atch es an ad ro m o u s, eu ry h alin e or riv erine
clupeoids. In some cases a basket is used rather than a net.
B arriers - estuarine barriers o f stakes, reeds, etc., w ith w ings to direct the fishes; they often catch inshore
a n d m a in ly eury h alin e clu p eo id s, p articu larly in tro p ical areas. R iver barriers or w eirs w ith non-return or
funn el-sh ap ed baskets are used for an ad ro m o u s c lu p eo id s such as shads (e.g. T en u alo sa ascen d in g In d ian
rivers).
Hook-and-line : A lthough not im portant econom ically, it is o f interest that some clupeoids will take a hook,
as fo r exam ple larg e p ilc h a rd s in th e n o rth e a st P a cific ( S ard in o p s c a e ru le a ) or shads in E u ro p e (A lo sa sp e c ie s).
P um ps : U sually operated in conjunction w ith a light to attract schools.
chilka (C lupeonella species).
E xp losives :
tow ers.
Em ployed on the Caspian Sea for
A lth o u g h illeg a l, ex p lo siv es have been used in stead o f the liftn e ts from the Java Sea bagan
P oison s : Some freshw ater clupeoids occur in the more general catches made by poisons in tropical rivers.
9
1.1.3
U tiliza tio n o f clu p eo id s
Clupeoid fishes contribute to world food resources in two ways: directly, through actual consum ption (fresh,
frozen or processed) and indirectly, by providing products used for animal feeds and fertilizers or by serving as
bait to catch other fishes further up the food chain. Very small clupeoids may be consum ed w hole (e.g. w hitebait
S prattus sp rattu s in E u ro p e, sm all S to lep h o ru s in so u th east A sia):
slightly larg er clupeoids (e.g. p ilchards,
sardines. S ard in a. S a rd in o p s) m ay be pro cessed so th at all but the head is p alatab le: m ost m edium and large
species, however, are valued alm ost entirely for their epaxial and hypaxial body m uscles, the head, guts, fins and
axial skeleton being rem oved and the interm uscular bones largely ignored. Parts that are consum ed or m arketed
separately are the gonads (e.g. the eggs o f the Pacific herring Clupea pallasii from the Pacific coast o f Canada)
and oils or body fluids extracted from the liver and other tissues (e.g. from the m enhaden B revoortia tyrannus).
From earliest tim es the need to store clup eo id catches for later co n su m p tio n n ecessita ted som e form o f
preservation.
N ow ad ay s a large p art o f the w o rld clup eo id catch is p rocessed in one w ay or an o th er, m ost
particu larly in the case o f the enorm ous single species fish eries o f tem p erate w aters (fo r S a rd in o p s. E n g ra u lis.
C lupea, etc.). The tropical clupeoids, on the other hand, are more frequently eaten fresh (or after short storage in
ice), not only because industrialization is generally less advanced in such areas, but also because catches are
relatively sm aller and the species com position is much more diverse: nevertheless a substantial proportion o f the
catch may be cured (o ften su n-dried or sm oked), esp ecially the sm aller species.
A s clupeoid fish eries have
developed (e.g. the Sardinella fisheries o f W est A frica), the investm ent in m echanized exploitation has had to be
m atched by th at for sto rag e and p ro cessin g . The co n co m m itan t rise in price o f the product has usually meant
m arketing in the developed countries.
Clupeoid fishes are utilized in ten m ain ways. For the main categories an approxim ate percentage is given
below, although this can vary from year to year, depending on availability and market factors (especially prices).
The collapse o f the Peruvian anchoveta fishery led to a drop in fishm eal and fish oil production (Peru becam e a
major fish oil im porter in 1983), but price rises (e.g. by 51% for fish oil in 1983) probably stim ulated meal and oil
p ro d u ctio n from o th er species. N ew processing techniques, as also alternative sources (e.g. jojoba for vegetable
oil), also vary patterns o f utilization.
(Direct)
Fresh (17% ): all species, a lth o u g h fresh
co n su m p tio n
is m inim al
in som e
(e.g.
m enhaden).
Frozen (12%): most species, depending on the need to store or transport.
Cured (12%): num erous species, especially Clupea in northern w aters and small clupeoids in the tropics:
salted (dry or brine so lu tio n ), sm oked, sun- an d /o r w in d -d ried :
also allow ed to ferm en t to produce a
condim ent (Roman garum . G reek halec: southeast A sia nowadays) or a fish silage as an animal feed additive.
Canned (14%): industry dom inated by scom brids and clupeoids, the latter including species o f C lupea.
Sprattus. Sardinella (chiefly the S^ aurita com plex). S ardina. S ardinops. B revoortia and Engraulis (R
en c ra sic o lu s, also the A rg en tin e R a n c h o ita ): sardines and p ilch ard s form alm o st h a lf the canned sm all
p elagios; packed mainly in vegetable oils, but also in tom ato sauce, brine or water.
Minced : chiefly applied to gadoid fish es, but trials have been m ade w ith C lu p e a. S p rattu s.
S ard in ella.
Sardinops. B revoortia. Engraulis and Thryssa: the flesh is separated in a com inuted form from skin, bones,
scales and fins: various clupeoids used for fish paste (especially anchovies).
Fish flour : refinem ent o f techniques used to produce anim al feeds from whole degutted fishes, resulting in
a useful protein additive for hum an consum ption.
Oil : over 80% o f the U SA fish oil p ro d u ctio n has in som e y ears com e from the m en h ad en B rev o o rtia
tyrannus (yielding 8 to 20 gallons o f oil per 1000 fishes, depending on how far north the fishes are caught):
body oils (used in paints, soap, shortenings and m argarine) are also extracted from Sardinops and E ngraulis:
vitam in rich condensed fish solubles, serving as an animal feed additive, result from w et reduction o f fishes
such as menhaden.
(Indirect)
Animal feeds (44%) : ch iefly C lu p ea . S ard in o p s. B rev o o rtia and E n g ra u lis: the p rin cip al p ro d u ct from
reduction plants, using either w hole fish (w ith or w ithout extraction o f oil) or offal: used for supplementing
feeds for cultured fishes (trout), poultry, cattle, sheep and pigs.
F e rtiliz e rs : herring in Scandinavia and m enhaden on the eastern coast o f N orth A m erica have been used
since earliest tim es, but the practice is relatively unim portant nowadays.
Bait : Sardinops and E ngraulis used for ‘chum m ing’ tuna: species o f Stolephorus used as live-bait, especially
in southeast Asia.
click fo r n e x t p a g e
click fo r p re v io u s p a g e
10
1.2
P la n o f th e C a ta lo g u e
Four clupeoid fam ilies are presented: Chirocentridae, Clupeidae, Pristigasteridae and Engraulididae. For each
fam ily are given general descriptive rem arks, notes on biology and fisheries, and a key to the subfam ilies and
genera. Thereafter, the genera are given in the same sequence as in the fam ily (or subfamily) key, but the species
w ithin them are arranged alphabetically. For each genus a b rief diagnosis is given w hich supplements the data in
the key: also references to junior synonym s (some o f w hich may stillbe in current
use or entrenched in the older
literature), notes on biology and fisheries, and a b rief reference
to the
num ber o f species, particular taxonom ic
problem s, and any major revisional studies. Keys are not given to species since in many cases existing keys are
unw ieldy (especially in large genera such as Sardinella. Thryssa and A nchoa) or in need o f further refinem ent:
how ever, the species diagnoses contain com parisons w ith m ost species likely to be confused in the area. The
inform ation given for each species is arranged in the follow ing paragraphs:
( 1)
Scientific N am e : R eference
is given to the first valid nam e applied to the species and its
accom panying description: also, the type locality. E arlier but invalid names appear in the synonyms
(w here n e cessary ).
( 2)
Synonyms : All names that have been applied to the species are given, as well as some different name
com binations (if significant) w ith author, date, page number and usually a com m ent in parenthesis. The
complete synonym ies (i.e. all reference in the literature) o f some clupeoid species are enorm ous and no
purpose w ould be served in citing them in full: in any case, there is often grave doubt w hether the
material studied was really indentified correctly. W here possible, one or more references at the end o f
each synonym give the correct name and the author(s) w ho correctly described the species, discussed
its synonym s or provided distributional or other data.
(3)
FAO N am es : O nly E nglish
nam es have been chosen for use w ithin FA O and to serve as the
recom m ended names for fishery, m arketing and other purposes. Some French and Spanish equivalent
names for clupeoid fishes have already been given on FA O Identification Sheets, but it is considered
prem ature to attem pt this for all clupeoid species until fishery and other specialists in countries using
French and Spanish can be consulted.
(4)
D iagnostic Features : D istinctive features are given, accom panied w here necessary by diagram s, as
an aid to identifying species w ithin a genus (after using the keys to fam ilies, subfam ilies and genera).
I f the species is more fully described by on FAO Species Identification Sheet, reference is made at the
end to the alpha-num erical FA O code o f the species and the Fishing A rea (e.g. ‘see CLUP Spratel 2,
Fishing A rea 51’ in the case o f Spratelloides g ra c ilis). In principle, the species is diagnosed to separate
it from sim ilar sym patric species and not from all other species in the genus (i.e . those that occur in
other areas).
( 5)
G eographical D istribution : The general range o f the species is given in the text and in the map (areas
betw een scattered records may m erit a question m ark if a continuous range seems
doubtful). W here
appropriate, the list o f species given after each genus is arranged geographically.
(6)
H abitat and B iology : The true identity o f the species is often in doubt, especially in the older
literature, so that much apparently useful biological inform ation has had to be omitted here. For very
many o f the tropical species, how ever, alm ost nothing is known o f feeding habits, spawning reasons,
m igrations, tolerances and preferences for particular tem perature or salinity ranges, etc.
( 7)
Size : The maxim um know n size and w here possible the usual adult size are given (as standard length,
in cen tim e tre s). A s a rule th e av erag e ad u lt size
is g e n e ra lly a b o u t 1/4 less than the m axim um .
(8)
Interest to Fisheries : Partly based on the statistics by species and by Fishing A rea given in the FAO
Y earbook o f Fishery Statistics for 1983. Since only 47 clupeoid
species (out o f over 300 described
species) are given individual statistics, and since identifications are sometimes doubtful (i.e. more than
one species probably included), only a general im pression o f the relative im portance o f a species is
possible in many cases, or reference to a particular locality w here significant catches w ere seen or
reported. H ow ever, from fisheries literature for particular areas or particular species considerable
inform ation on size o f fishery, gear and utilization is available. This has been included w henever
identifications can be trusted.
(9)
Local Names : These are only occasionally given, usually w hen a particular species is the basis for a
m ajor fishery and its identity is certain. In very many cases, how ever, a local name is applied to
several clupeoid species and m eans no more than that the fish is a kind o f sprat, sardine or anchovy.
( 10 )
Literature : R eference is made to the m ost im portant (not only taxonom ic) books or papers on the
species, w ith a b rief indication o f their relevance in parentheses. Some or all may already have been
cited under the synonym s, w here the page reference is also given. See page 14 for list o f m ajor
works.
TO CONTINUE
>
11
(11) R em arks : Taxonom ic or other problem s are outlined here, e.g. explanations for unexpected name
changes, doubtful status o f species, indications o f further w ork in progress or needed, and presence o f
subspecies (w hich are given a b rief diagnosis and geographical range).
1.3
P rob lem s
of
Id en tifica tio n
M any clupeoid sp ecies, esp ecially those o f tro p ical and subtropical w aters w here d iv ersity is greatest, are
difficult to identify, partly because the distinctions require a m icroscope or good hand lens (for gillraker numbers,
shape o f second supra-m axilla, form o f scale striae, etc.), and partly because the taxonom y o f some genera is not
su fficien tly w ell understo o d (e.g. the clupeid genera S ard in ella and H e rk lo tsich th y s, or the en g rau lid genera
Thryssa. A nchoviella and especially A nch o a).
The keys to fam ilies, subfamilies and genera have been based, where possible, on the most obvious external
ch aracters, but the lo n g er g en eric keys (e.g. for the C lu p ein ae) may pose p roblem s. H o w ev er, since ra th e r few
genera are shared betw een the Indo-W est Pacific and the N ew W orld, the distribution o f the genera has often been
introduced into the key. The elim ination o f genera (and species) not found w ithin the area studied can often save
tim e and side-step d iffic u ltie s over u n fam iliar characters. For this reason the occurrence o f genera and species
w ithin the 19 marine and 9 freshw ater F A O Fishing A reas is given (see Part 2). H aving arrived at a particular
genus by using the key, its occurrence in the area studies should be checked. This list will also help in narrowing
down the num ber o f species diagnoses that should be consulted, although this can also be done from the list o f
species given after each generic diagnosis.
A ssum ing that the correct genus
has been found, a quick glance at the illustrations may suggest one or more
likely species, especially if the shape
or colour pattern is distinctive. The species diagnoses are designed to lead
to the correct species or even genus, based on com parisons w ith sim ilar species or genera sharing even a small
part o f the geographical range. H owever, the sim ilarities o f preserved specimens (on w hich many o f the diagnoses
are based) may be quite different from the sim ilarities o f fishes still in the net. It is as w ell, therefore, to check
all the characters in the diagnosis o f the com pared species (or genus), w here additional characters will be found.
S hort-cuts to clupeoid id en tificatio n s are reg rettab ly few.
C olour is gen erally o f little help, but som e
clupeoids have characteristic black spots along the flanks ( Sardina. S ard in o p s. A m bligaster sirm , some H e rk lo tsichtliys. some shads), and others have black tips to the tail (e.g. Sardinella m elanura or a bright silver stripe on
the flank (Lile and some anchovies).
A m ong clupeids the num ber o f pelvic finrays is often diagnostic o f a genus or subfamily (but in anchovies
there are alw ays 7 finrays, ex cep t in C oilia ram carati w ith 10). In the tab le b elo w the co u n ts are g iven for
all clupeoids. A few ch aracters may instantly diagnose certain species or genera, such as the canine teeth o f
C hirocentrus. Lycengraulis and L ycothrissa, the light organs o f Coilia dussum ieri, the rat-tail body o f all Coilia or
the enorm ously long upper ja w (m a x illa ) in T h ry ssa s e tir o s tr is , but for the m ost part one must rely on tedious
counts o f fin ra y s , gillrakers and scales, m easurem ents o f body proportions, and exam ination o f small body parts.
M u ch o f th e c o n fu s io n in c lu p e o id lite r a tu re is b e c a u s e w o rk e rs d id n o t a lw a y s do th is in th e p a s t.
Family
Ge n e r a
Pelvi c finrays
C HIRO CEN TRIDA E
C h iro cen tru s
CLU PEID AE
(D ussum ieriinae)
D ussum ieria. E trum eus.
Spratelloides.
Jenkinsia
(C lupeinae)
7
7 or :
P la ta n ic h th y s. R a m n o g a ster. E scu alo sa
S p rattu s sp rattu s
A ll o th er S p ra ttu s: H a re n g u la . H e rk lo tsic h th y s.
O pisthonem a. L ile , Sardina. Sardinops. R hinosardinia.
C lu p e o n e lla . A m b ly g a ste r. S tran g o m era, m ost
S ardinella
S ard in ella a u rita . £L b ra s ilie n s is . S_. le m u ru . £L longc ep s. R n e g lec ta: C lupea (o ccasio n ally 8 or 10)
(Pellonulinae)
7
7 or !
S ie rra th rissa . C lu p eich th y s perak en sis
T h rattid io n
A ll other Clupeichthys
all other genera
(A losinae)
B re v o o rtia . E th m idiu m
E thm alosa. H ilsa. Tenualosa. Gudusia
A losa
(D orosom atin ae)
All genera
TO CONTINUE
>
12
Fam ily
Pelvic finrays
Genera
6
PR ISTIG ASTERIDA E
C h iro c e n tro d o n ; P ello n a harrow eri
Ilisha
P ello n a
P lio ste o sto m a , O d o n to g n a th u s, N e o o p isth o p te ru s,
O p isth o p te ru s, R a c o n d a ; som e specim ens o f
P ris tig a s te r (if present, then 5)
6 or 7
7
(ab sen t)
ENG RAU LIDID A E
(E ngraulidinae)
All genera
(C ollinae)
7
10
C oilia (except one species)
C oilia ram carati
1.4 T ech n ica l T erm s, M ea su rem en ts and C ou n ts
A typical herring and a typical anchovy are shown in Figs 8 and 9. In many taxonom ic w orks on clupeoids a
large num ber o f external features are given, either as percentages or in the older literature as proportions o f
standard length, or o f head length; in some cases one body part can be expressed as x tim es larger than another
(or contained x tim es w ithin it). O f such proportional m easurem ents, the follow ing are the m ost useful and have
been used here (Fig. 10).
unbranched soft rays
d orsal filam ent
p resen t in
som e species
p red o rsal scales
2nd
su p ram ax illa
u n b ran ch ed soft rays
sub-o p ercu lu m
po stp elv ic
scutes
pre p e lv ic scutes
typical h errin g -lik e fish
bran ch ed
soft rays
unbranched soft ray:
axillary scale
low er ]aw
un d erslu n g
2nd
supram axilla
'
m axilla
often long
prepelvic
| |i
postpelvic
branched
soft rays
typical anchovy-like
unbranched
scutes often present
so^ raYs
fish
13
standard length (SL)
pre-dorsal
d orsal fin base
head
snout
eye
d orsal fin
o rig in
upper
ja w
hind tip
o f maxilla
anal fin base
pectoral fin
o rig in
p elv ic u n
in sertio n
m easurem ents used for clupeoid fishes
F ig .10
S ta n d a r d L e n g th - tip o f snout to end o f body, i.e., to base o f caudal fin (w h ere th e finrays re a c h th e
hypurals). Bending the tail produces a w rinkle at this point.
B od y d ep th - measured at deepest point, usually under origin o f dorsal fin. Fishes are described here as
slender w hen the depth is less than about 25% o f standard length; m oderately slender at 25 to 30% o f standard
length; m oderately deep at 30 to 35"..: deep at 35% or more.
Head length - the longest possible m easurem ent,
alw ays a horizontal measurem ent).
from tip o f snout to hind border o f gilicover (thus
Snout versus eye - tip o f snout to front border o f eye, com pared w ith the horizontal diam eter
M ost usefully expressed as the one larger than the other.
U pper ja w - from tip o f snout to hind tip o f m axilla, but m ost often expressed
maxilla, e.g. to beyond hind border o f eye, to hind border o f gili cover, etc.
as
not
o f the eye.
the point reached by
the
Post-orbital - from hind border o f eye to hind border o f gili cover, the longest measurem ent (thus not always
horizontal).
Fin positions - the origins o f the dorsal and anal fins, and the insertions o f the pectoral and pelvic fins (their
vertical position along the axis o f the body) are either compared relative to each other, e.g. pelvic fin insertion in
advance o f dorsal fin origin, anal fin origin behind dorsal fin base); or given as a pre-pectoral, pre-dorsal, p re pelvic or pre-anal distance (percent o f standard length or for the dorsal origin often given as before or behind
midpoint o f body
Fin b ases - the distance betw een the first and last finray bases. R ather than give fin ray numbers, it is often
easier to state that the anal fin base is longer (or shorter) than the dorsal fin base
F in rays - in the dorsal and anal fins the first tw o, three or four finrays are unbranched
(the first very small
and easily m issed), the rem ainder being branched (the last som etim es branched near its base, thus appearing as
tw o, but. counted as one). The first pectoral and pelvic finrays are also unbranched. The counts are given in the
form iii 12 to 15 or i 17. etc.. to indicate unbranched and branched finrays
This elim inates doubts about
the first
often m inute unbranched finray since the first branched finray is always obvious.
B ra n ch io ste g a l ra y s- these support the gili m em ­
brane th at seals the un d ersid e o f the gili cavity. Since
the m em branes on each side are not jo in e d in the middle
(except in the anchovy C etengraulis), they tend to cling to
the gili cover when this is opened up and so must be pulled
inward to expose the branchiostegal rays. H owever; most
clu p eo id s have ab o u t 6 or 7 b ran ch io ste g al rays, so that
the high num ber in round h errin g s (11 to 17) is fairly
obvious (Fig. 11).
brachiostegal rays
underside o f head
Fig. 11
TO CONTINUE
>
14
Gillrakers - unless stated otherwise, these are the
num bers o f gillrakers on the low er h a lf o f the first gili
arch, i.e. on the ceratobranchia!, w hich are much easier
to c o u n t th a n th o s e on th e u p p e r h a lf, i.e . on th e
epibranchial bone. A t the angle o f the arch the curved
bases o f the gillrakers show w hether they belong to the
upper or the lower series (Fig. 12).
Scales - there are no pored lateral line scales
along the flank, but a count o f scales along the midline
o f the flank (in lateral series) is sometimes diagnostic.
Flow ever, since clu p eo id s often lose th e ir scales, this
has b een a v o id e d w h e re v e r p o ss ib le and re fe re n c e
m erely m ade to th e p re s e n c e o f v e ry sm all or v ery
large scales.
Scutes - all clupeoids (except C hirocentrus) have a
pelvic scute im m ediately in front o f the pelvic fins. In
m ost clupeoids the scutes continue forw ard to the gili
o p e n in g to fo rm th e p re -p e lv ic se rie s (in c lu d in g th e
pelvic scute); behind the pelvic scute is a sm all scute
betw een the pelvic fins and this is the first o f the p o st­
pelvic series. Counts are usually given in the form 13
or 14 + 10 to 12, or som etimes 13 or 14 + 10 to 12 = 23
to 26 to in d ic a te p re -p e lv ic , p o s t-p e lv ic and to ta l
scutes (Fig. 13). In a few species scutes are present on
the back, from the origin o f the dorsal fin forward to
the head.
M any other and often small characters are used in
th e k ey s an d d ia g n o s e s, b u t are illu s tra te d at th e
a p p ro p ria te p lace .
W h erev er p o ssib le, non -tech n ical
term s are used, e.g. gili cover, gili opening, upper jaw ,
low er part o f gili arch, e tc., unless specific reference
m ust be m ade to the op ercu lu m , m ax illa, ep ib ran ch ial
g illra k e rs , etc.
S im ilarly, term s like antero d o rsal or
posteroventral have been avoided.
1.5
upper g illrakers
low er g illrakers
//
y /
' ■' ' /
— ÙL 1 7 / / / 7
gili filam ents
dorsal scutes
p re-p elv ic scutes
(pelvic scute black)
p o st-p elv ic scutes
(including small scutes
betw een pelvic fins!
L it e r a t u r e
The taxono m ic, b io lo g ical and fishery lite ra tu re on clupeoid fishes is enorm ous. F or ex am ple, over five
hundred books, papers and reports up to 1984 contain data on merely the 11 species o f clupeoids occurring o ff the
w estern coasts o f A frica. I f this is m ultiplied up for more than three hundred species o f clupeoid fishes in the
w orld, then nearly seventeen thousand books, articles and reports have made some contribution to our knowledge
o f these fishes. In several geographical areas, how ever, attempts have been made to sum m arize these data. These
major reference w orks have been cited in the synonym ies in order to show w here the taxonom y has now changed
and different names are applicable; such w orks often contain valuable biological inform ation, w hich may not have
been significantly updated in the last tw o or three decades. In addition to these w orks, the biologist or fishery
w orker can also use the F A O Id en tific a tio n Sheets in five m ajor fish in g areas. Since id en tifica tio n o f clupeoid
species is simplified by elim ination o f those not found in the area under study, the m ajor reference works and
Identification Sheets are listed here by geographical area.
A.
I N DO- WE S T PACI FIC
1.
General (excluding cool-w ater
species o f Japan and A ustralia)
W ongratana, T., 1980.
S y stem atics o f clupeoid fishes in the In d o -P acific re g io n .
Ph.D.
thesis. Im perial College o f Science and Technology, London, 2 vols. K eys, diagnoses,
line
draw ings, m aps, m eristic counts for 154 species. The clupeoid w o rk er’s vade m ecum . but
still unpublished.
The 24 new species and one new name were published in Jap.J.Ichthyol..
29(4):385-407, 25 figs (1983).
2.
W estern Indian Ocean (Area 51)
FAO Species Identification Sheets for A rea 51. 5 vols, 1984. Clupeoids by P.J.P. W hitehead
and T. W ongratana. K eys, diagnoses and biological and fishery notes for 42 clupeoids (all
illu strated ).
15
SFSA.
Sea Fishes o f southern A frica (in press). A totally revised edition o f J.L.B. Sm ith’s
classic w ork. K eys, d iagnoses, line draw ings
for 18 clupeoid species (all illu strated ),
by
P.J.P. W hitehead and T. W ongratana.
3.
Northern and Central Indian Ocean (Areas 51, 57 - in part)
W hitehead, P.J.P ., 1973. A synopsis o f the clup eo id fishes o f India. J.m ar.b io l.A ssn I n d ia ,
14(1): 160-256. K eys, line draw in g s, literatu re for 68 clup eo id species; still u seful, but in
part superceded by W ongratana’s thesis (see item A 1 above).
4.
E astern Indian ocean. W estern Central
Pacific (A reas 57, 71)
FA Q S p e c ie s I d e n tific a tio n S h e e ts fo r A re a s 57, 7 1 , 4 vols., 1974.
C lu p eo id s by P.J.P.
W hitehead.
K e y s, d ia g n o s e s and b io lo g ic a l and fish e ry
n o te s fo r 40 c lu p e o id s (all
illustrated); much o f it superceded by W ongratana’s thesis (see item A 1 above).
5.
N orthw est Pacific (A rea 61)
W arm -w ater species included in W ongratana’s thesis, but there is no m odern w ork devoted to
the clupeoids o f China, the K oreas and Japan.
Svetovidov, A .N ., 1963. Fauna o f the U SSR . Fishes, 2(1), Clupeidae. Zool. Inst. A cad. Sei. US SR .
new series No. 48, published for the N ational Science Foundation by the Israel Program for
S c ie n tific T ran slan tio n s, J e ru s a le m , 4 2 8 pp. (E n g lis h tr a n s la tio n fro m 1952 R u s s ia n
original). K eys, half-tone figures, line draw ings, literature for 6 clupeoids from A rea 61.
6.
Southw estern Pacific (A rea 81)
N o m odern w ork on all the clupeoids o f N ew South W ales and N ew Zealand. M unro
(1956)
described and figured 40 A ustralian clupeoids, but in many cases the names given m ust
be
updated from W ongratana’s thesis (see item A 1 above).
B.
E A S T E R N PACI FIC
1.
G e n e ra l.
No single modern work on the
2.
N ortheastern Pacific (A rea 67)
clupeoids o f this region.
C lem en s, W .A . & G .V. W ilby, 1967.
F is h e s o f th e P a c ific c o a s t o f C a n a d a , 2 n d e d .,
B ull.F ish.R es.B d C an., (68): 1-443. D e scrip tio n s, p h o to g rap h s, b io lo g ical data for the four
clupeoid species from this area (pp. 99 to 105).
H a rt, J.L ., 1973.
P a c ific fish e s o f C a n a d a .
B ull.F ish.R es.B d C a n ., (180):l-740. K e y s ,
d escrip tio n s, line draw in g s, b io lo g ic a l d a ta and b ib lio g ra p h y (e x te n s iv e ) fo r th e fo u r
clupeoid species from this area (pp. 94-106). One o f the best works o f its kind.
3.
E astern Central Pacific (A rea 77)
(C alifo rn ia)
M ille r, D .J. & R .N . L e a , 1 9 7 2 .
G u id e to th e c o a sta l m arin e fish e s o f C a lifo rn ia .
F ish B u ll.C alif.. (157): 1-235. Illu stra ted keys, b rie f d escrip tio n s and d istrib u tio n notes for
the 12 clupeoids o f this area (pp. 54-57).
(M exico to Panama)
A non., 1976. C a ta lo g o de p e ce s m arinos M exicanos.
S e c re ta ria de In d u stria y C om ercio,
Instituto N atio n a l de Pesca, M exico, 462 pp. K eys, distribution notes and photographs for
8 Pacific coast clupeoids o f M exico (pp 67-70, figs 60-75).
M eek, S.E. & S-F. H ildebrand, 1923.
The marine fishes o f Panama. Pubis Field M us.nat.H ist.
(zo o l.S er.), 15(215), Pt 1:1-330. K e y s, d e s c rip tio n s and a few illu s tra tio n s o f 22 P a c ific
coast clupeoids o f Panam a (pp. 179-214). N om enclature outdated, but useful in conjunction
w ith Peterson (see below).
P e te rso n , C .L ., 1956.
Observations on the taxonom y, biology, and ecology o f the engraulid
and clupeid fish es o f the G u lf o f N icoya, C o sta R ica. B u ll-in te r-A m .tro p .T una C o m m n,
1(5): 137-280.
K eys, d istrib u tio n and biolo g ical notes for 23 P acific co ast clupeoids o f
Costa Rica. The best general w ork for this region, but lacks illustrations.
16
(E cuador to Peru)
C obo, M. & S. M assay, 1969. L is ta de Ios peces m arinos del Ecuador. B oln cient.tecn.Inst.
n a c .P e se .E c u a d o r. 2(1): 1-68. L ist o f 22 clupeoids (7 illustrated) from Ecuador co asts; the
n o m e n c la tu re u se fu lly u p d ate s th a t o f M eek & E lildebrand (Panam a) and Elildebrand (Peru).
Elildebrand, S.F., 1946. A d escrip tiv e catalo g u e o f the shore fish es o f Peru. B u ll.U .S .n atn .
M us., (189): 1-530.
K eys and d escrip tio n s o f 14 clu p eo id s (6 illu stra ted ) from P eru v ian
coasts (pp. 80-105). Still the most useful w ork for this region.
4.
Southeastern Pacific (Area 87)
M uch o f the older Chilean literature is assembled by D elphin (1901:39-42 for clupeoids) and
by Fowler (1945:1-6 for clupeoids), but there are no keys and the nom enclature is outdated;
the same applies to Mann (1954).
B o re , R. & F. M a r tin e z , 1981.
C h ile a n fis h e rie s re s o u rc e s c a ta lo g u e . C o r p o r a c ió n de
F o m en to de la P ro d u cció n , Instituto de F o m en to P e s q u e ro , C h ile , u n p a g e d . D e s c rip tio n s
and biological and fishery notes for 4 clupeoids (all w ith excellent colour photographs) o f
this region.
D e B uen, F., 1958.
P e c es de la superfam ilia clupeoidae en las aquas de Chile. R evta Biol.
M ar.. 8( 1-3 ):83-110. K eys and d escrip tio n s o f all five clupeoids (no illu stratio n s) o f this
region.
L e ib le , M . & E. A lveal, 19 8 2 .
C a ta lo g o de p e c e s q u e h a b ita n la s a g u a s co steras y
d u lcea cu ico las en la p ro v in c ia de C o n cep tio n . P o n tific ia U n iv ersid ad C ató lica de Chile,
D epartam ento de B iología y Tecnología d e l M ar, Talcahuano, 104 p. K eys, descriptions,
biological and fishery notes for four clupeoids (all illustrated) o f this region (pp. 12-21).
C
W E S T E R N ATLANTIC
1.
G eneral. N o single m odern w ork on the clupeoids o f this region.
2.
N orthw estern A tlantic (A rea 21)
FW NA Fishes o f the w estern N orth A tlantic, 1964.
Sears Foundation for M arine Research,
Yale U niversity, M em oir 1 (3), 630 p. K eys, diagnosis and biological and fishery notes for
79 clupeoids (all illustrated) for this region and the northern part o f A rea 31 (pp. 148-454,
authors S.F. E lildebrand, L.R . R ivas and R .R . M iller).
In p a rts o u td a te d , b u t still an
e x cellen t guide.
Liem , A .Fl. & W .B. Scott, 1966. Fishes o f the A tlantic coast o f Canada. B u ll.F ish.res.
Bd C an., (155): 1-485. K eys, descriptions and biological and fishery notes for 7 clupeoids
(all illustrated) o f this region.
3.
W estern Central A tlantic (A rea 31)
FWNA, 1964 (see item C 2 above), which covers also the northern part o f this area.
C erv ig ó n , F., 1966. Los peces de V enezuela, 2 vols, F ondo de C u ltu ra C ie n tífic a , C aracas.
K eys and descriptions for 28 clupeoids (12 illustrated) from this region (pp. 114-149 and
925-926).
W hitehead, P.J.P ., 1973. The clupeoid fishes o f the G uianas. B ull.B r.M us.nat.H ist.(Z ook).
Suppl. 5:1-227. K eys and descriptions o f 37 clupeoids (all but one illustrated) from the
southern part o f A rea 31.
F A Q Species Identification Sheets for A rea 3 1 , 6 v o ls, 1978. Clupeoids by P.J.P. W hitehead.
K eys, diagnoses and biological and fishery notes for 31 clupeoids o f this area (all illustrated).
4.
Southw estern A tlantic (A rea 41)
Figueiredo, J.L. & N.A. M enezes, 1978. M anual de peixes m arinhos do sudeste do B rasil, 2(1).
M useu de Z oología, U niversidade de Sào Paulo, B razil. K eys and descriptions o f 21
clupeoids (all but one illustrated) from this region (freshw ater species not included) (pp. 21
to 32).
17
R ingueleti R .A ., R.H. A ram buru & A.A. A rám buru, de, 1967.
Los peces A rgentinos de agua
d u lce. C om isión de In v e stig a c ió n Científica, Buenos A ires, 602 pp. Keys and descriptions
o f 7 freshw ater or euryhaline clupeoids (4 illustrated) o f A rgentina (pp.52-65).
M enni, R .C ., R .A . R in g u elet & R .A . A rám b u ru . 1984.
P eces m arin h o s de la A rg en tin a y
U ru g u a y . E d ito rial H em isferio Sur. B uenos A ires. 395 pp. R eco rd s o f 18 c lu p eo id s (13
figures o f this region (pp. 107-110)
B ellisio , N .B ., R .B . L ópez & A . T o rn o . 1979.
P e c e s m a rin o s P a tá g o n ic o s. S ecretaria de
Estado de In te r e s e s M a r ítim o s , M inisterio de E c o n o m ía , Buenos A ires, 279 pp. D escrip­
tions and biological and fishery notes for 2 clupeoids (both illustrated) o f this region (pp.
53-58).
D.
EASTERN
ATLANTIC
1.
G eneral. N o single m odern work on all the clupeoids o f these
2.
N ortheastern A tlantic and M editerranean (A reas 27.
two areas.
37)
Svetovidov, A.N., 1963 (see item A 5 above). K eys, descriptions and a great deal o f biological
and fishery data for 18 m arine and fresh w ater clu p eid s (all illu stra ted ) o f the n o rth east
A tlantic and Black Sea.
B anarescu, P., 1964. Fauna R epublicii Populare Romine. P isc es-O steich th y e s (pesti ganoizi si
o so si), 13, E d itu ra A c a d e m i e i R ep u b lic ii P o p u la re R o m in e. B u c h a re st. 959 pp. K eys,
descriptions and biological notes for 8 clupeoids (all illustrated) o f the Black Sea (pp. 223251).
W heeler, A .C ., 1978. Key to the fishes o f northern E urope. Frederick W arne. London. 380 pp.
K eys, d escrip tio n s, and b rie f no tes for 6 clu p eo id s all illu strate d ) o f no rth ern E uropean
coasts (pp. 66-71)
F A Q Species Identification Sheets for Area 37. 2 vols., 1973. C lu p eo id s by A .N . S vetovidov.
Descriptions and biological and fishery notes for 6 clupeoids (all illustrated) in this area.
C L O FN A M C h eck list o f the fish es o f the n o rth -eastern A tlan tic and M e d ite rra n ea n . 2 vols..
1973.
C lupeoids m ainly by A .N . S v eto v id ov , also P.J.P . W hitehead & A. B en-T uvia.
Virtually complete com pilation o f taxonom ic literature, including that for 13 clupeoids o f
the northeastern A tlantic and M editerranean (pp. 99-112).
FNAM . Fishes o f the north-eastern A tlantic and M editerranean. 1st vol.. 1984. Clupeoids by
P.J.P. W hitehead. K eys, diagnosis and notes on biology for the 14 clupeoids (all illustrated)
for the CLO FN AM area.
3.
Eastern Central A tlantic (A reas 34 and
47 in part)
F A Q Species Identification Sheets for A reas 34. 47 (in part). 6 vols, 1981. Clupeoids by P.J.P.
W hitehead.
K eys, d iagnosis and biolo g ical and fishery notes for the 11 clu p eo id s (all
illustrated) o f this area.
C LO FE TA . C h eck -list o f the fishes o f the eastern tro p ical A tlan tic (in press). C lupeoids by
P.J.P . W hitehead. V irtually co m plete co m p ilatio n o f taxonom ic litera tu re , in clu d in g th at
for the 11 clupeoids o f this area.
4.
Southeastern A tlantic (A rea 47)
All clupeoids are covered by the two works above (item D 3)
E.
FRESHWATERS
1.
A frica (Area 01)
CLOFFA. C hecklist o f the freshwater fishes o f A frica. 1st vol., 1984. Clupeoids by M. Poll.
G.G. Teugels & P.J.P. W hitehead. V irtually co m plete co m p ilatio n o f tax o n o m ic literatu re
for 39 freshw ater or euryhaline clupeoids o f A frica (pp. 41-56)
2.
North and South America (Areas 02, 03)
N o sp e c ia l w o rk , b u t fr e s h w a te r or e u ry h a lin e c lu p e o id s in c lu d e d in th e lite r a tu re c ite d
alre a d y .
TO CONTINUE
■>
18
3.
Asia (Area 04)
A ll tro p ical or su btropical fresh w ater or eury h alin e clu p eo id s included by W ongratana (see
item A 1 above).
4.
Europe (Areas 05, 07)
All freshwater or euryhaline clupeoids included by Svetovidov (see item A 5 above; see also
item D 2).
5.
A ustralia (Area 06)
No special work, but some species included by Wongratana (see item A 1 above).
1.6 A cronym s
The follow ing six m ulti-authored w orks are more readily re c o g n ized by their titles than by their editor(s) or
even th e ir authors.
To save space, how ever, esp ecially in the synonym ies, th e s e w o rk s are c ite d by th e ir
acronym s.
This seem s m ore im m ed iately in fo rm ativ e, esp ecially w hen th ere may be num erous o th er papers by
the same author. The six w orks are:
CLOFETA (in press). C heck-list o f the fishes o f the eastern tropical A tlantic. U nesco, Paris.
J.-C . Quéro.
Edited by
C L O FFA , vol. 1, 1984. C h eck -list o f the fresh w ater fish es o f A fric a . M usée R oyale d ’Afrique C e n tra le ,
B russels & ORSTOM, Paris. Edited by J. Daget, J.P. Gosse & D.F.E. Thys van den A udenaerde.
C LO FN AM , 1973.
C h ec k -list o f the fishes o f th e n o rth -eastern A tla n tic
U nesco, Paris. Edited by T. M onod & J.C. Flureau.
and o f the M e d ite rran e an .
FN A M , vol. 1, 1984. F ish es o f the n o rth -ea ste rn A tlan tic and M e d ite rra n e a n . U n esco , P aris. E d ited by
P.J.P. W hitehead, J.C. H ureau, J. N ielsen & M .-L. Bauchot.
FW N A, part 3, 1964. Fishes o f the w estern N orth A tlantic. Sears Foundation for M arine Research, Yale
University. Edited by H.B. Bigelow et ah
SFSA (in press). Sea fishes o f southern A frica. G rahamstown. Edited by P. Heemstra.
A c k n o w le d g e m e n ts
Taxonomy is a creative and not merely an applied science, but only in its application does it finally justify
the labour expended.
It has been a privilege, therefore, to participate in w hat m ust now qualify as one o f the
m ost im portant program m es for the application o f fish taxonom y.To an overall appreciation o f the role played by
D r W alter Fischer (FAO, Rom e) in initiating the Catalogue series and related w orks in this program m e, I m ust add
three personal debts: first for persuading me that a synopsis o f clupeoid fishes w as not prem ature, second for
doing so w ith such irresistible drive and enthusiasm , and third for providing his considerable editorial skills during
the g estatio n o f th is w ork.
N o less a d ebt is ow ed to D r T h o sap o rn W o n g ratan a (C h u lalo n g k o rn U n iv e rsity ,
Bangkok), w ho graciously allowed me to use all his draw ings o f Indo-Pacific clupeoids - the finest such drawings
ever made; his revision o f these fishes (w ith occasional differences o f opinion) underpins the taxonom y adopted
here. The rem aining draw ings, except for a few by m yself, w ere occasionally taken direct from the literature, but
mostly w ere carefully redrawn from various sources by M r Oliviero Lidonnici (F A O , Rome), w ho also redrew all
my small sketches; M r Paolo Lastrico is also thanked for his valuable assistance in various aspects o f the artwork.
Help w ith fishery statistics, types o f gear and modes o f utilization w as kindly given by F A O staff, in particular
Mrs Fiorenza de L uca, M r J. Prado and Ms Helga Josupeit. Ms Claire Cuerden and her staff in the F A O Fisheries
B ranch L ib rary , and M s G loria A . Soave gave assistan ce w ith the b ib lio g rap h y .
F in ally , the ty p in g o f the
m anuscript (literally hand-w ritten) and the com plex setting o f text and figures on the pages, w ere undertaken w ith
great skill, speed and rem arkable cheerfulness by M rs Giulia S ciarap p a-D em u ro (FAO, Rome).
click fo r n e x t p a g e
click fo r p re v io u s p a g e
19
2.
O R D E R CLUPEIFORMES- S U B O R D E R
CLUPEOIDEI
M oderate-sized o r small o r very small fishes (2 to 100 cm standard length) w ith no spines in the fins, the
dorsal fin single and short (11 to 23 finrays) and usually near the m idpoint o f the body (further back in
C hirocentridae and some Pristigasteridae, the latter including
Raconda w ithout a dorsal fin); the Pelvic fins small
(6 to 10 finrays), a little before, under or a little behind the dorsal fin
base
(but absent in some Pristigasteridae)!
the anal fin usually short or m oderate (10 to 36 finrays), but longer in some Pristigasteridae (34 to 93) and some
Engraulididae (14 to 81, or about 100 in C oilia)! the cau d al fin forked (except rhom boid in C oilia).
The body is usually fusiform , som etimes alm ost round in cross-section (some D ussum ieria, E trum eus, also
E ngraulis), but more often com pressed, som etim es highly com pressed (C hirocentridae, some Pristigasteridae).
Typically, there is a pelvic scute w ith ascending arm s ju s t in front o f the pelvic fins (absent in Chirocentridae!
W -shaped in the D ussum ieriinae, and a series o f sim ilar scutes in front o f the pelvic fins and behind them , but
absent in the D ussum ieriinae, some Pellonulinae, E ngraulis, some specimens o f Stolephorus purpureus and all N ew
W orld Engraulididae! the scutes do not reach forward to the isthm us in
some
Engraulididae. Fossil clupeoids (e.g.
the Eocene K nightia) had a series o f scutes from the occiput to the dorsal fin origin! such a complete dorsal series
occurs in the Pellonulinae (Potam alosa. Elyperlophus) and the A losinae (E thm idium ), w hile the D orosom atinae
(Clupanodon). Clupeinae (H aren g u la and others) and Pristigasteridae (Pristigaster) include species w ith one, two or
a few pre-dorsal scutes! in the Engraulididae, some Stolephorus and all other Indo-Pacific genera have a single
spine-like scute ju st before the dorsal fin.
ribs
pelvic fin
w -shaped
pelvic scute
pelvic bone
pelvic fin
pelvic scute
D ussum ieria. E t r u m e u s ,
S p ra te llo id e s. Jenkinsia
dorsal finrays
pre-dorsal scutes
P o ta m a lo sa . H y p e rlo p h u s.
E th m id iu m
spine-like
pre-dorsal scute
T h ry ssa . S etip in n a, etc.
The m o u th is sm a ll, w ith th e lo w e r ja w d eep an d tr ia n g u la r in th e C h iro c e n trid a e , C lu p e id a e and
Pristigasteridae, but slender and long in m ost Engraulididae. The p re-m ax illae are trian g u lar (b u t re ctan g u lar in
D ussum ieriinae) and the m axillae usually have an anterior (first) and posterior (second) supra-m axilla along the
20
upper edge.
Sm all conical te eth are ty p ically p resen t in the ja w s and on the vom er, p alatin es and endo-and
ectopterygoids (i.e. the ro o f o f the m outh), but some or all may be absent, or the jaw s may bear canine teeth
(C hirocentridae, also C hirocentrodon o f the P ristigasteridae and both
L ycengraulis and L ycothrissa o f the
Engraulididae). The gut may be short (carnivores) or long and coiled (phytoplankton feeders, filter feeders) and in
some the stomach is m uscular like a gizzard (D orosom atinae; partially so in some Clupeinae); food is collected in
a bolus by pharyngeal pouches in the D orosom atinae and some Clupeinae. A swim bladder is present, sometimes
double-cham bered (some E ngraulididae), w ith a pneumatic duct joined to the oesophagus or stom ach blind-sac; in
the Pristigasteridae (Ilisha) some species have one or a pair o f post-coelom ic tubes extending from the hind end o f
the sw im bladder back into the muscles o f the body.
1st
supra-m axillae
2nd
1st
supra-m axillae
2nd
E n g rau lid ae
C lupeidae
sw im bladder
C h iro cen tro d o n
extension o f swim bladder
beyond body cavity
p n eu m alic d u c t (connecting
sw im bladder w ith stomach)
A lm o s t a ll s p e c ie s h a v e a c o m p le te
covering o f cycloid scales on the body (mostly
lo s t in T h ra ttid io n an d S ie r ra th r is s a o f th e
Pellonulinae), w hich are freq u en tly d eciduous;
sm all scales occasio n ally cover the bases o f
the dorsal, anal and/or cau d al fins, and one or
sometimes several axillary scales lie above the
bases o f the first pectoral and pelvic finrays.
There is no lateral line canal w ith pored scales
a lo n g th e fla n k s (o c c a s io n a lly one or tw o
behind the gili opening).
A b ran ch in g and
mainly cutaneous sensory canal system covers
th e to p and sid e s o f th e h e a d ;
th e su p ra­
o rb ital, in fra-o rb ital, p re -o p ercu lar and pterotic canals all meet in the re cessu s la te ra lis, a
sp e c ia l c h a m b e r c h a ra c te ris tic o f c lu p e ifo rm
fishes, its inner wall being a m em brane sealing
th e p e rily m p h a tic sp ace th a t s u rro u n d s th e
inner ear.
pectoral axillary
scales
pelvic axillary
scales
21
Two im portant
the C lupeoidei:
(a)
(b)
internal
features
h e a d can al
(w ith p o re &
neuromast)
characterize
A pair o f fine tubes from the front o f
the sw im bladder penetrate the skull and
expand inside bony capsules (bullae) in
the pro-otics and (except in Sprattus and
C lupeonella) also in the pterotic bones.
A m em brane w ithin each pro-otic bulla
separates gas (from the sw im bladder)
from perilym phatic liquid (surrounding
the
inner ear). This
system , together
w ith
the head canal
system and the
recessus la te ra lis.
probably
m onitors
inform ation necessary for schooling and
other swimming activities, as also detec­
tion o f predators and hazards.
recessus
lateralis
ut r i cul ar
ma c u l a
pr o- ot i c bul l a
"
endolimph
swimbladder
perilymph
coelom ic ducts
dorsal view (schem atic) o f connexions betw een
sw im bladder, inner ear and canals on head! black-gas!
stippled- e n d o - and perilym oh; wavy lines - w ater
The small bones supporting the caudal
fin
have a ch aracteristic arrangem ent
and series o f fusions
betw een particular
e le m e n ts .
utricular
m aculae
Most species o f Clupeoidei are m arine, coastal
and sc h o o lim g fishes, but some enter brackish- or
freshw aters and some live perm anently in fresh­
w ater (rivers or lakes). In this catalogue are given
over 300 species o f 80 genera, placed in 4 families!
in most earlier literature the fam ily Pristigasteridae
is given as a subfamily o f the C lupeidae, but recent
w ork suggests that it is distinct (G rande, in press).
PU1 - U N I
fused
V - r
~—
endolmph
fenestra
elastic th re a d
perilym ph -
p r e - c o e lo m ic d u c t
fro m s w im b la d d d ïs
gas
m e m b ra n e o f
p ro -o tic b u lla
lateral view (cross-section) o f connexion betw een prootic bulla (below) and inner ear (above)! variations in gas
presscure in the sw im bladder are transm itted to the
m em brane o f the pro-otic bulla! a fine elastic thread
signals m ovem ents o f the m em brane to the inner ear
typical clupeoid caudal fin skeleton! the first uroneural
(U N I) and first pre-ural v e rte b ra (PU1) are fu se d ! th e
first o f the six hypural plates (H P1) is free! the second
(HP2) is fused at its base to the first u r a l vertebra (U l)!
and th e h y p u ra l (P H ) is free at its base
K ey to the C lu p eo id F a m ilies
la .
A r t i c u l a t i o n o f lo w e r ja w u n d e r o r o n ly
ju s t behind eye, low er ja w deep (Fig. 1)
low er ja w
deep
a rtic u la tio n
o f lo w er ja w
click fo r n e x t p a g e
click fo r p re v io u s p a g e
22
canine teeth
2a.
No scutes along belly (even pelvic
sc u te
ab sen t);
tw o f a n g - lik e
canine teeth in upper jaw , pointing
forward (Fig. 2); body highly compressed , elongate
. Chirocentridae
C hiro cen trid ae
2b.
Scutes usually presen t along belly
(pelvic scute alw ays p resen t) (Fig.
3);
c a n in e t e e t h r a r e , n e v e r
pointing directly forw ard in upper
jaw ; body usually oval or round in
cro ss-sectio n
3a.
A nal fin m o d erate, less than
30 finrays (unless m outh infe­
rior, e.g. D orosom a) .................. C lu p eid ae
3b.
A n a l fin lo n g , a t le a s t 30
finrays; low er ja w projecting,
m outh d irected m ore or less
upward
Pristigasteridae
p elvic scute
p re-p elv ic scutes
lb.
C lu p eid ae,
p o st-p elv ic scutes
P ristig a ste rid a e
F ig . 3
A rtic u la tio n o f lo w e r ja w w ell b eh in d
eye, low er ja w usually slender (Fig. 4);
snout ‘pig-like’ and projecting, low er jaw
‘underslung’ ......................................................... Engraulididae
low er ja w
slender
a rticu latio n o f
lower jaw
Engraulidae
2.1
Im a
FAM ILY CHIROCENTRIDAE
Fig- 4
C H IR O C
FAO Names : En - W olf-herrings.
D iagnostic Features : E lo n g ate, highly co m p ressed , silvery fish es resem b lin g the C lupeidae (h errings,
sardines), but w ithout scutes along the belly. Head strongly com pressed, w ith two fang-like canine teeth pointing
forward in the upper jaw , a series o f canine teeth in low er ja w (larger at the front). Dorsal fin short, set well
behind m idpoint o f body; pectoral fins set low; pelvic fins very small; anal fin longer than dorsal fin, beginning
below about dorsal fin origin; caudal fin deeply forked. Scales num erous, sm all, usually lost; no lateral line down
flank. Back bright blue (fading to grey), flanks bright silver.
Biology, H abitat and D istribution : C h iro c e n trid s are m arin e c o a sta l fish e s w id e ly d is trib u te d in the
warmer parts o f the Ind o -P acific region, from the w estern Indian Ocean (Red Sea, East A frica south to D urban) to
the w estern P acific (Jap an , the P h ilip p in e s south to n o rth ern A u stralia ). They are pelagic inshore predators on
sm all fishes and are said to ex h ib it a sort o f feeding frenzy am ong sm all h errin g s and anchovies du rin g night
fishing w ith lights. Few studies have been made o f breeding, but like most other clupeoids they probably scatter
pelagic eggs from w hich planktonic larvae hatch. A m ong the largest o f the clupeoids, they reach 100 cm standard
len g th (Fow ler, 1959:30, gave 3.66 m, but this is not correct; followed by Smith, 1953:87). I f they are schooling
fish es, then the schools are p robably o f m od erate size (at least com pared to schools o f o th er clu p eo id s). No
special fisheries exist and catches are small (50 083 tons in 1983).
A single genus. C hirocentrus, w ith 2 species.
23
C H IR O C C hiroc
Chirocentrus Cuvier, 1816
C h iro cen tru s C u v ie r, 1816, R ègne anim al, 1st ed., 2:178 (type: C lupea dorab F o rssk ál, 1775) (fo r co rre c t
dating o f Cuvier, see Whitehead, 1967, emended by Cowan, 1969 to 7 December 1816).
Biology, Habitat and Distribution : See family.
Species ! In the past, m ost w orkers have assumed a single species, C. dorab, occasionally w ith a second
species CL h y p selo so m a . W h iteh ead , B o esem an & W heeler (1966:27) showed that Bleeker’s hypselosom a was C.
d o ra b , w hile L u th er 1968) resu rrected an early Swainson n a m e , nudus, for a second species in Indian waters.
A lthough variations in body depth are not yet satisfactorily accounted for, m odern w orks reco g n ize two species:
C dorab (Forsskál, 1775), In d o -W est Pacific
C. nudus Swainson, 1839, Indo-W est Pacific.
C h irocen tru s dorab (F o rs s k á l 1775)
Clupea dorab Forsskál, 1775,
C H IR O C C h iro c
1
P e se r.anim .: xiii, 7 (Djedda and M ocha ,Red Sea).
Synonym s : C lu p ea den tex
S ch n eid er,
1801:428; E so x ch iro cen tru s L apède, 1803:296; C h iro cen tru s
h y p se lo so m a B leeker, 1852:71;
C h ir o c e n tr u s d o ra b - W hitehead et a h , 1966:27 (Bleeker's CL h y p s e lo s o m a );
Luther, 1968:194; W h iteh ead , 1973b: 167, fig. 2; SFSA, in press (southern Africa).
FAO Names : E n - D orab w o lf-h errin g .
D ia g n o stic F ea tu res : The slightly shorter pecto­
ral fin (11 to 13% o f stan d ard len g th ; cf. 13 to 18%)
and the black marking o f the upper part o f the dorsal
fin are the only satisfacto ry ch aracters sep aratin g this
species from CL nudus; there is also some black on the
anterior part o f the anal fin. V ariations in body depth
may represent sexual dim orphism , but this needs study.
See CHIROC Chiroc 1, Fishing Areas 57, 71 and also 51.
G eographical D istribution : Probably throughout
the w arm er coastal w aters o f the Indo-Pacific, from
the A rab G ulf, R ed Sea, east A frican co ast south to
D urban, eastward to Japan, the Philippines and south to
n o rth ern A u stralia. H ow ever, it is not known to w hat
extent CL nudus may contribute to these records.
H a b ita t an d B io lo g y :
P e la g ic ,in s h o re fish e s;
feeding mainly on small fishes, but perhaps also crusta­
ceans, etc.; no precise inform ation on breeding.
S ize :
To about 100 cm o f standard length.
Interest to F isheries
Separate statistics for
Chirocentrus (alm ost certainly including a proportion o f
C. n u d u s) are rep o rted by T an zan ia, P ak ista n , In dia,
Thailand, Indonesia,
M a la y s ia , S in g a p o r e a n d th e
Philippines, w ith a total o f 50 083 tons in 1983. The
24
catches in Palk Bay and around Ram esw aram I in the G u lf o f M annar (southern India) are the only ones where the
two species o f Chirocentrus w ere positively separated (Luther, 1968); G dorab contributed only 20% to the catch,
the rest being G nudus. C aught w ith gillnets, seines, shallow trawls and traps. M arketed fresh or frozen.
Local Nam es : INDIA: D orab;
Calcutta: Chela, K handa, Samudrik; RED SEA: D orab, Sam ak abu sayf.
Literature : L u th e r (1968 - sep aratio n o f G
synonym s, references, fig.).
dorab from G
C h irocen tru s n u d u s (Swainson, 1839)
n u d u s);
W h iteh ead (1973 - key to species,
C H IR O C
C h iro c 2
C h iro cen tru s ru ssellii S w ain so n , 1838. N a t.h is t.a n im .. 1:289 (on Wallah o f R ussell, 1803) (nomen oblitum ).
Chirocentrus nudus Sw ainson, 1839, Ibid, 2:294 (also on Walla ).
S yn on ym s :
C h iro cen tru s dorab by m any au th o rs; C h iro c e n tru s nudus - Luther, 1968:194; W h ite h e a d ,
1973b: 168, fig. 3; Talwar. 1976:324.
FAO Names :
E n - W hitefin
w o lf-h e rrin g .
D iagnostic Features : The slightly lo n g er p ecto ­
ral fin (13 to 18% o f standard length; cf. 11 to 13%)
and the absence o f black m arkings on the dorsal fin tip
are th e o n ly sa tis fa c to ry c h a ra c te rs se p a ra tin g th is
species from the otherwise virtually identical G dorab;
also, there is no black on the anterior part o f the anal
fin. A gain, v ariatio n s in body d epth need m ore study.
See CHIROC Chiroc 2, Fishing Areas 57, 71 and also 51.
Geographical D istribution :
P robably sim ilar to
that o f G dorab, but not alw ays distinguished from that
species.
C e rta in re c o rd s are fro m M o m b a sa , th e
“G u lf’, India (east and west coasts), Sri Lanka, the IndoA ustralia archipelago and Canton.
H abitat and B iology :
for C. dorab.
Size :
A p p a re n tly th e sam e as
To about 100 cm standard length.
In te r e st to F ish e r ies
A s for C. dorab. It is o f
interest that G nudus predom inated (80%) in the C hiro(L u th e r, 1968), and co m p rised the to ta l C h iro cen tru s
catch o ff the Orissa coast (Talw ar, 1976); in both cases
these w ere gillnet fisheries.
Local Names :
As for C. dorab.
Literature : L u th er (1968 - sep aratio n from G d o rab );
references, fig.); Talw ar (1976 - diagnosis).
W h ite h e a d (1973b - key to sp ecies, synonym s,
click fo r n e x t p a g e
click fo r p re v io u s p a g e
25
2.2
F A M IL Y
CLUP
C LU PEID A E
FAO Nam es : En - H errin g s, S ardines.
D ia g n o s tic F e a tu r e s : Typically, clupeids are fusiform fishes, oval in cross-section, w ith a com plete series
o f scutes along the belly (pelvic scute alw ays present)! the mouth is term inal, there are 2 supra-m axillae, and the
jaw teeth are small or m inute; the dorsal fin is short and near the m idpoint o f the body. The pelvic fins are just
in front of, below or ju st behind the dorsal fin base, and the anal fin is short and its origin is well behind the last
dorsal finray; the scales are adherent and o f m oderate size (about 40 to 50 in lateral series).
Elowever, there is great variation in body shape and depth (round bodied to strongly com pressed and deep),
scutes (some or all absent along belly, but a few or a complete series o f pre-dorsal scutes occasionally present),
m outh shape (lo w er ja w p ro m in en t to m outh fully in fe rio r in the g izzard shaps), su p ra-m ax illae (one or both
absent), teeth (absent in some, canines in others), and scales (deciduous in some, minute in others). The fam ily as
at present constituted is probably an artificial assem blage, being defined largely on the shared absence o f those
special characters that define the other clupeoid families.
Biology H abitat and D istribution : Clupeids are typically marine coastal and schooling fishes found in all
seas from 70° N to about 60° S, feeding on small planktonic anim als (m ainly crustaceans), form ing large schools and
scattering large num bers o f p elag ic eggs from w h ich p lan k to n ic larvae hatch. A dults are usually 10 to 20 cm
standard length. It is m ainly the co o l-w ater genera (C lupea. S ard in a. S ard in o p s. S p rattu s) th a t dom inate the
clupeoid catches.
A s w ith m orphology, so there is a great range in the biology and ecology o f clupeids. Some enter freshwater
to feed, some are anadrom ous (shads), and some live perm anently in freshw ater (W est A frican pellonulines), some
are partial or full-tim e filter-feeders (e.g. gizzard shads), some are predators on fishes (and probably form only
loose and small schools as adults), and some produce only two hundred eggs or less (pygmy species) or attach their
eggs to the substrate (C lupea). Great variation is found in size, from the shad Tenualosa ilisha (to about 60
cm) to
certain pygmy riverine species w hich may be mature at under 2 cm (Sierrathrissa. T h ra ttid io n , etc.).
Interest to Fisheries : Individually, few o f the w arm -w ater species dom inate clupeoid catches (exceptions
are Sardinella aurita. S^ longiceps and related species), but m ulti-species clupeid fisheries may account for as
much as a third o f the total fish catch in some areas.
Some 180 clupeid species are listed here, placed in 56 genera, but more species will probably be described!
the true total may be close to two hundred, and the number o f genera could rise to about sixty.
The clupeids can
be conveniently placed in 5 subfam ilies,although w ork in progress w ill probably show th at the present
arrangem ent is artificial and does not reflect true relationships.
K ey to the C lu p eid S u b fa m ilies
a
Pelvic scute W -shaped (Fig. la); no
o th er sc u te s along belly.........................
D ussum ieriinae
W -shaped pelvic scute
lb.
P elvic scute w ith ascen d in g arm s
(F ig . lb )! sc u te s u su a lly p re s e n t
before and behind pelvic fins
p elvic scute
a. D u s s u m ie riin a e
pelvic fin
b. o th e r su b fa m ilie s
click for n e x t p a g e
click fo r p re v io u s p a g e
26
2a.
2b.
U pper ja w rounded w hen
from front (Fig. 2a)
seen
3a.
A nterior as w ell as posterior
supra-m axilla
pre­
sent (Fig. 3a); worldwide . . . C lupeinae
3b.
No anterior supra-m axilla
(Fig. 3b); not in N ew
W orld .......................................
rounded,
no
Pellonulinae
U pper ja w w ith a d istin c t
m edian notch or cleft w hen
seen from front (Fig. 2b)
4a. L ow er jaw norm al, m outh
term inal (Fig. 4a); last
dorsal finray n o m ial.....................
4b.
Alosinae
L ow er ja w flared o u t­
w ard, mouth usually infe­
rior (Fig. 4b); last dorsal
finray
filam en tou s
in
many species ...................
Dorosomatinae
1st (anterior)
supram axilla
2nd (posterior:
supra-m axilla
only 2 n d (p o ste rio r)
su p ra -m a x illa
a.
A losinae
2 .2.1
b.
b. D o ro so m a tin a e
F ig -4
SU BFA M IL Y DUSSUMIERIINAE
FAO Names :
En - Round herrings.
Diagnostic Features : Small or m oderate-sized herring-like fishes im m ediately distinguished from all other
clupeids by their W -shaped pelvic scute and absence o f any other scutes (some pellonulines have only a few or no
ventral scutes, but the pelvic scute always has lateral arms - see p.)
Biology, Habitat and Distribution : R ound herrings are m arine coastal and schooling fishes w idely
distributed in the Indo-Pacific region, from the w estern Indian O cean (the “G u lf’ to Cape Town) to the w estern
Pacific (Japan to northern A ustralia, also Samoa); in the southern part o f the eastern A tlantic (Cape Town north
to Cunene R iver); in the w estern A tlantic (N ew York to Guyana); and in the eastern Pacific. They are pelagic
inshore fishes, feeding m ainly on Zooplankton. Like other clupeoids, they scatter pelagic eggs from w hich
planktonic larvae hatch. The largest round herrings
(E trum eus) reach 25 cm standard length; the sm allest are
mature at about 5 or 6 cm.
Interest to Fisheries : All are schooling fishes and thus exploited by fisheries, especially in Japan, South
Africa and Indonesia. Recorded catches in 1983 were 150 578 tons.
Remarks : In earlier w orks, the round herrings have been placed in a separate fam ily. D ussum ieriidae (e.g.
by W hitehead, 1963; given as Stolephoridae by F o w ler, 1941:561). There
are tw o pairs o f genera, w hich can be
placed in separate tribes, and 12 species.
27
K ey to the Genera :
la.
B ran ch io steg al rays num erous
(11 to 18) (Fig. 1); p r e ­
m a x illa e rectangular
(Fig.
2a)
T rib e D u ssu m ieriin i
2a.
2b.
lb.
Pelvic fins under dorsal
fin base; anal f in r a y s 14
to 18; isthm us pointed
an terio rly
(Fig.
3 a );
I n d o - P a c ific o n l y
D u s s u m ie r ia
b r a n c h io s te g a l ra y s
Pelvic fins behind d or­
sal fin base; anal f i n ­
rays 10 to 13; isthm us
w ith
‘sh o u ld ers’
an te­
riorly (Fig. 3b ); w orld­
w ide....................................................Etrumeus
u nderside o f head
T rib e D u ssu m ie riin i
Branchiostegal rays few (6 or
7); p re - m a x illa e tr ia n g u la r
Fig. 2b) ....................T r ib e
S p r a t e l lo i d in i
3a.
3b.
F ia i
tr ia n g u l a r
r e c t a n g u la r
Indo-Pacific only; 2 supram a x illa e (F ig . 4a) — S p r a te llo id e s
W e s te rn A tla n tic
1
su p ra-m a x illa
4b)
o n ly ;
(Fig.
J en k in sia
a. T ribe D u ssu m ieriin i
isth m u s
p o in te d
w ith
b.
D u s s u m iria
u n d e r s id e
1st
of
fro m
S p r a t e ll o id i n i
F ig ^
above
isth m u s
‘s h o u l d e r s ’
E tru m e u s
head
F ig J
i.
2 n d s u p ra -m a x illa
a.
b. Tribe
p r e - m a x i lla e , s e e n
S p ra te llo id e s
2nd
s u p r a - m a x i ll a
b.
4
Jen k in sia
28
D ussum ieria V alenciennes,
C LU P
1847
D uss
D u s s u m ie ria V a le n c ie n n e s , 1 8 4 7 , H is t.n a t.p o is s ., 2 0 :6 4 7 (ty p e : D u s s u m ie ria a c u ta V a le n c ie n n e s , 18 4 7 ).
M ontalbania Fow ler, 1934, Proc.A cad.nat.Sci.Philad.. 8 5 :2 2 4 (type: E tru m eu s (M o n ta lb an ia) alb u lin a F o w ler, 1934)
(m is s p e lt as M o n ta lb ia n a by B e rtin , 1943).
S yn onym s : N one.
D ia g n o s tic F e a tu r e s :
B o d y sle n d e r, b elly ro u n d e d , w ith o u t p re- and p o s t-p e lv ic sc u te s; th e W -sh a p ed
pelvic
scute,
num erous
b ran ch io steg al
rays
(12
to
17)
and
rec tan g u la r
p re-m ax illae
d istin g u ish
D ussum ieria from all other clupeids except E trum eus, w hich has fewer anal f i n r a y s , the pelvic fins less advanced
and the isthm us (sternohyoideus m uscle) w ith lateral flanges or ‘sh o u ld ers’ (see key).
Biology, Habitat and Distribution :
M arin e p elag ic and sch o o lin g fish e s o f th e In d o -P a c ific reg io n , from
the w e ste rn In d ian O cean (the “G u l f ’ to p e rh ap s M ad a g a sc a r) to the w este rn P a cific (C h in a to P a p u a N ew G uinea
and S olom on Islan d s).
Interest to Fisheries :
C o n tr ib u te to g e n e r a l c lu p e o id c a tc h e s , s i g n if i c a n tl y so in I n d o n e s ia a n d th e
Philippines. The total catch in 1983 w as 35 239 tons.
Species : H ith e rto u su a lly c o n sid e re d m o n o ty p ic (W h ite h e a d , 1963, 1973), b u t now k n o w n to in c lu d e at le a st
tw o s p e c ie s (W o n g ra ta n a , 1 9 8 0 ), w h o se ra n g e s o v e rla p in th e “ G u l f ’, a ro u n d In d ia n c o a sts an d M a la y s ia and
In d o n e s ia :
D. acu ta V a le n c ie n n e s, 1847, In d o -W e st P a cific
D. elo p so id e s B le e k e r, 1849, In d o -W est P ac ific .
D ussum ieria acuta V alenciennes,
CLU P
1847
Duss
D u ssu m ie ria a c u ta V a le n c ie n n e s, 1847, H is t.n a t.p o is s ., 2 0 :4 6 7 (B o m b a y , C o ro m a n d e l).
Synonyms : C lupea flosm aris R ichardson, 1846:305 (suppressed); Elops javanicus V alenciennes, 1847:271;
Etrum eus (M ontalbania) albulina Fow ler, 1934:244 (the Philippines); D ussum ieria acuta - W hitehead, 1963:312,
fig. 5; W hitehead 1973b: 170, fig. 4; W ongratana, 1980:88, pis 11, 12 (revision).
FAO Names
En - Rainbow sardine.
wimi
Diagnostic Features :
C lo s e ly re s e m b le s EE e lo p s o id e s , b u t
body d e e p e r (u s u a lly 22 to 29% o f sta n d a rd le n g th ) lo w e r g illra k e rs
few er (19 to 2 6 ), b ra n c h io ste g a l ray s fe w e r (1 2 to 15) and p o ste rio r
p a rt o f s c a le s m a rk e d w ith n u m e ro u s tin y r a d ia tin g s tria e . B la c k
ir rid e s c e n t b lu e w ith a sh in y g o ld /b ra s s lin e b e lo w (q u ic k ly fa d in g
a fte r d e a th ); h in d m a rg in o f ta il b ro a d ly d a rk . S ee C L U P D u ss 1,
Fishing A reas 57, 71 and also 51.
striae
29
Geographical Distribution
W arm er w aters o f
Indo-Pacific, f r o m th e “ G u l f ” ( p e r h a p s s o u t h to
Somalia), along coasts o f Pakistan, In d ia’and M alaysia
to In d o n esia (K alim an tan ) and the P h ilip p in es. E arlier
records included D elopsoides.
Habitat and Biology :
M a rin e , p e la g ic , m ain ly
inshore fishes; studies on bionom ics, general biology,
food, breeding, eggs and larvae, juveniles, tem perature
tolerance o f larvae, the gut and seasonal variations in
fat content are listed by W hitehead (1973:171), but may
equally have referred to D. elopsoides.
Size !
To about 20 cm standard length.
Interest to Fisheries
Separate statistics only
recorded from Hong K ong, Indonesia and the Philippines
(the last tw o m ost likely w ith
ET elopsoides also
included). Good catches are made in Palk Bay and G ulf
o f M annar (southern India) (Sam uel, 1968). C aught will
gillnets (rolavalai o f southern India), also seines.
Local Names :
Literature :
key, diagnosis).
INDIA: Rainbow sardine.
W hitehead (1963 - com bined w ith D
D ussum ieria elopsoides B leeker, 1849
elopsoides, however;
1973b - same); W ongratana (1980
CLUP
D u ss
2
D ussum ieria elopsoides Bleeker, 1849, V erh .b atav .G en o o t.K u n st.W et.. 22:12 (Madura Strait, Java Sea, etc.).
Synonyms :
D u ssu m ieria h asseltii B leek er, 1850:422;
D u ssu m ieria p ro d u ctissim a C h ab an au d , 1933:4,
figs 3-6 (Suez); D u ssu m ie ria a c u ta :C L O F N A M , 1973:110 (e a s te rn M e d ite rra n e a n ); F N A M , 1 9 8 4 :2 7 5 , fig. (sam e);
D ussum ieria elopsoides :W ongratana. 1980:85, pis 9, 10 (revision).
FAO Names :
En - S lender rain b o w sardine.
D iagnostic Features : M ore slen d er th an ET ac u ta (d ep th u su ally 16 to 22% o f stan d ard len g th ), low er
g illrak ers m ore (21 to 32), branchiostegal rays more (13 to 17) and no striae on posterior part o f scales; colour
apparently similar.
30
G eographical D istribution :
From Suez and
w estern Indian O cean (the “G u lf’ to M om basa; possibly
to M adagascar) along coasts o f Pakistan, and India and
M alay sia to w estern P acific (C h in a to about Solom on
Islands).
Im m igran ts
in
eastern
M ed iterran ean
(reported as D. acuta).
H abitat and B iology
Presum ably sim ilar to
th o s e o f FF a c u ta , b u t th e s p e c ie s n o t d is tin g u is h e d in
earlier studies.
Size :
D.
To about 20 cm standard length.
Interest to Fisheries :
acuta.
Local Names :
C om bined w ith data for
As for D. acuta .
40°
60°
80°
100°
120°
140°
160°
180°
Literature :
F ir s t p ro p e rly se p a ra te d fro m D.
acuta by W ongratana (1980).
Remarks : Even more slender specimens (depth 16.7 to 17.2% o f standard length) have been examined from
Fiji w hich have a low er gillraker count o f only 20 to 21 (cf. 21 to 23 in FF elopsoides). Possibly a third species is
present.
Etrumeus Bleeker, 1853
C LU P Etru
E tru m e u s B le e k e r, 1 8 5 3 , V e rh .b a ta v .G e n o o t.K u n s t.W e t., 25:48 (type: Clupea m icropus Schlegel, 1 8 4 6 ).
P e rk in sia E ig en m an n , 1891, A m e r.N at. P h il a d ., 25:153 (tipe: P e rk in sia otho no p s E ig en m an n , 1891). H alecula
Jo rd an ,1925, Stanford Univ. Puhi.B iol. S ei., 4:41 (type: H alecula acum inata Jordan, 1925) (p re-o ccu p ied).
Parahalecula Fow ler, 1958, N otul.N aturae P h ila d ., (310):5 (replacem ent for H alecula).
Diagnostic Features : B ody slender, belly rounded, w ith o u t pre- and p o st-p elv ic
scu tes; th e W -sh a p e d
p e l v i c sc u te ,
n u m e ro u s b ra n c h io s te g a l
rays
(11
to
15)
and
r e c ta n g u la r p r e - m a x illa e d is tin g u is h
Etrum eus from all other clupeids except D ussum ieria, w hich has more anal fin ray s, the pelvic fins more advanced
and the isthmus (stern o h y o id eu s muscle) tapering evenly forward (see key).
Biology,
H abitat and D istribution : M arine pelagic and schooling fishes o f both the
N ew W orld (Atlantic and
P acific co asts) and the In d o -P ac ific (ab sen t b etw een 25°N and 25°S, h o w e v e r); also, so u th ern p art o f eastern
A tla n tic .
Interest to Fisheries : O f im portance mainly o ff Japan and southern A frica,
the Red Sea. The total catch in 1983 was 110 084 tons.
Sp ecies :
(1983):
but a significant fishery
also in
A single species rec o g n ized by W hitehead (1963), but a second sp ecies found, by W o n g ratana
E. teres (De Kay, 1842), Indo-W est Pacific, w estern A tlantic, eastern Pacific
E. w h iteh ea d i W ongratana, 1983, southern A frica.
Etrumeus teres (De Kay, 1842)
CLUP Etru 1
Alosa teres De Kay, 1842, Nat.Hist.New York, pt. 4, Fishes:262 (New York region).
Synonyms :
C lu p e a m icropus S chlegel, 1846:236 ( J a p a n ) ;
E tru m e u s ja c k s o n ie n s is M a c le a y , 1879:36
(A ustralia);
E tru m e u s a c u m in a tu s Gilbert, 1891:56 (e a ste rn P a c ific ); P e rk in sia oth o n o p s E ig en m an n , 1891:153
(eastern Pacific); E trum eus sadina:FW NA, 1964:263 (n o t C lu p ea sad in a M itch ill, 1814, w hich w as p robably
S ard in ella au rita ); E tru m eu s teres - W h iteh ead , 1963:321, fig. 11; Chirichigno, 1968:399, fig. 7 (Lobos de A fuera
Islands, Peru, about 6°30’S); W ongratana, 1980:83, pis 5, 6 (revision); SFSA, in press (southern Africa).
31
FAO Names :
E n - R e d -e y e
ro u n d
D iagn ostic F eatu res :
See genus
m eristic c h a ra c te rs (e sp e c ia lly g illra k e rs),
(W h iteh ead , 1963). D istin g u ish e d from R
(30 to 35) and the pelvic fin base about
F is h in g A re a 51.
Geographical Distribution :
O ff southern
A frican
coasts
(M ozambique to Durban, perhaps
fu rth er south, but p ro b ab ly re ­
p la c e d by R w h ite h e a d i o n
w estern coast), w estern Indian
O cean (o ff Cape G ardefui, in
Red Sea, w ith im m igrants into
the
eastern
M ed iterran ean ),
Japan and southern coasts o f
A ustralia; in w estern A tlantic
(Bay o f Fundy south to Florida,
G u lf o f M exico, V enezuela and
the G uianas); in eastern Pacific
( C a l i f o r n i a , Galapagos
Islands,
ITawaii, Peru at 6°30’S). The
single
eastern
M editerranean
specimen reported by B en-Tuvia
(1963 - see also C LO FN A M ,
1973:110) was probably Spratel­
loides delicatulus.
h e r r in g .
for m ain features.
Some variatio n occurs in body depth and certain
b u t ran g es o v erla p b e tw een p o p u la tio n s from w id e ly se p a ra ted reg io n s
w h iteh ea d i o f so u th e rn A fric a n w ate rs by h av in g few er lo w er g illra k e rs
1/3 eye diam eter behind base o f last dorsal finray. See CLUP Etru 1,
¡WST
8Ö1-
160°
140°
120°
100‘
H a b ita t and B io lo g y : M a rin e , p e la g ic , m ain ly in sh o re fish e s; gen eral b io lo g y in w e ste rn A tla n tic g iv en by
Hildebrand (1963:264-7) and in A ustralia by B lackburn (1941); data from southern A frica included R whiteheadi
also.
Size : To 25
cm
standard
length.
Interest to Fisheries : Separate statistics mainly reported from southern A frica and Japan (105 638 tons in
1983), but E. whiteheadi included in former. C a u g h t m a in ly w ith p u rse se in e s.
L ocal N am es : A U STR A LIA : M aray; JAPAN: Iw ashi; PERU: Sardina redonda; SO U TH A FRICA :
R o n d eh arin g , R ooioo g ; U SA:
A tla n tic ro u n d h errin g (ea ste rn c o ast), M ak iaw a, M ik iaw a, O m aka (H aw aii), R ound
h erring (A FS list).
Literature :
I n d o - P a c if i c ) .
W h ite h e a d (1 9 6 3 - re v isio n ); H ild e b ra n d (1 9 6 4 - e a ste rn c o a st o f U S A ); W o n g ra ta n a (1 9 8 0 -
R e m a r k s : E le c tro p h o re tic s tu d ie s o n p ro te in s m ay w e ll p o in t to d if fe re n c e s b e tw e e n th e v a rio u s
populations that will justify separation o f subspecies or even species. I f it is assumed that geographically isolated
populations o f Sardinops probably represent different species or subspecies, then the same is surely true o f
E tru m e u s.
32
E tru m eu s w h iteh ea d i W ongratana, 1983
CLUP Etru 2
Etrum eus w hiteheadi W ongratana, 1983, J a p .J .Ic h th y o l., 29 (4):387 (South Africa).
Synonym s i E tru m eu s teres o f au th o rs on so u th ern A frica n sp ecim en s,
W o n g ratan a, 1980:84, pis 7, 8 (revision); SFSA, in press (southern Africa).
FAO N am es :
e s p e c ia lly o n w e ste rn c o a sts;
E n - W hitehead’s round herring.
D iagnostic Features : See genus for main features. D istinguished from R teres in southern A frican w aters
by having more low er gillrakers (36 to 39) and the pelvic fin base below or just before base o f last dorsal finray.
G eographical D istribution : W alvis Bay to D urban, pos­
sibly o v erlap p in g w ith R te res from ab o u t C ape Tow n eastw ard .
H abitat and B iology : M arine pelagic, m ainly inshore
fis h e s ; p ro b a b ly in c lu d e d in th e d a ta g iv e n fo r R te re s by
w orkers in southern Africa.
Size :
To 20 cm standard length.
Interest to F ish eries
Separate statistics for South
A frica (63 009 tons in 1983) m ust refer largely to this species,
alth o u g h g iven as R te re s . M uch sm aller c atch es, also p re ­
sumed to be th is sp e c ie s, are re p o rte d by N a m ib ia , and in
earlier years by P oland and the G erm an D em o cratic R epublic
working off this coast (6 653 tons in 1983).
Local Names :
SO U TH A FR IC A : S u id afrik aan se
ro n d e-
harig.
Literature : W ongratana
(1983
- key, diagnosis).
Remarks : M uch m ore w ork is needed to determ ine the
overlap o f this species w ith R teres in southern A frican waters
and to explore possible differences in their biology.
click fo r n e x t p a g e
click fo r p re v io u s p a g e
33
B leeker,
Spratelloides
1851
C LU P S pratel
S p ratello id es B leek er, 1851, N atu u rk .T id sc h r.N e d .-In d ië , 2 :2 1 4 (type: C lu p ea a rg y ro ta e n ia ta B le e k e r, 1849 =
£L gracilis).
Stolephorus:Fowler, 1941,
B ull.U .S.natn.M us., 13(100):561 (not Stolephorus Lacepède, w hich is an
anchovy).
D ia g n o s t ic F e a tu r e s :
T he W -s h a p e d p e lv ic s c u te an d fe w b ra n c h io s te g a l ra y s (6 o r 7) d is tin g u is h
Spratelloides from all other clupeids except Jenkinsia, w hich has only a single (posterior) supra-m axilla and does
not occur in the Indo-Pacific re g io n .
B iology, H ab itat and D istrib u tion :
Sm all m arine pelagic fishes o f the Indo-P acific region, from the
w e s te rn In d ia n O ce an (th e “ G u lf” so u th to a b o u t D u rb a n ) to th e w e s te rn P a c ific (J a p a n so u th to n o rth e rn and
w e ste rn A u s tr a lia , a lso S am o a).
Interest to Fisheries :
Contribute to general clupeoid catches, but
around Japan (5 159 to n s in 1983) and F iji (9 6 tons in 1983).
Species :
£L
£L
£L
S.
separate
statistics
only
given
for
W hitehead (1963: 1973) recognized only two species, but W ongratana (1980, 1983) adds two more:
delicatulus (Bennett, 1831), Indo-W est Pacific
gracilis (Temm inck & Schlegel, 1846), Indo-W est Pacific
rohustus Ogilhy, 1897, Indo-W est Pacific.
lewisi W ongratana, 1983, Indo-W est Pacific
S p r a te llo id e s d e lic a tu lu s
C lu p e a
(B e n n e tt, 1831)
delicatula B e n n e tt,
C L U P S p ratel 1
1831, Proc.zool.Soc.Lond.. 1:168 ( M a u r it iu s ) .
Synonym s
Clupea m acassariensis B leeker, 1849:72:
A lausa alburnus K ner & Steindachner, 1866:387
(V alparaiso given, but in fact Samoa): Spratelloides delicatulus - W hitehead, 1963:345 (including £L robustus):
Idem . 1973b:172, fig. 6 (same): FN AM , 1984279, fig. (single eastern M editerranean specimen: given’& Etrum eus
teres in CLOFNAM , 1973:110): W ongratana, 1980:105, pis 39, 40 (revision: £L robustus rem oved): SFSA, in press
(s o u th e rn A fric a ).
FAO Names :
En - Delicate round herring.
c o n tin u o u s striae
Diagnostic Features :
N o b rig h t silv e r
band alo n g fla n k s , m a x illa to o th le s s , se c o n d
su p ra -m a x illa p ad d le -sh a p e d and sy m m e tric a l,
lo w e r g illr a k e r s 26 to 3 2 , v e rtic a l s tria e on
s c a le s m e e tin g a t c e n tre , p o s te rio r m a rg in o f
sc a le s sm o o th , p re -d o rs a l sc a le s 8 to 13, and
35 to 41 sc a le s in la te ra l s e rie s: tw o p ro m i­
n e n t d a r k s t r e a k s o n c au d a l fin b a s e . See
C L U P S p ratel 1, F is h in g A re a 51.
2 n d su p ra m a x illa
scale
34
G eographical D istribution :
Indo-P acific, fro m
w estern In d ian O cean (R ed Sea south to D u rb an , also
M ad ag ascar and M au ritiu s) to w estern P ac ific (Japan
south through the Philippines to northern A ustralia, also
e a stw a rd to S o c ie ty Is la n d s , b u t n o t T u a m o tu and
M arquesas Islands).
A single sp ecim en from eastern
M editerranean (Tel-Aviv, Israel).
H abitat and B iology : M arine pelagic and usually
in sh o re schooling fishes. M ore d ata needed.
Size :
2o°
To 7 cm standard length.
In terest to F isheries
Separate statistics are
not recorded, but probably makes a significant contri­
bution to clupeoid fisheries. Used as a tuna b a itfish in
the P acific.
'
L ocal N am es :
20°
0°
20°
40°
60°
80°
100°
120°
140° 160°
180°
160°
140°
Literature : W hitehead (1963, 1973b - review); W ongratana (1980 - key, diagnosis); Lewis, Smith & Ellway
(1983 - tuna baitfish).
CLUP
S p ra tello id es g r a c ilis (Tem m inck & Schlegel, 1846)
C lupea gracilis Temm inck & S ch leg el, 1846, F au n a Jap o n ica. P oiss.
Nagasaki).
S p ra te l 2
pt. 5, inst. 13:238 (so u th e rn co ast o f
Synonym s
C lupe a
argyrotaeniata
B leeker, 1849:72;
S pratelloides
atrofasciatus
Schultz, 1943:8;
Stolephorus japonicus:F ow ler,
1941:567 (not Sk i aponicus L acepède, 1803, an anchovy);
S p ra tello id e s g racilisW h iteh ead , 1963:338, fig. 18; Id e m . 1973b:171, fig. 5; W ongratana, 1980:102, pis 35, 36 (revision).
F A O Names
E n - S ilver strip e round herring.
c o n tin u o u s stria e
D iagn ostic F eatu res : Im m ed iately
d istin g u ish ed from oth er species by the
b rig h t s ilv e r b an d a lo n g th e fla n k s (a
fain ter band w hich fades an terio rly in S^
lew isi).
A ls o , m a x illa to o th e d , seco n d
su p ra -m a x illa a sy m m e tric a l (lo w e r p art
larger than upper), low er g illrak ers very
v a r ia b le (2 7 to 37), v e r tic a l s tr ia e on
scales not m eeting at centre and 42 to 48
scales in lateral series. See CLUP Spratel
2, Fishing Area 51.
2 n d su p ra m a x illa
scale
35
G eographical D istribution :
Indo-Pacific, from
w estern Indian O cean (Red Sea south to Z anzibar) to
w estern Pacific (Japan south through the Philippines to
Sam oa, not Cook, Society or M arquesas Islands, but
reappears in the T uam otu Islands, and southeastern
A u s tr a lia , also o f f w e ste rn A u stra lia ).
H ab itat and B iology :
M arine, p elagic
u sually in sho re schoolin g fish es. M ore d ata needed.
and
Size : T o 9 .5 cm s ta n d a r d le n g th .
Interest to F isheries
Separate statistics only
recorded for Fishing A reas 61 and 71 (5 255 tons in
1 9 8 3 , w h ic h m ay in c lu d e o th e r s p e c ie s ), b u t p ro b a b ly
c o n tr ib u te s m o re to c lu p e o id c a tc h e s th a n th e re c o rd s
show.
U sed as a tuna b a it f i s h in the Pacific.
L ocal N am es : L iterature : W hitehead
E llw a y (1 9 8 3 - tu n a b a i t) .
(1963,
1973b
W ongratana
- review );
(1980
- key, diag n o sis); L ew is, Sm ith &
R em arks: W hiteh ead (1 9 6 3 , 1973b) co n sid ered the so u th ern and w estern A u stra lia n form a su b sp ecies, but
W ongratana (1980) convincingly show ed it to be a d istin ct sp ecies, Sk ro b u stu s.
S p r a te llo id e s le w isi W o n g ra ta n a , 1983
C L U P S pratel 3
S p r a t e l l o i d e s lewisi W o n g r a t a n a , 1 9 8 3 , Jap.J.Ichthyol., 2 9 (4 ):3 8 9 (S o lo m o n Is la n d , c o a sts o f P a p u a N ew
G u in e a ).
S y n o n y m s : P ro b a b ly in c lu d e d by a u th o rs in re c o rd s o f
G u in e a are a ; W o n g r a ta n a , 1 9 8 0:104, pis 37, 38 (rev isio n ) .
g ra c ilis or
d e lic a tu lu s fro m th e P a p u a N e w
FAO Nam es : E n - L e w is’ round herring.
tí/ fi à
t ffv' Jt
p y y y V . V 'A
tS/\, ^
w 1.
^
1 *
> '- 'V
^ KA
K'K~^ /-W «-4
J•t t ' w*“icv)s?
I’ ‘
'
'
t
> k I
A K K >{ ) A a T À K
■-4
\
.
t
I
l i i . »
TlV
* >■
d isc o n tin u o u s striae
D ia g n o s t ic F e a t u r e s : D i s t i n ­
g u is h e d fro m o th e r sp e c ie s m a in ly by
th e s i l v e r b a n d on th e f la n k s w h ic h
fad es a n te rio rly (at a b o u t tip s o f p e c to ­
ral fin s). A lso , m a x illa to o th e d , second
su p ra -m a x illa asy m m e tric a l (lo w e r p art
la rg e r th a n u p p e r), lo w er g illra k e rs 28
to 3 2 , v e r t i c a l s t r i a e o n s c a l e s n o t
m e e tin g at c e n tre , and 39 to 43 sc a le s
in lateral series.
2 n d su p ra m a x illa
scale
36
Geographical Distribution :
W e s te rn P a c if ic
(Solom on Islands, coasts o f Papua N ew G uinea, o ff Irian
Jaya).
w
Habitat and Biology : M arin e, pelag ic and usually
inshore schooling
fishes. M ore data needed.
Size :
To 6 cm standard length.
20
°
0
°
Interest to F ish eries : No separate statistics.
L o ca l
N am es :-
Literature :
W ongratana
(1983 - key,
diagnosis).
Remarks :
The anterior fading
o f the silver lateral 2QO
stripe along the flanks appears to be a good field charac­
te r .
40°
80°
Spratelloides robustus Ogilby,
100°
120°
140°
160°
180°
C LU P S pratel 4
1897
Spratelloides robustus O gilby, 1897, Proc.L inn.Soc.N .S.W ., 22:64 (New South Wales).
Synonyms : S p ra te llo id e s d e lic a tu lu s ro b u s tu s :W h iteh e ad , 1 9 6 3 :3 4 8 ; S p ra te llo id e s ro b u stu s - W o n g ratan a,
1980:108, pis 41, 42 (revision)
FA O
Names:
En - F rin g e-scale round herring.
■Vr r r p r f t t { J f f f f f ]
i\ ) 1 1 , 1 . 11. ( K y y r y y y ’/ y Y y y y y Y V y y Y Y Y y y y y y v y
7 7
M
n
* 6
1 1 ' *>^
1f U
' 7
1 ) ƒ >7
1 I I ! [ f t ■;
! lr
co n tin u o u s striae
D iagnostic F eatures :
C losely
resembles S^ delicatulus in lacking a sil­
very stripe-along the flanks, no teeth on
m axilla, second su p ra-m ax illa sym m etri­
cal and scale striae m eeting at centre;
d iffe rs in h a v in g a to o th e d p o s te rio r
m argin to scales, pre-dorsal scales 13 to
16, and 42 to 46 scales in lateral series.
2nd
supra-m axilla
scale
37
Geographical Distribution :
W e ste rn A u s tra lia
(Brenner B ank w estw ard and northw ard to D am pier A rc h i­
pelago), so u th eastern , A u stra lia (so u th ern p arts o f N ew
South Wales).
—
1 V t)
4
i
&
>
*"
%
»
Habitat and Biology : M arin e, pelagic and usually
inshore schooling fishes. M ore data needed.
Size
a
•Sa.**
To 7.5 cm standard length.
Interest to Fisheries :
S e p a ra te
recorded, probably o f little im portance.
s t a ti s ti c s n o t
40„
•
X)
L o ca l N a m es : Literature :
'
vr
C
e
W o n g ratan a (1980 - key, diagnosis).
Remarks : A pp aren tly overlaps w ith Sk delicatu lu s
along northw estern coast o f Australia.
C L U P Jenk
J e n k in sia Jo rd an & E v erm a n n , 1896
Je n k in s ia Jo rd a n & E v e rm a n n , 1 8 9 6 , B ull.U .S .natn.M us.. 50:418 (type:
G ilbert, 1884.
D u ssu m ieria
sto lifera Jo rdan &
D iagnostic Features : The W -shaped pelvic scute and few branchiostegal rays (6 or 7) distinguish Jenkinsia
from all other clupeid s ex cep t S p ra tello id e s, w hich has tw o su p ra-m ax illae (an terio r m issing in Jen k in sia )
confined to the Indo-P acific region.
Biology, H abitat and
(Florida south to V enezuela
Interest to Fisheries :
D istribution :Sm all, m ainly coastal m arine pelag ic fish es o f the w estern A tlan tic
and the Guianas; also Bermuda).
N o separate statistics; perhaps o f sm all local interest.
Species ! H ildeb ran d (1964:268, i.e. FW NA) recognized two species, the w idespread .L lam protaenia and the
Berm udan J. viridis, the latter considered merely a synonym by W hitehead (1963:351), who described a second
species, E m a i u a . C e r v i g ó n & Velasquez (1978) split E lam protaenia into three apparently distinct species, the
other tw o being E stolifera and E parvula. G illraker num bers suggest that the situation is more com plex and can
only be resolved using large co llectio n s th ro u g h o u t the C arib b ean area. F or the m om ent, four species can be
recognized:
E
E
E
E
lam p ro taen ia (G o sse, 1851) w estern A tlan tic
m ajua W h iteh ead , 1963, w e ste rn A tla n tic
parv u la C erv ig ó n & V elasq u ez, 1978, w e ste rn A tla n tic
sto lifera (Jo rd a n & G ilb e rt, 1884), w e ste rn A tla n tic .
Jenkinsia lam protaenia (G osse, 1851)
C L U P Jen k 1
C lupea lam protaenia G osse, 1851, N atu ralist’s soiurn in Jam aica:291 (Jamaica).
Synonym s
Stolephorus viridis
B ean, 1912:122 (B erm uda); Jenkinsia berm udana R ivas, 1946:1 (viridis
overlooked); Jenkinsia lam protaenia -FW N A, 1964:268, fig. 64; W hitehead, 1963:351, fig. 23; C ervigón
1966:116; W hitehead, 1973a:18, fig. 3; C ervigón & V elasq u ez, 1978:5, figs, 1-54 (descr., osteology).
38
F A O Names :
En - D w arf round herring (D w arf herring in FA O Sheet, A rea 31).
D iagnostic Features : P re-m ax illae to o th ed , low er g illrak ers 20 to
30, but som e co rre la tio n w ith g eo g rap h ical area (see R em arks b elow ),
isthm us slender, but w ith slight ‘shoulders’ anteriorly, pectoral finrays 12 to
15 (usually 13), vertebrae 40 to 43 (usually 42 - V en ezu ela), silv er lateral
band not becom ing narrow er or fading near gili opening. See CLUP Jenk 1,
Fishing A rea 31.
G eographical D istribution :
W e ste rn c e n tra l
A tlan tic (F lorida, G u lf o f M exico and C arib b ean south
to Tobago; also Bermuda).
Habitat and Biology : M arine pelagic and usually
inshore schooling fishes, feeding on Zooplankton.
Size :
To 6.5 cm standard length.
Interest to F isheries :
S e p a ra te s ta tis tic s n o t
recorded, probably o f little im portance.
L o ca l N am es : USA:
V ENEZUELA: Canalera.
D w a rf h errin g (AFS list);
Literature :
F W N A (1 9 6 4 - k e y , d ia g n o s is,
b iology); W hitehead (1 9 6 3 , 1973a - revision); C erv ig ó n
& Velasquez (1978 - osteology).
Remarks :
L o w e r g illr a k e r c o u n ts , a lth o u g h
show ing som e o v erlap , sh o w som e c o rr e la tio n w ith
g e o g ra p h ic a l a re a , su g g e s tin g th a t th e sp e c ie s m ay
comprise three races or subspecies:
(a)
(b )
(c)
gillrakers 20 to 26, usually 22 to 24; Mexico,
Ja m a ic a , D ry T o rtu g a s , B a h a m a s, V irg in
Islands, M artinique, Tobago;
g illra k e rs
2 3 to 2 9 , u s u a l l y 2 4 to 2 7 ;
Colom bia, Venezuela;
g illr a k e r s 2 7 to 3 0 , u s u a lly 2 9 ; B e rm u d a
(v irid is).
isthm us
underside o f head
39
Jen k in sia m ajua
C L U P Jen k 2
W hitehead, 1963
Je n k in sia m aju a W h ite h e a d , 1963, B u ll .B r .M u s .n a t .H i s t.Z o o l., 1 0 (6 ):3 5 5 (C a m p e c h e B a n k ; S w an Is la n d ;
B ah am as).
Syn onym s : Je n k in sia m ajua w oo d si W h iteh ead , 1963:356, fig. 24.
FAO Names :
En - Little-eye round herring.
D iagnostic F eatures : No teeth on pre-m axillae, low er gillrakers 21
to 28, but c o rre la te d w ith g e o g ra p h ic a l a re a (see rem ark s b elo w ), isth m u s
w ith d is tin c t b lu n t ‘s h o u ld e rs ’ a n te rio rly , p e c to ra l fin ra y s 12 to 14, v e rte ­
brae 38 (one fish ), silv er lateral band no t fad in g aw ay an terio rly .
G e o g r a p h ic a l D is t r ib u tio n :W e s te r n c e n tra l
A tla n tic ( G u lf o f C a m p e c h e ; B a h a m a s , n o rth w e s te rn
c o ast o f C aribbean).
isth m u s
underside o f head
H a b ita t a n d B io lo g y :M a r in e , p e la g ic and
usually inshore schooling fishes, feeding on Zooplankton.
S iz e : T o 5 .5 c m s t a n d a r d l e n g t h .
I n t e r e s t to F is h e r i e s : S e p a r a te s t a t i s t i c s n o t
reco rd ed , pro b ab ly o f little im p o rtan ce.
L o c a l N a m e s :U S A :
L ittle -e y e h e rrin g (A FS
li s t ) .
Literature : W hitehead (1963 - key, revision).
R e m a r k s : T W O s u b s p e c ie s w e re re c o g n iz e d by
W hitehead (1 9 6 3 :3 5 4 ), based on low er g illraker counts:
(a )
(b )
L m a iu a m a i u a : lo w e r g il lr a k e r s 2 4 to 28
(m ode 26); G u lf o f M exico.
L m aju a w o o d s : lo w e r g illr a k e r s 21 to 24
(m o d e 2 3 ); B a h a m a s , n o r th w e s te r n C a r ib ­
bean.
40
J e n k in s ia
C LU P Jenk 3
p a r v u la C ervigón & V elasquez, 1978
Jenkinsia parvula C e r v ig ó n & V elasquez, 1978, C u a r d e r n .o c e a n o g r .U n iv .d e O riente V e n e z ., (7 ):9 (V enezuela).
Synonym s : Jenkinsia stolifera - C ervigón, 1966:925, flg. 376.
FAO Names :
E n - S h o r t - s t r ip e d
ro u n d
h e r r in g .
D ia g n o s tic F e a tu r e s : P re - m a x illa e to o th e d , lo w e r g illr a k e rs
2 0 to 24 ( u s u a lly 21 o r 2 2 ) , is th m u s b r o a d ly tr ia n g u l a r , p o in te d
an te rio rly , p ecto ral f i n r a y s 10 to 13 (u su a lly 11 or 1 2), v e rte b ra e 38
o r 39 (u s u a lly 3 9 ), silv e r la te ra l band b e co m in g n a rro w e r and fad in g
anteriorly.
G e o g r a p h ic a l D is t r ib u tio n
A tla n t ic ( V e n e z u e la ) .
W e s te r n c e n tr a l
H a b ita t an d B io lo g y : M a rin e , p e la g ic sch o o lin g
n e a r c o ra l re e fs , fe e d in g on Z o o p lan k to n .
Size
To 5.5 cm sta n d a rd len g th .
I n t e r e s t to F is h e r i e s : S e p a r a te s t a t i s t i c s n o t
re c o rd e d , p ro b a b ly o f little im p o rta n c e .
L ocal N am es
Literature
diagnosis).
C e rv ig ó n & Velasquez (1 9 7 8 - key,
R e m a r k s : S p e c im e n s w ith th e sa m e c h a r a c t e ­
ristic anterior narrow ing o f the silver band on the
fla n k s, but w ith a w id e ran g e o f lo w er g illra k e rs (18 to
2 5 ), h a v e b e e n re p o r te d f r o rn F lo r id a , th e B a h a m a s ,
J a m a ic a , S e rra n a B a n k an d P u e rto R ic o ( W h ite h e a d ,
1 9 7 3 a:2 0 ). M o re ta x o n o m ic w o rk is n e ed ed .
isth m u s
underside
o f head
41
C LU P Jenk 4
J en k in sia sto lifera (Jo rd an & G ilb ert, 1884)
D ussum ieria stolifera Jordan & G ilbert, 1884, B ull.U .S .natn.M us., 7:25 (Key W est, Florida).
S y n o n y m s : In c lu d e d in L la m p ro ta e n ia in F W N A , 1 9 6 4 :2 7 1 : W h ite h e a d , 1 9 6 3 :3 5 1 : J e n k in s ia s to life ra W h ite h e a d , 1973a: 18 (k ey ): C ev ig ó n & V e la z q u e z, 1 978:9, 12 (V e n e z u e la ).
F A O N a m e s : E n - F lo r id a ro u n d h e r r in g .
D ia g n o s t ic F e a tu r e s : P re - m a x illa e to o th e d , lo w e r g illr a k e rs
28 to 24 (u s u a lly 22 o r 2 3 ), is th m u s b ro a d ly tr ia n g u l a r , p o in te d
a n te rio rly , p ecto ral fin ray s 11 or 12, v erteb rae 38 to 39 (u su ally 39 V e n e z u e la ). S ilv e r la te r a l b a n d n o t b e c o m in g n a rr o w e r o rf a d in g
anteriorly.
isth m u s
G e o g r a p h ic a l D is t r ib u t i o n : W e s te r n c e n tr a l
Atlantic (Florida, British Honduras, Venezuela).
H a b ita t a n d B io lo g y : M a rin e , p e la g ic and
usually inshore schooling fishes, feeding on plankton.
S ize : To at le a s t 4 cm sta n d a rd le n g th , p ro b a b ly
to 5 or 6 cm.
40°
I n t e r e s t to F is h e r ie s : S e p a r a te s t a t i s t i c s n o t
recorded, probably o f little importance.
Local Names :
USA: S h o rth an d h errin g (A FS
20
list).
L ite r a tu r e : W h ite h e a d (1 9 7 3 a - k ey ): C e rv ig ó n
& V elazquez (1978 - key, d ia g n o sis, on V en ez u e la n
specimens only).
o°
Remarks :
The broadly triangular isthmus and
th e c o m p le te s ilv e r la te ra l b an d seem to c h a r a c t e r i z e
the species, but more taxonom ic w ork is needed.
20„
1 20 °
100°
80 °
60 °
40 °
click fo r n e x t p a g e
click fo r p re v io u s p a g e
42
2 . 2.2
SU BFA M ILY CLUPEINAE
F A O Names :
E n - H e rrin g s, S ard in es, Sprats.
D iagnostic Features :
Small or
m o d e ra te -siz e d
h errin g -lik e
fishes
w ith a norm al pelvic scute (i.e. w ith
a sc e n d in g arm s) an d scutes p r e s e n t
b e fo r e an d b e h in d th e p e lv ic fin s .
U pper ja w round ed and not notch ed
w hen seen from the front; tw o su p ram a x illa p re s e n t, a u su a lly e lo n g a te
a n te rio r and a p a d d le -sh a p e d p o s te ­
rior; mouth term inal, lower ja w som e­
tim es projecting slig h tly , tee th sm all,
conical. Dorsal fin at about m idpoint
o f body, short (13 to 21 finrays); anal
fin short (12 to 23 finrays), its origin
usually well behind the last dorsal finra y
p elv ic finrays 7 to 10 (m ostly 7
o r '8).
notch
rounded
2nd
supramaxilla
1st
C lupeinae
b.
supramaxilla
Alosinae
Biology, H abitat and D istribution : The Clupeinae are mainly marine coastal and schooling fishes, but some
enter brackishw ater and a few are confined to fresh- or brackishw ater (R hinosardinia, Platanichthys).
They occur
in the In d o - P a c ific region, on both sides o f the A tlantic and in the eastern Pacific. M ost o f the species are found
in tropical or subtropical w aters, but genera such as C lupea, S p ra ttu s, Sardina and S ardinops occur in cool waters
and high latitudes, extending the range o f this subfam ily to about 70°N and 55°S. These cool-w ater genera
contribute to som e o f the m ost im p o rtan t o f all clup eo id fish e ries, but considerable tropical and subtropical
catches come from species o f Sardinella, H arengula, H erklotsichthys, etc.
There are 15 genera (4 w orldw ide, 6 N ew W orld, 3 Indo-Pacific, 2 European) and 72 species, thus the largest
o f the 5 subfamilies o f Clupeidae.
Key to the G enera o f C lupeinae :
N o bony capsule (bulla) in pterotic bone: pelvic
finrays i 6 to i 7; tem perate w aters only
2a.
2b.
lb.
Last tw o anal rays norm al, not enlarged;
low er g illrak ers 30 to 41; Europe, South
A m erica, A u stralasia ......................................
FiS- 1
enlarged
Sprattus
L ast tw o anal finrays enlarged (Fig. 1);
lower gillrakers 39 to 67; Europe only .... C lupeonella
pre-epiotic fossa
B o n y c a p su le (bulla) p resen t in p tero tic bone
(Fig.' 2)
3a.
O p e rc u lu m w ith b o n y ra d ia tin g stria e
(Fig. 3); last two anal finrays e lo n g a te d ;
g illrak ers ab sen t on hind face o f third
e p ib ra n c h ia l; fle sh y ‘r a k e r s ’ on u p p e r
edge o f ceratohyal (Fig. 4)
top o f
pterotic bulla
(fossa and bulla visible
w hen m uscles rem oved)
~
bony
striae
F i^ A
‘rakers’ on upper
edge o f ceratohyal
Fig. 4
43
4a.
4b.
3b.
Scales on flank o f uneven size; low er gillrakers
not shortened at angle o f first gili arch (Fig.
5a);
m a x illa n o t r e a c h in g to ey e c e n tr e ;
eastern A tla n tic , M e d iterran ean only ......................Sardina
/
Scales o f equal size along flanks; low er g il l­
ra k e rs sh o rte n e d at a n g le (F ig . 5b); m a x illa
re a c h in g o r a lm o st re a c h in g to eye c e n tre ;
eastern N , S P ac ific , so u th ern A frica, A u stra ­
lia, N ew Zealand ........................................................... Sardinops
O perculum sm ooth;
g illrak ers usually p resen t on
hind face o f third epibranchial (Fig. 6); upper edge
o f ceratohyal smooth
5a.
angle ot arch
Sardina
a.
shortened
Flind border o f gili opening w ith tw o d istin ct
fleshy outgrowths (Fig. 7)
6a.
T o o th ed h y p o -m a x illa ry b o n e b e tw e e n
ore-m axilla tio and bulge o f maxilla (Fig.
8); w estern A tlan tic ....................................... H aren gu la
6b.
No hypo-m axillary bone
7a.
L ast dorsal finray a long filam ent;
w e s te r n A tla n t ic , e a s te r n P a c i ­
fic ....................................................... O pisthonem a
7b.
L a s t d o rs a l finray
P a c ific
Sardinops
F ig. 5
gili arches
1 2 3
n o rm a l; Indo-
8a.
F ro n to -p arietal striae on top
o f head few, about 3 to 7
(F ig . 9a); lo w e r p a rt o f se ­
cond
(p o sterio r)
su p ra ­
m a x i l l a l a r g e r th a n u p p e r
(F ig . 10a); last tw o anal fin ­
rays not enlarged
H erklotsichthys
8b.
F ro n to -p arietal striae on top
o f h e a d n u m e ro u s , 7 to 14
( F i g . 9b); s e c o n d s u p r a ­
m a x illa la r g e r th a n u p p e r
1 0 b ); la s t tw o a n a l fin ra y s
enlarged
gillrakers on
hind face o f
3rd epibranchial
F is- 6
fleshy lobes
to oth ed
hypom axilla
F ig. 8
H aren g u la
2nd
a. H erklotsichthys
2nd
supra-m axilla
supra-m axilla
b. Sardinella
fronto-parietal striae
Fig- 9
a.
H erklo tsich ty s
b
S ardinella
Fig. 10
44
5b.
9a.
Low er gillrakers 26 to 43; a m edian series o f
p re -d o rsa l sc a le s (Fig. Ila)
.............................. A m blygaster
9b.
Low er gillrakers rarely less than 40 (m ostly 45
to 9 0 , b u t o v e r 2 0 0 in som e s p e c ie s ); p re­
dorsal scales usually paired (Fig. lib )................. Sardinella
Hind border o f gili opening evenly rounded, w ithout
fleshy outgrowths
10a.
P e lv ic finrays i 8 (rarely i 7 or i 9); belly not
sharply keeled; pelvic in sertio n behind dorsal
fin origin; North Atlantic, North P a c ific ........................ C lupea
10b.
P elvic finrays i 6 to i 7
lia.
A m b ly g aster
pre-dorsal
b.
scales
Sardinella
Fig. 11
Indo-Pacific;
pelvic finrays i 6; second
supra-maxilla rectangular (Fig. 1 2 )..............E scu a lo sa
lib. N ew W orld only
12a. P elvic finrays i 6
13a.
13b.
S ilv e r s trip e a lo n g fla n k s ;
p o s te rio r fro n ta l fo n ta n e lle s
2nd supra-m axilla
re ta in e d
in
a d u l t s (Fig.
1 3 ) ........................................ P latanichthys
Escualosa
FiS- 12
F lanks
s i l v e r , no
strip e ;
p o s te rio r fro n ta l fo n ta n e lle s
occluded in a d u lts ................ R am nogaster
12b. P elvic finrays i 7
14a. Sharp
b ackw ard
p ointing
sp in e n e a r fro n t o f m a x illa
(Fig. 1 4 ) ....................................R hinosardinia
14b. N o sharp spine on m axilla
15 a. S i l v e r s t r i p e a l o n g
flanks; e a s te rn c e n tra l
P a c if ic , C a rib b e a n and
so to B r a z i l ................................ L ile
15b. F lan k s silv er, no stripe;
so u th e rn tip o f S o u th
America...............................Strangom era
p o sterio r
fro n ta l
fo n tan elles
top o f head
P la t a n ic h t h y s
spine
R hinosardinia
Fig. 13
45
S p ra ttu s G irg en so h n ,
C LU P S p ratt
1846
Sprattus G irgensohn, 1846, M ém .savants étrangers A cad .Sei.P étersb., 5:534 (type: Sprattus haleciform is
G irgensohn, 1846). Spratella V alenciennes, 1847, H ist.nat.poiss., 20:356 (type: Spratella p u m ila V alenciennes,
1846).
M eletta V alenciennes, 1847, H ist.n at.p o iss.. 20:366
(type: M e le tta v u l g a r i s V a le n c ie n n e s, 1847).
M augeclupea W h itley . 1932. R e c .A u s tr.M u s.. 18:332 (type: C lupea bassensis M cC ulloch,1911). A ntu de Buen,
1958, R e v .B io l.m a r.V a lp a ra iso , 8 :8 7 (type: C lu p ea fu eg en sis Jen y n s, 1846).
D iagnostic Features : A bsence o f a pterotic bulla
(bony dome on floor o f pre-epiotic fo ssa) distinguishes
Sprattus from all other clupeine genera except C lupeo­
nella (w hich has the pelvic fin origin distinctly behind the
dorsal fin origin and has the last tw o anal finrays e n ­
larged). From other C lupeinae that occur sym patrically,
Sprattus differs in having only i 6 or 7 pelvic finrays
(usually i 8 in C lupea), sharply keeled sc u te s (rather
rounded in S tran g o m era) and no radiating bony striae on
the opercu lu m ( S a rd in ia , S ard in o p s).
A p a rt fro m th e
absence o f a pterotic b u ll a , how ever, Sprattus hardly
differs from the South A m erican R am nogaster (U ruguay
to T ierra del F u e g o ).
no p te ro tic b u lla
/
B io lo g y , H a b ita t an d D istr ib u tio n : M arin e p elag ic
and schooling fishes, especially o f coastal w aters, often
close inshore; som etim es tolerating very low salinities; distribution essentially antitropical, occurring in the
N orthern H em isphere (Europe) and the Southern Hem isphere (southern parts o f South A m erica and A ustralia, also
in N ew Z ealand).
Interest to Fisheries : Im portant fisheries in the N orth Sea, o ff N orw ay and in the B altic (Sk sprattus), but
perh ap s u n d erex p lo ite d in the S o u th ern H em isp h ere.
Species
A single representative in the N orthern H em isphere
(Sk sprattus) and 4 in the Southern
H em isphere, the 2 N ew Z ealand species only recently d istinguished ( W h ite h e a d , Sm ith & R obertson, 1985):
R
R
R
R
R
antipodum (Hector, 1872), N ew Zealand
fuegensis (Jenyns, 1842), Southern South A m erica
muelleri (Klunzinger, 1880), N ew Zealand
novaehollandiae (Valenciennes, 1847), Southern A ustralia
sprattus (Linnaeus, ,1758) N ortheast A tlantic, M editerranean.
R em arks : A bsence o f a pterotic bulla can only be determ ined by dissection. R em oval o f the muscles near
the corner o f the operculum reveals a cavity, the pre-epiotic fossa. I f the pterotic b u ll a is present, it appears as a
rough-surfaced dom e on the floor o f the fossa.
Sprattus antipodum (H ector, 1872)
CLUP
S p ra tt
4
C lupea sprattus
var. antipodum H ector, 1872, N otes edib.fish. in H utton, C at.Fish.N .Z., Colon.M us. & Geol.
Surv.Dept., Puhi. 18:133 (Foveaux Strait, N ew Zealand).
S yn on ym s : C lu p e a sp ra ttu s var. a n tip o d a ru m :C o le n so , 1 8 7 9 :5 7 2 (u n ju stifie d em e n d a tio n ); C lu p e a a n tip o d a
H utton,
1904:51; C lupea holodon R egan, 1916:5 (Stew art 1.); C lupea antipodum :Regan. 1917:227;
S p rattu s
antipodum - W hitehead, Smith & R obertson, in press (key, diagnosis, biology, synopsis); name applied to
muelleri in most literature.
FAO Names :
En - N ew
Z ealan d
blueback s p r a t
46
ïàëèr
fine
striae
D iagnostic Features : Body slender, its
d e p th 16 to 26% o f sta n d a rd le n g th (m e a n
22%); gili cover w ithout bony radiating striae;
toothplate on tongue broad, alm ost oval, width
about 3 tim es in len g th , lateral tee th strong:
scales w ith fine rad iatin g or h o rizo n tal striae
on exposed posterior part; vertebrae 48 to 51;
back steel blue when fresh.
*
■
i
toothplate on tongue
scale
G eographical D istribution : N ew Zealand coasts
(east coast o f the N orth Island, Cook Strait, apparently
all coasts o f the S outh Islan d south to S tew art Island
and F o veaux Strait).
H ab itat and B iology
M arine pelagic and
sc h o o lin g fish e s o f c o a sta l w a te rs ; b io lo g y p ro b a b ly
sim ilar to that o f Sk m u elleri, from w hich it has not
usually been distinguished in earlier studies.
Size :
10 cm .
To 12 cm stan d ard len g th , u su ally 8 to
Interest to Fisheries :
No
but perhaps o f local interest.
L ocal
se p a ra te
s ta tis tic s ,
N am es : -
L iterature : W hitehead,
p ress - synopsis).
Sm ith
& R obertson
(in
S p rattu s fu eg en sis (Jenyns, 1842)
CLU P
S p ra tt
2
C lupea fuegensis Jenyns, 1842, Z o o l.v o y .B eag le , fishes:133 (Tierra del Fuego).
Synonym s
C lupea fuegensis: R egan, 1916:4; N orm an, 1937:37, fig. 14 (T ierra d e l Fuego, F alklands/
M alvinas); Fow ler, 1945:1, fig (all records); Svetovidov, 1952:117; Idem , 1963:122; Clupea (A ntu) fuegensis:de
B uen, 1958:88; S p rattu s fu eg en sis - W h iteh ead , 1964:326 (key).
FAO Names :
E n - C h il e a n sprat.
47
D iagnostic Features : Low er ja w slightly projecting, gili cover w ithout bony
present on vomer, or only 1 or 2; belly w ith strong keel o f scutes; pelvic fínrays i 7,
behind dorsal fin origin, last two anal finrays not enlarged. No dark spots on flanks.
radiating striae, teeth rarely
insertion o f fin under or just
G eograp hical
D istrib u tion
W estern
South
A tla n tic (fro m a b o u t 40°S to T ierra del F u e g o , also
Falklands/M al vinas); records from the Pacific coast
(e.g. to V alparaiso, Chile, by M ann, 1954:130) may refer
to S tran g o m era.
H ab itat and B iology
M arine pelagic and
schooling fishes o f coastal waters. More data needed.
Size
15 cm.
To
18
cm
standard
length,
usually
to
Interest to F isheries : O f perhaps considerable
im portance, but separate statistic s not y et reported.
L ocal
quichay.
Nam es
L iteratu re
CHILE:
Pechu
chalw a,
Sardina
De Buen (1958 - synonymy only).
R em arks : Perhaps confused in the field w ith the
possibly sym patric S tran g o m era b e n tin c k i, from w hich
it can be distinguished by its more sharply keeled belly
and fewer low er gillrakers (less than 50, cf. more than
60) w hich do not overlap the upper gillrakers.
C LU P S pratt 5
S p r a ttu s m u elleri (K lu n z in g e r, 1880)
Clupea m u elleri K lunzinger, 1880,
collection.
Sitzber.A kad.W iss.W ien. 80(1):416 (New Zealand, Ferdinand von M ü lle r
Synonym s : C lu p ea m uelleri:Regan, 1916:228;
Sprattus m uelleri:W hitehead, Smith & R obertson, in press
(key, d ia g n o sis, b io lo g y , sy n o p sis); g iv en as Sk antip o d u m in m ost lite ratu re after R egan.
FAO Names
En - N e w Z ealand sprat.
s tr ia e
D iagnostic Features : Body m oderately
deep, its depth from 24 to 31% o f standard
length (m ean 26% ); gili cover w ithout bony
radiating striae: toothplate on tongue narrow,
pointed at each end, its w idth about 4 o r 5
tim es
in length, lateral
teeth
m oderate!
scales w ith o u t fine striae on ex p o sed p o ste rio r
parti vertebrae 43 to 47; back green/grey
w h e n fr e s h .
¿ flC
. ■t>- -
I'ljJ
-,
-if'fv.l' S 1. . . .
a
to o th p la te on tongue
Geographical D istribution : N ew Zealand (east­
ern and w estern coasts o f the N orth and the South
Islands, possibly south to Foveaux Strait and even to
A u k lan d Islan d , nearly 51 °S).
H ab itat and B iology : M arin e, pelagic and
schooling fishes o f coastal w aters, from beaches down
to 110 m or more. A n apparently long spawning season
(July to January) around the South Island, but not a
fractional spaw neri eggs present in m id-sum m er near
Clutha R iver m outh (but sam ples may also have in­
c lu d e d S^ a n tip o d u m ). M o re d a ta n ee d ed .
Size :
10 cm.
To ab o u t 13 cm stan d ard len g th , u su ally to
In terest to F ish eries :
b u t p e rh a p s o f lo c a l in te re s t.
N o s e p a r a te s ta ti s ti c s ,
100 °
120 °
140°
160°
180°
L ocal N am es : L ite r a tu r e :
press - synopsis).
S m ith & R o b e rtso n (1981 - sum m ary o f b reed in g d ata)i
Sprattus novaehollandiae
W h iteh ead , S m ith
(V a le n cie n n e s, 1847)
M eletta n o v ae-h o lla n d ia e V alen cie n n e s, 1847, H ist.n a t.p o iss.
&
R obertson
(in
C L U P S p ra tt 3
20:376 (Port Jackson, A ustralia).
S y n o n y m s:
C lu pea (P o m o lo b u s) b asse n sis M cC u llo ch , 1911, Z o o l.R e s.E n d e a v o u r, 1:16, pi. 4, tig. 2 (Bass
S tra it and T a sm a n ia ): C lu p e a b a s s e n s i s : R e g a n . 1916:5; M u n ro , 193824, sp. 166: S p ra ttu s b a sse n sis S v eto v id o v ,
1952:106: Idem . 1963:106 in E n n g lish ): S p rattu s n o v a e h o lla n d ia e - W h ite h e a d , 196721 (ty p e ): I d e m .1964:327 (key,
synonymy).
FAO N am es:
E n - A u stra lia n sprat.
49
Diagnostic Features : Lower ja w slightly projecting, gili cover w ithout bony radiating striae, teeth absent
on vom er; belly not sharply keeled , scutes rounded; p elvic finrays i 7, insertion o f fin a little before dorsal fin
origin, last two anal fin ra y s not enlarged. Overlaps ranges o f Sardinops (series o f spots along flank, bony striae on
gili cover) and Hyperlophus (scutes present on back before dorsal fin).
G eographical
D istrib u tion
T asm ania,
Strait, southeastern A ustralia north to Sydney.
B ass
H abitat and B iology :
M arine pelagic and
s c h o o lin g f is h e s o f c o a s ta l w a te r s , a p p e a r in g o f f
Tasmanian shores in large schools (especially in A ugustN ovem ber) and often entering estuaries (e.g. o f Tamar
and D erw ent rivers; said to ascend the form er as far as
Launceston in M arch fide B lackburn, 1941). M ore data
needed.
S iz e : T o a b o u t 14 cm s ta n d a r d le n g th .
40°
In te r e s t to F is h e r ie s :
but schools seasonally large.
No
se p a ra te
s ta tis tic s ,
L o c a l N a m e s : A U S T R A L IA : S p ra t ( T a s m a n ia ) .
Literature :
Blackburn (1941 - as Clupea
S p rattu s sp rattu s (Linnaeus,
100°
120°
140°
160°
180°
bassensis, taxonomy, biology summarized briefly).
1758)
C LU P S p ratt 1
C lupea sprattus Linnaeus, 1758, S y s t.n a t.. 10th ed.:318
(Europe).
Synonym s
C lupanodon phalerica R isso, 1827:425; C lupea latulus C uvier, 1829:318; C lupea papalina
B o n a p a rte , 1845:34; C lupea schon ev eld ii Kroyer, 1846:193; S p ra te lla p u m ila V a le n c ie n n e s, 1847:357; M e le tta
v u lg a ris V a le n c ia n e s, 1 8 4 7 :3 6 6 ; C lu p e a sp ra ttu s b a ltic u s G. S c h n e id e r, 1 9 0 4 :6 6 ; C lu p e a su lin a e A n tip a , 1 9 0 6:38;
Spratella serdnica N ikolsky, 1923:2; Sprattus sprattus - Svetovidov, 1952:107, pi. 1, fig. 3; Id em , 1963:111 (in
English); D em ir, 1965:unp. (synopsis); CLOFNAM , 1973:104 (full synonymy); F N A M , 1 9 8 4:224, fig. (sy n o p sis).
FAO N am es :
E n - E u ro p ean
sprat.
50
D ia g n o stic F ea tu res : L ow er ja w slightly p ro jectin g , gili cover w ith o u t bony rad iatin g striae, tee th rarely
p resent on vom er; belly w ith a strong keel o f scutes; p e lv ic fínrays i 6 (rarely i 7), in sertio n o f fin u n d er or
before the dorsal fin origin, last two anal finrays not enlarged. No dark spots on flanks. See C L U P S p ra tt 1,
Fishing Area 37.
G eographical D istribution : N ortheast A tlantic
(from N o rth Sea and B altic south to M oro cco ;
also
M editerranean, A driatic, B lack Sea).
H abitat and Biology :
M a r in e p e la g ic an d
usually inshore schooling fish es, som etim es entering
estu aries (especially the ju v e n ile s) and to leratin g sali­
n itie s as lo w as 4°/oo;
stro n g m ig ra tio n s b etw een
w inter feeding and sum m er spaw ning grounds. F eed s
on p la n k to n ic c ru s ta c e a n s .
S om e sp aw n in g alm o st
throughout the year, near to the coast or up to 100 km
out to sea, m ainly in spring and sum m er, the young
drifting inshore. M ove to the surface at night.
Size :
length.
To 16 cm , u su ally to 12 cm stan d ard
In terest to F ish eries : O f im portance in N orth
Sea, B altic and o ff N orw egian coasts, w ith a total o f
327 420 to n s in 1983; m ore th a n tw o th ird s o f th is w as
fished by D enm ark and N orw ay. M ed iterran ean and
Black Sea catches in 1983 were 43 193 tons, mostly by Yugoslavia and Bulgaria. C aught in traw ls or driftnets, or
driven up N orw egian Ijords by nets and penned until needed by the canning factories (sold as ‘brislings’); ju v e n iles
sold as ‘white bait’ (often mixed with juvenile herrings).
L ocal N am es :
1970:56).
M any v arian ts on th e nam es S prott or S prat, E sp ad in ,
P ap alin a and B rislin g (see Bini,
L itera tu re : Svetovidov (1952, 1963 - Russia); D em ir (1965 - biology, synopsis); B anarescu (1968 - Black
Sea); W heeler (1969 - UK); Bini (1970 - M editerranean); FNAM (1984 - synopsis).
R em a rk s :
subspecies:
(a)
(b)
(c)
Sm all d ifferen ces, m ainly in num bers o f p o st-p elv ic scutes, have been used to d efine three
R
sprattus sprattus: average post-pelvic scutes more than 11.5; A tlantic and N orth Sea coasts.
R sprattu s p h a le ric u s: av erag e p o st-p elv ic scutes no t m ore th an 11.3; M ed iterran ean , A d ria tic ,
Sea
S. sprattu s b a ltic u s: average post-pelvic scutes less than 11.5; B altic Sea.
B lack
click for n e x t p a g e
c lic k fo r p r e v io u s p a g e
51
Clupeonella Kessler, 1877
CLU P Clupnla
C lupeonella K essler, 1877, Ryby A ralo-K aspiisko-Ponti-cheskoi O blasti:187, pi. 6, fig. 24 (type: larva o f C.
grim m i K essler, 1877).
D iagnostic Features : A bsence o f a pterotic bulla
(bony dom e on floor o f pre-epiotic fossa), distinguishes
C lu p e o n e lla fro m
a ll o th e r c lu p e in e g e n e ra e x c e p t
Sprattus (w hich has the pelvic fin origin below or in front
o f the dorsal fin origin, and the last two anal finrays not
enlarged).
F ro m o th e r c lu p e in a e th a t o c c u r sym patrically, C lupeonella species differ in having no notch at the
c e n tre o f th e u p p e r ja w ( d is tin c t n o tc h in sp e c ie s o f
A losa).
B iology,
H abitat and
D istrib u tio n
P elagic,
schooling, in b ra c k is h w a te r and e u ry h a lin e or p u re ly
fresh-w ater, some species anadrom ous. Found only in Sea
o f M arm ara, B lack Sea, Sea o f A zov, C aspian and certain
rivers afflu en t to these w aters: ap p aren tly not entering
M ed iterranean.
pre-epiotic fossa
no pterotic bulla
In terest to F ish eries :
O ne sp ecies (C. c u ltiv e n tris) is am ong the top co m m ercial fish es o f the Sea o f
A zov: other species o f seasonal or local interest.
S p ecies :
C.
C.
C.
C.
Following Svetovidov (1952, 1963), there are 4 species:
abrau (M alyatskii, 1930), M editerranean region
c u ltriv e n tris (TSfordmann, 1840), M e d iterran ean reg io n , C asp ian Sea
engrauliform is (Borodin, 1904), Caspian Sea
grimm i K essler, 1877, C aspian Sea.
Clupeonella abrau (Malyatskii,
1930)
H arengula abrau M alyatskii,
Novorossiisk.
1930,
CLUP Clupnla
Trudy azov.-cherno-m orsk ryb.K hoz.Stantsii, 6:65
1
(Lake A brau, near
Synonym s : C lu p eo n ella m u h lisi N eu , 1934: C lu p eo n ella ab rau - S v eto v id o v , 1952:208, pi. 7, fig. 3: Id e m .
1963:228, pi. 7. fig. 3 (in English).
FAO N am es :
E n - A b rau sprat.
D iagnostic Features : Body m oderately slender, its depth about 19 to 23% o f standard length: head short
and narrow , in ter-o rb ita l w id th not m ore th an 17.5% o f stan d ard len g th : belly sharply keeled , w ith 23 to 26
scutes G illrakers 39 to 50. Pectoral fin tips pointed.
52
G eographical D istribution :
Lake A b ra u (fresh­
w ater, lan d lo ck ed , at 70 m above sea lev el, near to
N ovorossiisk);
not L ake A p o ly o n t (T u rk ey ), w hich is
linked to Sea o f Marmara (M eric, 1984).
H abitat and B iology : Freshw ater, pelagic and
schooling.
Feeds on cru stacean s (ju v en iles feed on
copepods, r o t if e r e g g s , p la n ts ) .
B re e d s M a y to
O ctober, eg g s p e la g ic , d eveloping r a p id ly b e fo re
sinking to bottom . M atu rity after one y ea r, at 3.5 to
4.5 cm.
Size :
To 9.5 cm standard length (or 6 cm in
Lake A buliond).
In terest to F ish eries : N il.
Local Nam es :
Sardel’ka
L iterature :
USSR:
A b ra u sk a y a ty u l’ka,
Svetovidov (1952, 1963).
C lupeonella cultriven tris (N ordm ann, 1840)
C LU P C lupnla 1
C lupea cultriventris N ordm ann, 1840, Faune pontique, 3:522 (northern coast o f Black Sea).
Synonym s : C lu p ea d e lic a tu la N o rd m an n , 1840:524 (p re -o c c u p ie d by C lu p ea d elic a tu la B en n e tt, 1831
Spratelloides delicatulus); C lupeonella d e li c a tu l a :Svetovidov. 1952:194, pi. 8, figs 1, 2, 3; Id e m . 1963:212, pi.
figs 1, 2, 3 (in E nglish); CLO FN A M , 1973:101 (full synonym y); FN A M , 1984:274, fig. (synopsis)
FAO Names :
E n - B lack Sea sprat.
D iagnostic Features : Body m oderately deep, its depth about 21 to 27% o f standard length; head short and
w ide, inter-orbital w idth at least 17.5% o f standard length; belly sharply keeled, w ith 24 to 29 scutes. G illrakers
49 to 62. Pectoral fin tips pointed.
53
G eographical D istribution :
B lack Sea (n o rth ­
w e ste rn p a rts ), S ea o f A z o v and C a sp ia n S e a , also
most o f the affluent rivers o f the area, reaching as far
as 60 km inland.
A lso , L ake P alaeo sto m i (B ulgaria)
and in Bay o f Feodosiya (Rom ania), also Lake A polyont
(Turkey).
H abitat and B iology :
P elag ic,
eu ry h aline,
e s s e n tia lly a b ra c k is h w a te r sp e c ie s (to le ra tin g s a li­
nities up to 34°/oo), but w ith sem i-anadrom ous and
purely freshw ater forms in rivers and lakes; migratory
b e tw e e n w in te r o r a u tu m n fe e d in g a n d s u m m e r
spaw ning grounds. Feeds on Z ooplankton. B reed s in
early sum m er in Sea o f A zov (peak in M ay), and from
about M ay in the low er reaches o f rivers (D n e ip e r,
Dneister).
Size :
10 cm.
To 14.5 cm standard length, usually to
Interest to Fisheries : O f m a jo r im p o r t a n c e in
Sea o f A zo v (o n e o f th e to p c o m m e rc ia l fish e s);
sto c k s d e p le te d in s o u th e rn p a rt o f C a sp ia n S ea (Coad, 1980:71 - C d e lic a tu la). The to tal catch in 1983 w as
396 731 tons.
Local Names
(a)
(b)
T z a tz a ;
Svetovidov (1952, 1963);
Literature
R em ark s
B U L G A R IA :
ROMANIA:
G ingirica;
U S S R : Tyul’ka.
M eric (1984 - gro w th , rep ro d u ctio n , d istrib u tio n ).
: Svetovidov (1952:194; 1963:212) re c o g n ized 2 subspecies, based on pectoral and pelvic fin length:
CL cu ltriv e n tris c u ltriv e n tris : p ecto ral and p elv ic fins long (17.5 to 21.5% and 11.5 to 14% o f stan d ard
length respectively); Black Sea, Sea o f Azov.
CL c u l t r i v e n t r i s c a s p i a : p ecto ral and p elv ic fin s sh o rte r (15.5 to 19% and 8.5 to 12.5% o f stan d ard
length resp ectiv ely ); Caspian Sea and basin.
C lupeonella engrauliform is (Borodin, 1904)
Clupea
engrauliform is
CLU P C lupnla 3
Borodin, 1904, V est.R y b o p ro m y sh le n n o sti. 19(6):335 (Buinaksh, central part o f Caspian
Sea).
Syn onym s :
English).
FAO N am es
C lu p eon ella
en g rau lifo rm is
: En - A n c h o v y
- Svetovidov, 1952:205, pi. 7, fig. 2; Idem . 1963:225, pi. 7, fig. 2 (in
sp rat.
* 1
7"
D iagnostic Features : Body slender, its depth about 16 to 19% o f standard length; head short and w ide,
in ter-o rb ital w idth 16 to 18.5% o f stan d ard len g th ; belly ro u n d ed , w ith 23 to 31 scutes. G illrak ers 56 to 67.
Pectoral fin tips pointed.
54
Geographical Distribution : Caspian Sea (middle
and southern parts).
H abitat and Biology : M ainly in open sea, only
occasionally
ap pro ach in g
th e
shore,
so m e tim e s
m assing in large schools, risin g to the surface in the
spring m onths, but descending to as m uch as 78 m in
la te s u m m e r ( a p p a r e n tly r is in g a g a in in O c to b erN o v e m b e r, b u t d e s c e n d in g o n ce m o re in w in te r).
B reed from end-A pril to N o v em b er, m ostly in July,
interm ittently and in the open sea.
Size : To 15.5 cm standard length, usually 11.5
to 12.5 cm (females a little larger than males).
Interest to F ish eries : C aught together w ith
C. c u ltriv e n tis c a sp ia by c o a sta l fis h e rie s or s e p a ­
ra te ly by lig h t fish in g in o p e n sea.
M ay rival or
e x ceed c a tc h e s o f CL c u ltriv e n tris c a s p ia , a lth o u g h
sto c k s cla im e d to be d e p le te d in s o u th e rn p a rt o f
C aspian Sea (C oad, 1980:71).
L ocal N am es : USSR: A nchousovidnaya ty u l’ka.
L iteratu re : Svetovidov (1952, 1963).
C lu p e o n e lla grim m i K e ssle r, 1877
C LU P C lupnla 4
C lu p e o n e lla grimm i K e ssle r, 1877, R y b .A ra lo -K a p iis k o i-P o n tic h e s k o i O blasti:187, pi. 6, fig. 24 (larv ae,
central part o f C aspian Sea).
Synonym s :
C lu p eo n ella grimmi - Svetovidov, 1952:209, pi. 7, fig. 1; Idem , 963:230, pi. 7, fig. (in English).
FAO N am es :
E n - S ou th ern C asp ian sprat.
D ia g n o stic F eatu res : Body m oderately slender, its depth about 17 to 22% o f standard length; head lo n g
and narrow, inter-orbital w idth 13 to 15% o f standard length; belly sharply keeled, w ith 26 to 32 scutes. G illrakers
42 to 51. Pectoral fin tips rounded.
55
G eographical D istribution
in south, absent in north).
Caspian Sea (m ainly
H a b ita t and B io lo g y
In open sea, not
approaching shores; in w inter, found mainly in south­
ern part, m igrating to m iddle parts o f sea in spring
( e s p e c ia l ly th e e a s te r n h a lf ) , r e tu r n in g s o u th in
N ovem ber-D ecem ber; occurs in upper layers in M archA p ril, b u t d e sc e n d to 16 to 32 m in su m m e r and
autum n.
F e e d s m ain ly on c o p e p o d s, also p e la g ic
m ysids
and
sm all
fish es.
B reeds
in
JanuarySeptem ber, interm ittently and in the open sea.
Size ! To 14.5 cm standard length, usually 10.5
to 11 cm (females a little larger than males).
Interest to F isheries : O f potential value, but
n o t c a u g h t by c o a sta l fis h e rie s ;
sto c k s said to be
d e p le te d in s o u th e rn p a rt o f C a sp ia n S ea (C o a d ,
1980:71).
Local Names
Literature :
: USSR: B olsheglazaya ty u l’ka.
Svetovidov (1952, 1963).
C L U P S ard i
S a r d in a A n tip a , 1906
A rengus C ornide, 1788, E n s .h is t.p e c e s ... G a lic ia :91 (suppressed, O pinion
Sardina A n tip a, 1906, D enkschr.A kad.W iss.W ien, 78:54 (after Sardina A n tip a,
A ntipa, 1906).
D ia g n o stic F ea tu res :
Resem bles Sardinops (non-European)
in having low er part o f gili cover w ith distinct bony striae radiating
dow nw ard (in ad d itio n to the norm al fleshy rad iatin g can als), but
low er g illrak ers not shortened at angle o f first arch, m ax illa not
reaching to eye centre and scales on flank o f uneven sizes (sm aller
scales hidden beneath the larger ones).
799, Int. C om m . Z ool. N o m e n c l.).
1904) (type: Sardina dobrogica
gillrackers
angle o f
gili arch
B iology, H abitat and D istribution :M arine, pelagic and
schooling fishes, esp ecially o f coastal w aters; restricted to N o rth ­
ern H em isphere (northeastern A tlantic and M editerranean region).
Interest to F ish eries
A bundant in some areas and o f
considerable comm ercial im portance, both as adults (pilchards) and
as juveniles (sardines).
Sp ecies :
A single sp ecies r e c o g n i z e d :
bony
striae
Sk p ilchardus (W albaum , 1792), n o rth eastern A tla n tic , M ed i­
te rra n e a n .
Sardina pilchardus
(W alb au m ,
1792)
C LU P Sardi 1
C lupea pilchardus W albaum , 1792, in Artedi, G en.pise.:38 (C ornw all, on P ennant, 1769).
Synonym s
A rengus m inor C ornide, 1788:91 (suppressed by O pinion 799, Int. Com m .Z ool.N om encl.);
C lupanodon sardina R isso, 1827:452 (N ice); C lupea sardina (cheironym ?):L ow e, 1837:189 (M adeira); C lupea
laticosta Low e, 1843:90 (M adeira); C lupea pilchardus var.2 sardinia:Günther, 1868:440 (Madeira); Sardina
dobrogica A ntipa, 1906; Sardina pilchardus - Svetovidov, 1952:188, pi. 6, fig. 2; Idem , 1963:205, pi. 6, fig. 2;
C LO FN A M , 1973:102; FNAM, 1984:276, fig. (synopsis); CLOFETA, in press.
56
FAO N am es
En - E u ro p ean pilchard.
w ï/m
m
D iagnostic Features : Body subcylindrical, belly rather rounded (but body more com pressed in juveniles).
Blind margin o f gili opening sm oothly rounded (w ithout fleshy outgrowths); 3 to 5 distinct bony striae radiating
dow nward on low er part o f operculum ; low er gillrakers 44 to 106, not becom ing shorter at angle o f first gili arch,
the upper series not overlapping the lower. Pelvic fin insertion well behind dorsal fin origin; last two anal finrays
enlarged. A series o f dark spots along u p p er flan k s, som etim es w ith a second or even th ird series below . See
CLUP Sardi 1, Fishing Areas 34, 47 (in part):
G eographical D istribution : Coasts o f eastern
N o rth A tla n tic , fro m Ic e la n d (ra re ) and N o rth S ea,
southward to Bay de G o ré e Senegal (14°43’N ) (ex ten ­
sion o f southern limit by about 15° m the last decade);
also M ed iterran ean (com m on in w estern p art and in
A d riatic, rare in eastern p art), Sea o f M arm ara and
B lack Sea.
H abitat and B iology : C oastal, pelagic, usually
at 25 to 55 or even 100 m by day, rising to 10 to 35 m
at night, schooling, m igratory, Feeds mainly on p la n k ­
tonic crustaceans, also larger organisms. B reeds at 20
to 25 m, near the shore or as much as 100 km out to
sea fro m A p ril (E n g lis h C h a n n e l), Ju n e to A u g u st
(N orth Sea, also B lack Sea), S eptem ber to M ay (o ff
E uropean coasts o f M ed iterran ean ) and N o v em b er to
June (o ff A frican coasts o f M editerranean).
Size :
20 cm.
To 25 cm stan d ard len g th , u su ally to
Interest to F isheries : The total catch for 1983
202 599 tons; A rea 34, eastern central A tlantic, 484 061
p ilchardu s w as the fo u rth m ost im p o rtan t co m m ercially
purse seines and lam paras (lig h t fish in g ), also g illn ets,
bottom trawls (French M editerranean coast).
w as 930 310 tons (A rea 27, eastern N orth A tlantic,
tons; A rea 37, M editerranean, 243 650 tons). Sardina
ex p lo ited clupeoid species in th at year.
C augth w ith
beach seines, trap nets and occasionally high opening
Local Nam es : V ariations on Sardele, Sardina, Sardine, etc. (see CLU P Sardi
G óm ez L a rra ñ e ta , 1960:140).
1, Fishing A rea 37; also
L iteratu re : Svetovidov (1952, 1963 - Russia); Gómez Larrañeta (1960 - biology, synopsis);
- B lack Sea); W heeler (1969 - U K); Bini (1970 - M editerranean), FN A M (1984 - synopsis).
B anarescu (1968
R em arks :
A uthors (e.g. Svetovidov, 1952, 1963) have often re c o g n ized tw o subspecies, based m ainly on
gillrakers counts and head length, but the separation is not satisfactory:
(a)
(b)
p ilc h a r d u s p ilchardus: lo w er g illrak ers m ore th a n 60, head len g th 20 to 23% o f standard length;
A tlantic (B ergen to G ib raltar).
R p ilch ard u s sardina: low er gillrakers 44 to 70 (but som etimes to 106), head length :18.5 to 2L 0% o f
stan d ard len g th ; M ed iterran ean , B lack Sea, A tlan tic (G ib ralta r to C. B lan c). H ow ever, specim ens
from G orée Bay, Senegal, have 59 to 90 low er gillrakers and head length 24.514 to 28.2% o f standard
length (Freon & Stequert, 1978).
57
O ther races, form s, v arieties and p o ssib le su b sp ecies have been suggested fo r p o p u latio n s in the w estern
M ed iterran ean (review ed by G óm ez L a rra ñ e ta , 1960), o ff w estern S ahara (F u rn estin , 1955) and o f f M au ritan ia
(Maurin, 1968).
Sardinops Flubbs, 1929
CLU P Sardop
Sardinops Flubbs, 1929, Proc.C alif.A cad.Sei. (4)18(11):264 (type: M eletta caeru lea G irard , 1854). N o te th at
Article 30(a)(ii) o f the International Code considers all genera ending in -ops to be m asculine, hence caeruleus, not
cae ru le a , etc.
D iagnostic Features : R esem bles Sardina o f Euro­
pean w aters in having low er part o f gili c o v e r(o p e rc u lu m )
w ith d istin ct bony striae rad iatin g dow nw ard (in ad d itio n
to the normal fleshy radiating canals), but low er gillrakers
are shortened at angle o f first arch and o v erlap p ed by
upper g illrak ers, m axilla reach es or alm o st reaches eye
centre and scales on flank o f even size (no sm aller ones
hidden beneath). No other clupeoid fishes have bony striae
on the operculum .
Biology, H abitat and D istribution :
M arin e,
p ela­
gic and schooling fish es, e sp e c ia lly o f c o a sta l w a te rs;
a n titro p ical d istrib u tio n (e astern N o rth and S outh P acific,
southern A frica, A ustralia, N ew Zealand).
bony
striae
Interest to Fisheries :
The five species (or sub­
species) o f Sardinops contributed about 1/4 o f all clupeoid
fish e s c a u g h t, m ak in g th is th e m o st p ro d u c tiv e o f all
clupeoid genera (8 250 201 tons in 1983); Sardinops was
exceeded only by the Peruvian anchovy during the peak o f
the latter’s exploitation (1962 to 1971).
Species :
Svetovidov (1952:178;
1963:193) considered th at the published differences betw een the five
‘species’ o f Sardinops were not significant, except perhaps in the case o f the A ustralian pilchard; he preferred to
recognize fiv e s u b s p e c ie s o f S a rd in o p s s a g a x , ,at le a s t u n til m o re d e ta ile d c o m p a ra tiv e s tu d ie s w ere m ade.
R ecent (unpublished) electrophoretic studies o f proteins by D r Stuart Grant have tended to show that at least
ÍE
c a e ru le u s , S. n eo p ilc h a rd u s, ÍE m elan o stictu s and ÍE o cellatu s hardly d iffer g en etically . N ev erth e less, the five
populations are so widely s e p a r a te d geographically th a t any
exchange o f genetic material is either im possible or
highly unlikely. For this reason 5 species are given here:
ÍE
ÍE
ÍE
ÍE
ÍE
caeruleus (Girard, 1854), eastern N orth Pacific
m e la n o s t ic t u s ( T e m m in c k & S c h le g e l, 1 8 4 6 ), w e s te r n N o r th P a c if ic
neopilchardus (Steindachner, 1879), western South Pacific
ocellatus Pappe, 1854), southern A frica
sagax (Jenyns, 1842), eastern South Pacific.
Sardinops caeruleus (G irard, 1854)
M eletta
caerulea G irard,
C L U P S ardop 4
1854, Proc.A cad.nat.Sei.Philad., 7:138 (San Francisco).
Synonyms : A la u sa califo rn ica Gili, 1862:33;
S ardina sagax (p art):R e g an , 1916:13 (com bined w ith sagax.
m e la n o s tic ta and o c e lla ta ) ; C lem ens & Wilby, 1961:101, fig . 35 C anada, P acific); H art, 1973:100, fig. (C anada
P acific, synopsis); Sardinops caeruleus - H ubbs, 1929:265 (do ub ted species statu s, h o w ever); A h lstro m , 1960:417
(syno psis); Fitch & L av en b erg , 1971:54, fig. 16 (California); M iller & Lea, 1972:54.
FAO Names :
E n - C a lifo rn ia p ilc h a rd .
58
D iagnostic Features : The radiating bony striae on the operculum distinguishes this species from all other
clupeids in the area; in addition, Clupea pallasii has the pelvic fin insertion before the dorsal fin origin and lacks
spots on the flanks; Etrum eus te re s lacks scu tes along the belly; and A losa sapidissim a is deeper-body and has a
distinct m edian notch in the upper jaw.
G eographical D istribution :
A laska (southeastern coast)
south to C. San Lucas and throughout the G ulf o f California.
Habitat and Biology :
C oastal, pelagic, in large (or in the 60o
past very large) schools (up to 10 m illio n indiv id u als estim ated in
tim e s o f a b u n d a n c e );
m ig ra to ry ,
w ith a d e fin ite n o rth w a rd
m o v e m e n t b e tw e e n C a lifo rn ia an d B ritis h C o lu m b ia w a te rs in
su m m er and re tu rn (a u tu m n , w in te r).
F eeds on Zooplankton*»”
(chiefly sm all cru stacean s), also p h y to p lan k to n , m ainly by filte rfeeding; also by ‘pecking’ at individual animals. B reeds m ainly
o ff the southern C alifo rn ia co ast ab o u t 80 km o ffsh o re b e tw e e n 200
Point C onception and San D iego; January to June, but a peak in
A pril to M ay (at night); eggs pelagic; some individuals spawn in
th eir first year, b u t m ost in th e ir
second; scale studies suggest
that some fishes live 20 to 25 years.
20°
Size
!
To about 36 cm standard length; usually to 25 cm.
Interest to Fisheries :
L andings (variously 70 to 100%
from C alifornia) reached a peak in 1936 (791 100 tons), but from
1944 declined until the C alifo rn ia fishery co llap sed and a m o rato riu m w as declared
1983 was 31 601 tons.
(1967). The total catch for
USA: Pacific sardine (AFS list).
L ocal N am es
Literature :
A hlstrom (1960 - biology, fishery, synopsis); M arr (1960 - fishery); R adovich (1960 - fishery);
C lem ens & Wilby (1961 - Canada, synopsis); Fitch & Lavenberg (1971 - California, synopsis), H art (1973 - Canada,
synopsis, biology an excellent summary).
Remarks :
H ild eb ran d (1946:87) compared Peruvian and Californian pilchards and failed to
differences.
O f all the p ilch ard s, th ese tw o are m ost lik ely to be the sam e sp ecies, in w hich
P acific pilchard should be g iven to both.
N e v e rth e le s s , M ille r & Lea (1972:212) stated th a t
discovered significant otolith and other differences suggesting that Sk caeruleus and S. sag ax are
and not subspecies.
Sardinops melanostictus (Schlegel, 1846)
find significant
case the nam e
J.E. F itc h had
distinct species
C LU P S ardop 5
C lupea m elan o sticta T em m inck & Schlegel, 1846, Fauna Japonica, P oiss., pt. 5, inst. 13:237, pi. 108, fig. 3
(Japan).
Synonym s : Sardina sagax (part): Regan, 1916:13 (com bined w ith sagax, ocellata and caerulea); Sardinops
m elan o sticta - H ubbs, 1929:265; A non., 1960:213 (sy n o p sis); S ardinops sag ax m e la n o s tic ta : S v e to v id o v 1952:178,
pi. 6, fig. 1; Idem . 1963:193, pi. 6, fig. 1.
FAO N am es :
En
Ja p a n e s e p ilc h a rd .
59
D iagnostic Features : The radiating bony striae on the operculum distinguish this fish from all other
clupeoids in the area; in addition, species o f Sardinella and H erklotsichthys have two fleshy outgrowths on the
hind margin o f the gili opening and no series o f dark spots along the flank (Am blygaster sirm
has similar spots, but
seems to be rare north o f Taiwan Island, possibly to O kinawa); Clupea pallasii has the pelvic fin insertion before
the dorsal fin origin and lacks spots on the flank; and E tru m eu s teres and D u ssu m ieria elo p so id es lack scutes
along the belly, also no spots.
___. J Ä T ’
G eographical D istribution : U SSR (w estern coast o f
Sea o f Japan, rarely to Sea o f O khotsk near eastern coasts
^ c
jo f S a k h a lin and in B e rin g S ea to c o a sts o f K a m c h a tk a
i\
Peninsula, absent in w inter), Japan (all coasts), the K oreas
Ac*
(east coast only), China (Taiwan Island, Hong Kong).
JfWM*
H ab itat and B iology : C oastal p elagic, form ing
large schools, m igratory, m oving northward in sum m er and
tending also to move more inshore, the reverse as tem pera- ;
tures begin to drop. F eeds m ainly on Z ooplankton, e sp e­
c ia lly c o p e p o d s, b u t a lso p h y to p la n k to n . B re e d s fro m
D ecem ber to the beginning o f M ay, earlier in the southern
than the n o rth ern parts o f ran g e, in bays and in coastal
parts o f open sea; fishes mostly mature in second year.
Size
20 cm.
To
24
cm
standard
length,
usually
15 to
\
V '
V?
»
«»asfcj.
Í
.
5
«
Interest to F ish eries : One o f the m ost im portant
com m ercial fishes o f Japan. The total catch for 1983 was
4 219 677 tons. R ecord catches in 1934-39 (over 2 000 000 tons) w ere succeeded by a sharp decline around 1941,
w ith only partial recovery from 1945 and then further decline.
Fluctuations in the catches and their relationship
to the Kuroshio current were reviewed by Kawasaki (1979) and Watanabe, Honjo & O kutani (1979).
L ocal N am es : JAPAN:
local vernacular names.
M aiw ashi;
THE
KOREAS:
Chong-o-ri;
USSR:
Ivasi;
see A non.
(1960:217)
for
L iterature : Svetovidov (1952, 1963 - full synopsis o f biology); also, see A non. (1960 - biology, synopsis);
N akai (1960 - fisheries); K urita (1960 - fisheries); U da (1960 - fish eries); H ayashi (1960 - fisheries). A great deal
o f Japanese literature is available, sum marized to 1959 by the papers cited here.
Sardinops neopilchardus
C lu p e a
A ustralia).
n e o p ilc h a rd u s
(Steindachner, 1879)
S te in d a c h n e r,
C LU P Sardop
1
1879, D e n k s c h r.A k a d .W is s .W ie n . 41(1): 12 (Hobson’s B ay, V ictoria,
Synonym s :
C lupea lata R ichardson & G ray, 1843:221 (nom en nudum ); Sardina n e o p ilc h a rd u s :R egan.
1916:14, pi. 1, fig. 1 S ardinops neopilch ard u s - H ubbs, 1929:265; B lack b u rn ,
1941 (biology); Id em . 1949:9
(biology); Idem , 1960:247 (synopsis); Baker, 1972:15 (New Zealand).
FAO N am es : En - A ustralian pilchard.
60
D iagnostic F eatures
The radiating bony striae on the operculum distinguish this fish from all other
clupeids in the area;in addition, species o f Sardinella and H erklotsichthys have two fleshy outgrow ths along the
hind m argin o f the gili opening and no series o f dark spots along flank (A m blygaster sirm has sim ilar spots, but
occurs o ff th e n o rth ern co asts o f A u stra lia ); E tru m eu s teres lacks scutes along the belly. See C l-U P Sardop 1,
Fishing Areas 57, 71.
Geographical D istribution : A ustralia
(southern
coasts, to 24° S on the w estern side and to 25° S on the
eastern side, reachin g so uthw ard to 43°S o ff eastern
Tasm ania) and N ew Z ealand (north to 34°S, down
entire eastern and m ost o f w estern coasts, also Cook
Straight, southward to A uckland Island at 57°S).
H abitat and B iology : Coastal pelagic, forming
large schools (at surface during breeding season and in
southern N ew South W ales at surface also w hen ju v e ­
niles leave bays to jo in the adult schools). Feeds
mainly on crustaceans (copepods, ostracods, eu p h au siids, m ysids), also mollusc larvae and phytoplankton
(diatom s). Breeds in spring and sum m er in southern
part o f range, and in w inter in northern part, appa­
rently related to seasonal m ovem ent o f the lim iting
14° and 21 °C isotherm s. M ature at 8 to 13 cm, mostly
at 12 cm or above in New Zealand.
Size
: To 21.3 cm standard length, usually to 18 cm.
Interest to F isheries : Insignificant, but som etim es used for bait. C aught w ith various form s o f seine net.
Local N am es : AUSTRALIA: Pilchard; N EW ZEALAND: Pilchard.
L iterature : B lackburn (1941, 1949, 1950, 1951,
breeding, growth).
1960 - biology, fisheries); B aker (1972 - N ew Zealand,
R em arks : In N ew Zealand the species appears to grow larger (21.3 cm standard length; cf. 19.7 cm), has
slightly larger eggs and a higher mean num ber o f vertebrae (50.52; cf. 49 to 50.08 in various sam ples) (Baker,
1972:17).
Sardinops ocellatus
(Pappe, 1854)
C LU P
S ardop 2
Clupea ocellata Pappe, 1854, Svn.edible fish Cape o f Good Hope : 20 (Cape o f Good Hope).
Synonym s : Sardinops sagax (part) R egan, 1916:13, pi. 1, fig. 1 (com bined w ith sagax, m elanosticta and
c a e r u le a ); Sardinops ocellata - H ubbs, 1929:265; Smith, 1953:92, pi. 5 (as A rengus s a g a x in Smith, 1969:92);
M o n teiro , 1960:1105 (A n g o la ), CLOFETA, in press; SFSA, in press (southern Africa)
FAO Names :
En - S outhern A fric an pilchard.
61
%
" " ít
D iagnostic Features
: The radiating bony striae on the operculum distinguish this fish from all other
clupeids in the area; in addition, species o f Sardinella and H erklotsichthys have two fleshy outgrowths along the
hind margin o f the gili opening and no series o f dark spots along flank (A m b ly a s te r sirm has similar spots, but not
recorded from South A frican coasts); H ilsa k e le e has spots on flanks but is deeper-bodied and has a distinct
median notch in the upper jaw ; and species o f Etrum eus and D ussum ieria lack sc u te s along the belly. See CLUP
Sardop 2, Fishing A reas 34 and 51.
G eographical D istrib u tion : A ngola (B aia dos Tigres,
Porto A le x a n d re ) southward to Cape Town and north to N atal
(D urban); possibly to M auritius.
H abitat and B iology : C oastal pelagic, form ing large
schools, m igratory (but northern stocks perhaps rarely moving
south o f O range R iver, thus South A frican stocks separate; on
east coast, seasonal m igration northw ard to D urban in
June/July). Feeds on Zooplankton (m ainly copepods) when
young, but on phytoplankton (m ainly diatom s) from about 10 cm
standard length; a non-selective filter feeder. B reeds mainly
from Septem ber to February.
S i z e : To 28 cm standard length, usually to 25 cm.
Interest to F ish eries : The total catch for 1983 w as
110 229 tons (South A frica 61 814 tons; N am ibia 44 014 tons),
virtually all from the w estern and southern coasts. Small
catches by foreign nations, chiefly Poland.
L ocal
pelser.
N am es
SO U T H
A FR IC A :
Sud
A frik aan se
L iterature : D avies (1957 - biology, synopsis); Du Plessis (1960 - fishery); D e Jager (1960 - biology);
M athews (1960, 1964 - biology); King & M acleod (1976 - feeding).B iology and fishery data are given in many o f
the Investigational Reports o f the Sea Fisheries B ranch o f the South A frican D epartm ent o f Industries from 1954
onward.
Sardinops sagax (Jenyns, 1842)
C LU P
Sardop
3
C lupea sagax (Jenyns, 1842, Z o ol.voy.B eagle. fish es: 134 (Lim a, San Lorenzo Island, Peru).
Synonym s : A lo sa m u sic a G irard, 1854:199 (C hile); C lupea advena Phillippi, 1879:161, pi. 10 (Chile);
A rengus sa g a x :Fowler. 1945:3 (all Chile re fs );
Sardinops sagax - M ann, 1954:131; H ildebrand, 1946:86; de Buen,
1960:265 (synopsis); Bore & M artinez, 1981:unpaged (sy n o p sis) ; L eib le & A lv ea l, 1982:18 (synopsis).
FAO Names
En - South A m erican pilchard.
62
D iagnostic Features : The radiating bony striae on the operculum distinguish this species from all other
clupeids in the area; in addition. Ethm idium m aculatum has dark spots on the flank, but is deeper-bodied and has
a distinct m edian notch in the upper jaw ; Strangom era bentincki has a slender body, but lacks the spots and last
two anal finrays not enlarged.
Geographical D istribution : Peru and Chile (Sechura Bay at
5°S southward to M ocha Island, Chile, at 3 8 :3 0 ’S;
also Galapagos
Islands).
H abitat and B iology : C oastal pelagic, caught dow n to
depths o f about 40 iii, in sum mer at 16° to 23°C,
in w inter at 10°
to 18°C .
Form s large schools in the Peru Current. Feeds mainly
on planktonic crustaceans. B reeds tw ice in one year (at least o ff
C hile), from July to Septem ber and a lesser spaw ning from
February to M arch thro u g h o u t the area betw een A frica and
southw est o f A ntofagasta (eggs
also recorded o ff V alparaiso);
mature from about 24 cm.
Size
20 cm.
To
about
30
cm
standard
length,
usually
around
In terest to F ish eries
The total catch for 1983 w as
3 888 694 tons
(Peru
1 064 448 tons,
C h ile
2 823 424 tons).
C aught w ith purse seines; in 1979, there were 3 ships operating in
the northern zone o f Chile (A ric a , Iquique, A ntofagasta) and 41
from T alcahuano; and o f the total catch (1 618 937 tons), 90%
w as used for fishm eal or oil, 4% w as canned and the rest
marketed fresh or frozen.
Local N am es
CHILE:
Sardina española;
ECUADOR:
Sardina; PERU:
Sardina.
L iterature
H ildebrand (1946
- Peru); de B uen (1960 - biology, synopsis); A lberti
Antofagasta and Iquique); Bore & M artinez (1981 ; Chile, synopsis, fishery data).
(1960
-
fishery,
Rem arks :
De Buen (1958, 1960:270) recognized two subspecies and suspected that the G alapagos population
represented at least an isolated stock:
S^ sagax sagax: head length 27
or 28.5% o f standard length, pre-dorsal distance
length, pectoral fin length 53 to 55.5% o f head length; Peru.
47
to 50% o f standard
sagax m usica: head length 25.5 to 26.1% o f standard length, pre-dorsal distance 45.1 to 47.6% o f standard
length, pectoral fin length 51.5 to 58% o f head length; Chile.
c lic k for n e x t p a g e
c lic k fo r p r e v io u s p a g e
63
H arengula
C L U P H ar
V alenciennes, 1847
H aren g u la V ale n c ie n n e s, 1847, H ist.n a t.p o iss., 20:227 (type: H aren g u la
clu peola C u v ier, 1829).
H y rtlin u s F o w ler, 1958, N o tu l.N a t., (310):5 (type:
H arengula jaguana Poey).
D iagnostic Features : The presence o f a sm all toothed hypom axilla betw een the hind tip o f the p re-m ax illa and the expanded
blade o f the m axilla distinguishes H arengula from Sardinella and all
other clupeoid
genera, except the pristigasterids Pellona and P lioste­
ostom a, w hich have a long anal fin w ith more than 30 finrays (about
16 to 19 in H arengula), its base much longer than that o f the dorsal
fin (about equal in H arengula). Low er gillrakers less than 45 (m or
than 45 in adult Sardinella o f N ew World).
latu lu s V alenciennes = C lupea
H yrtlinus
a ltifo rm a F o w le r =
to o th ed
hypo-m axilla
Biology, H abitat and D istribution : M arine, pelagic and schooling fishes, especially
N ew W orld only (Atlantic and Pacific coasts), mainly in tropical and subtropical waters.
Interest to Fisheries
Species
H.
H.
H.
H.
o f coastal waters.
N o major fisheries, but contribute to clupeoid catches.
The m ost recent revision (Rivas, 1964) recognized 3 Atlantic and 1 Pacific species:
clupeola (Cuvier, 1829), w estern A tlantic
hum eralis (Cuvier, 1829), w estern Atlantic
jaguana Poey, 1865, w estern A tlantic
thrissina (Jordan & Gilbert, 1882), eastern P acific.
H arengula clupeola
CLU P H ar 1
(C uvier, 1829)
C lupea clupeola C uvier, 1829,
R é g n e anim .. 2nd ed., 2:318 (footnote, on C ailleu o f D uham el,
Traité gén, pêches, pt. 2, (3): pi. 31, fig. 3: reproduced by Storey, 1938: fig. 16).
1776,
Synonym s :
C lu p ea m ac ro p h th alm a R an zan i, 1 8 4 2:320, pi. 23, fig s 1-4 (? B ra z il); H a ren g u la latu lu s
V alenciennes, 1847:280, pi. 595 (? locality); A losa bishopi M üller & Troschel, 1847:675 (Barbados); Schomburgk,
1848:19 (repeat); H arengula clupeola - L- Storey, 1938:29; R ivas, 1950:289; FW N A , 1964:390, fig. 96; C ervigón,
1966:124, fig. 52; W hitehead, 1967:26 (H, latulus types, not from northern France, but from w estern A tlantic);
Idem , 1973a:38, fig. 10 (G u ian as); W hitehead & Bauchot, in press (types o f latulus).
FAO Names
E n - F a ls e h errin g .
ton gu e
D iagnostic Features : Tooth-plate on tongue and toothplate behind
it (b asih y al and basibranchial toothplates) broad, their w idth about 3 to 5
tim es in their com bined length, the basibranchial toothplate w ith a distinct
bulge in front. Low er gillrakers 28 to 34, usually 30 to 32. Scales fairly
strongly attached, not easily lost. N o dark pigm ent on dorsal fin tip. See
CLUP Har 1, Fishing Area 31.
to o th ­
p la t e s
g ill­
ra k e r s
to o th p lates on
floor o f m ooth
64
G eographical D istribution :
G u lf o f M exico,
southeastern Florida, B aham as, entire C aribbean and
W est Indies and southward to Brazil.
H abitat and B iology : Coastal, pelagic, estuaries
and lagoons, tolerating quite low salinities, schooling,
small individuals often occurring in num bers along
sandy beaches.
Size :
10 cm.
To 15 cm standard length, usually about to
Interest to F isheries : M ost frequently caught
for bait w ith seines and cast nets in estuaries and bays
(Rivas, 1 9 5 0 :2 9 2 ). C ontributed to the 543 tons o f the
‘H arengula’ catch for 1983.
Local N am es : USA: False pilchard (AFS list):
see also FW NA (1964:390) : VENEZUELA: C arapa­
cho na.
L iterature
H arengula h u m eralis
See under synonyms.
(C uvier. 1829)
C LU P H ar 3
C lupea hum eralis C uvier,
1829, R ègne a n im .. 2nd ed. 2 :3 1 8 (footnote, on Sardine de la M artinique o f
D uham el, 1776, Traité gén.pêches, pt. 2 (3) : pi. 31, fig. 4: reproduced by Storey, 1938: fig. 16).
Synonym s
: H arengula m aculosa V alenciennes, 1847:292 (M artinique): A losa apicalis M üller & T roschel,
18 47:675 (B a rb a d o s ): H a re n g u la sa rd in a P o e y , 1 8 6 0 :3 1 0 (C u b a ): H a re n g u la c a llo le p is G o o d e , in G o o d e & B ean ,
1 8 7 9 :1 5 2 (B e rm u d a ): H a re n g u la h u m e ra lis - S to re y , 19 3 8 2 8 :R iv a s , 1 9 5 0 :2 8 2 : F W N A , 1 9 6 4 :3 8 7 , fig . 95:
C erv ig ó n , 196 6 :1 2 2 , fig. 51: W h ite h e a d , 1 9 67:27 (ty p e: no ty p es found fo r IT m a c u lo sa ): Id e m . 1 9 7 3 a :3 5 , fig. 9
(G uianas): W hitehead & B auchot, in press (type o f m aculosa lost).
FAO N am es : En - R edear herring.
«'•y
“****
.■j
...... .............
to o th ­
p la te s
D iagnostic Features : Toothplate on tongue and toothplate behind it
(basihyal and basibranchial toothplates) very narrow , their w idth about 10
tim es in com bined length, the basihyal toothplate not bulging in front.
L ow er gillrakers usually 27 to 31. Scales thin, loosely attached, easily lost.
T ip s o f a n te rio r d o rsa l fin ra y s d ark .
g ill­
ra k e rs
to o th p late s on
floor o f m ooth
65
G eographical D istribution
B erm uda, Florida,
B aham as, C arib b ean , G uianas (b u t no reco rd s to the
south o f this; no B razilian specimens in the extensive
c o ll e c tio n s o f th e M useu de Zoología, S ao P aulo,
B razil).
H abitat and Biology
schoolin g.
C o a sta l,
p e la g ic ,
Size : To 17.2 cm standard length, usually about
10 cm.
In terest to F ish eries
C o n trib u ted
543 tons o f the ‘H arengula’ catch for 1983.
L ocal N am es : USA:
VENEZUELA: M anzanillera.
L iterature
H arengula jaguana
See
under
See
FW N A
to
the
(1964:387);
synonyms.
C LU P H ar 2
Poey, 1865
H arengula jaguana Poey, 1865, R e p e rt.F isic o -N a t.C u b a .
1:189 (Bahia de Jaqua, Cuba).
Synonyms : H arengula hum eralis V alenciennes, 1847:293 (St. D om ingo, Surinam, B razil;
by H arengula h u m e ra lis C uvier, 1829 fide W hitehead, 1967:30); H arengula pensacolae G oode
(Pensacola, Florida); Storey, 1938:33; R iv as, 1950:292 (6 s u b s p e c ie s; see re m a rk s); F W N A ,
Cervigón, 1966:125, fig. 53; H aren g u la m ajo rin a S torey, 1938:32; H y rtlin u s altifo rm a Fow ler,
H arengula jaguana - W hitehead, 1973a:41, fig. 11 (G uianas).
FAO Nam es
name preoccupied
& B ean, 1879:152
1964:393, fig. 97;
1958:6 (skeleton);
E n- Scaled herring.
tongue
to o th ­
p la te s
D iagnostic Features : Toothplate on tongue and toothplate behind it
(b asih y al and basibranchial toothplates) very broad, their w idth about 3 to 5
tim es in their com bined length, the basihyal toothplate w ith a very distinct
bulge in front. L o w er gillrakers 30 to 40, usually 32 to 39. Scales fairly
strongly attach ed , not easily lost. N o dark p igm ent on dorsal fin tip. See
CLUP IFar 2, Fishing Area 31.
g illrak ers
to o th p late s on
floor o f mouth
66
G eograph ical D istrib u tion
N ew Jersey (U SA ), also
Bermuda, southward to Florida, G ulf o f Mexico (abundant). Carib- 40°
bean to southern Brazil.
H abitat and B iology : C oastal, pelagic, schooling, perhaps
also in estuaries and bays, but often confused w ith H. clupeola 20°
(with w hich it occurs over most o f its range). Juveniles o f 2.9 to
4.4 cm standard length appeared in G uanabara B ay, R io de
Janeiro, in M ay and m igrated out in D ecem ber or M arch at 7.5 to
8.0 cm (Hubold & M azzetti, 1982). Breeds February to July (peak °°
May and June) around Miami.
Size : To 15 cm standard length, usually to 12 cm.
Interest to F isheries : C ontributed to the 543 tons o f the
‘Harengula’ catch for 1983.
L ocal N am es
FW NA (1964:393).
: USA:
Scaled
sardine
(AFS
list):
see also
L iterature : M atsuura (1972 - as H. pensacolae. B razil, eggs): H oude, Richards. & Saksena (19 - eggs,
larvae): H ubbold & M azzetti (1982 - B razil, ju v en iles .
R em arks : R ivas (1950:280, key) recognized the follow ing 6 subspecies: pensacolae, m ajorina and his own
proposed names floridana, caribbaea and pinensis. Subsequently (Rivas, 1963:395, i.e. FW N A) he accepted that
these forms intergrade in their defining characters -and separating on geographical grounds w ould be arbitrary.
N evertheless, specimens show differences in m orphom etric and m eristic features over the rather extensive range
o f the species and at least some subspecies may eventually be recognized.
H arengula thrissina
C LU P H ar 4
(Jordan & G ilbert, 1882)
Clupea thrissina Jordan & Gilbert, 1882, P ro c .U .S .n a tn .M u s.. 5:353 (Cape San Lucas, Mexico).
Synonym s
: H arengula peruana Fow ler & B ean, 1923:2 (Callao, Peru): R ivas, 1950:287, fig. 39: Peterson,
1956:177 (G u lf o f N ic o y a , Costa Rica):
H arengula thrissina - Flildebrand, 1946:88 (Peru): Rivas, 1950:287 (G ulf
o f C alifornia to A capulco, M exico): FW N A, 1964:387 (key: peruana kept separate).
FAO N am es : En - Pacific flatiron herring.
to n g u e
to o th ­
p lates,
D iagnostic Features : Toothplate on tongue and toothplate behind it
(basihyal and basibranchial toothplates) broad, th eir w idth about 3 to 5
tim es in their com bined lengths. L ow er gillrakers 28 to 34. L ast dorsal
finray not elongate (cf. O pistnonem a): anal finrays 15 to 17 (cf. m ore than
30 in species o f Pristigasteridae).
-»-gillrakers
toothplates on
floor o f mouth
c lic k for n e x t p a g e
c lic k fo r p r e v io u s p a g e
67
G eographical D istribution
Peru (at least to Callao).
Habitat and B iology
More data needed.
G u lf o f California to
Coastal, pelagic, schooling.
Size : To 8 cm standard length, usually to 5 cm.
Interest to
F isheries
: Probably
little.
Local N am es : USA: Flatiron herring (A F S list).
L iterature : See under synonyms.
Rem arks : Rivas (1 9 5 0 :2 8 6 -7 , also 1964:387, i.e.
FW NA) recognized H. peruana as a distinct species, but
it seem s more likely a subspecies or perhaps m erely a
southern form:
H. thrissina thrissin a: 42 to 44, usually 43 scales
in lateral series, 15 to 19 pre-dorsal scales: G u lf
o f California to Panama
H. thrissina peruan a: 38 to 42, usually 39 to 41
scales in lateral series, 11 to 14 pre-dorsal scales:
Panama to Peru.
Opisthonem a
C L U P O pi
Gili, 1861
O p isth o n em a G ili, 1861, P r o c .A c a d .n a t.S c i.P h ila d .:37 (type: O isthonem a thrissa Gili = M egalops
LeSueur). Filialosa F ow ler, 1944, M onagr.A cad.nat.Sci.Philad.. (6):207 (type: M eletta lib e rta te Günther).
oglina
D iagnostic Features : The filam entous last dorsal fin ra y distinguishes O pisthonem a from all other genera
o f the C lupeinae: it otherw ise resem bles Sardinella (but pelvic finrays i 7, cf. i 8 in all N ew W orld Sardinella )
and H arengula (w hich has a small toothed hypo-m axilla between the hind tip o f the pre-m axilla and the expanded
blade o f the m axilla). A filam entous last dorsal fin ray occurs in D orosom a
and certain other gizzard shads, but
mouth inferior.
Biology, H abitat and D istribution : M arine, pelagic and schooling. N ew W orld only (A tlantic and Pacific
coasts), mainly in tropical and subtropical waters.
Interest to
1983).
F isheries : M oderate catches o f both A tlantic
and P acific
species (45
745 tons com bined in
Species : The m ost recent revision (Berry & B arrett, 1963), recognized 1 A tlantic and 4 Pacific species, to
w hich Rivas (1972) added another A tlantic species (CF captivai) w hich is here considered merely a subspecies (see
remarks):
O.
O.
O.
O.
O.
oglinum (LeSueur, 1818), w estern A tlantic
berlangai Berry & Barrett, 1963, eastern Pacific
bulleri (Regan, 1904), eastern Pacific
libertate (Günther, 1867), eastern Pacific
medirastre Berry & Barrett, 1963, eastern Pacific.
R em arks :
The 4 eastern P acific species are only distinguished
by num bers o f low er gillrakers, w hich
increase w ith size o f fish: by excluding the Galapagos species (CFberlangai), there is little or no overlap in adult
gillraker counts.
68
Berry & Barrett, 1964
Opisthonema berlangai
CLUP
Opi 2
O pisthonem a b erlan g ai B erry & B arrett, 1964, B ul L in te r-A m . tro p . Tuna C o m m , 7(2): 119, fig. IB (G alapagos
Is la n d s).
Synonym s :
FAO Nam es
O pisthonem a lib e rta te : all G alap ag o s accounts.
En - G alap ag o s th read herring.
1■
"•*=•«
V V
\
D iagnostic Features : The filam entous last dorsal fin ra y distinguishes it from m em bers o f any other eastern
P acific clupeoid genus. L o w er g illrak ers 50 to 87 in fish es over 14 cm standard len g th (in term ed iate b etw een
O. medirastre with 41 to 69 and O. libertate with 63 to 110, but neither o f these occur in the Galapagos Islands).
0. li b e r ta t e
O .m e d ira s lre
-S T A N D A R D
LE N G T H -
Geographical D istribution : Galapagos Islands only.
H abitat and B iology : C oastal, pelagic, schooling.
Size : To 26 cm standard length, usually to 18 cm.
Interest to Fisheries : O f little interest.
Local N am es : N ot distinguished from other eastern Pacific species.
L iterature : Berry & B arrett (1964
- taxonom y only).
69
(Regan, 1904)
O pisthonem a bulleri
CLUP
Opi 3
C lupea (O pisthonem a) b u lle ri R e g a n , 1904, A n n .M a g .n a t.H is t., (7)8:255 (Las P eñ a s (P u e rto V a lla r ta ) , J a l is c o ,
M exico - but mixed w ith a specimen o f Cf libertate).
Synonym s : Probably included in at least some accounts o f CL libertate.
FAO N am es : E n - S le n d e r th r e a d
h e r r in a .
D iagnostic Features
The filam entous last) dorsal fin r a y s distinguishes it from m em bers o f any other
eastern Pacific genus. Separated from other eastern Pacific
Opisthonem a by having only 25 to 36 low er gillrakers
in fishes over 14 cm standard length (41 to 69 in CL medirastre. 63 to 110 in CL libertate: in CL berlangai there are
50 to 87 and it occurs only in the G alapagos Islands.
Q .lib erf a te
0 .b e r la n gai
O .m e d ira s tre
O .buU eri
-S T A N D A R D
LE N G T H -
Geographical D istribution :
P eru ).
E a ste rn P a c ific (M a z a tla n , S inaloa, M exico southw ard to Pta Sal and Pta P ic o s ,
H abitat and B iology :
C oastal, pelagic, schooling. A pparently the least abundant o f the eastern Pacific
sp e c ie s o f O p is th o n e m a . F e e d s on c ru s ta c e a n s an d p te ro p o d s.
Size
: To
19.6
cm
standard
In terest to F ish eries :
40 081 tons in 1983.
Local Nam es :
Literature :
length,
Presum ably
usually
to
18 cm.
contributes to the total O pisthonem a catch in the eastern Pacific o f
N ot distinguished from other eastern Pacific species.
Berry & B arret (1 9 6 4 - ta x o n o m y ), o th e rw ise u n c e rta in sin ce n o t se p ara te d fro m CL lib e rta te .
70
C LU P Opi 4
O pisthonem a libertate (G ünther, 1867)
M eletta lib e rta tis G ü n th e r, 1867, P ro c .z o o l.S o c .L o n d .' (3 ):6 O3 (L a L ib e rta d , El S alv ad o r; p a rt 3 issu ed A p ril
1867, not in 1866).
Synonym s : O pisthonem a libertate - Berry & B arrett, 1964:120, fig. l e (full synonym y; references to O.
b erla n g a i. O. bulleri and Cf m ed irastre exluded); Cobo & M assey, 1949:7, tig. 5 (E cuador, listed); Peterson,
1956:178 (G ulf o f N icoya, C o sta Rica); Chirichigno, 1963:14, fig. 6 (Puerto Pizarro and Puerto Rico, Peru).
FAO N am es : En - Pacific thread herring.
D iagnostic Features : The filam entous last dorsal finray distinguishes it from m em bers o f any other eastern
P acific clupeoid genus. S eparated from o th er eastern P acific O pisthonem a species by having 63 to 110 low er
g illrak ers in fish es over 14 cm standard len g th (41 to 69 in Q. m e d ira stre , only 25 to 36 in Q. b u lle ri; in O
b erlan g ai there are 50 to 87, but it occurs only in the G a lap ag o s Islands.
O .iifrertate
O .b e rla n qai
O .m e d ira s tre
O.bulleri
-S T A N D A R D
LE N G T H -
G eographical D istribution : E astern
southward to Pta Sal and Pta P ico s, Peru).
Pacific
(Santa
R osalita,
P acific
H abitat and B iology : C oastal, pelagic, schooling. A pparently the
species o f O pisthonem a. Feeds on crustaceans and pteropods.
Size : To 25 cm standard length, usually 12 to 18 cm.
coast
o f B aja
C alifornia,
M exico,
m ost abundant o f the eastern Pacific
71
Interest to F isheries : O f some im portance as a live bait until introduction o f purse seines for tuna.
Contributed to the total
O pisthonem a catch in the eastern Pacific o f 40 481 tons in 1983 (the m ajor part from
Ecuador).
L ocal N am es :
See
C erv ig ó n & Fischer (1979).
L iterature : Berry & B arrett (1964 - taxonom y only); earlier w orks probably com bined data for CF bulleri
and CF m edirastre, e.g. P eterson (1956 -b io l., etc.).
C LU P O pi 5
Berry & Barrett, 1964
Opisthonem a m edirastre
O p isth o n e m a m e d ira s tre B e rry & B arrett, 1964, B u ll.in ter-A m .tro p .T u n a
Panam a, also various localities from Los A ngeles to Sechura Bay, Peru).
C o m n i.. 7(2): 118, fig. la (G ulf o f
Synonym s : Included in at least some accounts o f CF bulleri and especially CF libertate (the only Pacific
species formerly recognized).
FAO
N am es
: En - M iddling thread herring.
aga
D iagnostic F eatures
The filam entous last dor­
sal fin ray distinguishes it from m em bers o f any other
eastern Pacific clupeoid genus. Separated from other
eastern Pacific O pisthonem a by having 41 to 69 lower
gillrakers in fishes over 14 cm standard length (63 to
110 in CF lib ertate, only 25 to 36
in CF b u lle ri:
in
0 . b e rla n g a i there are 50 to 87, but it occurs only in
the Galapagos Islands.
O .lib erfrate
O.berlanqai
60 \ g
O .m e d ira tirt
I
5
0 .b u tle ri
------------- STANDARD
I
!
2
1
I
1
6
1
6
1
10
LENGTH-------------
1
12
1
14
1
16
1
18
1
20
1
22
:+
24
•—
26
72
Geographical D istribution : E astern Pacific (Los
A n g e le s B ay , C a lifo rn ia so u th w a rd to S e c h u ra B ay ,
Peru).
H ab itat and B iology :
C oastal, pelagic,
schooling. Feeds on crustaceans and pteropods.
Size : To 22 cm standard length, usually
20 cm.
15 to
Interest to F isheries : O f some im portance as a
live bait until in tro d u ctio n o f purse seines for tuna.
C o n tr ib u te d to a to ta l O p is th o n e m a c a tc h in th e
eastern Pacific o f 40 481 tons in 1983.
Local N am es : N ot distinguished from the other
eastern P acific species.
L iterature
O. lib e rta te .
: U ncertain
Opisthonem a oglinum
since
not separated
from
120 °
100 °
80 °
(L eS ueur, 1818)
60 °
40 °
C LU P Opi 1
M egalops oglinum LeSueur, 1818, J.A cad.nat.Sci.Philad., 1:359 (N ewport, Rhode Island).
Synonym s : M egalops notata LeSueur, 1818:361 (G uadeloupe): Chatoessus signifer D eKay, 1842:264, pi. 41,
fig. 132 (N ew York): M eletta thrissa V alenciennes, 1847:380 (N ew Y ork, Santo D om ingo, G uadeloupe, M artinique:
not C lupea thrissa L innaeus, 1758): A lausa striata V alenciennes, 1847:429 (G uadeloupe): C hatoessus eum orphus
G osse, 1851: 290 (Jam aica): O p isth o n em a cap tiv ai R iv as, 1972:
(C o lo m b ia): O p isth o n em a og lin u m - FW N A ,
1964:381, fig. 94: Berry & B arrett, 1964:123 (re v isio n ); C e rv ig ó n , 1966:118, fig. 49: W hitehead, 1967:73, 75
(types o f W stria ta , M. th riss a ): Idem , 1973:46, fig . 12 (G uianas);
F ig u e ired o & M en ezes, 1978:22, fig. 25;
W hitehead & Bauchot, in press (type o f notata thrissa).
FAO N am es : En - A tlantic thread herring.
D iagnostic Features : The filam entous last dorsal fin ra y distinguishes it from all other w estern A tlantic
clupeoids except D orosom a, w hich has an inferior mouth. O therw ise, su p erficially resem bles species o f S ardinella
(but i 8 pelvic finrays in w estern A tlantic species, cf. i 7 in O pisthonem a) and H arengula (but small toothed hypomaxilla in upper jaw ). Lower gillrakers increasing initially w ith size o f fish, but stable at 28 to 46 after 8 cm
standard length.
c lic k for n e x t p a g e
c lic k fo r p r e v io u s p a g e
73
G eographical D istrib u tion : W estern A tlantic (G u lf o f
M ain e, B erm u d a, th ro u g h o u t G u lf o f M e x ic o , C arib b e a n and W est 40Indies to Santa C aterina, Brazil).
H abitat and B iology : Coastal, pelagic, schooling (but soli­
ta ry in d iv id u a ls re p o r te d ) , p ro b a b ly n o t e n te rin g w a te r o f lo w
sa lin ity . F e e d s by filte rin g p la n k to n ( c o p e p o d s ) , b u t a lso ta k e s
sm all fishes, crabs and shrimps. B reeds in M ay-June o ff N orth
C arolina, possibly M arch-July o ff V enezuela.
20
-
o°
Size : To 25 cm standard length, usually to 20 cm.
Interest to F ish eries : A lthough locally abundant, not
greatly valued for food although appearing in m arkets (W est 20
Indies, Panama). The total catch w as 5 264 tons in 1983 (m ostly
U SA , Cuba and V enezuela).
L ocal N am es
FW N A (1964:381).
BRAZIL:
Sardinha
bandeira: USA:
See
L iterature :
Berry & B arrett (1964 - full revision, but alm ost no biological
synonym s, again w ith rather little data on food and breeding).
data): other references
under
R em arks : Rivas (1972) distinguished a C olom bian form as a distinct species, w hich he claim ed occurred
with Q. oglinum , but the distinguishing characters mostly overlap or are very close: possibly this is a subspecies:
O. o g lin u m o g lin u m : b o d y d e e p e r, 32 to 37% o f s ta n d a rd le n g th : m o re lo w e r g illr a k e r s , 30 to 3 7 : m o re
dorsal and anal fin ra y s , 19 to 22 and 22 to 25: few er scales, 32 to 35: G u lf o f M aine to B razil, but not
C olom bia.
O. oglinum captivai: body more slender, 27.9 to 31.1% o f standard length: few er low er gillrakers, 25 to 28:
few er dorsal and anal f i n r a y s , 19 or 20 and 18 to 21: m ore scales, 34 to 37: C olom bia only.
H erklotsichthys
H erklotsella
H erklotsella H erre,
(type: H arengula
1941:584 (adopted
C LU P H erk
W hitley, 1951
Fow ler, 1934, Proc. Acad.nat. Sci.Philad., 8 5 :2 4 6 (type: H erengula dispilonotus Bleeker) (n o t
1933 - a siluroid genus). Herklotsichthys
W hitley, 1951, Proc.R .zool.Soc.N .S.W ., (1 9 4 9 -5 0 ):6 7
dispilonotus Bleeker: replacem ent name since H erklotsella preoccupied). H arengula: Fow ler,
by numerous authors until H arengula Valenciennes w as finally restricted to N ew W orld species).
" fleshy
outgrowths
fronto-parietal
striae on top
o f head
2nd supra-maxilla
D iagnostic F eatures : The presence o f tw o fleshy outgrow ths on the hind m argin o f the g ili opening
distinguishes H erklotsichthys from all other clupeoid genera except H arengula and Opisthonem a (both restricted
to N ew W orld: also c h a r a c t e r i z e d by a toothed hypo-m axilla and a filam entous last dorsal finray respectively) and
Sardinella and A m blygaster o f w hich some species closely resem ble H erklotsichthys, but have more fronto­
parietal striae on top o f head (7 to 14: cf. 3 to 7), a sym metrical second supra-m axilla (cf. lower part larger than
upper) and the last two anal fin ray s enlarged .
Biology, H abitat and D istribution : M arine pelagic and schooling, but some A ustralian species apparently
entering freshw ater, or at least estuaries.
In d o -P acific reg io n only, m ainly in tropical or subtropical w aters.
Earlier biological studies are o f limited value because o f poor taxonomy.
Species : In the most recent revision, W ongratana (1 9 8 0 ) r e c o g n iz e d 8 species and subsequently described 2
o f these as new (W ongratana, 1983). Further studies on the A ustralian H erklotsichthys increases the total known
species to 12 (o f w hich 3 A ustralian species have yet to be named):
74
W idespread
H. quadrimaculatus (Rüppell, 1837), Indo-W est Pacific
W estern
Indian
Ocean
H. lossei W ongratana, 1983, the “G u lf’
H. punctatus (Rüppell, 1837), Red Sea.
H. spilurus (Guichenot, 1863), East A frica, Réunion Island.
W estern
C entral
Pacific
H. dispilonotus Bleeker, 1852, Indonesia, G ulf o f Thailand
A u stra lia
H. blackburni (W hitley, 1948), w estern A ustralia
H. castelnaui (Ogilby, 1897), eastern A ustralia
H. gotoi W ongratana, 1983, northern A ustralia, N ew Guinea
H. koningsburgeri (W eber & de Beaufort, 1912), w estern Australia.
Elerklotsichthys Species A , w estern A ustralia
Elerklotsichthys Species B, w estern A ustralia
Elerklotsichthys Species C, eastern Australia.
Herklotsichthys blackburni (W hitley, 1948)
CLUP
H erk
6
M acrura blackburni W hitley, 1948, A u st.Z o o l.. 11:266 (Port H edland, W estern A ustralia).
Synonyms :
FAO Nam es :
H aren g u la b lack b u m i:M unro. 1956:23 flg. 163 (Port H edland, W estern A ustralia).
En - B la c k b u rn ’s herring.
D iagnostic F eatures : B ody m oderately deep, its depth 30% o f standard length o f more. D istinguished
from m ost other A ustralian H erklotsichthys by lacking spots on the flank (H. koningsbergeri. H erklotsichthys
Species A) and lacking elongate w ing-like scales underneath the normal paired pre-dorsal scales (the previous two
species, also H. quadrim aculatus and H erklotsichthys Species B).M ost closely resembles H. castelnaui o f eastern
A u s tr a lia , b u t h as fe w e r lo w e r g illr a k e r (38 to 4 2 ; cf. 42 to 47, 51 or 52). S e p a r a te d from EL gotoi and
H erklotsichthys Species C (neither found in W estern A ustralia) by its black tips to cau d al fin (cf. at m o st grey) and
three distinct dark lines along upper flank.
75
W estern
G eographical D istribution
(Port Hedland, Onslow).
A ustralia
&
V
C o astal,
H ab itat and B iology
schooling. More data needed.
p elagic,
' ¿ t i
v,< y>
— / (jC
\
To 10.5 cm standard length, usually
Size
about 8 cm.
Interest
to
F isheries
No
separate
L ocal
N am es
: A U S T R A L IA
yellowfm herring (Munro, 1965).
4f
^
w*
statistics.
%
/
B la c k b u rn ’s
o -
r ?
-v T
V
■
Literature : See under synonyms.
R em arks : The w estern c o u n terp art
castelnaui and possibly merely a subspecies.
Heklotsichthys
K ow ala
of
H.
castelnaui (Ogilby, 1897)
castelnaui
Ogilby,
0
<
C L U P H e rk 7
1897, Proc.Linn.Soc.N.S.W., 22:66 (New South Wales).
Synonym s :
H arengula abbreviata:Munro, 1956:24, fig. 164 (not H arengula abbreviata Valenciennes,
1847,
w hich is considered a nomen dubium by W hitehead, 1967:69);
H erklotsichthys castelnaui - W ongratana, 1981:145,
pis 89, 90 (but excluding the specimens o f H. blackburni).
FAO N am es :
En - C astelnau’s herring.
D iagnostic Features : Body m oderately deep, its depth 30% o f standard length or more. D istinguished
from m ost other A ustralian H erklotsichthys by lacking spots on the flank (H. koningsbergeri. H erklotsichthys
Species A ) and lacking elongate w ing-like scales underneath the norm al paired pre-dorsal scales (the previous two
species, also H. quadrim aculatus and H erklotsichthys Species B). M ost closely resembles IT blackburni o f W estern
A ustralia, but has more low er gillrakers (42 to 47, some specimens 51 or 52; cf. 38 to 4 2 ). S e p a ra te d from IT gotoi
and H erklotsichthys Species C by having three distinct dark lines along upper flank (also, low er gillrakers only 34
to 58 in IT gotoi!. U pper caudal tip black, dorsal fin w ith distinct pale band separating grey base from black tip;
a dark line along upper edge o f caudal peduncle.
76
G eographical D istribution
E astern A ustralia
(Queensland southward to N ew South Wales).
H abitat and B iology :
schooling. M ore data needed.
Size
10 cm.
: To
Interest
14
to
standard
F isheries
Local Names
(Munro, 1956).
L iterature
cm
C o astal,
length,
No
: AUSTRALIA :
separate
o°>
pelagic,
usually
to 20,
statistics.
Southern
herring
40"
: See under synonyms.
R em arks : Possibly is m erely an eastern sub­
species, o f w hich IT b lack b u rn i is the w estern c o u n terp art; h ow ever, it seem s equally close to H erk lo tsich th ys
Species C.
C L U P H e rk
H erklotsichthys dispilonotus (B leeker 1852)
5
H arengula dispilonotus Bleeker, 1852, N a tu u rk .T ijd sc h r.N e d .-In d ie . 3:456 (B anka, Indonesia).
Synonym s : H arengula dispilonotus :Fow ler. 1941:585
W o n g ratan a, 1981:151, pis 98, 99 (description, illustrations).
FAO
(full
synonym y);
H erklotsichthys
dispilonotus:
N am es : En - B lacksaddle herring.
D iagnostic F eatures : Body
distinguished from all other species
back, at the hind p art o f the
H. quadrim aculatus, w hich lacks the
pre-dorsal scales.
m oderately deep, its depth 30 to 35% o f standard length. Im m ediately
o f H erklotsichthys by the presence o f two dark saddle-like blotches on the
dorsal fin base and a short distance behind this. O verlaps range o f
black saddles and has elongate w ing-like scales underneath the normal paired
77
---------i--------- ----------!....... ...T
G eographical D istribution : G u lf o f Thailand, the
Philippines, Indonesia (not eastern Indian Ocean).
i—
---\—
H abitat and B iology : C oastal, pelagic,
More data needed.
_
schooling.
À
Size : To 8.5 cm. standard length,usually to 7 cm.
2
In terest to F ish eries : N o separate
enters artisanal fisheries, but uncommon.
s ta tis tic s ;
\J
----
a
L ocal N am es :
1/-
Synonym s :
FAO N am es :
\t
C L U P H e rk
H erklotsichthys gotoi W ongratana, 1983
H erklotsichthys gotoi W ongratana,
fs:
-
8
1983, Jap.J.Ichthyol.. 29(4):391, fig. 6 (M imiko R., Papua N ew Guinea).
N one.
E n - Goto’s h e r r in g .
D iagnostic Features : Body m oderately deep, its depth 38 to 41% o f standard length. D istinguished from
most other A ustralian H erklotsichthys by lacking spots on the flank, except for a single dark spot behind gili cover
(H. koningsbergeri. H erklotsichthys Species A ) and lacking elongate w ing-like scales underneath the norm al paired
pre-dorsal scales (the p rev io u s tw o sp ecies, also IT q u ad rim acu latu s and H erk lo tsich th y s S pecies B). R e se m b le s
H. blackburni and IT castelnaui, but lacks the three distinct dark lines along upper flank (also only 34 to 38 low er
gillrakers; cf. 38 to 4 2 and 41 to 51). D istinguished from H erklotsichthys Species C by the few er gillrakers (cf. 45
to 4 8 ), presen ce o f a dark spot behind the gili cover, but no dark tip to dorsal fin.
78
G eographical D istribution : New Guinea (M imika
R iver) and northern A ustralia (G ulf o f Carpenteria).
H ab itat
and
B io lo g y
C o astal,
pelag ic,
schooling; appears to enter estuaries, if not to ascend
some way up rivers. More data needed.
Size : To at least 9 cm standard length, usually
to 8 cm.
Interest to F isheries : Possibly enters artisa­
nal fisheries, but does not appear to be common.
Local N am es : -
C lupea (H arengula)
C L U P H e rk 9
(W eber & de Beaufort, 1912)
H erklotsichthys koningsbergeri
koningsbergeri W eber & de B eaufort, 1912, V erhand.A kad. W et.A m st.. 17:14 (Ara Island).
Synonym s : H arengula
1956:23, fig. 162.
m accullochi
W hitley,
1931:143,
fig.
2
(Port
H edland,
W estern
A ustralia);
M unro,
FAO N am es : E n - K o n in g sb e rg e ri herring.
».
D iagnostic Features : Body fairly deep, its depth 37 to 40% o f
standard length.
D istin g u ish ed from all o th er A u stralia n H erklot­
sichthys by the series o f large and alm o st oval black spots on the
flank, som etim es w ith a second partial row below . H erk lo tsich th y s
Species A has m uch sm aller, round spots and is m ore slen d er, its
depth 29 to 35% o f standard length; also, hind edges o f its scales are
not denticulated and low er gillrakers 33 to 36; cf. 28 to 31 in H.
koningsbergeri. Elongate w ing-like scales present underneath n o r m a l
p aired p re-d o rsal scales.
p re-dorsal
scales
w ing-like
scale
79
Geographical D istribution
(Carnarvon to about Broome).
W estern A ustralia
H a b ita t
and
B io lo g y
C o astal,
p elagic,
schooling; enters inlets and low er reaches o f rivers
(Munro, 1956:23). More data needed.
Size
10 cm.
: To
13 cm
standard
Interest to F isheries : N o
perhaps not sufficiently abundant.
L ocal N am es :
herring (M unro, 1956).
length,
usually
separate
A U ST R A L IA :
to
statistics;
M cC u llo ch ’s
L iterature : M unro (1956 - as H. m accullochi,
description, b rie f notes).
H erklotsichthys
lossei
W ongratana,
Herklotsichthys lossei Wongratana,
C L U P H e rk 3
1983
1983,
J a p .J .I c h th y o l.. 29(4):392, fig. 7 (the “G u lf’).
Synonym s : Sardinella m elanura: B legvad & L oppenthin, 1944:66 (the
H e rk lo tsich th y s p u n c ta tu s: W h iteh ead . 1965244 (th e “G u lf” m aterial only).
FAO
N am es
En
“G u lf’); Fow ler,
1956:67
(sam e);
- G u lf herring.
D iagnostic F eatures : B ody slender, its depth 26 to 28% o f standard length. C losely resem bles H.
punctatus o f the Red Sea in shape, but the latter lacks a dark blotch on the dorsal fin and has small dark spots
along the back (either side o f dorsal fin) and not on the flank. D istinguished from H. spilurus (east A frican coast)
by its more slender body (cf. depth 28 to 35% o f standard length) and spots along flank. Possibly overlaps range o f
H. quadrim aculatus, w hich lacks spots on the flank and has elongate w ing-like scales underneath the normal paired
pre-dorsal scales. Flank w ith small dark spots, tip o f dorsal fin w ith a dark blotch.
click fo r n e x t p a g e
c lic k fo r p r e v io u s p a g e
80
G eographical
D istribution
The
“G ulf” , possibly
also
Red
Sea.
H abitat and B iology : C oastal, pelagic, schooling. M ore data
needed.
Size : To 8 cm standard length, usually to 7 cm.
Interest to Fisheries : No separate statistics, but perhaps o f
p otential interest.
Local N am es : THE “G U LF” : H ashineh.
H erklotsichthys punctatus
(R üppell, 1837)
Clupea punctata R üppell,
C L U P H erk 1
1837, N eue W irbelth., F isch e:78, pi.21, fig. 2 (Red Sea).
Synonym s : H aren g u la arabica V alan cien n es, 1847:298 (M ohila, R ed Sea, on E hrenberg M S): S pratella
erythraea K lunzinger, 1870:599 (Red Sea): H erklotsichthys punctatus - W hitehead, 1965:244 (Red Sea records
only): W ongratana, 1988:149. pis 96, 97 (revision): FN AM , 1984:221 (M editerranean). Note: the name punctata
or punctatus has been w idely m isuded in the literature for IT quadrim aculatus.
FAO N am es :
E n - Spotback h e r r in g .
D iagnostic Features : Body slender, its depth 24 to 30% o f standard length. Closely resem bling IT lo ssei o f
the “G u lf’ in shape, but the latter has a dark blotch on the dorsal fin and small dark spots along the flank and not
on the back. D istinguished from IT spilurus by its more slender body (cf. depth 28 to 35% o f standard length) and
presence o f spots on back. O verlaps range o f H. quadrim aculatus, w hich lacks spots on the back and has elongate
w ing-like scales underneath the normal paired
pre-dorsal scales. Flank silvery, w ithout spots, but a series o f very
small dark spots on back on either side o f dorsal fin base or behind it. See CTUP Herk 1, Fishing Area 51.
81
G eographical D istribution : Red Sea, possibly G u lf o f Aden;
immigrant into eastern Mediterranean (FNAM, 1984:221).
H abitat and B iology : C oastal, pelagic, schooling. More data
needed.
Size : To 8.5 cm standard length, usually to 7 cm.
Interest to
F isheries
: No
separate
statistics.
L ocal N am es : —
H erklotsich th ys q uad rim acu latu s
(R ü p p e ll, 1837)
CLU P H erk 2
C lupea quadrim aculata R ü p p e ll, 1837, N eue W irb e lth .,F isc h e :78. pi. 21, fig. 3 (Bay o f M assaw a, R ed Sea).
Synonym s :
H arengula bipunctata Valenciennes, 1847:98 (on Ehrenberg M S; M assawa); Sardinella lineolata
V alenciennes, 1847:272 (Trincom alee, B ourou Island);
C lupea fasciata V alenciennes, 1847:349 (Saint-D enis de
Bourbon); M eletta obtusirostris
V alenciennes, 1847:375 (S ey ch elles); M e le tta v en en o sa V alen cie n n e s, 1847:377
(Seychelles); A lausa schrammi B leeker, 1849:11 (Bali); H arengula m oluccensis B leeker, 1853:609 (Ternate,
A m boina, Ceram); H arengula kunzei Bleeker, 1856:209 (Ternate); Clupea (H arengula) dubia Bleeker, 1872:108 (on
R lineolata); H arengula stereolepis
O gilby, 1897:759 (Torres St., D arnley Island, southeastern N ew Guinea);
Clupea mizun K ishinouye, 1907:98, pi. 20, fig. 3 (Okinawa); Clupea rechingeri Steindachner, 1908:1424 (Samoa);
H arengula lippa W hitley, 1931:142, fig. 1 (northw estern A ustralia); H erklotsichthy punctatus Form B :Losse,
1968:88 (E ast A frica).
H erklo tsich th y s quadrim aculatus - W ongratana, 1980:139, pis 85, 86 (full synonym y,
illustrations); SFSA, in press (southern A frica). Note: This species is widely given as H arengula or H erklotsichthys
punctatus in the literature, although Fowler (194 1 :5 8 9 ) used the equally incorrect name Harengula ovalis.
FAO N am es :
En - B luestripe
herring.
tik
iff/m.
) )
/ y ; ; y V A ) ’r }
'■'X:X.XKA.X¥) y V ) > ')' x -a
D iagnostic F eatures : Body slender, its depth 18 to 30% o f
standard length. The presence o f elongate w ing-like scales underneath
the normal paired pre-dorsal scales separates it from all o th er species
except the tw o A u stralia n species w ith pro m in en t black spots on the
flank (H. koningsbergeri and H erk lo tsich th y s Species A ) and the u n ­
spotted A ustralian K erklotsichthvs Species B), w hich has dusky tips to
dorsal and caudal fins and more lower gillrakers (36 to 42; cf. 33 to 36 in
H. quadrimaculatus). Flank silvery with an electric blue line preceded by
tw o o range spots. See CLUP Flerk 2, Fishing A rea 51.
p re-dorsal scales
wing-like
scale
82
G eographical D istribution : W idespread in Indian
O cean and w e s te rn P a c ific (e n tire e a s te rn c o a s t o f
A frica, M ad agascar, M au ritiu s eastw ard to Japan, e a st­
ern A u stralia, Sam oa). In tro d u ced into H aw aii, ap p a­
re n tly by a c c id e n t, and n o w a b u n d a n t (W illia m s &
C larke, 1983).
H a b ita t
and
B io lo g y
C o astal,
p elag ic,
schooling; daytim e in a few m etres o f clear w ater,
n ig h t-tim e into deep er w ater and fu rth er o ffsh o re (at
le a s t th e a d u lts).
F e e d s on Z o o p lan k to n , m a in ly at
night, chiefly co p ep o d s in juvenile stages, but larger
prey as ad u lts (ch aeto g n ath s, p o ly ch aetes, shrim ps and
sm all fishes); at no tim e is p h y to -p lan k to n tak en (data
from W illia m s & C la rk e , 19 8 3 ;
M a ric h a m y , 19 7 0 ).
Breeds during its first year and probably survives only a
few m onths after m atu rity (W illiam s & C lark e, 1983:
595).
M ost p reviou s data p u b lish ed u n d er the nam e
H. punctatus.
Size
: To 14 cm stan d ard le n g th , u su ally ab o u t 10 cm ..
Interest to F isheries : U sually not separately
Locally o f some im portance in artisanal fisheries.
reported,
but probably
com bined
w ith
Sardinella
catches.
Local N am es : USA: G old spot herring (H awaii).
L iterature : W illiam s & Clarke (1983 - H aw aii, biology, but spawning not observed). A lso Losse (1968 - as
H punctatus Form B, E ast A frica); M aricham y (1970, 1974 - as IL punctatus, in Indian w aters, food, condition
factor).
Harklotsichthys spilurus
(Guichenot, 1863)
C LU P H erk 4
H arengula spilura G uichenot, 1863, In M aillard, Ile de Réunion p o iss.:16 (Bourbon).
Synonym s : H erk lo tsich th y s p u n ctatu s F orm A :Losse, 1968:88 (K enya, Z an zib ar); W h iteh ead , 1973b: 176 (on
L osse); H erklotsichthys spilurus:W ongratana. 1981:149, p is 94, 95; W hitehead & B auchot, in press (type o f
sp ilu ra).
FAO
N am es
: E n - R eunion herring.
D iagnostic Features : Body m oderately deep, its depth usually 28 to 35% o f standard length. R esem bles
H. lossei, w hich is more slender (depth 26 to 28% o f standard length) and has sm all dark spots on the flank.
O verlaps range o f IL q u ad rim a cu latu s, w h ich has elo n g ate w in g -lik e scales u n d ern eath the norm al p aired p re ­
dorsal scales. Flank silv ery , w ith o u t dark spots; a bright yellow or orange patch behind gili opening; dark blotch
on tip o f dorsal fin.
83
G eographical D istribution : Zanzibar,
Island, presum ably also M adagascar.
K enya
coast,
Réunion
H abitat and B iology : C oastal, pelagic, schooling. More data
needed.
Size : To 8.5 cm standard length, usually to 7 cm.
Interest to F isheries
enters artisanal fisheries.
Local N am es :
sim ilar clupeids).
Herklotsichthys
: No
separate
ZA N ZIB A R :
statistics,
D agaa
(but
but
used
probably
for
other
Species A
C LU P H erk 10
(A W AITING D ESC R IPTIO N A N D N AM E)
Synonym s
: H arengula
ko n in g sb erg eri:M unro,
1956:23, fig. 161 (n o t IE k o n in g sb erg e ri o f W eb er &
deBeaufort, w hich M unro in error called IL
m a c c u llo c h i W hitley); ? H aren g u la o v a lis:M unro. 1956:23, fig. 160
(sensu Fow ler, 1941:589, i.e. H. quadrim aculatus).
FAO Names :
En - Australian spotted herring.
D iagnostic Features : B ody m o derately slender, its depth 29
to 35% o f standard length. D istinguished from all other A ustralian
H erklotsichth s by the series o f small round black spots on the flank
( H. koningsbergeri has much larger oval spots and is deeper-bodied,
its depth 37 to 40% o f standard length; also, hind edges o f its scales
denticulated arid lower gillrakers 28 to 31; cf. 33 to 36 in Species A).
Elongate w ing-like scales underneath normal paired pre-dorsal scales.
A dark spot behind gili cover.
p re -d o rsal scales
w in g -lik e
scales
84
G eographical D istribution : W estern A ustralia
(Exm outh G u lf to G ulf o f C arpenteria, Cape Y ork and
A ra Islands).
H ab itat
and
B io lo g y
schooling. More data needed.
Size
14 cm.
: To
16 cm
:
standard
C o astal,
length,
pelag ic,
usually
to
Interest to F isheries : N o separate statistics;
possibly more abundant than IT koningsbergeri.
L ocal N am es :
Literature : M unro (1956 - as IT koningsbergeri,
description, brief notes).
H erklotsichthys
100°
120°
140°
160°
C TU P H erk 11
Species B
(A W A IT IN G D E S C R IP T IO N A N D N A M E )
H.
Synonym s : ?
quadrim aculatus!.
H arengula lippa o f authors, e.g. M unro,
195623, fig.
159 (not H. lippa W hitley, w hich is
FAO N am es : En -
D iagnostic F eatures : B ody slender, its depth 27 to 35% o f
standard length. D istinguished from m ost other A ustralian H erklot­
sichthys by the presence o f elo n g ate w in g -lik e scales ben eath the
normal paired pre-dorsal scales and absence o f spots along the flank.
Closely resem bles IT quadrim aculatus in these features, but has more
low er gillrakers (36 to 42; cf. 32 to 35 in IT quadrim aculatus). Three
fine dark lines along upper flank and dorsal fin w ith distinct pale band
separating grey base from black tip.
pre-dorsal scales
w ing-like
scale
85
G eographical D istribution
(Shark Bay to Onslow).
Habitat and B iology :
schooling. More data needed.
Size
10 cm .
: To
Interest
to
13 cm
C oastal,
standard
F isheries
: W estern A ustralia
: No
length,
pelagic,
usually
separate
L ocal N am es : A U STR A LIA : O ily
(Munro, 1956 - if it was indeed this species).
to
statistics.
herring
H erk lo tsich th y s S pecies C
C L U P H erk 12
(A W AITING D ESC R IPTIO N A N D NAM E)
Synonym s
: None.
FAO N am es : En
D iagnostic Features : Body m oderately slender, its depth about 30% o f standard length. D istinguished
from m ost o ther A u stra lia n H e rk lo tsic h th y s by lack in g spots on the flan k (H. k o n in g sb e rg e ri. H erk lo tsich th y s
Species A ) and lacking elongate wing-like scales beneath the normal paired pre-dorsal scales (the previous two
species, also IE quad rim acu latu s and H erk lo tsich th y s S pecies B).
R esem bles IL b lackburni and especially
H. castelnaui (same range), but lacks the three distinct dark lines along upper flank. D istinguished from IT gotoi
by having more low er gillrakers (47 to 48: cf. 34 to 38) and no dark spot behind gili cover: also dorsal fin with,
distinct pale band separating grey base from black tip and a dark line along upper edge o f caudal peduncle.
G eographical D istribution : Eastern A ustralia
(Warrell Creek at Scott’s Head, N ew South Wales).
H abitat and B iology : More data needed.
Size
: To 9 cm
standard length.
Interest to Fisheries : Probably nil.
Local N am es : L iterature : R em arks : V ery sim ilar to IT castelnaui in
num ber o f low er gillrakers and c o lo u r p a tte rn , except
lacking the lines along upper flank. M ore specimens
may show it to be that species.
100°
'2 0 °
140°
io o °
180°
click fo r n e x t p a g e
c lic k fo r p r e v io u s p a g e
86
A m b ly g aster
CLUP
Bleeker, 1849
A m bl
A m blygaster B leeker, 1849, J.Ind.A rch., 3:73 (type. A m blygaster clupeoides Bleeker). Fusiclupea W hitley,
1940, A ust.Z ool., 9(4):401 (type:Sardinops (F u sic lu p e a ) dakini W hitley = W s im i). Sardinella: w idely used until
recently for the 3 species o f A m blygaster, also many authors acknow ledged A m blygaster as a subgenus.
D iagnostic Features : The presence o f two fleshy outgrowths on the hind m argin o f the gili opening distin­
guishes A m blygaster from all other clupeid genera except H arengula and O pisthonem a (both restricted to N ew
W orld: also characterized by a toothed hypo-m axilla and a filam entous last dorsal finray respectively) and
H erklotsichthys and S a r d in e lla .
A m blygaster has 7 to 14 f r o n to ­
p a rie ta l stria e on to p o f th e h e a d , a sy m m e tric a l seco n d su p ram axilla and the last tw o anal f i n r a y s enlarged (3 to 7 striae, an
asym m etrical second sup ra-m ax illa and final anal finrays norm al in
H e rk lo tsic h th y s).
A m b ly g aster m ost closely resem bles S ardinella
and w as form erly com bined w ith it, b u t has only 26 to 43 low er
fleshy
g illrak ers and a m edian series o f p re-d o rsal scales (cf. 40 to 100
outgrowths
low er gillrakers, or up to 200 or more in some species, and a median
series o f pre-dorsal scales (usually paired p re-dorsal scales in Sar­
d inella). In general body shape species o f A m blygaster resem ble
Sardinops (but bony striae on operculum).
X
r
/
C
O x
,XX X X ) i
striae
a. S ardinella
2nd supra-maxilla
sy m m e tric a l
b A m blygaster
pre-dorsal scales
fro nto-parietal striae
on top o f head
Biology, H abitat and D istribution
tropical or subtropical waters.
Interest to F isheries : C ontributes
statistics from general clup eo id catches.
M arine, pelagic and
to
num erous
local
schooling. Indo-Pacific region only, mainly in
artisanal
fisheries,
but usually
not
separated
in
Species : The m ost recent revision, by W ongratana (1980), re c o g n iz e d 3 species:
A. clupeoides Bleeker, 1849, Indo-W est Pacific
A. leiogaster (V alenciennes, 1847), Indo-W est Pacific
A. sirm (W albaum, 1792), Indo-W est Pacific.
Am blygaster clupeoides
Bleeker, 1849
C LU P A m bl 1
A m b ly g aster clupeoides B leek er, 1849, J.In d .A rch ., 3:73 (M acassar).
Synonym s : Sardinella
clupeoides:Fow ler. 1941:619; B ennet, 1965:1 (India); C han, 1965:115, fig. 14
(d e s c rip tio n , sy n o n y m s); w h ite h e a d et aL , 1966:67 (ty p e ); W h ite h e a d , 1973b:188 (Indian Ocean); N air, 1973:62
(India); A m blygaster clupeoides - W ongratana, 1980:137, pis 81, 82 (revision).
FAO N am es : En - B leeker’s sm oothbelly sardinella.
87
D iagnostic Features : Body m oderately slender, belly rather rounded, scutes not prom inent. D istinguished
from A. sirm by the absence o f spots along the flank (gold in life, black on preservation) and few er low er
g illra k e rs (2 6 to 31; cf. 33 to 43 in W sirm ). Closely resembles W leiogaster, w hich has more lower gillrakers (31
to 33). Round-bodied Sardinella species (£L longiceps, etc.) have i 8 pelvic finrays (i 7 in A . clupeoides) and more
than 100 lower gillrakers. See CLUP Ambl 1, Fishing Area 51.
G eographical
D istrib u tio n
:
E astern
Indian
Ocean (southern coasts o f India, perhaps Bay o f Bengal)
and Indonesia eastward to Fiji.
H a b ita t
and
B io lo g y
:
C o a sta l,
p elag ic,
schooling. Feeds on copepods, M ysis and other ele­
ments in the Zooplankton (B ennet, 1965).
Size
15 cm.
To
17
cm
standard
length,
usually
to
Interest to F ish eries : N o separate statistics,
but evidently enters som e local artisan al fish e rie s, at
least in small numbers.
Local N am es : —
A m blygaster leiogaster
Sardinella
leiogaster
(V alenciennes, 1847)
V alenciennes,
CLU P A m bl 2
1847, H ist.nat.poiss., 20:270 (Indian O cean, Trincom alee).
Synonym s:
C lupea okinaw ensis K ishinouye,
1907:96 pi. 19, fig.2, pi. 21, fig.5 (O kinaw a); Sardinella
l e i o g a s t e r : C h a n , 1965:117, fig. 15 (d escrip tio n ); W h iteh ead , 1967a:68 (ty p e);Id em , 1973b:188, fig . 18 (Indian
O cean); Losse, 1968:101 (E a st A fric a );
M enon & T a lw a r, 1974:41 (In d ia ); W o n g ratan a, 1980:138, pis 83, 84
(revision).
FAO N am es
: En - Sm oothbelly sardinella.
D iagnostic Features : Body m oderately slender, belly rather rounded, s c u te s not prom inent. D istinguished
from Æ sirm by the absence o f spots along the flan k (gold in life, black on p reserv atio n ) and few er low er
gillrakers (31 to 35; cf. 33 to 43 in A sirm ). Closely resem bles A clupeoides, w hich has few er low er gillrakers
(26 to 31). R ound-b o d ied S ard in ella species (Sk lo n g ice p s, etc.) have i 8 pelvic finrays (i 7 in A. leio g aster) and
more than 100 lower gillrakers. See CLUP A nibi 2, Fishing Area 51 (also CLUP Sardl 10, F ish in g A re a s 57, 71).
G eographical D istribution : In d o -W e st Pacific
(coasts o f A frica eastw ard to O kinaw a, also W estern
A ustralia).
H abitat and Biology :
schooling. More data needed.
Size
18 cm.
:
To
23
cm
C o astal,
standard
length,
pelag ic,
usually
to
Interest to Fisheries : N o
separate
statistics,
but evidently enters som e local artisan al fish eries, at
least in small numbers.
Local Names :
(Okinawa).
L ite r a tu r e
:
JA PAN :
Y am ato-m izun
L o sse (1 9 6 8 - E a st A fric a ).
40 »
A m blygaster sirm
(Walbaum, 1792)
60 »
80 »
100 »
120 »
140 »
160 »
180 »
C LU P A m bl 3
Clupea sirm W albaum , 1792, in A rtedi, G en.pise.:38 (on Forssksâl, 1775, D escrip t.an im . :17 - Arabia).
Synonym s : Sardinella leiogastroides B le e k e r, 1854255 (M anado, Sulaw esi); C lupea pinguis G ü n th er,
1872:425 (M isol);
Sardinops dakini W hitley, 1937:114; Sardinella sirm :Fow ler. 1941:616; C han, 1965:113, fig. 13
(description, synonym s W hitehead et ah, 1966:69 (type o f leiogastroides); Losse, 1968:99 (East A frica; Raja &
Hiyam a, 1969:105 (O k in aw a); W h iteh ead , 1973b: 187, fig. 17 (Indian O cean); N air, 1973:58 (India); A m b ly g aster
sirm - W ongratana, 1980:135, pis 79, 80 (revision).
FAO N am es :
E n - S potted
sardinella.
89
D iagnostic Features : Body slender, belly rather rounded,
sc u te s not pro m in en t. D istin g u ish e d fro m A ,
clupeoides and A. leiogaster by the presence o f a series o f 10 to 20 gold (in life) or black (on preservation) spots
dow n the flank (but som etim es m issin g )'
also, low er g illrak ers 33 to 43 (26 to 33 in the o th er tw o species).
Sardinops species are also round-bodied and have spots on the flanks, but possess bony radiating striae on the gili
cover. R ound-bodied S ard in ella species (R longiceps, etc.) have i 8 pelvic fin ray s (i 7 in W sirm) and more than
100 lower gillrakers, also no spots. See CLUP Ambl 3, Fishing Area 51 (also CLUP Sardl 9, Fishing Areas 57, 71).
Geographical Distribution :
Indo-W est P a c if ic
(coasts o f A frica, in clu din g R ed Sea and M adagascar,
eastward to the Philippines, Taiwan Island,
Okinawa, 40°
N ew G uinea, Fiji and northern coasts o f A ustralia).
H abitat
and
B iology
C o a stal,
p e la g ic , 20 schooling. Feeds mainly on copepods, nauplii and zoea
larvae, larval bivalves and g astro p o d s. P erid in iu m and
C eratiu m (C h ack o , 1956);
ju v e n i le s a ls o o n p h y to - QO
plankton
(B a sh e e ru d d in & N ay ar, 1962). B reeds
N o v e m b e r-D e c e m b e r, F e b ru a ry -M a rc h , M a y -Ju n e and
A ugust-S eptem ber in Indian w aters (N a ir, 1973:59). 20„
M ature at 14 to 15 cm standard length (G n an am ek alai,
1962).
Size ! To 23 cm standard length, usually to about
20 cm.
40 °
60 °
80 °
100 °
120°
Interest to F isheries : N o separate statistics,
but enters som e local artisan al fish erie s, e.g. in T u tico rin B ay in N o v em b er-M arch (C h ack o ,
(Lewis, Smith & Ellway, 1983).
L ocal N am es : IN DIA
L iterature
140 °
160 °
180 °
1956), and in Fiji
(Tam il): K eerim een chalai.
: N air (1973 - good summary for Indian Ocean, biology, fisheries);
Losse (1968 -
East Africa).
Remarks : The num ber o f spots on the flanks shows som e variation (10 to 20), but this has not yet been
correlated w ith
lo cality , sex or o th er factors. Raja & Fliyama (1969:106) noted “more than 30 dark spots” w hich
w ere above the
in terfa ce b etw een silv er flan k and dark back (norm ally they lie along th is in terfa ce); they also
noted that the spots w ere faint or absent in some specimens.
click fo r n e x t p a g e
c lic k fo r p r e v io u s p a g e
90
Sardinella Valenciennes, 1847
CLU P Sardai
S ardinella V alen cien n es, 1847, H ist.n a t.p o iss., 20:18 (type:
S ard in ella au rita V alen cien n es). C lupeonia
V alen cien n es, 1847, H ist.n a t.p o iss., 20:345 (type: Clupeonia jussieui Valenciennes).
Kowala V alen cien n es, 1847,
H ist.nat.poiss., 20:362 (type: K ow ala albella V alenciennes). C lu p alo sa B leek er, 1849, V e rh .b a ta v .G en o o t.K unst.
W et.. 22:12 type: C lupalosa bulan Bleeker = K ow ala alb ella V a le n c ie n n e s ). Sardinia Poey, 1860, M em .Hist.nat.
C u b a . 2:311 (ty p e: S a rd in ia
pseudohispanica
Poey = Sardinella aurita Valenciennes).
P aralo sa B le ek e r, 1868,
V erst.M ed ed .K .A k ad .w e t.A m st.. 2 (2 ) :300 type: H arengula (Paralosa) valenciennesi B leek er = C lupea m elanura
C uvier).
W ilkesina F ow ler & B ean , 1923, P ro c .U .S .n a tn .M u s.. 63:3 (type: H arengula fijiense Fow ler & Bean).
F im briclupea W hitley, 1940, A u st.Z o o l.. 9(4):399 (type: Fim briclupea dactylolepis W hitehead
= C lupea gibbosa
Bleeker).
Fiscina W hitley, 1940, A ust.Z ool.. 9(4):400 (type: Fiscina posterus W hitley = Sardinella lem uru Bleeker).
D iagnostic Features : The presence o f tw o fleshy outgrowths on the hind margin o f the gili opening distin­
guishes Sardinella from all other clupeoid genera except H arengula (New W orld: a toothed hy p o -m ax illa p resen t).
Opisthonem a (new W orld: last dorsal finray filam entous).
H erklotsichthys (Indo-W est
Pacific: only 3 to 7 frontparietal striae on top o f head, second sup ra-m ax illa asy m m etrical,
final tw o anal finray s n o t e n la rg e d ) and A m b ly g a s te r.
S ardinella
closely resem bles A m b ly g aster (the tw o w ere form erly com bined) in
having 7 to 14 fro n to -p a rie ta l stria e , a sy m m e tric a l p a d d le -sh a p e d
seco n d s u p ra -m a x illa and th e la s t tw o an al fin ray s d is tin c tly e n ­
larged, but has m ore, often many m ore, low er gillrakers (40 to 1Ö0, or
up to 200 or more in ro u n d -b o d ied sp ecies: cf. 26 to 43 in Am bly­
fleshy
g a s te r )also, all round-bodied species o f Sardinella have paired pre­
outgrowths
dorsal scales (a single m edian series in A m b ly g a ste r). S up erficially ,
the round-bodied Sardinella resem ble Clupea and Strangom era, whose
ranges they som etimes overlap in the N orth A tlantic and perhaps also
in th e N o rth and S o u th P a c if ic ,, b u t th e c h a ra c te rs lis te d ab o v e
d istin g u ish them (gili o p ening, fro n to -p arietal striae, enlarged anal
finrays).
striae
fronto-parietal striae
on top o f head
2nd supra-maxilla
sy m m e tric a l
Sardinella
A m blygaster
pre-dorsal scales
Biology, H abitat and D istribution
M arin e, pelag ic and schooling:
som e sp ecies en ter lagoons and
estu aries, usually as ju v en ile s. Species o f the subgenus Sardinella occur on both sides o f the A tlantic, in the
M e d ite rra n e a n and th ro u g h o u t th e tro p ic a l and su b tro p ic a l p a rts o f th e Indo-W est P a c if ic : m e m b e rs o f the
subgenus Clupeonia are found mainly in the Indo-W est Pacific, but 2 species are in the eastern A tlantic (o f which
one extends into the M editerranean).
Interest to F isheries : A num ber o f species make a very significant contribution to artisanal and fully
m echanized fisheries, w hile numerous others are collectively im portant in the Indo-W est Pacific region (o f much
more im portance than are species o f H erklotsichthys, as can be seen in the fish m arkets o f India and southeast
Asia).
The total recorded catch for Sardinella in 1983 was 1499 437 tons (about 60% unidentified,: the major
species were IT longiceps and IT aurita.
Species : In the last revision o f Indo-W est P acific sp ecies, W o n g ratan a (1 9 8 0 ) re c o g n ized 18 sp e c ie s and
subsequently described one o f these as new and gave a replacem ent name to another (W o n g ratan a, 1983). To these
can be added 4 fu rth e r species from the A tla n tic and M ed iterran ean . Id en tificatio n o f som e sp ecies, especially
the sm aller m em bers o f the sub g en u s C lu p e o n ia , is o fte n e x tre m e ly d if fic u lt and in th e ca se o f ju v e n ile s
som etim es im possible. As an aid to identification, the geographical distribution o f the species should serve as a
first step once the correct subgenus has been determ ined:
91
Subgenus Sardinella (pelvic fin ra y s i 8;mostly rather round-bodied species)
S.
S.
S.
S.
S.
aurita V alenciennes, 1847, A tlantic (both sides and M editerranean), also Japan
brasiliensis (Steindachner, 1879), w estern A tlantic (? also eastern A tlantic and M editerranean)
lemuru Bleeker, 1853, Indonesia, W estern and northern A ustralia, N orth China Sea
longiceps Valenciennes, 1847, Indian Ocean (northern part)
neglecta W ongratana, 1983, Indian Ocean (East A frican coast)
Subgenus C lupeonia (pelvic fin ra y s i 7;m ostly more com pressed species)
W id esp rea d
albella (Valenciennes, 1847)
brachysoma (Bleeker, 1852)
S. fimbriata Valenciennes, 1847)
R gibbosa
(Bleeker, 1849)
S. m elanura (Cuvier, 1829)
W estern Indian Ocean
if jussieui (Valenciennes, 1847) (southern India, M adagascar, M auritius)
S. sindensis (Day, 1878) (northern part only)
W estern Pacific, northern part
R, hualiensis (Chu & Tsai, 1958) (Taiwan Island)
S. richardsoni W ongratana, 1983 (South China Sea)
R taw ilis (Herre, 1927) (South China Sea)
R zunasi (Bleeker, 1854) (China, Japan)
W estern Pacific, southern part
S. atricauda (Günther, 1868) (Indonesia)
S. fijiense (Fowler & Bean, 1923) (New Guinea, Fiji)
R m arquesensis Berry & W hitehead, 1968 (M arquesas Islands)
Eastern
A tlantic
S^ maderensis (Lowe, 1839), eastern A tlantic, M editerranean
R rouxi (Poll, 1953), eastern A tlantic (western Africa).
Sardinella albella
(Valenciennes, 1847)
C L U P Sardl 6
K ow ala albella V alenciennes, 1847, H ist.nat.poiss., 20:362 (Pondicherry).
Synonym s :
Clupalosa bulan Bleeker, 1849:12 (Indonesia); Kowala lauta Cantor, 1850:297 (Penang); Clupea
perforata C antor, 1850:1276 (Penang to Sum atra and Singapore); Spratella kow ala B leeker, 1851:492 (R io,
B atavia); C lupea (H arengula) sundaica B leeker, 1872:105 (B atavia, etc.); H arengula dollfusi Chabanaud, 1933:1,
figs 1-2 (G ulf o f Suez); Sardinella perforata:Fowler. 1941:605 (the Philippines); Chan, 1965:16, fig 7B (Thailand to
Taiwan Island); Sardinella b ulan:W hite head, 1963:33-41 (types o f bulan, perforata,
kow ala); W hitehead et ah,
1966:53, 55, 61 (types o f bulan, kow ala, sundaica); Sardinella alb e lla :L osse, 1 9 68:96 (E a st A frica); W hitehead,
1973b:183, fig. 13; W o n g ratan a, 1980:121, pis 58, 59 (revision); SFSA, in press (southern A frica); W hitehead &
Bauchot, in press (types o f albella, dollfusi).
FAO Nam es :
E n - W hite
sardinella.
92
discontinuous
y striae
D iagnostic F eatures : Body som ew hat com pressed but variable,
from slender to moderately deep, its depth 25 to 40% o f standard length;
total number o f scutes 29 to 33 (usually 30 to 32). Lower gillrakers 41 to
68 (at 4 to 15 cm standard length, increasing a little w ith size o f fish).
V ertical striae on scales not m eeting at centre, hind p art o f scale w ith a
few p e rfo ra tio n s and so m e w h a t p ro d u c e d p o s te rio rly . A dark spot at
dorsal fin o rigin. Closely resem bles Sk fim briata and Sk dayi w hich have
more low er gillrakers (54 to 82, usually more than 55 in Sk fim briata; 88 to
126 in S. dayi) and can be confused w ith Sk gibbosa and R. sindensis which
have more scutes (31 to 34, usually 32 or 33). O ther sim ilar species have
m ore scutes or m ore g illrak ers or o v erlap p in g scale striae or no spot at
dorsal fin origin or caudal tips black (or a com bination o f some o f these
features). See CLUP Sardl 6, Fishing A rea 51, also A reas 57, 71.
perforations
scale
G eographical D istribution
Indo-W est Pacific
(Red Sea, the “G u lf’, E ast A frican coasts, M adagascar
eastward to Indonesia, north to Taiwan Island, south to 40°
Papua N ew Guinea).
H ab itat and B io lo g y
C o astal, p elag ic,
schooling. M isid en tificatio n s m ake p ublished biological
data potentially unreliable. The biology is sum m arized
by N air (1960, 1973).
Size :
To 14 cm standard
20
«
0„
length, usually to
Interest to Fisheries :
In c lu d e d in th e g en eral
statistics for S a rd in e lla , but o f som e im portance along
th e w estern coasts o f In d ia (alth o u g h fish ery sta tistic s
probably include Sk fim briata and Sk gibbosa).
40°
Local Names
gibbosa.
60°
80°
100°
120°
140°
160°
180°
M o s tly th e sam e as fo r £L
L iterature : N air (1960, 1973 - India, synopsis o f biology and fisheries); Losse (1968 - E ast A frica).
Sardinella atricauda
(G ünther, 1868)
C LU P Sardl 16
C lu p e a a tric a u d a G ü n th e r, 1868 (14 M a rc h ), C a t.F is h e s B r it.M u s ., 7 :4 2 6 (C e ra m ; B leeker’s specimen o f
H arengula m elanurus).
Synonym s :
H arengula m elanurus B leek er,1853:245 (not C lupea
(P a ra lo sa ) v alen c ien n e si B le e k e r, 1868 (p resu m ed a fte r 14 M arch): 300
H are n g u la m e la n u ru s); W ongratana, 1980:134, pis 77, 78 (revision). Note:
(1965:5); W hitehead et ah (1966:63); W hitehead (1973b:180) and others in the
FAO Names
En - B leeker’s blacktip sard in ella.
m elanurus C uvier, 1829); H arengula
(W aig o u ; replacem ent nam e for his
included by F o w ler (1 9 4 1 :615), Chan
synonymy o f Sk m elanura.
93
D iagnostic Features : Body slender, its depth 23 to 27% o f standard
length;
to ta l sc u te s 32 to 35 (p re -p e lv ic sc u te s u s u a lly 19).
Lower
gillrakers 39 to 43, w ith many asperities on them (probably not increasing
in num ber in larger fishes). V ertical striae on scales not meeting at centre,
num erous small perforations on hind part o f scale.
No dark spot at dorsal
fin origin; tips o f caudal fin black. M ost closely resembles Sk m elanura,
w hich also has black caudal fin tip s, but few er scutes (27 to 31). O ther
S ardinella species lack the black caudal tip s or have m ore g illrak ers or
few er scutes or occur outside the area.
discontinuous
striae
small
perforations
G eographical D istribution : W estern Pacific (Java
Sea from Jak arta to A m b o in a, b u t p erh ap s m ore w id e ­
spread in Indonesia).
Habitat and Biology
More data needed.
Coastal, pelagic, schooling.
about 10 cm.
Interest to F isheries : Presum ably
local S ardinella catches.
contributes to
L ocal N am es : L iterature : See under synonyms.
R em arks
Occurs w ith
S. m elanura in the Java
Sea and must have been confused with it frequently.
Sardinella aurita
V a len cie n n e s, 1847
Sardinella aurita V alenciennes, 1847, H ist.n a t.p o iss..
Synonym s :
(a)
(b)
C LU P
Sardl
2 0 2 6 3 (M essina to G ulf o f M orée, G reece).
These are separated for the two m ajor areas o f distribution:
Eastern A tlantic : C lupea allecia R afinesque, 1810:57 (Sicily; potential nom en oblitum , but not yet
form ally suppressed as such); A losa senegalensis B ennett, 1831 (Senegal; overlooked and should be
d e sig n a te d a nomen o b li tu m );
C lupea aurovittata Sw ainson, 1838 (nom en nudum ) and 1839:385
(P alerm o; o v erlo o k ed , an o th er p o ten tial nomen o b litu m ); M eletta m editerranea V alenciennes, 1847:
369 (Toulon, M arseille); Sardinella euxina A n tip a , 1906:46, pi. 3, figs 12to 18 (Black
Sea); Clupea
v e n u lo s a S te in itz , 1927:3230 (Haifa); S ard in ella a u rita terrasae L ozano y R ey, 1950:14, pi. 3, fig. 2
(W estern Sahara, C anaries); Sardinella aurita var. m editerranea:R ossignol, 1959:215 (M editerranean).
S a rd in e lla a u rita - S v e to v id o v , 1952:172, pi. 5, fig. 2 (R ussian w aters); Id em , 1963:189, pi. 5, fig. 2
(s a m e ); C L O F N A M , 1 9 7 3 :1 0 3 (fu ll sy n o n y m y fo r M e d ite rra n e a n an d e a s te rn A tla n tic ) ; F N A M ,
1 9 8 4 2 2 2 f ig . ; C LO FE TA , in press (eastern central A tla n tic, full synonym y);
SFSA , in press
(southern A frica); W hitehead & Bauchot, in press (types o f aurita, m editerranea).
W estern A tlantic
fig. 99 (synopsis
p s e u d o h is p a n ic a
1964:400, fig. 98
: Sardinella anchovia V alenciennes, 1847:269 (Rio de Janeiro); FW N A, 1963:401,
o f taxonom y and b iology);
C erv ig ó n , 1966:120, fig . 50 (V e n e z u e la );
Sardinia
P o e y , 1860:311 (C u b a ).
S ard in ella p in n u la B ean, 1912:122 (B e rm u d a );
FWNA,
(synonym y, biol.); S ard in ella a u rita :W hitehead, 1973a:27, fig. 6 (Guianas).
FAO N am es : En - Round sardinella.
1
94
D iagnostic Features : Body elongate, usually subcylindrical, but som etim es a little com pressed; belly
rath er rou n d ed , b u t scutes a p p a re n t.
Low er gillrakers fine and num erous, more than 80 (162 to 248 in W est
A frican specim ens o f 23 to 28 cm standard length); anterior gillrakers on low er lim bs o f second and third gili
arches lying more or less flat (strongly curled in Sk brasiliensis). The pelvic fin ray count o f i 8 distinguishes Sk
aurita from all other species o f Sardinella, also H a re n g u la . O pisthonem a, H erklotsichthys and A m blygaster that
occur w ith it. R esem bles C lupea, but has two fleshy outgrowths along outer margin o f gili opening (smooth in
C lupea) and num erous fine fro n to -p arie ta l striae on top o f head. Flanks silvery, w ith a faint golden midlateral
line, preceded by a faint golden spot behind gili opening; a distinct black spot at hind border o f gili cover (absence
o f silver pigment). See CLUP Sardl 1, Fishing Areas 37, 34 and 47 (in part), 31.
fla t
curled
fleshy
outgrowth
S. aurita
S. b rasiliensis
fro n to -p a rie ta l stria e
on top o f head
lower limbs o f gili arch es
G eographical D istribution
E astern
A tla n tic
(M e d ite rra n e a n ,
b u t ra r e in B la c k S e a ;
A frican
coasts from G ib raltar southw ard to
Saldanha Bay, South A frica (see also
under rem arks);
w e ste rn A tla n tic
(Cape Cod to A rgentina).
H abitat and B iology : Coastai, pelagic, preferring clear saline
w ater w ith a m inim um tem perature
below 24°C ; from inshore and near
surface to edge o f shelf and down to
350 m (W e st A fric a ), or p e rh a p s
even deeper; sch o o lin g and strongly
m igrato ry , often risin g to surface at
night and dispersing. F eeds m ainly
on Zooplankton, especially copepods,
but som e p h y to p lan k to n (esp ecially
by ju v en iles). B reeds perhaps at all
tim e s o f th e y e a r (e .g . o f f W e st
80<
eo-
40<
20„
20
°
40°
60°
160°
140°
120°
100°
80°
60°
40°
20°
0°
20°
40°
60°
80°
100°
120°
140°
160°
1
95
A f r ic a ) , b u t w ith d is ti n c t p e a k s , e .g . m id -J u n e -e n d o f S e p te m b e r, i.e . th e su m m e r m o n th s , in th e
M editerranean and perhaps o ff N orth A m erica; from about M ay o ff Senegal (but again in O ctober-N ovem ber)
through to July-A ugust o ff M auritania; around July or A ugust o ff Ivory C oast and G hana; but apparently
September to February, i.e. the w inter months, in the G ulf o f M exico (Houde & Fore, 1973), January-February o ff
V enezuela (Sim pson, 1969), and, if it spawns w ith
brasiliensis, then Septem ber to M arch o ff Brazil.
The
breeding pattern is
extremely complex, with two principal spawning periods in some areas (linked with upwelling
regim es o ff W est
A frica). The ju v e n ile s tend to stay in n ursery areas (m ain n u rseries o ff M au ritan ia and
Senegal/G am bia for the W est A frican region), but on m aturity rejoin adult stocks in the colder offshore w aters.
Size : To 30 cm standard length, usually to 25 cm.
Interest to F isheries : M ajor fisheries o ff W est A frica,
although catches not always distinguished from those o f other
western Atlantic).
The total catch for 1983 was 702 775 tons,
with S. maderensis). 401 039 tons for W est A frica (mixed with
A tlantic (m ixed w ith
brasiliensis).
L ocal N am es : USA (Florida):
in M editerranean and o ff V enezuela and Brazil,
Sardinella species (especially o f
brasiliensis in
being 15 209 tons for the M editerranean (mixed
S. maderensis) and 286 527 tons for the western
Spanish sardine.
L iterature : For eastern A tlantic area, Svetovidov (1952, 1963 - M editerranean, B lack Sea); B en-Tuvia
(1960 - general synopsis); Postel (1960 - synopsis, W est Africa); Banarescu (1964 - Black Sea);
Fagetti & Marak
(1972 - synopsis o f spawning o ff W est Africa); Anon. (1979 - same, also growth, fecundity); Boely & Freon (1980 sam e); FN A M (1984 - synopsis); CLOFETA (in press - virtually com plete bibliography for W est A frica). For
w estern A tlantic area, H ildebrand (i.e. FW N A, 1964).
R em arks : The separation o f S^ aurita from S^ brasiliensis by m eans o f gillrakers shape (see diagnostic
features) is tentative; it appears to reinforce the difference in gillrakers numbers (more in
brasiliensis, but in
both cases they increase as the fish grows) w hich w orkers have hitherto used.
The ‘brasiliensis’ curled form o f
gillrakers has been found also in both M editerranean and W est A frican fishes. Possibly this is a racial difference,
w hich might account for the double peaks in spawning in some areas. The species has been best studied o ff W est
A frica, w here there are three quite distinct ‘stocks’:
(i)
N orth Transitional
(ii) Central U pw elling
(iii) South Transitional
Sardinella brachysom a
Zone (M auritania to G uinea; also, o ff Sierra Leone - perhaps a separate stock)
Zone (Ivory Coast, also Ghana)
Zone (southern Gabon to southern A ngola).
B leeker, 1852
CLUP
S a rd l 5
Sardinella brachysom a B leeker, 1852, V erh.batav.G enoot.K unst.W et., 24:19 (Batavia).
Synonym s : H arengula hypselosom a Bleeker, 1855:427 (A m boina); M eletta schlegelii Castelnau, 1873:93
(D arw in); S ardinella albella:F ow ler, 1941:605 (incorrect synonym y); Sardinella brachysom a - C han, 1965:19, fig. 9
(India; but China, Hong Kong specimens probably
hualiensis); W hitehead et ah, 1966:49 (types o f brachysom a,
hypselosom a); W hitehead, 1973b: 182, fig. 12; W ongratana, 1980:127, pis 67, 68 (revision).
FAO N am es :
E n - Deepbody s a r d in e lla
96
D iagnostic Features : Body deep and com pressed, its depth 30 to
39% o f standard length; total scutes 29 to 32. Low er gillrakers 48 to 67,
hardly increasing w ith size o f fish. Vertical striae on scales overlapping or
som etim es continuous at centre o f scale, num erous small perforations on
hind part o f
scale.
A dark spot at dorsal fin origin. Closely
resem bles Sk
hualiensis o f
China, w hich has dark dorsal and caudal fin tips.
D iffers from
other deep-bodied species w ith overlapping
or continuous scale
striae
chiefly in its lower gillraker count (87 to 134 in Sk filíense, 88 to 126 in Sk
dayi) and presence o f a dark spot at dorsal fin origin (absent in Sk
richardsoni and Sk zunasi). See CLUP Sardl 5, Fishing A reas 57, 71.
overlapping
striae
scale
G eographical D istribution
Indo-W est Pacific
(M adadagascar, but apparently not elsew here in the
w estern Indian O cean; M adras, Indonesia, northern
A ustralia; C han’s Hong Kong and Taiping specimens
were most likely S^ hualiensis).
Habitat and B iology :
M ore data needed.
Size :
12 cm.
To
Coastal, pelagic, schooling.
13 cm standard length, usually to
Interest to F isheries : Included in the general
statistics for Sardinella; probably o f some local im por­
tance in parts o f Indonesia.
20" '
Local Names : -
Sardinella brasiliensis
(Steindachner, 1879)
C L U P S a rd l 11
Clupea brasiliensis Steindachner, 1879, Sber.Akad.Wiss.W ien, 80:182 (Rio de Janeiro).
Synonym s : Clupea janeiro Eigenm ann & Bray, 1849, A nn.N .Y .A cad.nat.Sci.:626 (replacem ent name in view
o f the hom onym C lupea brasiliensis Schneider, 1801 = A lbula vulpes L innaeus,1758; the name laneiro seems never
to have been adopted and although corect should be rejected on grounds o f - usuage); S ard in ella b rasilien sis FW N A , 1964:407, fig. 101; W h iteh ead , 1973a:30 (G uianas).
FAO N am es
En - B ra z ilia n sard in ella.
D iagnostic Features : See Sardinella aurita, from w hich it differs in having the anterior gillrakers on the
low er lim bs o f the second and third gili arches distinctly curled dow nw ard (m ore or less flat in Sk au rita. The
pelvic finray count o f i 8 d istin g u ish es Sk b rasilien sis from all o th er species o f S a rd in ella, also H aren g u la and
O pisthonem a. R esem bles Strangom era o f A rgentina and C hile, but has tw o fleshy appendages along outer m argin
o f gili opening (smooth in Strangomera) and numerous fine fronto-parietal striae on top o f head. See CLUP Sardl
11, Fishing A rea 31.
97
fla t
curled
fleshy
' outgrowths
S. brasiliensis
S. aurita
low er limbs o f gili arches
fro n to -p arietal striae
on top o f head
G eographical D istribution :
W estern A tlantic (G u lf o f
M exico, Caribbean, W est Indies southward to Brazil and northern
U ruguay, but identifications not alw ays reliable).
See also
rem arks.
H abitat and B iology : Coastal, pelagic, schooling. Probably
sim ilar to £L a u rita , but i f ind eed th is sp ecies is d istin ct from £L
a u rita th e n it m ay a cc o u n t fo r one o f th e a p p a re n t d o u b le
spawning peaks o f that species.
Size : To 25 cm standard length, usually around 20 cm.
Interest to F isheries : L arge fishery in V enezuela, but
statistics do not separate if from Sk aurita.
Local N am es : USA:
O rangespot sardine (Florida).
L iterature : Included in many o f the w estern A tlantic
references to Sk aurita and perhaps m ixed w ith Sk aurita even in studies purporting to deal only w ith S. b rasiliensis
(e.g. M atsuura, 1975, 1977 - southern Brazil, spawning, eggs, larvae: Figureido & M enezes, 1978:23, fig. 27).
Rem arks : Separation o f S. brasiliensis from the w idespread Sk aurita on the basis o f gillrakers shape and a
higher gillraker count is tentative, the more so since fishes w ith the brasiliensis curled form o f gillrakers occur
o ff W est A frica and in the M editerranean.
Sardinella fijiense
(Fow ler & B ean, 1923)
C L U P S a rd l 17
H arengula fijiense Fow ler & B ean, 1923, P roc.U .S .natn.M us.. 63:63 (Fiji).
Synonym s : H arengula fijiensis Fow ler,
Fow ler, 1941:599, fig. 15 (his IL fijiensis
W ongratana, 1980:125, pis 61, 62 (revision).
1928:31 (unjustified em endation o f name): H arengula nym phaea:
in the synonym y: his Fiji m aterial only): Sardinella fijie n se:
FAO N am es : En - Fiji sardinella.
click fo r n e x t p a g e
f t
overlapping or
continuous striae
D iagnostic Features : Body fairly slender, its depth usually less than
30% o f standard length; total scutes 29 or 30. Low er gillrakers slender,
sm ooth and num erous, 87 to 134 in fishes 6 to 11 cm standard length (but
more data needed). Vertical striae on scales continuous or overlapping, but
no perforations on hind part o f scale. No dark spot at dorsal fin origin;
tips o f dorsal and caudal fins dusky or black. Im m ediately distinguished
from all other Sardinella species in the area by its high gillraker count, also
the overlapping or continuous scale striae and absence o f dorsal spot.
G eograph ical D istrib u tion
(Papua N ew Guinea, Fiji).
W estern
Pacific
^V
scale
^ ¿A*? &
r1
»4
H a b ita t
and
B io lo g y
schooling.
More data needed.
C o astal,
■ ft
r
pelag ic.
r
*
«*
*
Size : To 11.5 cm standard length.
Interest to F isheries
Presum ably contributes
to S ard in ella catc h e s, b u t ap p ears no t to be abu n d an t.
Local N am es : -
H,
V
Sardinella fim briata
(V alenciennes, 1847)
k rÇ 7
C
C LU P
9
Sardl
Spratella fim briata V alenciennes, 1847, H ist.n a t.p o iss., 2 0 :3 5 9 (M alabar).
Synonym s : Sardinella fim briata - Fow ler, 1941:609 (the Philippines); C han, 1965:14, fig. 7a (India,
Thailand, the Philippines); W hitehead, 1967:50 (type); W hitehead, 1973b: 184, fig. 14 (but not the W estern Indian
Ocean records); W ongratana, 1980:119, pis 56, 57 (revision); W hitehead & Bauchot, in press (types o f S. fim briata).
FAO N am es
: En - Fringescale
sardinella.
7
99
D iagnostic F eatures : Body som ewhat com ­
pressed but variable, from slender to m oderately
deep, its depth 25 to 34% o f standard length; total
num ber o f scutes 29 to 33 (usually 30 to 32). Lower
gillrakers 54 to 82 (at 5 to 12 cm standard length,
increasing w ith size o f fish). V ertebral striae on
scales not meeting at centre, hind part o f scale with
a few perforations and (in Indian Ocean specimens)
som ew hat produced posteriorly. A dark spot at
dorsal fin origin. C losely resem bles £L alb ella,
w hich has few er low er gillrakers (41 to 68, most
specimens less than 60) and many more perforations
on the scales. Often confused with £L gibbosa, w hich
has more scutes (32 to 34) and few er gillrakers (45
to 59). Other sim ilar species have more scutes, or mo
fin base or caudal tips black (or a com bination o f some
discontinuous
, X striae \
,
perforations
S.
scale
gillrakers or overlapping scale striae or no spot at dorsal
these features). See CLUP Sardl 7, Fishing A reas 57, 71.
G eographical D istribution : Indo-W est Pacific (not
in w estern Indian O cean, but from southern India and Bay
o f B engal to the P h ilip p in es, also eastern tip o f Papua
N ew Guinea).
H abitat and B iology : C oastal, pelagic, schooling.
M isidentifications (especially w ith
£L gibbosa in Indian
w aters and £L albella in the w estern Indian O cean) make
published biological data potentially unreliable. The bio­
logy is summarized by Nair (1960, 1973).
Size :
11 cm.
S. albella
fim b riata
To 13 cm standard length, usually around
Interest to F ish eries : Included in the general
statistics for Sardinella, but o f some im portance in south­
e rn p a rts o f In d ia (a lth o u g h fis h e ry s ta tis tic s p ro b a b ly
include £L gibbosa and/or £L albella).
L ocal N am es
: INDIA: C halam athi (M alayalam );
Charree addee (H indi); Erebai, Pedi (K annada); H oira,
K haira (Bengali); Noone kavallu (Telegu); Pedw a, W ashi
(M arathi); Saudai (Tam il). These nam es are probably
equally applied to similar species in many areas.
L iterature : N air (1960, 1973 - India, synopsis o f biology and fisheries).
100
(B leeker,
Sardinella gibbosa
C LU P
1849)
Sardl
8
Clupea gibbosa B leeker, 1849, J.In d .A rc h ., 3:72 (M acassar).
Synonym s : Clupanodon jussieu L a c e p è d e , 1803:469, ? pi. 11 (M auritius; nomen dubium );
Spratella tembang
Bleeker, 1851:214 (nam e only to replace Clupea gibbosa); C lupea im m aculata K ishinouye, 1907:96, pis 19, fig. 1,
21, fig. 4 (Saga, K yushu, A m oy, Swatow, China; types destroyed); Sardinia im m aculata:Chyung, 1961:135 (Korea);
Fim briclupea dactylolepis W hitley, 1940:399, fig. 5 (northw estern A ustralia); Sardinella dactylolepis :M u n ro .
1956:23, fig. 157; S ard in ella taiw an e n sis R aja & F liy am a, 1969:90, p i 2 6 (Taiwan); Sardinella ju ssie u :Fow ler.
1941:611 (the Philippnes);
W hitehead, 1965:252 (Gulfs o f Aden and O m an); Sardinella iu s sie u i:C han. 1965:9, fig.
20 (Thailand to Flong Kong); W hitehead et ah, 1966:56, 58 ( gibbosa, tem bang ty p es); W h iteh ead 1967:54 (ju ssieu
problem );
Sardinella gibbosa - L o s s e , 1968:98 (E ast A fric a ); W h ite h e a d , 1973b:185, fig . 15; W o n g r a ta n a ,
1980:117, pis 54,55 (revision, im m acu lata identified); SFSA, in press (southern Africa).
E n - G oldstripe
FAO Nam es
sardinella.
D iagnostic Features : Body m oderately slen­
discontinuous
der, its depth u sually 24 to 30% stan d ard length;
striae
total num ber o f scutes 32 to 34. Low er gillrakers 45
to 59 (at 6 to 17 cm standard length, not increasing
w ith size o f fish after 6 cm standard length).
V er­
t i c a l s t r i a e o n s c a l e s n o t m e e t in g a t c e n t r e ,
num erous sm all p erfo ratio n s on hind p a rt o f scale.
A golden midlateral line down flank (at least in G ulf
o f Thailand); dorsal and caudal fin m argins dusky; a
dark spot at dorsal fin origin. Closely resembles S.
perforations
sinden sis (A rabian Sea and th e “ G u lf ’), w h ich has
fe w e r p e rfo ra tio n s on th e sc a le s.
M o st o fte n
S_. gibbosa
S. sindensis
confused w ith S. fim b ria ta w hich has few er scutes
scale
(usually 29 to 32) and m ore g illrak ers (54 to 82).
Other sim ilar species have few er scutes or more gillrakers or overlapping scale striae or deeper bodies or no spot
at dorsal fin base or caudal tips black (or a co m m atio n o f some o f these features). See CLUP Sardl 8, Fishing A rea
57. (Sheet for Fishing A reas 57, 71 not-accurate).
G eographical D istribution : Indo-W est Pacific
(the “G u l f ’, but ap p aren tly not R ed Sea; E ast A frican
coasts, M adagascar eastw ard to Indonesia, north to
Taiwan and K orea south to northern A ustralia, possibly
also w estern coasts o f A ustralia). One o f the m ost
abundant Sardinella species in the Indo-W est Pacific
region.
H abitat and B iology : Coastal, pelagic, schooling.
Possible or even probable confusion w ith other species
( e s p e c ia lly £L f im b r ia ta in I n d ia n w a te r s ) m a k e s
p u b lish e d b io lo g ic a l d a ta p o te n tia lly u n re lia b le . The
biology is summarized by Nair (1973:39 to 48).
Size
15 cm.
: To
17
cm
standard
length,
usually
to
101
Interest to Fisheries :
Included in the general statistics for Sardinella, but o f some im portance in southern
parts o f India (A ndhra Pradesh, Tamil N adu, K erala) and entering m arkets throughout southeast Asia. It is usually
caught in association w ith other species o f Sardinella.
L o ca l N am es : H O N G KONG: H w ang lum ; IN D IA : C halam athi (M alayalam ); Choodai, N onalai (Tam il);
Erebai (Kannada);
K avallu (Telugu); IN D O N E SIA : Tem bang; TA IW A N ISLA N D :
Ju shi sh a-tin; TH A ILA N D :
Pia lang keo. These names are probably equally applied to sim ilar species in many areas.
L iterature : N air (1960, 1973 - India, synopsis o f biology and fisheries); Losse (1968 - East A frica);
(1974 - East Africa).
Sardinella hualiensis
C LU P
(Chu & Tsai, 1958)
H aren g u la h u alien sis C hu & T sai,
Taitung, east coast o f Taiwan Island).
1958, Q u a rt.J.T a iw a n M u s .. 11(1-2): 16, pi.
Sardl
O kera
18
3, figs 2, 3, 4 (H ualien,
Synonym s : Sardinella brachysom a:Chan. 1965:19 (the China and perhaps Hong Kong specimens); Sardinella
hualiensis :W ongratana. 1980:126 pis 65, 66 (revision).
FAO Nam es :
E n - T aiw an sardinella.
y
T A «S*
Wi- r
__,>T'
overlapping or
continuous striae
D iagnostic F eatures : Body m oderately deep and com pressed, its
d ep th 29 to 34% o f stan d ard len g th ;
to ta l
scu tes 30 to 32.
Lower
g illrak ers 51 to 66, hardly in creasin g w ith size o f fish. V ertical striae o f
scales overlap p in g or som etim es co n tin u o u s at cen tre o f scale, num erous
sm all p erfo ratio n s on hind p art o f scale. Tips o f caudal fin lobes black.
Closely resembles £L brachysom a from Indonesia, etc., w hich does not have
black caudal fin tips. F rom o th er deep -b o d ied species in the area w hich
have overlapping or continuous scale striae it differs in having a dark spot
at dorsal fin origin (absent in £L richardsoni and S. zunasi!.
p e rfo ra tio n s
scale
..............
------ !......... .... .. “Y..
G eographical D istribution : Western Pacific (Taiwan
Island, possibly south to Hong Kong).
1
H abitat and B iology : Coastal, pelagic, schooling.
M ore data needed.
S ize : To 12.5 cm standard length, usually
10 cm.
Interest to F ish eries : Probably
local contribution to Sardinella catches.
to about
m akes a
20
"
'
•
sm all
k
^
Local Nam es : -
s.
Or
tS1
k
b
C lupeonia
CLUP
(V alenciennes, 1847)
Sardinella jussieui
jussieui
Sardl
19
V alen cien n es,1847, H ist.n at.p o iss., 20:346, pi. 599 (M auritius).
S y n o n y m s : C lupeonia ju ssieu i:Sauvage, 1891:495 (M adagascar); Sardinella dayi R egan, 1917:381
(India);
C h a n ,1 9 6 5 :1 2 , f i g . 2 2 ( C e y 1o n ) ; W hitehead, 1973b: 181, fig. 10; R a ja & L a z a ru s , 1975:114, figs 1,2 (Karwar,
T u tic o r in , V iz h in ja m ) ; W o n g ra ta n a , 1 9 8 0 :1 2 4 ,pi. 60 (rev isio n );
S ardinella ju ssie u i:W hitehead, 1967:59 (type);
Idem , 1973b: 181 fig. 11; W o n g ratan a, 1980:114, pis 50, 51 (revision).
note:
the C alicut specim ens o f Fow ler
(1941:604, fig. 16) are H ilsa kelee; W hitehead & B auchot, in press (type o f jussieui).
FAO Names
En - M auritian sardinella.
AVw\A/v\AAyW
/; X1 >% ñ I
discontinuous
striae
D iagnostic Features : Body com pressed and deep, its depth 28 to
37% o f standard length; total numbers o f scutes 31 or 32. Lower gillrakers
88 to 126 (in fishes 9.5 to 11 cm standard length, strongly increasing with
size o f fish).
V ertical striae on scales not m eeting at centre. The only
o th er sp ecies o f S a rd in e lla w ith such a h ig h g illra k e r c o u n t (m ore th a n 87)
is S. fijiense (black caudal tips, western Pacific).
103
G eograp hical D istrib u tion
Indo-W est Pacific
(w estern
coasts o f southern India, from Bombay south to Sri Lanka; also to
M adagascar, M auritius).
H abitat
data needed.
Size
and
To
B iology
12 cm
C oastal,
pelagic,
schooling.
M ore
standard length, usually to about 10 cm.
Interest to F isheries : Enters m arkets in southern India
(e.g.
at
C ochin), but probably not sufficiently
abundant to make any
significant contribution to Sardinella catches.
L ocal N am es : L iterature : See under synonyms.
R em arks : A ll m eristic counts o f Sk dayi o f southern India
overlap those o f S. jussieui
o f M auritius. W ongratana (1980:45 -key)
separated the tw o on scute counts, but his counts do not support this
(30 to 32 versus 31 or 32). There is a great variation in body depth,
even in specimens from the same batch, and a sim ilar v ariatio n in
gillrakers numbers. M ore material should be studied.
Sardinella lem uru
Bleeker,
( h
C L U P Sardl 14
1853
Sardinella lem uru Bleeker, 1853, N a tu u rk .T ijd sc h r.N e d .-In d ië . 4:500 (B atavia).
Synonym s : Clupea nymphaea Richardson, 1846, I c h th y o l.C h in a Japan:304 (China Sea) (name suppressed by
International C om m ission in 1970, O pinion 901, B ull.Z ool.N om encl.. 26(5-6):217);
A m b ly g aster p o steru s W hitley,
1931:144 (W estern A u stralia); A m b ly g a s te r p o ste ra :M unro. 1956:22, fig . 154;
S ard in ella sam arensis R o x as,
1934:275, pi. 2, fig. 11 (Sam ar, th e P h il ip p i n e s ) ;
S a rd in e lla lo n g ic e p s:F ow ler, 1941:603 (Philippine m a te ria l);
C h an , 1965:3, fig. 17 (Philippine material); Sardinella aurita R aja & H iyam a, 1969:78 (the Philippines, Hong Kong,
Taiwan Island; the Hong Kong specimens designated Sk aurita terrase i.e. terrasae Lozano y Rey, a supposed W est
A frican subspecies); S ardinella le m u ru :W ongratana. 1980:111, pis 47 , 48 (rev isio n ). See R em arks for fu rth er
w estern Pacific S. aurita references.
FAO Names
En - Bali sardinella.
V r , y „ , . V '. v
AAVm
A R ' >)
D iagnostic Features
Body elongate, subcylindrical, its depth less than 30% o f standard length, belly
rounded. The pelvic finray count o f i 8 distinguished Sk lem uru from all other clupeids in the eastern Indian Ocean
and w estern P acific. Very closely resembles R lo n g ice p s (w h o se range it may overlap in the A ndam an Sea), but
head shorter (26 to 29% o f standard length; cf.79 to 35% in R longiceps) and low er gillrakers few er (77 to 188 in
fishes o f 6.5 to 22 cm standard length; cf. 150 to 253 in R lo n g icep s o f 8 to 15.5 cm, usually more than 180). A
fain t golden spot behind gili o p ening, follow ed by a fain t-g o ld en m idlateral line; a d istin ct black spot at hind
border o f gili cover (absence o f pigment).
104
G eographical D istribution : E astern Indian Ocean
(P h u k et, Thailand, southern coasts o f East Java and Bali;
W estern A ustralia) and w estern Pacific (Java Sea north to
the P h ilip p in es, H ong K ong, T aiw an Islan d to so u th ern 40*
Japan - see Remarks).
H abitat and B iology : C oastal, pelagic, schooling,
strongly m igratory.
F e e d s on p h y to p la n k to n , a lso Z o o ­
p lankton (chiefly copepods). In the B ali reg io n spaw ning
probably occurs at the end o f the annual rainy season
m igration
into the B ali
Strait (u sually
S eptem berF ebruary, but a peak m ainly in D ecem b er-Jan u ary , at
least ju d g in g by num bers caught), but the spaw ning
grounds are not knwon. A s in the case o f the related
species (SF a u rita ,
SF lo n g iscep s), spaw ning and m ajor
m igrations appear closely linked w ith hydrological condi­
tions (especially tem perature), but sudden appearances or
d is a p p e a ra n c e s o f sh o a ls h av e no re a d y e x p la n a tio n .
Spaw ning in the E ast C hina Sea reach es a peak in late
M arch to M ay, but continues through August.
Size : To 23 cm standard length, usually to 20 cm .
ta
80°
100°
120°
140°
160°
180°
Interest to F ish eries : O f m ajor in terest in the
E ast C hina Sea (100 000 to n s in 1971) m ainly o ff so u th ern F u jian and e a s te rn G u a n d o n g p ro v in c e s ; and in
Indonesia, the bulk o f the catch being from the Bali Strait.
The total catch in 1983 was 59 980 tons (all from
Indonesia).
L ocal N am es
: HONG KONG: H wang tseih; IN DONESIA: Lem uru; TAIW AN ISLAND: H wang
sha-tin.
Literature : Soerjodinoto (1960 - biology, fishery); Li Kwan-M ing (1960 - biology, fishery); Ritterbush (1974
- Bali, population biology); Chiu & Tsongchion (1982 - East China Sea, biology fishery).
R em arks : Regan (1917:378) included two Japanese specimens in his w ideranging SF aurita,while Fow ler
(1931:116) included C hin ese specim ens (Amoy, S w atow , C an to n ; he later added a Jap an ese specim en (Fow ler,
1941:602 - as S. a lle c ia ). It seem s m uch m ore lik ely th a t £F lem u ru ex ten d s its range no rth w ard from the
Ph ilip p in es to C h in a a n d so u th ern Japan.
M u c h m o re w o rk is needed before it can be d efin itely stated that
S. lem uru is not merely an eastern form o f SF aurita, from w hich, given the variation in body shape and gillraker
num bers in A tla n tic S^ a u rita , it c an n o t be d istin g u ish ed as y et on m o rp h o lo g ical g rounds.
Sardinella longiceps
Valenciennes,
C L U P S a rd l 3
1847
Sardinella longiceps V alenciennes, 1847, H ist.nat.poiss., 20:273
(Pondicherry).
Synonym s : Sardinella neohowii Valenciennes, 1847, H ist.nat.poiss., 20:274 (Cannanore); A lausa scom brina
V alenciennes, 1847, H ist.nat.poiss., 20:442 (Cannanore); Sardinella longiceps - W hitehead, 1965:249 (Gulfs o f Aden
and O m an, not Red Sea); C han 1965:3, fig. 17; W hitehead, 1967:44-46 (types o f longiceps, neohow ii, scom brina);
L o sse , 1968:96, pi. I d (E ast A frica); W h i t e h e a d , 1973b:177, fig . El (N . In d ia n O c e an re c o rd - N a ir, 1973:4
(India, synopsis); W ongratana, 1980:108, pts 43, 44 (revision); W h iteh ead & Bauchot, in press (types o f longiceps,
neohow ii, scom brina).
FAO N am es :
En - Indian oil sardine.
105
D iagnostic Features : Body elongate, subcylindrica!, its depth less than 30% o f standard length, belly
rounded. The pelvic finray count o f i 8 distinguishes S. longiceps
from all other clupeids in the northern Indian
Ocean.
Very closely resembles Sk neglecta o f East A frican coasts, but head longer (29 to 35% o f standard length;
cf. 26 to 29% in Sk neglecta) and more low er gillrakers (150 to 253 in fishes o f 8 to 15.5 cm standard length,
usually more than 180; cf. 143 to 188 in Sk neglecta o f 9.5 to
13 cm standard length, usually less than 185).
D istinguished in the same way from Sk lem uru
(whose range it may overlap in the A ndam an Sea), but the latter has
even few er gillrak ers (77 to 188 in Sk lem uru o f 6.5 to 22 cm standard length). A faint golden spot behind gili
opening, followed by a faint golden m idlateral line; a distinct black spot at hind border of-gill cover (absence o f
pigment).
Geographical D istribution : Indian O cean (northern and w est­
ern parts only. G ulf o f A den, G u lf o f Oman, but apparently not Red
Sea or the “G u lf’ eastw ard to so u th ern p art o f In d ia , on eastern
coast to A ndhra; possib ly to Andam ans).
H abitat and B iology : C oastal, pelagic, schooling, strongly
m igratory. Feeds m ainly on phytoplankton (especially diatom s),
both as ju v e n ile s and ad u lts, b u t also on Z ooplankton (esp ecially
copepods by th e ju v e n ile s ). B re e d s o n ce a y e a r, th e sp a w n e rs
arriving o ff w estern coasts o f India in June-July when tem peratures
and s a lin ity are lo w d u rin g th e so u th w e s t m o n so o n m o n th s; an
extended spaw ning season, b u t m ost in ten se in A u g u st-S ep tem b er;
e x act spaw ning g ro u n d s n o t lo cated (N a ir, 1973:18 - w ho su m m a­
rized the biology o f the species).
2QO
o°
Size : Perhaps to 23 cm standard length, usually to 20 cm.
20°
Interest to F ish eries : The m ost im portant and abundant
clupeoid in Indian w aters (up to 30% o f all marine fishes), although
w ith c o n s id e ra b le v a ria tio n s in c a tc h e s fro m y e a r to y e a r, e.g.
7 400 tons in 1956, but 189 000 tons in 1960 (Jhingran, 1982:527).
The total catch for 1983 402 586 tons, o f which 350 260 tons came
from India.
L ocal N am es : INDIA: M athi (M alayalam ), Boothai (Kannada), Taralai, Haid (M arathi).
L iterature : N air (1960, 1973 - the best sum mary o f biological data); other references to biological studies
sum marized by W hitehead (1973b: 178).
click for n e x t p a g e
c lic k fo r p r e v io u s p a g e
106
Sardinella m aderensis (Lowe, 1839)
C L U P S a rd l 2
C lupea m aderensis Low e, 1839 (June), Trans.zo o l.Soc.L ond., 2(3):189 (M adeira) (w rongly dated 1841 in some
publications, e.g. CLOFNAM).
Synonym s : Sardinella granigera Valenciennes, 1847:267 (Egypt): A lausa eba Valenciennes, 1847:417 (Corée,
Senegal): P ello n u la m o d e s ta Fischer, 1885 (Elobey, Equatorial Guinea): Sardinella cam eronensis R egan, 1917:380
(Cam eroon): Sardinella m aderensis:Ben-Tuvia, 1960:499 (synopsis): W hitehead, 1967:47, 49 (types o f g ran ig era,
eba): CLOFNAM , 1973:103 (synonymy): FNAM , 1984:223 (synopsis): CLO FETA , in press (com plete b ib lio g rap h y ;
SFSA, in press (southern A frica): W hitehead & Bauchot, in press (types o f granigera, eba).
FAO N am es : En - M adeiran sardinella.
D iagnostic Features : Body elongate, but variable in depth, belly fairly sharply keeled: total
scutes 31 to
34. Low er gillrakers 70 to 166 (in fishes 6 cm standard length or more). U pper pecto ral finrays w hite on outer
side, the m em brane betw een black. R esem bles £L aurita, but pelvic finrays i 7 (i 8 in R. aurita) and no black spot
on hind part o f gili cover (but faint gold or black area ju s t behind gili opening). D istin g u ish e d from S. rouxi by
having m ore low er g illrak ers (only 30 to 40 in S. rouxi) and the caudal fin grey, its tips alm ost b la c k (yellow in
R ro ux i). See CLUP Sardl 2, Fishing Area 34.
G eographical D istribution : M editerranean (southern and
eastern parts, also p en etratin g Suez C an al): eastern A tla n tic 40°
(G ilb raltar southward to A ngola and a single recorded specimen
from W alvis Bay, Namibia).
H abitat and B iology : C oastal, pelagic, but tolerant o f
low salinities in estuaries, schooling, preferring w aters o f 24°C , 20°
at su rfa c e or at b o tto m d o w n to 50 m , stro n g ly m ig ra to ry .
Feeds on a variety o f sm all plank to n ic in v erteb rates, also fish
larvae and phytoplankton. B reeds only once in the year, during
the w arm season (Ju ly -S ep tem b er), in co astal w aters. Ju v en iles
and a d u lts sh o w c le a r n o rth -s o u th m ig ra tio n s in th e G abon- °°
C o n g o -A n g o la se c to r o f th e ir ra n g e and a lso in th e S ie rra
L e o n e -M a u rita n ia se c to r, e a c h a re a h a v in g n u rs e rie s :
th e
m ovem ents are co rrelated w ith the seasonal u p w elling. M ig ra­
tions in the central part (Ivory C oast-Ghana) are not so marked. 2QO
Size : To 30 cm standard length, usually to 25 cm.
Interest to F isheries : O f considerable im portance o ff
W e st A fric a n c o a s ts , b u t c o m b in e d w ith —
S. -------a u rita in m o st 40°
statistics, partly because the tw o are often caught together.
Local Names
:-
L iterature : See under synonyms.
R em arks : M any more W est A frican references in the literature (right up to the present tim e) give this fish
as R e b a ; a substantial num ber use the name R cam eronensis for fishes caught in the Cam eroon area. A ttem pts
to distinguish three species, how ever, rem ain u n c o n v in cin g in view o f the considerable variation in body depth and,
most likely, gillraker numbers.
107
Sardinella m arquesensis
Berry
C L U P S a rd l 20
& W hitehead, 1968
S ard in ella m arq u esen sis B erry & W h iteh ead , 1968, P ro c. bi ol. S oc. W a s h ., 8 1 :2 0 9 -2 2 2 , figs 1-6 (M arq u esas
Islands).
Synonym s :
FAO N am es
Sardinella m arquesensis:W ongratana. 1980:131, pis 73, 74 (revision).
: E n - M arquesan sardinella.
d isco n tin o u s
striae
D iagnostic Features : Body m oderately slender, its depth 25 to 30%
o f standard length;
to ta l sc u te s 2 8 to 30.
L ow er g illrak ers strongly
increasing with size o f fish, about 45 at 5 or 6 cm standard length, 50 to 80
at 6 to 10 cm and a m axim um o f about 80.
V ertical striae on scales not
m eeting at centre, no perforations on hind part o f scale. No dark spot at
dorsal fin origin, no dark tips to caudal fin. There appear to be no other
Sardinella species in the M arquesas area; it can be sep arated from other
species by a com b in atio n o f ch aracters (slen d er body, few scutes, scale
striae interrupted, many gillrakers, no dark dorsal spot and plain caudal fin
tips).
G eographical D istribution :
W estern Pacific (M ar­
quesas Islands only; m ore recen tly (1955) in tro d u ced into
Haw aiian w aters - Murphy, 1960).
H abitat and B iology
Size
10 cm.
Coastal, pelagic, schooling.
To 12.5 cm standard length, usually about
Interest to F ish eries : U sed as a b a it f i s h in the
M arquesas, but w hen transported to H awaii never becam e
su fficien tly ab undan t nor m ade any sig n ifican t c o n trib u ­
tion.
Local N am es : L iterature : B ro ck (1960 - H awaii); M urphy (1960 Hawaii);
Hida & M o rris (1963 - H a w a ii); N a k a m u ra &
Wilson (1970 - biology).
no p erfo ratio n s
108
Sardinella m elanura
C L U P S a rd l 4
(C uvier, 1829)
C lu p ea m elan u ra C u v ier, 1829, R ègne a n im ., 1st ed ., 2:318 (f o o tn o te , o n L acep è d e : “CL m e l a n u r a , N .,
L acép. V, xi, 3, sous le nom de C lu p an o d o n J u s sie u , m ais la description se rapporte à la fig. xi, 3, nommé v a rié té
du C lupanodon chinois”) (See discussion in Whitehead, 1967:62).
Synonym s : Clupeonia comm ersoni Valenciennes, 1847:350 (near Bom bay); Clupea otaitensis Valenciennes,
1847:351 (on Solander M S n a m e , in synonym y o f CL c o m m erso n i, th u s a nomen nudum );
C lupeonia v i t t a t a
V a le n c ie n n e s , 1847:352 (V a n ik o ro ); H a re n g u la v an ic o ris Jo rd an & S e a le , 1906:187 (on A lau sa m elan ura o f
V alenciennes);
S a rd in e lla n ig ric a u d a ta C h a n , 1965:7, fig. 19 (M alak u la Islan d );
H e rk lo tsic h th y s v itta tu s :
W h iteh ead , 1965247 (G u lf o f A den): Sardinella m e la n u ra :F o w ler, 1941:614 (M au ritiu s, the P h ilip p in e s, Sam oa,
Tahiti; D elagoa Bay record needs to be checked); Chan, 1965:5, fig. 18 (Fiji, N ew H ebrides); W hitehead, 1967:62,
64, 65, 66 (types o f m elanura, com m ersoni, otaitensis, vittata); Id e m .
1973b: 180, fig. 9; W ongratana,
1980:132,
pis 75, 76 (revision); W h iteh ead & B auchot, in press (types o f m elanura, com m ersonii, otaitensis, vittata).
FAO N am es
: En - B la c k tip
sardinella.
V
D iagnostic Features : Body som ewhat com pressed, its depth usually not much more than 30% o f standard
length. The je t b lack c a u d a l tip s d is tin g u ish SL m elanura from all other species o f Sardinella except S. atricauda
(s c u te s 32 to 35; cf. 27 to 31, usually 28 to 30 in SL m elanura), S. h u alien sis (dark spot present at dorsal fin origin,
scales fimbriated and w ith small perforations, re c o r d e d so far only from Taiwan Island) and SL fijiense (gillrakers
87 to 134; c f only 38 to 74 in SL m elanura). See CLUP Sardl 4, Fishing A rea 51, also 57, 71.
G eographical D istribution : Indo-W est Pacific
(w e s te rn In d ia n O c ea n fro m G u lf o f Aden so u th to
M a d a g a s c a r and M a u ritiu s , a p p a re n tly n o t so u th o f
Bombay nor in northern Bay o f B engal; Indonesia, not
South China Sea; eastward to Samoa).
H abitat and B iology : Coastal, pelagic, schooling.
M ore data needed.
Size : To 12.2 cm standard length, usually about
10 cm.
Interest to F ish eries : Included in the general
statistics for S ard in ella; perhaps o f local im portance,
but appears not to be very abundant.
L ocal N am es:
Islands).
IN D IA : Sapta t a r n i
(A ndam an
109
Sardinella neglecta
CLUP
W ongratana, 1983
Sardinella neglecta W ongratana,
1983,
Sardl 15
Jap.J.Ichthyol., 29(4):390, fig. 4 (Kenya, Tanzania).
Synonym s : Sardinella longiceps (w estern Indian Ocean only) o f authors.
FAO N am es : En - E ast A frican sardinella.
D iagnostic Features :
Body elongate, subcylindrical, its depth less than 30% o f standard length, belly
rounded. The pelvic finray count o f i 8 distinguishes £L neglecta from all other clupeids along the E ast A frican
coast. Very closely resem bles £L longiceps (whose range it may overlap), but head shorter (26 to 29% o f standard
length; cf. 29 to 35% in £L longiceps and few er low er gillrakers (143 to 188 in fishes o f 9.5 to 13 cm standard
length, usually less than 185; cf. 150 to 253 in £L longiceps o f 8 to 15.5 cm standard length, usually more than
180). A faint golden spot behind gili opening, followed by a faint golden
midlateral line;
a distinct black spot at
hind b o rd er o f-g ill co v er (ab sen ce o f silv er p igm ent).
G eographical D istribution
K enya and Tanzania).
: W estern Indian O cean (coasts o f
H abitat and B iology : C oastal, pelagic, schooling, in w aters
o f 1 to 60 m, in bays and over shallo w banks. F eeds presum ably
mainly on phytoplankton like S. longiceps and its breeding biology is
perhaps similar to that species too.
Size : To 13 cm standard length, but probably grows larger.
Interest to F isheries : A pparently not abundant, occurring
mostly during the northeast m onsoon period (N ovem ber-January).
Local N am es : L iterature : Losse (1968 - taxonom y, biological notes).
110
Sardinella richardsoni W ongratana, 1983
C LU P Sardl 22
S ard in ella (C lupeonia) richardsoni W ongratana, 1983, Jap.J.Ichthvol., 29(4):390, fig. 5 (replacem ent name for
C lupea isingleena Richardson).
Synonym s :
C lu p e a is in g le e n a R ic h a rd s o n , 1846, Ich th v o l.S eas C hina Ja p a n :304 (C h in a seas) (nam e
su p p ressed b y Int.Com m .Zool.N om encl.
in 1 9 7 0 , O p in io n 9 0 1 , B ull.zool.N om encl.. 26(5-6):217);
S ardinella
fim b ria ta :W hitehead . 1966:24 p i. 1, fig . 2 ) ( t y p e ) ; S a rd in e lla ric h a rd soni W o n g r a ta n a . 1980:129, pis
69,70
(revision).
FAO Names :
En - R ic h a rd so n ’s sardinella.
im m
D iagnostic Features : Body fairly com pressed and deep, its depth 32
to 36% o f standard length; total scutes 30 to 32. L o w er g illra k ers 63 to
74, p ro b a b ly n o t in c re a s in g g re a tly a fte r a b o u t 7 cm sta n d a rd len g th .
Vertical striae on scales continuous or overlapping, num erous small perfo­
rations on hind part o f scale.
N o dark spot at dorsal fin origin. M ost
closely resem bles £L z u n a si, w h ic h has o n ly 4 2 to 58 lo w e r g illra k e rs
(usually less than 55 in fishes up to 12 cm standard length) and is more
slender (depth 24 to 33% o f standard length, usually 31% or less). O f other
species in the area, Sk hualiensis also has overlapping or continuous scale
striae, but black tip s to dorsal and cau d al fins; while Sk fim briata and £L
a lb ella have the scale striae interrupted at the centre o f the scale.
Geographical Distribution
Island, Hong Kong, China).
H abitat and B iology :
M ore data needed.
Size :
10 cm.
continuous
W stria e
perforations
: Western Pacific (Hainan
C o astal, p elag ic, schooling. 4°°
To 12 cm standard length, usually to about
20
°
Interest to F isheries :
P ro b a b ly m ak es a sm all
local contribution to Sardinella catches.
o°
Local N am es :
H ON G KONG:
lun) (fide Richardson, 1846:304).
T sin g lein (lin or
20°
40°
80°
100°
120°
140°
160°
180°
Ill
S a rd in ella rou xi (P o ll, 1953)
C L U P Sardl 12
H arengula rouxi Poll, 1953, R es.Exped.beige A ti.S ud., 4(2):46, fig. 16 (Cape Lopez Say, 0°42'S, 8°49’E).
Synonyms : H aren gu la m acro p h th alm a:R oux & C o llig n o n , 1950:27 (key); H aren g u la ro u x i:Fagetti & M arak,
1972:5 (eggs, larvae);
Sardinella rouxi:W hitehead, 1964:273
(not H arengula but Sardinella); CLOFETA, in press
(com plete bibliography).
FAO Nam es :
E n - Y ellow tail
sardinella.
D iagnostic Features : Body moderately deep, belly fairly shrply keeled. Lower gillrakers 30 to 40. A black
or golden spot behind gili opening; upper part o f pectoral fins dusky; caudal fin pale yellow w ith dusky hind
margin. R esem bles sm all
m aderensis, w hich has more gillrakers (70 to 166) and the caudal fin grey, its tips
alm ost black. D istinguished from small Sk aurita by having i 7 pelvic finrays (i 8 in R aurita) and no black spot on
hind part o f gili cover. See CLUP Sardl 12, Fishing Area 34.
E astern
A tla n tic
(S enegal
Habitat and Biology :
C o a sta l, p e la g ic , in in sh o re
w aters and along beaches, schooling. M ore data needed.
Size : To 16 cm stan d ard len g th , u su ally around 13 cm.
In terest
to
F ish eries:
C ontrib u tes
to
S ardinella
Literature : This species is m entioned in less than tw enty
w orks (CLOFETA, in press) and more studies are needed. Eggs
and la rv a e are d e sc rib e d by M a rc h a i (1 9 6 5 ); K rz elj (1 9 7 1 );
20°
0°
20°
40°
60°
112
C L U P Sardl 13
S ard in ella sin d en sis (D ay, 1878)
C lupea sindensis Day, 1878, Fishes o f India, pt. 4:638, pi. 163, fig. 2 (Karachi).
Synonyms : S ardinella ju ssieu :W hitehead. 1965:252 (Gulfs o f Aden and O m an); Sardinella sin d en sis:Fow ler,
1941:611 (com piled, m islead in g
B le g v a d & L o p p e n th in , 1944:65 (G u lf o f O m an); W h iteh ead & Talwar, 1971:61
(type); W h ite h e a d , 1973b:186, fig. 16 (Karachi and Bombay records only); W o n g ra ta n a , 1980:115, pis 52, 53
(revision).
N o te : the referen ces to £L sin d en sis by C han (1965:11, fig. 21) and others probably referred to Sk
gibbosa, but the two are very close.
FAO Names
E n - Sind sardinella.
D iagn ostic
F eatu res
: B ody
m oderately
slender, its depth rather variable (21 to 35% o f
standard length); total numbers o f s c u te s 31 to 34,
usually 32 or 33. Low er gillrakers 38 to 77 (more
th an 50 in fish es o v er 5 cm stan d ard len g th , in ­
c re a s in g w ith size o f fish ).
V e rtic a l stria e on
sc a le s n o t m e e tin g at c e n tre , o n ly a fe w sm all
perforations on hind part o f scale.
C losely resem ­
bles the w idespread R g ib b o sa, w hich has m ore
p erfo ratio n s on the scales. O f species o v erlapping
the range o f R sin d e n sis, the black cau d a l tips o f
R m elanura are distinctive, while R albella has fewer
sc u te s 29 to 33, but usally 30 to 32. S ee C L U P
Sard 13, Fishing Area 51.
discontinous
/
stria e \
p e rfo ra tio n s
S. sindens
S. g i b b o s a
scale
G eographical D istribution :
Indo-W est P acific (A rabian Sea,
from G u lf o f A den to the “G u lf’ and Bombay).
H abitat and B iology : C o astal, pelag ic, schooling. M ore data
needed.
in
Size
To 17 cm standard length, common to 14 cm.
Interest to F ish eries : In clu d ed in the general statistics for
Sardinella; perhaps o f some im portance in the G u lf o f O man and the
“G u lf’.
L ocal N am es : THE “GULF”: Mashineh, Moomagh (Arab G ulf B legvad & lo p p e n th in , 1944:65).
Literature : R andall, A llen & Sm ith-Vaniz (1978- the “G u lf’).
113
Sardinella taw ilis (H erre, 1927)
CLU P Sardl 23
H a re n g u la ta w ilis H e rre , 1927, P h ilip p .J .S e i., 3 4 :2 7 3 , 2 9 6 , pi. 3, fig s 1-6 (L ak e B o m b o n ,
L u zo n ).
S y n o n y m s : S ard in ella taw ilis: W ongratana, 1980:126, p ls 64 (r e v is io n ),
FAO Names
: En - F re s h w a te r
s a rd in e lla .
ÍSfh
^
h
/
L-yy yy p,
Kr
'
i t
i
i
i
h i l i i '
’ I
i
h
i
i
i
)
i
i
> i
1
> '
i
>
L
-
m '
•'
■êi®
D iagnostic Features : Body fairly slender, its depth 29 to 31% o f standard length; total scutes 28 to 30.
Lower gillrakers 61 to 74 (at 7 to 10 cm standard length). No dark markings on dorsal or caudal fins. Im mediately
separated from all other species o f Sardinella by its only known locality (Lake Bom bon, Luzon, the Philippines).
G eographical D istribution :
Bombon, Luzon).
The Philippines (Lake
H abitat and B iology :
F re s h w a te r, p re s u m a b ly
schooling.
This is the only species o f S ard in ella th at is
a p p a re n tly fo u n d o n ly , o r e v en m a in ly , in fre sh w a te r.
M ore data needed.
Size :
To 10 cm standard length.
In terest to F ish eries : O f m inor local interest.
L ocal N am es : -
h
Sardinella zunasi (B leek er, 1854)
H arengula zunasi (B leeker, 1854, N a tu u rk .T jd sc h r.N e d .-In d ië . 6:417
CLU P Sardl 24
(Nagasaki).
Synonyms : H arengula zunasi:F ow ler. 1941:597 (Japan, the K oreas); Sardinella zu n a si:Chan.
(Japan); W h iteh ead , 1973b:179 (in key only); W o n g ratan a, 1980:130, pis 7 1 ,7 2 (rev isio n ).
FAO Nam es :
E n - Ja p a n e s e s a rd in e lla .
1965:21, fig. 10
114
overlapping or
continuous striae
D iagnostic F eatures
Body m oderately slender, but variable, its
depth 24 to 33% o f standard length' total scutes 29 to 32 (usually 30 or
31). Low er gillrakers 42 to 58 (usually less than 55 in fishes up to 12 cm
standard length). V ertical striae on scales overlapping or continuous, only
a few small perforations on hind part o f scale. N o dark spot at dorsal fin
origin. M ost closely resem bles Sk richardsoni, w hich has more gillrakers (63
to 74) and has a slightly deeper body (32 to 36% o f standard length, at least
at 10 to 12 cm standard length). O f other species in the area, Sk hualiensis
also has overlapping or continuous scale striae, but black tips to d o rs a l and
caudal fins; w hile Sk fim briata a n d Sk a lb e lla h a v e th e s c a le s tria e
interrupted at the centre o f the scale.
G eographical D istribution : W estern Pacific (south­
ern coasts o f Japan south to about Taiw an Island; Chan
(1965:22) included Hong Kong, but it is suspicuous that he
had no specimens from there).
H abitat and
More data needed.
Size : To
10 cm.
B iology
13 cm
: C oastal,
pelagic,
schooling.
standard length, usually to about
Interest to Fisheries
C ontributes to local catches
of
_________
Sardinella. R eported as H arengula zunasi (10 606 tons
from the K oreas in 1983, but probably m ixed w ith other
species).
Local Names
JAPAN:
Z unasi, Zunashi.
..-y——-p—.....i»
\i
Ny
perforations
scale
,
r
z
c -
\\
Í
r.j* ? ::'
*
V .,
*
Í S * Z -'
l~ d \
"V-
h
c lic k for n e x t p a g e
c lic k fo r p r e v io u s p a g e
115
C lu p e a
1847,
L in n a e u s , 17 5 8
Clupea Linnaeus, 1758, Syst.nat., 10th ed .:3 1 7 (type:
Clupea harengus Linnaeus).
H ist.n at.p o iss., 20:340 (type: Rogenia alba Yarrell = C harengus).
C LU P C lup
R ogenia V alenciennes,
D iagnostic Features : M oderately slender fishes, w ith the belly fairly rounded and the scutes not forming a
strong keel: mainly characterized by lacking various specialized features o f other genera. O perculum smooth and
without bony radiating striae (cf. Sardinops, w hich has spots along the flanks). H ind border o f gili opening smooth
and w ithout a pair o f fleshy outgrowths (cf. Sardinella). Pelvic finrays i 8 (i 7 in Sprattus and C lupeonella, w hich
lack a bony capsule (b u lla) in the pterotic boned 8 in A lo sa , but a distinct m edian notch in upper jaw : i 8 in some
species o f Sardinella, but strong fronto-parietal striae on top o f head, also gili opening different and last two anal
finrays enlarged): pelvic fin insertion behind dorsal fin origin (below or in front in Sprattus). N o dark spots along
flanks, at dorsal fin origin, on gili cover or tips o f c a u d a l fin.
Biology, H abitat and D istribution : M arine, pelagic and schooling, down to about 200 m, mainly offshore
(but some populations entering or even confined to brackishw ater in bays or saline lakes): juveniles in shallow
w ater.Eggs dem ersal, adhesive on the sea bed or on marine vegetation. Confined to cooler or cold w aters o f the
N orthern H em isphere.
Interest to Fisheries : One o f the prime northern genera, w ith a long history o f exploitation and, in recent
years, over-exploitation in both A tlantic and Pacific waters. The total catch o f Clupea in 1983 was 1418 078 tons
(but up to tw ice that catch in earlier years before im position o f restrictions, e.g. 2 520 800 tons in 1973).
Species : A uthors have mostly followed Svetovidov (1952:120, 1963:126), w ho advocated recognition o f two
subspecies o f Clupea harengus, an A tlantic (and B altic) and a Pacific (and W hite Sea eastw ard), the latter with
few er vertebrae. The A tlantic and Pacific herrings, as Svetovidov also noted, have rather different spawning
patterns: in addition, the Pacific herring is said to have a thinner egg mem brane. N o detailed analysis has been
made o f this problem since S v e to v id o v i studies. W ere it not for the presence o f Pacific-type herrings in the
W hite Sea and eastwards, and an overlap w ith A tlantic-type herrings in the W hite Sea, it w ould be much sim pler to
recognize a distinct Pacific species w idely separated geographically from an A tlantic species. H ow ever, the
W hite Sea and associated forms apparently rem ain genetically distinct and could be regarded as a relict from the
tim e w hen a Pacific species inhabited all the A rctic w aters o f the Soviet Union. For this reason it is preferred
here to follow A m erican authors and to recognize two species:
C. harengus Linnaeus, 1758, N orth A tlantic to W hite Sea, B altic
C. pallasii V alenciennes, 1847, N orth Pacific, W hite Sea and eastward.
Remarks :
Differences between the two species can be summarized as follows:
(a)
U harengus: vertebrae 51 to 60 (usually 55 to 57): spawning throughout year (spring, sum mer, autum n
and w inter herrings): spaw n usually some distance from shore, at 25 to 200 m: spaw n usually at
relativ ely high tem p eratu res (10 to 15°C ); spaw n at high salin ities: egg masses dense, deposited on
sea bed.
(b)
C p allasii:
vertebrae 49 to 57 (usually 52 to 55): spawning in spring (i.e. spring herrings only): spawn
in coastal w aters, often close to shore, at 0 to 20 m (usually less than 15 m): spawn at relatively low
te m p e ra tu re s (0 to IO C); sp aw n at lo w s a lin itie s (10 to 21%o optim um ) egg m asses more sparse,
in one or two layers only, deposited on marine vegetation.
C lupea haren gu s L in n aeu s, 1758
CLUP Clup
1
C lupea harengus L innaeus, 1758, S yst.n at., 10th ed.:317 (northern Europe).
Synonyms : Clupea harengus B m em bras L innaeus, 1761:128 (Baltic): Cyprinus esca W albaum , 1792:36 (on
Pennant, 1769):
Clupea elongata LeSueur, 1818:234 (M assachusetts):
Clupea alba Y arrell, 1829:137, 465, pi. 5,
fig. 2 (Tham es estuary; C lupea leachii Y arrell, 1832:277, pi. 12 (T h am es, M edw ay estu arie s): C lupea m inim a
Storer, 1839:113 (N ew H a m p s h ire L C lu p e a h aren g u s - S vetovidov, 1952:117, pi. 2, figs 1, 2, pi. 3 fig. 2 (eastern
A tlan tic, B altic): Idem . 1963:123, sa m e p la te s :
F W N A , 1964:275, fig. 66 (w estern A tla n tic ): A n d riy ash ev,
1964:73 (northern seas o f USSR): Lieni & Scott, 1966:94 (w estern A tla n tic ): W h iteh ead, 1967:17 (Rogenia alba o f
V alenciennes):
C L O F N A M , 1973:99 (eastern A tlan tic, full synonym y):
F N A M , 1984:219 (e a s te rn A tla n tic ,
synopsis).
FAO Names :
E n - A tla n t ic herring .
116
D iagn ostic F eatu res :
B ody elo n g ate and fairly slen d er, belly rath er ro u n d ed , scutes w ith o u t pro m in ent
keel.
N o m edian n otch in upper ja w (cf. A lo sa ).
G ili cover (op ercu lu m ) w ith o u t
rad iatin g bony striae (cf.
Sardinops,
w hich has dark spots along the fla n k ). Hind border o f gili opening evenly rounded (w ith two fleshy
outgrowths in Sardinella). Pelvic fin insertion behind vertical from dorsal fin origin (below or in front in S p rattu s);
pelvic finrays i 8 (rarely i 7 or i 9) (only A losa and some species o f Sardinella have this count; all other clupeids
have i 6 or i 7). N o d istinctive dark spots on body or fins. O verlaps C7 pallasii in W hite Sea, but d istinguished by
more vertebrae and post-pelvic scutes (usually 55 to 57 and 12 to 16; c f. usually 52 to 55 and 10 to 14).
G eographical D istrib u tio n :
E a s te rn A tla n tic (n o rth e rn B ay o f
B is c a y n o rth w a r d to I c e la n d an d
s o u th e r n G r e e n la n d , e a s tw a r d to
S pitzbergen a n d N o v a y a Z e m ly a ,
a ls o B a ltic ) ;
w e s te r n A tla n tic
(so u th w estern G reen lan d , L ab rad o r,
southward to South Carolina).
o
H abitat and B iology :
C o a s­
tal, pelagic dow n to 200 m, school­
ing, w ith c o m p le x fe e d in g an d
sp a w n in g m ig ra tio n s , w h o se tim e s
and
e x te n t
c o rre la te
w ith
the
various m ore or less d istin ct races
which can be reco g n ized on m orpho­
logical grounds (m ainly num bers o f
v e rte b ra e , finrays, sc a le s and gillrakers). F eeds on sm all p lanktonic
copepods i n th e f i r s t y e a r , a n d
t h e r e a f t e r m a in ly copepods ( e s p e ­
c ia lly
C a lan u s fin m a rc h ic u s and
T em ora longicornis), but also hyperid a m p h ip o d s, e u p h a u s id s , m y sid sh rim p s , sm all fis h e s , arro w -w o rm s,
cten o p h o res and p tero p o d s). A t least one population is spawning in any one m onth o f the year, each race having a
different spaw ning tim e and place (spring, sum m er, autum n and w inter herrings; in 0 to 5 m o ff G reenland down
to 200 m in autum n (bank) herrings o f the N orth Sea; eggs laid on the sea bed, on rock, stones, gravel, sand or
beds o f algae or phanerogam s (see also data under genus). N o te: it is im possible to summarize briefly the wide
range o f spaw ning strateg ies o f A tla n tic herrin g ; the best rev iew s are those o f Svetovidov (1952, 1963) for the
eastern A tlantic and FW NA (1964) for the w estern Atlantic.
Size :
To 40 cm standard length, usually 20 to 25 cm.
In terest to F ish eries : A lthough stocks have been badly over-fished and depleted in recent years, Clupea
harengus still ranks as the third most heavily exploited clupeid fish.
The total catch in 1983 was 1141
736 tons,
o f which 976 041 tons came from the eastern N orth A tlantic, and 23 253 tons from the w estern N orth A tlantic.
L ocal N am es : N um erous local names have been given, not only to the species, but to all the various forms
(subspecies, races, etc., o f authors).
L iteratu re :
M ore has been p u b lish ed on C lu p ea h aren g u s th an on any o th er fish. This lite ra tu re is
sum marized by Svetovidov (1952, 1963) and FW NA (1964, w ithsom e additional references in CLO FN AM (1973), a
succinct account for Canadian A tlantic coasts by Liem & Scott (1966), and a sim ilar account for the northern seas
o f the U SSR by A ndriyashev (1964); many further and more recent papers are given by B laxter & H unter (1982).
A s many as a hundred or more papers appear annually on all aspects o f herring biology and fisheries.
117
C lu p ea p a lla sii V ale n cie n n e s,
1847
C LU P
C lup
2
C lupea pallasii V alen cien n es, 1847, H is t.n a t.p o is s ., 20:253 (Pallas specim en, K am chatka).
Synonym s : ? C lupea lineolata V alenciennes, 1847:256 ( P a lla s specim en, no locality);
G irard, 1854:138 (San Francisco); Clupea inerm is B asilew sky, 1855:242 (China); Spratelloides
1873:25 (A laska); C lupea harengus p a lla si:S vetovidov, 1952:145, pis 3, fig. 2 and 4, figs 1, 2
Idem , 1963:155, same plates;
A ndriyashev, 1964:77 (northern seas o f U SSR); Shm idt, 1965:19
Flart, 1973:96, fig. (Canada, Pacific coasts, synopsis); Clupea pallasii:Clem ens & W ilby, 1967:99,
P acific coasts; m isspelt); W h iteh ead & Bauchot, in press pallasii and lineolata types).
FAO Names :
C lupea m irabilis
bryoporus Cope,
(w estern Pacific);
(Sea o f O khotsk);
fig. 34 (Canada ,
E n - P acific herring.
D iagnostic F eatures :
B ody elo n g ate and fairly slen d er, belly rath er ro u n d ed , scutes w ith o u t pro m in en t
keel. N o m edian notch in u p p er ja w (cf. the in tro d u ced A lo sa sap id issim a o f the eastern P acific). G ili cover
(operculum ) w ithout radiating bony striae (cf. Sardinops, w hich has dark spots along the flank). Pelvic fin ra y s i 8
(as in S ardinella lem uru o f southern Japan, w hich has strong fronto-parietal striae on top o f head and a pair o f
fleshy outgrow ths on the hind border o f the gili opening). N o distinctive dark spots on body or fins. O verlaps C.
harengus in W hite Sea, but distinguished by few er vertebrae and post-pelvic scutes (usually 52 to 55 and 10 to 14;
cf. usually 55 to 57 and 12 to 16).
G eographical D istribution :
A rctic Sea (W hite Sea eastward to
Ob inlet);
w estern P acific (from
A n a d y r B a y , e a s te r n c o a s ts o f
K am chatka, possibly the A leu tian
Is la n d s so u th w a rd to Ja p a n and
w estern coasts o f K o rea); eastern
P acific (K ent P en in su la at 107°W
a n d B eaufort S e a s o u t h w a r d to
n o rth ern B aja C alifo rn ia and San
Diego).
H ab itat and B io lo g y : Coas­
ta l, p e la g ic , sc h o o lin g , m ig ra tin g
inshore to b reed, but w ith o u t any
strong n o rth -so u th m ig ratio n s, the
p o p u la tio n b ein g localized. A p p a ­
rently
landlocked
pop u latio n s
(races) ex ist in the lakes o f South
S a k h a lin , e a s te r n H o k k a id o a n d
eastern H onshu.
Feeds on euphausids, also copepods, m ysids, am phipods and zo ea o f crabs. B reed s from D ecem b er to July, d ep en d in g on the
latitude, coming into shallow w ater and depositing eggs on marine vegetation (m ainly e elg rass and seaw eeds) or
solid m aterials (see also data under genus). Spawning fishes will enter estuaries.
Size :
To 33 cm, usually to 25 cm standard length.
118
Interest to Fisheries :
O f prim e im p o rtan ce on both sides o f the P ac ific , but stocks dep leted by o v e r­
fishing in recent years. The total catch in 1983 w as 276 342 tons, o f w hich 179 721 tons came from the w estern
N orth Pacific (mainly USSR, China and Japan) and 88 643 tons from the eastern North Pacific (Canada and USA).
L o ca l N am es
:-
Literature :
Sum m arized for w estern Pacific by Svetovidov (1952, 1963) and for the Canadian Pacific by
H art (1973 - excellent synopsis o f biology and fishery, w ith over sixty selected references).
See also Ye (1980,
1983) and Ye, Tang & Qin (1981) on the Yellow Sea form and its fishery.
E scualosa W hitley, 1940
CLUP
E sc
L e p to g a s te r B le e k e r, 1872, A tla s Ich th y o l.In d .N é erla n d ., 6: pi. 26 2, fig. 2 (type: C lu p ea
(L ep to g a ste r)
arg y ro taen ia - nomen o b li tu m f id e W h it e h e a d e t ah, 1966:70). K ow ala (not o f V alenciennes: Regan, 1922,
A nn.M ag.nat.H ist., (9)10:587 (type:
K ow ala thoracata V alenciennes, designated by R egan, but overlooking earlier
designation o f ÍC albella by Gili -s e e Sardinella synonymy). Escualosa W hitley, 1940, A u st.Z o o l..9(4):402 (type:
C lupea m acrolepis Steindachner = ÍC thoracata Valenciennes).
D iagnostic F eatures : Small clupeid fishes, strongly com ­
pressed, the belly keeled. R esem bling ju v e n ile S ard in e lla , A m b ly ­
gaster and H erk lo tsich th y s, but hind m arg in o f gili o p ening evenly
rounded (cf. w ith tw o flesh y o u tg ro w th s;
also, tip o f m axilla and
second supra-m axilla en larg ed , the la tte r m ore or less rectangular.
P e lv ic finrays i 6. Anal fin short (iii to iv 14 to 18 finrays; cf. iii to
iv 31 to 49 in the pristigasterids Ilisha and P ello n a ). A b rig h t silver
stripe along flank.
Biology, H abitat and D istribution :
C o a sta l,
p e la g ic ,
schooling, m ainly shallo w w aters; ju v e n ile s found in low er parts o f
rivers.
Interest to F isheries :
C o n trib u te s to c lu p e o id
especially on the w estern coast o f India.
Species : Following the revision by W ongratana
c a tc h e s ,
2nd supra-maxilla
(1981), two species are now recognized:
R elongata W ongratana, 1983, G u lf o f Thailand
R thoracata (V alenciennes, 1847), northern Indian O cean, w estern Pacific.
Escualosa elongata W ongratana,
1983
E scualosa elongata W ongratana,
Synonym s
1983, Jap .J.Ichthyol., 2 9 (4 ):3 9 2 , fig. 8 (east co ast o f G u lf o f T hailand).
: None.
FAO N am es :
C LU P E sc 2
En - Slender w hite sardine.
119
D iagnostic F eatures :
B ody slender, its d epth 25% o f standard len g th , belly strongly keeled. S eparated
from sim ilar fishes in the area by the ch aracters g iven u n d er ÍC th o ra c a ta , from w h ich it d iffers by its m ore
slender body (cf. 27 to 37% o f standard length) and narrow er silv e r stripe along the flank (about h a lf eye diameter:
cf. about equal to eye diameter).
Geographical Distribution :
K n o w n o n ly f r o m 2
specim ens-from the east coast o f the G u lf o f T hailand
(but discovered at the Sunday M arket in Bangkok). More
specimens needed.
H ab itat and B io lo g y : P resum ed m arine and c o a s­
tal pelagic. M ore data needed.
Size :
To 6.7 cm standard length.
Interest to Fisheries :
not abundant.
No
d a ta ,
b u t p re s u m a b ly
L ocal N am es : -
E scualosa thoracata (V alenciennes, 1847)
K ow ala thoracata V alenciennes,
CLUP Esc 1
1847, H ist.nat.poiss., 20:363 (Pondicherry).
Synonyms :
? C lu pea co v al C u v ier, 1829:318 (f o o tn o te ; on Kowal o f R ussel, 1803 - nomen dubium fide
W hitehead, 1967:701: M eletta lile V alenciennes, 1847:378 (Pondicherry, C orom andel); A lausa
champil (Cantor,
1850:1284 (Penang);
R ogenia
arg ijrotaenia
B leeker,
1852:457
(M untok,
Jakarta); C lupea
(L ep to g aster)
a rg y ro ta e n ia :B leeker. 1872:pl. 262, fig. 2; C lu p ea m acro lep is S tein d ach n er, 1879:13 (T o w n sv ille, Q u een slan d ; ?
C lu p e a h u a e T ir a n t, 1929:29 (V ie t N am
K ow ala co v al:Fow ler, 1941:638 (B orneo); E scualosa thoracata W hitehead et alii, 1966:71 (argijrotaenia types); W hitehead, 1967:70, 71, 72 (coval, also thoracata and lile types);
Idem, 1973b: 189, fig. 20 (Indian Ocean, synopsis); N air, 1973:74, fig. 18 (India); W ongratana, 1980:154,pis 100,
101 (revision); W hiteh ead & Bauchot, in press (coval, lile ).
FAO Names :
En - W hite sardine.
120
D iagnostic Features : B ody fairly deep and com pressed, its depth 27 to 30% o f standard length, belly
strongly keeled.
The a lm o st re c ta n g u la r sec o n d s u p ra -m a x illa and th e b rig h t s ilv e r strip e a lo n g th e fla n k
distinguish it from juveniles o f S a rd in e lla . A m blygaster and H erklotsichthys (see also under genus). R esem bles
some pellonulines w ith a silver stripe (species o f C lupeoides, Cupleichthys), but they lack a first (anterior) supram axilla. D istinguished from E scualosa elongata o f the G u lf o f Thailand by its d e e p e r body (m ore than 25% o f
standard length) and broader silver stripe (about eaual toeye diam eter: cf. about h a lf eye diam eter). See CLU P
Esc 1, Fishing A rea 51.
G eographical D istribution :
N o rth e r n In d ia n
O cean (K arachi eastw ard to R an g o o n ), In d o n e sia (Java
Sea), the Philippines southward to Papua N ew G uinea,
A u stralia (Q ueenslan d at T ow nsville and W estern A u s­
tralia at Onslow).
H abitat and Biology : Coastal, pelagic, schooling,
in shallow w aters, the juveniles apparently entering the
low er parts o f rivers (e.g. Canning River, eastern coast
o f India), but returning later to the sea. Feeds on both
Z ooplankton (copep o d s, crab zo ea, larvae o f b ivalves
and fish eggs) and phytoplankton. B reeds from October
to February (m ainly N ovem ber to January) o ff w estern
c o a st o f In d ia , u s u a lly in re la tiv e ly sh a llo w in sh o re
waters. The eggs and larvae are well illustrated by N air
(1973).
Size
To 10 cm stan d ard len g th , com m on at
8 cm.
In terest to F ish eries :
A n im p o rtan t elem en t in clup eo id fish eries o ff w estern coasts o f In d ia (over
1000 tons in some years fide N air, 1973:table 17). Separate statistics not reported.
L ocal N am es :
L itera tu re :
INDIA: Chooda, Veloori (M alayalam ), Swadi balanjil (Kannada), Bhitgi (M arathi).
N air (1973 - excellent synopsis for India, including illustrations o f eggs and larvae).
121
P latan ich th ys W hitehead,
1968
C L U P P lat
P la tan ich th y s W h iteh ead , 1968, J.n a t.H is t., 2 :4 7 7
(type: L ile p latan a R egan).
D iagnostic Features : Small clupeid fishes o f South A m erican fresh- or brackishw aters, the body strongly
com pressed and the belly sharply keeled. P o ste rio r fro n tal fo n ta n e lle s on top o f head reta in ed in a d u lts (o cclu d ed
in R am nogaster).
A n terio r (first) su p ra-m ax illa very sm all or absent; no sharp backw ard-pointing (retrorse)
spine near front o f m ax illa (cf. R h in o sa rd in ia ). Pelvic fin ra y s i 6 or rarely i 5 (cf. i 7 in Rhinosardinia. L ile.
S tra n g o m e ra ). V ery clo se ly re se m b le s S p ra ttu s, b u t a bony cap su le (b u lla ) p re s e n t in p te ro tic and a v ery d istin c t
silver lateral stripe down flank (stripe absent in Sprattus and R am nogaster).
pre-epiotic
posterior
— frontal
fo n tan elles
fossa
p terotic b ulla
1st sup ra-m ax illa
2nd
su p ra-m axilla
1
(m uscle removed to expose
pre-epiotic fossa)
top o f head
Biology, H abitat and D istribution
from A rgentina, U ruguay and Brazil.
Interest to Fisheries :
Species :
Estuaries and low er parts o f rivers, presumably schooling; know n only
P ro b ab ly o f little im portance.
A sin g le sp ecies reco g n ized :
P latan ich th y s platan a (R eg an , 1917), w estern South A tlan tic.
P la ta n ich th y s p la ta n a (R eg an , 1917)
C LU P P lat 1
Lile platana Regan, 1917, A nn.M ag.nat.H ist.. (8)19:394 (La Plata).
Synonym s:
S pratella pallida de Buen, 1952: (N u e v a P a lm ira , m o u th o f U ru g u a y R iv e r);? S to le p h o ru s
o tid u s R in g u e le t, 1942:435 (L ag u n a C h a s c o m ú s , A rg e n tin a ); C lu p ea m elan o sto m a lim n o ica A ram b u ru , 1961:2
(Laguna C hascom ús and o th ers, A rg e n tin a );
P la ta n ic h th y s p la ta n a - W h iteh ead , 1968:479, fig. 1 (d e sc rip tio n ,
re la tio n sh ip s); F ig u eired o & M en ezes, 1978:25, fig. 29 (B razil)..
FAO Names :
E n - R iver Plate sprat.
122
D iagnostic Features : Body m oderately deep and strongly com pressed, its depth about 30% o f standard
length, belly sharply keeled. A n te rio r (first) su p ra-m ax illa m inute or absent.
C losely resem bles species o f
R am n o g a ste r, w hich lack a d istin ct silv er stripe along flank and have a w ell-d ev elo p ed first supra-m axilla.
D istin g u ish ed from species o f R h in o sard in ia by the absence o f a sharp b ack w ard -p o in tin g (retro rse) spine near
front o f maxilla and only i 5 or 6 pelvic finrays (i 7 in R hinosardinia)' from Sprattus fuegensis by the pelvic count
(cf. i 7) and presence o f silv er lateral stripe; and from small Sardinella and H arengula by the absence o f two
fleshy outgrowths on the hind margin o f the gili opening.
G eographical D istribution :
W e s te rn S o u th
A tla n tic (la g o o n s, e s tu a rie s , lo w e r p a rts o f riv e rs o f
A rg en tin a (R io de la P la ta ), U ruguay (U ruguay R iv er),
northward to ju st north o f Rio de Janeiro, Brazil).
Habitat and Biology :
P ro b a b ly c o n fin e d to
fresh- and brackishw aters o f lagoons, estuaries and the
low er reaches o f rivers. More data needed.
Size :
To 6.7 cm standard length, usually around
5 cm.
In terest to F ish eries :
co ntribution locally.
P erh ap s m akes a sm all
Local Names : Literature :
De B uen (1952 - Uruguay, as Spra­
te lla pallida); W hitehead (1964 - descrip tio n ).
c lic k for n e x t p a g e
c lic k fo r p r e v io u s p a g e
123
Ramnogaster W hitehead,
CLU P R anino
1964
R am n o g aster W h iteh ead , 1964, A n n .M a g .n a t.H ist., (1 3 )7 :3 2 4 (type: C lupea arcu ata Jen y n si.
D iagnostic Features :Small clupeid fishes o f South A m erican freshw aters, the body strongly com pressed
and the belly sharply keeled. Posterior frontal fontanelles on top o f head reduced in size and finally occluded in
fishes o f 7 or 8 cm standard length (retained in Platanichthys, see page 121). N o sharp backw ard -p o in ting
(retrorse) spine near front o f m axilla (cf. R h inosardinia). Pelvic finrays i 6 (cf. i 7 in R hinosardinia. L ile.
Strangom era). Very closely resem bles Sprattus, w hich lacks a bony capsule (bulla) in the pterotic bone. Flanks
silvery, w ithout a bright silver stripe (cf. Platanichthys).
B iology,
H abitat
and
Interest to Fisheries :
D istribution
: L ittoral
areas,
estuaries
and
rivers
from
U ruguay
to
Patagonia.
Probably o f little im portance.
Species
De B uen (1952) recognized three species.w ith som e overlap
in proportional and m eristic
ch aracters. O ne o f th ese (R. p a llid a ) is pro b ab ly P la ta n ic h th y s p la ta n a , b u t the o th er tw o seem distin ct:
R am n o g aster arcu ata (Jenyns, 1842), western South Atlantic
R am n o g aster m elanostom a (Eigenmann, 1907), western South Atlantic.
R am n ogaster arcu a ta (Jen y n s, 1842)
C L U P R anino 1
Clupea arcuata Jenyns, 1842, Z o o l.v o y .B e ag le, fishes: 134 (Bahia Blanca, Argentina).
Synonym s : C lupea arcuata R egan, 1917:228 (B ahia B lanca, M ontevideo):N orm an, 1937:39, fig. 16:
Fow ler, 1945:2, fig. (all Chile refs); Ringuelet, A rám buru & A rám buru, 1967:56 (M ar del Plata, good synonymy);
Spratella arcuata:de B uen, 1952:5 (La Palom a, Uruguay):
R am nogaster arcuata - W hitehead, 1964:324 (type,
review ).
FAO Names :
En - Jenyns’s sprat.
D iagnostic Features : Body m oderately deep and com pressed, its depth about 30% o f standard length, belly
sharply keeled. D istinguished from the very sim ilar R. melanostom a o f freshwaters by having more dorsal finrays
(18 or 19: cf. 15 or 16), more pectoral finrays (16: cf. 13 or 14) and more gillrakers (total 41 to 45: cf. 36 to 38).
R esem bles Platanichthys platana, w hich has a distinct silver stripe along the flank, and the first (anterior) supramaxilla minute or absent: separated from small Sardinella and H arengula by the evenly rounded hind margin o f
the gili opening (cf. with two fleshy outgrowths), and from Sprattus fuegensis by having i 6 pelvic finrays (cf. i 7).
124
G eograp hical
D istrib u tio n
W estern
South
A tlan tic (coasts o f South A m erica, from U ruguay and
southern Brazil (estuary o f the Lagoa dos P ato s) so u th ­
w ard to at least B ahia B lanca, A rg en tin a; som e o f the
m ore southerly reco rd s (ev en to the B eag le C hannel)
m ust refer to Sprattus fuegensis.
Habitat and Biology : Coastal, pelagic, schooling,
ap p aren tly inshore but no t en terin g fresh w ater.
M ore
data needed.
Size : To 9 cm standard length, usually about 7
to 8 cm.
Interest to Fisheries :
P erh ap s o f local im por-
ta n c e .
L ocal N am es : Literature :
De Buen (1952 - review).
R am n ogaster m ela n o sto m a (Eigenm ann, 1907)
C LU P Ram no 2
Pom olobus ? m elanostom us Eigenm ann, 1907, Proc.W ash.A cad.Sei., 8:452, pi. 23, fig. 6 (Buenos Aires).
Synonym s : Clupea m elanostom a:Regan, 1917:229 (Rio de la Plata); N orm an, 1937:39 (same); Ringuelet,
A rám b u ru & A rám b u ru , 1967:58 (Rio de la Plata, P a ra n á d e lta , go o d sy n o n y m y ); C lu p e a m e la n o sto m a lim n o ic a
A rám buru, 1961:2, pi. 1 (Buenos A ires); R inguelet et ah, 1967:59, fig. 1 B (sam e); Spratella m elanostom a:de
B uen, 1952:6 (N ueva Palm ira, m outh o f U ruguay River); R am nogaster m elanostom a - W hitehead, 1964:326
(m ention).
FAO N am es :
E n - U ru g u ay riv er sprat.
D iagnostic Features : Body moderately deep and com pressed, its depth about 30% o f standard length, belly
sharply keeled. D istinguished from the very sim ilar IE arcuata o f coastal w aters by having few er dorsal finrays
(15 o r 16; cf. 18 or 19), few er pectoral finrays (13 or 14; cf. 16) and few er gillrakers (total 36 to 38; cf. 41 to
45).
R esem bles the fresh and brackish w ater Platanichthys platana, w hich has a distinct silver stripe along the
flan k and the first (an terio r) su p ra-m ax illa m inute or absent. Separated from small Sardinella and H arengula by
the evenly rounded hind m argin o f the gili opening (cf. w ith two fleshy outgrowths).
125
Geographical D istribution :
W e s te rn S o u th
A tla n tic (U ru g u a y , A rg e n tin a , in e stu a ry an d lo w e r
reaches o f R io de la P lata and asso ciate d riv ers, e.g.
P araná and U ruguay rivers).
Habitat and Biology :
A p p aren tly confined to
freshw ater at mouths o f rivers and their low er reaches,
presum ably schooling. M ore data needed.
Size :
To
10 cm stan d ard length.
Interest to F isheries :
co n trib ution locally.
Local Names :
Literature :
P erh ap s m akes a sm all
A R G E N T IN A :
M andufia.
See under Synonyms.
Rem arks : A cco rd in g to R in g u elet, A rám b u ru &
A rám b u ru (1 96 7), tw o su b sp ecies can be reco g n ized :
(a)
R. m elanosto m a
der, its depth 27
d o rsal finrays 16
middle and low er
Rio Uruguay.
m ela n o sto m a :
body slen ­
to 31% o f standard length;
or 17; Rio de la Plata and
reaches o f Rio P a ra n á and
(b)
R. m elanostom a lim n o ic a :
body deep, its
depth 33 to 41% o f stan d ard len g th ; dorsal
finrays 13 or 14;
b rackish lagoons around
B uenos Aires.
120°
100°
80°
60°
40°
C LU P R hino
R hinosardinia Eigenmann, 1912
H erin g ia F ow ler, 1911, P ro c .A c a d .n a t.S e i.P h ila d ., 63:207 (type: C lu p ea am azo n ica S tein d ach n er; nam e p re ­
occupied in D iptera).
R h in o sa rd in ia E ig en m an n , 1912, M e m .C arn eg ie M us.. 5:445 (type:
R h in o sa rd in ia serrata
E ig en m an n = R amazonica!.
D iagnostic F eatures :
Sm all fresh w ater clu peid s o f South A m erica, the
body fairly strongly com pressed and the belly
sharply keeled.
A sharp backwardp o in tin g (retro rse) spine on upper part o f m axilla at about level o f eye centre.
T his retro rse spine is unknow n in any oth er clu p eid fish and th u s im m ediately
d istin g u ish es the genus. A n te rio r (first) su p ra-m ax illa norm al (m in u te or absent
in P latanichthys. H ind border o f gili opening evenly rounded (w ith two fleshy
outgrowths in Sardinella and H arengula). Anal fin short (about 15 to 18 finrays;
cf. 30 or more in the pristigasterids Pellona. Ilisha, etc.). Flanks
w ith or w ithout
a distinct silver stripe.
Biology, H abitat and D istribution :
F resh- and b rack ish w a ters, p erh ap s
presum ably schooling; from the Orinoco south to Salvador, Brazil.
Interest to Fisheries :
spine
to quite
high salin itie s,
P ro b ab ly o f little im portance.
Species : Three species are given in FW NA (1964:411, key), but E igenm ann’s R. serrata is undoubtedly
R.
am azonica.The status o f IE bahiensis is probably secure, but its distinction relies alm ost wholly on its higher
vertebral and scale counts (b ased on rather few specim ens); the presence o f a silver lateral stripe in R. bahiensis
but not R. amazonica, used in the key o f FWNA and by Whitehead (1973a:53), is doubtful (see under species):
R. am azo n ica (S te in d a ch n e r, 1879), w estern South and central A tlantic.
R. bahiensis (Steindachner, 1879), w estern South and central A tlantic.
126
R h in o sa rd in ia a m a zo n ica (S te in d a c h n e r, 1879)
C lupea am azonica Steindachner,
Ic h th y o l.B e itr., (8 ):65 (rep eat).
1879,
C LU P R hino
S b er.A k a d .W iss.W ie n , 80:183 (A m azo n at P ará);
1
Id e m , 1880,
Synonym s
H eringia am azonica:Fow ler, 1911:207 (S u rin am ); Regan, 1917:394 (Guyana); Rhinosardinia
serrata Eigenm ann, 1912:445,fig. 39 (m axilla), pi. 62, figs
3,4 (G uyana); FW NA, 1964:413, figs 102, 103 (maxilla);
R h in o sard in ia am az o n ica - FWNA, 1964:412 (com piled); W hitehead 1970:12 (types o f G am azonica; R. serrata a
synonym); Idem . 1973a:53, fig. 15 (Guyanas); C ervigón, 1982212, fig. 1A ( m a x illa ) (O rin o c o ).
FAO N am es :
En - A m azon spinejaw sprat.
D iagnostic Features : Body m oderately slender, but com pressed, its depth about 25 to 28% o f standard
length, belly strongly keeled. D istinguished from the very sim ilar R. bahiensis mainly by having few er vertebrae
(38 or 39; cf. 43 in R. bahiensis) and fewer scales in lateral series (39 to 41; cf. 45); other characters apparently
overlap. Easily separated from all other clupeids in the area by the sharp backw ard-pointing (re tro rse ) spine on
the upper part o f the m axilla at about the level o f the eye centre (see under genus). A silver stripe along flank in
some specimens fide C ervigón (1983:213). Two distinct parallel striae on scales.
G eographical D istribution : W estern central and
South A tlantic (low er reaches o f the O rinoco, rivers o f
the G uianas, A m azon at P ará , but undoubtedly else­
w here in its lo w er part).
H abitat and B iology : Freshw aters o f rivers, but
also tolerating
brackishw ater, presum ably
schooling.
Feeds on Zooplankton. More data needed.
Size : To 8 cm standard length, usually around
5 cm.
Interest to Fisheries
: Probably o f little im por­
tance.
L ocal N am es : Literature :
Cervigón (1983 som e notes on bio lo g y).
O rinoco
m aterial,
R em arks : C ervigón (1983:213) noted an increase
in gillrakers w ith size o f fish in his 17 Orinoco speci­
m ens, but not in all cases. M uch more material should
be studied before the species can be properly diagnosed
and its d istrib u tio n confirm ed.
120°
100°
80°
60°
40°
127
R h in o sa rd in ia b ah ien sis
(S tein d ach n er,
C LU P R hino 2
1879)
Pellonula b a h ie n s is S te in d a c h n e r, 1 8 7 9 , Sber.A kad.W iss.W ien, 80:181, pi. 3, fig. 2 (B ah ia, B razil); Idem .
1880, Ic h th y o l.B e itr., (8):63, pi. 3, fig. 2 (repeat). N o te : the figure seems based in part on Lile piquitinga.
Synonym s : Sardinella pernam bucana Schreiner & R ibeiro, 1903:72 (P ern a m b u c o ); H eringia bahiensis:
R eg an , 1917:395 (compiled); FW NA 1964:411 (key only); W hitehead, 1970:14 (type o f R bahiensis - but the chosen
le c to ty p e and p a ra le c to ty p e are L ile p iq u itin g a , n o t R h in o s a r d in ia );
C erv ig ó n , 1983:213, fig. IB (m ax illa)
(O rinoco ); W hitehead & Nelson, 1984:65 (identity o f lectotype reported, new lectotype proposed).
FAO Names :
E n - B ah ia sprat.
D iagnostic Features : Body m oderately slender, but com pressed, its depth about 25% o f standard length,
belly strongly keeled. D istinguished from the very sim ilar IE am azonica mainly by having more vertebrae (43; cf.
38 or 39 in IE am az o n ica) and m ore scales in lateral series (4 5 ; cf. 39 to 41 ); o th er c h aracters ap p aren tly
overlap. Easily separated from all other clupeids in the area by the sharp backw ard-pointing (re tro rse ) spine on
the upper part o f the m axilla at about the level o f the eye centre (see under genus). The silver stripe along the
flank, reported in FW N A (1964:411) and by W hitehead (1970, 1973a), may merely have resulted from inclusion o f
Lile piquitinga in Steindachner’s type material and his figure o f the species. Two distinct parallel striae on scales.
G eographical D istribution : W estern central and
South A tlantic (low er reaches o f Orinoco fide C erv ig ó n ,
1983; Surinam fide N elson, in l i t t . : Pernam buco, Brazil
- Schreiner & R ib e iro ’s £L pernam bucana; Bahia, Brazil
-the types).
H abitat and B iology : Freshw aters o f rivers, but
p ro b a b ly also to le ra tin g b ra c k ish w a te r.
M o re d ata
needed.
Size :
To 8 cm standard length, perhaps more.
Interest to Fisheries
: Probably
o f little
im por­
(1983
O rinoco,
single
tance.
L o ca l N am es : L iterature
specimen).
: C e rv ig ó n
-
R em arks : The high vertebral count is surprising
in v ie w o f th e a p p a re n t o v e rla p o f a lm o st all o th e r
c h a ra c te rs ;
u n fo rtu n a te ly , only the ty p es and V en e­
zu ela n and S urinam sp ecim en s have been c o u n ted , so
th at m any records o f th e species are u n co n firm ed and
its real distribution is not certain.
128
Lile Jordan & E verm ann,
C LU P
1896
L ile
Lile Jordan & E v erm a n n , 1896, B ull.U .S.natn.M us., 47(1):428, 429 (type: Clupea stolifera Jordan & Gilbert).
D iagnostic Features : Small marine fishes o f central and South A m erica, the body elongate to m oderately
deep, com pressed, belly strongly keeled.
N o b a c k w a rd -p o in tin g (retro rse) spine on upper part o f m axilla (cf.
R hinosardinia): a w ell-developed first (anterior) supra-m axilla (cf. minute or absent in Platanichthys). H ind border
o f gili opening evenly rounded (w ith tw o fleshy o u tgrow ths in S a rd in e lla . H aren g u la and O p isth o n em a). P elvic
finrays i 6 (as in R hinosardinia, but i 7 or i 8 in all other clupeid genera in this area). A very distinct silver stripe
along flank.
Biology, H abitat and D istribution : C oastal, pelagic, schooling,
w aters; w estern South and central A tlantic, eastern central Pacific.
Interest to Fisheries :
probably
entering
som ewhat b r a c k is h -
Probably o f local interest, perhaps more as a b a itfish than for food.
Species : The presence o f a distinct A tlantic and Pacific species does not appear to have been questioned,
although the tw o are extrem ely sim ilar.
R e g a n ’s L ile p latan a is P la ta n ic h th y s. T here are thus tw o species
recognized:
L. piquitinga (Schreiner & Ribeiro, 1903), w estern South and central A tlan tic
L. stolifera Jo rd an & Gilbert, 1881), eastern central P acific.
Lile piquitinga (Schreiner & R ibeiro, 1903)
CLU P L ile 1
Sardinella piquitinga Schreiner & Ribeiro, 1903, A rchos Mus.nac.Rio de J., 12:72 (Salvador, Brazil).
Synonym s : Piquitinga o f M arcgraf, 1648:159, figure (R ecife); C lu p e a a rg e n ta ta S w a in so n , 1839:386
(Pernam buco - nom en o b litu m ); S ard in ella s a r d i n a :S ta r k s . 1913:8 (L. P ap ary , n ear N a tal, B raz il); L ile p iq u itin g a
- R egan, 1917:393 (Pernam buco); C ervigón, 1966:132, fig. 56 (V enezuela); W hitehead, 1973a:48, fig. 13
(Pernam buco, key, S w a in so n ’s U arg e n ta ta d iscu ssed , also M a rc g ra f’s P iq u itin g a );
G asp ar, 1976:4 ( F i g .) , 5
(osteology).
F A O Names :
E n - A tlan tic
piquitinga.
D iagnostic Features : Body fairly slender, quite strongly com pressed, belly sharply keeled. The very bright
silver stripe along the flan k d istin g u ish es it from all sim ilar clu p eid s in the area ex cep t P latan ic h th y s p la ta n a ,
w hich has i 6 pelvic finrays (cf. i 7 in L ile) and first supra-m axilla minute or absent; and R hinosardinia species
(sharp b ackw ard-p o in tin g spine on upper p art o f m ax illa on a level w ith centre o f eye). C losely resem bles the
Pacific L. stolifera, but cau d al fin tips not black. See CLUP Lile 1, Area 31.
129
1 Ä 1 u ...
G eographical
D istrib u tion
W estern
central
A tla n tic (Nueva E s p a rta , V e n e z u e la ), w e s te rn S o u th
A tlantic (coasts o f Brazil from R ecife and Lake Papary
near N atal south to B ahia and Espirito Santo); possibly
it also occurs betw een V enezuela and Brazil.
—!
ri
s'
H abitat and Biology : Coastal, pelagic, schooling,
in s h o re and b e a c h e s;
p ro b a b ly e n te rin g so m e w h a t
b ra c k is h w a te r ( e s tu a rie s ,
lagoons
and
m angrove
swamps). M ore data needed.
t\
f
I
i
Size : To 15 cm standard length, com m on around
12 cm.
Interest to F isheries : P erhaps
local contribution to clupeoid catches.
Local Names :
pitinga.
BRAZIL:
L iterature : W hitehead
h istorical aspects).
m akes
a
small
'
y t
Cri
n
P iq u itin g a , Pitim
(1973a
-
nom enclature,
R em arks : It w as the search for the original
illu stra tio n o f M arc g ra v e’s P iq u itin g a th a t led to the
rediscovery o f the holograph score o f M ozart’s ‘Cosi fan
tu tte ’ and other m usical tre a su res lo st since the w ar
(W hitehead, 1982a).
it
Y i
120°
100°
80°
L ile sto life r a (Jo rd an & G ilb ert, 1881)
60°
40°
C LU P L ile 2
C lupea stolifera Jordan & G ilbert, 1881, Proc.U .S.natn.M us.. 4:339 (M azatlan, M exico).
Synonym s : Sardinella stolifera:Jordan & Everm an, 1896:431;
G ilbert & Starks, 1904:39 (P a n a m a B ay );
M eek & H ild eb ran d , 1923:182 (Panam a); H arengula stolifera:Everm ann & R adcliffe, 191721 (Capon, Peru); Lile
stolifera - Regan, 1917:393 (M exico); H ild eb ran d , 1946:90 (G ulf o f G uayaquil, Peru); P e te rso n , 1956:176 (G ulf of
N icoya, Costa R ica); C hirichigno, 1963:13, fig. 5 (Pta Pizzaro, Pta R ico, Peru).
FAO Names :
E n - P acific
piquitinga.
D iagnostic Features : B ody m oderately slender, quite strongly com pressed, belly sharply keeled. The
bright silverstripe along the flank distinguishes it from all sim ilar clupeids in the area. H ind border o f gili opening
evenly rounded (w ith tw o fleshy o u tg ro w th s in H aren g u la and O p isth o n e m a , the la tte r w ith a filam en to u s last
dorsal finray).
130
G eographical
Distribution
E astern
central
Pacific (G ulf o f California southward to G u lf o f G u ay a­
quil, Peru).
Habitat and Biology : Coastal pelagic, schooling,
inshore
and b e a c h e s ; p o s s ib ly e n te r s
som ew hat
brackishw aters like its A tla n tic counterpart.
Size
10 cm.
To 13 cm standard length, usually around
Interest to Fisheries : Perhaps
local contribution to clupeoid catches.
m akes
a
sm all
L ocal N am es : PERU : P elad a.
Literature : M eek & Hildebrand (1923 - Panam a);
H ildebrand (1946 - Peru).
Strangom era
W hitehead,
1964
Strangom era W hitehead,
CLU P Strang
1964, A nn.M ag.nat.H ist., (13)7:323 (type: C lupea bentincki Norman).
D iagnostic Features :
Closely resem bling the N orthern Hem isphere
herrings (C lu p e a ).
but more low er gillrakers (65 to 95; cf. 40 to 51),
gillrakers absent on posterior (inner) face o f third epibranchial (upper part
o f third arch, and ep ib ran ch ial (u p p er arch) g illrak ers o f first arch curled
upward. Pelvic f in r a y s i 7 (cf. i 6 in R am nogaster, P latanichthys; i 8 in
N ew W orld S ard in ella).
The h ig h g illra k e r count d istin g u ish e s it from
S p ra ttu s.
N o dark spots on flan k (cf. S a rd in o p s, w h ich also has bony
radiating striae on the gili cover).
Biology, H abitat and D istribution :
eastern South P acific.
Interest to Fisheries :
Species :
C o astal,
pelag ic,
gili arches
1 2 3
schooling;
A pparently supports a m oderate fishery in Chile
A single species, usually placed in Clupea in the literature:
Sk bentincki (N orm an, 1936), eastern South Pacific.
Strangom era bentincki (N orm an, 1936)
C lupea bentincki N orm an, 1936, A nn.M ag.nat.H ist., (10)17:491
no gillrakers on
hind face o f
3rd epibranchial
CLU P Strang
1
(Talcahuano, Chile).
Synonyms : C lu p ea b e n tin c k i:N orm an, 1937:38, fig. 15 (Talcahuano, Chile); F o w le r, 1945:2 (ref. only);
C lupea (A n tu) bentincki:De B uen, 1958:89, fig. 1 (subspecies c u g a , V alparaiso)
C lupea b en tin c k i:L e ib le & A lveal,
1 9 8 2 :1 4 ,fig . 1 (synopsis);
C lupea ( Strangom era) bentincki:W hitehead. 1964:323 distinction from species o f
C lu p ea, key); B ore & M artinez, 1981 :unpaged, colour photo (synopsis).
131
FAO N am es :
En - C h ilean herring.
y
3
D iagnostic Features : Body slender, rather rounded in cross-section, belly not strongly keeled. R esem bles
Sardinops sagax, but no series o f dark spots along flank and gili cover sm ooth (cf. bony radiating striae in
Sardinops)' also resem b les S prattus fu e g e n sis, but many more and finer lower gillrakers (65 to 95; cf. less than
50.
Geographical D istribution : E astern South P aci­
fic (C o q u im b o at 2 9 :3 1’S, V a lp a ra is o and so u th to
T alcahuano, C hile;
p erh ap s reach es fu rth e r south to
M ocha Island, but more southerly records may refer to
Sprattus fuegensis).
H abitat and Biology : C oastal, pelagic, schooling
at or near surface, o ccu rrin g at 0 to 70 m. F eed s on
p lankton, especially diatom s (filter-feed in g ).
B reeds
m ainly June-N ovem ber, from ab o u t 10 cm ; ap p aren tly
re le a s e s p la n k to n ic eg g s (cf. th e d e m e rsa l e g g s o f
N orthern H em isphere C lupea). M ore data needed.
Size :
To about 15 cm standard length.
Interest to F isheries : A significant fishery o ff
C hile betw een V alparaiso and T alcah u an o , m ainly at
C o q u im b o and T a lc a h u a n o (35 000 tons in 1981 fide
L eib le & A lveal, 1982). In 1983 a total o f 18 557 tons
was recorded (Chile only). A round 80% o f the catch is
used for fishmeal.
L ocal
N am es
C H ILE :
S ard in a
Sardina,
Sardina
anchoa,
S a rd in a da
(Talcahuano).
com ún;
invierno
Literature : L eible & A lv e a l (1982 - synopsis);
B ore & M arlines (1981 - synopsis).
R em arks : De Buen (1958:89) recognized a northern and a southern subspecies:
R bentincki b en tin c k i:
southward.
R bentincki c u g a :
lo w er g illrak ers 75 to 95;
eye 25 to 28.6% o f head len g th ;
Talcahuano
and
low er gillrakers 65 to 75; eye 21.7 to 26.1% o f head length; V alparaiso to Talcahuano.
The anatom ical differences from the N orthern Hem isphere Clupea and the probability that this fish does not
lay demersal adhesive eggs seem to show that it is not more closely related to Clupea than to some other clupeid
genus; however, its actual relationships have yet to be determined.
c lic k for n e x t p a g e
c lic k fo r p r e v io u s p a g e
132
2.2.3
SUBFAM ILY PELLONULINAE
FAO N am es :
En - Freshw ater herrings, double-arm oured herrings.
Diagnostic Features : Small or very sm all h e rrin g ­
like fishes (usually about 5 to 10 cm standard length, but
some larger species to 20 cm and some pygm y species
mature at only 1.8 to 2 cm standard length)! pelvic scute
normal (i.e. w ith ascending arm s), but pre- and post-pelvic
scutes reduced or absent in certain general scutes
present before dorsal fin in the A ustralian Potam othrissa
and Elyperlophus.
A nterior supra-m axilla absenti poste­
rior supra-m axilla reduced or absent in some general
m outh term inal, low er ja w projecting in some genera,
teeth usually small and conical, but
caniniform in both
jaw s in Cynothrissa and O daxothrissa.
Posterior
frontal
fontanelles (on top o f head behind level o f eyes) rem ain
open in adults. D orsal fin at about m idpoint o f body,
short (12 to 18 finrays); anal fin short (16 to 27 finrays),
its origin well behind last dorsal fin ray ; pelvic finrays
usually i 7 (but i 6 in some genera)i branchiostegal rays
usually 6 or 7 (but 2 to 4 in some genera). Scales present
in most, but highly reduced in some genera.
no 1st (anterior)
supra-m axilla
2nd (posterior)
supra-m axilla
p o s t e r io r y
frontal
fo n ta n e lle s
top 0f ileacj
B iology, H abitat and D istrib u tion
The P e llo n u lin a e are m ostly freshw ater fishes
o f tropical or
subtropical rivers and lakes, but some enter brackishw ater or school in the fully saline w aters o f bays and inlets.
Those o f W est A frica show varying degrees o f reduction o f scutes, scales, branchiostegal rays, the posterior
second supra-m axilla, and also sizei species such as Sierrathrissa leonensis mature at a size where other species
are still larvae or small juveniles, and the term progenetic can be applied to them. Pellonulines occur in w estern
and southern A frica, o ff Indian coasts, in southeast A sia and in Australia! they are absent from the N ew W orld.
Interest to F ish eries
A
Stolothrissa in Lake Tanganika).
few
species contribute significantly
to fisheries
(e.g.
L im nothrissa
and
Remarks : There are 23 genera (13 W est A frica, 3 southern A frica, 5 India and the In d o -A u stra lia n
A rchipelago, and 2 A ustralia) and about 44 species, the biggest concentration o f species being in W est A frica
(about 26). Future w ork will probably separate the ‘double-arm oured’ A ustralian genera (P o ta m o th rissa . H y p erlo ­
phus) as yet another subfamily and the W est A frican may be sim ilarly separatedfrom the rest.
Key
la .
to
A frica
2a.
the
and
G enera
M ad ag ascar,
fresh w ater
W est and central A frican rivers,
lakes
3a.
pre-pelvic scutes
P re-p elv ic
scutes
easily
visible and keeled, even if
on ly w eak ly so an te rio rly
(Fig. 1)
4a.
P r e - m a x illa e
w ith
strong
canine teeth
w ithin or set behind
no rm al
se rie s
of
sm all
teeth i
low er
ja w teeth enlarged or
canine-like; adults to
13
cm
sta n d ard
length or m ore (Fig.
canine
teeth
keeled
canine teeth
n pre-maxillae
C y n o th rissa
2)
canine teeth
on pre-maxillae
O d ax o th rissa
(upper jaw seen from below)
F i^
133
4b.
5a.
P r e - m a x illa e w ith 2 to 4 la r g e b a c k w ard pointing can in es on each side b ehind norm al
o u te r se rie s o f sm a lle r te e th ; sc u te s b e g in
before base o f first pectoral f i m a y .......................C ynothrissa
5b.
P re -m a x illa e w ith one (o r 2 ) c a n in e s a b o u t
halfway along normal outer series, curved and
pointing back; scutes begin behind base o f last
pectoral fin ray .................................................
Odaxothrissa
P re -m a x illa e w ith o u t c a n in e te e th (s o m e tim e s e n ­
larged, but not fan g -lik e); sm all fish es, rarely ex ­
ceed 12 cm standard length, usually 5 to 7 cm
6a.
6b.
maxilla blade deep
M axilla blade deep (depth 3 or less in length),
u p p e r ed g e o f m a x illa n o t fla re d o u tw a rd ;
blade o f second su p ra-m ax illa diam ond-shaped
(Fig. 3a)
P re -m a x illa e to o th e d ;
pelvic w ith i 7
fin ray s, its in s e rtio n u n d e r o r b e h in d
d o rsal fin o rig in ;
ad u lts 5 to 12 cm
standard len g th .............................................. P ellonula
7b.
P re -m a x illa e to o th le s s ; p e lv ic w ith i 6
finrays, its i n s e r ti o n u n d e r o r b e fo r e
dorsal fin origin; adults to 4 cm sta n ­
dard length ................................................. Nannothrissa
Blade
m ore
flared
often
o f m axilla long and slender (depth 3 or
tim es in len g th ), u p p er edge w ith ridge
outw ard; blade o f second su p ra-m ax illa
small, spatulate (Fig. 3b)
8a.
B lad e o f m a x illa n o rm a l, on ly a little
longer than its shaft; low er gillrakers 14
to 24; Ghana and Zaire system.
9a.
9b.
8b.
Pellonula, N annothrissa
a.
7a.
2nd
supra-maxilla
m axilla blade long
2nd
su p ra-m axilla
P o e c ilo th riss a , etc.
b.
F iS -3
L o w e r ja w n o rm a l, risin g ste e p ly
w ith in m outh, te e th at fro n t only
(Fig. 4a)
10a.
B o d y sle n d e r (d e p th 20 to
2 4 % o f s t a n d a r d length);
scales large, 27 to 34 in late­
ral s e r i e s ..........................Poecilothrissa
10b.
Body deep (depth 22 to 37%
o f s ta n d a r d le n g th ) ; s c a le s
sm all, 36 to 42 in lateral
s e r i e s .......................................Microthrissa
te e th at
front o f jaw
Lower ja w not rising steeply w ithin
m outh, te eth co n tin u ed at side o f
ja w ( F ig .4 b ) ...................................
Potam othrissa
B la d e o f m a x illa v e ry lo n g , a t le a s t
tw ic e th e le n g th o f its sh a ft;
low er
gillrakers 22 to 26 (Lake M w eru) or 31 to
42 (Lake Tanganyika)
lower jaw -deep
P o e cilo th rissa , M icro th rissa
a.
te e th at
side o f jaw
lower j aw- slender
b.
P o tam o th rissa
F is- 4
134
M a x illa b la d e n o rm a l, to o th e d lo w e r
edge not extending tow ard to tip o f p r e ­
maxilla (Fig. 5 a ) ...........................................Stolothrissa
11 b.
3b.
M a x illa b la d e lo n g , its to o th e d lo w er
edge reaching to tip o f pre-m axilla (Fig.
5b) ................................................................. Limnothrissa
P re -p e lv ic sc u te s e ith e r a b se n t, or w ith o u t
keels, or w ithout lateral arms, and difficult to
see; belly rounded (Fig. 6)
12a. P re-p elv ic scutes p resen t (m inute o r h id ­
den by scales)
13a.
2nd
supra-m axilla
tip o f
p re-m axilla
blade o f
m axilla
Low er gillrakers 10 to 16; anal fin
origin below dorsal fin base; dw arf
fish es, ad u lt at about 2 cm sta n ­
dard length
S tolothrissa
14a. Second
s u p r a - m a x illa p r e ­
sent;
g illr a k e rs 13 to 16;
p re -p e lv ic
sc u te s
w ith o u t
a rm s ...........................................Sierrathrissa
14b. N o
second
supra-m axilla;
g illra k e rs 10 to 11;
p re pelvic scutes w ith arms. . . . Thrattidion
13b.
Low er gillrakers 21 to 26; anal fin
o r i g i n b e h in d d o r s a l f i n b a s e ;
a d u l t a t a b o u t 3 cm s ta n d a rd
l e n g t h ............................................ L aeviscutella
12b. N o p re -p e lv ic or p o s t-p e lv ic sc u te s; no
second supra-maxilla; low er gillrakers 10
to l2 ; adult at about
3 c m ............ Congothrissa
no gap
L im n o th rissa
Fig-5
2b.
15b.
6a. A n a l fin e n ti r e , la s t tw o finrays n o t
. Sauvagella
se p a ra te
16b. L a s t tw o an a l finrays
rest o f tin (F ig .7)
belly rounded
pelvic scute
s e p a r a te d fro m
Spratellomorpha
pre-pelvic scutes
hidden or absent
anal fin
S pratellom orpha
separate
fmlet
135
Ib.
Indo-W est Pacific (southern Indian O cean, southeast
A sia, A ustralia)
17a. No scutes in midline before dorsal fin
18a.
18b.
B e lly ro u n d e d , no p o s t- p e lv ic s c u te s ;
low er gillrakers 24 to 30; southern India
19a.
Pre-pelvic scutes 5 to
base ju s t before dorsal
sec o n d s u p ra -m a x illa
length o f maxilla blade
19b.
Pre-pelvic scutes 1 to 4; pelvic fin
base ju s t behind dorsal fin origin;
second su p ra-m ax illa at least h a lf
length o f maxilla blade (Fig. 8b). . D a y e lla
2nd
8; pelvic fin
fin origin;
sm a ll, h a lf
(Fig. 8a) . . . E h ir a v a
maxilla blade
E h ira v a
2nd
B elly keeled , scutes p resen t before and
behind pelvic fin base; low er gillrak ers
12 to 27; southeast A sia, northern A u s­
tralia, Papua N ew Guinea
20a. A n a l fin e n ti r e , last tw o f i n r a y s
n o t s e p a ra te ; low er g illrak ers 12
to 2 4 .................................................... C lu p e o id e s
20b. L a s t tw o a n a l finrays
from rest o f fin (Fig. 7)
21a.
21b.
supra-m axilla
short
supra-m axilla
long
b. D ayella
s e p a r a te d
T eeth
in
jaws
small
or
ja w tee th
m in u te, a b s e n t a t s id e s o f
enlarged
low er jaw ;
low er g illrakers
19 to 27 .......................................... C orica
teeth at side
o f jaw
/
Teeth in jaw s enlarged, espe­
c ia lly in lo w e r ja w and on
p re -m a x illa e ;
p re s e n t on
sid es o f lo w e r ja w (F ig . 9);
low er
gillrakers
13
to
1 9 .............................................C lu p eich th y s
17b. A series o f pre-dorsal scutes present (Fig. 10);
A ustralia only
22a.
P e lv ic finrays i 7; second sup ra-m ax illa
not as deep as m ax illa blade;
8 b ra n ­
chiostegal r a y s .......................................
P o ta m a lo sa
22b.
P e lv ic finrays i 6; second sup ra-m ax illa
deeper th an m ax illa blade; 4 b ra n c h io s­
te g a l r a y s ................................................. H yp erlop h u s
C lupeichthys
Fis- 9
pre-dorsal scutes
\
P o tam a lo sa , H y p erlo p h u s
Fig. 10
136
C y n o th r is s a
C lu p
R e g a n ,1917
C y n o th rissa R eg an , 1917, A n n .M a g .n a t.H ist., (8)19:203 (type:
C yno
C y n o th rissa m ento Regan, design, by Jordan,
1920).
D iagnostic Features : A m ongst the largest o f W est A frican pellonulines,
reaching about 14 cm standard length. B elly w ith strong keel o f scutes, the
a n te r io r scute b e g in n in g b e fo re b a se o f firs t p e c to ra l finray.
L o w er ja w
prom inent, w ith strong canines at symphysis; pre-m axillae w ith 2 to 4 b a c k w a rd p o in tin g c a n in e s on e a c h sid e b e h in d n o rm a l o u te r se rie s o f te e th .
Lower
gillrakers 16 to 35, usually shorter than corresponding gili filam ents in adults.
C losely resem bles O d a x o th rissa, w hich has only one large b ack w ard -p o in tin g
canine in the upper ja w and anterior scutes beginning under or behind base o f last
p e c to r a l finray;
resem bles P e llo n u la v o rax , w hich has g illrak ers as long as
corresponding gili filam ents and enlarged but not canine-like teeth in jaw s.
can in e te eth
behind outer series
Biology, H abitat and D istribution : Freshw ater, in W est A frican rivers
from Senegal to A ngola.
P red ato ry on sm aller fish es, also p robably aquatic
insects and perhaps crustaceans.
Interest to Fisheries : E nter artisanal fisheries, but catches small.
Species
Since the synopsis by Regan (1917), two species have been consistently recognized, but there is
now evidence o f a third, rather slender species from Ivory Coast:
C. ansorgii (B oulenger, 1910), W est A frican freshwaters
C. m ento R egan, 1917, W est A frican freshwaters
C y n o th rissa species o f D aget & litis, 1965, W est A frican freshwaters.
R em arks : The presence o f fang-like or canine teeth on the pre-m axillae behind the norm al outer
series is highly unusual in clupeid fishes.
Such te eth are o th erw ise only found in the so u th east A sia n genus
C lu p e ic h th y s.
C ynothrissa ansorgii (Boulenger,
1910)
C L U P C yno
1
O d ax o th rissa an so rg ii B o u len g er, 1910, A n n .M a g .n a t.H ist., (8)6:541 (Q uanza R iv er at C am bam be, L u calla
River at Kalenge, Bengo River at Cambiri, all Angola).
Synonym s :
O daxothrissa anso rg ii:B oulenger, 1916:172, fig. 112 (A ngola, excellent figure o f a sy n ty p e);
Cynothryssa ansorgii - R egan, 1917:204 (lower Congo and A ngola); Poll, 1974:145 (synopsis); CLOFFA, 1984:42
(com plete synonymy).
FAO N am es :
En - A n so rg e ’s fan g to o th pello nu lin e.
137
D iagnostic Features : Body m oderate, often deep (depth 25% o f standard length or more). Pre-pectoral
scutes present. Low er jaw strongly projecting, w ith strong canines at sym physis; 2 to 4 strong backw ard-pointing
canines on each side behind o u ter row o f p re-m ax illary teeth.
L o w er g illrak e rs 23 to 2 6 , short (less than
corresponding gili filaments). A n a l finrays ii to iii, 14 to 16. S ilver stripe along flank. R esem bles Ç7 m ento,
w hich has few er gillrakers (16 to 19); also, Pellonula vorax, w hich lacks canines in the upper ja w and has longer
gillrakers (equal to gili filaments). Species o f O daxothrissa have only one large canine on each side o f the upper
jaw and the first scute is behind the pectoral fin base.
G eographical D istribution
(Senegal to Angola).
H abitat and B iology
needed.
Size :
: W est
A frican
freshw aters
: R ivers and stream s. M ore data
To 13 cm standard length.
Interest to
catches small.
F isheries
: E nters
artisanal
fisheries,
but
species
are
L ocal N am es : L iterature : F ourteen
given in CLOFFA (1984:42).
references
to
this
Remarks : M eristic characters probably vary through­
out the wide range o f this species, but it rem ains very clearly
separated from O m ento on gillraker numbers.
20°
C yn oth rissa
mento R egan, 1917
0°
20°
40°
60°
C LU P Cyno 2
Cynothrissa m ento R egan, 1917, A nn.M ag.nat.H ist., (8)19:204, fig. 1 (head only) (Agberi, southern Nigeria).
Synonyms : Cynothrissa m en to - Poll, 1974:146 (synopsis); CLOFFA, 1984:42 (com plete synonym y, refs to
an ato m y , fo o d , b reed in g , m ig ra tio n s, fish e rie s).
FAO N am es :
En - N ig eria n fa n g to o th pello n u lin e.
D iagnostic Features : Body m oderate, som etim es deep (depth 25% o f standard length or more). P r e ­
p ectoral scutes present.
L ow er ja w strongly p ro jectin g , w ith strong canines at sym physis;
2 to 4 strong
backw ard-pointing canines on each side behind outer row o f pre-m axillary teeth. Lower gillrakers 16 to 19, short
(less than correspondin g gili filam ents). A nal finrays ii to iii, 16 to 19. S ilv er stripe along flank. R esem bles
C ansorgii, which has more gillrakers (23 to 26); also, Pellonula vorax which lacks canines in the upper jaw and has
longer gillrakers (equal to gili filaments). Species o f O daxothrissa have only one large canine on each side o f the
upper jaw and the first scute is behind the pectoral fin base.
138
Geographical D istribution : W est A frican freshw aters
(lower parts o f Volta and N iger basins, also low er Benue and
rivers o f Cam eroon; apparently not in Zaire system).
H abitat and B iology : R ivers and stream s, also the
m an-made Lake V olta (but only in the m ost riverine parts, e.g.
Yeji and Kete K rach i). Feeds on small fishes (Pellonula, also
sm all cyprinids, cy p rin o d o n ts, schilbeids and even b a g rid catfish es), including its ow n ju v e n ile s, also on aqu atic insects.
P ossibly m igrates up stream to breed. G row th in L ake V olta
has been estimated at 7.3 mm a month.
Size :
To 13 cm standard length.
Interest to F isheries
catches are small.
L ocal N am es
: E nters
artisanal
fisheries,
but
:-
L iterature : R eynolds (1966, 1967, 1969 - fisheries;
1970 - food;
1 9 7 1 - m ig ra tio n ; 1974 - b re e d in g ); P e tr &
R ey n o ld s (1969 - fisheries); 23 referen ces to th is sp ecies in
CLOFFA (1984).
C y n o th rissa species D aget & litis, 1965
(AW AITING DESCRIPTION A N D
C LU P Cyno sp.
NAME)
Cynothrissa species D aget & litis, 1965, M ém .In s t.fr.A fr.n o ire . (74):50, fig. 27 (Lake E brié, Ivory Coast).
Synonym s :
C ynothrissa species - CLOFFA, 1984:42.
FAO N am es :
E n - Lake E brié fan g to o th p ellonuline.
D iagnostic Features : Body slender (20 to 23% o f standard length). Pre-pectoral s c u te s present. Low er
jaw strongly projecting, w ith strong canines at symphysis; 2 to 4 strong backw ard-pointing canines on each side
behind outer row o f pre-m axillary teeth. Lower gillrakers 28 to 33, long (equal to or more than corresponding gili
filam ents). S ilver stripe along flank. D istin g u ish e d from C7 m ento and C. ansorgii by having more and longer
g illrak ers (cf. only 16 to 19 and 23 to 26 resp ectiv ely , sho rter th an gili filam e n ts ). R esem bles Pellonula until
about 70 mm standard length, w hen canine teeth develop.
139
G eographical D istribution :
W est A frican fresh w aters
(Ivory Coast, in Lake E brié, but perhaps elsewhere also).
Habitat and Biology : A p p a re n tly able to to lerate sali­
nity flu ctu atio n s o f 0.6 to 2 4 .6 ° loo (Lake E brié). M o r e d a ta
needed.
Size !
To at least 10.1 cm standard length, usually 6 to
8 cm.
Interest to Fisheries :
P re s u m a b ly
fisheries, but catches w ould be small.
L o c a l N a m es
Literature
e n te rs a rtis a n a l
:
See under Synonyms.
Remarks : D a g e t & litis (1 9 6 5 ) h e sita te d to nam e this
sp e c ie s b e c a u se th e sm all and sle n d e r ty p e s o f P e llo n u la
le o n e n s is B o u le n g e r, 1916 m ig h t p e rh a p s be its ju v e n ile s .
H owever, the latter lack pre-pectoral scutes and if such scutes
are c o n s is te n tly p re s e n t in j u v e n i l e C y n o th r i s s a , th e n
R leo n en sis belo n g s in th e rath er v ariab le co m p lex th a t in clu d es
R afzeliusi and P. miri (see u n der R le o n e n s is).
Odaxothrissa Boulenger,
CLU P Odaxo
1899
O daxothrissa B oulenger, 1899, A nn.M us.C ongo, Z o o k , (1)1:64 (type: O daxothrissa lo s e ra Boulenger).
D iagnostic Features : A m ongst the largest o f the W est A frican p e llo n u lines, reaching about 15 cm standard length. B elly w ith strong keel o f scutes,
th e a n te r io r scute b eginning behind base o f last p ectoral finray.
Low er jaw
p rom inen t, w ith strong can in es at sym physis; pre-m ax illae w ith one (rarely 2)
c a n in e te e th ab o u t h a lfw a y a lo n g n o rm a l o u te r s e rie s , c u rv e d and p o in tin g
backward. Low er gillrakers 18 to 24, shorter than corresponding gili filam ents in
adults.
C losely resem bles C y n o th rissa, w h ich has 1 to 4 b ack w ard -p o in tin g
c a n in e s b e h in d n o rm al o u te r p re -m a x illa ry s e rie s, a lso p re -p e c to ra l sc u te s
present.
R esem bles P ello n u la v o rax , w h ich has g illrak ers as long as c o rre s­
ponding gili filam ents and enlarged but not canine-like teeth.
can in e te eth
w ithin outer series
Biology, H abitat and D istribution : F resh w ater, in W est A frican rivers
(C ongo system ). P red ato ry on sm aller fish es, also p robably aqu atic in sects and
perhaps crustaceans.
Interest to Fisheries
Enter artisanal fisheries, but catches small.
Species :
Since the synopsis by R egan (1917), tw o species have been co n sisten tly re co g n ized (but the
distinction not very secure - see under (A lo s e ra ):
O. lo sera B oulenger, 1899, W est A frican freshw aters
O. vittata R egan, 1917, W est A frican freshwaters.
CLU P O daxo 1
O daxothrissa losera Boulenger, 1899
O daxothrissa lo s e r a
M bandaka).
B oulenger,
1899,
A n n .M u s.C o n g o . Z ool.. (1)1:64, pi.
31, fig.
1 (C oquilhatville
=
Synonyms : O d ax o th rissa lo se ra - B oulenger, 1909:160, fig. 128 (good figure o f a syntype); Regan, 1917, 205
(ju venile syntype, Coquilhatville);
poll, 1974:146 (sy n o p sis); C L O F F A , 1984:47 (com plete synonym y, refs to
h ab itat, food).
FAO N am es :
En - L osera
fa n g to o th
p e llo n u lin e .
140
D iagnostic Features :
B ody slen d er or m o d erate (d ep th ab o u t 22 to 29% o f stan d ard len g th ). Scutes
beginning behind base o f last pectoral finray. L o w er ja w strongly p ro jectin g , w ith strong canines at sym physis;
one large canine to o th about halfw ay along norm al p re-m ax illary series (rarely a sm aller one beside or a little
behind it), curved and pointing backward. Low er gillrakers 18 to 21, short, less than corresponding gili filam ents
and the an terio r ones reduced to stum ps in large adults (m ore and lo n g er g illrak ers in ju v e n ile s). S ilver stripe
along flank. R esem bles CL v itta ta , w hich has slig h tly m ore g illrak ers (20 to 23 in adults); also Pellonula vorax,
w hich lacks canines in the upper jaw and has one or more pre-pelvic sc u te s. Species o f C ynothrissa have 1 to 4
canine teeth behind the normal outer pre-m axillary series.
Geographical Distribution :
W est A fric an fresh w aters
(low er parts o f Z aire R iver, from ab o u t M b an d ak a to B raazav ille, also the riv ers o f G abon, C ongo and p erh ap s A n g o la).
Habitat and Biology : R iv ers and stream s. M ore data
needed (but see notes by M atthes, 1964; also H ulot, 1950 and
M arlier, 1958 on food).
Size :
!
40°
20°
To 13 cm standard length.
Interest to Fisheries :
catches sm all.
Local Nam es :
L uhelele.
E n ters a rtisan al fish e ries, but
0
CONGO:
Losera;
L ake
Tumba:
20”
Literature :
(1984:47-48).
T w e n ty - th r e e
re fe re n c e s
in
CLO FFA
Remarks :
O n ly th e slig h tly lo w e r g illra k e r c o u n t
distinguishes this species
from O. v ittata, but in both species 40„
the g illrak ers appear to becom e few er and sho rter in adu lts,
the an terio r rakers d eg en eratin g into m ere stum ps.
Possibly
O. vittata is m erely an upper C ongo form (su b sp ecies ?) o f
O. losera Poll (1974:146) gave 26 to 27 gillrakers for
Q . losera: a syntype o f 7.7 cm standard length has 27 lower
gillrakers, but larger specim ens have 18 to 21 only (but 21 and 24 in two fishes o f 11 cm standard length from
C am eroon).
O daxothrissa vittata R egan,
1917
C LU P O daxo 2
O daxothrissa vittata R egan, 1917, A nn.M ag.nat.H ist.. (8)19:205 (Banzyville = Y asanyam a, Oubangui River).
Synonym s :
ecology).
O d axo th rissa v itta ta - Poll, 1974:146 (synopsis); C LO FFA , 1984:48 (com plete synonym y, ref. to
FAO N am es :
En -R e g a n ’s
fan g to o th
pellonuline.
141
D iagnostic F eatures : B ody slender or m oderate (depth about 22 to 26% o f standard length). Scutes
beginning behind base o f last pectoral finray. Lower ja w strongly projecting, w ith strong canines at sym physis;
one large canine tooth about halfway along normal pre-m axillary series (rarely a sm aller one beside or behind it),
curved and pointing backward. Low er gillrakers 20 to 24, short, less than corresponding gili filam ents and the
anterior ones reduced to stumps in large adults (more and longer gillrakers in juveniles). Silver stripe along flank.
R esem bles Q. losera, w hich has slightly few er gillrakers (18 to 21 in adults); also P e llo n u la vorax, w hich lacks
canines in the upper ja w and has one or more pre-pelvic scutes. Species o f C ynothrissa have 1 to 4 canine teeth
behind the normal outer pre-m axillary series.
G eographical D istribution
W est A frican freshw aters
(upper parts o f Zaire R iver and its tributaries, e.g. Ubanqi and
H abitat and
Size : To
Interest
: R ivers
and
stream s.
M ore data
13 cm standard length, perhaps more.
to
Literature
R em arks
B iology
F isheries
: Enters
artisanal
fisheries,
: Fifteen references in CLOFFA
: N o t certainly
distinct from
O.
but
(1984:48).
losera
(see
20 "
0°
20 °
40°
60 °
142
C LU P Pellón
P ellonula G ü n th er, 1868
Pellonula G ünther, 1868, C a t.F ish .B rit.M u s., 7:82, 452 (type: Pellonula vorax Günther).
D iagnostic Features
Small or m edium -sized W est A frican
pello n u lin es, reaching ab o u t 12 cm (u su ally 6 to 8 cm). B elly w ith
strong keel o f scutes, the anterior scute som etim es b eg in n in g before
base o f first pectoral finray. L ow er ja w slightly or rath er distin ctly
p ro je c tin g , w ith stro n g b u t n o t c a n in e te e th at sy m p h y sis;
p re ­
m axillae w ith a single (outer) row o f teeth, small or well-developed
but never can in e-lik e; m axilla blade deep, its depth about 3 tim es in
its length, second su p ra-m ax illa paddle-sh ap ed .
Low er gillrakers 20
to 36, lo n g er than co rresp o n d in g gili filam ents.
Scales sm all, in
lateral series 38 to 44. D iffers from Cynothrissa and O daxothrissa in
the lack o f canine teeth in the jaw s; d iffe rs from N a n n o th rissa and
S ierrath rissa in having i 7 p elvic finrays (cf. i 6) and the pelvic fin
insertion below or behind the dorsal fin origin (cf. in front); P o e c ilo ­
thrissa has fewer scales (27 to 34).
Biology, H abitat and D istribution : Freshw ater, in W est A fri­
can rivers from Senegal to A ngola, but in some cases enters b ra c k ish w ater o f estuaries and may even appear along beaches.
m axilla
blade
2nd
su p ra-m axilla
Interest to Fisheries : Enter artisanal fisheries, but catches small.
Species : R evisional w ork is urgently needed, but tw o species (or species com plexes) can be
distinguished, based on the form o f the pre-m axillary teeth and presence or absence o f pre-pectoral scutes:
clearly
P. leonensis R egan, 1917, W est A frican freshw aters
R vorax G ünther, 1868, W est A frican freshwaters.
CLU P P ellón 1
P ello n u la leo n en sis Boulenger, 1916
Pellonula leonensis Boulenger,
1916, C at.f.-w .Fish.A frica, 4:172, fig. I l l
(Sherbo D istrict, Sierra Leone).
Synonym s : P ellonula afzeliusi Johnels, 1954:351 (G am bia R iver); P oll, 1974:146 (G am bia, L ake Ebrié);
poll, Teugels & W hitehead, 1984:48 (com plete synonym y); M icro th rissa
m iri D aget, 1954:67, fig. 12 (D iafarabe,
14°N, upper N iger, M ali); Pellonula miri: Poll. 1974:146 (sy no p sis);
Poll, Teugels & W hitehead, 1984:49 (com plete
synonymy);
P ello n u la leo n en sis - C L O F F A , 1984:49 (com plete synonym y, S ierra L eone only; P. m iri and P
afzeliu si reco g n ized as d istin c t sp ec ie s).
FAO N am es
En - S m alltoothed
p ellonula.
143
D iagnostic Features : Body slender to m oderately deep (depth 17 to
30% o f standard length).
Scutes b eginning behind base o f first (u sually
behind last) pectoral finray, 10 to 13 + 6 to 10 = 17 to 23. Lower ja w very
s lig h tly p ro je c tin g , te e th at sy m p h y sis o n ly slig h tly e n la rg e d ;
p re m axillary teeth sm all, curved inw ard, only the tip s usu ally ap p aren t, no
strongly m arked indentation at centre o f jaw . Low er gillrakers 20 to 30,
long (equal to or lo n g er th an corresp o n d in g gili filam en ts). S ilver stripe
along flank. R esem b les R v o ra x , w hich has larg er and stra ig h te r teeth ,
often pointing forw ard , and scutes b efo re p ectoral fin bases. A b sen ce o f
la rg e c a n in e te e th in u p p e r ja w (w ith in o r b e h in d n o rm al o u te r p re m axillary series) d istin g u ish es it from O d ax o th rissa and also C y n o th rissa
(w hich has pre-pectoral scutes).
M icrothrissa species are deep-bodied and
have a much more slender maxilla. See CLUP P ellón 1, Fishing Area 34.
p re-m ax illary
te e th sm all
G eographical D istribution
W est A frican freshw aters
(Senegal to Z aire, upper reaches o f N iger and low er parts o f
Benue, but apparently not in upper parts o f Congo system).
H abitat and B iology : C hiefly rivers and stream s, also
m an-m ade and natural lakes and lag o o n s; ap p aren tly able to
tolerate m oderate or even quite high salinities (reported from
th e b e a c h e s at H a n n n e a r D a k a r and n e a r P o in te N oire,
Congo). Breeds July to Septem ber in Lake Volta.
A ccording
to D aget (1954 - as M icrothrissa m iri ) in the upper N iger the
fishes do not leave the m ain river to enter floodwater pools;
n e v e rth e le s s th e sp e c ie s a p p e a rs w e ll-e s ta b lis h e d in L ak e
Volta and to some extent in Kainji Lake (Nigeria). Feeds on
terre strial and aquatic in sects, esp ecially P o v illa nym phs in
L ake V olta, also o straco d s and en to m o stracan s;
stom achs
c o n ta in in g c lu p e id fish sc a le s m ay re p re s e n t c a n n ib a lis m .
Breeds July to September in Lake Volta.
Size !
To 8.5 cm standard length.
Interest to
catches small.
F isheries
L o ca l N am es
: E nters
artisanal
fisheries,
but
: -
L iterature : T eugels & Thys van den A vdenaerde (1979 - anatom y); all
(separately as afzeliusi, leonensis and m iri) to about 1981 are given in CLOFFA (1984).
references
to
this
species
R em arks : G reat uncertainty surrounds the nam e o f this species because the tw o slender syntypes o f R
leonensis are juveniles. Rather sim ilar slender fishes, again w ithout pre-pectoral scutes but w ith gillraker counts
up to 35 (27 to 28 in the sy n ty p es) o ccu r in Iv o ry C o ast, C am eroon, G abon and C ongo. U n fo rtu n a te ly , th ese
frequently occur w ith apparently sim ilar fishes that possess pre-pelvic scutes (but
not w ith P. vorax-type
teeth);
am ong the latter are som e w ith can in e te eth b eg in n in g to fold inw ard and also larg er adults w ith c o m p le te
C yn o th rissa-ty p e canines. The la tte r, b eing slen d er fish e s, are assum ed here to be the C y n o th rissa sp ecies o f
L ake E brié; those w ithout canines are assumed to be either Cynothrissa in w hich canines do not alw ays develop,
or else another species o f Pellonula that shares the c a ra c te rs o f both R leonensis (pre-m axillary teeth small and
curved) and o f R vorax (pre-pectoral scutes present).
Until this is resolved it is assumed that there are three subspecies o f R leonensis :
R leonensis le o n e n sis: body slen d er (d ep th 17.7 to 18% o f stan d ard len g th ); lo w er g illra k e rs 27 to 28;
syntypes o f R leonensis
from Sierra Leone, coastal.
R leonensis afzeliusi: body m oderate or deep
to 2.9; Senegal to N iger River, coastal.
(depth 21.7 to 30.2% o f standard length); low er gillrakers 21
R leonensis m iri: body m oderate or deep (depth 23.8
23; upper parts o f Niger and Benue.
to 28.5% o f standard length); low er gillrakers 20 to
144
C L U P P ellón 2
P ellonula vorax G ü n th e r, 1868
Pellonula vorax G ü n th er, 1868, C at.Fish.B rit.M us., 7:452 (“W est A frica”).
Synonym s : Pellonula stanleyana R egan, 1917:201 (Stanley Falls, but in fact Cette C arna); Pellonula
vorax :B oulenger, 1909:156, fig. 124 from Steindachner (poor) (N iger to Congo, but some specimens m isidentified);
R eg an , 1917:200 (B oulenger specimens re-identified, but R leonensis material included); Poll, 1974:146 (sy no p sis);
C LO FFA , 1984:49 (com plete synonym y, refs to breeding).
FAO N am es :
En - B igto o th ed p ellonula.
D iagnostic Features :
B ody m oderate or fairly deep (d ep th 23 to
30% o f stan dard len g th ).
Scutes beg in n in g at isth m u s, the first usually
long, 11 to 15 + 9 to 11 = 20 to 25 total. L o w er ja w slightly pro jectin g ,
teeth at sym physis large in large fish es; pre-m ax illary tee th strong, fairly
straight, pointing forward in some fishes and an indented toothless gap at
centre o f jaw . Low er gillrakers 25 to 36, long (usually equal to or longer
than corresponding gili filam en ts). S ilver stripe along flank. R esem bles R
le o n e n sis, w hich has sm all and in w ard -cu rv in g te e th and no p re-p ecto ral
sc u te s. A bsence o f large canine teeth in the upper ja w (w ithin or behind
norm al outer pre-m ax illary series) d istin g u ish es it from C ynothrissa and
also Odaxothrissa (w hich lack pre-pectoral scutes).
p re-m ax i
te e th p o in t
o u tw ard
Geographical Distribution :
W est A fric an fresh w aters
(lo w er parts o f riv ers from Ivory C o ast to A n go la);
th e
Stanley Falls record (R stanleyana') was an error.
H a b ita t and B io lo g y : R iv ers and stream s. M ore data
needed.
Size :
to 10 cm.
To 12 cm standard length, perhaps more; usually
Interest to Fisheries :
catches sm all.
E n ters artisan a l fish e ries, but
L ocal N am es : L iterature :
N u m ero u s refe re n ce s to th is species;
literature to 1981 is given in C LO FFA (1984).
the
Remarks :
In all sam ples the pro m in en t and alm ost
straig h t pre-m axillary te e th are co rrelated w ith the presence
o f p re -p e c to ra l sc u te s.
T he ra n g e o f th e sp e c ie s is no w
extended to Ivory coast, th u s b roadly ov erlap p ing th at o f the
less easily defined R leonensis.
c lic k for n e x t p a g e
c lic k fo r p r e v io u s p a g e
145
N a n n o th rissa Poll, 1965
C L U P N ann
N an n o th rissa P o ll, 1965, R ev .Z o o l.B o t.a fr ., 7 2 (3 -4 ):3 0 9 (type: M ic ro th rissa p a rv a R egan).
D iagnostic Features : D w a rf p ello n u lin es, reaching at m ost
2.2 cm in one species). Belly w ith very strong keel o f scutes, the
pectoral finray. L ow er ja w p ro m in en t, te e th ab sen t: no
teeth on
tongue. L ow er g illrak ers 20 to 31. Resem bles Pellonula, w hich has
about 4 cm standard length (m ature at only
anterior scute b eginning before base o f first
pre-m axillae, low er edge o f maxilla, vom er or
pre-m axillary teeth and i 7 pelvic finrays (cf.
i 6).
Biology, H abitat and D istribution : Freshw ater, in
W est A frican rivers and lakes (m iddle Zaire River and
associated lakes): both species are know n from acid w aters (pH 4-5).
Interest to F ish eries : Probably none.
S p ecies :
Poll & R o b erts (1 9 7 6 ) reco g n ized tw o sp ecies w hich ap p ear to be sep arab le on num bers o f
gillrakers, anal finrays and scales: how ever, preserved specimens are few:
N. parva (R egan, 1917), W est A frican freshwaters
N. stewarti Poll & R o b e rts, 1976, W est A fric a n fresh w aters.
N annothrissa parva (Regan, 1917)
C L U P N an n 1
M icro th rissa p a rv a R eg an , 1917, A n n .M a g .n a t.H is t.. (8)19:202 (Zaire River at Coquilhatville = M b a n d a k a ,
also Lake Tumba).
Synonym s :
M icro th rissa parva:M arlier. 1958:375 (L ake T um ba): M atth e s, 1964:21 (L ake T um ba, ecology):
N an n o th rissa p a rv a :Poll. 1974:150 synopsis): CLOFFA, 1984 (com plete synonym y, refs to habitat, food).
FAO N am es :
E n - L ake T um ba d w a rf sprat.
D iagnostic Features : Body slender (depth about 20% o f standard length). Scutes beginning
at isthm us (or
ju st behind), very strongly keeled (even at only 1.7 cm standard length), 13 to 15 + 6 to 8. Low er ja w projecting,
to o th less: no teeth on p re-m ax illae or m axillae: second supra-maxilla as deep as maxilla blade and w ith long
slender anterior shaft. Low er gillrakers 24 to 31. Pelvic fin w ith i 6 finrays, its insertion about h alf eye diam eter
b efo re d o rsa l fin o rig in : an al finrays 20 to 22.
Scales in lateral series 35 to
40. D itters trom FL stew arti in
having more gillrakers (cf. 20 to 23) more anal finravs (cf. 17 to 19) and more scales(cf. 34 to 35).
R e se m b le s
ju v en iles o f P ellonula v o ra x , also o f C y n o th rissa . O b a x o th rissa. M ic ro th rissa. P o ecilo th rissa and P o ta m o th rissa,
but even small individuals o f these have pre-m axillary teeth.
146
G eographical D istribution : W est A frican freshw aters
(Zaire R iver at M bandaka, Lake Tumba; Ruki and Oubangui
rivers fide P o ll, 1974:150 and the Tshuapa River at Ikela fide
M atthes, 1964).
H abitat and B iology : R ivers and lakes, apparently in
b o th ru n n in g and still w a te r, o fte n in h u g e sc h o o ls (L ak e
Tumba).
F eeds on p lan k to n (u n ice llu la r algae, diatom s and
e sp e c ia lly co p e p o d s, b u t a ls o h y d r a c a r ia n s an d a q u a tic
insects). B reeds in Lake Tumba, ripe fishes being caught in
late A ugust and ripe and some spent fishes occurring in about
m id-Septem ber (M atthes, 1964).
Size :
To 4.2 mm standard length.
Interest to F isheries : H ardly o f m uch
although occurring in som etim es dense shoals.
significance,
Local Names : ZAIRE: M em pesu, Isandja (Lake Tumba).
Literature : Fourteen references in CLOFFA (1984).
C L U P N an n 2
N a n n o th rissa ste w a r ti Poll & R oberts, 1976
N anno thrissa stew arti
S yn on ym s :
Poll & R o b erts, 1976, R e v .Z o o l.a fr.. 90(1):236 (Lake M ai-ndom be,
N a n n o th rissa stewarti - CLOFFA, 1984:47
FAO N am es :
Zaire system).
(com plete synonymy).
E n - M ai-ndom be d w a rf sp rat.
'95
D iagnostic Features : Body slender or m oderate (depth about 25% o f standard length). S cu tes a little
b ehind isthm us, very stro n g ly keeled (even at 2 .2 cm stan d ard len g th ), 9 + 7 to 9.
L ow er ja w very slightly
p rojecting, to o th less; no te eth on p re-m ax illae or m axillae; second supra-m axilla as deep as maxilla blade and
w ith long slender anterior shaft. Low er gillrakers 20 to 23. Pelvic fin w ith i 6 finrays, its insertion ju st below
dorsal fin origin; anal finrays 17 to 19. Scales in lateral series 34 or 35. D iffers from FL parva in having few er
gillrakers (cf. 24 to 31), few er anal finrays (cf. 20 to 22) and few er scales (cf. 35 to 40). Resem bles juveniles o f
P ello n u la v o rax , also o f C ynothrissa, O daxothrissa, M icrothrissa, Poecilothrissa and Potam othrissa, but even small
individuals o f these have pre-m axillary teeth.
147
G eographical D istribution : W est A frican freshw aters
(Lake M ai-ndom be or Inongo near Ipeke, Zaire system).
H abitat and B iology : Lacustrine in acid w aters (pH
M ore specimens and data needed.
S ize :
<o°
4).
To 2.25 cm standard length.
20
°
0
°
In terest to F ish e rie s : L o ca l N am es
:-
Literature : A pparently not m entioned since its original
d escrip tio n in 1976.
20
°
40°
20°
0°
20°
40°
P o ecilo th rissa Regan, 1917
Poecilothrissa R egan,
1917, A nn.M ag.nat.H ist.. (8)19:201 (type:
D iagnostic Features : Small pellonulines, reaching about 5 cm
standard length. Belly w ith fairly strong keel o f scutes, the an terio r
scute beginning under or before the base o f the last pectoral finray.
L ow er jaw only slightly or not at all prom inent, rising steeply in
m outh, w ith small teeth at front; pre-m axillae w ith a single row o f
small teeth; m axilla blade slender, its depth more than 3 tim es in its
length, its upper edge w ith a distinct flange against w hich the lower
edge o f the an to rb ital bone curves o u tw ard; second sup ra-m ax illa
sm all, spatulate, w ith a long anterior shaft. Low er gillrakers 17 to 24.
P elvic finrays i 7, its insertion a little before dorsal fin origin. Scales
large, in la te ra l se rie s 27 to 34.
R esem b les M icro th rissa and
Potam othrissa, w hich have more scales (36 to 42), also M icrothrissa is
d eep-bodied and P o tam o th rissa has alm o st no a n te rio r shaft to the
seco n d su p ra -m a x illa . P ellonula has a deeper maxillary blade and a
more diam ond-shaped second supra-maxilla.
Biology, H abitat and D istribution : Freshw ater, in W est A fri­
can rivers (m ost parts o f Zaire system) also lakes (Lake Tumba, Lake
Mweru); schooling.
60°
CLUP P o ec
Poecilothrissa congica Regan).
antorbital
flange
m axilla
blade
2nd
supra-m axilla
In terest to F ish eries : Little or none.
Species : In the review by Poll (1974), three species w ere recognized, but Poll, Teugels & W hitehead (i.e.
C LO FFA , 1984:50) added P. eupleura (L öhnberg & Rendalii, 1920) from the ‘low er C ongo’; how ever, it seems more
likely that the four known specimens are P. congica:
P. centralis Poll, 1974, W est A frican freshwaters
P. congica Regan, 1917, W est A rican freshw aters
P. m oeruensis Poll, 1948, W est A frican freshwaters.
148
CLUP P oec 1
P o ecilo th rissa cen tr a lis Poll, 1974
Poecilothrissa centralis Poli, 1974, B u ll.A c a d .r.B e lg .C I.S c i., (5)60:151, fig. 1, map 1 (Lake Tumba region and
R uki R iv e r, Z a ire sy stem ).
Synonym s : P oecilothrissa co n g ica:Regan 1917:202
cen tralis - CLOFFA, 1984:50 (co m p lete synonym y).
FAO N am es :
(one
of
the
syntypes
m isidentified); P o ecilo th rissa
En - C entral Z aire p ellonuline.
D iagnostic F eatures : Body slender (depth 20 to 24% o f standard length).
S cutes stro n g ly k eeled,
beginning under pectoral fin base, 10 + 10 or 11. Snout rather acute. Lower ja w slightly projecting, with small
te e th ; pre-m axillae with a single row o f small teeth; maxilla slender, its blade a little over 3 tim es as long as
deep, upper edge slig h tly ridged (flared o utw ard); second su p ra-m ax illa slen d er, sp atu late, w ith d istin ct an terio r
shaft. Low er gillrakers 21 to 24. D iffers from P. congica mainly in having a sharper, more acute snout and more
gillrakers (cf. only 17 to 19, rarely 20 in R co n g ica). N a n n o th rissa p a rv a from the same area has no teeth on the
p re -m a x illa e .
G eographical D istribution : W est A frican freshwaters
(restricted to the reg io n o f L ake T um ba and th e R uki R iv er,
also the Zaire River at M bandaka and its nearby tributary the
B u sira as far up as B o k u n g u on the T sh u ap a b ranch).
H abitat and B iology : R ivers and lakes, apparently in
both running and still water; like N annothrissa p a rv a , it seems
to occur in rather acid w aters. M ore data needed.
Size :
To 5.7 cm standard length.
In terest
to
F ish eries
L ocal N am es
: Probably none.
:-
L itera tu re : A p p a re n tly n o t m e n tio n ed sin ce its d e s c rip ­
tio n in 1974.
149
P o ecilo th rissa co n g ica Regan, 1917
C LU P Poec 2
P oecilothrissa congica R egan, 1917, A nn.M ag.nat.H ist., (8)19:202 (C o q u ilh atv ille = M bandaka and M onsem be,
Zaire system).
Synonym s : Poecilothrissa congica - Poll, 1974:147 (synopsis); CLOFFA, 1984:50 (com plete synonym y, refs
to habitat and anatom y). Possibly M icrothrissa eupleura L öhnberg & R endahl, 1920 is this species, in w hich case
the name should be P. eupleura.
F A O Names :
E n - B igscale p ellonuline.
D iagnostic Features : Body slender (depth about 20 to 24% o f standard length). S cu tes strongly keeled,
beginning under pectoral fin base, 11 or 12 + 9 or 10. Snout rather blunt. Low er ja w not projecting, included in
upper w hen m outh closed, w ith small teeth; pre-m ax illae w ith a single row o f sm all teeth; m ax illa slender, its
blade more than 3 times as long as deep, upper edge quite strongly ridged (flared outward); second supra-m axilla
slender, spatulate, w ith
distinct anterior shaft. Low er gillrakers 17 to
19 (rarely 20). D iffers from R centralis
and R m oeruensis in having a distinctly blunter snout, the low er jaw
not projecting; also few er gillrakers in R
centralis (cf. 21 to 24). S pecies o f P o tam o th rissa have little or no an terio r shaft to the second su p ra-m ax illa,
species o f M icrothrissa are m uch deeper-bodied, and other pellonulines have a deeper m axilla blade w ithout a
dorsal flange.
G eographical D istribution : Freshw ater, in W est A fri­
can rivers (m ost o f Z aire system , b u t ap p aren tly ab sen t from
the Lake Tumba area (R c e n tra lis) or in the L u ap u la-M w eru
sy stem (P. m oeruensis);
known from alm ost the m outh o f the
Zaire (around Boma) to the upper parts o f the Lualaba).
H a b ita t and B io lo g y :
M ore data needed.
Size :
In terest
Rivers, but perhaps not in lakes.
To about 6 cm standard length.
to
F ish eries
: Probably none.
L o ca l N am es : L itera tu re :
CLOFFA (1984).
Tw enty
references
to
the
species
in
20°
40 °
60 °
150
CLU P Poec 3
P o ecilo th rissa m oeru en sis P o ll, 1948
P o e c ilo th rissa m o eru en sis P o li,
Islan d).
Synonyms :
P o e c ilo th rissa
F A O Names :
1948, B u ll.M u s.r.H is t.n a t.B e lg ., (24):21:15, fig. 6 (Lake M w e ru o f f K ilw a
m o eru en sis - CLOFFA, 1984:50 (complete
synonymy).
En - Lake M w eru sprat.
■_? > ? v ï i
D iagnostic Features : Body slender (depth about 20 to 25% o f standard length). Scutes
strongly keeled,
beginning ju s t behind base o f
last pectoral finray, about 10 + 8. Snout rather
pointed. Low er ja w slightly
projecting, w ith small teeth;
pre-m axillae w ith a single row o f
sm all
teeth; m axilla
slender, its blade a little
more than 3 tim es longer than
deep, upper edge a little ridged (flared outw ard);
second supra-m axilla slender,
spatulate, w ith distinct anterior
shaft. L ow er gillrakers 18 to 20. Resem bles small R congica, w hich has a much
blunter snout and m ore anal
finrays (18 to 21; cf. 16 to 18). C onspecific
in Lake M w eru w ith Lim nothrissa
stappersi, which has more gillrakers (22 to 25) and no distinct keel before the pelvic fins.
G eographical D istribution
Freshw ater, in one
A frican lake and perhaps its tributaries (Lake M weru).
W est
H abitat and B iology :
L acustrine, but perhaps also in
sm all stream s or riv e rs feed in g L ak e M w eru.
M ore d a ta
needed.
20°
Size
To
about
3.5
cm
standard
length.
Interest to F ish eries
C lupeoid catches in Lake
M weru have increased since 1980 and may now yield 3 500 tons
an n u ally ; R m oeruensis m ust contribute to this.
L o ca l N am es : Literature : N ine references to the species in CLOFFA
(1984).
2o°
40°
20°
20°
40°
60°
151
M icroth rissa B o u len g er.
M icrothrissa B oulenger, 1902, A nn.M us.Congo, Z ool., (1)2:26 (type: M
D iagnostic Features :
S m all or m e d iu m -siz e d fa irly d e e p ­
b o d ie d W est A fric a n p e llo n u lin e s , re a c h in g a b o u t 10 cm in one
species.
B e lly w ith s tro n g k e e l o f sc u te s, th e a n te r io r scute (s)
beginning before base o f first pectoral finray in some. Low er ja w not
projecting, w ith small teeth at sym physis; pre-m axillae w ith a single
(outer) row o f small teeth and a few small teeth on maxilla; m axilla
blade long and slender (depth 3 or more tim es in length), upper edge
w ith a ridge flared outward; blade o f second supra-m axilla small and
spatulate. Low er gillrakers 14 to 22. Scales sm all, 36 to 42 in lateral
series. Resem bles
Poecilothrissa, w hich has few er scales (27 to 34)
and a m ore slender body (22 to 24% o f standard length), and
P otam othrissa, w hich has teeth at the sides o f the low er ja w and
alm ost no shaft to the second supra-maxilla. Other pellonulines have
a deeper maxilla blade (e.g. Pellonula) or are less trenchantly keeled.
B io lo g y , H a b ita t and D istr ib u tio n : Freshwater, in W est A fri­
can rivers (Congo/Zaire system, also Ghana).
Interest to Fisheries
CLU P M icro
1902
royauxi Boulenger).
antorbital
maxilla blade
2nd
supra-m axilla
Enter artisanal fisheries, but catches small.
Species : Poll (1974:147-148) re co g n ize d 2 species in the Congo/Zaire system, but a G hanaian species has
now been found:
M. minuta Poll, 1974, W est A frica, freshwaters
M. royauxi B oulenger, 1902, W est A frica, freshwaters
M icrothrissa Species A , W est A frica, freshwaters.
M icrothrissa m inuta P oll, 1974
CLUP
M icro
M ic ro th rissa m in u ta P o ll, 1974, B u ll.C l.S e i.A c a d .r.B e ig e , (5)60(2):154, fig. 2, m ap 2 (G aram b a R iv er and
Dungu River at G angala na Bodio, upper Zaire system).
Synonym s : N one.
F A O Names :
En - D u n g u sprat.
1
152
D iagnostic Features : Body m oderately deep (depth about 22 to 3 0 % o f standard length). Scutes strongly
keeled, 1 or 2 before base o f first pecto ral fin ray , 11 or 12 + 8. Snout p o in ted . L o w er ja w n ° t p ro jecting,
included in upper when m outh closed, w ith small teeth at sym physis; m axilla blade slender, more than 3 times as
long as deep, upper edge w ith a ridge flared outward;
second supra-m axilla small, shaft as long as blade, the
la tte r sp a tu la te . L ow er g illrak ers 19 to 21.
Total anal fin ray s 18 to 21.
Scales in lateral series 37 to 40.
R esem bles M. royauxi, w hich has few er gillrakers but more anal fin ray s (14 to 16 and 23 to 27), and M icrothrissa
sp. A w hich lacks scu tes before the pectoral fin base. Species o f Poecilothrissa have larger scales (only 27 to 34
in lateral series) and are more slender (depth 20 to 24% o f standard length); species o f Potam othrissa have teeth
at the sides o f the low er ja w and alm ost no shaft to the second supra-maxilla.
G eographical D istribution :
W est A fric a n fresh w aters
(upper Z aire system in D ungu R iver and its tributary the
Garamba River; also low er Zaire).
H a b ita t and B io lo g y
mens needed.
40°
: R iverine. M ore data and sp eci­
20°
Size :
To 3.5 cm standard length.
I n te r e s t to F is h e r ie s : L ocal N am es : Literature :
O n ly R o b e rts & S te w a rt (1 9 7 6 ) h a v e
m entioned the species since its orig in al d escrip tio n in 1976
(see CLOFFA, 1984:46).
40°
20°
M icrothrissa royauxi B oulenger,
0°
1902
F A O Names :
E n - R oyal sprat.
40°
60°
C L U P M ic ro 2
M icrothrissa royauxi B oulenger, 1902, A nn.Congo M us.. Z ool.. 2:26, pi. 8, fig. 1
Synonyms :
P e l l o n u l a (M ic ro th rissa ) ro y au x L P o ll, 1933:115;
(synopsis); CLOFFA, 1984:46 (full synonym y, distribution).
20°
(Oubangui, Zaire system).
M ic ro th ris s a ro y a u x L P o ll, 1974:147-148
153
Diagnostic Features : Body rather deep (depth about 30 to 37% o f standard length).
S cutes strongly keeled,
1 or 2 before base o f first pectoral fin ray , 13 or 14 + 6 or 7. S nout fairly pointed. L o w er ja w n o t p ro jectin g ,
included in upper w hen m outh closed, w ith small teeth at sym physis; m axilla blade slender, more than 3 tim es as
long as deep, upper edge w ith a ridge flared outw ard;second supra-m axilla sm all, shaft as long as or longer than
blade, the latter spatulate. L o w er g illrak ers 14 to 16. Total anal fin ray s 23 to 27. Scales in lateral series 36 to
40.
R e se m b le s ÁL m in u ta , w h ic h has m o re g illr a k e rs b u t fe w e r an al finrays (19 to 21 and 18 to 21), and
M ic ro th ris s a Species A ., w h ic h la c k s scu tes b e fo re th e p e c to ra l fin b ase.
S p e c ie s o f P o e c ilo th r is s a and
Potam othrissa are more slender, the latter also w ith teeth at the sides o f the low er jaw and alm ost no shaft to the
second supra-maxilla. Pellonula vorax has a deep m axilla blade (depth less than 3 tim es in length) and the second
supra-m axilla paddle-shaped.
G eographical D istribution
W est A frican freshw aters
(m ost o f Congo/Zaire system , including Oubangui R iver but
apparently not in the Kasai and its tributaries).
H a b ita t and B io lo g y : Riverine (see Gosse, 1963).
Size :
To 8 cm standard length.
Interest to
riv erin e catches.
F isheries
: A
sm all
contribution
to
local
L o ca l N am es : L iterature
T w enty-six
CLOFFA (1984:46-47).
M icrothrissa
references
to
the
species
in
Species A
(A W A IT IN G
D E S C R IP T IO N A N D
C L U P M icro 3
NAM E)
Synonym s : N one (but perhaps reported as P ello n u la sp.).
F A O Names :
E n - M u ffin sprat.
154
D iagnostic Features : Body m oderately deep (depth 29 to 32% o f standard length). Scutes m o d e ra te ly
keeled before pelvic fin base, strongly keeled behind; no scutes before pectoral fin base, 9 or 10 + 8 or 9. Snout
pointed. Low er ja w not projecting, included in upper w hen mouth closed, w ith small teeth at sym physis; maxilla
slender, a little over 3 tim es as long as deep, upper edge w ith a sharp ridge slightly flared outward; second supram axilla small, shaft about as long as blade, the latter spatulate. Lower gillrakers 21 to 23. Total anal finrays 18
to 20. Scales in lateral series 35 to 37. Resem bles M royauxi and M m inuta, but no pre-prectoral scutes (also,
only 14 to 16 gillrakers in M royauxi). Species o f P o e c ilo th riss a have larger scales (only 27 to 34 in lateral series)
and are more slender (depth 20 to 24% o f standard length); species o f Potam othrissa have teeth at the sides o f
the low er ja w and alm o st no shaft to th e second su p ra-m ax illa. Pellonula vorax and R leonensis have a deep
m axilla blade (depth less than 3 tim es in length) and the second supra-m axilla paddle-shaped.
G eographical D istribution
W est A frican freshw aters
(only known from the Birrim River near Kade, Ghana).
H abitat and
Size :
: R iverine.
B iology
M ore
data
needed.
To 7.3 cm standard length.
Interest to Fisheries :
riv erin e catches.
P robably co n trib u tes a little to
L o ca l N am es : L it e r a t u r e
R em arks : This represents a considerable extension in
the range o f M ic ro th rissa , b eing the first record o u tsid e the
C ongo/Z aire system .
P otam othrissa Regan, 1917
Potam othrissa R egan,
C LU P Potam o
1917, A nn.M ag.nat.H i st., (8)19:203 (type:
Pellonula obtusirostris B oulenger).
D iagnostic F eatures
Small pellonulines, reaching about 7 cm
(usually about 5 cm).
B e lly w ith a m o d e ra te k eel o f sc u te s b e g in n in g
behind the p ectoral fin b ase, or a strong keel begin n in g befo re it. L ow er
ja w not p ro m in en t, not risin g steeply w ith in m outh, te eth at sym physis
continued back on e ith er side o f ja w ; p re-m ax illae w ith a single row o f
m o d e ra te te e th ; m ax illa blade slen d er, to o th less, its d epth m ore th an 3
tim es in its length, its upper edge w ith a distinct flange against w hich the
low er edge o f the a n to rb ital curves o u tw ard; second su p ra-m ax illa sm all,
w ith a short or v estig ial an terio r shaft. L o w er g illrak ers 14 to 18. P elvic
finrays i 7, its insertion slightly behind origin o f dorsal fin. Scales small,
36 to 42 in lateral series. D iffers from all other W est A frican pellonulines
in having a slen d er low er ja w w ith saw -like te e th reaching back on each
sid e (s im ila r b u t m in u te te e th o c c u r in L im n o th ris s a m io d o n o f L ak e
Tanganyika, but the low er ja w is norm al, i.e. deep).
Biology, Habitat and Distribution :
Interest to Fisheries :
Species :
R
R
R
teeth at
side o f jaw
\
lower jaw
slender
Freshw ater, in W est A frican rivers (Congo/Zaire system).
L ittle or none.
In the review by Poll
(1974), 3 fairly distinctive species were reco g n ized :
acutirostris (B oulenger, 1899), W est A frican freshwaters
obtusirostris (B oulenger, 1909), W est A frican freshwaters
w hiteheadi Poll, 1974, W est A frican freshwaters.
Rem arks : The presence o f teeth at the side o f the low er jaw is highly unusual in clupeid fishes. Such teeth
also occur in the southeast A sian freshw ater genus C lu p eich th y s (but lower jaw deep). As noted above, similar but
much sm aller teeth occur in the A frican genus L im n o th rissa (jaw deep).
155
P o ta m o th rissa a cu tir o str is (B oulenger, 1899)
P e llo n u la a c u tiro s tris
M b a n d a k a , Z a ire sy ste m ).
B o u le n g e r,
1 8 9 9 , A n n ls
C LU P P otam o 1
M u s .r.C o n g o
S yn on ym s :
P e llo n u la a c u tiro s tris :B oui e n g er, 1909:159, fig .
a c u tiro s tris - R eg an , 1917:203 (u p p e r Z a ire R iv e r);
M i c r o th r is s a
(synopsis); C LO FFA , 1984:51 (com plete synonym y, ref. to habitat).
F A O Names :
B elg e, 1:63, pis 35, fig. 7 (C o q u lh atv ille =
127 (Z a ire , O u b a n g u i);
a c u tiro s tris :P o ll, 1948:21;
P o tam o th rissa
P o ll, 1974:148
En - S h arp n o sed saw to o th pellonuline.
D ia g n o stic F ea tu res : Body slender (depth 19 to 27% o f standard length).
P re-p elv ic sc u tes fairly strongly
keeled, beginning under or ju s t befo re p ecto ral fin b ase, 11 to 13 + 1 0 to 12. Snout p o in ted . L o w er ja w not
projecting, included in upper when m outh closed, deepest at m id-point o f jaw , w ith small saw-like teeth on either
side; pre-m ax illae w ith rath er peg-like teeth , those near m id -lin e p ointing forw ard; m ax illa very slen d er, its
blade over 3 tim es as long as deep, upper edge ridged (flared outw ard), low er edge w ithout denticulations; second
supra-m axilla very small, spatulate, its shaft about as long as its blade. Low er gillrakers 16 to 18. D iffers from
R obtusirostris and R w hiteheadi in its pointed snout, also more gillrakers (cf. 14 to 16 and 14 to 17). Other
pellonulines lack saw -lik e teeth at the sides o f the lower jaw.
G eographical D istribution : W est A frican freshw aters
(m ost o f Zaire system up River from Brazaaville, including the
O ubangui, Lualaba, Kasai and other tributaries).
H abitat and B iology : R ivers and lakes, apparently in
both running and still w ater; the d istrib u tio n o f the species
suggests that it tolerates a wide variety o f w ater conditions.
20
°
0
°
Size : To 7 cm standard length.
Interest to F ish eries : Probably little or none.
L o ca l N am es : L iterature
: Twenty-seven references to the species in
CLOFFA (1984).
40°
20 °
0°
20 °
40°
60 °
156
Potamothrissa
CLUP Potam o 2
o b t u s ir o s t r is (Boulenger, 1909)
Pellonula obtusirostris B oulenger, 1909, C a t.A fr.f .-w .f is h e s , 1:158 (Aruwimi River, upper Zair system).
Synonyms : P e llo n u la ten u is N ich o ls & Griscom, 1917:675, fig. 3 (Stanleyville = K isangani); S to lo th rissa
tem b essen sis F ow ler, 1949:239, figs 5-8 (Oka, Z aire system ); Potam othrissa obtusirostris - Regan, 1917:203, fig.2
(B oulenger’s types); Poli, 1974:148, map 3 (synopsis); CLOFFA, 1984:51 (com plete synonym y, refs to habitat,
food).
F A O Names :
En - B lu n tn o sed saw to o th p ello n u lin e.
D iagnostic Features : Body slender (18 to 27% o f standard length). Pre-pelvic scu tes not strongly keeled,
beginning under or behind base o f last pectoral fin ray , 7 to 10 + 10 to 12, belly rounded. Snout bluntly rounded.
L ow er ja w not p ro je ctin g , exactly m eeting tip o f upper w hen m outh clo sed , deepest at m id-point o f jaw , with
sm all saw -like teeth on each side; pre-m axillae w ith small teeth pointing downward; m axilla very slender, its
blade over 3 tim es as long as deep, upper edge ridged (flared outw ard), low er edge w ithout denticulations; second
supra-m axilla very small, spatulate, its shaft about as long as its blade. Low er gillrakers 14 to 16. Silver stripe on
flank, narrow er anteriorly. D iffers from P. whiteheadi in having a distinct shaft to the second supra-m axilla and
lacking derm al d en ticles on the snout, and from P. acutirostris in its blunt snout and few er gillrakers (cf. 16 to
18). Other pellonulines lack saw -like teeth at the sid es o f the lower jaw.
Geographical D istribution
W est A frican freshw aters
(upper parts o f Z aire system , i.e. n o rth ern and eastern trib u taries).
40°
H abitat and B iology : R ivers and stream s, perhaps not
in lakes; feeds on aquatic insects (e.g. chironomid larvae) also
h y d racarian s (M atth es, 1964:21). M ore data needed.
20°
Size :
To about 6 cm standard length.
In terest to F ish eries
: Probably little or none.
o°
L ocal N am es :
L iterature
Z A IR E : Isa n d ja .
Twenty-six
references
to
the
species
in
CLOFFA (1984).
40°
20°
20°
40°
60 °
157
C LU P P otam o 3
Potam othrissa w hiteheadi Poli, 1974
P o tam o th rissa w h iteh ead i P o li, 1974, B u ll.A c a d .r.B e lg .C l.S c i., (5)60:156, lig. 3, map 3 (Hombo R iver, eastern
part o f Zaire system).
Synonym s :
P o tam o th rissa w h iteh ea d i - CLOFFA, 1984:52.
F A O Names :
En - W h iteh e a d ’s saw to o th p ello n u lin e.
D iagnostic Features : Body slender (depth 22 to 25% o f standard length). P re-p elv ic scutes not strongly
keeled, beginning u n d er or behind base o f last pecto ral fin ray , 8 to 10 + 7 to 9, b elly rounded. S nout b luntly
rounded. Low er jaw not projecting, exactly meeting tip o f upper jaw w hen m outh closed, deepest at m id-point o f
ja w , w ith sm all saw -like te e th on each side: p re-m ax illae w ith sm all te e th p ointing dow nw ard: m axilla very
slender, its b lad e o v e r 3 tim e s as lo n g as d e e p , upper edge ridged (flared o u tw ard ), low er edge w ith o u t
denticulations: second supra-m axilla m inute, virtually w ithout anterior shaft. Small dermal denticles on snout and
bo rd er o f m outh. L o w er g illra k e rs 14 to 17. S ilver stripe along flan k , n arrow er anterio rly . C losely resem bles
R obtusirostris, w h ic h la c k s d e rm a l d e n ti c le s o n th e s n o u t a n d h a s m o re p o s t - p e lv ic s c u te s (10 to 12):
R acu tiro stris has a p ointed snout and m ore g illrak ers (16 to 18). O ther pellonulines lack saw-like teeth at the
sides o f the lower jaw.
Geographical Distribution :
W est A fric a n fresh w aters
(Flombo R iver, affluent o f Luhoho R iver, about L 30’S 28°30’E,
Zaire system).
H abitat and B iology :
Size :
40<
R iverine, but no other data.
To 4.7 cm stan d ard length.
In terest to F ish eries : P ro b ab ly none.
L ocal N am es : Literature :
tion in 1974.
A pparently not mentioned since its descrip­
c lic k for n e x t p a g e
c lic k fo r p r e v io u s p a g e
158
S to lo th rissa
CLU P Stolo
Regan, 1917
Stolothrissa Regan, 1917, A nn.M ag.nat.H ist. (8)19:206 (type:
S to lo th rissa tan g an icae R egan).
D ia g n o stic F ea tu res :
Small o r m o d e r a te p e ll o n u li n e s ,
reaching about 8.5 cm , know n only from L ake T anganyika.
Belly
rath er rounded, pre-p elv ic scutes not strongly keeled , but p o st-p elv ic
scutes w ith sharp spines. L o w er ja w very slig h tly p ro tru d in g w hen
m outh closed, risin g steeply w ith in m outh, te e th sm all: pre-m ax illae
w ith very small teeth: maxilla blade long, more than tw ice length o f
its shaft, its upper edge w ith a distinct flange: second supra-maxilla
som ew hat paddle-sh ap ed , as deep as m axilla blade. L ow er gillrak ers
long, slender and num erous. P e lv ic
fin ray s i 7, its insertion below
first third o f dorsal fin base. Scales small, 36 to 46 in lateral series.
R esem bles L im nothrissa, w hich has an even longer m axilla blade (to
tip o f p re -m a x illa ) and lo w e r p a rt o f sec o n d su p ra -m a x illa m u ch
deep er th an upper.
A ll oth er W est
A frican p ello n u lin es have the
maxilla blade only a little longer than its shaft.
B iology, H a b ita t an d D istrib u tio n :
tral A frican lake (Lake Tanganyika).
F resh w ater, in one cen
m axilla
shaft '
tip o f
pre-m axilla
maxilla blade
supra-m axilla
In terest to F ish eries :
O f c o n sid e ra b le lo c a l c o m m e rc ia l im p o rtan ce and fa r m ore ab u n d an t than
L im n o th rissa : combined catches were 65 129 tons in 1983.
S p ecies :
A single species reco g n ized :
R tanganicae
R eg an ,
1917, c en tral A fric an
fresh w aters.
S to lo th rissa ta n g a n ica e Regan, 1917
Stolothrissa tanganicae R egan, 1917, A nn.M ag.nat.H ist.
CLU P Stolo 1
, (8)19:206, fig. 2 (2) (Lake Tanganyika).
Synonym s :
P ello n u la m iodon :B oulen g er, 1906:546 (some o f his specimens, the rest Lim nothrissa m iodon):
B o u le n g e r, 1909:157 (sam e)
S to lo th ris s a ta n g a n ic a e - Poll, 1974:149, m ap 3 (s y n o p s is ): C L O F F A , 1984:54
(com plete synonym y, refs to habitat, food breeding, population biology, growth, m igrations, fisheries).
F A O Names :
En - L ake T anganyika sprat.
D iagnostic Features : Body slender (depth about 17 to 22% o f standard length). Pre-pelvic scutes not
strongly keeled, beginning behind base o f last pectoral fin ra y M axilla blade about 2 1/4 times as long as its shaft,
but not co ntinu ed forw ard to hind tip o f p re-m ax illa:
second su p ra-m ax illa d iam ond-shaped or m ore or less
rhom boidal, appro x im ately sym m etrical. L ow er g illrak ers long and slender, 36 to 42. A d istin ct silver stripe
along flanks, b ro ad est over tips o f p elvic fins.
R esem bles Lim nothrissa m iodon o f Lake Tanganyika, w hich is
slightly deeper-b o d ied , has a larg er eye (ab ou t equal to len g th o f head behind and a m axilla blade continued
forward to hind tip o f pre-maxilla.
159
G eographical
(Lake Tanganyika).
D istribution
One
central
A frican
lake
H abitat and B iology : L acustrine, form ing very large
schools, the juveniles tending to stay closer to the shore than
th o se o f 5 cm stan d ard len g th or m ore;
a p p e a r to sp en d
daylight hours below about 60 m, rising up to 8 to 15 m at
night, esp ecially on dark nights. F eeds on p lan k to n (praw ns,
shrimps, a lso c o p e p o d s, c h iro n o m id s, d ia to m s and a lg a e ).
B reeds at about 6 cm standard len g th , w ith ripe in d iv id u als
p resen t alm o st th ro u g h o u t the y ea r, but m ajo r spaw ning in
M ay/June and again in D e ce m b er/Jan u ary , the ad u lts m oving
inshore to breed. The eggs sink slowly.
Size : To about 8.5 cm standard length, usually about
7 cm.
Interest to F isheries : The more abundant o f the two
L ake T anganyika clu p eid s, but catch statistics com bined w ith
those for L im nothrissa m iodon to total 65 129 tons in 1983.
C aught w ith seine nets, often using lights.
Local Names : N dakala, D agaa (Lake Tanganyika).
L iterature
CLOFFA (1984).
F ifty -six
references
to
the
species
in
C LU P
L im nothrissa Regan, 1917
L im nothrissa Regan,
L im no
1917, A nn.M ag.nat.H ist., (8)19:207 (type: Pellonula m iodon B oulenger, in part).
D iagnostic Features : Small or fairly large pellonulines, one
species reaching about 13 cm, known only from Lake Tanganyika and
Lake M weru. B elly rath er rounded, pre-p elv ic scutes not strongly
keeled, but po st-p elv ic scutes w ith sharp spines.
L ow er ja w very
slightly pro tru d in g w hen m outh clo sed , risin g steeply w ith in m outh,
with minute teeth at sides (not saw-like, however); pre-m axillae with
very sm all teeth;
m ax illa blade very long, m ore th an four tim es
length o f its shaft, its upper edge w ith a slight ridge, its low er edged
toothed and meeting tip o f pre-m axilla; second supra-m axilla p a d d le ­
shaped, low er part larg er th an u p p er, not quite as deep as m axilla
blade.
Low er gillrakers long, slender and in one species numerous.
P e lv ic finrays i 7, its insertion below first third o f dorsal fin base.
Scales sm all, 40 or more in lateral series. Resem bles Stolothrissa o f
L ake T anganyika, w h ich has a sh o rter m ax illa blade (n o t reach in g
forward to tip o f pre-m axilla) and a more sym metrical second supram axilla. A ll other W est A frican pello n u lin es have the m ax illa blade
only a little longer than its shaft.
Biology, H abitat and D istribution
M w eru); introduced into other lakes.
: Freshw ater,
in tw o
m axilla
shaft
no gap
central A frican
maxilla
blade
lakes
(Lake
2nd
supra-m axilla
T anganyika, Lake
Interest to Fisheries : O f some local com m ercial im portance in Lake Tanganyika and w ith potential value
w hen intro d u ced into o th er lakes.
C atch es o f th is sp ecies and the m ore ab u n d an t S to lo th rissa in 1983 w ere
65 129 tons.
Sp ecies :
Long believed monotypic, but a second species was subsequently discovered in Lake Mweru:
U m iodon (B oulenger, 1906), cen tral A frican fresh w aters
U stappersii (Poll, 1948), central A frican freshw aters.
160
CLU P Lim no 1
L im n o th rissa m iodon (B oulenger, 1906)
Pellonula m iodon B oulenger, 1906, T ra n s.z o o l.S o c.L o n d ., 17(6):546, pi. 30, fig. 2 (Lake Tanganyika;
some specimens were Stolothrissa tanganicae Regan).
but
Synonym s :
P ellonula m io d o n :B oulenger, 1909:157, fig. 125 (but some specim ens R tan g an icae); L im no­
th riss a m io d o n - R eg an , 1917207, fig. 2(1) (L ak e T a n g an y ik a); P o ll, 1 9 7 4:149, m ap 2 (sy n o p sis); C LO FFA
1984:45 (com plete synonym y, refs to habits, food, breeding, grow th, behaviour, m igrations, fisheries).
F A O Names :
E n - L ake T an g an y ik a sardine.
D iagnostic Features : Body fairly slender (depth about 22 to 24% o f standard length). Pre-pelvic scutes
not strongly keeled, beginning behind base o f last pectoral fin ra y M axilla blade over 4 times as long as its shaft,
its low er toothed edge contin u ed forw ard to m eet hind tip o f p re-m ax illa; second su p ra-m ax illa asym m etrical,
low er h alf larger. Low er gillrakers long and slender, 35 to 38. A d istin ct silver stripe along flank. R esem bles
Stolothrissa tanganicae o f Lake Tanganyika, w hich is more slender, has a small eye (less than head behind eye) and
a shorter maxilla blade not reaching forward to hind tip o f pre-maxilla. D istinguished from its co ng en er in Lake
M w eru, Lim nothrissa stappersii, chiefly by having more gillrakers (cf. only 22 to 25).
G eographical D istribution :
One central A frican lake
(L ake T anganyika), but in tro du ced into several oth er lakes
(Lake K ivu, Lake K ariba, Cahora Bassa reservoir).
H abitat and B iology :
L a c u s tr in e , f o r m in g la r g e
schools. F e e d s on p la n k to n (e s p e c ia lly a ty id sh rim p s , also
copepods, prawns, etc.), but larger individuals apparently take
larval S to lo th rissa . B reed s close to the shore th ro u g h o u t the
rainy seasons, b u t w ith p e a k s in M a y /Ju n e and D e c e m b e r/
January.
Size :
To ab o u t 13 cm stan d ard length.
Interest to Fisheries : O f less im p o rtan ce th a n S to l­
o th rissa tan g an icae in L ake T ang an y ik a, alth o u g h reach in g a
slightly larger size. C atch statistic s com bine the tw o sp ecies,
w ith a total o f 65 129 tons in 1983.
In tro d u ced into Lake
K ariba, it may support a m oderate but locally valuable fishery.
Local Nam es :
(Lake Kariba).
Literature :
CLOFFA (1984).
D a g a a (L a k e T a n g a n y ik a ), K a p e n ta
S ix ty -fo u r re fe re n c e s to th e sp e c ie s in
2°°
161
L im n oth rissa
sta p p e r sii (Poll, 1948)
CLU P Lim no 2
M icrothrissa (Potam othrissa) stappersii Poll,
Lukonzolwa).
S ynonym s :
L im n o th rissa
1984:46 (com plete synonymy).
F A O Names :
1948, B u ll.M u s.r.H is t.n a t.B e lg ., (24)21:17, fig. 7 (Lake Mweru at
stap p ersii - Poll, 1974:149, map 2 (synopsis, m isspelt as sta p p e rsi):
CLO FFA ,
E n - S tap p e r’s sprat.
D iagnostic Features : K nown only from small and perhaps juvenile specim ens o f 2.8 cm total length, but
appears sim ilar to U m iodon o f Lake Tanganyika except for few er low er gillrakers (22 to 25; cf. 35 to 38), few er
anal finrays (15 to 17; cf. 17 to 18) and absence o f scutes (a juvenile character) ?
G eographical
(Lake Mweru).
D istribution
One
central
H abitat and B iology : L acustrine,
large schools. More data needed.
A frican
apparently
lake 40„
form ing
Size : Specim ens o f 2.8 cm total length (about 2.3 cm
standard length) recorded, but perhaps grows larger.
2°°
Interest to F isheries : Little or none.
0°
L o ca l N am es : L iterature
(1984).
: Six references to this species in CLOFFA
20°
R em arks
Possib ly th is is m erely the ju v e n ile o f
U m iodon, although adult anal finray counts are usually fixed
by about 2.5 cm standard length and scutes are visible.
40°
20°
20°
40°
60°
162
S ierra th rissa Thy s van den Audenaerde, 1969
S ierrath rissa Thys van den A u d en aerd e, 1.969, R e v .Z o o l.B o t.a fr ., 80(3-4):386
Thys van den Audenaerde).
C LU P Sierr
(type: S ierrath rissa leonensis
D ia g n o stic F ea tu res : Very small p e llo ­
p elvic scute
nulines, adult at 1.8 to 2 cm standard length.
Scutes p re s e n t b e fo re and b e h in d p e lv ic fin
b a s e , w ith o u t la t e r a l arm s (e x c e p t fo r th e
pelvic scute itself), but bearing a low m e m b ra ­
nous keel. Jaws about equal, w ith small teeth;
second su p ra-m ax illa sp atu late, w ith m oderate
an terio r shaft. L ow er g illrak ers less th an 20.
B ran ch io steg al rays usu ally 3. D orsal fin o ri­
gin behind midpoint o f body; pelvic fin in s e r­
arm less scutes w ith
tio n w ell before dorsal fin, w ith i 6 finrays;
m em b ran o u s keel
anal finrays 17 to 2 1 , its o rig in b elow dorsal
fin base.
S c a le s a p p a re n tly a b se n t e x c e p t n e a r ta il.
R esem bles ju v en ile s o f o th er p ello n u lin es (esp ecially
Pellonula), but other genera have i 7 pelvic fin ray s, the pelvic fin insertion below or only a little before the dorsal
fin base and 6 branchiostegal rays (except Thrattidion and Congothrissa w ith 3, but no second supra-maxilla).
B iology, H abitat and D istribution
m an-made lakes (K ainji and Volta).
In terest to F ish eries
S p ecies :
: Freshw ater, in W est A frican rivers from Senegal to Cam eroon, also
: Little or none.
A single species reco g n ized :
R leonensis Thys van den A udenaerde, 1969, W est A frican freshwaters.
Sierrathrissa leo n en sis Thys van den Audenaerde, 1969
near
C LU P Sierr 1
Sierrathrissa leonensis Thys van den A udenaerde, 1969, R ev.Z ool.B ot.afr., 80(3-4):386, figs 1,2 (W aanje ferry
Pujehun,
Sierra).
Synonyms :
P ello n u la afz e liu si:
J o h n e ls , 1954:351 (G am bia R iv er, som e o f the sm aller syntypes o f
afzeliu si);
M icro th rissa m iri:D aget, 1954:67 (Bam ako, D iafarable, upper N iger; some o f the sm aller paratypes);
P o tam o th rissa m iri:T u rn er, 1970:11 (some juveniles); Sierrathrissa leonensis - Poll, 1974:150 (synopsis); Teugels &
T h ys v an den A u d e ria e rd e , 1979:523, figs 2,4,6,9 (c o m p a riso n w ith P e llo n u la a f z e liu s i);
C L O F F A , 1984:54
(com plete synonymy); W h ite h e a d & T eu g els, 1985 (d e s c r.d is trib ., o ste o lo g y ).
FAO Names :
En - W est A frican pygm y herring.
D iagnostic Features : Small, slender fishes, easily m istaken for the juveniles o f P ellonula. Belly rounded,
pre-pelvic scutes m inute, 4 to 10, w ithout arm s, supporting a low m em branous keel, post-pelvic scutes similar, the
keel more apparent. Low er gillrakers 13 to 16. Pelvic insertion well before dorsal fin, w ith i 6 fin ray s; anal fin
beginning under dorsal fin base. Comm only occurs w ith P ellonula, w hich has i 7 pelvic finrays, normal scutes and
the pelvic fin insertion more or less below the dorsal fin origin.
163
G eographical D istrib u tion
R ivers o f W est A frica
(S enegal to C am eroon, ap p aren tly in both low er and u p p er
re a c h e s , b u t th e la tte r o n ly c o n firm e d in th e case o f th e
N iger); also the m an-m ade lakes K ainji (N ig eria) and V olta
(Ghana).
H abitat and B iology : R iverine, but also in m an-m ade
lakes, schooling in open waters and at about 2 to 8 m in Kainji
Lake; rising to about 30 cm from the surface at night in Volta
Lake. Feeds on plank to n , esp ecially cladocerans. B reeds at
about 1.8 to 2 cm standard length, in K ainji Lake in February
M a rc h and O c to b e r/N o v e m b e r,
b u t w ith rip e in d iv id u a ls
occurring throughout the year.
Size :
8^
40°
2QO
o°
To 3 cm standard length, usually 2 to 2.5 cm.
In terest to F ish eries : Little or none.
20
-
L ocal N am es : L itera tu re :
F ifte e n re fe re n c e s to th is sp e c ie s in
CLOFFA (1984);
W h ite h e a d & T eugels (1985 - synopsis o f
biol.; also anatomy).
40°
20"
T h ra ttid io n
0°
40°
60°
C LU P Thrat
R oberts, 1972
T hrattidion
20°
R oberts, 1972, B reviora, (382):2 (type:
Thrattidion
noctivagus
Roberts).
D iagnostic Features : A m ong the sm allest o f the pellonulines, not yet know n to exceed 2.14 cm standard
length. Scutes present before and behind pelvic fin base, w ith lateral arms, the pre-pelvic scutes unkeeled. Jaw s
about equal, w ith small teeth; no second supra-maxilla. Low er gillrakers few (at m ost 11). B ranchiostegal rays 4
or 5.Pelvic fin with i 6 or 7 fin ray s, its insertion before dorsal fin origin; anal fin with 23 to 25 fin ray s, its origin
below dorsal fin base. Scales present behind gili opening and on either side o f belly. R esem bles juveniles o f other
pellonulines, but all have a second supra-m axilla (except
C ongothrissa, w hich has no pre-pelvic or post-pelvic
scutes and only 3 branchiostegal rays).
Biology, H abitat and D istribution : Freshw ater, in at least one W est A frican river (S an ag a, C am eroon).
In terest to F ish eries : S p ecies :
A single species reco g n ized :
T h rattid io n
T hrattid ion
n o ctiv ag u s R o b e rts,
1972, W est A fric a n , fresh w aters.
noctivagus Roberts, 1972
CLU P
T h ra t
Thrattidion noctivagus R oberts, 1972, B reviora, (382):4, figs 1-14 (Sanaga R iver at Edea, Cameroon).
Synonyms :
FAO
Thrattidion noctivagus
N am es:
- Poll, 1974:150
E n - S anaga pygm y herring.
(synopsis);
CLOFFA, 1984:55 (com plete synonymy).
1
164
D iagnostic Features : Small and slender or m oderately deep-bodied fishes (depth 20 to 28% o f standard
length), perhaps not reaching much over 2 cm standard length. Belly rounded, pre-pelvic scutes 4 to 7, w ith
lateral arm s, but w ithout keels; pelvic scute w ith 2 or 3 lateral arm s (i.e. branched); post-pelvic scutes keeled
and w ith arms. Low er gillrakers 10 or 11. Pelvic insertion well before dorsal fin, w ith i 6or 7 finrays; anal fin
beginning under dorsal fin base. The presence o f scutes distinguishes it from Congothrissa gossei and the absence
o f a second supra-m axilla distinguishes it from all other W est or central A frican pellonulines (even at 2 cm
standard length).
G eographical D istrib u tion
O nly recorded
single W est A fric a n riv e r (S an ag a at E dea).
from
a
40°
H abitat and B iology : R iverine, collected only at dusk
and daw n in swiftly flowing w ater o f about 1 m ju st above the
hydro-electric dam at Edea. Feeds on plankton and some
terrestrial insects. N o data on breeding; R oberts (1972) 2o°
assum ed his largest specim en (2.14 cm standard length) was
m ature.
Size :
To 2.14 cm stan d ard length.
o°
In terest to F ish e r ies : L o ca l N am es : L iteratu re :
Subsequent references m erely repeat the
data o f R o b erts (1972).
20
40°
20°
0°
20°
L a e v isc u te lla P o ll, W h ite h e a d & H opson, 1965
L a e v isc u te lla P o ll, W h iteh ead & H opson, 1965, B u ll.A c a d .r.B e lg .C l.S e i., (5)51(3):279 (ty p e:
dekim pei P o ll, W h iteh ead & Hopson).
40°
60°
C L U P L aev
L aeviscutella
D ia g n o stic F ea tu res :
Sm all p e llo n u ­
lines reaching about 4 cm standard length.
p o st-p elv ic scutes
Scutes present before pelvic fin base, w ith
w ith o u t arm s
lateral arm s, but not keeled, those behind
w ithout arm s, but supporting a low m em bra­
nous keel. Low er ja w a little prom inent, both
jaw s w ith small teeth; second supra-m axilla
paddle-shaped. L ow er gillrakers 21 to 26.
B ranchiostegal rays 6. P elvic fin w ith i 7
p re-p elv ic scutes
m em branous
finrays, its insertion below about dorsal fin
not
keeled
keel
origin; anal fin origin behind dorsal fin base.
Scales present and visible on w hole body. D is­
tinguished from other pellonulines by its armed but unkeeled pre-pelvic scutes and arm less post-pelvic scutes.
165
B iology, H a b ita t and D istrib u tio n :
F resh w a ter, in W est A frica n riv ers (Iv o ry C o ast to G abon).
In terest to F ish eries : Little or none.
S p ecies :
D
A single species reco g n ized :
dekimpei Poll, W hitehead & H opson, 1965, W est A frican freshwaters.
L aeviscutella
CLUP L a e v i 1
dekim p ei Poll, Whitehead & Hopson, 1965
L a e v is c u te ll a d e k im p e i P o ll, W h ite h e a d & H opson, 1965, B u ll.A c a d .r.B e lg .C l.S e i., (5)51(3):280, figs 1-4
(G hana near Tema, N igeria near Lagos, D ahom ey in Lake N okoué, Gabon at Agile and Congo in the Loum é River).
Synonyms :
Laeviscutella dekimpei
F A O Names :
- Poll, 1974:150
(synopsis);
C LO FFA , 1984:45 (com plete synonymy).
E n - R o u n d b elly p ello n u lin e.
D iagnostic Features : Small and fairly slender fishes (depth 22 to 25% o f standard length), probably not
exceeding about 4 cm standard length. Belly rounded, pre-pelvic scutes 7 or 8, w ith lateral arm s but w ithout keels
and hidden by scales; post-pelvic scutes w ithout arms, but supporting a low mem branous keel. Low er gillrakers
21 to 26. In all other W est A frican p ello n u lin es w ith scutes, the p o st-p elv ic scutes have lateral arm s, except
Sierrathrissa (in w hich the pre-pelvic scutes also lack arms and the pelvic fin insertion is well before the dorsal fin
origin).
G eographical D istribution : Low er parts o f rivers o f
W est A frica (Ivory Coast to Congo, perhaps also to Zaire), also
lagoons (Lake E brié, Ivory Coast; Lake N okué, D ahom ey).
H abitat and B iology : R iverine, chiefly near the coast
and entering lagoons, where salinities vary w ith the tides; the
species is presum ably euryhaline, although it may escape high 2QO,
salinities. No data on feeding. R ipe fem ales o f 3.5 to 4 cm
standard len g th w ere reco rd ed in L ake Ebrié by Daget & Iltis
(1965:54).
Size : To ju s t over 4 cm standard length.
In terest to F ish eries : Little or none.
L ocal N am es : Literature : Only five references to this species found
by CLOFFA (1984), adding little to the original description.
20°
20°
40°
60°
166
C o n goth rissa Poll, 1964
C o n g o th rissa P o ll,
CLU P Congo
1964, B u ll.A c a d .r.S c i.O u tre-M e r
C l.S c i.n a t.m é d ., NS 15(2):8 (type: C ongothrissa gossei
Poll).
D ia g n o stic F ea tu res : U nique among W est A frican p ello n u lin es in lacking pre-pelvic and post-pelvic scutes.
Small fishes not exceeding about 3.5 cm standard length, with a reduced number o f lower gillrakers (10 to 12) and
bran ch io ste g al rays (3 o r 4) and th e second su p ra -m a x illa absent.
pelvic scute
3 brachiosstegal rays
no scutes
underside o f head
B iology, H a b ita t an d D istr ib u tio n :
Freshw ater, in W est A frica (Zaire system only).
In terest to F ish eries : Little or none.
S p ecies :
C.
A single sp ecies reco g n ized :
gossei P o ll, 1964, W est A fric a n fresh w aters.
C o n g o th rissa g o sse i
P oll, 1964
C o n g o th rissa
gossei
Y aekela, Zaire River.
Synonym s :
P o ll,
C LU P Congo 1
1964,
B u ll.A c a d .r.S e i.O u tre -M e r C l.S c i.n a t.m é d ., NS 15(2):8 (Y a n g a m b i, n e a r
Congothrissa gossei - Poll, 1974:151
F A O Names :
E n - S m oothbelly pellonuline.
(synopsis); Poll & Teugels, 1984:56 (com plete synonymy).
167
Diagnostic Features : Small, fairly slender fishes (depth around 22 to 24% o f standard length), probably not
exceeding 3.5 cm standard length. Belly rounded, smooth, no trace o f scutes before or behind pelvic fins (pelvic
scute its e lf present, how ev er, the tip s o f its lateral arm s bifurcated.
Jaw s ab o u t eq u al, w ith fine teeth ; no
second supra-m axilla. L o w er g illrak ers 10 to 12. Pelvic fin insertion just behind dorsal fin origin, with i 7 finrays.
Scales large, about 22 in lateral series. O ccurs w ith species o f M ic ro th rissa . P o e c ilo th rissa . P o ta m o th rissa , etc.,
but the smoothly rounded belly, absence o f spiny post-pelvic scutes and lack o f a second supra-m axilla makes it
unmistakable.
G eographical D istrib u tion
C entral part o f Zaire
system (around K isangani area and also U bangui R iver at
Bangui and upstream).
H abitat and B iology : R iverine, apparently in the main
stream .
F e e d s on sm all sh rim p s , m o s q u ito la rv a e and
chironomid larvae (Poll, 1964). M ore data needed.
Size :
To 3.5 cm standard length.
Interest to F isheries : Little or none.
L ocal N am es : L iterature : O nly
Poll & Teugels (1984).
seven
references to
this
species
in
R em arks : Poll (1964) proposed a distinct fam ily for
this species, the C ongothrissidae, and on additional anatomical
g ro u n d s T a v e rn e (1977) agreed and w as follow ed by P oll &
Teugels (1984). H ow ever, C o n g o th rissa only carries p ellonuline trends o f red u ctio n a stage fu rth e r (scu tes and second
supra-m axilla now com pletely ab sen t) and shows no characters
that differ fundam entally from those o f all other pellonulines;
even the recognition o f a separate tribe, the C ongothrissini o f
Poll (1974), seems unnecessary.
Fowler, 1935
Gilchristella
C LU P Gil
G ilch ristella F ow ler, 1935, P ro c .A c a d .n a t.S c i.P h ila d ., 87:365, fig. 4 (type: Spratelloides aestuarius Gilchrist,
1914).
D iagnostic Features : Small pellonulines o f southern A frica reaching about
6.5 cm standard length. Thin
scutes, unkeeled and barely visible, w ith slender lateral arms present before pelvic fins, none behind. Lower jaw
slightly projecting; second su p ra-m ax illa p ad dle-shaped. L o w er g illra k e rs 42 to 76 (at 4 to 6.3 cm standard
length); gillrakers small but present on posterior face o f third epibranchial. B ranchiostegal rays 6 or 7. Pelvic fin
w ith i 7 finrays, its insertion below or ju s t before dorsal fin origin; anal fin origin ju st behind last dorsal finray.
Scales present, about 40 in lateral series. D istin g u ish ed from all oth er clu p eid s in the area by the presence o f
unkeeled pre-pelvic scutes.
gili arches
1 2 3
\,/ \
/ \
, / \ , / \ ,A , ,/ \
unkeeled pre-pelvic scutes
(partly hidden by scales)
gillrakers on hind face
o f 3rd epibranchial
B io lo g y , H a b ita t a n d D is tr ib u tio n : Rivers, estuaries, lakes and lagoons o f the eastern and w estern coasts
o f southern Africa (Lake P iti, M ozam bique to Saldanha Bay and perhaps O range R iver mouth).
168
In terest to F ish eries : L ittle or none.
Sp ecies :
1971:69):
G.
A single species recognized, but a second species has been suspected (Talw ar & W hitehead,
aestu ariu s G ilc h rist,
1914, S outh A fric a n fresh - and b rack ish w aters.
G ilc h r iste lla a e stu a r iu s (G ilc h rist, 1914)
C LU P Gil 1
Spratelloides aestuarius G ilchrist, 1914, M a r.b io l.R e p .S .A fr.,
(1):55 (S w artkops River, Port Elizabeth).
S y n o n y m s:
S to le p h o ru s a e s tu a riu s :F o w ler, 1941:569 (Blue Lagoon, N atal); E hirava madagascariensis:
W hitehead, 1963:333 (only the 11 Buffalo R iver specimens); G ilchristella aestuarius - Whitehead, 1963:334, fig. 13
(St. L ucia south to K nysna estuary); Talw ar & W hitehead, 1971 (key, B uffalo R iver specim ens); W hitehead
1973:194, fig. 24 (key, synonymy, refs); CLOFFA, 1984:44 (synonym y); SFSA, in press (key, descr.).
F A O Names :
En - G ilc h rist's round herring.
D iagnostic Features : Body rather com pressed, belly rounded, w ith 6 to 9 thin unkeeled pre-pelvic scutes
(difficult to see unless scales rem oved) followed by an equally thin pelvic scute, all w ith slender ascending arms;
no post-pelvic scutes. Snout pointed, low er ja w a little prom inent, w ith a single row o f fine teeth in each jaw ;
second supra-m axilla paddle-shaped. G illrakers 39 to 45 (but to 76 in B uffalo River fishes); gillrakers small but
present on posterior face o f third epibranchial. Pelvic fin w ith i 7 finrays, under or a little before dorsal fin
origin. R esem bles S pratelloides spp or ju v en ile E trum eus and D ussum ieria, but pre-pelvic scutes present; other
clupeids in area have keeled pre-pelvic and post-pelvic scutes.
40 °
G eographical D istribution : Lake
P iti, M o z a m b iq u e
along southern A frican coast to Saldanha B ay, possibly to
m outh o f Orange R iver (but range perhaps confused by pre­
sence o f a second undescribed species - see below).
H abitat and Biology
R iv ers, lakes, estuaries and
lagoons, apparently able to tolerate brackishw ater. M ore data
needed.
Size :
To 6.5 cm stan d ard length.
20°
o°
In terest to F ish eries : L ittle or none.
L ocal N am es : 20 -
Literature : Twenty-one references in CLOFFA (1984).
R em arks : The B uffalo R iver specim ens, placed in
E hirava m adagascariensis by W hitehead (1963:333), had 40 to
56 gillrakers, w hile specim ens from other lo calities, given as 40°
G ilc h riste lla a e s tu a riu s , had only 39 to 45 g illra k e rs; la te r
w ork suggested other differences betw een the tw o (T alw ar &
W hitehead, 1971:68,69) and the gillraker count for ‘aberrant’ specimens w as found to be as high as 76. A second
species o f G ilchristella is thus probable.
169
S a u vagella B ertin , 1940
C LU P Sauv 2
S a u v a g e lla B ertin , 1940, B u ll.M u s.n a tn .H ist.N a t.P a ris , (2)12(6):300 (ty p e:
S p ratelloid es m ad ag ascarien sis
S a u v a g e , 1883 - but ex clu d in g £3. m ad ag ascarien sis bian alis B ertin , 1940, w hich is S p ratello m o rp h a b ia n a lis).
Pellonulops Sm ith, 1949 (June), A n n .M a g .n a t.H is t., (12)2:98 (type: S p ratello id es m ad ag ascarien sis S auvage, 1883);
Idem, 1949, S e a fish .S .A fric a :91 (? published after June).
Diagnostic Features : Small elongate pellonulines o f M adagascar reaching about 4.6 cm standard length.
N o pre-p elv ic scutes, b u t pelvic scute w ith slen d er vertical arm s (W -shaped in S p rate llo id es, D u ssu m ie ria,
E trum eus).
Jaw s ab o u t equal or lo w er slightly pro jectin g . L o w er g illrak ers about 20; g illrak ers p resen t on
po sterio r face o f th ird epib ran ch ial (see G ilc h riste lla ). Pelvic fin w ith i 7 fin ray s, its in sertio n a little before
dorsal fin origin; anal fin w ithout last two finrays separate (cf. S p ratello m o rp h a).
B iology, H a b ita t and D istrib u tio n : F resh w aters o f M ad ag ascar.
In te r e st to F ish e r ie s : None.
S p e c ie s :
S.
A single sp ecies reco g n ized :
m adagascariensis (Sauvage, 1883), M ad ag ascar, fresh w aters.
Sauvagella m ad agascarien sis (Sauvage, 1883)
Spratelloides
m adagascariensis
(part)
Sauvage,
1883,
Bull. Soc.philom ath.P aris, (7)7:160
excluding the form bianalis B ertin , 1940, based on 12 o f the syntypes = Spratellom orpha bianalis).
C LU P Sauv 1
(M adagascar;
Synonym s
Sauvagella m adagascariensis longianalis and
brevidorsalis
B ertin , 1 9 4 0 ;
Pellonulops
m a d a g a s c a rie n s is :S m ith , 1949a:98 (in part on South A frican specimens o f G ilchristella aestuarius, but diagnosis
mainly on Sauvage, 1883); Ehirava m adagascariensis:W hitehead, 1963:333 (type, but 11 Buffalo River (L aestuarius
included in error); Sauvagella m adagascariensis - Talw ar & W hitehead, 1971:67 (key); W hitehead, 1973:193, fig. 22
(key, synonymy, refs); CLOFFA, 1984 (synonymy); Whitehead & Bauchot, in press:types).
F A O Names :
En - M ad ag ascar round herring.
D iagnostic Features : Body elongate, a little com pressed, belly rounded, w ithout pre-pelvic scutes, but a
pelvic scute presen t w ith slen d er v ertical arm s.
G illrak ers pro b ab ly few (19 at 4 cm standard len g th in one
syntype). Pelvic fin with i 7 fin ray s, under or a little before dorsal fin origin; last two anal finrays not separate
from others (cf. S p rate llo m o rp h a b ia n a lis). D istin g u ish ed from o th er no n -scu ted clu p eid s in area ( S p ra te llo id es,
D ussum ieria, E trum eus) by presence o f normal pelvic scute (cf. W -shaped).
170
G eo g ra p h ica l D istr ib u tio n : F re sh w ate rs o f M adagascar.
H abitat and B iology
Freshw aters, possibly
b rack ish w ater. M ore specim ens and data needed.
tolerating
Size : To ab o u t 5 cm stan d ard len g th , p erh ap s m ore.
In terest to F ish eries : L ittle or none.
L ocal N am es : L itera tu re :
rep etitiv e.
O nly 11
references in CLOFFA (1984), mostly
C LU P S pratam
S p ra tello m o rp h a B e rtin . 1 9 4 6
S p r a te llo m o r p h a B e rtin , 1946, in A n g e l, B e rtin & G u ib e. B u ll.M u s .n a tn .H is t.n a t.P a ris . (2)18:473-474 (type:
S auvagella m ad a g a sc a rie n sis b ia n alis B ertin , 1940).
D iagn ostic F eatu res : Sm all p ello n u lin es o f East A frica and
M ad ag ascar reach in g ab o u t 4.5 cm standard length. N o p re-p elv ic
scu tes, but pelvic scu te w ith sle n d e r v e rtic a l arm s (W -sh a p e d in
S p ra tello id es. D u ssu m ie ria. E tru m e u s).
S nout p o in ted , lo w er ja w
p rojecting slightly. L ow er g illrak ers 26 to 31: g illrak ers ab sen t on
p o sterio r face o f th ird epib ran ch ial. Pelvic fin w ith i 7 fin ray s, its
in s e rtio n w ell b e fo re d o rsa l fin o rig in :
la s t tw o a n a l finrays
separate, form ing a d istin c t little fin le t (o th erw ise know n only in
the Indian and Southeast A sian genera C o rica and C lupeichthys).
Biology, H abitat and
M adagascar, also Kenya.
D istribution
M arine
or estuarine in
no gillrackers on
hind face o f
3rd epibranchial
Interest to Fisheries : None.
Species : A single species
gili arches
1 2 3
reco g n ized :
£L b ia n a lis (B ertin , 1940), M adagascar, K enya, marine or
e stu arin e .
S p ra tello m o rp h a bianalis (B ertin , 1940)
C L U P S pratam
1
S p ra te llo id e s m a d a g a sc a rie n sis ( p a r t) S a u v a g e , 1 8 8 3 , B u ll.S o c .p h ilo m a t.P a ris . (7)7:160 ( M a d a g a s c a r ,
excluding the form s longianalis and brevidorsalis o f B ertin 1940, based on two o f the syntypes = Sauvagella
m ad ag a scarien sis).
Synonyms :
S auvagella m ad ag ascarien sis b ian alis B e rtin , 1940 (ty p es):
S auvagella b ia n a lis:W hitehead.
1963:336, fig . 14 ( ty p e s , d escr.): S p ratello m o rp h a b ian a lis - L o s s e , 1968:84 (M o m b a s a , d e s c r .) : T alw ar &
W h iteh ead , 1971:68 (key): W h ite h e a d , 1973:194, fig. 23 (key, synonym y, refs): C LO FFA , 1984:54 (sy n o n y m y ):
W hitehead & Bauchot, in p ress:ty p e s).
F A O Names
:
En - Two-finned round herring.
click fo r n e x t p a g e
c lic k fo r p r e v io u s p a g e
171
D iagnostic Features : Body elongate, a little com pressed, belly rounded, w ithout pre-pelvic scutes, but a
pelvic scute present w ith slender vertical arms. Gillrakers 26 to 31. Pelvic fin with i 7 finrays, well before dorsal
fin origin: last tw o anal finrays separate, form ing a distinct little fin let. This last feature identities the species
(found in no other clupeoid o f the w estern Indian Ocean).
G eographical Distribution: M adagascar (no locality given in
original description) and Kenya coast (stake traps at entrance to
Port Tudor, M om basa: probably more widespread).
H abitat and B iology
Presum ed
more specimens and data needed.
Size :
m arine
or
estuarine,
but
To 4.5 cm standard length.
Interest to F isheries : A pparently rare.
L ocal Nam es
: -
L iterature : Losse (1968
references in CLOFFA (1984).
best description to date): only five
C L U P E h ir
Ehirava D eran iy ag ala , 1929
Ehirava
D eraniyagala, 1929, Spolia zeylan., 15:34 (type:
Ehirava fluviatilis D eraniyagala,
1929).
Diagnostic Features :
Small slender-bodied pello­
nulines o f southern India
and Sri Lanka reaching
about 5 cm standard length.
Thin scutes, unkeeled and
2nd supramaxilla
barely visible before pelvic
fins, none behind.
Jaw
teeth very w eakly deve­
loped. G illrakers 24 to 30:
pre-pelvic scrutes
no gillrakers on posterior
(unkeeled,
w ithout arms,
face o f third epibranchial
hidden by scales)
(see
S p ratello m o rp h a).
maxilla blade
B ranchiostegal rays 5 or 6.
Pelvic fin w ith i 7 finrays,
its insertion a little before
dorsal fin origin: anal fin origin well behind last dorsal finray. Scales m oderate, 36 to 40 in lateral series,
D istinguished from all other clupeids in its area by the presence o f unkeeled pre-pelvic scutes and no post-pelvic
scutes.
172
Biology, H abitat and
India and Sri Lanka.
D istribution
: R ivers
and
estuaries, tolerant
o f brackishw ater;
coasts
o f southern
Interest to Fisheries : O f local importance.
Species : A single species re c o g n iz ed (W hitehead, 1963:333 wrongly included Spratelloides
Sauvage, 1883, now placed in Sauvagella):
R
madagascariensis
fluviatilis D eraniyagala, 1929, southern India.
Ehirava
fluviatilis
D eraniyagala,
1929
C LU E L h ir 1
Ehirava fluviatilis D eraniyagala, 1929, Spolia zeylan., 15:35, pi. 14 (western province o f Sri Lanka).
Synonyms : E h ira v a m a la b a ric u s (part):W hitehead, 1963:331 (8 types o f R fluviatilis, but m ixed w ith 5
M alabar specim ens o f D ayella m alabarica);E hirava fluviatilis - T alw ar & W hitehead, 1971:68 (key); W hitehead,
1973:196, fig. 25 (key, synonymy, refs); W ongratana, 1980:90, pis 13, 14 (revision).
FAO Names
E n - M alabar sprat.
D iagnostic Features : Body slender, belly rather rounded, w ith 5 to 8 thin unkeeled pre-pelvic scutes
(difficult to see unless scales rem oved), follow ed by an equally thin pelvic scute, all w ith slender ascending arms;
no post-pelvic scutes. Snout pointed, low er ja w a little prom inent, w ith barely visible teeth in each jaw ; second
supra-m axilla short, about h alf length o f m axilla blade. G illrakers 24 to 30. Pelvic fin insertion ju st before dorsal
fin origin. Closely resem bles G ilchristella o f southern A frican and the sym patric D ayella (but gillrakers present
on posterior face o f third epibranchial; also, pelvic insertion ju s t behind dorsal fin origin in D ayella, cf. just
before in Ehirava). See CLUP Ehir 1, Fishing A rea 31.
G eographical Distribution : Southern India and Sri Lanka.
H abitat and B iology:
stuaries, lagoons and as
m uch as
10 km up rivers, presumably tolerating a wide range o f salinity.
Size :
To 5 cm standard length, usually about 4 cm.
Interest to F isheries : O f local importance,
seines and liftnets, m arketed fresh or dried salted.
Local Names :
SRI LANKA: Ehirava.
caught
with
173
D a y ella
T alw ar & W hitehea, 1971
C LU P D ay
D ayella T alw ar & W h ite h e a d , 1 9 7 1 , B u ll.B r.M u s.n a t.H ist.(Z o o l.), 22(2):63
Day, 1873).
(type:
Spratelloides
m alabarica
D iagnostic Features :
gili arches
Sm all
p ello n u lin es
of
1 2 3
so u th e rn In d ia , re a c h in g
about
6
cm
stan d ard
length.
Sm all
irregular
scutes (1 to 4) underlying
scales o f belly, w ith only
rudim entary lateral arms;
pelvic scute norm al, but
2nd supra-m axilla
w ithout keel. L ow er jaw
slightly projecting; second
supra-m axilla rather spatulate. Low er gillrakers 24
gillrackers on hind face
to 27; about 7 short gill­
o f 3rd epibranchial
rakers on posterior face o f
third
epibranchial. B ran­
m axilla blade
chiostega! rays 5. Pelvic
fin w ith i 7 finrays, its
insertion som ewhat behind
dorsal fin origin; anal fin origin well behind last dorsal finray. Closely resem bles E hirava, w hich lacks gillrakers
on the posterior face o f the third epibranchial and has the pelvic insertion before the dorsal fin origin.
Biology, H abitat and D istribution :
Rivers and estuaries o f southwestern coasts o f India.
In terest to F ish eries : P ro b ab ly none.
Species :
A single
species reco g n ized :
D. malabarica (Day, 1873), southwestern India, fresh- and brackishwaters.
Dayella m alabarica (Day, 1873)
Spratelloides
C LU P D ay 1
m alabaricus D ay ,1873, Proc.zool.Soc.Lond.:240 (M alabar, India).
Synonyms :
Ehirava m a la b a ric u s:W h iteh ead , 1963:331 (confused w ith E. flu v ia tilis); D ayella m alabarica
Talwar & W hitehead, 1971:63, fig. lb (scu tes) (M alab ar, C añ ara, descr., new genus); W h iteh ead , 1973:195 (key,
sy n o n y m y , refs); W ongratana, 1980:91, pis 15, 16 (revision).
FAO Nam es :
En- D ay’s round herring.
174
D iagnostic Features : Body slender, belly rather rounded, w ith 1 to 4 thin, unkeeled and irregular scutes
(hidden by scales) w ithout vertical arm s; pelvic scute w ith vertical arm s (not W -shaped, cf. S pratelloides.
D ussum ieria); no post-pelvic scutes. Snout pointed, low er ja w a little prom inent, small but distinct teeth in each
jaw ; second supra-m axilla long, about 3/4 length o f m axilla blade. G illrakers 24 to 27. Pelvic insertion behind
dorsal fin origin. Closely
resem bles the sym patric E hirava fluviatilis, w hich has the pelvic insertion ju st before
dorsal fin origin and no gillrakers on the posterior face o f the third epibranchial.
G eographical D istribution
H abitat and B iology
and data needed.
Size :
: Southwestern India.
: R ivers and estuaries; more specimens
To about 6 cm standard length.
Interest to F isheries : Probably none
Local Nam es : -
CLUP C lu p e o i
C lupeoides B leeker, 1851
Clupeoides Bleeker, 1851, N a tu u rk .T ijd sc h r.N e d .-In d ië , 1:27 (type: C lupeoides borneensis Bleeker, 1851).
D iagnostic Features : Southeast A sian freshw ater pellonulines reaching about 8.6 cm. Prom inently keeled
scutes both before and behind pelvic fin base, belly compressed. Low er gillrakers 12 to 24. Pelvic finrays i 7, its
in sertio n a little before dorsal fin origin;
anal fin en tire, last tw o finrays not fo rm ing a sep arate fin le t (cf.
C orica and C lupeichthys).
B iology, H a b ita t and D istrib u tio n :
Interest to F isheries
S p ecies :
C.
C.
C.
C.
Rivers o f southeast A sia and Papua N ew Guinea.
: Probably rather little.
Four species re c o g n iz e d by W ongratana (1980):
borneensis B leeker, 1851, southeast A sia, freshw ater
hypselosom a B leeker, 1866, K alim antan, freshw ater
papuensis Ram sey & Ogilby, 1886), Papua N ew G uinea, freshw ater
venulosus W eber & d e B e a u fo rt, 1912, Papua N ew G uinea, freshw ater.
C lu p eoid es
b o rn een sis
CLU P C lupeoi 1
M eeker, 1851
C lu p e o id e s b o rn e e n s is B le e k e r,
In d o n e s ia ).
1 8 5 1 , N a tu u rk .T i|d sc h r.N e d .-In d ië , 1:275
( B a n je r m a s in ,
K a lim a n ta n ,
Synonyms:
C lupeoides exilis Fow ler, 1935:92, fig .12 (B angkok); F o w le r, 1941:637 (re p e a t); K o w a la
born een sis:Fow ler. 1941:636 (on B leeker); C lupeoides b o rn een sis: W hitehead, B o e se m a n & W heeler, 1966:73, pi. 8
(2) (types); W ontagrana, 1980:92, pis 17,18 (revision).
F AO
N am es:
En - Borneo river sprat.
pre-m axillary
teeth
D iagnostic Features : Body depth increasing w ith size o f fish
(from 21 to 34% o f standard length), belly sharply keeled, w ith 9 to
12 + 6 to 9 scutes. Pre-m axillae long and toothed; second supram axilla as long as or longer than blade o f maxilla. Low er gillrakers
18 to 24. Pectoral axillary scale at least 1/3 length o f fin. N o grey
or silvery stripe
along flank. C losely
resem bles C. hypselosom a,
w hich has only 12 to 19 low er gillrakers
and a silver stripe down the
flank. O ther
Clupeoides spp. have a much shorter second supramaxilla, no pre-m axillary teeth and a shorter pectoral axillary scale.
G eographical D istribution : Indonesia
(K alim an ­
tan), Thailand (Ayudhya and N akornsw an, 100 to 260 km
up river), Cambodia, Viet N am (southern part).
w
Habitat and Biology
d ata needed.
■- - i- - 1-- r . . . » ■ .. . —1
i. A
*i>1
Freshw ater, in rivers. More
\n
Size : To 7 cm standard length.
Interest to F isheries
contribiution to river fisheries.
m axilla
blade
: P erhaps
m akes
a
sm all
o°
L o c a l N a m es : L iteratu re : C hevey & Poulain (1940 - Cambodia);
Taki (1975 - Viet N am ).
20"
i .—
2nd supra-m axilla
*
•1 *
-k %
k
^^ V ex* L
<2==*-.
jT 4
t )— -
N ed.Tijdschr.D ierk., 3:293 (B anjerm asin,
K alim antan,
A tla sIc h th .In d .N e e rla n d .. 6:101 (on C hypselosoma).
Synonym s: K ow ala h y p se lo so m a :F ow ler, 1941:637 (on B leeker); W hitehead, B oesem an
pi. 8 (3) (holotype); Wongratana, 1980:93, pi. 16 (revision).
F A O Names :
V
C L U P C lu p eo i 2
C lupeoides hypselosom a Bleeker, 1866
Clupeoides hypselosom a Bleeker, 1866,
Clupea (clupeoides) potamophilus
Bleeker, 1872,
a
En - K alim antan river sprat.
Indonesia).
& Wheeler, 1966:74,
176
i
D iagnostic Features : Body m oderately deep, belly
sharply keeled, w ith 7 + 6 scu tes. P re-m ax illae long and
to o th ed ; second supra-m axilla as long as or longer than
blade o f m axilla. L o w e r g illra k e rs 12 to 19. P e c to ra l
a x illa ry scale at le a st 1/3 le n g th o f fin .
A pro m in en t
silver strip along flank. C losely resem bles CL b o rneensis
(s y m p a tric in K a lim a n ta n ), w h ic h has more low er g ill­
rakers (18 to 24), but no silver stripe along flank. O ther
C lu p e o id e s sp p . h a v e a m u c h s h o r te r s e c o n d supram a x illa , no p re -m a x illa ry te e th and a sh o rte r p e c to ra l
axillary scale.
pre-m axillary
teeth
m axilla
blade
supra-m axilla
Geographical
K apuas Rivers).
D istribution
H abitat and Biology
specimens and data needed.
Indonesia
(Barito
and
Freshw ater, in rivers. M ore
Size : To 4.2 cm standard length, perhaps more.
Interest to F ish eries
contribution to river fisheries.
P erhaps
m akes
a
small
L ocal N am es : -
C lu p eoid es p ap u en sis (Ramsey
& Ogilby, 1886)
C L U P C lu p eo i 3
C lupeoides papuensis R am sey & O gilby, 1886, P roc.L inn.S oc.N .S .W ., ( 2 ) l : 19 (Strickland R iver, Papua N ew
G uinea).
Synonym s
C lupeoides m ultispinus H ard en b erg ,1941219 (low er part o f D igoel river, P apua N ew
G uinea); K ow ala p a p u e n sis:F ow ler, 1941:636 (on R am sey & O g ilb y );
M u n ro , 1967:51, pi. 4 (51) (com piled);
Clupeoides papuensis - W ongratana, 1980:94, pis 19, 20 (revision).
FAO N am es :
E n - P ap u an riv er sprat.
177
D iagnostic F eatures
B ody m oderately elongate, its
depth 17 to 23% o f standard length, belly sharply keeled, with
(9) 10 or 11 (12) + (7) 8 or 9 (10) scutes. S n o u t b l u n t , p r e ­
m axillae short and toothless, but prom inent teeth in low er jaw ;
second supra-m axilla short, about h a lf length o f m axilla blade.
L ow er gillrak ers 15 to 19. P ectoral axillary scale sm all, less
th a n 1/3 length o f fin. A silver stripe along flan k , tapering
anteriorly. Closely resem bles the deeper-bodied sym patric C.
venulosus (depth 26 to 29% o f standard length; also, pre-dorsal
scales 14 or 15, cf. 17 to 19 in C7 papuensis). O ther C lupeoides
Spp. have a long and toothed p re -m a x illa , also pectoral axillary
scale 1/3 length o f fin or more.
pre-m axilla
toothless
m axilla
blade
2nd su p ra -m ax illa
G eographical D istribution
Southern Papua N ew
G uinea (Strickland, Fly and Digoel Rivers;
possibly more o°
w idespread).
H abitat and Biology : Freshw ater, in rivers. More
specim ens and data needed.
20°
Size :
To 7.7 cm standard length.
Interest to F ish eries
contribution to river fisheries.
L o ca l N am es
P erhaps
m akes
a
:-
40
C lupeoides venulosus W eber & D eB eaufort, 1912
Clupeoides venulosus W eber & D eBeaufort,
sm all
°
C L U P C lu p eo i 4
1912, V erh.K .A kad.W et.A m st., 17:3 (L o re n tz River, Irian Jaya).
S y n o n y m s:
K o w a la v e n u lo su s:F ow ler, 1941:638 (on W eber & D eB e au fo rt); M u n ro , 1967:50 (in key only);
C lupeoides venulosus - W ongratana, 1980:96, pis 21, 22 (revision).
F A O Names :
E n - W est Iria n river sprat.
178
D iagnostic Features : Body m oderately deep, its depth
26 to 29% o f standard length, belly sharply keeled, w ith 9 to 12
+ 7 to 9 scu tes. Snout blu n t, p re-m ax illae sh o rt and to o th less,
but prom inent teeth in low er jaw ; second supra-m axilla short, a
little over half length o f m axilla blade. Low er gillrakers 16 or
17. Pectoral axillary scale small, less than 1/3 length o f fin. A
silver stripe along flank, tap erin g anterio rly . C losely resem bles
the m ore slender-b o d ied sym patric CL p ap u en sis (d ep th 17 to pre-m axilla
23% o f standard length; also, p re -d o rsal scales 17 to 19, cf.
14 toothless
or 15 in CL venulosus). O th er C lu p eo id es spp. have a long and
to o th e d p re -m a x illa , also pectoral ax illary scale 1/3 len g th o f
fin or more.
m axilla
blade
2nd su p ra -m ax illa
G eographical D istribution : Indonesia (in Lorentz
R iv er, Irian Jaya) and P a p u a N e w G u in e a (F ly R iv e r);
probably in other southwest-flow ing rivers in area.
H abitat and B iology : Freshw ater, in rivers. More
specimens and data needed.
Size :
To 8.6 cm standard length.
Interest to F ish eries
contribution to river fisheries.
L o ca l N am es : -
P erhaps
m akes
a
sm all
179
C orica H a m ilto n -B u c h a n an , 1822
Corica H am ilton-B uchanan,
1822,
CLUP Coric
Fishes o f G anges:253, 283 (type:
Corica soborna
H am ilton-Buchanan,
1822).
D iagnostic Features : Small Indian and southeast A sian
fresh w ater p ello n u lin es reaching about 5 cm stan d ard length.
K eeled scutes both before and behind pelvic fin base, belly
com pressed . Jaw teeth m inute or ab sen t, none along sides o f
low er jaw . L ow er g illrak ers 19 to 27. P elv ic fin ray s i 7, its
insertion in advance o f dorsal fin origin; last two anal finrays
separated from rest o f fin, form ing a distinct fin let. R e se m b le s
Clupeichthys in this latter feature (but ja w teeth prom inent and
continued along sides o f lower jaw).
B iology, H a b ita t and D istrib u tio n :
(O rissa) and southeast Asia.
Interest to F isheries
R iv e rs o f In d ia
separate
finlet
: Perhaps make a sm all contribu­
tion.
S p ecies :
Two species re c o g n iz e d by Wongratana (1980):
anal fin
C. laciniata Fow ler, 1935, southeast A sia, freshw ater, estuarine
C. soborna H am ilton-Buchanan, 1822, India, southeast A sia, freshwater.
C orica
la cin ia ta
Fow ler, 1935
CLUP Coric 1
Corica laciniata Fow ler, 1935, P ro c .A c a d .n a t.S c i.P h ila d .. 87:92. fig. 11 (B angkok, Paknam).
Synonym s : Corica laciniata - Fowler,
24 (revision).
F A O Names :
1941:646 (B an g k o k , P ak n am , T h ailan d ); W o n g ratan a, 1980:96, pis 23,
E n - B an g k o k riv er sprat.
D ia g n o stic F ea tu res :
scutes. Teeth minute or absent
23 to 27. Last two anal fin ray s
has few er g illrak ers (19 to 21),
have a separate anal fin le t, but
B o d y m o d e ra te ly e lo n g a te , b e lly k e e le d , w ith 10 to 12 t 6 to 8 (usually 11 +7)
in jaw s; second supra-maxilla at least as long as m axilla blade. Lower gillrakers
form ing a separate fin le t. Resem bles C soborna (sy m p a tric in K alim an tan ), w hich
but more pre-dorsal bones (12, cf. 9 in C. lacinata). Species o f Clupeichthys also
jaw teeth prom inent and present on sides o f low er jaw.
180
G eographical D istribution :
(B angkok, P aknam , S ongkla L ak e);
R iver at B anjerm asin and A luhuluh).
G u lf o f T h a ila n d
K a lim a n ta n (B arito
---------1--------- T--------- 1--------- 1--------- !
*
n.
li
H a b ita t an d B io lo g y : Freshwater, in rivers, also in
estuaries (but salinities not recorded). M ore data needed.
Size :
V
r\
To 5 cm standard length.
In terest to F ish eries : P ro b ab ly none.
\ l
i
A
L ocal N am es : -
)
¡S # -
!(
I
•
.
c= 3 i „
:
j T
4
V
j
C o rica sob orn a H a m ilto n -B u c h a n an ,
1822
C LU P
C o ric
2
Corica soborna H am ilton-B uchanan, 1822, Fishes o f G an g es:253, 383 (Mahanadi River, Orissa, India).
Synonyms :
C o rica arg e n ta ta S w ain so n , 1839 (on CL sob o rn a H am ilto n -B u c h an a n , 1822);
Spratella
p s e u d o p te r u s B leek er, 1 8 5 2 P a m a n g k a t, K a l i m a n t a n
lectotype
only, p a r a le c to ty p e = CL la c i n ia t a fid e
W o n g ratan a, 1980:98); Corica so b o rn a :F ow ler, 1941:644 (m ostly on D ay, 1878); W hitehead, Boesem an & W heeler,
1966:76, pi. 9, fig. 1 (types, Pam angkat); W hitehead, 1973:192, fig. 21 (key, synonym s, but excluding CL perakensis
and CL laciniata, refs); W ongratana, 1980:97, pis 25, 26 (revision).
F A O Nam es :
E n - G anges riv er sprat.
D iagnostic Features : Body fairly elongate, belly keeled, w ith 11 + 6 or 7 (usually 7) scu tes. T eeth m inute
or absent in ja w s; second su p ra-m ax illa at least as long as m ax illa blade. L o w er g illrak ers 19 to 21. L ast two
anal finrays forming a separate f i n l e t R esem bles CL laciniata (sym patric in K alim antan, but see R em arks), w hich
has more gillrakers (23 to 27), but fewer pre-dorsal bones (9, cf. 12 in CL soborna). Species o f Clupeichthys also
have a separate anal fin le t, but ja w teeth prom inent and present on sides o f jaw .
click for n e x t p a g e
c lic k fo r p r e v io u s p a g e
181
G eographical D istribution : India (M ahanadi River,
O rissai
p e rh a p s a lso riv e rs o f B e n g a l) and In d o n e s ia
(P a m an g k at, K alim antan - but see Remarks).
H abitat and B iology : Freshw ater, in rivers, pos­
sibly also in estuaries. M ore specimens and data needed.
Size : To 4.1 cm standard length, perhaps to 5 cm.
: Probably none.
Interest to Fisheries
L ocal N am es : Literature :
See under synonyms.
Remarks :
The K alim an tan record is based solely
on the lectotype o f S p ratella p seu d o p teru s (R M N H 7116),
w hich has damaged gili arches.
CLUP C lu p ei
C lu p e ic h th y s B leek e r, 1855
C lu p e ic h th y s B le e k e r, 1855,
B leeker, 1855).
N a tu u rk .T ijd sch r.N ed .-In d ië,
9 (260)274
(ty p e: C lu p e ic h th y s g o n io g n a th u s
D iagnostic Features
Southeast A sian freshw ater pellonulines reaching about 6.5 cm standard length.
K eeled scu tes both before and behind pelvic fin base, belly compressed.
L o w er ja w te e th large and contin u ed
along side o f jaw . Low er gillrakers 13 to 19. Pelvic fin ray s i 6 or 7, its insertion below or a little behind dorsal fin
origin: last two anal fin ray s separated from rest o f fin, form ing a distinct fin le t. R e se m b le s C o rica in th is la tte r
feature (but teeth small, low er gillrakers 19 to 27).
can in e te eth
behind o u ter series
low er ja w teeth
at sides
separate
fin let
m axilla blade
supra-m axilla
Biology,
Habitat and Distribution
Interest to F isheries
S p ecies :
C.
C
C.
C.
: Rivers o f southeast
Asia.
: Probably rather little.
Four species re co g n ized by W ongratana (1980):
aesarnensis W ongratana, 1983, Thailand, freshwater
bleekeri (H ardenberg, 1936), K a lim an tan , fresh w ater
goniognathus B leeker, 1855, Thailand, Sumatra, freshwater
perakensis (H erre, 1936), M alaysia, freshwater
182
R em arks
T eeth rarely o ccur at the sides o f the low er ja w in clupeids (see P otam o th rissa and
L im n o th rissa ); e q u a lly ra re are fa n g -lik e te e th on th e p re -m a x illa e b e h in d th e n o rm a l o u te r se rie s (see
C y n o th rissa).
CLU P Clupei 1
C lupeichthys a esa r n e n sis W ongratana, 1983
Clupeichthys aesarnensis W ongratana, 1983, J a p a n .J .Ic h th y o l., 29(4):388, fig. 2 (U bonrat reservoir, K onkhan,
also H ualuang near U don Thani and Lam poa reservoir, K arasint, Thailand).
: None.
Synonym s
F A O Names :
E n - Thai riv er sprat.
j
/
-«eïgyT »« f i «
D iagnostic Features : Body moderately elongate, b e lly k e e le d , w ith
8 to 10 + 6 to 8 scu tes. S n o u t b lu n t,
pre-m axillae small and toothed, prom inent teeth at sym physis and along sides o f low er jaw ; second supra-m axilla
spatulate, about h a lf len g th o f m ax illa blade. L o w er g illrak ers 17 to 19. P ectoral ax illary scale less than h a lf
length o f fin; pelvic fin ray s i 7; last two anal fin ray s form ing a separate fin le t. R esem b les CL g o n io g n ath u s (but
lo w e r g illra k e rs o n ly 15 or 16), and CL b leekeri (b u t p ectoral ax illary scale m ore th a n h a lf len g th o f fin);
C. perakensis has only i 6 pelvic fin ra y s - Species o f C o rica also have a separate anal fin le t, but the ja w teeth
are small or minute and there are more lower gillrakers (19 to 27).
G eographical D istribution
Thailand
n o rth easte rn p art, M ekong drainage).
H abitat and B iology
p re s u m a b ly also in riv e rs;
needed.
Size :
To 4.6 cm
In terest
to
L ocal N am es : -
in
Freshw ater, in reservoirs,
m o re sp e c im e n s and d a ta
stan d ard
F ish eries
(reservoir
length.
40°
20°
: Perhaps none.
o°
20°
40°
183
C lu p eich th ys
b leek er i (Hardenberg,
1936)
CLU P Clupei 2
Corica bleekeri H ardenberg, 1936, T reubia, 15(3):229 (Kapuas River, K alim antan).
Synonyms :
C o ric a b le e k e ri:F o w ler,
1980:99, pis 27, 28 (revision).
F A O Names :
1941:646 (on H ard en b erg );
C lu p eich th y s bleekeri
W o n g ra ta n a ,
E n - K apuas riv er sprat.
D ia g n o stic F eatu res : Body m oderately elongate, belly keeled, w ith 9 or 10 + 7 or 8 scu tes. S nout b lu n t
pre-m axillae small and bearing up to 4 teeth behind the outer series, prom inent teeth at sym physis and along sides
o f low er jaw ; second su p ra-m ax illa sp atu late, about h a lf
------------- 1-------------- 1-------------- T “ ..... .....“ Ï ................
le n g th o f m a x illa b lad e.
L o w e r g illra k e rs 16 to 18.
*
P e c to ra l a x illa ry scale m o re th a n h a lf le n g th o f fin ;
p e l v i c finrays i 7;
la s t tw o a n a l finrays f o r m in g a
-,
/
€
se p a ra te fin let.
R esem b les CL ae sarn en sis (b u t p ectoral
axillary scale less than h a lf le n g th o f fin) and C. gonio­
gnathus
(same; also only 15 or 16 low er gillrakers); CL
perakensis has only i 6 pelvic finrays. Species o f C o ric a
■■
also hav e a s e p a ra te an al fin le t, bu t the ja w te e th are 20 small or minute and there are more low er gillrakers (19 to
*i
27).
yr r
»X
;
s )
G eographical
D istribution
w estern K alim antan, K apuas River).
Indonesia
(south-
■«o
p a s i*
3
H ab itat and B iology
F resh w ater, in rivers;
know n from rather few specim ens, more data needed.
20,
Size :
k
^
-
A .\
\_
\
To about 6 cm standard length.
In terest to F ish eries : Perhaps none.
40‘
(
L o ca l N am es : -
C lupeichthys goniognathus Bleeker, 1855
C LU P C lupei 3
Clupeichthys goniognathus B leeker, 1855, N atu u rk . T ijd sehr. N ed .-In d i ë , 9:275 (Lahat, southern Sumatra).
Synonym s : C o ric a g o n io g n a th u s:W eber & D e B e a u fo rt,1913:55, fig. 21 (m ixed w ith CL b lee k e ri); F ow ler,
1941:647 (on W eber & D e B e a u fo rt ; C lu p eich th y s g o n io g n ath u s - W ongratana, 1980:101, pis 31,32 (revision).
F A O Names :
E n - S um atran riv e r sprat.
184
D ia g n o stic F ea tu res : B ody m oderately elo n g ate, belly keeled , w ith 9 or 10 + 7 or 8 scu tes. S n ou t blunt,
pre-m axillae small and toothed, prom inent teeth at sym physis and along sides o f lower jaw ; second sup ra-m axilla
sp a tu la te , a b o u t h a lf le n g th o f m a x illa b lad e.
lo w e r
g illrak ers
15 or 16. P ecto ral axillary scale less th an h a lf
len g th o f fin; p e lv ic fin ray s i 7; last tw o anal finrays
*
form ing a separate fin le t. R esem b les C7 b leekeri (but
V —
_
r
f
p e c to r a l
a x i l l a r y s c a le m o r e th a n h a l f l e n g t h a n d w
C. aesarnensis (but low er gillrakers 17 to 19); Q p e ra k e n ­
sis has only i 6 pelvic finrays. Species o f Corica also have
a separate anal fin le t, b u t th e ja w te e th a re sm all or
minute and there are more lower gillrakers (19 to 27).
’
G eographical D istribution
Indonesia (southern
Sum atra at Lahat, upper reaches o f eastward flowing Musi
system ) and T hailand (at A yudhya).
P ro b ab ly in other
riv ers, b u t th e K a p u a s records
seem to
refer to
C. bleekeri.
\ r "
\ i
it
I
•¿ Q
^
•
L
°°
?
H ab itat and B iology
F resh w ater, in rivers;
know n from rather few specim ens, more data needed
20°
Size :
ƒ
V ...
y
h
% *■
r ' -
To 6.6 cm standard length.
In terest
to
F ish eries
: Probably none.
40„
1
S
L
S ____
_______i
L ocal N am es : -
C lu p eich th y s p era k en sis (H e rre ,
1936)
C L U P C lu p e i 4
Corica perakensis H erre, 1936, B u ll.R a ffle s M u s., (12):5, pi. 1 (P erak River, M alaysia).
Synonyms :
C o ric a
p e ra k e n s is :F ow l er, 1941:645 (o n H erre);
1980:102, pis 33, 34 (rev isio n ).
F A O Names :
E n - P erak riv er sprat.
C lup eich th y s
p erak en sis - W o n g ra ta n a ,
185
D iagnostic Features : Body m oderately elongate, belly keeled, w ith 7 to 9 + 4 to 6 scutes. Snout blunt
pre-m axillae small and toothed, prom inent teeth at sym physis and along sides o f low er jaw : second supra-m axilla
paddle-shaped, about h a lf length o f m axilla blade. Low er
g illrak ers 13 to 15.
P ecto ral a x illa ry scale m in u te or
absent; pelvic finrays i 6; last tw o anal finrays form ing a
separate finlet. O ther C lupeichthys spp. have i 7 pelvic
finrays, usually 15 or m ore total scutes (11 to 14 in
C. perakensis) and a pectoral axillary scale at least 1/4
length o f fin. Species o f Corica also have a separate anal
finlet, but the jaw teeth are small or minute and there are
more low er gillrakers (19 to 27).
G eographical D istribution
: M alaysia (Perak River).
H ab itat and B iology
: F resh w ater, in rivers;
known from only a few specimens, more data needed.
Size :
To
about 3
cm
standard
larger.
Interest to Fisheries : Probably none.
L ocal N am es : -
length,
perhaps
186
C LU P Potam l
P otam alosa Ogilby, 1897
P o tam alo sa O g ilb y, 1897, P ro c.L in n .S o c.N .S .W ., 21:504 (type: Potam alosa novaehollandiae o f Ogilby, 1897
(not M eletta novaehollandiae V alenciennes, 1847, w hich is Sprattus) =Clupea richm ondia M acleay, 1879).
D iagnostic
or more. K eeled
dorsal fin origin.
rays 8. P e lv ic
branchiostegal rays
Features : Relatively large A ustralian freshw ater pellonulines reaching 20 cm standard length
scutes both before and behind pelvic fin base and a complete series o f dorsal scutes from head to
Jaw teeth minute or absent; second supra-m axilla not as deep as m axilla blade. Branchiostegal
finrays i 7. Ju v en iles resem b le H y p e rlo p h u s, w hich has a d eep er second su p ra-m ax illa, only 4
and i 6 pelvic finrays.
p re-d o rsal
scutes
m axilla
blade
Biology, H abitat and D istrib u tion :
Interest to F isheries :
2nd
sup ra-m axilla
Freshw ater, in rivers o f N ew South W ales and V ictoria (Australia).
O f small local interest only.
S p ecies : A single species reco g n ized :
P richm ondia
(M acleay, 1879), southwestern A ustralia.
P otam alosa richm ondia (Macleay, 1879)
Clupea
W a le s ).
C LU P Potam l
1
richm ondia M acleay, 1879 (1 D ecem ber), Proc.L inn.Soc.N .S.W ., 4(3):380 (R ichm ond R iver, N ew South
Synonyms : C lu p ea n o v a e h o lla n d ia e :G ü n th er, 1868:431 (H aw k esb u ry R iv er, N ew South W ales; not novae­
hollandiae o f V alenciennes, 1847, w hich is Sprattus); Potam alosa antiqua O gilby, 1897 (N epean River); Potam alosa
n o v a e h o lla n d ia e :R o u g h le y , 1916:17 (b io l.,d is tr.);
P o ta m a lo s a ric h m o n d ia
M c C u llo c h , 1 9 2 9 :4 0 (sy n o n y m y );
Fowler, 1941:641 (N epean, H unter and Richm ond R ivers, N ew South W ales; 2 Fiji specimens questionably labelled
as such); M unro, 1956:24, fig. 167 (N ew South W ales and V ictoria); W h ite h e a d & B auchot, in press (syntype of
P o tam alo sa an tiq u a ).
F A O Names :
En - A u stra lia n fresh w ater herring.
187
Diagnostic Features : Body fairly elongate, belly keeled, w ith 16 to 18 + 14 or 15 scutes; about
14 dorsal
scutes w ith low keel from head to dorsal fin origin. M inute or no teeth in jaw s; second su p ra-m ax illa slender.
B ran ch io steg al rays 8.
P e lv i c fin ray s i 7; anal fin o rig in far behind dorsal fin base.
F o r d istin ctio n from
H yperlophus vittatus and IF translucidus, see genus.
G eographical D istribution
Southeastern A us­
tra lia (H unter, H aw k esb u ry , N ep ean and o th er rivers
o f New South Wales and Victoria).
H abitat and B iology : F reshw ater, in m iddle
and upper parts o f rivers, migrating dow n into estua­
ries to breed in July/August. Feeds on praw ns, worms
and insects.
S iz e :
To 20 cm standard length, perhaps more.
Interest to F ish eries
Taken in nets during
sp a w n in g se a so n (J u ly /A u g u s t); w ill a lso ta k e b ait.
Catches small.
L ocal N am es :
L itera tu re :
R oughley (1 9 1 6 - general biology).
H yp erlop h u s Ogilby,
C L U P H yper
1892
H y p e rlo p u s O g ilb y , 1892, R e c .A u s t.M u s .,2 (2 ):2 6 (ty p e: C lu p ea s p ra te llid e s O g ilb y , 1892
= M eletta v ittata
C astelnau, 1875). O m ochetus O gilby, 1897, P roc.L inn.S oc.N .S .W ., 22:72 (type: H yperlophus copii O gilby, 1897
M e le tta v itta ta C a s te ln a u , 18 7 5 ).
H y alo sp rattu s W h itley , 1936, Mem.Od M u s., 11(1) :25 ty p e : H y p e rlo p h u s
translucidus M cCulloch, 1917).
Diagnostic Features :
Sm all A u stra lia n coastal or estu arin e p ello n u lin es reach in g about
8 cm standard
length. K eeled scutes both before and behind pelvic fin base and a com plete series o f dorsal scutes from head to
dorsal fin origin. Jaw te e th m inute or ab sen t; second su p ra-m ax illa deeper th an m ax illa blade. B ran ch io steg al
rays only 4. P elvic fin ray s i 6. Resem bles small Potam alosa, w hich has a more slender second supra-m axilla, 8
branchiostegal rays and i 7 pelvic finrays.
pre-dorsal
scutes
I
blade
m axilla
2nd
sup ra-m ax illa
B iology, H ab ita t and D istr ib u tio n :
Bays and estuaries o f the southern coasts o f Australia.
Interest to Fisheries : Sold as bait.
S p ecies !
Two species, overlapping in part o f their ranges:
H. translucidus M cC ulloch, 1917, southern A ustralia
H. vittatus (C a ste ln a u , 1875), so u th ern A u stralia.
188
H yperlophus translucidus M cC ulloch, 1917
CLU P H yper 1
H yp erlo p h u s tra n slu c id u s M cC u llo ch , 1917, R e c .A u st.M u s., 11(71:165, pi. 29, fig. 3 (Sans Souci, Botany Bay,
New South Wales).
Synonyms : H y p erlo p hu s tran slu c id u s - M cC ulloch, 1929:40 (synonymy); Fow ler, 1941:643 (on M cCulloch);
M unro, 195624, fig. 169 (N ew South W ales, Q ueensland); Y abum oto & Uyeno, 1981:69, figs 3, 4, 7 (skeleton).
F A O N am es:
E n - T ran sp aren t sandy sprat.
wm i» »«m i -Am ;»i
--------*
D iagnostic Features : Body fairly elongate, belly keeled, w ith 17 + 9 scutes; about 19 dorsal scutes with
low keels from head to dorsal fin origin. No teeth in jaw s; second su p ra-m ax illa p ad d le-sh ap ed , low er portion
larger. B ran ch io steg al rays 4. Anal fin origin under or only slightly behind base o f last dorsal finray (well behind
in H. v itta tu s). S cales d eciduous; body translucent, w ith silver band along flank. Dorsal scutes also present in
Potam alosa richm ondia, but anal fin much further back.
Geographical Distribution :
N ew South W ales
and Q ueensland (26°48’S to 3 3 °5 9 ’S).
Habitat and Biology : S hallow sandy parts o f
b ay s and e s tu a rie s , p e rh a p s to le ra tin g lo w e re d s a li­
nities.
Size !
To about 6 cm standard length.
In terest to F ish eries : N o n e .
L ocal N am es : 40°
100°
H yperlophus vittatus (C a ste ln a u , 1875)
M e le tta v it ta t a C a s te ln a u , 1 8 7 5 ,
M elbourne, (2):46 (M elbourne, Victoria).
120°
140°
160°
180°
CLU P H yper 2
R e s .F is h .A u s tr.P h ila d .C e n t.E x h ib ., In terco l.E x h ib .E ssa y s, V ie t.D e p t.,
Synonyms :
C lupea sp ratellid es O gilby, 1892:24 (P a rra m a tta R iv er, N ew South W ales);
H yperlophus
( O m o c h e tu s ) copii Ogilby, 1897:72 (M aro u b ra, N ew S outh W ales); H y p erlo p h u s v itta tu s - M c C u llo c h , 1929:48
(synonym y); F ow ler, 1941:642 (N ew South W ales); M u n ro , 1 95624, fig. 168 W estern A ustralia to N ew South
W ales, also Q ueensland);
S c o tt, G lo v e r & S o u th co tt, 1973:69, fig. (all A u stra lia n states ex cep t T asm ania);
Y a b u m o to & U yeno, 1981:69, fig s 1, 2, 4 -7 (sk ele to n ); H u tch in s & T h o m p so n , 1983:18, 75, fig. 53 (K albarri,
W estern A ustralia to N ew South W ales); W hitehead & Bauchot, in press (syntypes in Paris).
F A O N am es:
E n - Sandy sprat.
189
D iagn ostic F ea tu res : Body fairly elongate, belly keeled, w ith 19 to 24 - 10 to 13 scutes; 23 to 27 dorsal
scutes w ith low keels from head to dorsal fin origin. No teeth in jaw s; second sup ra-m ax illa p ad d le-sh ap ed , low er
po rtion larger. B ran ch io steg al rays 4. Anal fin origin an eye diam eter or more behind base o f last dorsal finray
(under or only just behind in H translucidus). Scales more or less firm on body; a silvery band along flank. Dorsal
scutes also present in P o ta m a lo sa richm ondia, but pelvic fin ray s i 7 (i 6 in H yperlophus) and branchiostegal rays 8.
G eographical D istribution : Southern coasts o f
A u s tra lia (K albarri at 2 7 :3 0 ’S in W estern A u stralia to
South A ustralia, N ew South W ales and M oreton Bay,
Q ueen slan d ; not T asm ania).
H abitat and Biology : Schools in large numbers
in shallow sandy areas o f bays and estuaries.
Size :
To about 10 cm standard length.
Interest to F isheries
Sold as w hitebait and
used as a popular bait by anglers in W estern Australia.
L ocal N am es :
A U STR A LIA : W hitebait.
L itera tu re :
Scott, G lover & S o u th co tt (1973);
H utchins & Thom pson (1983 - general notes, figures).
100°
120°
140°
160°
180°
click for n e x t p a g e
c lic k fo r p r e v io u s p a g e
190
2.2.4
SU B F A M IL Y A L O SIN A E
FAO
N am es
: En - S h ad s.
notch
Diagnostic Features :
M o d e ra te to la rg e or (fo r
clupeoids) v ery
large h errin g -lik e fish es (to 60 cm stan d ard len g th , usu ally ab o u t 20 to
30 cm;
fully scu ted along b elly , scutes also p resen t along back before
dorsal fin in some (E thm idium ). U pper jaw not evenly rounded in front, but
w ith a distinct notch into w hich the sym physis o f the lower ja w fits;
ja w
teeth reduced or absent. D orsal fin at about m idpoint o f body, pelvic fins
below , anal fin short and well behind dorsal fin base; pelvic finrays i 6 to
8. S cales usually w ell attach ed . S tom ach m u scu lar in som e. O ften a dark
spot posterior to gili opening and in some species a series o f sim ilar spots
along flank (or even a second row below this).
Biology, H abitat and D istribution : The A losinae are m arine, pelagic, estuarine or freshw ater fishes, with
some species (or subspecies or races) anadrom ous, sem i-anadromous or totally freshw ater (rivers and lakes). All
are schooling fishes, and m ost or all are migratory (except w here landlocked).
Food includes fishes and various
in v erteb rates, bu t a few are filte r-fe ed e rs on p h y to p lan k to n (e.g. E thm alosa ). A num ber o f shads c o n tribute
significantly to fisheries, especially species o f B revoortia, Alosa and Tenualosa.
R em arks : Regan (1917) and some later authors have strongly deplored separation o f the shads from the
gizzard shads (subfam ily D orosom atinae)
and there is
some evidence that neither subfamily is in itself homogenous
(i.e . th a t all sp e c ie s h av e a c o m m o n
a n c e sto r).
F o r th e p re s e n t, h o w e v e r, it is c o n v e n ie n t to re ta in the
trad itio n al classificatio n .
There are 7 genera (2 Europe and w estern A tlantic, 1W est A frica, 3In d o -P ac ific
and 1eastern Pacific) and
31 species, the larg e st concentration o f species (also subspecies)
being the European and w estern A tlantic Alosa
(15 species or nearly half o f all the shads);
Key to the Genera :
la .
U pper g illrak ers o f first gili arch (w hen
n u m e ro u s) o v e rla p p in g th e lo w e r g il l­
rakers at angle o f arch (Fig. 1); pelvic
finrays i 6 or i 8; Europe, western Atlan­
tic, eastern P acific
2a.
upper gillrackers
overlapping ----low er gillrackers
No
pre-dorsal
scutes;
g illrakers
p resen t on p o sterio r face o f third
epibranchial (Fig. 2)
3a.
3b.
P e l v i c finrays i 8;
norm al
scales along back before dor­
sal fin; body scales norm al,
no t
deeply
overlap pin g ;
E u ro p e, w e ste rn and eastern
A tla n tic ..........................................
gili arches
1 2 3
Alosa
P e l v i c finrays i 6;
p a ire d
scales along back before dor­
sal fin enlarged, fringed (Fig.
3); b o d y sc ales d e e p ly o v e r­
lapping;
w este rn
A tla n tic
o n ly ................................................... Brevoortia
gillrakers on
hind face o f
3rd epibranchial
enlarg ed scales
Brevoortia
F ig ^ l
191
2b.
lb.
P ie-d o rsal scutes p resen t (Fig. 4);
no g illrak ers on p o sterio r face o f
th ird ep ib ran ch ial;
p e lv ic finrays
Ethmidium
i 6; eastern Pacific o n ly ...............
pre-dorsal scutes
U p p e r g illra k e rs o f firs t g ili arc h n o t
overlapping the low er gillrakers at angle
o f arch; pelvic finrays i 7; W est A frica
and Indo-Pacific
4a.
4b.
W est A fric a o n ly ; g illra k e rs very
num erous, upper gillrakers strongly
b en t, V -s h a p e d ...................................... Ethm alosa
E thm idium
In d o -P ac ific only; upper g illrakers
normal or curled outward
5a.
S c a le s la rg e , e v e n ly a rra n ­
ged, 37 to 47 in lateral
series;
m a r in e o r an ad ro mous
6a.
6b.
5b.
F ro n to -p a rie ta l
stria e
(on top o f head) many,
8 to 14 (F ig . 5a); g il l­
ra k e rs on in n e r a rc h e s
d is tin c tly c u rle d o u t­
w ard;
s c a le s p e r f o ­
ra te d .....................................
F ro n to -p a rie ta l
stria e
w eakly
developed,
usually h idden by skin
(F ig . 5b); g illra k e rs on
in n e r a rc h e s s t r a ig h t ;
scales not p erfo rated
T enualosa
Scales
except
91 in
w ater
A lo sa L in c k , 1790
H ilsa
sm all, series irreg ular
on upper flank, 77 to
la te ra l se rie s; fr e s h ­
o n l y .......................................
a.
H ilsa
b
fro n to -p a rie ta l
stria e
Tenualosa
F is- 5
Gudusia
C L U P A lo s
A lo sa Linck, 1790, N e u e ste s M ag.P hys.N aturgesch.. 6(3):35 (type: no species included, but C lupea alosa
Linnaeus, 1758 by tautonomy). Pom olobus Rafinesque, 1820 (April) W estern Review M isc .M a g .. 2(3): 170 and 1820
(D ecem ber), Ic h th y o l.O h ie n s is :38 (type: Pom olobus chrysochloris R afin esq u e,1820, but see com m ent by
H ildebrand, 1964:315, footnote). A lausa V alenciennes, 1847, H ist.nat.poiss., 20:389 (type: A lausa v u lg a ris
V alenciennes, 1847 = Clupea alosa Linnaeus, 1758) (apparently a variant spelling o f A losa). A lausella Gili, 1861,
P ro c.A cad .n at.S ei.P h ilad ., Suppl.:54 and I b i d .:36 (ty p e:
C lupea parvula M itchill, 1814 = probably C lupea
pseudoharengus W ilson, 1811, juvenile). C asp ia lo sa B erg, 1915, M ater.p o zn .ru ss.iy b o l.. 4(6):4 (type: Clupea caspia
Eichw ald, 1838).
Paralosa Roule, 1925, P o iss.E a u x d o u ce France :73 (type:
C lupea fallax L a c e p è d e , 1803)
(preoccupied by Paralosa Bleeker, 1868 = Sardinella; also by Paralosa Regan, 1916 = Hilsa).
D iagnostic F eature
M oderate or large herring-like fishes (to 60 cm standard length), som ew hat
com pressed, w ith a fairly prom inent keel o f scutes along the belly. U pper ja w w ith a distinct m edian notch.
U pper gillrakers, when num erous, folding down over low er gillrakers at angle o f first arch; total gillrakers (upper
plus low er) from 30 to as many as 130. Pelvic fin ra y s i 8.Scales norm al, hind borders sm ooth, no enlarged and
fringed scales along back before dorsal fin (cf. B re v o o rtia ). The notch ed u p p er ja w and long u p p er g illrak ers
separates A losa from sim ilar sym patric genera (C lupea, Sardina, Sardinella, etc.).
Biology, H abitat and D istribution :
M arin e, p elag ic, m ostly anadrom ous or sem i-an ad ro m o u s, but som e
purely fresh w ater species (su b sp ecies or races); sch o o lin g , m igratory (ex cep t w here land lo ck ed ). F eed on sm all
fishes and invertebrates, also phytoplankton in species w ith numerous gillrakers.M editerranean, B lack Sea, Sea o f
A zov, C aspian, and eastern and w estern N orth A tlantic (introduced into N orth Pacific).
192
Interest to Fisheries
waters (82%).
: Species o f A losa contributed 25 663 tons in 1983, mainly from A m erican A tlantic
Species : E arlier authors co n sid ered A lo sa , C a sp ialo sa and P o m o lo b u s as d istin c t gen era; S v eto v id o v (1 9 6 4)
un ited th em , but k ep t P o m o lo b u s as a su b g en u s o f A lo sa (sy m p atric in w estern N o rth A tlan tic). S pecies o f
‘C aspialosa’ are Ponto-C aspian in d istribution and only m eet ‘tru e ’ A losa in the associated rivers and lakes o f the
northern part o f the A egian Sea. To help identification, the 15 species o f A losa are listed geographically:
E a stern A tla n tic , M ed ite r r a n e a n
A. alosa (Linnaeus, 1758), eastern A tlantic, M editerranean
A. fallax (L acepède. 1803). eastern A tlantic M editerranean
P onto-C aspian
A.
A.
A.
A.
A.
A.
A.
pontica (E ichw ald, 1838), B lack Sea, Sea o f Azov
m aeotica (Grimm, 1901), Black Sea, Sea o f Azov
caspia (Eichwald, 1838), Caspian, also Black Sea, Sea o f Azov
brashnikovi (Borodin, 1904), Caspian
saposhnikovi (Grimm, 1887), Caspian
sphaerocephala (Berg, 1913), Caspian
kessleri (Grimm, 1887), Caspian (anadromous)
W estern A tla n tic
A.
A.
A.
A.
A.
A.
aestivalis (M itchill, 1814), w estern N orth and central Atlantic
alabamae Jordan & Evermann, 1896, western central Atlantic
chrisochloris (Rafinesque, 1820), w estern central A tlantic
mediocris M itc h ill, 1814), w estern N orth and central Atlantic
pseudoharengus (W ilson, 1811), w estern N orth and central Atlantic
sapidissim a (W ilson, 1811), western N orth and central Atlantic
Rem arks : The taxonom y o f the European shads, and especially the Ponto-C aspian species, still requires
much work. In the absence o f fully diagnostic m orphometric or meristic characters, biochem ical analysis may be
the best solution, especially in resolving questions o f subspecies.
Alosa a e stiv a lis
(M itc h ill, 1814)
CLUP Alos
1
Clupea aestivalis Mitchill, 1814, Fishes N ew York:21 (New York).
Synonym s
? C lupea fasciata LeSueur, 1818:233 (Sandw ich, M assachusetts); ? Pom olobus cyanonoton
Storer, 1848:242 (presum ed M assachusetts); Pom olobus a e stiv a lis:FW N A . 1964:324, fig. 80, also fig. 81 (growth
stages) (full synon., d e sc r., b iol.); A losa aestivalis - Liem & Scott, 1966:87, fig. (poor) (synopsis).
FAO Nam es : En - B lueback shad.
193
D i a g n o s t i c F e a t u r e s : Body fusiform , m oderately compressed, belly
with distinct keel o f scutes. Low er ja w rising steeply w ithin mouth; minute
teeth present at front o f jaw s (disappearing w ith age), no teeth on vomer.
L ow er gillrakers 41 to 52 (few er in fishes under 10 cm stan d ard len g th ),
slender. B ack dark blue, sometimes bluish-grey; a dark spot on shoulder.
R esem bles W p seudo h aren g u s in num ber o f g illrak ers and steeply rising
low er jaw , but that species has the eye usually larger than snout length, the
peritoneum silvery (cf. dark) and the back greyish-green. Other shads have
a gently rising low er ja w and more (59 to 73) or few er (18 to 24) lower
gillrakers or occur only in the G u lf o f M exico (W a lab a m ae). See C LU P
Alos 1, Fishing Area 31.
low er jaw
rising steeply
G eo g ra p h ica l D istrib u tio n
: North America (Atlan­
tic coasts from Cape B reton, N ova Scotia south to the St.
John’s River, Florida, also in lower parts o f rivers).
H abitat and B iology : E uryhaline,
a n a d ro m ó u s
p o s s ib ly w in te rin g n e a r th e b o tto m an d o u t fro m th e
coast, appro achin g the shore in late spring.
F eeds on
sm all fis h e s , copepods, sm all sh rim p s.
S p a w n s in
b ra c k ish - or fre s h w a te rs o f riv e rs , h a v in g a rriv e d in
coastal w aters a m onth or so later than W pseudoharengus
(C hesapeak e B ay in A p ril, ap p aren tly w hen the w ater is
above 70°C ; la te r fu rth e r n o rth ); the young d escen d in
autum n (C hesapeake Bay in O ctober/Novem ber).
Size : To about 38
about 25 to 30 cm.
cm
standard
length,
usually
Interest to Fisheries : Probably not distinguished
from W p seu d o h aren g u s in n o rth ern p art o f ran g e, but
c a tc h e s in south negligible (128 tons in 1974; not recorded
in 1983).
L ocal N am es : C A N A D A : B lueback herring; USA:
Blueback herring (AFS list).
Literature :
A lo sa
120 °
H ildebrand (i.e. FW NA, 1964 - USA, biol., etc.);
alab am ae Jordan & Evermann, 1896
A lo sa alabam ae Jordan & E v erm a n n , in E v e rm a n n ,
River, Tuscaloosa, Alabama).
10 0 °
80°
60°
40°
Lieni & Scott (1966 - Canada, synopsis).
C LU P A los 2
1896, R ep.U .S.C om m nr F ish , (21):203 (Black W arrior
Synonyms : M eletta suoerii Valenciennes, 1847:375 (W abash R iver, Illinois; nomen oblitum ); A losa ohiensis
E v e r m a n n , 1902:fig 1,2 (O hio R iver, L o u isv ille, K entu ck y ); A lo sa alabam ae - Regan, 1916:7 (key), (d e s c r.,
subspecies W alabamae ohiensis); FW N A , 1964:308, fig. 75 (full synon., descr., biol.); W h ite h e a d , 1967:84 (ty p e s
o f M eletta suoerii); W hitehead & B auchot, in press (same; see Remarks).
FAO N am es
: En - A labam a shad.
194
D iagnostic Features : Body fusiform , m oderately com pressed, belly
w ith distinct keel o f scutes. Low er ja w not rising steeply w ithin mouth; no
teeth present at front o f jaw s, no teeth on vomer. Low er gillrakers 41 to 48
at 30 cm stan dard len g th or m ore.
The only o th er shad o f the G u lf o f
M exico, W ch ry so ch lo ris, has only 20 to 24 gillrak ers. C losely resem bles
A. aestivalis and ¡V p seu d o h aren g u s o f A tla n tic co asts, but in them the
low er jaw rises v e ry steeply in th e m outh. See CLUP Alos 2, Fishing Area
31.
G eographical
ern p a rt, fro m
C hoctaw hatchee
to Arkansas and
lower
rising gently
D istribution
G u lf o f M exico (north­
th e M is s is s ip p i d e lta e a stw a rd to th e
River in Florida; also in rivers from Iowa
across to W est Virginia).
H abitat and B iology
E uryhaline, anadrom ous.
Food (no data given in FW N A , 1964). Spaw ns in fre sh ­
w ater in spring or early sum m er, ascending riv ers and
stream s, the young presum ably descending in autumn.
Size : To 51 cm standard length, usually about 40
to 45 cm.
Interest to F isheries
: O nly
small catches in the
past.
Local N am es : USA: G u lf shad, Ohio shad.
L iterature
: H ildebrand (i.e. FW N A,
1964 - biol.,
e tc .).
R em arks
The lectotype o f M eletta suoerii (a
ju v e n i l e ) h a s 33 lo w e r g il lr a k e r s , th u s
c an n o t be
A. ch ry so ch lo ris as supposed by H ild eb ran d (1 9 6 4 , i.e.
F W N A ). H ow ever, the name has not been used for either
species.
A losa alosa
(L innaeus,
1758)
C lupea alosa Linnaeus, 1758 Svst.nat.. 10th e d .:318 (in O cé a n o
C LU P A lo s 5
E uropaeo’).
Synonyms : A losa co m m u n is Y a rre ll, 1836:136 (Thames and Severn Rivers); A lausa v u lg a ris V alenciennes,
1847:391, pi. 604 (Europe, N orth A frica); A lo sa cuvieri Maim, 1877:154 (Bordeaux); A losa alosa - S v e to v id o v ,
1952:312, pi. 25, fig. 1 (but not A alosa bulgarica = A. caspia); Id em , 1963:351, pi. 25, fig. 1 (same): W h e e le r,
1969:70, fig. (synopsis); CLOFNAM , 1973:105 synonymy); W heeler, 1978:70, fig. (synopsis); FNAM , 1984:269, fig.
(synopsis); CLOFETA in press (synonymy).
F A O Names :
E n - A llis shad.
195
D iagnostic Features
Body som ew hat com pressed, fairly deep
(depth at pectoral fin more than head length), scutes apparent along belly.
U p p e r ja w n o tc h e d , lo w e r ja w fittin g in to it;
no te e th on v o m e r.
G illrak ers long, th in and num ero u s, to ta l 85 to 130, lo n g e r th a n g ili
filam ents.
A d a rk sp o t p o s te rio r to g ili o p e n in g (s o m e tim e s a b se n t;
occasionally 1 or 2 more spots). R esem bles W fallax, w hich has few er and
shorter gillrak ers (total 30 to 80) and 7 or 8 black spots along flank. See
CLU P Alos 5, Fishing Areas 34, 47 (in part).
G eographical D istribution : C oasts o f Europe,
from about B ergen to northern M auritania (but not in
Baltic), also w estern part o f M editerranean.
g illrak ers 85 to 130
gili filam en ts
H abitat and B iology :
M a rin e , p e la g ic ,
schooling and strongly m igratory, p en etratin g far up
rivers (but not into small tributaries). Enters rivers in
spring to spawn in May, usually at night and where the
c u rre n t is sw ift, th e a d u lts re tu rn in g to th e seas
afterw ard; ju v e n ile s pass dow n to the sea in autum n.
F e e d s on a w id e ran g e o f p la n k to n ic c ru s ta c e a n s;
la rg e r a d u lts feed on sm all sc h o o lin g fish e s. L ess
com m on than W fallax, but both have su ffered from
pollution and w eirs or other obstructions.
Size :
To 60 cm standard length and 2.7 kg.
Interest to F ish eries
Palatab le, but bony.
N o t a lw a y s s e p a r a te d fro m W
fallax in
fishery
records; the com bined catc h for both species in 1983
w as o n ly 971 to n s ( M o ro c c o , F r a n c e ,
Turkey).
Local Names :
DENM ARK:
M a js ild
GERMANY:
M aifisch;
NORWAY:
M aifisk;
THE
NETHERLANDS: Groote m eifisch; UK: M ayfish.
Literature :
W heeler (1969, 1978 - synopsis);
FNAM (1984 - synopsis).
Rem arks : A losa a f r ic a n a Regan, 1916 from A lgeria is not a distinct subspecies, but a hybrid betw een
A. alosa and A. fallax (Furnestin& V incent, 1958). A losa m ac e d o n ic a V inciguerra, 1921, is also not a subspecies
o f A. alosa (see under Alosa caspia).
A losa b ra sh n ik o v i (B o ro d in , 1904)
C LU P A lo s 8
C lupea casio-pontica var. brashnikovi B orodin, 1904, V estnik R yboprom yshlennosti, 19(3): 176, 180, 182-183,
184-185, fig. 2 (in part: A leksandrovsk N os 1-15, K rasnovodsk N os 25-26, 28-33; Fort Shevenko chosen).
Synonyms:
C lupea caspio-pontica var. g rim m i B o ro d in ,1904:180, 184 (A shur-ade, near A strabad Bay,
C aspian);
C lupea
(A lo sa) curensis Suvarov,1907:165, 192-194, fig. 4 (opp. m outh o f K ura R iver, and near
K ziligach Bay, Caspian); Clupeonella leucocephala Berg, 1913:27, pi. 14, figs 1, 2 (S u m g ait and Gyurgenchai at
Belidzhi Station, Caspian); Caspialosa brashnikovi autum nalis Berg, 1915:5, 6, pi. 1, fig. 2, pi. 2, fig. 3 (C h elek e n
Island, K rasnovodsk, L enkoran, K izilagach Bay opposite Sam ur R iver m outh at Tyulenii Island, C aspian);
Caspialosa caspia nigra K isselevitch, 1923:115, 139-140 (Bay or M ertvi K ultuk, Caspian; also included specimens
o f W saposhnikovi); Caspialosa kisselevitshi B ulgakov, 1926:27, pi. 2 (G a sa n -u li, Caspian); C aspialosa nirchi
M orosov, 1928:73 K rasnovodsk Bay, Caspian); C aspialosa brashnikovi kenderlensis B udam shin, 1938:36 Kendyrli
Bay, Caspian); Caspialosa brashnikovi orientalis M ikhailovskaya, 1941:560 (eastern coast o f southern Caspian north
to K enderyli Bay); C aspialosa brashnikovi agrachanica M ikhailovskaya,1941:559 (southern
part in w inter,
spaw ning in northern part at T yulenii and D arga Islands in sum m er, C aspian; earlier d escriptions on
M ikhailovskaya’s data by M eisner, 1932:52 and D ekhtyareva, 1940:47, but authorship given to M ikhailovskaya by
Svetovidov, 1952, 1963); C aspialosa brashnikovi sarensis M ikhailovskaya, 1941:560 (w estern coast o f southern and
central C aspian); C aspialosa brashnikovi derzhavini T arasevich, 1946:48 (biom etrics);
A losa brashnikovi
Svetovidov, 1952:251 et seq., pis 15-20 (synopsis;subspecies brashnikovi, grimm i, nirchi, autum nalis,
orientalis,
sa re n sis, ag rach an ica, k isse le v itsh i; also m ae o tic a, considered a species here; also A losa curensis, considered a
subspecies o f W brashnikovi here ; Idem . 1963:280 et seq., pis 15-20 (same).
FAO Nam es : En - C aspian m arine shad.
196
D iagnostic Features : Body fairly elongate, more ‘herring-like’ than ‘shad-like’. Total gillrakers 18 to 47,
thick and coarse, shorter, equal to or a little longer than gili filam ents, either straight or curved, pointed or blunt,
occasionally even bifu rcated at tips. Teeth well developed in both jaw s. O ther C aspian shads have deeper, more
shad-like bodies (A^ caspia, w hich has more gillrakers, 50 to 180;
saposhnikovi and A^ sphaerocephala').
Geographical Distribution :
(throughout, but m ainly in south).
C a s p ia n
S ea
t o
H abitat and B iology
B rackishw ater, n o n an ad ro m o u s, and n o t e n t e r i n g f r e s h w a t e r s , b u t
strongly m igratory. F eeds on sm all clu p eid s, gobies,
a th e rin e s , a lso c ru s ta c e a n s and o c c a s io n a lly in s e c ts
and m olluscs. V ariou s spaw ning p attern s am ong the
su b s p e c ie s (m o stly in sp rin g and su m m er a fte r an
inshore m igration and m ovem ent northward).
Size : To alm ost 50 cm standard length, usually
around 30 cm (mature at about 20 cm).
Interest to Fisheries : Som e o f the subspecies
are o f considerable im portance (esp ecially
b ra s h n i­
kovi b rash n ik o v i, A. brash n ik o v i ag rach an ica and A.
brashnikovi k is s e le v i ts h i). The total catch for all
Caspian A lo sa spp. (including A^ brashnikovi) in 1983
w as 3 561 tons. C oad (1980:70) claim ed stocks
depleted in Iranian freshwaters.
Local Names : USSR:
K asp iisk o -C h ern o m o rsk ie
1963) gives further R ussian names for the subspecies.
Literature :
M ikhailovskaya (1941 - subspecies);
morskie
sel’di, brazhnikovskie
sel’di.
Svetovidov
(1952,
Svetovidov (1952, 1963 - synopsis).
Rem arks : Svetovidov (1952, 1963) re c o g n iz e d nine subspecies, mainly separated on the number and form o f
the gillrakers. Following B anarescu (1964:244), Svetovidov’s
m aeotica o f the B lack Sea is recognized as a distinct
species (thus A^ brashnikovi becomes entirely Caspian), while his A^ curensis is included here as a subspecies o f
A. brashnikovi (which he seems to have suspected). The follow ing n in e su b sp ec ies occur:
A.
brashnikovi agrachanica: total gillrakers 20 to 46 (M 33); w estern Caspian, or southeast in winter
A. brashnikovi autumnalis Total gillrakers 21 to 37 (usually 26 to 33); southern Caspian
A.
brashnikovi brashnikovi: total gillrakers 27 to 47 (M 34); throughout Caspian, spawning in
A.
brashnikovi curensis: total gillrakers 32 to 38 (M 34); w estern part o f southern Caspian
A.
brashnikovi
grim m i: total
A.
brashnikovi
kisselevitshi:total gillrakers 30 to 49 (usually 34 to 42); southern Caspian, mainly southwest
A.
brashnikovi
nirchi: total
A.
brashnikovi
orientalis: total gillrakers 24 to 34 (M 28); eastern part o f southern
gillrakers 18 to 27
gillrakers 20 to 30
north
(M 22); eastern coasts o f southern Caspian
(M 25); eastern part o f CaspianLinck, 1790,
Caspian
A. brashnikovi sarensis: total gillrakers 23 to 32 (M 27); w estern part o f southern Caspian.
197
C L U P A lo s 7
A losa caspia (E ichw ald, 1838)
C lupea caspia E ichw ald, 1838, B u ll.Soc.N at.M oscou, 11(2): 134 (C aspian, tow ard the south).
Synonyms : C lupea tanaica Grim m, 1901:59, 62, 67 (Sea o f Azov); Alosa nordmanni A ntipa, 1906:28, pi. 2,
figs 6-10 (Black Sea, northw estern part); C lupea m acedonica V inciguerra, 1921:4 (Lake V olvi); C aspialosa
k n ipow itscha I l ’in, 1927:69, fig. (E nzeli B ay); C asp ialo sa casp ia var. p ersica I l ’in, 1927:72 (on data by Suvarov,
1907); A losa bulgarica D rensky, 1934:79, figs 1, 2 (B lack Sea, southw ard from Burgas); C aspialosa tanaica
palaestom i S adow sky, 1934:139 (P aläeostom e lagoon); A lo sa alo sa b u lg a ric a :S vetovidov, 1952:313, pi. 25, fig. 1
(synopsis); Id em , 1963:352, pi. 25, fig. (sam e); C aspialo sa caspia salina S v etovidov, 1963:51 (C aspian Sea,
n o rth east part); A losa alo sa m a ce d o n ic a :S vetovidov, 1952:313 m ention); Idem , 1963:352 (m ention); A losa caspia
- Svetovidov, 1952:229 et seq., pis 10-15, fig. 1 (synopsis; subspecies caspia, knipow itschi, salina, persica, tanaica,
nordm anni, alaeostom i); Idem , 1963:254 et seq., pis 10-15, fig. 1 (same); Bânârescu, 1964:245, fig. 107 (Black Sea;
as A losa) (C aspialosa) caspia nordm anni); CLO FN AM , 1973:106 (synonym ies; includes subspecies bulgarica,
nordm anni, palaeostom i, tanaica); FNAM , 1984:270, fig. (synopsis; subspecies as in CLOFNAM , diagnosed).
FAO Names :
En - C asp ian shad.
/ '
D iagnostic Features :
Body ‘shad-like’, usually rather deep and compressed. Total gillrakers 50 to 180,
long and thin, distinctly longer than gili filam ents. Teeth poorly developed in jaw s, sometimes barely detectable.
A black spot posterior to gili opening and occasionally further black spots on flank in some cases. Resem bles A.
k e ssleri in gillraker num bers, but that species is more slender and ‘herring-like’; other C aspian shads have few er
g illrak ers (A brashnikovi 18 to 47; A. sapashnikovi 25 to 41; A. sphaerocephala 26 to 42), while other B lack Sea
shads also have few er gillrakers and are more slender and ‘h e rrin g -lik e ’ A. p o n tic a 47 to 69, thus a slight overlap;
A. m aeo tica 33 to 46).
G eo g ra p h ica l D istr ib u tio n : Caspian Sea, B lack
Sea, Sea o f Azov.
H abitat and B iology :
E u ry h a lin e , m ig ra to ry ,
chiefly in b rack ish w aters, b u t en terin g fresh w aters to
sp aw n ; one o f th e m o st w a rm -lo v in g A lo sa o f th e
Caspian.
F eeds on plankton. D ep en d in g on the su b ­
species some are sem i-anadrom ous and spawn in fresh
or slightly saline w aters, mainly in the lower reaches,
w hereas others are p ractically m arine and spaw n in
brackish or even fully saline oceanic waters.
Size :
To 28 cm standard length, usually 18 to
22 cm (C aspian) or to 20 cm standard length, usually
only 14 to 16 cm (Black Sea, Sea o f Azov).
Interest to Fisheries :
O n ly A lo s a c a s p ia
caspia o f the C aspian Sea is o f im portance, the other
subspecies co ntributing little.
The to ta l c a tc h f o r
C a s p ia lo s a spp. (in c lu d in g A c a s p ia ) in 1983 w as
3 561 tons.
198
Local Nam es
1963:257 et seq .).
Literature :
: USSR:
K aspiisko
puzanok
(num erous
Svetovidov (1952, 1963 - synopsis);
other nam es for the
subspecies
- see
Svetovidov,
B anarescu (1964, Black Sea).
Remarks : Svetovidov (1952, 1963) re c o g n iz e d 7 subspecies, mainly separated on body shape, gillraker
num bers, pectoral fin length and distribution, adding a further subspecies subsequently (his previous W alosa
b u lgarica - S vetovidov, 1973:106, i.e . C L O F N A M ). E conom idis (1973) has show n that S vetovidov’s W alosa
m acedonica (V inciguerra, 1921, o f Lake
Volvi, Greece), is a true ‘C aspialosa’ (teeth on vom er and palatines) and
thus not W alosa; since its gillraker countis within the range for the widespread W caspia is and its body is deep
and ‘shad-like’, it seems best regarded as a subspecies o f A. caspia. Econom idis & Sinis in press) have described
yet another lacustrine subspecies, from Lake V istonis (Greece). This m akes 10 subspecies altogether:
Caspian
subspecies
A. caspia caspia: total gillrakers 70 to 150 (usually 100 to 140); C aspian (m ainly w estern half, but to
south and east in winter).
A.
caspia knipow itschi: total gillrakers 120 to 180 (usually 130 to 160); Caspian Sea (southern part,
mainly w estern half).
A. caspia p e r s ic a : total gillrakers 50 to 110 (usually 60 to 90); C aspian Sea (southern part, mainly in
east).
A. caspia salina: total gillrakers 60 to 120 (usually 80 to 100); C aspian Sea (eastern half).
Black Sea, Sea o f Azov subspecies
A.
caspia bulgarica: total gillrakers 100 to 107; B lack Sea (southw estern part).
A.
caspia m acedonica: total gillrakers 106 to 128; Lake Volvi only (M acedonia).
A. caspia nordm anni:
total gillrakers 66 to 88 (average about 80); B lack Sea (w estern part).
A.
caspia palaeostom i: total gillrakers 73 to 87 (average about 76); B lack Sea (south eastern part).
A.
caspia ta n a ic a : total gillrakers 62 to 85 (average about 76);
parts o f B lack Sea.
A. caspia subsp. A o f Economidis & Sinis
Sea o f A zov
and adjacent
eastern
total gillrakers 78 to 97; Lake V istonis only (M acedonia).
The scheme adopted here is com piled from the literature and merely represents a synopsis o f all the forms
o f A lo s a that have a deep, ‘shad-like body, numerous gillrakers (50 or more) and teeth on the vomer and palatines.
The true relationships, in spite o f Svetovidov’s painstaking studies, are yet to be discovered; a separation o f
C aspian from B lack Sea form s m ight be expected, but no m orphom etric or m eristic basis for this exists at present.
A lo sa c h r y so c h lo r is
(R a fin e sq u e , 1820)
C L U P A lo s 3
Pom olobus chrysochloris R afinesque, 1820, W est.R ev.M isc.M ag., 2:171 and Ichthyol.O h ien sis:39 (Ohio River,
but seldom ascending to Pittsburgh; see Rem arks for identity o f R afinesque’s species
Synonyms :
FAO Names :
P o m o lo b u s chrysochlorisJ W N A . 1964:315, fig. 78 (full synon., descr., biol.).
En - S kipjack shad.
8
«
YYYY
199
D iagnostic Features : Body fusiform , m oderately com pressed, belly
witn a d is tin c t keel o f scutes
Low er ja w not rising steeply w ithin mouth;
teeth present in jaw s, prom inent at front o f low er jaw , but none on vomer.
L ow er gillrakers 20 to 24, slender. Back bluish green, abruptly changing to
silver on flank; no dark spot at shoulder. The only other shad o f the G ulf
o f M exico, W alab a m ae , has 41 to 48 g illrak ers.
C losely resem bles A.
m ediocris o f A tlantic coasts, which has no upper and weak lower jaw teeth,
a dark shoulder spot and the body deeper than head length.
low er jaw
rising
gently
G eographical D istribution : G u lf o f M exico (from
C orpus C hristi in Texas eastward to Pensacola in Florida;
also in rivers, e.g. M ississippi and Ohio Rivers to M inne­
sota, W isconsin and Pennsylvania).
H a b ita t
and
Biology
E u ry h a lin e ,
enterin g
brackish- and freshw aters, but perhaps not always or not
c o n s is te n tly a n a d ro m o u s, a lth o u g h stro n g ly m ig ra to ry
w ithin rivers, m ostly in fast-flow ing w ater where they are
renow ned for leaping (hence the com m on name). Feeds on
small fishes, the juveniles on insects. Spawning tim es and
p la c e s n o t c e r ta in (FWNA, 1964:317). A d u lts se rv e as
hosts to the larvae (glochidia) o f the economically valuable
pearly mussel (Eusconaia ebena) o f the M ississippi basin.
Size
45 cm.
: To
50 cm
standard
length;
usually
30 to
In terest to F ish e r ies : Little or none.
L ocal N am es
USA: B lue or G reen h erring,
G olden shad, R iv er h errin g , S kipjack, S kipjack herring
(AFS list).
Literature : H ildebrand (i.e. FW N A, 1964 - b io l.,
etc.).
R em arks : H ildebrand (i.e. FW NA, 1964:315, footnote) noted that in the original description o f Pom olobus
chrysochloris. Rafinesque described the jaw s as toothless, the flesh as ‘esteem ed’ and the name as Ohio shad, all o f
w hich point to W alabamae and not W chrysochloris; how ever, to reverse the nom enclature w ould simply cause
confusion.
A lo s a f a lla x (L acep èd e , 1 8 0 3 )
C L U P A lo s 6
Clupea fallax L a c e p è d e, 1803, H is t.n a t.p o is s ., 5:452 (Seine).
Synonym s : Clupea rufa L a c e p è d e , 1803:452 (Seine); Clupea nilotica
E. G e o ffro y Saint-Hilaire, 1808:pl.
10
(N ile); C lupea fin ta C u v ier, 1829:399 (M ed iterran ean ); A lo sa fin ta var. lacu stris F atio , 1890:51 (L a k e s C o m o ,
M aggiore and Lugano); A lo sa b en acen sis B arb ie ri, 1907 (L ake G arda); A lo sa finta a lg e rie n sis, k illarn en sis and
g ra c ilis Regan, 1916:9, 10 (see Rem arks); A losa finta rhodanensis Roule, 1924:263 (Rhône) A losa fallax bolivari
Lozano y Rey, 1929:660, pi. 30, fig. 3 (co stas del Rif); A losa fallax - S vetovidov, 1952:310, figs 51, 52 (gillrakers),
pi. 24, fig. 2 (synopsis); Id e m . 1963:346, figs 51, 52, pi. 24, fig. 2 (s a m e ); W h eeler, 1969:128, fig. (synopsis);
C L O F N A M , 1973:107 (synonymy); W heeler, 1978:68, fig. (synopsis); E iras, 1980:7 (Portugal, m eristics); F N A M ,
1984:271, fig. (synopsis); CLOFETA, in press (synonymy); W h ite h e a d & B auchot, in press (some types o f A.
vulgaris Valenciennes, 1847 = A. fallax, not A. alosa; types o f Clupea rufa and A. f. rhodanensis).
FAO Names :
E n - T w aite shad.
200
gillrakers 30 to 80
D iagnostic Features : Body som ewhat com pressed, m oderately
deep (but depth at pectoral fin less than head length), scutes ap p aren t
along belly. U pper ja w notch ed , lo w er ja w fittin g into i f no teeth
on vom er. G illrak ers fairly short and stout, to tal 30 to 80, shorter
than gili filaments. A dark spot posterior to gili opening, followed by
7 or 8 sim ila r sp o ts a lo n g fla n k (b u t so m e tim e s fa in t or a b se n t).
Resem bles A. a lo s a , w hich has more and longer gillrakers (total 85 to
130) and at m ost only 3 dark spots on flank. See C LU P A los 6,
Fishing Areas 34, 47 (in part).
gili filam en ts
G eographical
D istribution : Coasts o f Europe,
from southern coast o f Iceland, B ritish Islands, Baltic;
south to M orocco; also w hole o f M e d iterran ean (b u t
not Black Sea - B an arescu , 1964).
H a b ita t
and
B io lo g y
M arin e,
p elag ic,
schooling and strongly m ig rato ry , but app aren tly not
p en etratin g far up rivers. E n ters tid al parts o f rivers
in M ay or early June to spawn there or a little above;
eggs dem ersal, scattered over gravel or sand, the fry
m oving dow n as they d evelop. F eed s on sm all fishes
and crustaceans, the young taking the fry o f herrings,
sprats and gobies. M ore com m on and w idespread than
A. alosa, but similarly suffers from river pollution and
to some extent from river barrages.
Size :
To 55 cm standard length and 1.5 kg.
Interest to F ish eries : P alatab le, but bony.
N ot alw ays separated from M alosa in fishery records
but probably m akes up the major part o f A losa catches
reported from France, Portugal and M orocco (267 tons
in 1983).
Local Names : Very many names are given by Palom bi & Santarelli (1961:9), including variants on Ceppa,
F int, Lacia, Losa, Savel, etc.
Literature :
Svetovidov (1952, 1963 - synopsis);
W heeler (1969, 1978 - synopsis);
FNAM (1984 -synopsis).
Rem arks : R egan (1916:7) re c o g n iz e d six subspecies and subsequent authors (e.g. Svetovidov, 1952, 1963
and W heeler, 1969) have accepted at least some o f these, based mainly on gillraker counts:
A n ad rom ou s su b sp ecies
A. fallax fallax: total gillrakers 37 to 42; A tlantic coasts.
A.
fallax n ilo tica : total gillrakers 34 to 37; M editerranean, A driatic (com m on),
Seas ( if so, th en rare).
perhaps M arm ara
and Black
Landlocked lacustrine subspecies (high gillraker counts)
A.
fallax benacensis: Lake Garda, Italy (30 to 40 or more low er gillrakers
gracilis, overlooking the earlier name benacensis o f Barbieri, 1907).
fide Regan, 1916
in his
W fallax
c lic k for n e x t p a g e
c lic k for p r e v io u s p a g e
201
A. fallax k illarn en sis: total gillrakers 43 to 53: landlocked in K illarney lakes, Ireland (to 23 cm standard
length only: feeds on C ladocera).
A.
fallax la c u stris: to tal g illrak ers to 76:
gillrakers fide R egan, 1916).
L akes C om o, M ag g io re and L u g an o , Italy (30 to 34 low er
R e g a n ’s subspecies a lg erien sis and R o u le ’s rh o d an en sis have both been placed in
CLOFNAM . 1973:107. 108).
fallax n ilo tica (e.g.
C LU P A los 15
A lo sa k e s s le r i G rim m , 1887
Clupea kessleri Grim m, 1887, A strakhanskaya seledka, St. P e te r s b u rg :7 ,
16 (Volga River delta).
Synonym s : C lupeonella caspia volgensis B erg ,1913:34, pi. 5 (near m outh o f Sulak R iver, w est coast o f
central Caspian):
Caspialosa volgensis b e r g i and im itans Tanasiichuk, 1938:33:
A losa k e s s le r i:Svetovidov.
1952:280, pi. 21, tig. 2 and pis 22, 23 (synopsis, subspecies k essleri, volgensis and pontica, the
latter here
considered distinct): Id em . 1963:314, pi. 21, fig. 2 and pis 22, 23 (same).
FAO Names :
En - Caspian anadromous shad.
Diagnostic Features : Body
thick, coarse and shorter than gili
(i.e. A. k e ssleri volgensis). Teeth
that species is deep-bodied, more
saposhnikovi. A. sphaerocephala).
fairly elongate, more ‘herring-like’ than ‘shad-like’. Total gillrakers 59 to 155,
filam ents in some, long, thin and equal to or longer than gili filam ents in others
well developed in both jaw s. R esem bles A^ caspia in num ber o f gillrakers, but
‘shad-like’: other Caspian shads have less than 50 gillrakers (A. brashnikovi. A.
G eographical D istribution : C aspian Sea (in sea
and along both shores o f cen tral and n o rth ern parts,
but in south and especially southeast in winter).
Habitat and Biology : E uryhaline, anadrom ous,
fro m su rfa c e d o w n to 85 m or m o re:
no rth w ard
m igration in spring (M arch /A p ril, but som etim es as
early as February or even January), a little distance
from the shore. Feeds chiefly on small fishes (C lupeo­
n e lla . a th e rin e s. etc.), less freq u en tly on cru stacean s
and in sect larvae (bu t m ainly on c ru stacean s in A .
k e ssleri volgensis): no feeding by A^ k e ssle ri k e ssleri
during migration up riv e rs. Spawns from mid-M ay to
A u g u st in the V olga, V yatk a, K am a, Oka and o th er
rivers, some entering w ith ripe gonads and spawning in
the low er reaches or even delta (A. k e ssle ri v o lg e n sis),
others entering unripe and reaching as much as 500 km
upstream : th e y o u n g d e sc e n d in la te su m m e r and
autum n.
202
Size ! To 52 cm standard length, usually 36 to 44 cm (or to 40 cm standard length, usually 27 to 29 cm in
A. k e ssleri volgensis').
Interest to Fisheries : The V olga subspecies is the more abundant and occupies second place among the
C aspian herrings. The total catch for all C aspian A losa spp. (including A^ kessleri) in 1983 was 3 561 tons.
Coad
(1980:70) claim ed stocks depleted in Iranian freshwaters.
L ocal Nam es
USSR: C hernospinka
(1952, 1963) gives further Russian names.
L itera tu re :
(A^ k e s s le r i
kessleri):
V olzhskaya
(V olga
subspecies).
Svetovidov
Svetovidov (1952, 1963 - synopsis).
R em arks : This is the only anadrom ous Caspian shad. The differences in gillraker form and in spawning and
feeding habits, as well as size, well justify Svetovidov’s recognition o f two subspecies:
A. k e ssleri k e ssle ri: total gillrakers 59 ta 92, thick, coarse and short; enters rivers unripe, spawns high and
does
not feed en route; usually 36 to 44 cm standard length.
A. k e s s le ri volgensis: total gillrakers 99 to 155, long, thin; enters rivers ripe, spawns in low er parts, feeding
en ro u te ; usually 27 to 29 cm standard length.
A lo sa m a eo tica (G rim m , 1901)
CLUP
Alos 9
Clupea m aeotica G rim m , 1901, V estnik R yboprom yshlennosti, 16(2):67 (Black Sea, Sea o f Azov).
Synonym s : Caspialosa brauneri N ik o lsk i, 1923:5 (D neister lim an at A kkem an and O vidiopol, B lack Sea);
C aspialosa brauneri m orph elongata Isachenko, 1925:128 (O chakov, B lack Sea); A losa brashnikovi m aeotica:
Svetovidov, 1952:254, pi. 15, fig. 2 (synopsis); Idem , 1963:284, pi. 15, fig. 2 (same); A losa pontica pontica var.
?:C L O F N A M , 1973:108 (see Remarks); A losa m aeotica - B a n arescu , 1964:244 (synopsis).
FAO N am es :
E n - B lack Sea shad.
¡sr-as-v
D iagnostic Features : Body fairly elongate, more ‘herring-like’ than ‘shad-like’. Total gillrakers 33 to 46,
thin and straight, often closely packed and pointed, usually shorter than gili filam ents. Teeth well developed in
both jaw s. O ther B lack Sea A losa have more gillrakers (TL caspia 50 to 80 and A^ pontica 47 to 66). Sardinella
aurita is more slender, has many more gillrakers and i 8 p elvic fin ray s (i 7 in A. m a e o tic a ) .
203
Geographical D istribution
o f A zov.
: B lack Sea and Sea
H abitat and Biology : Brackish- and saltw ater,
non-anadrom ous, entering lim ans and low er parts o f
river deltas, but only occasionally in freshw ater; a
cold-loving species, tolerating 3 or 4 ° C . Feeds mainly
on small fishes (m ainly sprats and anchovies) also on
shrim ps, gam m arids and other large crustaceans.
Spaw ns in spring and early sum m er.
Size : To 31 cm standard length, usually 16 to
20 cm .
Interest to F ish eries
O f little im portance;
com bined catches w ith A. pontica in the B lack Sea in
1983 were 1 839 tons (R om ania and B ulgaria).
L ocal Nam es : ROM ANIA: Scrum bie de mare;
USSR: Chernomorsko-azovskaya morskaya se l’d.
20»
0»
20»
40»
60»
Literature : Svetovidov (1952, 1963 - synopsis); B a n a re sc u (1964 - synopsis, B lack Sea).
Remarks : Svetovidov (1 9 7 3 :1 0 8 , i.e. CLOFNAM ), having earlier r e c o g n iz e d A^ m aeotica as a subspecies o f
A. brashnikovi, eventually placed it as a ‘v ar.’ (unnamed) o f A^ pontica, in spite o f the apparent difference in
gillrakers counts (33 to 46, cf. 47 to 69 in A^ pontica).
A lo sa m e d io c r is (M itc h ill, 1814)
C L U P A lo s 13
Clupea m ediocris M itchill, 1814, Fishes N ew Y o r k :20 (New York presumed).
Synonym s : ? C lu p e a p u s ilia M itchill, 1814:20 (New York presumed); C lupea m attow aca M itchill, 1815:451
(Long Island, N ew Y ork); Pom olobus m ediocris: FW5ÍA, 1964:319, fig. 79 full, synon., descr., biol.); Alosa
m ediocris - M ansueti, 1962:173, figs 3 (eggs), 4 (em bryos), 5-9 (larvae and juveniles) (M aryland; breeding, eggs,
larvae).
F A O Names : E n - H ic k o ry
sh a d .
D iagnostic Features : Body fusiform , m oderatley com pressed, belly
w ith a distinct keel o f scutes. Low er ja w very prom inent, but not rising
steeply w ithin m outh; teeth present in jaw s, reduced or in upper ja w
absent in larger fishes (over 23 cm standard length), none on vomer. Lower
gillrakers 18 to 23. A dark spot on shoulder, several obscure dark spots
along flank (m issing sometimes). Closely resembles A^ chrysochloris o f the
G u lf o f M exico, w hich has stronger ja w teeth, no shoulder spot and the
body depth less than head length. A losa aestivalis, A. pseudoharengus and
A. sapidissim a have more low er gillrakers (41 to 51, 38 to 44 and 59 to 73
respectively).
low er ja w
rising
gently ,
204
G eographical D istribution : N orth A m erica (A tlan­
tic coasts from M aine to the St John’s R iver, Florida and
in rivers).
H ab itat
and
B iology
E u ry h alin e,
entering 40°
b ra c k ish - and fre s h w a te r, a n a d ro m o u s. F e e d s on sm all
fish es, also squid, sm all crabs and o th er cru stacean s, as
well as fish eggs. S paw ns in tid al fresh w ater (P atu xen t
River, Chesapeake Bay in May fide Mansueti, 1962).
20 °
Size : To 60 cm standard length, usually 30 to
38 cm.
Interest to F isheries : O f m inor im portance, the
recorded catch for 1983 being only 34 tons.
C aught w ith
seine nets, pound nets, and in lesser quantities in gili and
fyke nets.
L ocal N am es
USA: B onejack, F all
Freshw ater taylor, Hickory Jack, Shad herring.
etc.);
herrin g ,
Literature : H ildebrand (i.e. FW N A, 1964 - b io l.,
Mansueti (1962 - spawning).
A lo sa p on tica (Eichw ald,
20
°
1 20 °
1 00 °
80 °
1838)
60 °
CLU P A los 10
C lupea pontica E ichw ald, 1838, B ull.S oc.N at.M oscou. 11(2): 135 (O dessa, B lack Sea).
Synonyms:
? A losa im m aculata B ennett, 1835 (Trabizon, B lack Sea); C lupea eichw aldi G rim m , 1901:67
(Roste on D on R iver); A losa pontica var. russae A ntipa, 1906:78:22, pi. 2, figs 1-5 (Danube delta) A losa pontica
var. danubii A ntipa, 1906:17, pi. 1, figs 4-8 (R om anian coast, Danube to and above Braila); A losa pontica var.
nigrescens A ntipa, 1906:21, pi. 1, figs 1-3 (D anube delta); A losa pontica var. moriae
A ntipa
1909; C aspialosa
pontica var. chtam alocephala Isachenko, 1925:110, pi. 2 (D neiper delta at O chakov); C aspialosa pontica var.
hypselocephala Isachenko, 1925:112, pi. 3; A losa kessleri pontica:Svetovidov, 1952:282, pi. 21, fig 1 (synopsis);
Idem , 1963:316, pi. 21, fig. 1 (sam e); A losa (C aspialosa) p o n tica:Banarescu, 1964:239, fig. 106 (synopsis);
subspecies borysthenis and issattschenkovi designated for the north-central B lack Sea, but not diagnosed; A losa
pontica - CLOFNAM , 1973:108 (full synonym y, but excluding W pontica pontica var. ? - see R em arks; FNAM ,
1984:272, fig. (synopsis, but included A. m aeotica in error).
FAO Names :
E n - Pontic
shad.
D iagnostic Features : Body fairly elongate, more ‘herring-like’ than ‘shad-like’. Total gillrakers 47 to 69,
rather thin, usually equal to or a little shorter than gili filam ents. Teeth well developed in both jaw s. Resem bles
A. caspia, w hich usually has more gillrakers (50 to 180, much longer than gili filam ents), poorly developed teeth
and a deeper, ‘shad-like’ body; A. m aeotica has few er gillrakers (33 to 36).
205
G eographical D istribution : B lack Sea and Sea
o f A zov (in sea and in th e D on, D anube and oth er
rivers, as m uch as 567 km up the D on and as far as
Kiev on the Dneiper before the dani was built).
60 °
H abitat and B iology : Euryhaline, anadrom ous,
m igratory, m o v in g n o rth w a rd in th e B la c k S ea in
spring and early summer. Feeds mainly on small fishes
(E ngraulis. C lu p eo n ella. S p ra ttu s), but also cru stacean s
(C rangon. U p o g eb ia. Id o th e a , g am m arids). S paw ns in
riv e rs fro m m id -M ay to m id -A u g u st, as m u ch as 40°
500 km or more upstream , but also in low er reaches;
the young descend quickly, but rem ain in or near the
deltas or estuaries until winter.
Size : To 39 cm standard length (D n ie p e r), or
30 cm, usually 14 to 18 cm (Don); a small form o f this
species reaches only 21 cm.
20°
Interest to F isheries : The m ain com ponent o f °°
the B lack Sea A losa catches, the com bined total (with
some W m aeotica) for 1983 being 1 839 tons (Romania
and B u l g a r i a ) .
Local N am es
G ERM ANY: D onauhering; ROM AN IA:
sel’d (further Russian names in Svetovidov, 1952, 1963).
Scrum bie
de
Literature : Svetovidov (1952, 1963 - synopsis;1973:108, i.e. CLOFNAM
synopsis. Black Sea).
D unàre;
USSR:
C hernom orskaya
synonym y); B an a re sc u (1964
R em arks : Svetovidov (1952, 1963) recognized a
large and a sm all form (m orphologically
identical
o th erw ise), but rejected d istin c tio n o f the ‘v a rie tie s’ ru ssa e . n ig re sc e n s, d a n u b ii, ch tam alo cep h ala and hypselo­
c ep h ala; B an arescu (1964239) added two more nam es, b o ry sth en is and issatsch en k o v i from the n o rth ern -cen tral
part o f the B lack Sea, but did not diagnose them. In his earlier w ork Svetovidov placed the present species as
merely a subspecies o f A. kessleri, but finally he recognized it as distinct, although combining it w ith his earlier
A. brashnikovi m aeotica (see R em arks under that species). There w ould appear to be much more w ork needed to
clarify the taxonom y o f this difficult genus.
A losa pseudoharengus (Wilson, 1811)
C lu p ea pseu d o h aren g u s W ilso n ,
(probably Philadelphia).
CLU P A los 14
1811, R ees’ N ew C yclopedia (A rt.C lu p ea). A m erican ed. only, 9 :unpaged
Synonyms : C lupea parvula M itchill, 1814:21 (N ew Y ork p resu m ed ); C lu p e a v e rn a lis M itch ill, 1815:454
(N ew York); Clupea m egalops R afinesque, 1818:206 (D elaw are R iv er); C lupea v irescen s D eKay, 1842:252, pi. 13,
fig. 37 (New York Bay); M eletta venosa V alenciennes, 1847:374 (New Y ork
P o m olobus p se u d o h are n g u s:FW N A .
1964:332, fig. 82 (full synon., descr., biol.); A losa pseudoharengus - W hitehead, 1967:82 (types o f M eletta venosa);
L eim & Scott, 1966:88, fig. (Canada, synopsis); Roy, 1968:16, figs (C anada, sy nopsis); W h iteh ead & B au ch o t, in
press (venosa types).
FAO Names
: En - Alewife
206
D iagnostic Features : Body fusiform, moderately com pressed, belly
w ith a d istin ct keel o f scutes. L o w e r ja w ris in g ste e p ly w ith in m o u th ;
minute teeth present at front o f jaw s (disappearing w ith age), no teeth on
vomer. Low er gillrakers 38 to 44, increasing w ith age. Back greyish green
on capture; a dark spot on shoulder. R esem bles W aestivalis in number o f
gillrak ers and steeply rising low er ja w , but that species has the eye not
longer than snout, the peritoneum dark (cf. silvery) and the back blue on
capture. O ther shads have a gently rising low er ja w and more (59 to 73) or
few er (18 to 24) lo w er g illra k e rs, or o ccur only in the G u lf o f M exico
(A. alabama ).
G eo g ra p h ica l D istrib u tio n : N orth A m erica (A tlan­
tic coasts from the G u lf o f St. Lawrence and N ova Scotia
to N o rth C aro lin a and in stream s and riv ers; in tro d u ced
into Lake O ntario, and now landlocked there and in Lakes
Erie (1931), H uron (1933), M ich ig an (1 9 4 9 ) and S uperior
(1954); also occurs in Lakes Seneca and Cayuga).
low er jaw
rising
steeply
"Sir-1-or
H abitat and B iology :
E u ry h alin e ,
entering
brackish- and fresh w aters, anadrom ous. F eed s on shrim ps
and sm all fis h e s , th e y o u n g o n d ia to m s , copepods and
ostracods w hile in the rivers (see especially Janssen, 1976,
1978). Spaw ns in M arch in C h esap eak e B ay area, late
April or May in Maine and in Canada, the adults migrating
up rivers and even small stream s, spawning in lakes and
q u ie t s tre tc h e s o f riv e rs ;
lan d lo ck ed pop u latio ns also
ascend afflu en t rivers and stream s; th e fry descend in
su m m er an d a u tu m n or e v e n as la te as N o v e m b e r or
D ecem ber.
Size :
To about 38 cm standard len g th , usually
about 25 to 30 cm.
Interest to Fisheries : Probably not always distin­
g u is h e d fro m W a e s tiv a lis ; th e re c o rd e d c a tc h o f A
p seu d o h a re n g u s in 1983 was 18 385 tons, being the highest
for a N orth A m erican shad.
Local Names :
CANADA: G asp ereau , G asp ero t;
USA:
herring, Gray herring, Grayback, Kyak, Sawbelly, W hite herring.
Literature : H ildebrand (i.e. FW NA, 1964 - USA, b io l., etc.);
Roy (1968 - Canada, biol., etc.).
Alosa sa p id issim a (W ilso n , 1811)
B igeye h errin g , B ran ch h errin g , F resh w ater
L eim & Scott (1966 - Canada, biol., etc.);
C LU P A los 4
Clupea sapidissim a W ilson, 1811, R ees’ N ew C y clo p ed ia (A rt.C lu p e a ). A m erican ed. only, 9:unpaged (probably
P h iladelphia).
Synonym s : ? Clupea indigena Mitchill, 1814:22 (N ew York presum ed); A lo sa p raestab ilis D eKay, 1842:255,
pi. 15, fig. 41 (New York); Mansueti, 1955:1, figs (M aryland, summary o f biol.) FW N A , 1964:295, fig. 71 and figs
7 2 - 7 4 (devel.) (full syn o n ., d escr., biol.);
L iem & S cott, 1966 (C an ad a, sy n o p sis); R oy, 1968:3, fig s (C anada,
synopsis).
FAO N am es :
En - A m erican shad.
207
D iagnostic Features : Body fusiform , m oderately com pressed, belly
w ith a d istin ct keel. L o w er ja w n o t risin g steeply w ith in m outh; teeth
absent in jaw s (m inute in juveniles to 15 cm standard length), no teeth on
vom er. L ow er g illrak ers 59 to 73, long and slen d er (few er in young). A
d a rk sp o t on sh o u ld e r, so m e tim e s fo llo w e d by several m ore, or even a
second row. R esem bles W pseudoharengus, w hich has the low er ja w rising
s t e e p l y w i t h i n m o u th and fe w e r lo w e r g il lr a k e r s (38 to 43), as also
A. aestiv alis and W m ed io cris (41 to 51 and 18 to 23 resp ectiv ely ). In
addition, W
pseudoharengus has a larg er eye (about equal to snout), but
shorter jaw s not reach in g behind hind m argin o f eye). See CLUP A los 4,
Fishing Area 31.
low er jaw
rising
gently (
G eographical D istribution
N orth A m e­
ric a (A tla n tic c o a s ts fro m th e S t. L a w r e n c e
R iv e r an d N o v a S c o tia s o u th w a r d to c e n tr a l
F lo rid a and in stream s and riv ers). In tro d u ctio n s
in th e G u lf o f M e x ic o a p p a re n tly fa ile d , b u t
th o s e o f f P a c ific c o a sts o f N o rth A m e ric a and
U SSR succeeded, the sp ecies now b eing found
fro m K a m c h a tk a and B ritis h C o lu m b ia to B a ja
C alifornia.
H abitat and B iology
E u ry h alin e, en ­
terin g brack ish- and fresh w aters, an ad ro m o u s; in
the sea, down to about 100 m, strongly migratory
(in d iv id u als have been cau g h t up to 3 000 km
from w here they w ere tag g e d ), in riv ers m ig ra­
tin g up as fa r as M o n tr e a l, b u t u s u a lly n o t
ascending as far u p stream as W p se u d o h a re n g u s.
F e e d s m ain ly on p la n k to n , m ainly copepods and
mysids, occasionally on small fishes (smelt, sandlance);
no feedin g during m ig ratio n up rivers;
the young feed on copepods and insect larvae in the rivers. Spawns at five years, som etimes at four, apparently in
their home river, on sandy or pebbly ground in the evening; enters rivers as early as N ovem ber in Florida and as
late as M ay or June in the north, depending on w ater tem perature (peak runs at about 18.5°C ), d escen d in g again
shortly after spawning; the young descend in the autumn.
Size :
To 60 cm standard length, common to 50 cm.
Interest to F isheries : N ow adays o f m oderate im portance, chiefly in rivers
for 1983 w as 2 712 tons (Canada and USA; Pacific coast catches very small).
Local Names :
of
the U SA;
the total catch
USA: A tlantic shad, Comm on shad, W hite shad.
Literature : M ansueti (1955 - M aryland, biol., etc.); H ildebrand (i.e. FW NA, 1964 - U SA, biol., etc.); Liem
& Scott (1966 - C an ad a, b io l., etc.); C le m e n s & W ilby (1967 - Canada, Pacific, synopsis); Roy (1968 - C anada,
biol., etc.); L e g g e tt & W h itn ey (1 9 7 2 - tem p eratu re and m ig ratio n s, A tla n tic
and P acific co asts);
D od so n &
Leggett (1974 - homing).
208
C LU P A los
A losa saposhnikovi (G rim m , 1887)
C lupea saposhnikovi Grim m, 1887, A strakanskaya seledkha, St. P e te r s b u rg ^ ,
S y n o n y m s:
(same).
A lo s a s a p o s h n ik o v i-S v e to v id o v , 1 9 5 2 :2 2 3 ,
FAO Names :
11
16 (Volga delta).
p i.9 , fig . l( s y n o p s is ) ; I d e m , 1 9 6 3 :2 4 7 , p i.9 f i g . 1
En - S aposhnikovi shad.
D iagnostic Features :
Body ‘shad-like’, ra th e r d e e p and c o m p re sse d ;
upper and lo w er head p ro files
straight. T otal g illrak ers 25 to 4 1 , m oderately th ick and short, tip s p o in ted , usu ally sh o rter th an gili filam ents.
Teeth well developed in both jaw s. Resem bles A^ sphaerocephala, w hich has a blunter head, w ith round upper and
lower head profiles; A. caspia has more gillrakers (50 to 180) and A brashnikovi is more slender and ‘herring-like’.
G eographical D istribution : Caspian Sea (through­
o ut, in c lu d in g lo w e r p a rts o f V o lg a R iv e r; m o re
common in north).
Habitat and Biology :
E u ry h a lin e , m ig ra to ry
but non -an ad ro m o u s;
one o f th e m o st c o ld -lo v in g
A losa o f the Caspian, being one o f the first to begin
its spring m ig ra tio n to th e n o rth .
F e e d s on la rg e
c ru s ta c e a n s and sm a ll fish e s.
S paw ns in n o rth ern
C aspian, from end o f A p ril, w ith a peak in m id-M ay,
moving into shallow water and spawning at depths o f 1
to 6 m and salinities o f 0.07 to ll% o , th e y o u n g la te r
m igrating southw ard (ap p aren tly dow n both c o asts),
but som e perhaps rem ain in g in the n o rth e rn parts o f
the Caspian.
Size
28 cm.
To 35 cm standard length, usually 14 to
Interest to Fisheries :
M o d e r a te l y la r g e
c a tc h e s re p o rte d by S v e to v id o v (1963:252); the total
catch for C aspialosa spp. (in clu d in g
sap o sh n ik o v i) in
depleted in Iranian freshwaters.
L ocal Nam es :
Literature :
1983 was 3 561 tons.
C oad (1980:70) claim ed stocks
U SSR: B o l’sheglazyi iii saposhnikovskii puzanok.
Svetovidov (1952, 1963 - synopsis).
Remarks :
Specim ens o f
description o f C aspialosa caspia nigra
saposh n ik o v i (No. 15938, ZI L e n in g ra d ) ap p aren tly co n trib u ted
K isselevitch = A losa brashnikovi (fide Svetovidov, 1963:242).
to the
209
C LU P A los
A losa sp h a ero cep h a la (Berg, 1913)
12
Clupeonella sphaerocephala Berg, 1913, M ater.pozn.russ.rybol., 2(3):20, pi. 12, figs 1, la (A grakhan Bay, at
Tyulenii Island, Turali, Caspian Sea).
A losa
Synonyms:
(same).
FAO N am es :
sphaerocephala
-
Svetovidov,
1952:227,
pi. 9,
fig.2(synopsis):
Idem ,1963:252.pl. 9,fig.2
E n - A g rak h an shad.
Diagnostic Features : Body shape ‘shad-like’, rather deep and com pressed: upper and low er head profiles
strongly rounded. Total gillrakers 26 to 42, rather long and thin, equal to or usually longer than gili filaments.
Teeth well developed in both jaw s. R esem bles
saposhnikovi, w hich has a more pointed head, the upper and
lower profiles straight: A^ caspia has more gillrakers (50 to 180) and
A brashnikovi is more slender and ‘herringlike’.
G eograp h ical D istr ib u tio n : C aspian Sea (more
com m on in the northern and central parts).
H a b ita t and B io lo g y : C onfin ed to sem i-saline
w aters:
n o n -an ad ro m o u s, not en te rin g fresh w aters.
Food not recorded by Svetovidov (1963:254). Spawns in
eastern part at north end o f Caspian, from m id-M ay to
en d o f J u n e , m o stly w h e n te m p e ra tu re s are 18 to
20°C , salinities 8 to 10 %o and at dep th s o f around
3 m:
young d ep art southw ard late r th an any o th er
clupeids, late in autumn.
Size : To 25 cm standard length, usually 16 to
18 cm.
Interest to
tities only.
L ocal
A grakhanskii
F isheries
N am es
puzanok.
: C aught in
U SSR:
sm all
K ru g lo g o lo v y i
quan­
iii
Literature : Svetovidov (1952, 1963 - synopsis).
c lic k for n e x t p a g e
c lic k fo r p r e v io u s p a g e
210
C LU P B revo
B rev o o rtia G ili 1861
B rev o o rtia G ilL
Latrobe).
1861, P ro c .A cad .n at.S c i.P liila d .:37
(type:
B rev o o rtia m enhaden G ili = C lupea ty ra n n u s
D iagnostic Features : M oderately large herring-like fishes (to
50 cm standard length, usually 25 to 35 cm). F arly d eep-bodied and
c o m p re sse d , th e b elly fu lly k e e le d w ith s c u t e s . H ead la rg e , e s p e ­
cially the gili cover: mouth large, the upper ja w distinctly notched in
the m idline, the tip o f the lower ja w fitting into this notch: no teeth
in jaw s in adults. G illrakers long, fin and very num erous, increasing
pre-dorsal scales
w ith size o f fish, those on upper part o f arch overlapping those on
low er p art at angle o f arch. D o rsal and anal fins short, the latter
beginning under or behind base o f last dorsal finray; pelvic finrays i 6. D istinguished from all other shads by the
m odified pre-dorsal scales on either side o f m idline, form ing a ridge: other body scales deeply overlapping and
rath er irreg u la r, w ith the hind m arg in serrated or p ectin ate.
The pre-dorsal scales and the notched upper jaw
distinguish B revoortia from all other clupeids.
Biology, H abitat and D istribution : M arine pelagic and form ing large or extrem ely large schools, mainly
in shallow coastal waters, but also down to 60 m or more.
Feed by filtering small elem ents o f plankton. Spawn in
sea, eggs floating. W estern A tlantic, from Nova Scotia southward to northern A rgentina.
Interest to Fisheries : A fter Sardinops and E ngraulis, often the third m ost exploited clupeoid genus.
total catch o f Brevoortia in 1983 was 1 345 519 to fishing Areas 21, 31 and 41).
The
Species : H ildebrand (i.e. FW NA, 1964:345-346) re c o g n iz e d 7 species, but B. brevicaudata, known from only
8specimens collected in 1874 and never recorded again, is here regarded as possibly a variant o f B. ty ra n n u s:
B.
B.
B.
B.
B.
B.
au rea (Spix & A gassiz, 1829) w estern A tlantic (Brazil, A rgentina)
gunteri H ildebrand, 1948, w estern A tlantic (G u lf o f M exico)
patronus Goode, 1879, western Atlantic (G ulf o f Mexico)
pectinata (Jenyns, 1842), w estern A tla n tic (B razil, A rg en tin a)
sm ithi H ildebrand, 1941, w estern A tlantic (North C arolina to Florida, also eastern G u lf o f M exico)
tyrannus (L atro b e, 1802), w estern A tlantic (Nova Scotia to Florida).
Rem arks : G eographically, the 6 species fall into three pairs (A tlantic, G u lf o f M exico, B razil/A rgentina):
system atically, B. gunteri o f the G u lf pairs w ith R sm ith i o f the A tlantic coasts (high gillraker form s, no spots on
flank), while R p atro n u s o f the G ulf pairs with the Atlantic R ty rann us (low gillraker forms, spots along flanks).
B r e v o o r tia a u r e a
(Spix & A g assiz , 1829)
Clupanodon au re u s Spix & A g assiz,1829, Select.gen.spec.piscium B ras.. 1:52.
incorrectly given only to Spix by W hitehead & Myers, 1971:485 - see K ottelet, 1985).
C LU P B revo 4
pi.
21
(Brazil)
(authorship
Synonyms : B revoortia a u r e a - H ildebrand, 1948:25, fig. 5 (B razil): FW N A, 1964:346 (in key only).
Figueiredo & M enezes, 1 9 7 8 :2 4 (S â o Paulo, Rio de Janeiro, B razil): M enni, R inguelet & A rám b u ru , 1984:107;
(A rgentina, on. refs).
FAO Nam es
: E n - B razilian m enhaden.
211
D iagnostic Features
Body deep and com pressed, scutes apparent along
belly. U pper jaw w ith distinct
m edian notch, no teeth. Pectoral fin tip short o f pelvic fin base by w idth o f 3 to 7 scales; pelvic fin w ith oblique
and alm ost straight hind margin. Pre-dorsal scales m odified; scales in lateral series 48 to 56, those on back and
above base o f anal fin not m arkedly sm aller than rest. A black spot behind gili opening, but none along flank.
Closely resembles R pectinata, w hich has only 35 to 46 scales in lateral series and pectoral fin tips short o f pelvic
fin base by w idth o f only 0 to 3 scales. O ther B revoortia species occur only to north o f B razil.
O ther
clupeids lack the m odified pre-dorsal scales.
G eographical D istribution
B razil (Rio de Janeiro, Sâo
Paulo, Santa C atarina and Rio Grande do Sul), A rgentina (probably
to m outh o f R io de la Plata, but needs confirm ation; based on
B erg, 1895:20, follow ed by D evincenzi & Burattini, 1928:pl. 17, 40„
fig. 3 (Uruguay) and Pozzi & B órdale, 1935:155 (A rgentina, name
o n ly ).
H abitat and B iology : M arin e, p e la g ic , sch o o lin g . N o d ata
on food or breeding.
Size : To about 20 cm standard length (largest fish 26 cm
in M useu de Z oologia, Sâo Paulo).
Interest to F isheries : A pparently not form ing the e n o r ­
m ous schools o f R tyrannus and R patronus; catches in 1983
w ere 1 560 to n s (B ra z il).
L ocal
Nam es
Literature :
to
: BRAZIL:
Savelha;
A RG EN TIN A :
2QO
o°
20
°
Lacha.
See under synonym s.
Rem arks : More w ork is needed on this species, especially its separation from B. pectinata and relationship
the northern species.
B revoortia
gunteri H ildebrand, 1948
CLUP B r e v o
5
B revoortia gunteri H ildebrand, 1948, Sm ithson.m isc.C olins, 107(18):31, figs 7, 8 (scales), 9 (pelvic fin)
( L o u is ia n a , T e x a s, G u lf o f M e x ic o ).
Synonym s : B revoortia gunteri - Suttkus, 1958:402 (G ulf o f M exico, distr.); Christm as & G unter, 1960:339
(G ulf o f M exico, G ulf o f Campeche); FW NA, 1964:376, fig. 93 (key, descr.,biol., etc.); D ahlberg, 1970:107 (key,
descr., refs); A non., 1976:68 (M exico, so u th to G u lf o f C a m p e c h e , k ey ).
FAO Nam es
: En - Finescale m enhaden
212
D iagnostic Features : Body deep and com pressed, scutes apparent along
belly. U pper jaw w ith distinct m edian notch, no teeth. Pelvic fin w ith oblique
and alm ost straight hind m argin, the inner finrays m arkedly shorter than the
outer finrays when fin folded back. Pre-dorsal scales m odified; scales in lateral
series 60 to 77, small and num erous, those on back and above base o f anal fin
markedly sm aller than rest. A black spot behind gili opening, but none along
flank. W ell distinguished
from R patronus o f the G u lf o f M exico, w hich has
few er scales (42 to 48) and a rounded hind border to the pelvic fin (inner fin rays
equal or nearly
equal to outer w hen fin folded back). Very closely resem bles
B. sm ithi (A tlantic, also w estern coast o f Florida), w hich has 30 to 32 scutes (cf.
27 to 29, infrequently 30 in B. gunteri) and the pectoral fin tip short o f pelvic fin
base by w idth o f 3 to 5 scales (cf. only 1 or 2). Other clupeids lack the m odified
pre-dorsal scales.
pelvic fin
G eographical D istribution : G u lf o f M exico (C h a n d e le u r
S ound, L o u isian a, to the G u lf o f C am p ech e, M ex ico ), but no
certain records from Caribbean.
Habitat and Biology : M arine, pelagic, schooling in inshore 40
w aters, including bays (p ossibly euryhaline). F eeds on plankton.
No data on breeding.
Size :
To 26.4 cm standard length.
Interest to F isheries : O f local im portance, but not sepa­
rated from R patronus.
L ocal N am es : Literature :
H ild e b ra n d (i.e . F W N A , 1964 - sy nopsis); 20„
D ahlberg (1970 - taxonomic and biol. data, refs).
120°
Brevoortia patronus G oode,
100°
1878
80°
60°
40°
CLU P B re v o
1
B revoortia p a tr o n u s G oode, 1878, Proc.U .S.natn.M us.,1:39 (Brazos Santiago, Texas, but in part also “M outh
o f the Rio G rande”).
Synonym s :
B revoortia patronus - H ildebrand, 1948:13, fig. 3 (key, descr., biol., etc.); Suttkus, 1956:390,
figs 1-20 (Louisiana, early life hrstory Idem , 1958:402 (distr.); Christm as & G unter, 1960:338 (distr.); Suttkus &
S undararaj, 1961:177 (rep ro d .); FW N A, 1964:365, fig. 91 (key, descr., biol., etc.); D ahlberg, 1970:102, fig. 2
(photo) (key, descr., refs); Swift, Verger & Parrish, 1977:21 (lower Ochlockonee River, Florida).
F A O Names
E n - G u lf m enhaden.
D iagnostic Features : Body fairly deep and com pressed,
scutes apparent
along belly. U pper ja w w ith d istin c t m edian n o tch , no teeth . P elv ic fin w ith
rounded hind m argin, inner finrays equal or nearly equal to outer finrays when fin
folded back. P re-d o rsal scales m odified;
scales in lateral series 42 to 48, large,
pelvic fin
213
those on back and above anal fin base only
opening, followed by a series o f spots along
range, but that species has more scales (6 0
outer w hen fin folded back), also no line o f
coasts o f Florida). O ther clupeids lack the
a little sm aller and more irregular than rest. A black spot behind gili
flank and often further spots above and below. O verlaps B. gunteri in
to 77) and more pointed pelvic fins (inner rays m arkedly shorter than
spots on flank (the same characters separate it from B. s m ith i o f G ulf
m odified pre-dorsal scales.
See CLU P B revo 1, Fishing A rea 31.
Geographical Distribution :
G u lf o f M exico (Florida Bay,
G ulf o f Cam peche, M exico), but no certain records from Carib­
bean.
Habitat and Biology : Marine, pelagic, schooling, inshore in
s u m m e r, b u t a t le a s t so m e m o v in g o u t in to d e e p e r w a te r s fro m
O ctober (M ississippi delta area), although adults have been
recorded near shore in w inter (G u lf coast o f F lo r id a ); euryhaline,
the commercial catch mostly in salinities o f 5 to 24 %o , but also
in fully salt w ater and up to 60 %o in the hypersaline Laguna
M adre, Texas,and down to 0.1 %o in Grand Lake, Louisiana.
Feeds in dense schools, filtering phytoplankton, but probably also
feed s at b o tto m (m ud in sto m ach ). A p p a re n tly b re e d s in w in te r
(October to February suggested by Suttkus, 1956, based on larvae
entering Lake Pontchartrain, Louisiana in D ecem ber to March; a
peak in January fide Suttkus & Sundararaj, 1961, based on gonad
s ta te s o f f th e L o u is ia n a c o a st).
40°
20O
Size : 25 cm sta n d a rd le n g th , u s u a lly a b o u t 20 cm.
Interest to F isheries : Probably contributes the m ajor part o f the G u lf o f M exico m enhaden catches. The
total catch in 1983 w as 923 585 tons.
Local Names :
L iterature
s u m m a rie s .
R em arks:
USA:
L a rg e s c a le
m en h ad e n .
: See synonym y, o f w hich H ildebrand (i.e. FW N A ,
1964) and D ahlberg (1970) are the best
H ybrids w ith FL smithi w e re d e sc rib e d by D a h lb e rg (1970- c o m m o n ).
B r e v o o r tia p e c tin a ta (J e n y n s, 1842)
A losa pectinata Jenyns,
C L U P B rev o 6
1842, Z ool.voy.Beagle, fishes: 135, pi. 25 (B ahia Blanca, A rgentina).
Synonym s
B revoortia pectinata - H ildebrand, 1948:21, fig. 4 (B razil, U ruguay, A rgentina); FW N A,
1964:345 (in key only); Figueiredo & M enezes, 1978:24, fig. 28 (Sâo Paulo to A rgentina, probably on refs only);
M e n n i, R i n g u e l e t & A rám buru, 1984 (A rg e n tin a , on refs).
FAO
N am es
En
-
A rg en tin e
m enhaden.
214
D iagn ostic F eatu res :
B ody deep and com p ressed , scutes ap p aren t along belly. U pper ja w w ith d istin ct
m edian notch, no teeth. Pectoral fin tip to pelvic fin base or at most short by the w idth o f 3 scales; pelvic fin
w ith oblique and alm ost straight hind margin. Pre-dorsal scales modified; scales in lateral series 35 to 46, those
on back and above base o f anal fin not m arkedly sm aller than rest. A black spot behind gili opening, but none
along flank. C losely resem bles R au re a , w hich has more scales in lateral series (48 to 56) and pectoral fin tip
short o f pelvic fin base by width o f 3 to 7 scales.O ther Brevoortia species occur only to
north o f Brazil. Other
clupeids lack the m odified pre-dorsal scales.
G eographical D istribution : B razil (apparently o ff Rio
G rande do Sul, but needs confirm ation), U ruguay (U ruguay R iver 40°
and M ontevideo fide H ildebrand, 1948:24) and A rgentina (Buenos
A ires, R io de la Plata fide H ildebrand, loc.cit.. thus confirm ing
Berg, 1895:18).
20°
H abitat and B iology : M arine, pelagic, schooling, abundant
in w inter in La Plata region and entering rivers, but not above
brackishw ater (Berg, 1895). No data on food. In L a g ô a dos Patos
(southern Brazil) eggs were found mainly in spring and summer, in
high salinity w aters, but the la rv ae tolerate low salinities also.
Size :
o°
To 30 cm standard length.
20°
Interest to Fisheries : Perhaps o f some local im portance,
but recorded catches for 1983 were only 29 tons (Uruguay).
Local Names :
A R G E N T IN A : Lacha.
Literature : W eis & Krug (1977 - eggs and larvae in Lagôa dos Patos).
R em arks : M ore w ork needed to separate it from B. aurea and clarify if their ranges really overlap.
B revoortia
smithi H ildebrand, 1941
C L U P B re v o 2
Brevoortia sm ithi H ildebrand, 1941, C opeia, (4):224 (B eaufort, N orth Carolina).
Synonym s : B revoortia smithi - H ildebrand, 1948:28, fig. 6 (key, descr.,biol., etc.); C hristm as & G unter,
1960:341 (G ulf o f M exico westw ard to Louisiana); Reintjes, 1962:93 (Sebastian, Florida, eggs and larvae); FW NA,
1964:372, fig. 92 (key, descr., biol., etc.); D ahlberg, 1970:104, fig. 1 (photo) (key, descr., refs).
FAO Nam es
: En
Y ellow fin menhaden.
Diagnostic Features :
Body deep and com pressed, scutes apparent along
belly. U pper ja w w ith d istin c t m edian n otch, no teeth. P elv ic fin w ith oblique
and alm ost straight hind m argin, the inner finrays m arkedly shorter than outer
finrays w hen fin folded back. Pre-dorsal scales modified; scales in lateral series
54 to 80 (usually about 60 to 70) small and num erous, those on back and above
anal fin markedly sm aller than rest. A black spot behind gili opening, but none
along flank.In G ulf o f M exico resem bles R gunteri in scale count and shape o f
pelvic fin
215
pelvic fins, but that species has only 27 to 29, rarely 30 s c u te s (30 to 32 in R sm ith i) and the pectoral fin tip short
o f pelvic fin base by w idth o f 1 or 2 scales (cf. by 3 to 5 scales). O ff A tlantic coasts resem bles R tyrannus, w hich
has few er scales (usually about 45 to 52) and spots along flanks (the same criteria separate it from R p atro n u s o f
Gulf). Other clupeids lack the m odified pre-dorsal scales. See CLUP Brevo 2, Fishing Area 31.
Geographical Distribution : A tla n tic c o a s t (fro m B e a u fo rt,
N orth Carolina, to Indian River, Florida); G u lf o f M exico (Florida
Bay to Louisiana, w ith possible break betw een B iscayne Bay and
Florida Bay).
4oo
Habitat and Biology :
M arin e, p elag ic, schooling (but
perhaps not in n o rth ern p art o f ran g e), inshore and in bays and
estu aries;
occurs in b rack ish w ater.
F e e d s by filte rin g ph y to plankton. Breeds in w inter (eggs and larvae in plankton), probably
N o v em b er th rough to F eb ru ary or M arch (no data from G u lf o f
Mexico).
Size :
To 29 cm standard length; usually about 20 cm.
Interest to Fisheries :
than R tyrannus.
N ot o f much importance. Less oily
20
L o ca l N am es : Literature : R eintjes (1962 - eggs and larvae); H ildebrand (i.e. FW N A, 1964 - synopsis); D ahlberg (1970 synopsis); Houde & Swanson (1975 - eggs and larvae).
Remarks :
comm on).
H ybrids with B. tyrannus and w ith B. p a tr o n u s
B rev o o rtia ty ra n n u s (L atro b e,
Clupea tyrannus Latrobe,
missing).
were described by D ahlberg (1970 - both hybrids
CLUP B re v o 3
1802)
1802,
Trans.A m .Phil.Soc., 5:77, pi.
1 (C hesapeake Bay; no descr., dorsal fin
Synonyms:
Clupea m enhaden M itchill, 1814:21 (presumed N ew York); C lupea carolinensis G ray, 1854:140
(South C arolina,; on M s name and description by Gronovius); H ildebrand, 1948:7, fig (key, descr., bio l., etc.):
FW N A, 1964:346, figs 86, 87, also 85 (scale), 88 (eggs and larvae), 89 (young) (key, descr., biol., etc.); Liem &
Scott, 1966:93, fig. (Canada, infrequent); R eintjes, 1969:1, pi. (photo), fig. 1 (egg to juvenile) (synopsis o f biol.,
fishery); D ahlberg, 1970:99, fig. 1 (photo) (key, descr., refs).
FAO Names :
En - A tla n tic
m enhaden.
216
D iagnostic Features : Body deep and com pressed, s c u te s apparent along
belly. U pper jaw w ith distinct m edian notch, no teeth. Pelvic fin w ith rounded
hind m argin, inner fin ra y s equal or nearly equal to outer fin ra y s when fin folded
back. Pre-dorsal scales modied; scales in lateral series 40 to 58 (usually about
45 to 52), those on back, above base o f anal fin and at base o f tail much smaller
and irregularly placed. A black spot behind gili opening, followed along flank by
a variable num ber o f sm aller
spots form ing up to 6 approxim ate lines. Easily
distinguished from R s m ith i, w hich has 54 to 80 (usually about 60 to 70) scales
and no lines o f spots on flank. O ther B revoortia species do not overlap in range.
Other clupeids lack the m odified pre-dorsal scales. See CLUP Brevo 3, Fishing
A r e a 31.
p elv ic fin
Geographical Distribution :
A tlan tic coasts (N ova Scotia
southward to Indian River, Florida).
H abitat and B iology : M arine, pelagic, schooling, inshore
in sum mer, but at least some m oving into deeper w ater in w inter
(but perhaps not in south o f range); forms large and very compact
schools, both o f adults and ju v en iles (good aerial photo in
R eintjes, 1969:fig. 3); north/south m igrations (spring and su m m er40”
versus autum n) occur, as also short-term m igrations in and out o f
bays and inlets depending on tides, season and w eather. Feeds by
filtering phyto- and Zooplankton (diatom s, copepods, euph au sid s).
B reeding season apparently lim ited by high w ater tem peratures 20°
(20.5°C monthly mean m axim um ), spawning probably throughout
year: spring spaw ners (A pril/M ay) in Cape Cod and Long Island
w aters, autum n spawners (O ctober/N ovem ber) from Long Island to
N orth C arolina (plus som e spaw ning June to A ugust), probably
w inter spaw ners o ff Florida (D ecem ber to M arch); nursery areas
in estuaries.
Size : To about 38 cm standard length (reported to 50 cm
total length), usually about 18 cm and 28 cm fork length in
fisheries o f northern and southern parts respectively.
Interest to Fisheries : The m ost im portant o f
420 345 tons. C om m ercially caught by purse seines,
etc.); the schools are often located from the air
Fishes principally used for production o f oil and for
Local Names :
Literature :
R eintjes (1969).
Remarks :
USA: Bugfish, Bunker, F atb ack ,
the A tlantic coast m enhadens, w ith a total catch in 1983 o f
but small numbers taken by other gear (pound nets, gillnets,
(see R ein tjes,1969, for good sum mary o f fishing methods).
fertilizer and fishmeal.
M enhaden, M ossbunker.
Extensive (see R eintjes, 1964), but well sum m arized by H ildebrand (i.e. FW N A, 1964) and
Hybrids with B. sm ithi described by Dahlberg (1970).
217
E thm idium T h o m p so n ,
C LU P Ethm
1916
E thm idium Thompson, 1916, Proc.U .S.natn.M us.. 5 0 :4 5 8 (type:
A la u sa m a c u la ta V a le n c ie n n e s).
Clupea (A lo s a ) notacanthoides
D iagnostic Features : M oderate-sized herring-like fishes
(to about 30 cm standard length), fairly deep and com pressed,
w ith a keel o f scutes along belly; a com plete series o f scutes
also present on back, from occiput to dorsal fin. U pper jaw
w ith a distinct m edian notch. G illrakers long, fine and n u m e ­
ro u s, those o f upper arch folding down over lower gillrakers at
angle o f first arch. P elvic f i n r a y s i 6; anal fin short, well
behind dorsal fin base. Scales adherent, hind border denticu­
late, about 50 to 60 in lateral series. D ark spots on flanks,
usually vertically oval. Species o f Sardinops have spots on the
flanks, but are more slender, lack the notched upper ja w and
have radiating bony striae on the operculum ; Strangom era is
also more slender and lacks the notch, also no spots on flank
and p e lv ic fin ra y s i 7.
B iology,
H abitat
Interest
to
Species
subspecies:
E.
and
scutes
: See species.
F ish eries : See species.
: M odern authors have recognized a single species, at m ost considering H ildebrand’s R
m a c u la tu m
(V a le n c ie n n e s ,
Ethm idium m aculatum
A lausa
D istrib u tion
p re -d o rs a l
Steindachner
maculata
1847), e a s te rn
S o u th P a c ific .
(V alenciennes, 1847)
Valenciennes, 1847,
chilcae as a
CLUP Ethm 1
H ist.nat.poiss., 20:430 (Valparaiso, Chile).
Synonym s
? A lausa coerulea V alenciennes, 1847:432 (V alparaiso, C hile; in part on drawing); Clupea
notacanthus
G ünther,
1868:443
(Valparaiso); D elfin, 1901:39 (on G ünther); C lupea (A losa) notacanthoides
Steindachner, 1869:309, pi. 7 (M azatlan - error);
Clupea m aculata: D elfin, 1901:39 (Tome, Valparaiso, Iquiqui);
E thm idium chilcae H ildebrand, 1946:82, fig. 17 C hincha Island, Callao and Pisco Bay, Peru); B revoortia
(Ethm idium ) m aculata:De Buen, 1958:98, fig. 2 (gillrakers) (Valparaiso); Brevoortia m aculata
chilcae: Chirichigno,
1963:15, fig.70 (photo) (Pta Pizarro, Peru);
Ethm idium m aculatum - Fow ler, 1945:4 (full refs); M ann, 1954:133,
fig. (A rica to Talcahuano, Chile); W hitehead, 1967:88 et seq. (types o f m aculata, coerulea); Idem , 1970:18 (type
o f notacanthoides lost); B ore & M artin ez ,1981 :unpaged, fig. (photo) (A rica to Talcahuano, C hile); Leible &
A lveal, 1982:16, figs 1-3 (same range); W hitehead & B auchot, in press (types o f m aculata, coerulea, the
latter
lo st).
FAO
N am es
En
-
P acific
m enhaden.
218
D iagnostic Features : Body fairly deep, com pressed, scutes present along back (before dorsal fin) as well as
along belly. U pper ja w w ith m edian notch; no teeth. G illrakers fine and slender, about 130 to 150 in low er limb.
Pelvic fin ra y s i 6; anal fin short, w ith iii 12 to 15 fin ra y s , well behind last dorsal fin ra y . Scales w ith pectinated
hind border. A dark spot behind gili cover and up to 4 series o f vertically oval spots along flank. D istinguished
from all other clupeids in the area (O pisthonem a, H arengula, Lile in north;
Strangom era, Sardinops in south) by
the notched upper jaw and pre-dorsal scutes: soots are absent in all except Sardinops, w hich is also more slender
and has a striated operculum.
G eographical D istribution : Peru (Puerto Pizarro
at 3 °3 4 ’S in G ulf o f Guayaquil to C allao and P isco Bay;
presum ably to border w ith C hile, but no records) and
Chile (w hole coast south to Talcahuano at 36°40’S).
H abitat and B iology : M arine, pelagic, coastal,
form ing large schools. Feeds by filtering phytoplankton
(diatom s, etc.) and Zooplankton (especially copepods),
but w ill also take small fishes and crustaceans (fide
L eible & A lv e a l, 1982). B reeds in spring (C hile),
depositing its eggs in coastal w aters on sand, to w hich
the sticky eggs cling, the larvae rising and joining the
surface plankton on hatching.
Size ! To about 26 cm standard length, 35 cm
total length.
Interest to Fisheries : A m oderate contribution
to clupeoid catches in the area, the total catch in 1983
being 6 213 tons (Peru 2 100 tons, Chile 4 113 tons).
C aught by purse seines, both by com m ercial fleets and
by artisanal fisherm en. In 1978, 74.4% o f the catch
w ent for reduction, 21.8% was not processed, 2.7% was
canned and 0.1% was frozen (Bore & M artinez, 1981).
L ocal N am es : Literature :
L eible & A lv eal (1982 - synopsis).
Remarks : A uthors have often recognized a Peruvian subspecies (chilcae o f H ildebrand) as distinct from the
C hilean form (m a c u la tu m ); M ann (1954:134) stated that the geographic division betw een the subspecies w as at
A ntofagasta, Chile. B ecause o f size differences in the material studied by H ildebrand and others, W hitehead
(1970:19) doubted that body depth, pectoral length and scale form could separate the subspecies, although there
m ight be a slight difference in head length. M ore w ork is needed.
E th m a lo s a
R egan,
CLU P E th
1917
E thm alosa R egan,
B o w d ih ) .
1917, A nn.M ag.nat.H ist., (8)19:302 (type: Alausa d o rsalis V alenciennes = Clupea fim briata
D iagnostic Features : M oderate-sized herring-like fishes (to 35 cm standard length), fairly deep and
com pressed, w ith a keel o f scutes along the belly. U pper ja w w ith a distinct m edian notch. G illrakers long, fine
and num erous; upper gillrakers o f first arch not overlapping low er gillrakers at angle o f arch, the upper rakers
strongly bent, V -shaped. P elv ic finrays i 7; anal fin well behind dorsal fin base. Scales adherent, hind border
d e n tic u la te .
Species o f S ard in ella have the u p p er ja w ro u n d ed , not n o tch ed , in the m idline; also, the upper
gillrakers are not V-shaped.
Biology, Habitat and Distribution
Interest to Fisheries
Species :
: See species.
: See species.
A single species recognized:
E. fim briata (Bowdich, 1825), W est A frica.
219
E th m a losa
Cape
fim b ria ta (B o w d ic h , 1825)
Clupea fim briata Bowdich (S.), in Bowdich (T.E.), 1825, Excurs.M adeira:234 ,
V erde Islands, but m ost likely G am bia).
CLU P E th
1
fig. 44 (claim ed Praia, S. Tiago,
Synonyms : H arengula forsteri V alenciennes, 1847:299 (S. Tiago, Cape Verde Islands - probably on Bowdich);
M eletta senegalensis V alenciennes, 1847:418 (Senegal); A lausa dorsalis V alenciennes, 1847:418 (G orée, Senegal);
A lausa platycephalus Bleeker, 1863:123 (Ghana); C lupea setosa Steindachner, 1870:311, pi. 6 (L iberia, Gabon);
Ethm alosa dorsalis:Longhurst, 1971:353, 356 (W est A frica, synopsis, biol. fishery); Ethm alosa fim briata -Fowler,
1936:175, fig. 70 (G am bia, Congo River mouth); W hitehead, 1967:85 (types etc. o f forsteri, senegalensis, dorsalis);
Idem , 1970:17 (types o f setosa);
CLOFFA, 1984:42 (all refs in freshw ater); C LO FETA , in press all m arine and
freshwater refs); W hitehead & Bauchot, in press (types etc. o f forsteri, senegalensis, dorsalis).
FAO N am es :
En
- Bonga
shad.
D iagnostic Features : Body fairly deep, com pressed, sc u tes present along belly.
U pper ja w w ith distinct
m edian notch, into w hich tip o f low er ja w fits. Low er gillrakers long, fine and num erous, about 3 tim es as long as
gili filam ents, upper gillrakers bent sharply upw ard, V-shaped. Pelvic fin ra y s i 7; caudal
fin tips long and pointed.
A faint dark spot behind gili cover (som etim es follow ed by others); dorsal fin tip black; caudal fin deep chrome
yellow; golden tints on body. Resem bles S a rd in e lla aurita. Sk ro u x i and especially Sk m aderensis, but these are
more slender, have a rounded upper ja w (not notched) and the upper gillrakers are not bent upward like an elbow.
A losa species do not overlap R d o rsalis in the north o f its range (also upper gillrakers not bent, pelvic fin ra y s i 8).
See C L U P E th m 1, F is h in g A reas 34, 47 (in p a rt).
G eographical D istribution
Eastern central A tlantic
( D a k h la , w estern Sahara, to at least Lobito, A ngola - i.e. from
24° N to 12° S; d w arf population in Lake N o k o u é , Benin).
R ecords from Cape V erde Islands based on erroneous type
locality for R fim briata by B ow dich - follow ed by later
au th o rs.
H abitat and B iology : E uryhaline, inshore w aters, also
lagoons and more than 300 km up rivers (e.g. G am bia River,
w here they move dow n during flooding, but up again during
intrusion o f seaw ater in the dry season). Feeds by filtering
phytoplankton, chiefly diatom s (full analysis by B ainbridge,
1963). B reeds throughout year in w aters o f salinities 3.5 to
35% o,
but w ith peaks in at least some areas (M arch, June/
July and O ctober/N ovem ber at m outh o f G am bia R iver - see
Scheffers & C o n an d , 1976; July to Septem ber o ff Sierra
L e o n e ; N ovem ber to M ay/June o ff Ivory C oast and N igeria,
i.e. progressively later to south); spaw ns in the sea, in
estuaries and in rivers.
Size : To 35 cm standard length, usually about 20 to
25 cm.
220
Interest to F isheries
The largest fisheries are in Senegal, Sierra L eone, Ivory C oast, N igeria and
C am eroun, mainly in the dry season. Total c a rc h e s for 1983 w ere 88 121 tons (N igeria 31 622, Sierra Leone
21 127 tons, Ivory Coast 14 618 tons). C aught by canoe fisherm en using purse seines and encircling nets, also
seine nets in lagoons and estuaries. M arketed fresh, also sm oked and dried (the latter greatly preferred in
Cam eroun and perhaps elsew here, but the Senegam bian catch is mostly m arketed fresh).
Local Names
W EST AFRICA: Bonga.
Literature : B ainbridge (1963 - food); Scheffers & Conand (1976 - Senegam bian region - b io l.); W hitehead
(i.e. CLOFETA, in press - all refs to 1984).
H ilsa R eagan, 1917
C LU P Hi Is
Paralosa Regan, 1916, A nn.D urban M us., 1(3): 167 (type: Clupea durbanensis R egan = Clupea kelee Cuvier;
p re-o ccu p ied by P ara lo sa B leek er, 1868 = S ard in ella).
H ilsa Regan, 1917, A nn.M ag.nat.H ist., (8)9:303 (type:
C lu p e a durbanensis
Regan;
to replace
Paralosa Regan, 1916).
M acrura Fow ler, 1941,
Bull.U .S.nata M us.,
(100):626 (type: Clupea kelee Cuvier).
D ia g n o s tic F e a tu r e s : M oderate-sized herring-like fishes (to 25 cm stan­
dard length, but usually to about 20 cm), fairly deep and com pressed, w ith a keel
o f scutes along belly.
Top o f head w ith num erous lo n g itu d in al striae (fronto­
parietal striae); upper ja w w ith a distinct m edian notch. G illrakers fine and
num erous, those on inner arches curled outw ard. Pelvic f in r a y s i 7; anal fin
short, well behind dorsal fin base. Scales m oderate, 39 to 44 in lateral series,
perforated. R esem bles Tenualosa, w hich lacks striae on top o f head, has more or
less straight gillrakers on inner arches and no perforations on scales; species o f
Sardinella and H erklotsichthys have the upper jaw rounded, not notched, in the
midline;
gizzard shads (N em atalosa, etc.) have an inferior m outh and have a
filam entous last dorsal finray.
fro n to -p a rie ta l
stria e
Biology, Habitat and Distribution : See species.
Jnterest to Fisheries : See species.
Species :
A single species
top o f head
recognized:
H. kelee (C uvier, 1829), Ind o -W est P acific.
Hilsa kelee (C u v ier, 1829)
C LU P H ils 1
C lupea kelee C uvier, 1829, R ègne animal, 2nd ed., 2:320 (on k eelee o f R ussell, 1803; Vizagapatnam ).
Synonyms : Clupea sinensis:Bloch, 1795:38, pi. 405 (name taken from Linnaeus, 1758 = ? Tenualosa reevesii;
B loch’s specim en is IT kelee ex Tranquebar); C lupeonia blochii Valenciennes, 1847:353 (on Bloch’s sinensis); Alosa
brevis B leeker, 1848:638 (B im a, Sum baw a Island); A lausa kanagurta B leeker, 1852:13, 34 (Jakarta, M untok);
A lausa brachysom a B leeker, 1853:527 (Padang); A lo sa m a la y a n a B le e k e r, 1 8 6 6 :2 9 4 (J a v a , S u m a tra ); C lu p ea
platygaster G unther, 1868:448 (Sum atra); H arengula (Paralosa) zeynalica H ubrecht, 1879 (nom en nudum ); C lupea
durbanensis R egan, 1906:4, pi. 4 (D urban B ay ; M acru ra k e le e : F o w ler, 1941:627-629 (also M acrura b re v is, M.
d u rb a n e n s is ); M a c ru ra b r e v is :M u n ro , 1 9 6 7 :5 3 , p i .5, fig. 59 (P a p u a N ew G u in e a ); H ilsa kelee - W h iteh ead ,
1965:129 (revision); W hitehead et ah , 1966:79, 80, pi 10, fig. 1 (types o f zey lan ica, m alayana, brachysom a, b revis,
k a n a g u rta ); W h ite h e a d , 1 9 6 7 :9 0 , 91 (ty p es o f k e le e , b lo c h ii); Id e m , 1 9 6 9:267, pi. le (B lo c h 's sin e n sis); Id e m ,
1973b:197. fig. 26 (kev. synonym y, refs): W ongratana, 1980:156, pis 102, 103 (revision); SFSA, in press(South
A frica); Whitehead & Bauchot, in press (types o f kelee, b lo ch ii).
FAO Names :
En - Kelee shad.
221
D iagnostic Features : Body fairly deep and com pressed, belly w ith distinct keel o f scu tes. Top o f head
w ith num erous fronto-parietal striae; upper ja w w ith m edian notch. G illrakers about 100 to 175, those on inner
arches distinctly curled; outer row o f gili filam ents on first arch not more than h a lf length o f gillrakers. A series
o f small triangular scales above axil o f pectoral fin; hind part o f body scales perforated. A black spot behind gili
cover, usually followed by up to 10 spots along flank. D eep-bodied Sardinella species have no notch in m idline o f
upper jaw and no spots along flank; species o f Tenualosa
have no fronto-parietal
striae, more or less straight
gillrakers on inner arches and no perforations on scales; gizzard shads (N em atalosa, A nodontostom a)
have an
in f e rio r m o u th . See C L U P H ils 1, F ishing A rea 51, also 57, 71.
G eographical D istrib u tion
Indo-W est Pacific
(probably all coasts o f Indian Ocean, from G u lf o f Oman
and G u lf o f A d e n south to D urban and M adagascar,
across the Bay o f B engal, G u lf o f Thailand, Java Sea
and north to Hong Kong and east to Papua N ew Guinea
and p o ss ib ly fu rth e r).
H ab itat and B iology
M arine, pelag ic, but
entering estuaries and able to tolerate quite low sali­
nities (7 ° o o ) . Feeds chiefly on phytop lan k ton (m ainly
diatom s, also dinoflagellates),
but also copepods, m o llu s c a n an d c r u s ta c e a n la r v a e , p ra w n s , a m p h ip o d s and
p olychaetes
(G odavari
estuary,
B abu
R ao,
1966).
Spawns (at least in Godavari estuary) around February
(B abu R ao , lo c, c it .).
Size : To 24.4
15 to 18 cm.
cm standard length, usually
Interest to Fisheries :
Apparently
Godavari estuary, eastern coast o f India).
L ocal N am es :
Literature :
does
about
not
f o r m large
schools,
but enters
artisanal
fisheries
(e.g.
in
KEN YA : M akrange, P aw ah.
W hitehead (1973b - refs); B abu Rao (1966 - biol. in G odavari estuary).
click for n e x t p a g e
c lic k fo r p r e v io u s p a g e
222
Tenualosa Fowler,
C LU P T enu
1934
Tenualosa Fow ler, 1934, P ro c .A cad, nat. Sei.Phil ad., 85:246 (type: A losa reevesii R ichardson). M acrura
F ow ler, 1941, B ull.U .S .n atn .M u s. (100):626 (type: Clupea kelee Cuvier:
genus w rongly a ttrib u ted to van Flasselt,
1823 by Fowler). Hilsa (part): virtually all authors included the species o f Tenualosa in H ilsa until the latter was
recently restricted to IT kelee by W ongratana (1983:385).
D iagnostic Features : M oderate or large herring-like fishes (to 50 cm standard
length in one species), m oderately elongate but com pressed, w ith a keel o f s c u te s
along belly. Top o f head w ithout num erous longitudinal striae (fronto-parietal striae):
upper ja w w ith a distinct m edian notch. G illrakers fine and num erous, those on the
inner arches more or less straight, not curled outward. Pelvic fin r a y s i 7: anal fin
short, well behind dorsal fin base. Scales m o d erate, 37 to 47 in lateral series, hind
part w ithout perforations. R esem bles H ilsa, w hich has num erous striae on top o f head,
has the g illr a k e r s on the inner arches curled outw ard and perforations on scales:
gizzard shads (N em atalo sa, etc.) have an inferior m outh and m ost have a filam entous
last dorsal finray. O ther clupeids have no notch in the upper jaw .
top 0f ileacj
Biology, H abitat and D istribution : A nadrom ous or riverine shads o f the In d o -W e st Pacific region, from
“the G u lf’ to China.
Interest to Fisheries : A ll contribute to artisanal fisheries, but H ilsa ilisha and to a lesser extent H. toli
are o f considerable im portance (total catch for 1983 was 8 807 tons - recorded as Hilsa ilisha and H. toli).
Species :
T
T
T
T.
T
W ongratana (1980, 1983) reco g n ized 5 species:
ilisha (H am ilton-B uchanan, 1822), northern Indian Ocean
m acrura (Bleeker, 1852), Indonesia
reevesii (R ichardson, 1 8 4 6 ), China to A ndam an Sea
thibaudeaui (Durand, 1940), M ekhong River
toli (Valenciennes, 1847), India to Indonesia.
Tenualosa ilisha (H am ilton-B uchanan,
1822)
CLUP T enu 1
F o r m e r ly CLU P H ils 2
Clupanodon ilisha H am ilton-Buchanan, 1822, Fishes o f G anges:243. 382, pi. 19, fig. 73 (Ganges estuaries).
Synonyms : Clupea palasah Cuvier, 1829:320 (on
(com piled): H ilsa ilish a:Rosa & Pillay, 1963: unpaged
Idem . 1967:91 (types o f palasah): Idem . 1973:199, fig.
1980:164, pis 110, 111 (revision; W hitehead & Bauchot,
FAO Names
Palasah o f R ussell, 1803): M acrura ilisha :Fowler, 1941:633
(synopsis, biol.): W hitehead, 1965:134, fig. 9 (revision!:
27 (key, synonym y, refs):
Tenualosa ilisha - W ongratana,
in press (palasah).
223
D iagnostic Features : Body m oderately deep, com pressed, belly w ith 30 to 33 scutes. Head length 28 to
32% o f standard length; distinct m edian notch in upper jaw . G illrakers fine and num erous, about 100 to 250 on
low er part o f arch. C audal fin m oderate, 25 to 31% o f standard length. A dark blotch behind gili opening,
followed by a series o f small spots along flank. R esem bles T.to li, w hich has a shorter head (22 to 27% o f standard
length), a longer caudal fin (31 to 34% o f standard length), fewer gillrakers (60 to 100) and no spots along flank,
also scutes 28 to 30. The notched upper ja w distinguishes it from other sim ilar clupeids, except H ilsa kelee, w hich
has num erous longitudinal striae on top o f head. See CLUP Tenu 1, Fishing A rea 51; also, CLUP H ils 2, Fishing
Areas 57, 71.
G eographical D istrib u tion
N orthern part o f
Indian O cean (from “the G u lf ’ eastw ard to B urm a,
including w estern and eastern coasts o f India, also 40„
rivers).
H abitat and B iology
M arine, pelagic and
schooling in coastal w aters, euryhaline, anadrom ous,
ascending rivers for as much 1200 km, but usually about
50 to 100 km. Feeds on plankton, m ainly by filterfeeding, but apparently also by grubbing on muddy
bottom s;
diatom s,
protozoans,
crustaceans,
m olluscs
and tunicates are recorded, as well as young Tenualosa 2oo
(P illa y & R osa, 1963). Breeds mainly in rivers, in some
cases far up (about 400 km up the M ahanadi system, and
even to A gra and Delhi or over 1000 km up the G anges),
but elsew here only to about 50 km or less (younger 40°
fishes may breed in the tidal zone o f rivers). In some
rivers the m igration is restricted by barrages; there
40°
60°
80°
100°
120°
140°
160°
180°
some evidence that H ilsa far up the G anges and other
large rivers, although migrating upstream to spawn, are perm anent river populations that do not descend to the
sea. The m ain breeding season is during the southw est monsoon, w ith a shorter season from January to February
or March.
Size :
To 60 cm standard length, commonly to 35 to 40 cm.
Interest to F isheries
The m ost im portant o f the Indo-P acific shads, w ith a total catch in 1983 o f
6 277 tons (W est Pakistan 5 990 tons, K uwait 287 tons). The fishes are mainly caught w ith traps, fishing w eirs and
drifted or fixed g illn e ts in estuaries and rivers during the upstream spawning migration; fisherm en also use Seine
nets, bag nets, clasp nets and cast nets. The esteem in w hich the hilsa is held is reflected in Sanskrit and Bengali
literature, where the fish is described as
m atsyaraja (king o f fishes) and it is said that Illisah jitapiyusah (hilsa
su rpasses nectar).
L ocal Nam es
BURM A: N ga-tha-lauk; INDIA: Chaksi (N arbada R iver), H ilsa (B engali, H indi), Hilii
(A ssam ), Ilish, Ilisha (Bengal, O rissa), O olum, etc. (Tam il), Palasah (Telugu), Pala, Palo, Pulla (Sindhi), Paliyah,
P aluva,V alava (M alayalam ), Palla (M arathi), Paliya (C anarese), H ilsa, K oira (juveniles, W est B engal), Jodi
(juveniles, Chilka Lake); IRAN: Soboor IRAK: Shour; PAKISTAN: Ilish, Jatka, Palla.
L iterature : Pillay
references to biology).
& R osa
(1963
- best sum m ary
o f biology,
fishery);
W hitehead
(1973
- additional
R em arks : A uthors have (probably correctly) suspected that individual populations are often isolated and
that distinct races exist (reviewed by Pillay & Rosa, 1963). No overall study has yet been made the morphological
characteristics o f the various ‘races’.
T en u alosa
CLU P T enu 3
m acru ra (B leeker, 1852)
Form erly CLUP H ils 3
A lausa m acrurus Bleeker, 1852:31 (on macrura o f Kuhl & van Hasselt, 1823 - Jakarta, Java).
Synonyms : M acrura m acrura:Fowler, 1941:632 (com piled); H ilsa m acrura:W hitehead, 1965:141
W hitehead et a h ,1966:86 (neotype for m acrurus); T enualosa m acrura - W ongratana,1980:162, pis
(revision).
FAO N am es :
En - Longtail
shad.
(revision);
106, 107
224
D iagnostic Features : Body
m oderately deep, com pressed, belly w ith 30 to 31 sc u te s . Head length 22 to
25% o f standard length; a distinct median notch in upper jaw . G illrakers fine but not num erous, 60 to 75 on low er
part o f arch (barely increasing after 10 cm standard length). Caudal fin long, 40 to 42% o f standard length, the
lobes long and pointed. No series o f dark spots along flanks. Resem bles 17 to li, w hich has a longer head (25 to
27% standard length), but shorter tail (only 31 to 34% standard length) E reevesii has many m ore low er
gillrakers (80 to 250) and a larger head (27 to 33% standard length). The notched upper jaw distinguishes it from
other sim ilar clupeids, except H ilsa kelee, w hich has num erous longitudinal striae on top o f head, also spots along
flank. See CLU P Hils 3, Fishing Areas 57, 71.
G eographical D istribution : M alaysia and IndoK"rf’
nesia (Java Sea and Sarawak, also affluent rivers).
40°
^
H abitat and B iology
M arine, pelagic and
schooling in coastal w aters, euryhaline, anadrom ous,
ascending rivers to breed. Presum ably its biology is
sim ila r to th a t o f E ilis h a , b u t th e fe w e r g illra k e rs
suggest that it takes larger food organisms.
Size :
To 52 cm standard length.
Interest to F isheries : P erhaps m ore im portant
locally than the records imply. Indonesian records (as
Hilsa to li) gave a total o f 2 530 tons in 1983.
Local Names
Trubuk.
Literature :
IN D O N ESIA , M ALAYSIA:
3*
Trubu,
There seems to be no modern study o f the biology o f this species.
Tenualosa reevesii (R ichardson,
CLU P Tenu 4
1846)
Alosa reevesii R ichardson, 1846, Ichthyol.China Japan:305 (China, in fact C anton or Macao).
Synonym s
? C lupea sinensis Linnaeus, 1758:319 (identity
M acrura reevesii:Fow ler, 1941:630, fig. 17 (China); C hu, Tchang
reevesii :W hitehead, 1965:139 (revision); Idem , 1966:30, pi. ‘2, fig. 3
1970:208 (type o f reevesii, listed); Tenualosa reevesii - W ongratana,
FAO Names :
E n - R eeves shad.
uncertain, best considered a nom en dubium );
& C hen, 1963:101, fig. 76 (C hina); H ilsa
(type o f reevesii, the Reeves drawing); Idem ,
1980:163, pi. 108, 109 (revision).
225
AA
Diagnostic Features : B ody m oderately deep, com pressed, belly w ith 31 to 32 scu tes. Head large, its
length 27 to 33% o f standard length; a distinct m edian notch in upper jaw . G illrakers fine and num erous, 80 to
250 on low er part o f arch (increasing w ith size o f fish), each raker w ith scattered asperities. C a u d a l fin
m oderate, 25 to 31% o f standard length. A dark blotch behind gili opening and a series o f spots a lo n g flanks.
R esem bles 17 to li, w hich has a sm aller head (25 to 27% o f standard length), but longer c a u d a l fin (31 to 34% o f
standard length) and only 60 to 75 gillrakers; 17 m acrura o f Indonesia has a m uch longer tail (40 to 42% o f
standard length);
17 thibaudeaui o f Thailand has more gillrakers (more than 250 from 10 cm standard length). The
notched upper jaw distinguishes it from other sim ilar clupeids, except. H ilsa k elee.
w hich has num erous
longitudinal striae on top o f head.
Geographical Distribution
C hina
(to
about
3 0 ° N ) and possibly southward into South China Sea, but
records uncertain. Two specim ens from eastern part o f
Indian O cean (P huket Islan d , A ndam an Sea - see
W ongratana,
1980:164).
The species thus
overlaps
17 toli in range.
H abitat and B iology
M arine, pelagic and
schooling in coastal w aters, euryhaline, anadrom ous,
a sc e n d in g riv e rs to b ree d . M o re d a ta n e ed ed .
Size :
To at least 50 cm sta n d a rd le n g th .
Interest to Fisheries :
C a tc h e s n o t
b u t p ro b a b ly o f so m e im p o rta n c e in C h in a.
Local Names .
Literature :
C H IN A :
Sam
la i,
S an
r e p o r te d ,
le.
6o°
40°
so°
io o °
120o
1400
1600
1800
There appear to be no recen t studies.
R em arks
The P huket Island specim ens (A ndam an Sea)
eastw ard o f the closely related
17 ilisha (Indian Ocean only).
suggest that
17 reevesii
continues
CLUP
T en ualosa thibaudeaui (D urand, 1940)
the
range
T enu
5
thibaudeaui
-
C lupea (A losa) thibaudeaui Durand, 1940, Inst, océanogr.Indochine, (note 36):6, pi. 1 (Phnompenh).
Synonym s
H ilsa k an ag u rta: Taki, 1974:47,
W o n g r a ta n a , 1980:166, pis 112, 113 re v isio n ).
F A O Names :
En - L ao tian
shad.
fig.
48
(Laos
-
m isidentified);
Tenualosa
D iagnostic F eatures : B ody m oderately deep, com pressed, belly w ith 28 to 30
scutes. Head large, its
length 30 to 33% o f standard length; a distinct m edian notch in upper jaw . G illrakers fine and very num erous, 204
to 316 on low er part o f arch (in fishes o f 8.5 to 22 cm standard length, increasing w ith size o f fish); w ith mucosal
buds and not asperities on upper edges o f rakers. Caudal fin m oderate, about 25 to 30% o f standard length. A
dark blotch behind gili opening and a series o f spots along flank. Closely resem bles the Indian Ocean T. ilisha,
w hich has few er gillrakers (about 100 to 250) and 12 reevesii (probably present in South China Sea), w hich also has
few er gillrakers (about 80 to 250); the gillrakers increase w ith size o f fish,but even at 10 cm standard length
thibaudeaui has more than 250 rakers. The notched upper ja w and spots along flank distinguish it from any
sim ilar clupeids where it occurs, except Hilsa kelee, which has numerous longitudinal striae on top o f head.
G eograph ical
D istrib u tio n
M ekong
R iver
system (at Nongkai and at U bon-rajthani on M ooí River,
T hailand;
also
L uang
P rabang,
V ientiane,
P akse, 4oo
H atsalao, Tha N gon and Tha Bo, Laos; these localities
are up to nearly 2000 km from the sea, e.g. Luang
Prabang at about 20° N).
20°
H abitat and B iology : R iverine (at least judging
from the distance up the M ekong River). A ripe male
o f 12.7 cm w as recorded at
N o n g k ai, T hailand
(Wongratana, 1980:167). M ore data needed.
Size : To 26 cm standard
length (fide D urand,
1940 - specim ens lost; a fish o f 21.7 cm exam ined by
Wongratana).
°°
20
40°
Interest to Fisheries :
Perhaps o f local interest.
40°
Local Names :
Literature :
60°
80°
100°
120°
140°
160°
180°
LAOS: Pa mak pang.
See synonymy.
R em arks : I f truly confined to freshw aters, then resem bles the populations o f R ilisha in the upper parts o f
the Ganges, which are believed not to migrate back to the sea.
T en ualosa to li (Valenciennes, 1847)
C LU P Tenu 2
F o rm e rly C L U P Hils 4
Alausa toli Valenciennes, 1847, H ist.nat.poiss.. 20:435 (Bombay, Pondicherry).
Synonyms : A lausa argyrochloris V alenciennes,! 8 4 7 :4 4 0 (B om bay, but not M auritius specim ens); A lausa
ctenolepis B leek er, 1852:74 Jakarta, M untok, Singapore); M acrura sinensis:Fowler, 1941:631 (com piled, the name
sinensis considered to derive from C lupea
sinensis L innaeus, 1758:319, but that species probably com posite;
F o w ler’s C hina references w ill be R reevesii or for Thailand perhaps jR thibaudeaui); H ilsa to l i:W hitehead.
1965:143, fig. 10 (revision); W hitehead et ah, 1966:85 (type o f cte n o le p is);
W hitehead, 1967:93 (types o f to li,
argyrochloris); W hitehead, 1973b:200, fig. 28 (key, synonym y, refs);
Tenualosa toli - W ongratana, 1980:161, pis
104, 105 (revision); W hitehead & Bauchot, in press (types o f to l i, argyrochlorid).
227
FAO Names
En - Toli shad.
D iagnostic Features : Body m oderately deep, com pressed, belly w ith 28 to 30 scutes. Elead length 25 to
27% o f standard length; a distinct m edian notch in upper jaw . G illrakers fine but not num erous, 60 to
100 on
low er part o f arch (barely increasing after 10 cm standard length). Caudal fin short, 31 to 34% standard length.
A t m ost, a dark diffuse mark behind gili opening, but no other spots on flank. Resem bles E ilisha, w hich has a
longer head (28 to 32% standard length), shorter c a u d a l fin (25 to 31% standard length), more gillrakers (100 to
250, increasing in larger fishes) and spots along the flank, also scutes 30 to 33. Other Tenualosa species also have
more than 100 gillrakers in fishes over about 10 cm standard length. The notched upper jaw distinguishes it from
other similar clupeids, except Elilsa kelee, w hich has num erous longitudinal striae on top o f head and spots along
flank. See CLUP Tenu 2, Fishing Area 51; also CLUP H ils 4, Fishing Areas 57, 71.
G eographical D istribution
India (eastern and
w estern coasts, also rivers) to Java Sea and South China
Sea (Jav a,
T hailand, thus
o v erlap p in g
range
of
E m a c ru ra ).
H ab itat and B iology
M arin e, pelagic and
schooling in coastal w aters, euryhaline and perhaps
anadrom ous,ascending rivers to breed (but in some
areas fishery w orkers claim that it does not). P resu­
mably its biology is similar to that o f
E ilisha, b u t the
few er gillrakers suggest that it takes larger food orga­
nism s. M ore data needed, especially since it is not
alw ays distinguished from T. ilisha, especially at j u v e ­
nile stages.
Size :
To 50 cm standard length.
Interest to Fisheries : N ot alw ays distinguished
from T. ilisha in catches. C aught w ith the same gear as
E ilisha (g illn e ts , seines, etc.).
Local Names :
INDIA:
C handan-ilish (C alcutta), K ajli-ilish.
Literature : As given in synonymy.
228
G udusia Fow ler, 1911
C L U P Gud
G udusia Fow ler, 1911, Proc.A cad.nat.Sei.Philad: 207
incorrectly by some authors as G adusia.
(type: Clupanodon
chapra
Flamilton-Buchanan).
Given
D iagnostic Features : M oderate-sized herring-like fishes (to about 16 cm standard length), body quite deep
and com pressed, w ith a keel o f scutes along belly. Snout shorter than eye; upper ja w w ith a distinct m edian
notch. G illrakers fine and num erous, increasing w ith size o f fish (100 to 280 in specimens o f 4 to 16 cm standard
length). Pelvic fin ra y s i 7, its insertion usually ju st before dorsal fin origin; anal fin short, well behind dorsal fin
base. Scales sm all, the row s som ew hat irregular except on upper part o f body, 77 to 91 in lateral series.
R esem bles Tenualosa, w hich has few er scales (37 to 47), evenly arranged, and the snout usually longer than eye.
G iz z a rd shads have an inferior m outh and m ost have a filam entous last dorsal f i n r a y . O ther clupeids have no
notch in the upper jaw .
Biology, Habitat and Distribution :
Interest to Fisheries :
Species !
G.
G
C o n trib u te
R iverine
to
shads o f In d ia to Burm a.
a rtis a n a l
fis h e rie s
in
riv e rs.
A c c o rd in g to W o n g ra ta n a (1980) only 2 species:
c h a p ra (H a m ilto n -B u c h a n a n , 1822), In d ia n riv e rs
v a rie g a ta (D a y , 1869), B u rm e se riv e rs.
G udusia chapra (H am ilton-B uchanan, 1822)
C lupanodon chapra H am ilton-Buchanan, 1822, Fishes o f G a n g e s :2 4 8 . 383
C L U P Gud 1
(upper Ganges).
Synonyms : Clupanodon cagius H am ilton-Buchanan,1822:250, 383 (Ganges in northern Bihar); Clupea indica
Gray, 1834:pl. 91, figs 1, 2 (India, on H ardwicke draw ing); Clupea champil Gray, 1834:pl. 91, figs 5, 6 India, on
Hardwicke drawing); A lausa m icrolepis V alenciennes, 1847:439 (Bengal ; Clupea suhia Chaudhuri, 1912:436, pi. 38,
fig. 1 (G andak R iver at Saran, B ihar); G udusia godanahiai Srivastava, 1968:6, fig. 4a, b (G orakhpur, U ttar
Pradesh); Gudusia chapra - Fow ler, 1941:634 (Ganges); W hitehead, 1965:148, fig. 11 (revision); Idem . 1967:95
(types o f m icrolepis) ; Idem , 1973b:201, fig. 29 (key, synonym y, refs); W ongratana, 1980:167, pis 114, 115
(revision); Jayaram , 1981:40, fig. 25 (key only).
FAO Names :
En - Indian river shad
Diagnostic Features : B ody fairly deep, its depth 31 to 39% o f standard length; 26 to 29 scutes along belly.
A single triangular pectoral axillary
scale; depressed tip o f dorsal fin to behind vertical from anal fin origin.
Hind margin o f scales smooth. D ark blotch behind g ili opening, often followed by a series o f spots along flank.
R esem bles (L v a rieg a ta o f Burm ese rivers, w hich is much deeper (depth 42 or 43% o f standard length), has more
scutes (29 or 30, the depressed dorsal fin tip not reaching vertical from anal fin origin, and the hind m argin o f the
scales toothed. Species o f Tenualosa have larger scales (only 37 to 47 in lateral series; cf. 77 to 91).
229
G eographical D istribution : R ivers o f India and
B angladesh affluent to the Bay o f B engal (chiefly the
Ganges and B rahm aputra systems and the M ahanadi River
o f Orissa).
H abitat and Biology : Riverine in middle and upper
reaches. M ore data needed.
Size :
To 15 cm standard length.
Interest to F ish eries
C ontributes
artisanal fishes, but catches not recorded.
to
riverine
L ocal N am es : L iterature : W hitehead (1973b
papers that include notes on biology).
-
reference
to
5
C LU P G ud 2
Gudusia variegata (Day, 1869)
Clupea variegata D ay, 1869, P roc.zool.S oc.L ond.:623
(Irraw ady River, Burma).
Synonyms : G udusia variegata - Fow ler, 1941:635 (on Day): W hitehead, 1965:150, fig. 12 (revision): Idem .
1973b:201 (in key only): W ongratana, 1980:169, pis 116, 117 (revision): Jayaram , 1981:40 (key only).
F A O N am es :
E n - B u rm ese riv er shad.
D iagnostic Features : B ody deep.its depth 42 or 43% o f standard length: 29 or 30 scutes along belly.
Three small triangular pectoral a x illa r y scales: depressed tip o f dorsal fin not reaching to vertical from anal fin
origin. H ind m argin o f scales toothed. D ark blotch behind gili opening, follow ed by a series o f spots along upper
flank. R esem bles Ck chapra o f Indian rivers, w hich is more slender (depth 31 to 39% o f standard length), has
few er scutes (26 to 29), the depressed dorsal fin tip surpassing the anal fin origin, and the hind border o f the scales
smooth. Species o f Tenualosa have larger scales (only 37 to 47 in lateral series: cf. 77 to 91).
c lic k for n e x t p a g e
c lic k fo r p r e v io u s p a g e
230
Geographical Distribution :
R iv ers
(chiefly the Irraw addy, but perhaps others).
Habitat and Biology :
R iv e rin e in
upper reaches. More data needed.
Size :
m id d le and
To about 16 cm standard length.
Interest to Fisheries :
C o n trib u tes
artisanal fisheries, but catches not recorded.
L ocal
o f B u rm a
N am es
Literature :
to
riv e rin e
:-
See under synonyms.
Remarks :
R ath er few sp ecim en s seem to have
been studied (V ongratana, 1980, saw only three).
2.2.5
S U B F A M IL Y DOROSOMATINAE
F A O Names :
E n - G izzard shads.
D iagnostic Features : M oderate-sized herring-like fishes (to about 30 cm
standard length or a little more): fully scu ted along belly, sc u te s also present on
back before dorsal fin in some
(C lu p a n o d o n ). M outh inferior or subterm inal,
som etim es term inal, snout usually projecting:upper ja w not evenly rounded in
front, but w ith a distinct median notch into w hich the sym physis o f the low er jaw
fits; no teeth. G illrakers fine and num erous; a pair o f pharyngeal pouches
above 4th gili arch, apparently for collecting food seived by gillrakers. Dorsal
fin at about m idpoint o f body, last dorsal fin ra y filam entous and long (except in
the Indo-Pacific G onialosa and A nodontostom a)' anal fin m oderate, up to 38
finrays; pelvic fin under or a little before dorsal fin o rig in , w ith i 7 fin ray s.
Scales usually well attached, usually 38 to 55 in lateral series (but 43 to 71 scales
in G onialosa and up to 86 in some D orosom a).
Stom ach m uscular, gizzard-like.
A dark spot often present behind gili opening, in some species follow ed by a
series o f spots.
mouth
inferior
B iology, H abitat and D istrib u tion
The D orosom atinae are either m arine, pelagic and apparently
anadrom ous, or estu arin e or purely flu v ia tile, but m ore data is needed to show w h eth er all species spaw n in
freshwater. Food, together w ith mud and other m aterial grubbed from the bottom , is filtered by the gillrakers
and believed to be concentrated in the pharyngeal pouches, thereafter being e v e rte d as a bolus and passed down
the pharynx to the m uscular stomach.
oesophagous
loop
Interest to Fisheries :
G izzard shads con trib u te
rather little to fisheries (perhaps about 10 000 tons a
1st flexure
year), but may be locally important.
Rem arks : Separation o f the gizzard shads from the
shads (subfam ily A lo sin a e ) may be artificial; a gizzard is
found in the shads Brevoortia and Ethm idium , and a more
or less m uscular stom ach occurs in H ilsa.
Gudusia and
E thm alosa (as well as in the
clupeine
O pisthonem a).
N evertheless, only the gizzard shads have a unique
feature: the intestine is doubled into a characteristic loop
(the so-called ‘third prim ary fle x u re ’), either a simple
loop, or a loop that is coiled or doubled-back on itself
(N elson & R othm an, 1973).
There are 5 genera (1 A tlantic coasts and drainage
o f N orth and central A m erica, 4 Indo-W est Pacific coasts
and drainage) and about 22 species, the largest genus
being N em atalo sa (9 species, but more to be described).
2nd flexure
coiled loop
stomach
anus
flexures o f intestine
231
Key to the Genera
N ew W orld only; an terio r (first) supram axilla present (but thin and scale-like);
last dorsal finray p ro d u c e d , fila m e n to u s
D orosom a
( F i g - 1 ) ......................................................
lb.
Indo-P acific o n ly ;
m axilla
2a.
filam entous
/
no a n te r io r su p ra-
Last d o rsa l finray p ro d u c e d , f i la ­
m ento us
D orosom a
3a.
3b.
2b.
Fig.
4a.
Post-pelvic scutes 11 or
12; d o rsa l scu te s p re ­
s e n t...............................C lu p a n o d o n
4b.
Post-pelvic scutes 14 to
16; no dorsal scutes . . Konosirus
G illra k e rs o f firs t a rc h n o t
more than h a lf length o f cor­
responding
gili
filam en ts;
m o u th in f e rio r , u p p e r ja w
cu rv e d d o w n w a rd (F ig . 2b);
p re -d o rsa l sc a le s p a ire d and
overlapping in m idline (Fig.
3 b ) ............................................... N em a ta lo sa
m axilla
straig h t
a.
m axilla
turned dow n
C lupanodon
b.
b
N e m a ta lo s a
Fig. 2
pre-dorsal scales
not overlapping
L a s t d o rs a l finrays n o r m a l, n o t
filam entou s
5a
5b.
1
G illr a k e r s o f f i r s t a rc h at
le a s t 3/4 le n g th o f c o r r e s ­
ponding gili filam ents; mouth
su b term in al,
u p p er
ja w
straig h t (Fig. 2a); p re -d o rsa l
sc a le s p a ire d , b u t n o t o v e r­
lapping in midline (Fig. 3a)
A median series o f pre-dorsal
scales (Fig. 4); 38 to 45 sca­
les in lateral series (usually
40 to 43); marine ....... A nodontostom a
C lupanodon. K onosirus
pre-dorsal scales
o verlapping
P aired
pre-dorsal
scales,
overlapping in midline; 43 to
71 sc a le s in la te ra l se rie s;
fresh w ater, riv ers a fflu en t t.o
Bay o f B e n g a l...............................Gonialosa
b
N em atalosa
m edian pre-dorsal scales
•••••io
A n o d o n to sto m a
F
232
C LU P D oros
D orosom a R afinesque, 1820
D orosom a R afinesque, 1820,
W estern R eview & M ise.M ag ., 2 (3, A p ril): 171 (see F o w ler, 1945:6, 8) (type:
D o ro so m a n o tata R afin e sq u e = M eg alo p s c e p ed ian a L eS u eu r). C h ato essu s C u v ier, 1829,
Règne a n im a l, 2nd ed.,
2:320 (in part; restricted to M egalops cepediana by V alenciennes, 1847). Signalosa Everm ann & K endall, 1898,
B ull.U.S.Fish Com m n, 17:127 (type: Signalosa atchafalayae = M eletta petenesis Günther).
D iagnostic Features : M edium -sized gizzard shads (to 35 cm standard length, usually to about 20 to 25 cm),
the body com pressed and m oderately deep in some. Snout projecting, rounded, m outh small, term inal or inferior;
upper jaw w ith a thin, scale-like anterior (first) supra-m axilla, low er ja w included when m outh closed. G illrakers
fine and numerous. L ast dorsal fin ray filam entous (except in young); anal fin com paratively long, w ith 17 to 38
finrays, its origin below or ju st behind base o f last dorsal fin ray . Scales m oderate (less than 50 in lateral series)
and regular, or num erous (m ore than 50) and som ew hat irregular. A dark spot
behind gili opening in m ost
specimens.
B iology, H abitat and D istribution : C oastal saltw ater and brackishw ater to freshw ater, the latter either
for spawning or as a perm anent habitat. All species are filterfeeders. Found associated w ith the A tlantic drainage
o f North and central America and the Pacific drainage o f central America.
Interest to Fisheries :
Species :
O f lim ited local interest.
M iller (1960) recognized five species in two subgenera, accepted by N elson & Rothm an (1973):
Subgenus Dorosoma (mouth inferior; scales more than 50)
D.
D.
D.
D.
anale Meek, 1904, drainage o f western G ulf o f Mexico
cepedianum (LeSueur, 1818), Atlantic and G ulf drainage o f North and central America
chavesi Meek, 1907, Nicaraguan lakes
sm ithi Hubbs & Miller, 1941, Pacific drainage o f Mexico.
Subgenus Signalosa (mouth terminal; scales less than 50)
D . petenense (G ünther, 1866), Atlantic and G ulf drainage o f North and central America.
D o m so m a anale M eek, 1904
C L U P D oros 2
D o ro s o m a anale M eek, 1904, Pubis Field Mus., (93)( z o o l.S e r.). 5:93, fig. 26 (El H ule = R io Papaloapán,
M exico).
Synonym s:
D o ro so m a anale - Miller, 1950:395
1973:170, pi. 11B (photo) (bibliography).
F A O Names :
(revision);
En - M exican river gizzard shad.
Idem , 1960:373 (key only); N elson & Rothm an,
23 3
D iagnostic Features : Body m oderately deep, its depth 34 to 40% o f standard length, belly w ith 18+ 10 or
11 scutes. M outh sm all, in ferio r; low er jaw short, about 8 to 10% o f standard length. Last dorsal finray long,
about equal to distan ce snout tip to ab o u t m id -p ecto ral fin; anal fin long, w ith 29 to 38 finrays. Scales sm all,
som ewhat irregular, 70 to 82 in lateral series. A dark spot behind gili opening. Resem bles D. cepedianum , whose
southward range it continues, but that species has few er scales (52 to 70, usually 58 to 65) and a shorter anal fin
base (equals anal fin origin to hind third o f pectoral fin;
cf. to fro n t th ird in Ef anale); D. p eten e n se has few er
scales (41 to 48).
Species o f O p isth o n e m a a lso h av e a
filam entous last dorsal fin ray and dark spot behind gili
opening, but anal fin short and well behind dorsal fin base.
Other clupeids lack dorsal filament.
G eographical D istribution
M exico (Rio P a p a loapan in southern V eracruz and O axaca) southw ard to
northern G uatem ala (Rio U cum acinta basin).
H abitat and B iology : R iverine, apparently not yet
recorded in brackishw ater. M ore data needed.
Size :
Interest
in te re s t.
T o about 20 cm standard length.
to
F ish eries
: P erhaps
of
sm all
local
L o c a l N a m es : Literature : Only six references found by N elson &
R o th m a n (1973), none dealing w ith the biology o f the
species.
D orosom a cepedianum (L eSueur, 1818)
C L U P D oros
1
M e g a lo p s c e p e d ia n a LeSueur, 1818, J.A cad .n at.S ci.P h ilad .. 1:361 ( ‘P h ila d e lp h ia ’ = resid en ce o f L eSueur;
exact locality not known, presum ably Chesapeake and D elaw are Rays).
Synonyms :
C lu p ea h eteru ra R afin e sq u e , 1818:354 (O hio R iv er); D o ro so m a n o tata R afin e sq u e , 1820:172
(falls o f O hio R iver ; C h a to e ss u s e llip tic u s K irtla n d , 1838:169, 195 ( n o m e n n u d u m ); M e g a lo p s b im a c u la ta
V a le n c ie n n e s , 1847:104 (nomen n u d u m , in synonym y o f cep e d ia n u s); C hatoessus in so ciab ilis A b b o tt, 1861:365
(pond 2 m iles below Trenton, N ew Jersey); D o ro so m a c e p e d ia n u m e x ile Jo rd a n & Gilbert, 1883:585 (G alveston
Bay, Texas); D orosom a lacepedi B erry, 1958:318 (age and growth);
D orosom a cepedianum - M iller, 1950:392
(revision); Id em , 1960:373, fig. 1, also 2-4 (em bryos, larvae) (revision); FW NA, 1964:444, fig. 116 (review ); N elson
& R othm an, 1973:170, pi. Ile (p h o to ) (b ib lio g rap h y from 1960, rath er co m p lete);
W h iteh ead & Bauchot, in press
(types o f cepediana, status o f bim aculata).
F A O Names :
En - A m erican gizzard shad.
234
D iagnostic Features : Body m oderately deep, its depth about 30 to 40% o f standard
length, belly w ith 17 to
20 - 10 to 14 s c u te s . M outh sm all, inferior; low er ja w short, about 9 to 12% o f standard
length. L ast dorsal
fin ra y long, about equal to distance from snout tip to m id-pectoral fin or beyond; anal fin long, w ith 25 to 36
fin ra y s . Scales sm all, som ewhat irregular, 52 to 70 in lateral series (usually 58 to 65). A dark spot behind gili
opening. R esem bles D. a n a le , w hich continues its southward range, but that species has more scales (70 to 82) and
a longer anal fin b a s e (e q u a ls anal fin origin to front third o f pectoral fin; cf. to hind third in D. c e p ed ian u m );
D. petenense has few er scales (41 to 48). Species o f O pisthonem a also have a filam entous last dorsal fin ra y and
dark spot behind gili opening, but anal fin not longer than head
and well behind dorsal fin base. O ther clupeids
lack dorsal filam ent. See CLUP
Doros 1, Fishing A rea
31.
Geographical Distribution : A tlantic and G u lf o f
M exico drainage o f N o rth and central A m erica (so u th ­
eastern S outh D ak o ta and cen tral M in n eso ta, the G reat
Lakes drainage, i.e. in Lake Erie, southern parts o f Lakes
H u ro n and M ic h ig a n , L ak e O n ta rio b a sin ;
n o t L ak e
Superior; extrem e southern N ew Y ork southw ard to the
M ississippi system and sm aller rivers affluent to the G ulf
southward to the Rio P án u co , M exico.
20°
H a b ita t and B io lo g y :
M ainly in freshw ater in
large rivers, reservoirs, lakes, swamps, tem porary floodw ater pools, etc., but adults also found in brackish or
saline w ater o f estuaries or bays, preferring quieter open
w aters. Filter-feeding, alm ost entirely herbivorous; the
fo o d is stra in e d by th e n u m e ro u s fin e g illr a k e rs , th e n
presum ably transferred in a m ucus stream to the p h a r y n ­
geal pouches, concentrated and in some way ev erte d as a
b o lu s into the pharynx. Breeds near surface in fresh w ater
from late w inter (m id-M arch) thro u g h m ost o f the
sum mer (at least to about m id-A ugust) in ponds, lakes and
large rivers; eggs adhesive and sink (em bryonic and larval
developm ent figured and described by M iller, 1960).
Size :
a
°°
20°
120°
100°
80°
60°
40°
To about 35 cm standard length (20.5 inches total length recorded), usually about 20 cm.
Interest to Fisheries : Small catches only recorded (in 1983 a total catch o f only 461 tons). It is valued as
forage for various game fishes and has been used to some exent for fe r tiliz e r or cattle food.
Local Nam es :
USA:
E astern gizzard shad, G izzard shad (AFS list), Hickory shad, M ud
shad, Skipjack.
Literature : M iller (1960 - excellent review o f taxonom y and biology); N elson & R o th m a n (1973 - literature
from 1960, 158 references).
Remarks : H ybrids betw een this species and
K ru m holz (1960). The tw o species overlap over much
D orosom a ch avesi M eek, 1907
D orosom a chavesi M eek,
M anagua and N icaragua).
1907, Pubis Field
D. petenense were reported in the
Ohio R iver
o f their ranges and frequently occur together.
by M inckley &
C L U P D o ro s 3
M u s .(1 2 1 )(z o o l.S e r.). 7:112 (Laguna Jenicero, betw een Lakes
Synonym s : D o ro so m a c h a v e si - Miller, 1950:398 (revision); Idem . 1960:373 (key only); A storqui, 1971:29,
fig. 8 (descr., Lakes M anagua and N icaragua); N elson & R othm an, 1973:172, fig. 12A (photo) (bib lio g rap h y ).
FAO Names :
En - N icaragu a gizzard shad.
235
D iagnostic Features : Body m oderately deep, its depth about 34 to 40% o f standard length, belly w ith 15 to
18 + 9 to 12 scutes. M outh relatively large, inferior or subterm inal; low er ja w com paratively long, about 11 to
16% o f standard len g th , u p p er ja w reach in g to v ertical from eye cen tre. L a st d o rsa l fin ra y lo n g , b u t le ss th a n
d istan ce from snout tip to p elv ic fin base;
anal fin m oderately long, w ith 24 to 30 fin r a y s . Scales sm all,
som ewhat irregular, 72 to 86 in lateral series. A dark spot behind gili opening. No other D orosom a species found
so far south (D . a n a le o f northern G uatem ala and southern M exico has 29 to 38 anal fin ra y s ; D. s m ith i o f the
P a c ific d rain ag e o f c en tra l A m e ric a a sm a lle r m o u th ). N o o th e r c lu p e id in th e a re a re se m b le s EL c h a v e s i; T arpon
atlanticus has a dorsal filam ent, but lacks scutes, low er ja w pro jects, scales less than 50.
G eograp hical
D istrib u tion
N icarag u a
(lakes
M an agua and N icarag u a and a fflu e n t stream s or riv ers).
H abitat and B iology : R iverine and lacustrine.
M ore data needed, but presum ably feeding and breeding
sim ilar to o th er D o ro so m a species.
Size :
To
Interest
im p o r ta n c e .
18 cm standard
to
F ish eries
length.
: P erhaps
of
sm all
local
L ocal N am es
0°
L iterature :
(1 9 7 1 -d e s c r.).
M iller
(1950
-
revision);
A storqui
Rem arks : M iller (1 9 5 0 ) found differences between
h is sp e c im e n s fro m L ak e M a n a g u a an d L a k e N ic a ra g u a
and suspected that the falls betw een these lakes served to
isolate
the
M anagua population. A storqui (1971) con­
firm ed m any o f M ille r’s d iffe re n c e s, in p articu lar:
L ake M anagua: head len g th 32.6 to 3 5 .5 % , anal fin
sta n d a rd le n g th
20
°
base 24.6 to 27.1% , pre-pelvic distance 49.2 to 53% o f
Lake N icaragua: head length 29.2 to 3 2 .3 % , anal fin base 27.6 to 29.6%, pre-pelvic distance 43.7 to 49.5%
o f sta n d a rd le n g th .
H o w e v e r, n e ith e r a u th o r p ro p o s e d a su b s p e c ie s.
236
D orosom a p eten en se (G ü n th er, 1866)
M e le tta p e te n e n s is G ü n th er, 1866, P ro c.zo o l.S o c.L o n d .,
petenense by Jordan & E verm ann, 1896:417).
C LU P D oros 5
(3):603 (L ake P e té n , G u a te m a la ) ( e m e n d e d to
Synonym s
C hatoessus m exicanus G ü n th e r, 1868:409 (M exico, N ew O rleans, L ouisiana); Signalosa
atchafalayae E v e rm a n n & K endall, 1 8 9 8 :1 2 7 , pi. 7, fig. 4 (A tchafalaya R iver at M elville, L ouisiana; also
M ississippi);
Signalosa m e x ic a n a ca m p i W eed ,1925:143 (R esaca de la G uerra, on or near M edia L una Ranch,
B row nsville, Texas); S ig n a lo sa a tc h a fa la y a e v a n h y n in g i W e e d , 1925:145 (P rairie C reek , 6 m iles so u th east o f
G a in s v ille , F lo rid a ); D o ro so m a p e te n e n se - Miller, 1960:373 (key only); F W N A , 1964:448, fig. 117 (review);
N elson & R othm an, 1973:173, pi. 12C (photo) (bibliography from 1960, rather complete).
FAO Names :
E n - T h read fin shad.
D iagnostic Features : Body m oderately deep, its depth about 30 to 40% o f standard length, belly w ith 15 to
18 + 8 to 12 sc u te s . M outh sm all, term inal. L ast dorsal fin ray long, about equal to distance from snout tip to
m id-pectoral fin or beyond; anal fin relatively short, w ith 17 to 27 fin ra y s . Scales relatively large, regularly
arranged, 41 to 48 in lateral series. A dark spot behind gili opening. D istinguished from D. cepedianum and other
related species by the larger scales (more than 50 in other D orosom a species, som ewhat irregularly arranged) and
term inal m outh (cf. inferior or at m ost su b term in al in D^ chavesi o f N icaragua). Species o f O pisthonem a also have
a filamentous last dorsal finrav and dark spot behind gili opening, but anal fin not longer than head and well behind
dorsal fin base. Other clupeids lack d o rsal filament.
G eograp h ica l D istrib u tio n :
G u lf o f M exico
drainage o f N o r th a n d c e n tr a l A m e r ic a (M ississippi
system , from the O hio R iv er o f K en tu ck y and southern
In d ian a southw estw ard to O klah o m a, and south to Texas
and F lo rid a , a lso riv e rs a ro u n d th e G u lf to n o rth e rn
G uatem ala;
also Belize River, B ritish Flonduras). Inva- 60„
s io n o f th e O h io R iv e r b a s in a p p e a r s to h a v e b e e n
relatively recent; the species has been planted in V irgi­
nia, W est V irginia, G eorgia, P en n sy lv an ia, K ansas, A ri­
zona, N ev ad a, C alifo rn ia and N ew M exico (M in ck ley &
K ru m h o lz , 1960), and in tro d u ced in 1958 in to Flaw aiian 40°
w aters (Oahu, Kauai, Maui) (Brock, 1960).
H ab itat and B io lo g y :
P ela g ic , o ften sch o o lin g , 2o°
m ain ly in fr e s h w a te r in la rg e riv e rs , re s e rv o ir s , la k e s ,
swamps, tem porary floodwater pools, etc., but adults also
found in brackish or saline w ater o f estuaries and bays (up ^
to 30 °/oo salinity; ju v en iles to about 15% o). Filterfe e d in g , b u t n o t e n tire ly h e rb iv o ro u s sin c e c o p e p o d s,
cladocerans and even fish fry have been recorded; m ethod 2QO
o f feeding presum ab ly sim ilar to th at o f FT ce p e d ia n u m .
B reeds in spring and again in autum n, in open w aters near
or over plants or other objects; eggs slightly adhesive.
160°
140°
120°
100°
80°
60°
40°
237
Size :
To 18 cm standard length in southern part o f range, but only about 10 to 12 cm in northern part.
Interest to Fisheries : V alued as a food for larger game fishes, hence its introduction into other w aters;
also as a potential live bait (e.g. for tuna in Hawaii).
Local Names : Literature : M inckley & K rum holz (1960 - distribution, hybrids); M iller (i.e. FW NA, 1964 - review); N elson
& R othm an (1973 - literature from 1960, 103 references).
R em arks : M iller (1960, 1964) considered S ignalosa as m erely a subgenus o f D orosom a; M inckley &
K rum holz (1960) gave it generic status. N elson & R o th m a n (1973), the latest review ers, favoured subgeneric
status and are followed here. For hybridization w ith FF cepedianum , see that species.
D o r o s o m a smithi H ubbs & M ille r, 1941
C LU P D oros 4
D orosom a sm ithi Hubbs & Miller, 1941, Copeia, (4):232, fig. 1 (Rio Piaxtla near Piaxtla, Sinaloa, Mexico).
Synonyms : D orosom a sm ithi - M iller, 1950:401 (revision); Id e m . 1960:373 (key only); B ran so n , M cCoye &
Sisk, 1960:218 (S on o ra, M exico); A lvarez del Villar, 1970:41 (key); N elson & R o th m an , 1973:173, fig. 12B (photo)
(bibliography).
F A O Names :
E n - P acific gizzard shad.
.....
........
Diagnostic Features :
Body m oderately deep, its depth about 32 to 40% o f standard length, belly w ith 15 to
18 + 9 to 12 scutes. M outh small, inferior, low er ja w about 9 or 10% o f standard length. Last dorsal fin ray long,
but less than distance from snout tip to pelvic fin base; anal fin relatively short, w ith 22 to 29 fin ra y s . Scales
small, som ewhat irregular, 71 to 79 in lateral series. A dark spot behind gili opening. O pisthonem a species also
have a filam entous last dorsal fin ra y and a dark spot behind gili opening, but anal fin not longer than head and well
behind dorsal fin base. Other clupeids lack dorsal filament.
238
G eographical D istribution
Pacific drainage o f
northw estern M exico (Rio P iaxtla, R io del F uerte, Rio
Sinaloa system , Rio de M ocorito, Rio M uerto, R io Yaqui
system, i.e. betw een Sinaloa and Sonora).
H abitat and B iology
R iv erin e, perhaps a ls o 40
lacustrine. M ore data needed, but presum ably feeding and
breeding sim ilar to other D orosom a species.
Size :
To at le a st 14 cm stan d ard len g th , p erh ap s 20O
m ore.
Interest to Fisheries :
im portance.
P erhaps
of
sm all
local
o°
L ocal N am es : L iterature
rences).
: N elson
& R o th m a n
(1973
-
6 refe­
20
"
C LU P
Clupanodon L acepede, 1803
C lupan
Clupanodon Lacepede, 1803, H ist.n at.p o iss., 5:465 (type: C lupea thrissa Linnaeus; designated by Bleeker,
1872:112. Subsequent invalid designations were Clupeonia jussieui V alenciennes = Sardinella jussieui, by Jordan &
G ilbert, 1883:574; C lupea pilchardus W albaum = Sardina p ilchardus, by Jordan & Everm ann, 1896:422; and
M egalops oglina L eS u e u r = O pisthonem a oglinum ,
by
Jordan & S eale, 1905:771). Thrissa R afinesque, 1815,
A n a l.n a tu re :88 (type: Clupea thrissa Linnaeus since Thrissa proposed to replace C lupanodon).
D iagnostic
F eatu res
M e d iu m ­
sized m arine gizzard shads (to about
26 cm
standard
len g th ); belly
fully
sc u te d , also pre-dorsal scutes present.
M outh subterm inal; m axilla slender, not
turned dow nw ard at tip, w ith a single
spatulate
supra-m axilla.
G illrakers
fine
and
nu m ero u s (ab o u t
200
to
400,
increasing w ith size o f fish), at least 3/4
length o f corresponding gili filam ents on
first arch. L ast dorsal
fin ra y f i la m e n ­
to u s; anal fin shorter than head, w ith 22
to 28 fin ra y s . Pre-dorsal scales paired,
but not overlapping in m idline. D ark
spots on flanks. Closely resembles K o n o ­
sirus, w hich lacks pre-dorsal scutes and
has more ventral scutes (32 to 37; cf. 27
to 31 in C lupanodon).
Species o f N e m a ­
talosa have an inferio r m outh w ith its
edges flared outward.
Biology, Habitat and Distribution :
Interest to Fisheries
paired pre-dorsal scales
m axilla
p re-d o rsal scutes
See species.
: See species.
Species: N elson & R othm an (1973:164) included K onosirus punctatus in this genus, arguing that m onotypy is
infectious, thus inflating the rank o f all species w henever new differences are found. H owever, the gizzard shads
seem sufficiently well known to avoid this, and K onosirus is kept distinct here:
G thrissa (L innaeus, 1758), w estern N orth Pacific (also A ndam an Sea).
239
C lup an od on th rissa
(L innaeus,
1758)
C L U P C lu p an 1
C lupea thrissa Linnaeus, 1758, Syst.nat. 10th ed., 1:318 (nam e from O sbeck, 1757:257; diagnosis derived
from that for M ystus altus Linnaeus, 1754:26 - invalid!.
Synonyms : C lupea triza Linnaeus, 1759:251 (C h in a); C h ato essu s m acu latu s R ich ard so n , 1846:308 (V achell
specim en from Canton, lost); C hatoessus osbeckii V alenciennes, 1847:106 (China); C hatoessus haihoensis Oshim a,
1926:3 (H ainan); C lu p an o d o n th rissa - Fowler, 1941:557 (Japan, China, the Philippines, ? In d o n esia, but B om bay
incorrect); W hitehead, 1962:100 (key, diagnostic features); Id em , 1966:34, 37 (R ich ard so n ’s triza and m aculatus);
Idem , 1967:98 (types o f o sbeckii); N elson & R othm an, 1973: 169, pi. 11A (photo), map 4 (synon., descr., refs);
W o n g ratan a, 1980:170, pis 118, 119 (revision); W hitehead & Bauchot, in press (types o f osbeckii).
F A O Names :
E n - C hinese gizzard shad.
r e * ,:
'
■-
D iagnostic Features : Body m oderately deep, its depth 33 to 37% o f standard length, com pressed, belly
w ith 16 to 19 (usually 18) + 9 to 12 (usually 10 or 11) total 27 to 31 (usually 29 or 30) scutes; p re-d o rsal scutes
present, 17 to 26 (usually 20 to 25). M outh subterm inal; gillrakers o f first arch at least 3/4 length o f gili
filam ents. L ast dorsal fin ra y filam entous; anal fin ra y s 22 to 28. V ertebrae 43 to 46 (usually 44 or 45). A dark
spot behind gili open in g , follow ed by fu rth er spots on flank.
R esem b les K o n o siru s p u n c ta tu s, w h ich is m ore
slender (depth 28 to 33% o f stan d ard len g th ), lacks p re-d o rsal scutes and has m ore v entral scutes (32 to 37);
species o f N em atalosa have
an inferior m outh and gillrakers not more than h a lf length o f gili filam ents. Other
clupeids lack a dorsal filament.
G eographical D istribution : C oasts and rivers o f
n o rth w e s te rn P a c ific (C h in a to a b o u t 25°N , a p p a re n tly
south to Viet N am , but Philippine records o f Fow ler (1941)
appear doubtful); specim ens from P huket I, Thailand
(A ndam an Sea), were recorded by W ongratana (1980:171).
H abitat and Biology : Coastal and in rivers, presu­
mably tolerating brackishw ater, if not fully fresh condi­
tions. M ore data needed.
Size :
To about 26 cm standard length.
Interest to Fisheries : O f small local interest (Liu
& Shen, 1957 - Taiw an); the K orean catch in 1983 was
14 119 tons.
Local Names :
Yellow fish - Canton).
Literature:
ces).
CHINA:
Hwang yu, W ong u (=
N e lso n & R othm an (1973 - 76 referen­
240
Rem arks : The ranges o f Clupanodon thrissa and K onosirus punctatus overlap in the East China Sea (Eiong
Kong north to Taiwan); possibly the two have been confused in this area.
K o n o siru s Jordan & Snyder, 1900
C LU P K ono
K onosirus Jordan & Snyder, 1900, P r o c .U .S .n a tl.M u s ., 23:349 (type: C hatoessus punctatus Tem m inck &
Schlegel. Subsequent invalid designation Chatoessus nasus B loch = N em atalosa nasus, by Jordan, 1920) (m isspelt
K onoshirus by Jordan & Snyder, 1901:53). N ealosa H erre & M yers, 1931, Lingnan Sci.J.. (10):236 (type: Chatoessus
punctatus Tem m inck & Schlegel).
D iagnostic Features : M edium -sized marine gizzard shads (to about 24 cm standard length); belly fully
sc u te d , but no pre-dorsal scutes .M outh sub-term inal; m axilla fairly slender, not turned downward at tip, w ith a
single curved supra-maxilla. G illrakers fine and num erous (about 150 to 250, increasing w ith size o f fish), at least
3/4 length o f corresponding gili filam ents;anal fin shorter than head, w ith 19 to 27 fin ra y s . Pre-dorsal scales
paired, but not overlapping in midline. D ark spots on flanks. Closely resembles C lupanodon; w hich has pre-dorsal
scutes and few er ventral scutes (27 to 31; cf. 32 to 37). Species o f N em atalosa have an inferior m outh w ith its
edges flared outward.
Biology, Habitat and Distribution : See species.
Interest to F ish eries : See species.
Species : O pinions have been divided w hether this genus deserves recognition (see under C lupanodon), but
they are kept separate here, w ith a single species in each:
K. punctatus (Tem m inck & Schlegel, 1846), w estern N orth Pacific.
K onosirus punctatus (Tem m inck & Schlegel, 1846)
C LU P K ono 1
C hatoessus P u n cta tu s Temm inck & Schlegel, 1846, Fauna Japonica. P oiss. pt 5, inst. 13:240, pi. 109, fig. 1
(Japan, southern coast o f N agasaki).
Synonyms : C hatoessus aquosus R ich a rd so n .l846:307 (Hong Kong or Canton; R eeves specim en); N ealosa
p u n c ta ta :H erre & M y ers, 1931236 (H ong K ong); C lu p an o d o n p u n c ta tu s:F o w ler. 1941:559 (Jap an , K o rea); C hu,
Tchang & C hen, 1963:102, fig. 77 (E ast C hina Sea, revrew ; N elson & R othm an, 1973:164, fig. 10c (photo), map 4
(sy n o n ., d e s c r., re fs ); K onosirus punctatus
- W hitehead, 1962:100 (key, diagnostic features); Idem . 1966:33 (type
o f a q u o s u s ); W o n g r a ta n a , 1980:172, pis 120, 121 ( r e v is io n ); M a su d a e t ah, 1984:19, pi. 22B (c o lo u r p h o to )
(co m p iles).
F A O Names :
En - K o n oshiro gizzard shad.
c lic k for n e x t p a g e
c lic k fo r p r e v io u s p a g e
241
D iagnostic F eatures
B ody m oderately slender, its
depth 28 to 33% o f standard length, m oderately
com pressed, belly w ith 17 to 21 (usually 19 or 20) + 12 to 16 (usually 14 or 15), total 32 to 37 (usually 34 or 35)
scutes; no pre-dorsal scutes. M outh subterm inal, snout projecting slightly; gillrakers o f first arch at least 3/4
length o f gili filam ents. L ast
dorsal fin ra y filam entous; anal fin ra y s 19 to 27. V ertebrae 46 to 51 (usually 49,
also 48 or 51). A dark spotbehind gili opening, follow ed
by several lines o f dark dots along flank. Resem bles
C lupanodon thrissa, w hich has a deeper body (depth 33 to 37% o f standard length), pre-dorsal scutes and fewer
ventral scutes (27 to 31); species o f N em atalosa have a more inferior m outh and gillrakers not more than h a lf
length o f gili filam ents. Other clupeids lack a dorsal filament.
G eographical D istribution : Coasts and rivers o f
northw estern Pacific (coasts o f Japan southw ard from
a b o u t 38°N , i.e. not H okkaido;
Sea o f Jap an n o rth to
V ladivostok on mainland; also Y ellow Sea, including G ulf
o f P o -H ai and East China Sea south to Taiwan Island and
Hong Kong).
H abitat and B iology : M arine, o ff coasts and in
bays, but m igrating into shallow er brackishw ater for
breeding, at least in A riake Sound (south K yushu), where
the fishes spaw n far up the Sound in A pril and M ay,
individual fishes apparently spaw ning tw ice or more in
this season (Takita, 1978).
Size : To 25 cm standard length (at 2 years, but 4
year-old fishes occur and must be larger).
Interest to F ish eries
O ccupies an im portant
position in the fisheries o f A riake Sound (Takita, 1978)
and probably elsew here, but individual catches not
recorded by FAO. C aught by trap nets and beach seines
in Japan.
Local Names :
JA PAN : K onoshiro.
Literature : Uchida et ah (1958 - eggs and larvae, figures; in Japanese); Takita (1978 - breeding, w ith
to papers in Japanese on ecology); N elson & R othm an (1973 - w ith 98 references to the species).
N em atalosa
R egan,
1917
N em atalosa R egan, 1917, Ann.M ag, nat.Hi st.. (8)9:312 (type: Clupea nasus Bloch).
A u st.Z o o l.. 10(2): 170 (type: Chatoessus elongatus M acleay).
refs
CLUP N eili
F lu v ialo sa W hitley,
1943,
D iagnostic F eatures
M e d iu m sized m arine and riverine gizzard shads
(to about 33 cm standard length); belly
fu lly scuted.
M o u th u su a lly in fe rio r,
w ith
snout
rounded
and
pro jectin g
(n earer to subterm inal in som e species);
edges o f low er jaw flared outw ard, denm idline
tary bones m eeting at an obtuse angle,
som etim es form ing an alm ost transverse
cleft; pre-m axillae short, m axilla ex­
panded at tip and actually or apparently
turned dow nw ard, w ith a single sm all
supra-maxilla. G illrakers fine, num erous
(up to 500 in som e sp ecies), but fairly
short, not m ore th an h a lf le n g th o f corm axilla
dorsal scales overlapping
responding gili filam en ts on first arch.
Last dorsal finray filam entous; anal fin
shorter than head, w ith 17 to 26 fin ra y s .
Pre-dorsal scales paired and overlapping in m idline. One (or m ore) dark spots on flanks in som e species.
Resem bles C lupanodon and K onosirus, w hich have a more ‘norm al’ clupeid mouth (edges o f low er ja w not strongly
flared outw ard) and paired pre-dorsal scales not overlapping in midline. O ther Indo-Pacific gizzard shads and
shads (A losinae) lack a dorsal filament.
242
Biology, H abitat and D istribution : Some species m arine, but passing into brackish- or freshwater, others
purely fluviatile. Filter-feeders. Indo-Pacific region, from the eastern coasts o f A frica to A ustralia and Japan.
Interest to Fisheries
N ot recorded, but o f limited local interest.
Species : N elso n & R othm an (1973) reco g n ized 7 species, but W ongratana (1980) added two more; it seems
possible that the w ide-ranging FF nasus may be more than one species:
N.
N.
N.
N.
N.
N.
N.
N.
N.
arabica Regan, 1917, northwestern Indian Ocean
come (Richardson, 1846), A ustralia to East China Sea
erebi (Günther, 1868), A ustralia
Ilvensis W ongratana, 1983, Papua N ew Guinea
galatheae N elson & Rothm an, 1973, India to Viet Nam
japonica Regan, 1917, Japan to Taiwan Island
nasus (Bloch, 1795), northern Indian Ocean to South China Sea
papuensis (M unro, 1964), Papua N ew Guinea
vlaminghi (M unro, 1956), W estern A ustralia
N em a ta lo sa
a ra b ica R egan, 1917
CLUP N e m
2
N em atalosa ara b ic a R egan, 1917, A nn.M ag.nat.H ist.. (8)19:313 (M uscat).
Synonym s : N em atalosa arabica - Fow ler, 1941:554 (com piled); W hitehead, 1962:92 et seq.. figs lb, 2c, 3a
(k ey , d ia g n o stic fe atu re s); Id e m . 1965:261. (G u lf o f A d en , k ey , d e sc r.); Id e m . 1 9 7 3 b :2 0 5 , fig. 31 (k ey , refs);
N elso n & Rothman, 1973:149, fig. 5B (photo) (synon., descr., refs); Wongratana, 1980:177, pis 126, 127 (revision).
F A O Names :
E n - A rab ian gizzard shad.
Diagnostic Features : A marine gizzard shad;
body rather deep, its depth 34 to 40% o f standard length,
com pressed, belly w ith 17 or 18 + 10 to 15 scutes. A n te rio r arm o f p re-o p ercu lu m w ith a fleshy tria n g u la r area
above, not covered by th ird in fra-o rb ital bone (see FL co m e). M outh subterm inal, edge o f low er jaw som ewhat
flared outward. Last dorsal finray filam en to us. Hind edge o f scales not toothed. A dark spot behind gili opening.
R e se m b le s N nasus and N . g a lath e ae , w hich have no fleshy gap above the a n te rio r arm o f the p re-o p ercu lu m
(since the third infra-orbital bone comes down to m eet it); also, they have few er scutes (total usually not more
that 31; cf. 29 to 34 in N. a ra b ic a ). No other clupeids in the area have a dorsal filam ent and inferior mouth.
243
Geographical Distribution : N orthw estern Indian Ocean (G ulf
o f Aden to G ulf o f Oman; no records from Red Sea or the “G u lf’).
H abitat and B iology : M arine, pelagic.
Size :
u
r -
1
.....
■■
M ore data needed.
To 16.5 cm standard length.
[V
i* \
\jf
Interest to F ish eries : N ot recorded.
L ocal N am es : -
\
?
•
Literature :
T here ap p ears to be no p u b lish ed study on its
biology. N elson & R othm an (1973) found only five references.
•
(
U
N em atalosa com e (R ichardson,
1846)
C LU P N em 3
Chatoessus come R ichardson, 1846, Ichthyol.V oy.Erebus & T erro r:62.
pi. 38, figs 7-10 (Western Australia).
Synonyms : N em atalosa come - F o w le r, 1941:522 (com piled, but not all synonym s w ere this species, e.g.
rich ard so n i and ho rni b elo n g in FT e re b i, as also refs to th a t sp ecies); M u n ro , 195625, fig. 176 (A u stralia,
com piled); Id em , 1967:43, pi. 3, fig. 32 (P ort M oresby, P apua N ew G uinea); N elson & R othm an, 1973:150, pi. 5C
(photo), map 2 (synon., descr., refs); W ongratana, 1980:178, pis 128, 129 (revision).
F A O Names :
En - W estern P acific g izzard shad.
*
D iagnostic Features : A m arine gizzard shad; body rather deep, its
depth about 36 to 45% o f standard length, com pressed, belly w ith 18 or 19
(rarely 20) + 10 to 13 (usually 11 or 12), total 29 to 31 (usually 30) sc u te s.
A n te rio r arm o f p re -o p e rc u lu m w ith fle sh y tr ia n g u la r a re a a b o v e , no t
covered by third in fra-o rb ital bone. M outh subterm inal, edge o f low er ja w
strongly flared outw ard. L ast dorsal finray filam en to u s; pecto ral axillary
scale well developed. Hind edge o f scales not toothed. A dark spot behind
gili opening. Range in East and South C hina Seas and perhaps in Java Sea
overlaps that o f FL nasus and possibly also FL galatheae, neither o f w hich
has a fle sh y gap a b o v e a n te rio r arm o f p re -o p e rc u lu m (s in c e th e th ird
in fra o rb ita l b o n e co m es d o w n to m e e t it);
th is also d istin g u ish es the
fleshy gap
an terio r arm o f
pre-o p ercu lu m
3rd
in fra -o rb ita l
244
freshw ater N. erebi, N. papuensis and FT ilvensis (w hich in addition have only rudim entary or absent pectoral
axillary scales) ; in
FT japonica thehind edge o f the scales is distinctly toothed. N o other clupeids in the area
have a dorsal fila m e n t a n d inferior mouth.
Geographical D istribution : W estern central P aci­
fic (coasts o f Q ueensland, Papua N ew G uinea and Irian
Jay a (both on southern coasts only), Java Sea, the Philippines, north to E ast C hina Sea, i.e. R yukyu Islands at
about 23° N).
1-----------1
I----------- I-----------
40°
)
À à
H ab itat and B iology
needed.
Size :
to 18 cm.
*
y j
>
M arine, pelagic. M ore data
*9 ¿
*
To 21 cm standard length, usually about 15
0
I n t e r e s t to F i s h e r i e s
C o n t r i b u t e s to
clupeoid catches, but not o f importance.
L o ca l N am es : A U S T R A L IA : H a irb a c k
(Munro, 1956); JA PA N : R y u k yo-dorokui.
%i v
°
^ V
lo c a l
h e rrin g
20
°
, <f
•
Literature : There appears to be no published study
o n its b io lo g y . N e l s o n & R o t h m a n ( 1 9 7 3 ) fo u n d 85
re fe re n c e s .
V
80°
100°
120°
140°
160°
18C
C L U P N em 4
N em atalosa ereb i (G ü n th e r, 1868)
C hatoessus erebi G ü n th e r, 1868, C at.F ishes B rit.M u s., 7:407 (M ary R iver, Q ueensland).
Synon ym s : C h ato essu s rich ard so n i C asteln au , 1873:144 (M urray R iv er, A u stra lia ); C h ato essu s elongatus
M acleay, 1883:209 (M ary R iv er, Q ueen slan d , A u stralia );
C h a to e ssu s horni Z eitz, 1896:180, pi. 16, fig. 6 Finke
R iv e r, N o rth e rn T e rrito ry , A u s tr a lia ) ; N e m a ta lo s a e lo n g a ta :Fow ler. 1941:554 (c o m p ile d ); F lu v ia lo s a b u lle ri
W hitley, 1948:267 (Ord R iv er, W estern A u stralia); F lu v ialo sa aracom e W hitley , 1948:267 (Fitzroy River, W estern
A ustralia); Fluvialosa richardsoni:M unro, 1956:26, fig. 178 (com piled Scott, G lover & Southgate, 1974:71, fig.
(South A ustralia); Fluvialosa b ulleri:Munro, 1956:26, fig. 180 (com piled); Fluvialosa paracom e:Munro, 1956:26, fig.
181 (com piled); F lu v ialo sa e re b i:M unro, 1956:26, fig. 179 (com piled); N em atalo sa erebi - N elso n & R othm an,
1973:152, figs 6, 7, 8A (photos), map 3 (synon., descr., refs); W ongratana, 1980:173, pis 122, 123 (revision).
FAO Names
E n - A u stra lian riv er gizzard shad.
f >
I
{
)
t t
> i i i
in ii ii ii i
î
j
i
;
,
t
i
V
t
,
j
¡
> ■
y
i
;
u rrrr yA
■MXl n r , "ÍY"
■î i 1 ■kl %x\ XA ^
V'■■iiyi' Vïu V■:i ■(-ni1■AVïi1■n\W\'Ïi,
ïû'Y xnŸY
(h'XÏ i * »
mXK A
I V
C
(X'A À '/f/FV
J. AU*, , , A
' i
V y
U
V
1
,
n
"
')
V A .
D iagnostic Features : A riverine gizzard shad; body rather deep, its depth about 34 to 47% o f standard
length, compressed, belly with 14 to 18 (usually 16 to 17) - 11 to 14 (usually 12 or 13), total 25 to 31 (usually 20 to
30) sc u te s. A nterior arm o f pre-operculum w ith a fleshy triangular area above, not covered by third infra-orbital
bone (see FT com e). M outh inferior, edge o f low er jaw strongly flared outward.
Last dorsal fin ra y filamentous;
pectoral axillary scale ru d im en tary or absent. Hind edge o f scales not toothed. A dark spot behind gili opening.
O verlaps northw estern A ustralian range o f FL vlam inghi, w hich has hind edge o f scales toothed. Other species o f
N em atalosa are m arine, also pectoral ax illary scales well developed. No other clupeids in the area have a dorsal
filam ent and inferior mouth.
G eograp h ical D istrib u tio n : Rivers of Australia
and southw estern Papua N ew G uinea (from Fortesque
R iver near D am pier A rch ip elag o in W estern A u stralia,
eastward in rivers through the N orthern Territory and
Q ueensland south to the M urray - D arling system; also
the F inke R iver, afflu en t to L ake E yre; D igoel R iver
in Papua N ew Guinea).
H abitat and B iology : R iverine, often far from
sea, but also in estuaries; occurs in Lakes A lbert and
A lex an d rin a n ear m outh o f M urray R iver. M ore data
needed on food and breeding.
Size :
25 cm.
To 32.5 cm standard length, usually 20 to
Interest to Fisheries :
O f small local interest only.
100°
120°
140°
160°
180°
Local Names :
AUSTRALIA:
Bony bream (various adjectives supplied by M unro, 1956 for the species
which he reco g n ized ), Pyberry, Tukari.
Literature : There appears to be no published study o f the biology o f this species. Nelson
found 92 references.
& R othm an (1973)
Remarks :
N elso n & R othm an (1973) found only sm all d ifferen ce in m eristic co u n ts betw een specim ens
from d ifferen t A u stralian rivers.
N e m a ta lo s a
flv e n s is W ongratana, 1983
C L U P N em 5
N em atalo sa flven sis W o n g ratan a, 1 9 8 3 , Jap.J.Ichthyol., 29(4):393, fig. 9 (Toro Pass, Fly R iv e r, P a p u a
Guinea; based in part on a paratype o f N p a p u e n sis).
Synonyms :
N em atalosa
FA O N am es : En
flvensis:W ongratana.
- Fly river gizzard shad.
1980:184, pis 138, 139 (revision).
N ew
246
D iagnostic Features : A riverine gizzard shad; body rather deep, its depth about 32 to 40% o f standard
length, com pressed, belly w ith 16 to 18 (usually 17)+ 10 to 13 (usually 11 or 12), total 26 to 32 (usually 28 or 29)
scutes. A n terio r arm o f p re-o p ercu lu m w ith a fleshy tria n g u la r area above, not covered by th ird infra-o rbital
bone (see FL com e). M outh inferior, edge o f low er jaw strongly flared outward. Last dorsal finray filam en tous;
pectoral axillary scales absent. Hind edge o f scales toothed. No dark spot behind gili opening, but a faint band on
flank. Sympatric w ith FL papuensis, w hich has few er gillrakers (160 to 320 at 10 to 20 cm standard length; cf. 360
to 520), those on the low er arch being not more than 2/3 length o f corresponding gili filam ents (2/3 to equal in N.
flvensis). All other N em atalosa in the area are marine and have well developed pectoral axillary scales.
G eographical D istribution : Papua N ew Guinea
(only known from the Fly and Strickland rivers).
H abitat
tio n w h e th e r
needed.
o°
<y
and
B iology : R iverine, but no in d ic a ­
it d e sc e n d s to e s tu a rie s .
M o re d a ta
20°
Size !
To 2 2 .2 cm stan d ard len g th ,
p erhaps
m ore.
Interest to Fisheries
: Possibly m akes a local
contribution to river fisheries.
40°
L ocal N am es : 100°
N e m a ta lo s a
g ala th ea e N e ls o n & R othm an,
120°
140°
160°
C L U P N em 6
1973
N em atalosa galatheae N elson & R o th m an , 1973, Bull. A m .M us.nat.H ist..
(Ranong Province, A ndam an Sea, Thailand, also m outh o f Pakchan River).
180°
150(2): 158,
figs
8B
9
(photos)
Synonyms : N em atalosa n a s u s :R egan. 1917:313 (Cañara, India only); Herre & M yers, 1937:13 (Singapore);
N em atalosa c h a n p o le /W ongratana. 1980:176, pis 124, 125 (revision - see Rem arks for use o f chanpole as the name
for this species .
FA O Names :
E n - G alath ea g iz z a rd shad.
247
D iagnostic Features
A
m arine
gizzard
shad;
body
rather deep, its depth about 36
to 44% o f standard length,
com pressed, belly w ith 16 or 17
(usually 17) + 8 to 11 (usually
10), total 25 to 30 (usually 28)
s c u te s . A nterior arm o f p r e ­
operculum w ith the third i n f r a ­
grooves in skin
orbital bone im m ediately above
it, no fleshy gap betw een.
Mouth
in ferio r,
low er
ja w
strongly flared outward. A pair
o f grooves in the spongy skin on
top o f head, converg in g p o ste ­
riorly.
L a st d o rsal finray f i la ­
m en tou s;
p e c to ra l a x illa ry
a n terio r arm o f'
sc ale p re s e n t.
H ind edge o f
3rd
p re-o p ercu lu m
scales not to o th e d . A dark spot
in fra-orbital
behind gili opening, followed by
a series o f sm aller spots along flank.
Closely resem bles FL nasus, w hich lacks the pair o f grooves on top o f head
and has the hind edge o f the scales toothed; FL alatheae is the only N em atalosa w ith a series o f spots on flank
(at m ost a single spot behind gili cover in all others. N o other clupeid in the area has a dorsal filam ent.
Geographical D istribution :
In d ian O cean (so u th ­
w est coast o f India, presumably Bay o f Bengal, o ff A nda­
mana Sea co ast o f T hailand, Penang), G u lf o f T hailand,
Singapore, Viet Nam.
40°
Habitat and Biology : M arine, but penetrating and
presum ably breeding in freshw ater if H am ilton-B uchanan’s
C lupanodon chanpole is this species (see Remarks). M ore 2o°
specimens and data needed.
Size :
To 16.3 cm standard length.
Interest to F isheries :
Probably o f local interest.
Local Names : INDIA: Chanpole (Ganges delta, if
chanpole is this species - see Remarks).
20°
Literature :
biology.
T here appears to be no study o f its
Remarks :
W o n g r a ta n a
(1980:176) b elieved this 40s p e c ie s to be th e C lu p a n o d o n c h a n p o le o f H am ilto n B u c h a n a n (1822:249, pi. 18, fig. 74), a n a m e h ith e r to
placed in A nodontostom a (e.g. by Fow ler, 1941:549 and W hitehead, 1973b:208). H am ilton-Buchanan’s figure shows
a fish with 5 spots along the flank (as in N em atalosa g a la th e a e ), but no dorsal filam ent (as in A nodontostom a); the
dorsal view o f the head does not show the pair o f grooves characteristic o f FL galatheae (as W ongratana claimed),
but the freshw ater provenance o f chanpole (“ponds and ditches o f every part o f B engal” seems to hint more at the
freshw ater species o f G onialosa (w hich lack a dorsal filam ent, but also lack spots on the flanks). The problem is
left unresolved, w ith chanpole tem porarily put in the synonymy o f A nodontostom a chacunda.
Nematalosa
japonica R egan, 1917
C LU P N em
7
N em atalosa japonica R egan, 1917, A nn.M ag.nat.H ist., (8)19:313 (inland Sea o f Japan).
S y n o n y m s : N e m a ta lo s a ja p o n ic a - F o w le r, 1 9 4 1 :5 5 5 (c o m p ile d );W h ite h e a d , 1962:101 (key, as iap o n icus);
N elso n & R othm an, 1973:159, fig. 8C (photo) (synon., descr., refs); Masuda et ah, 1975:166, pi. 13C (colour photo)
(M atsushim a B ay so u th w ard ); W o n g ratan a, 1980:180, pis 133, 134 ( r e v is i o n ) ; M asuda et ah, 1984:20, pi. 22C
(colour photo) (southern Japan southward).
F A O Names :
En - Japanese gizzard shad.
248
D iagnostic Features : A marine gizzard shad; body generally more slender than in other species, its depth
33 to 38% o f standard length, belly w ith 16 to 19 (usually 18)+ 13 to 16 (usually 13 or 14), total 30 to 34 (usually
32 or 33) scutes. A n te rio r arm o f pre-o p ercu lu m w ith a fleshy trian g le above, n o t covered by th ird in fra-o rb ital
bone (see FL cornel. M outh inferior, edge o f low er jaw strongly flared outw ard. L ast dorsal fin ra y filam entous;
pectoral axillary scale w ell developed. Hind edge o f scales distinctly toothed. A dark spot behind gili opening. In
southern part o f range it overlaps FL nasus, perhaps also N. galatheae, both w ith no fleshy gap above the anterior
arm o f the pre-operculum (since the third infra-orbital bone comes down to meet it); also, scale edge sm ooth in
N. galatheae (as well as in N. come, which reaches to the Ryukyu Islands). B oth Clupanodon thrissa and K onosirus
punctatus o f Chinese and C hinese-Japanese w aters have a dorsal filam ent, but m outh subterminal and gillrakers at
least 3/4 length o f qill filaments.
------ 1-------l
i
l
i
------1r .....
1
G eographical D istribution : W estern P acific (from
M atsushim a Bay, Sea o f Japan, at about 36°N , so u th w ard 40°
to Taiwan Island, Hong K ong, the Philippines, and a single
record from G ulf o f Thailand).
H abitat and B iology :
M arin e, p referrin g w arm er
w ater th an K o n o siru s p u n ctatu s in Jap an , sch o o lin g n ear
shore over sand or m ud.
F ilte r-fe e d e r.
B reed s around
M ay in Japanese waters.
Size !
To about 19 cm standard length.
Interest to F isheries : O f local in te re st in Japan,
where it is caught by set net or g illn e t.
Local Names :
»
I
20
°
*
i'
\r 1
•
*
i
•> /» j
•
1
\
4
56
—
H
V
jT4
K
JA PAN : D orokui.
vl
<r
V
r
Literature :
re fe re n c e s .
N e ls o n & R othm an (1 9 7 3 ) fo u n d 23
N em atalosa n asu s (Bloch, 1795)
'V
C LU P N em
1
Clupea nasus Bloch, 1795, N atu rg esc h .a u sl.F isc h e . 9:116, pi. 429, fig. 1 (Malabar).
Synonym s : Clupanodon nasica Lacepède, 1803:468, 472 (M alabar; on C. nasus B loch); ? Chatoessus altus
G ray, 1834 (1 O ctober :pl. 91, fig. 2 (India, no exact locality); ? Chatoessus chrysopterus R ichardson, 1846:308
(C anton or M acao, on R eeves drawing); N em atalosa nasus - Fow ler, 1941:555 com piled; very com plete synon.);
W h ite h e a d , 1 9 6 2 :9 7 , fig . 4 b (k e y , in f ra -o r b ita l); Id e m , 1 9 6 5 2 6 2 (th e “ G u l f ’); Id e m , 1 9 6 6 :3 6 (id e n tity o f
chrysoptera); Idem , 1969272, pi. 2c (B loch’s pi.), fig. lb (type o f nasus); Idem , 1973b:204, fig. 30 (key, refs);
N elso n & R othm an, 1973:161, fig. IO A (photo) (synon., descr., refs); W ongratana, 1980:179, pis 130, 131 (revision).
249
F A O Names :
En - B loch’s gizzard shad.
1
•
tá * 1 AJl&) H ^ x) n
> i m V14 m i
-I'lvr-U.
-
”• i > í n
V
y J *, f ) '
t
ríͧPfi
^ ‘ m h ‘h V W /
^•K
D iagnostic F eatures : A m arine gizzard shad; body rather
deep, its depth about 34 to 41% o f standard length, belly w ith 17 to
20 (u s u a lly 18) + 9 to 13 (u su ally 11), to tal 28 to 32 (u su ally 30)
scutes. A n terio r arm o f p re-o p ercu lu m w ith the th ird in fra-o rb ital
bone im m ed iately above it, no fleshy gap betw een. M outh in ferio r,
low er ja w strongly flared outw ard. L ast dorsal finray fila m e n to u s ;
p e c to r a l axillary s c a le p r e s e n t.
H in d ed g e o f sc a le s d is tin c tly
too thed. A dark spot behind gili opening. R esem bles N. g ala th e ae ,
w hich has spots along the flank and a sm ooth hind edge to scales; in
w est o f range, overlaps IE ara b ic a , and in east o f range overlaps N.
com e and N . ja p o n ic a , b u t all o f th e s e h a v e a d is t i n c t fle s h y
tr ia n g u la r a re a above the an terio r arm o f the p re-o p ercu lu m (i.e.
third in fra-o rb ital bone does not cover th is area, but slopes b ack ­
w ard), also, none have toothed scale edges. No other clupeid in the
range o f FT nasus has a filam entous dorsal finray. See CLU P N em 1,
Fishing Area 51, also Fishing Areas 57, 71.
#
a n te rio r arm
o f pre-operculum
3rd in fra-o rb ital
G eographical D istribution
Indian O cean (the
“ G u lf” , c o a sts o f In d ia to A n d a m a n S e a ), G u lf o f
T hailand, S outh C hina Sea to the P h ilip p in es, H ong
K ong and no rth to so u th ern Jap an or so u th ern tip o f
K orea).
N ot South A fric a (W h iteh ead & W o n g ra ta n a ,
i.e. SFSA, in press).
H abitat and B iology : M arine, pelagic, probably
entering estu aries (e.g. specim en from B assein R iv er,
Burm a).
R ecords o f spaw ning in C hilka L ak e, east
coast o f India, in August (K ow tal, 1970).
Size :
15 cm.
To 21 cm standard length, usually about
Interest to F isheries : N o special fishery, but
co n trib u tes to clu p eid catches; in Saudi A rab ia it is
caught in set nets and traps, but com m ands only low to
medium prices in markets (W ray, 1979).
Local Names
SAUDI ARABIA:
Shroom.
Literature :
N e lso n & R othm an (1973) gave over a hundred references to this species; not included was
Kowtal (1970) on eggs and larvae from Chilka Lake.
Remarks : The very wide range o f this species suggests that it may be split into subspecies, although the
published studies o f N elson & R othm an (1973) and W ongratana (1980) show rather little variation.
N ev erth eless,
Nelson and Al-Hassan suspect one or more undescribed nasus- like species in the “G u lf’.
250
Nematalosa papuensis
CLUP Nem
(Munro, 1964)
8
F luvialosa papuensis M unro, 1964, Papua N ew G uin.agrie.J., 16:152, fig. 3 (Strickland R iver, N ew Guinea;
holotype only, the paratype being FL flvensis).
Synonyms : F lu v ialo sa p ap u en sis:M unro. 1967:43, pi. 3, fig. 33 (upper part o f Strickland River); N em atalosa
papuensis - W ongratana, 1980:182, pis 136, 137 (revision).
FAO Names :
En - S trickland river gizzard shad.
Diagnostic Features :
A riverine gizzard shad; body rather deep, its depth about 32 to 40% o f standard
length, belly w ith 16 to 18 (usually 17) + 10 to 13 (usually 11 or 12), total 26 to 31 (usually 28 or 29) scutes.
A nterior arm o f pre-operculum w ith a fleshy triangular area above, not covered by third infra-orbital bone (see
N. com e). M outh inferior, edge o f low er jaw strongly flared outward.
L ast dorsal finray fila m e n to u s ; p e c to ra l
axillary scales absent. H ind edge o f scales toothed. No dark spot behind gili opening, but a band on Hand.
Sym patric w ith FL flvensis, w hich has more gillrakers (360 to 520 at 10 to 20 cm standard length; cf. 160 to 520),
those on the lower arch being at least 2/3 length o f corresponding gili filam ents (about 1/5 to 2/3 in N. papuensis).
O ther species o f N em atalo sa are m arine, also pecto ral ax illary scales w ell developed. N o o th er clupeids in the
area have a dorsal filam ent and inferior mouth.
Geographical Distribution :
Papua N ew Guinea
(only known from Fly and Strickland rivers).
o°
H abitat and B iology : R iverine, but no indica­
tio n w h e th e r it d e sc e n d s to e s tu a rie s .
M o re d a ta
needed.
20°
Size :
To 20.7 cm standard length.
Interest to Fisheries :
P o ssib ly m akes a local
contribution to river fisheries.
Local Names : -
40°
100°
120°
140°
160°
180°
251
CLUP
N em atalosa vlam inghi (M unro, 1956)
N eili
9
F lu v ialo sa vlam inghi M unro, 1956, Fish N ew sletter, 25(12):25, fig. 177 (Sw an River, W estern A ustralia).
Synonyms : N e m a ta lo s a v la m in g h i - N elson & R othm an, 1973:163, lig. 10B (photo), map 2 (synon., descr.,
refs): W o n g ratan a, 1980:181, pi. 135 (revision): F lutchins & Thom pson, 1983:18, 75, fig. 55 (Bunbury to Broom e,
com piled).
F A O Names :
En - W estern A u stra lia n gizzard shad.
WwwWVYV y y yv)
YYYWn
D iagnostic Features : A marine and estuarine gizzard shad: body rather deep, its depth about 36 to 40% of
standard length, com pressed, belly w ith 16 to 19 (usually 18) + 12 to 14 (usually 12), total 28 to 32 (usually 31)
scutes. A n terio r arm o f p re-o p ercu lu m w ith a fleshy trian g u la r area above, not covered by th ird in fra-o rb ital
bone (see N. com e). M outh inferior, low er ja w strongly flared outward. Last dorsal finray filam en to u s: a sm all
pectoral axillary scale present. Flind edge o f scales toothed. A dark spot behind gili opening. Overlaps range o f
the riverine N. erebi and the marine FL com e, w hich have the hind edge o f the scales smooth. N em atalosa is the
only clupeid w ith a dorsal filam ent and inferior mouth in this area.
Geographical Distribution :
W estern A u stralia
(from Bunbury, south o f Perth, north and eastw ard to
Broome).
i
>
S ^
,e£>■
Habitat and Biology :
M arin e and estuarin e
(g iv e n as flu v ia tile by M u n ro (1956), b u t N elso n &
R othm an (1973) failed to record any tru ly fresh w ater
specim ens), abundant in estuaries, also found in c o a s­
tal em baym ents (H u tch in s & T h o m p so n , 1983). M ore
data needed.
\
'ïi'”
Ci
r
y.
Size :
To 19 cm standard length.
In terest to F ish eries : O f local interest only.
L ocal Nam es :
Perth herring.
Literature :
re fe re n c e s .
AUSTRALIA:
h
S? 0
VT
Í
B ony bream ,
T here appears to be no study o f its biology.
N e ls o n & R othm an
(1973) found only five
Remarks :
W o n g r a ta n a (1980:182) found g illrak e r d ifferen ces in d iffe re n t areas, su g g estin g th a t the
populations are isolated, but more material is needed.
click for n e x t p a g e
c lic k fo r p r e v io u s p a g e
252
A nodontostom a Bleeker.
C LU P A nod
1849
? G onostom a van H asselt, 1823, A lg em een e K o n st- en L e tte r-b o d e , 1(21):329 (ty p e:
none g iven:
not
G o n o sto m a R afinesque, 1810).
A n o d o n to s to m a B le e k e r, 1 8 4 9 , V e rh .b a ta v .G e n o o t.K u n s t.W e t., 2 2 :1 5 (type:
A n o d o n to sto m a h a s s e ltii B le e k e r = C lu p a n o d o n c h a c u n d a H a m ilto n -B u m .
D iagnostic Features :
S m a ll o r m o d e r a te - s iz e d
m arine gizzard shads (but
e n te rin g riv e rs ) (to a b o u t
18 cm
sta n d a rd
len g th ):
body v ery d eep , its d e p th
40 to 75%
o f standard
length in fishes over 9 cm,
m idline
b elly fully scuted. M o u th
su b te rm in a l or a little in ­
ferior, upper jaw slender at
tip and not strongly turned
2nd
dow n, a single slen d er or
supra-m axilla
even
splint-like
second
m edium p re-dorsal scales
su pra-m axilla:
lo w e r ja w
short, its e d g e s s tro n g ly
m axilla
fla re d o u tw a rd . G illra k e rs
fin e and n u m e ro u s (50 to
170 on lower arch). Last dorsal finray not filam entous: anal fin much shorter than head, with 17 to 25 (usually 19
to 21) finrays.
Pre-dorsal scales forming a single m edian series, post-dorsal scales w ith hind margin elongated.
Scales m oderately large, 38 to 45 in lateral series.
A dark spot behind gili opening.
Resem bles the freshw ater
G on ialosa in lacking a dorsal filam en t, bu t in th a t genus the p re-d o rsal scales are p aired , th ere are 22 to 28
(usually 24 to 26) anal fin ra y s and the upper jaw turns down strongly at its tip. Other In d o - P a c ific gizzard shads
have a dorsal filam ent: the mouth is terminal in the Alosinae (H ilsa. Tenualosa. G udusia) and other clupeids.
B iology, H abitat and D istrib u tion :
A ustralia, the Caroline Islands and N ew Caledonia.
Interest to Fisheries :
M arine, but entering freshw ater, from the “ G u lf ’ to northern
No specific fisheries, but o f lim ited local interest.
Species :
To the 2 species (chacunda and chanpole) mostly accepted by earlier authors, W ongratana (1983)
added a third, but identified H am ilton-B uchanan’s chanpole as a N em atalosa and resurrected B leeker’s se la n g k at:
chanpole is here re c o g n ized as a synonym o f chacunda:
A. chacunda (Hamilton-Buchanan, 1822), Indo-W est Pacific
A. selangkat (B le e k e r, 1852), eastern Indian Ocean and Indonesia
A. thailandiae W ongratana, 1983, Thailand, Indonesia.
A n o d o n to sto m a ch acu n d a
(Hamilton-Buchanan, 1822)
C LU P
A nod
1
Clupanodon chacunda H am ilton-B uchanan, 1822, Fishes o f G anges:246 (G anges estuaries).
Synonyms : Clupanodon chanpole H am ilton-Buchanan, 1822:249, pi. 18, fig. 74 (Ganges): ? Gonostom a sp.
van H asselt, 1823:329 (“die z e e r veel overeenkom st heeft met Meg. N aso Lacep. ", but w ithout dorsal filam ent: see
A lfred, 1961 for notes on van H asselt’s fishes): A n o d o n ta stom a hasseltii B leek er, 1849:15 (Madura Strait, Java
Sea): G onostom a jav arficu m H y rtl, 1855:49 (Java): A n o d o n to sto m a ch acu n d a - Fowler, 1941:549 (the Philippines,
‘B orneo’, Java, India: key, very large synon. d e sc r.); W h ite h e a d , 1962:101 (key): Id em . 1965:263 (the “G u lf” ):
W h iteh ead , B o esem an & W heeler, 1966:88, 89, pi. 11, fig. 1 (B leeker’s fig.) (types o f hasseltii, selangkat): N elson
& R othm an, 1973:141, figs 1C, 4A, B ( p h o t o s ) , m a p 1 (synon., d e sc r., v ery larg e lis t o f refs); W h i t e h e a d ,
1973b:207, fig. 33 (key, refs); W ongratana, 1980:188, pis 146, 147 (revision).
FAO N am es :
E n - C hacunda g izza rd shad.
253
D iagnostic Features : A marine gizzard shad; body very deep, its
depth increasing w ith size o f fish, 40 to 70% o f standard length in fishes
over 10 cm. M outh inferior; second su p ra-m ax illa a m ere splint. L ow er
g illrak ers 54 to 96; lo n g est g illrak ers on low er p art o f arch less than
corresponding gili filam ents (and much less in larger fishes). Last dorsal
finray not filam en to u s. H ind edges o f scales to o th ed , th e te eth th in n e r
th an gaps betw een them ; a m edian series o f p re-dorsal scales. A large
black spot behind gili opening. R esem bles A. selangkat, w hich has more
low er gillrakers (100 to 166) and teeth on scales w ider than gaps between,
and A. thailandiae, w hich has the longest gillrakers on low er part o f arch
at lea st as long as corresponding gili filam ents, also second supra-maxilla
paddle-shaped.
Species o f G o n ialo sa and N em ata lo sa have p aired p re ­
dorsal scales, also a dorsal filam en t in the latter. O ther clu p eid s have a
term in al m outh. See C L U P Anod 1, Fishing A rea 51, also Fishing A reas
57, 71.
scale
G eographical D istribution :
Indian O cean (the
“G u l f ’ to coasts o f In d ia and A n d am an S ea), w estern
P a c ific ( G u lf o f T h a ila n d to n o rth e rn A u s tr a lia , th e
Caroline Islands and N ew Caledonia; also Indonesia, the
Philippines and Viet Nam).
H abitat and B iology :
M arin e, pelag ic and in ­
shore, also in estuaries. Feeds on diatoms, radiolarians,
m olluscs, copepods and c ru s ta c e a n s (in th a t o rd e r o f
im p o rtan ce, at least in the G odavari estuary). B reeds
from N ovem ber to F eb ru ary , m ainly in the later part
(G odavari estuary).
Size :
To 17.5 cm stan d ard le n g th , com m on
around 10 to 14 cm.
In terest to F ish eries :
L ocal c o n trib u tio n s, e.g.
N ovem b er to June in G odavari estu ary , but no special
fishery.
Local Names : INDIA: Chacunda; INDONESIA: Selangkat, Trubala (M alay), Lakar, P enden, Slam at (Java),
B an d rin g , D jan g an (M adura), Belo (Sunda), Pias (Bagan Api-Api).
Literature : Chacko (1954 - breeding); Babu Rao (1965 - general biology); further references in W hitehead
(1973b) and over a h undred in N e lso n & R othm an (1973), alth o u g h at least som e o f th ese m ust refer to the
sympatric species W selangkat and W thailandiae.
Rem arks :
For identity o f H am ilton-B uchanan’s chanpole, see under N em atalosa galatheae
254
A nodontostom a selangkat
C LU P
(B leek er, 1852)
A nod
2
C hato essu s selan g k at Bleeker, 1852, N a tu u rk .T jd sc h r.N e d .-In d ië , 3:458 (Muntok, B atavia).
Synonyms : Chatoessus breviceps
Peters, 1877:838 (New H anover); A n ad o n to sto m a chacunda :W hitehead.
B o e se m a n & W heeler, 1966:89, pi. 11, fig. 1 (B leeker’s figure) (type o f selangkat); num erous authors w ho placed
selangkat in the synonym y o f chacunda; A nodontostom a selangkat - W ongratana, 1980:191, pis 148, 149 (revision).
F A O Names :
En - In d o n esian gizzard shad.
D iagnostic Features : A marine gizzard shad; body very deep, its
depth increasing w ith size o f fish, 40 to 75% o f standard length in fishes
over 9 cm. M outh in fe rio r; second su p ra-m ax illa a m ere splint. L ow er
g illrak ers 100 to 166; lo n g est g illrak ers on low er p art o f arch less than
corresponding gili filam ents (and much less in larger fishes). Last dorsal
finray not filam en to u s. H ind edges o f scales to o th ed , the te eth w ider
th an gaps betw een them ; a m edian series o f p re-dorsal scales. A large
dark m ark behind gili open in g , follow ed by lon gitu d in al streaks along
flanks. R esem bles W chacunda, w hich has few er low er gillrakers (54 to
96) and teeth on scales th in n e r th a n gaps b etw een th em , and W thailan­
d ia e , w hich has the lo n g est g illra k e rs on lo w er p art o f arch a t least as
long as correspon d in g gili filam en ts, also second su p ra-m ax illa p a d d le ­
shaped.
Species o f G onialosa and N em ata lo sa have p aired pre-dorsal
sc a le s, also a d o rsa l fila m e n t in th e la tte r.
O th e r c lu p e id s h a v e a
term in al m outh.
G eographical D istribution : Indian O cean (A n d a ­
m an Is la n d s - a single D ay sp ecim en ), Jav a Sea, the
P h ilip p in es and eastw ard to B ism ark A rch ip elag o (type
locality o f breviceps).
H abitat and B iology :
M arine, pelagic and in ­
shore (probably a similar biology to W chacunda).
Size :
To 18 cm standard length.
Interest to
Fisheries
Local Names :
cunda.
: No data.
N o t d istin g u ish ed from
L iterature : C onfused w ith
authors prior to W ongratana (1980).
W
W ch a­
chacunda
by
scale
255
A nodontostom a thailandiae W ongratana,
CLU P
1983
A nod
3
A nodontostom a thailandiae W ongratana, 1983, Jap .J.Ic h th y o l., 29 (4 ):394, fig. 11 (Songkhla, G u lf o f Thailand,
also Calcutta, Bangladesh, Phuket, northern Sumatra, Borneo, Sarawak).
Synonyms : Anodontostoma chacunda:numerous authors.
F A O Names : En - Thai gizzard shad.
Diagnostic Features : A marine gizzard shad; body very deep, its
depth increasing with size o f fish, about 40 to 68% o f standard length in
fishes over 8 cm. M outh inferior; second supra-m axilla paddle-shaped.
Lower gillrakers 46 to 140; longest gillrakers on lower part o f arch equal
to or longer than corresponding gili filam ents. L ast dorsal fin ra y not
filam entous.
H ind edges o f scales to o th ed , the teeth w id er th an gaps
betw een them; a m edian series o f pre-dorsal scales. A large dark mark
behind gili opening. Resembles W chacunda and W selangkat, w hich have
lower gillrakers shorter than corresponding gili filaments and the second
supra-m axilla a mere splint. Species o f G onialosa and N em atalosa have
p aired p re -d o rsa l sc a le s,a lso a d o rsal fila m e n t in th e la tte r.
O th er
clupeids have a term inal mouth.
scale
G eographical D istribution : Indian O cean (north­
ern
part o f Bay o f B engal, A ndam an Sea at P huket in
T hailand), G u lf o f T hailand, South C hina Sea, ? Java
Sea.
H abitat and B iology :
M arine, pelagic and in ­
shore, probably entering estuaries (probably a sim ilar
biology to A. chacunda).
Size :
14 cm.
To 18 cm standard length, usually
10 to
Interest to Fisheries : No data.
L ocal N am es :
chacunda.
N o t distin g u ish ed
Literature :
C o n fu sed w ith
authors prior to W ongratana (1983).
W
from A.
ch a c u n d a by
80»
100"
120"
140"
160"
180"
256
C LU P G on
G onialosa R egan, 1917
G onialosa Regan, 1917, A nn.M ag.nat.H ist., (8)19:315 (type: Chatoessus m odestus Day, designated by Jordan,
1920:560).
Indialosa H erre & M y e rs, 1 9 4 1 , L in g n a n S c i.J ., (10):238 (ty p e : C lu p a n o d o n m a n m in a HamiltonB uchanan).
D iagnostic Features : Small freshw ater gizzard shads (to about 12 cm standard length); belly fully scuted.
M outh usually inferior (sub-term inal in one), upper ja w actually or apparently turned down at tip, short, rarely
re a c h in g fro n t m a rg in o f e y e , a sin g le s u p ra -m a x illa ;
low er ja w short, its edges strongly flared outw ard.
G illrakers fine and num erous (90 to 180 on low er arch). Last dorsal finray not filamentous; anal fin equal to or a
little shorter than head, w ith 22 to 28 (usually 24 to 26) finrays.
P re-d o rsal scales p aired and o v erlapping in
m idline.
Scales m oderate or sm all, 43 to 71 in lateral series.
A dark spot behind gili opening. R esem bles
A nodontostom a in lacking a dorsal filam ent, but in that genus the pre-dorsal scales form a single m edian row,
th e re a re only 17 to 25 (usually 19 to 21) anal finrays and the upper jaw does not turn down at its tip. Other IndoP acific gizzard shads have a dorsal filam en t; the m outh is term inal in the A losinae (H ilsa, T e n u alo sa , Gudusia)
and other clupeids.
Biology, Habitat and Distribution :
Interest to Fisheries :
Species :
Freshw ater in rivers o f India, B angladesh and Burma.
No specific fisheries, but o f limited local interest.
To the 2 species accepted since the revision by Reqan (1917), W ongratana (1983) added a third:
G. m anm ina (H am ilton-B uchanan, 1822), Ganges
G. m o d esta (Day, 1869), B urm a
G. w h iteh ead i W ongratana, 1983, Burma.
Gonialosa manmina (H am ilton-B uchanan,1822)
C L U P G on 1
C lupanodon m anm ina H am ilton-B uchanan, 1822, Fishes o f G anges:247 (Ganges).
Synonym s : C lupanodon cortius H am ilton-B uchanan, 1822:249 (near G oyalpara, B ram aputra R iver);
G onialosa m anm ina - R egan, 1917:315 (India, A ssam ); F o w ler,1941:548 (m isspelt m anm inna, com piled);
W hitehead, 1962:101 (key); Idem , 1973b:206, fig. 32 (key, refs); N elson & Rothm an, 1973:147, fig. 4C (photo)
(synon., descr., refs); W ongratana, 1980:186, pis 142, 143 (revision).
FAO Names
En - G anges riv er gizzard shad.
257
Diagnostic Features :
A riverine gizzard shad; body fairly deep, its depth 33 to 38% o f standard length,
somewhat compressed, belly with 16 to 20 (usually 17 or 18) + 11 to 14 (usually 11 to 13), total 27 to 33 (usually 29
to 31) scutes.
M outh in ferio r, upper ja w slen d er at tip and d istin ctly turned dow n, second su p ra-m ax illa very
small. Scales small, 51 to 71 in lateral series. R esem b les G. m odesta o f Burm ese rivers, w hich is much deeper
(depth 41 to 45% o f standard length) and has larger scales (44 to 50 in lateral series); CL w hiteheadi o f Burm ese
rivers is also deeper-bodied, and has a subterminal mouth. A nodontostom a species u su a lly h av e only 19 to 21 anal
finrays (usually 23 to 26 in G. m anm ina) and are deeper-bodied. O ther clupeids in the G anges system either have
term in al m ouths (T en u alo sa. G u d u sia, etc.) or a dorsal filam ent (N em atalosa).
G eographical D istribution :
Rivers and associated
w aterbodies o f Sri L anka (fide D ay, 1878:633 - see
Rem arks), India (G anges and other rivers o f O rissa, U ttar
Pradesh, B engal, A ssam ), B angladesh; also A ndam an
Islands (fide H erre, 1940 - perhaps this w as A nodon­
tostom a selangkat).
H abitat and B iology
data needed.
Size :
: R iverine and in pools. M ore
20
°
To 11.5 cm standard length.
0
°
In te r e st to F ish e r ie s
:
C o n trib u te s to in lan d
catches, but no m ajor fishery reported.
Local Names : L iteratu re:
N e lso n & R o th m a n (1 9 7 3 ) g iv e 31
referen ces, m any o f them rep etitiv e. T here appears to be
no published study on the biology o f this species.
40°
80°
100°
120°
140°
160°
18<
Remarks : Day (1878) seems to have been the only
au th o r to cite Sri L anka (fo llo w ed by M unro, 1955), but
neither N elson & R othm an (1973) nor W ongratana (1980) found specim ens; D ay also gave the Indus system, but
again no specimens seem to be known. The A ndam an record by Herre (1940) is perhaps doubtful too.
Gonialosa m odesta (Day, 1869)
C h a to e s s u s m odestus D ay, 1869, P roc.zool.S oc.L ond.:622
CLU P G on 2
(B assein R iv er, B urm a, as high as E en -g ay -g y ee
Lake).
Synonyms :
G o n ia lo s a m odesta - F o w ler, 1941:548 (c o m p ile d ); R e g a n , 1917:315 ( B u rm a ); W h ite h e a d ,
1962:101 (key); Idem . 1971:73, pi. 2a (D a y ’s fig u re) (types o f m o d esta); N e ls o n & R othm an, 1973:149, fig. 5A
(photo) (sy n o n ., descr., refs); W hitehead & Talwar, 1976:155 (types o f m odesta listed ); W o n g ratan a, 1980:187, pis
144, 145 (revision).
FAO Names :
En - Burm ese river gizzard shad.
258
D iagnostic Features :
A riverine gizzard shad; body very deep, its depth 40 to 45% o f standard length,
strongly com pressed, belly w ith 16 to 19 (u su ally 17 o r i 8) +10 to 12 (usually 11), total 27 to 30 (usually 29)
scutes
M outh inferior, upper ja w a little expanded at tip, but distinctly turned down, second supra-m axilla small
(about h alf length o f pre-maxilla). Scales m oderate, 40 to 50 in lateral series.
R esem bles ÇL m anm ina o f Indian
rivers, w hich is much more slender (depth 33 to 38% o f standard length) and has smaller ecales (51 to 71 in lateral
series); sym patric w ith Ch w hiteheadi, w hich has a subterminal mouth and a large second supra-m axilla (longer
than pre-m axilla). A nodontostom a species usually have only 19 to 21 anal finrays (24 to 28 in Ch m odesta) and
m edian instead o f p aired p re-dorsal scales. O ther clupeids in B urm ese freshw aters either have term in al m ouths
(T enualosa. G udusia, etc.), or a dorsal filam ent (N em a talo sa).
G eographical D istribution :
R iv ers and asso ciated
w a te rb o d ie s o f B u rm a (S ittan g R iv er, B assein R iv er as
h ig h as E e n - g a y - g y e e L a k e , a ls o S e lw e in R iv e r a t w
M oulm ein, south to Tenasserim R iver, but presum ably also
in the Irrawady).
H abitat and B iology :
lakes. M ore data needed.
Size :
R iverine and in pools and
20°
To 10 cm standard length.
Interest to Fisheries :
C o n tr ib u te s
catches, but no special fishery reported.
to
in la n d o°
L ocal N am es : 20°
Literature :
N e ls o n & R othm an (1 9 7 3 ) g iv e 12
referen ces, m ost o f th em rep etitiv e. T here appears to be
no published study on the biology o f this species.
40 °
300
Gonialosa w hiteheadi W o n g ra ta n a ,
100°
120°
140 °
CLU P
1983
G onialosa w hiteheadi W ongratana,
1983,
160 °
J a p .J .I c h th y o l.. 29(4):394, fig.
10 (K o k a rie t,
180°
G on 3
Tenasserim River,
Burma).
Synon ym s : N one.
FAO
Names :
En - Southern Burm ese river gizzard shad.
click for n e x t p a g e
c lic k fo r p r e v io u s p a g e
259
D iagnostic Features : A riverine gizzard shad; body very deep, its depth at least 40% o f standard length
(44.6% at 6.8 cm in holotype), strongly com pressed, belly w ith 17 + 10, total 27 scutes. M outh su b term in al, upper
ja w expanded at tip , ap p earin g to tu rn dow n, second sup ra-m ax illa large (lo n g er th an pre-m ax illa).
Scales
m oderate, 43 in lateral series. R esem bles Ch m odesta o f B urm ese rivers, w hich has an inferior m outh and a small
second supra-m axilla (about h alf length o f pre-maxilla); Ch m anm ina o f Indian rivers is m uch more slender (depth
33 to 38% o f standard length) and has sm aller scales ( 5 1
to 71 in lateral series). A n o d o n to sto m a species usually
have o n ly 19 to 21 a n a l finrays (27 in Cf w h ite h ea d i).
O ther clupeids in Burm ese freshw aters either have term i­
nal m ouths (T enualosa. G udusia, etc.) or a dorsal filam ent
(N e m a ta lo s a ).
Geographical Distribution :
Tenasserim River).
H abitat and B iology :
and data needed.
Size :
B urm a (K o k a riet on
R iverine.
M ore
specim ens
To 6.8 cm standard length, perhaps more.
Interest to Fisheries : N ot known.
L ocal N am es :
Literature :
Probably m isidentified as G. m o d e s ta .
Remarks :
P e rh a p s no t u n com m on in B u rm ese
riv ers, but at presen t know n from a single specim en. It
may well have been confused w ith (T m odesta by fishery
w orkers.
2.3
FAMILY PRISTIGASTERIDAE
PRIST
F A O Names : En - P ristig asterid s.
D iagnostic Features :
M oderate or som etimes quite large clupeoid fishes (usually to about 20 to 25 cm
standard length, but som e
South A m erican P ello n a to ab o u t 50 cm ). B ody
co m p ressed , very deep in som e
(P ristig aster), more elongate in others; fully scu ted along belly. The m outh is term inal or more often the lower
jaw projects; there are 2 supra-m axillae, and the ja w teeth are small or minute (but canines in C hirocentrodon);
a toothed hypo-m axillary bone present in some genera, joining the hind tip o f the pre-m axilla to the blade o f the
m axilla.
The dorsal fin is short, its origin before the m idpoint o f the body in very deep form s, but behind in
elongate forms (or missing in R aconda); pectoral fins m oderate or large in the elongate forms; pelvic fins small,
with 6 or 7 finrays, but absent in some genera; anal fin long, with at least 30 finrays (60 or more in some species).
The scales may be adherent or easily lost, o f m oderate size (about 35 to 55 in lateral series).
U ntil recen tly , the p ristig aste rin e s w ere co n sid ered a subfam ily o f the C lupeidae. N elso n (1967:392) found
unique features in th e ir gili arches (to o th p lates fused to second b asib ran ch ial and to one or m ore pairs o f
hypobranchials, especially the second pair); he raised them to the rank o f super-fam ily. Grande (in press) did the
sam e, adding tw o fu rth er d istin ctiv e characters: p re-d o rsal bones eith er u p rig h t o r in clin ed forw ard (in clin ed
backward in all clupeids, except upright in R am nogaster), and no gap betw een the second and third hypural bones
o f the tail (upper and lowercau d al finrays separated by this gap in clupeids).
Biology, H abitat and D istribution :
P ristig aste rid s
are
m ostly m arine coastal and schooling fishes o f
tro p ical and som e subtropical seas.
They are found on both sides o f the P ac ific and A tlan tic O ceans and
throughout the Indian ocean, from about 30 N (southern Japan) to about 30°S (South A frica). Some species enter
estuaries and a few species in South A m erica appear to be restricted to freshw ater (P ellonai
Interest to Fisheries :
N o special fish eries ex ist for p articu lar species and the catches for only three
species are reported (Ilisha elongata. Ilisha african a and Pellona ditchela - 21 885 to n s in 1983). N e v e rth e le ss,
pristigasterids are common in tropical fish markets and probably make a useful contribution to clupeoid catches.
There are 9 genera (3 w o rld w id e , 5 N ew W orld and 1 Indo-Pacific) and 33 species (16 N ew W orld, 1 W est
A frica, 16 Indo-Pacific).
260
Key to the G enera
A.
WEST A F R I C A .................................................
B.
NEW WORLD
la .
Toothed hypo-m axilla present (Fig. la)
2a.
2b.
lb.
Ilish a
ligam ent (no toothed
hypo-m axilla)
P e lv ic fin s p r e s e n t;
A tla n tic
coasts and drainage ...............................
No pelvic fins; Pacific coasts .
P ellon a
P lio ste o sto m a
toothed
hypo-m axilla
b) Ilisha
a) Pellona
Fig. 1
No toothed hypo-m axilla (Fig. lb)
3a.
P elvic fins p resen t, body elongate
(if not, see 5b)
4a.
4b.
3b.
Jaw te e th sm a ll or m in u te ,
absent at centre o f upper jaw . . .
Ilisha
C a n i n e - l i k e t e e t h in b o th
ja w s, also at centre o f upper
ja w (Fig. 2 ) ............................. Chirocentrodon
No pelvic fins (or if present, then
body very deep indeed, see 5b)
5a.
B o d y e lo n g a te , d e p th up to
35% o f standard length; d o r­
sal fin origin behind midpoint
o f body
M axilla
re a c h in g
eye
7a.
7b.
6b.
sh o rt,
not
back beyond
C h iro cen tro d o n
Fis- 2
pre-m axilla
M outh
te rm in a l,
ja w s e q u a l; h in d
tip o f pre-m axilla
m e e tin g
m a x illa
b l a d e ( F i g . 3a);
a n a l finrays 5 0
no gap
o r le s s ;
A tla n ­
tic
and P a c if ic
c o a s ts
N eoopisthopterus
m axilla blade
L o w e r ja w p r o ­
je c tin g , a d istin ct
a. N eoopisthopterus
gap b etw een hind
tip o f pre-m axilla
and m axilla blade
(Fig.
3b);
anal
finrays
m ore
th an 50;
P a c ific
c o a sts ................. O pisthopterus
gap
O pisthopterus
Fig. 3
M a x illa lo n g , re a c h in g
to
or
beyond
gili
opening (Fig. 4); A tlan ­
tic
and
P a c ific
O dontognathus
c o a s t s .....................
m a x illa tip
O dontognathus
Fig- 4
261
5b.
B o d y v ery d e e p , d e p th a b o u t 50% o f
standard length (Fig. 5); dorsal fin origin
b efo re m id p o in t o f b o d y ; sm all p e lv ic
fins sometimes present; fresh w ater...................P riitigaster
C.
INDO-PACIFIC
la .
Toothed hypo-maxilla present (Fig. l a ) ....................... Pellona
lb .
No toothed hypo-m axilla (Fig. lb)
2a.
2b.
D orsal fin present
3a.
A nal fin m o d erate, 34 to 53
fin ra y s pelvic fins p re s e n t........................Ilisha
3b.
A nal fin lo n g , 51 to 65 fin ­
rays; no pelvic f in s ....................Opisthopterus
No dorsal fin; no pelvic fins; anal
fin very long, 81 to 92 f i n r a y s ...................
P ris tig a s te r
Fig. 5
Raconda
Ilish a R ich ard so n , 1846
F orm erly C LU P Ilish
Ilisha R ichardson, 1846, Ichthvol.Seas C hina Jap an :306 (type: Ilisha abnorm is R ichardson = A losa elongata
B ennec).
Platygaster Sw ainson, 1838, N at.H ist.F ish es. 1:278 (type: C lupea africana B loch, 1795) (preoccupied in
H ym enoptera). Zunasia Jordan & M etz, 1913, M em .Carnegie M ys., 6(1):7 (ty p e P ristig a ste r c h in en sis B asilew sk i
= A lo sa e lo n g a ta B e n n e tt). P se u d o c h iro c e n tro d o n M ira n d a -R ib e iro , 1923, P u b lc o e s L in h a s T e le g r.e s tra t.
M atto G rosso A m a z o n a s, (58): 8
(type: P seud o c h iro c e n tro d o n am azo n icu m M iran d a -R ib e iro ). E u p la ty g aster
F o w le r, 1934, P ro c .A c a d .n a t.S c i.P h ila d ., 8 5 :2 4 6 (ty p e : P e llo n a b ra c h y s o m a B le e k e r = C lu p e a m e la sto m a
Schneider).
D iagn ostic F eatu res :
M o d erate-sized m arine, estu arin e or fresh w ater clup eo id fish es (to ab o u t 40 cm
standard length), body m oderately deep in some, elongate in others, com pressed, w ith a sharp keel o f scutes along
belly. Eye usually large; low er ja w p ro jectin g , m outh d irected o b liquely upw ard; upper ja w reaching at m ost to
eye centre; no too th ed hy p o -m ax illa b etw een hind tip o f p re-m ax illa and blade o f m axilla; ja w teeth sm all or
m inute, usually w ith a d istin c t gap at centre or u p p er jaw . G illrakers fairly short and thick, few, 17 to 28 on
low er arch. D orsal fin before or behind m idpoint o f body; pelvic fins present, small, w ith 6 or 7 finrays, a little
or well in front o f dorsal fin origin; anal fin m oderately long, w ith 34 to 53 finrays, its origin under or a little
behind dorsal fin base. Scales m oderate, about 38 to 56 in lateral series. No distinctive colour patterns. The lack
o f a toothed hypo-m axilla distinguishes Ilisha from the otherwise sim ilar P ellona.
B iology, H ab itat and D istrib u tion :
M arine, p elagic, inshore, probably schooling, some or m ost species
entering bays and estuaries and presumably able to tolerate at least a slight lowering o f salinity, but a few appear
to be purely freshw ater species. W orldwide in tropical and subtropical w aters, occurring throughout the Indo-W est
Pacific, o ff W est A frica, and o ff A tlantic and Pacific coasts o f central and South A m erica
Interest to Fisheries : Only tw o species are individually reported: Ilisha elongata o ff coasts o f China and
K orea (in 1983 the catch w as 15 338 tons) and Ilisha a frican a o ff W est A frican coasts (4005 tons in 1983). Other
species contribute to general clupeoid catches, chiefly by artisanal gear.
262
Species : The 11 Indo-W est
W est African species is reviewed in
Pacific species w ere revised by W ongratana (1980), the literature for the single
CLOFETA, but more work is needed on the 2 New World
species.
Indo-West Pacific
L
L
L
L
L
L
L
L
L
L
L
elongata (Bennett, 1830), Madras to southern Japan
filigera (Valenciennes, 1847), Bombay to Sarawak
kampeni (W eber & de Beaufort, 1913), Bay o f Bengal to Indonesia
m acrogaster Bleeker, 1866, Indonesia
m egaloptera (Swainson, 1839), Bombay to Java Sea
melastoma (Schneider, 1801), India to Indonesia and Taiwan Island
novacula (V alenciennes, 1847), Rivers o f Burma
obfuscata W ongratana, 1983, India
pristigastroides (Bleeker, 1852), Indonesia
sirishai Seshagiri Rao,
1975, the “G u lf’ to Thailand
striatula W ongratana, 1983, A rabian Sea to northern Australia.
W est A frica
I. africana (Bloch, 1795), Senegal to Angola.
New W orld
L am azonica (M iran d a-R ib eiro , 1923), A tlantic drainage (A m azon o f B razil, Peru)
L furthii (Steindachner, 1875) Pacific coasts and drainage.
R em arks : M any o f the In d o -P a c ific species were seriously confused until fundam ental differences in the
sw im bladder w ere found b etw een su p erficially very sim ilar species (S esh ag iri R ao, 1974, 1976;
R am aiy an &
W h iteh ead , 1975;
R a m a iy a n & N a ta ra ja n , 1979; m ost co m p letely by W o n g ratan a, 1980).
T hree ty p es o f
sw im bladder occur:
A.
Two slender tubes from hind
end o f s w im b la d d e r p a ssin g
b a c k in to th e m u s c le s on
e it h e r s id e o f th e h a e m a l
s p in e s (I k a m p e n i, L m ela­
s to m a , L o b fu sc ata , I. stria tul ; also I africana o f W est
A fric a )
1st haem al
^ spine
sw im bladder
— tw o
p o st coelom ic
tu b es
body cavity
anus
A single long tube from hind
end o f s w im b la d d e r p a ssin g
back through m uscles o f right
side o f body (L elo n g ata,
L f ilig e r a , L m a c ro g a s te r,
L m e g a lo p te ra , L pristigas­
troides).
N o tube or a very sh o rt one
at hind end o f sw im bladder,
not passing back into muscles
(L novacula, L sirishai; this
is also th e
co n d itio n
in
Pellona species).
one
p o st coelom ic
tube
(right
side)
no
p o st coelom ic
tube
This ch aracte r is inclu d ed in the d iag n o ses, but w here p o ssib le o th er d iag n o stic featu res are also used to
separate the species.
263
P R IS T Iii sh 5
Ilisha african a (B loch, 1795)
F orm erly C LU P Ilish 5
C lupea african a Bloch, 1795, N atu rg e sc h .A u sl.F isch e, 9:45, pi. 407 (Accra, Ghana).
Synonyms : Pellona
iserti Valenciennes, 1847:307 (Accra); Pellona gabonica D um eril, 1861:259, pi. 23, figs
3, 3a (G abon); P ristig aster dolloi B o ulenger, 1902:271, pi. 30, fig.3(B anana, C ongo); Ilish a m elanota D erscheid,
1924:278 (Congo m outh ; Irvine, 1947:111 (A ccra); Ilisha africana - Fow ler, 1936:78, fig.71(C ongo mouth; synon.,
descr.); W h iteh ead , 1967:112 (ty p es o f is e rti, g a b o n ic a , d o lii, m e la n o ta ); Id e m , 1969:268 (ty p e o f a fric a n a );
CLO FETA in press (com plete synon.).
F A O Nam es : En - W est A frican ilisha.
D iagnostic F eatures :
B ody m oderately deep, com pressed,
belly w ith 25 to 27 + 7 or 8 sharp scutes.
Eye la r g e , lower jaw
projecting.
D orsal fin at or before m idpoint o f body; p elvic fins
sm all; anal fin lon g , w ith m ore th an 40 finrays, its o rigin below
dorsal fin base. Sw im bladder w ith two short tubes passing back in the
muscles on either side o f haemal spines. Other clupeoid fishes in the
area are m ore slend er, have a much shorter anal fin and the lower
jaw not strongly projecting. See CLUP Ilish 5, Fishing A reas 34, 47
(part).
G eographical D istrib u tion :
W est A frican coasts and
estu aries (n o rth ern parts o f Senegal south to B aie de SaintBras, A ngola, at 10°S).
H abitat and B iology :
M arine, pelagic, along beaches
and ju st o ff shore, down to about 25 m, also in lagoons and
e stu arie s, p enetrating into alm o st fresh w ater. F eeds on sm all
planktonic anim als (crustaceans, etc.).
Size
16 cm.
40-
20
-
To 20 cm standard length, usually about 12 to
Interest to F isheries :
N o special fishery, but co n tri­
b u te s to c lu p e o id fis h e rie s (a b o u t 4 0 0 to n s m a rk e te d at
A bidjan in some years). M ainly caught by trawl, but also in
p urse-seine.
The total catch in 1983 was 4005 tons.
Local N am es :
ANGOLA:
F an h ico (P o rtu g u ese);
D A H O M EY : A flém a (F on), K aflan (at G rand P o p o ), K aflanvi
(a t C o to n o u ); G H A N A :
K a n tia and v arian ts (F an te), also
T a m te m ire (F a n te );
GUINEA:
L ati (S o u sso u s); IV O R Y
COAST:
L a la (L ak e Ebrié), N a f r a n ( A p p o lo n ie n ) , Péténana 40°
ju a b a (B rignan);
SENEGAL:
R im bal (O v o lo f), T ialim aro
(M andique).
sw im bladder
bodv cav it\
tubes
264
L iterature :
W hitehead (i.e. C LO FETA , in press) gave 61 references up to
biological and fishery literature.
Ilisha amazonica (M iranda-R ibeiro,
PR IST Ilish
1923)
P seu d ochirocen tro d o n
am azonicum
A m azonas, (58): 8 (M anáos, A m azon).
1981 covering taxonom ic,
M iran d a-R ib eiro ,
14
1923, Publçoes L inhas T e le g r .e s tr a t.M a tto Grosso
Synonym s:
Ilish a iq u ite n sis N a k a sh im a , 1 9 4 1 :6 6 , fig. (n e a r Iq u ito s, P e ru v ia n A m azo n ); Ilish a ap apae
F lild e b ra n d , 1948:3, fig. 2 (A m azo n , b etw een M anáo s and P ará ); Ilish a a m az o n ica - M y ers, 1 9 5 0 :6 3 ( t y p e s o f
am azonicum , apapae, synon.); FW NA, 1 9 6 4 :4 2 1 , fig. 105 (synopsis); W hitehead, 1970:21 (key only).
F A O N am es : E n - A m azon ilisha.
D iagnostic Features : Body m oderately elongate, com pressed, belly w ith 25 or 26 scutes. Eye large, lower
jaw projecting; gap between hind tip o f pre-maxilla and blade o f maxilla a ligament, not a toothed bone; lower
g illra k e rs 17 to 22.
D o rsal fin o rig in w ell b efore
m idpoint o f body; pelvic fins present;
anal fin long,
w ith 47 to 52 finrays. R esem bles species o f P ello n a,
w hich have a distinct toothed hypo-m axilla betw een the
hind tip o f the pre-maxilla and the blade o f the maxilla.
O ther sim ilar clup eo id s are eith er m uch deep er-b o d ied
(P r is tig a s te r , depth about 50% o f standard len g th ), or
lack pelvic fins (O d o n to g n ath u s, O p isth o p te ru s, N e o o ­
pisthopterus) or have canine teeth (C h iro c e n tro d o n ).
G eographical D istribution :
and M anaos up to Iquitos in Peru).
A m azon (from Pará
H abitat and B iology : R iverine, apparently
entering sea. More specim ens and data needed.
Size :
not
To 17.3 cm standard length.
Interest to Fisheries :
N o data. The paucity o f
m useum specim ens may not reflect an actual rarity o f
the fish.
Local Names : BRA ZIL: Apapa, A suna nahui.
Literature :
descr., range).
H ildebrand (i.e. FW N A, 1964 -
265
PR IT
Ilish a elo n g a ta (B e n n e tt, 1830)
A losa elongata B ennett, 1830, M em .Life
July).
Ilsh
2
o f R affles:691 (Sum atra) (exact date not known, presum ed before
Synonyms :
C lupea affin is G ray, 1830 (15 July):pl. 96, fig. 2 (In d ia, on H ard w ick e draw ing; d atin g by
Saw yer, 1953); Ilish a abnorm is R ich ard so n , 1846:306 (Macao or C a n to n );
P ello n a lesch en au lti V alen cien n es,
1847:311 (Pondicherry);
P e llo n a g ra y a n a V a le n c ie n n e s , 1847:315 (on Clupea affinis Gray); P e llo n a v im b e lla
V a le n c ie n n e s , 1 8 4 7 :3 1 7 (M a c a o ) P e llo n a s c h le g e lii B le e k e r , 1 8 5 4 :4 1 8 N a g a s a k i) ; P r is tig a s te r c h in e n s is
B asilew ski, 1855:243 (G ulf o f Tschiliensi, China); P ris tig a s te r (P ristig aster) sin en sis Sauvage, 1881:107 (Sw atow );
Ilis h a e lo n g a ta - F o w ler, 1941:661 (B o m b a y ?; Jak arta, H ong K ong, K o rea, Japan; larg e synon.); S v e to v id o v ,
1952:321, pi. 26, fig. 2 (Peter the Great Bay, 1914to 1938); Idem , 1963:361, pi. 26, fig. 2 (sam e); Chu, Tchang &
C hen, 1963:104, fig. 79 (China); W hitehead, 1966:32, pi. 3, fig. 2 (R eeves draw ing) (type o f abnorm is); Idem .
1967:118, 119, 120, 121 (types o f leschenaulti,
grayana, vim bella); Id em . 1973b:211, fig. 36 (Key, sy n o n .);
R am aiyan & W hitehead, 1975:195 (key, d e s c r .);
W ongratana, 1980:204, pis 165, 166 (revision); Masuda et ah,
1984:20, pi. 22D (colour photo) (southern Japan); W hitehead & Bauchot, in press (types o f V alenciennes species).
F A O Names :
E n - E longate ilisha.
D iagnostic Features : B ody slen d er, its d ep th 27 to 31% o f
standard length, belly w ith usually 24 to 25 + 10 to 15, total 34 to 42
scutes. Eye large, low er ja w p ro jectin g ; low er g illrak ers 19 to 25.
swimbladder
\
Dorsal fin origin at about m idpoint o f body or a little behind; anal fin
1 tube
w ith 43 to 53 finrays, its origina a little b eh in d d o rsa l fin b ase.
on right
Sw im bladder w ith a single long tube passing back down right side o f
bodv cavit\
side
body a b o v e anal fin b a se . R e se m b le s
m e g a lo p te r a .
fi I i ge ra
E m acrogaster and E p ristig astro id es in th i s sw im bladder ch aracter,
b ut these are either deeper-bodied (depth 31 to 36% ) or have only 28
to 35 scutes (E m egaloptera). O ther Ilisha species have either tw in tubes from the hind end o f the sw im bladder
(and usually only 25 to 30 scutes) or no posterior tubes (and more or fewer scutes). Pellona species are d eep erbodied and have a distinct toothed hypo-m axilla. See CLUP Ilish 2, Fishing Areas 57, 71.
G eographical D istribution :
Indian
O cean
(Pondicherry, b a s e d s o le ly o n th e ty p e o f P e llo n a
le sc h e n a u lti; no other confirm ed Indian records) JavaSea (S in g ap o re), E a st C hina Sea (C an to n n o rth to the
K o re a s and s o u th e rn Ja p a n , as fa r as O sa k a on th e
P acific co ast and F ukuoka in Sea o f Jap an , also P eter
the Great Bay in Soviet waters).
Habitat and Biology :
M arin e, pelagic and in ­
shore, en tering estu aries and presum ably able to to le ­
rate lo w e re d sa lin itie s .
A w a rm -w a te r sp e c ie s th a t
does not reach the n o rth ern p art o f its range in cool
y ears.
Size
To 40.5 cm standard length.
266
Interest to Fisheries : The only Indo-Pacific species o f Ilisha for w hich individual statistics are reported.
In 1983 the catch was 15 338 tons from China and the Koreas.
L ocal N am es : L iteratu re :
See under Synonyms.
Rem arks : The westw ard extension o f the range o f
L elongata to India is based on a single specimen (see
above). H ow ever, befo re the d ifferen ces in sw im b lad d er form w ere noted and used to separate the species by
R am aiyan & W hitehead (1975), Indian specimens o f Ilisha may have been reported under other names.
Ilish a
filig e r a
(V alenciennes,
1847)
PRIST
Ilish 6
P ello n a filig era V alen cien n es, 1847, H ist.n a t.p o iss., 20:322 (Bom bay, Corom andel coast) (in part, the other
specimen being Ilisha o b fu sca ta ).
Synonym s : Pellona xanthoptera B le e k e r, 1851:439 (Sam bas, K alim antan, Indonesia); W eber & de B eaufort,
1913:94 (Java, K alim antan); Ilish a x an th o p e ra :Fow ler, 1941:658 (com piled); W hitehead et ah, 1966:96, pi. 12, fig.
1 (B le e k e r ’s fig .) (types o f x an th o p era ); Ilish a filig era - F o w ler, 1941:658 (co m p iled ); W h iteh ead , 1967:117 (types
o f filigera); Idem . 1973b:213 (in synon. o f L megaloptera); W ongratana, 1980:206, pis 167, 168 (revision); B auchot
& W hitehead, in press (types o f filigera).
FAO Names
E n - C orom andel ilisha.
D iagnostic Features : Body m oderately slender, its depth 31
to 35% o f standard length, belly with 23 to 26 + 11 to 13, total 34 to
sw im bladder
38 scutes. Eye large, low er jaw projecting; low er gillrakers 19 to 23.
Dorsal fin origin at about m idpoint o f body or a little behind; anal fin
with 46 to 52 finrays, its origin below dorsal fin base.
Sw im bladder
w ith a single
long tube passing back down right side o f body above
anal fin base.
R e s e m b le s four o th e r In d o P acific sp e c ie s o f Ilish a in
this sw im bladder character, but these have few er scutes (L m eg alo p ­
t e r a ). fe w e r gillrakers (L p r i s t i g a s t r o i d e s ), or m ore g illrak ers
(L m acro g aster), or are m ore slen d er (L elo n g ata ). O ther Ilish a species have eith er tw in tu b es from the hind end
o f the swim bladder (and usually only 25 to 30 scutes) or no posterior tubes (and few er scutes or a more slender
body). Pellona species have a distinct toothed hypo-maxilla.
267
G eograp h ical
D istrib u tio n
In d ian
O cean
(B o m b ay to e a s te rn c o a sts o f In d ia ; p e rh a p s e a st to
A n d a m a n Sea c o a s ts , b u t no c e rta in re c o rd s ), S o u th
C h in a S ea (n o rth w e s te rn c o a sts o f K a lim a n ta n , a lso
Sarawak).
H ab itat and B iology : M arine, pelagic and
c o a s ta l, a p p a r e n tly e n te r in g e s t u a r ie s (W eber & de
B e a u fo rt, 1913 - as P ello n a x a n th o p te ra ). M o re d a ta
needed, based on better identifications than in the past.
Size : To 22 cm standard length, perhaps more.
Interest to F isheries : C ontributes to
catches o f clupeoids, but no special fishery.
artisanal
Local N am es
IN DIA: Phansa (C alcutta, but
p ro b a b ly th e sam e n am e u sed fo r o th e r sp e c ie s o f
Ilisha).
Ilisha furthii (S teind ach n er,
1875)
PR IST Ilish 7
P ello n a furthii S tein d ach n er, 1 8 7 5 , Sber.A kad.W iss.W ien. 70:388 (B ay o f P an am a); Id em , 1 8 7 5 : I c h th y o l.
B e i t r .,( l ) : 14 (r e p e a t).
Synonyms :
P ello n a p an am en sis S tein d ach n er, 1875:389, (P a n a m a );
Idem . 1875:15 ( r e p e a t ) ; M e e k &
H ild e b ra n d , 1923:189
(brackishw ater at C orozal and from P an am a m ark et);
H ild e b ra n d , 1946:91 (o f f P u erto
Pizarro and m outh o f Tumbes River, G u lf o f G uayaquil, Peru; also G uayaquil, Ecuador); Peterson, 1956:180 (C hira
Island and B arranca Bay, G u lf o f N icoya, Costa Rica); Cobo & Massay, 1969:7 (E cuador - listed).
F A O Names :
E n - P acific ilisha.
D iagnostic Features : Body m oderately slender, com pressed, belly w ith 23 to 25 + 11 to 14 sc u te s . Eye
large, low er jaw projecting; gap between hind tip o f pre-m axilla and blade o f m axilla a ligam ent, not a toothed
bone; low er g illrak ers 20 to 25. Dorsal fin origin a little before m idpoint o f body; anal fin long, w ith 46 to 50
finrays, its o rig in b e lo w a b o u t la s t d o rsa l finrays.
O ther sim ilar clu p eo id s in the area have no p elvic fins
(P li o s t e o s to m a , w hich has a toothed h ypo-m axilla betw een pre-m axilla tip and blade o f m axilla; O d o n to g n ath us,
w hich has a long m ax illa reaching back beyond eye;
N e o o p isth o p te ru s, w hich like th e o thers is m uch m ore
elongate).
268
G eograp hical
D istrib u tion
E astern
central
Pacific (C osta R ica in G u lf o f N icoya to P anam a and
south to G ulf o f Guayaquil).
H ab itat and B iology
M arine, co astal, but
en tering estu aries and o th er b ra ck ish w ater, p erhaps to
spawn there.
Size : To 28.5 cm standard length.
Interest to Fisheries :
E n ters m ark ets (e.g. at
Panama) and is apparently a common food fish.
Local
N am es
ECU A D O R,
PERU:
M achete.
Literature : Peterson (1956 - notes on m aturity
and ecology).
I lis h a k a m p en i (W eber & de B e a u fo rt, 1913)
P e llo n a k a m p e n i W e b e r & de B e a u fo rt,
Balikpapan, K alim antan).
P R IST Ilish 8
191 3, F is h e s In d o -A u stralian A rc h ., 2:87 (J a k a rta , Ja v a and
Synonym s : Ilisha w hiteheadi Seshagiri Rao, 1974:861, figs 1-3 (K ak in ad a, Bay o f Bengal): Ilisha kampeni Fow ler, 1941:654 (c o m p ile d ): R a m a iy a n & W h iteh ead , 1975:196, fig. 2 a-e (key, descr., sw im b la d d e r: K a k in a d a ,
Porto Novo): W ongratana, 1980:208, pis 171, 172 (revision).
FAO Names
En - K a m p en ’s ilisha.
click fo r n e x t p a g e
c lic k fo r p r e v io u s p a g e
269
D iagnostic Features : Body
m oderately slender, its depth 24 to
32% o f standard length, belly w ith
striae not
usually 19 to 21 - 8, total 27 to 29
continuous
scutes. Eye large, low er ja w p ro ­
je c tin g ; low er g illrak ers 20 to 24.
2 tubes
Dorsal fin origin at about m idpoint
o f body;
anal fin w ith 38 to 46
sw im bladder
finrays, its orig in u n d er hind part
o f dorsal fin base. V ertical
striae
on scales not continuous, but w ith
bodv cavitv
a d is ti n c t g ap a c ro s s c e n tr e o f
scale. S w im bladder w ith tw o tubes
p a ss in g b ack in th e m u s c le s on
e ith e r
s i d e o f h a e m a l s p in e s .
scale
Resem bles three other In do-P acific
species o f I l i s h a in th i s sw im ­
bladder ch aracter, but th ese have m ore g illrak ers (L o b fu scata), scale
striae th at trav erse the scale, also body
deeper (I. m elastom a) or are deeper and have longer pectoral fins (L striatula; pectoral
fin 18 to 21% o f standard
len g th , cf. 15 to 17%). O ther Ilisha species have either a single long tube from the hind end o f the sw im bladder
(and m ore scutes), or no p o sterio r tu b es (and a deep er body or m ore scutes). P ello n a species have a d istin ct
toothed hypo-m axilla.
G eograph ical
D istrib u tio n
In d ian
O cean
(eastern coasts o f In d ia n orth to C alcu tta), In d o n esia
(Jak arta, Java; also K alim an tan at K o tab aru , T akisung
on southeastern coast and at Aluhaluh on B arito River).
Records o f this species from the A rabian Sea (Seshagiri
Rao, 1975) need confirm ation.
H abitat and B iology : M arine, pelagic, coastal,
but also entering riv ers and to le ra tin g w ater o f low
salinity.
F eeds on p lan k to n ic cru stacean s and fish es,
also amphipods.
Size : To 15 cm standard length, perhaps more.
Interest to F isheries : C ontributes to artisanal
clupeoid catches, but no special fishery.
C au g h t by
cast nets in the V ellar estuary, Porto N o v o , India.
L o ca l N am es : Literature :
R am aiyan & Whitehead (1975 - brief notes on distribution and biology).
R em arks : Long regarded as an Indonesian species, its presence o ff Indian coasts was unsuspected until
reported by R am aiyan & W hitehead (1975) and the revision by W ongratana (1980).
A n ap p aren tly d istin ct but
closely related species from the A rafura and Tim or Seas, G u lf o f Carpentera and o ff Papua N ew Guinea w ill be
d escribed by D r P atricia K ailola. It differs from L kam peni ch iefly in h aving the striae co n tin u o u s acro ss the
scale, slightly more pyloric caeca (19 to 24; cf. 15 to 19), few er gillrakers (18 to 20; cf. 20 to 24) and more
v ertebrae (43 to 45; cf. 41 or 42).
Ilisha macrogaster Bleeker,
P R IS T
1866
Ilisha m acrogaster B leeker, 1886, N e d .T ijd sch r.D ierk ., 3:300
Ilish 9
(Sam bas, K alim antan, Indonesia).
Synonym s : Pellona m acrogaster:W eber & de B eaufort, 1913:93 (on B leeker); Ilisha m acrogaster - F o w ler,
1941:660 (com piled);
W hehead e t ah, 1966:98, pi. 12, fig. 2 (B leeker’s fig.) (type o f m ac ro g aste r) W hitehead
1973b:211 (key only); W ongratana, 1980207, pi. 169 (revision).
F A O Names :
En
K alim an tan
ilisha.
270
Diagnostic Features : Body moderately deep, its depth 34 to
36% o f standard len g th , belly w ith 25 + 1 1 , total 36 scutes. Eye
sw im bladder
larg e, low er ja w p ro jectin g ; low er g illrak ers 23 to 25. D orsal fin
o rigin at about m idp o in t o f body;
an a l fin w ith 49 finrays, its
1 tube
origin under dorsal fin base. Sw im bladder w ith a single long tube
on right
body cavity
p a s s in g b a c k d o w n r i g h t sid e o f b o d y a b o v e a n a l fin b a s e .
side
R e se m b le s fo u r o th e r In d o -P acific species o f Ilisha in this sw im ­
b la d d e r c h a ra c te r, b u t th e s e a re m o re slender. (L e lo n g a t a , L
p ristig a stro id e s), have few er scutes and g illrak ers (L m e g a lo p te ra )
or fewer gillrakers (T filigera). Other Ilisha species-have either tw in tubes from the hind end o f the swim bladder
(and usually only 25 to 30 scutes) or no posterior tubes (and few er scutes or few er g illrak ers). P ello n a species
have a distinct toothed hypo-maxilla.
Geographical Distribution :
(w estern K alim antan, Sarawak).
S outh C hina Sea
Habitat and Biology : M arine, pelagic and coas­
tal, perhaps entering estuaries. M ore data needed.
Size :
To 12 cm standard length.
Interest to Fisheries : No data.
Local Names : Literature :
See under Synonyms.
Remarks :W o n g r a t a n a (1980)was able to exa­
mine only tw o specim ens; m ore m aterial m ay show
that this species is merely L filigera.
Ilish a m eg a lo p tera (Swainson,
1839)
PRIST Ilish 4
F orm erly C LU P Ilish 4
Platygaster
m egalopterus
Sw ainson, 1839, N at.H ist.A nim ., 2:294 (on Jangarloo o f Russell,
1803).
Synonym s : ? Clupanodon motius H am ilton-Buchanan, 1822:251 (Brahm aputra; nomen dubium ); Platygaster
m acrophthalm a Sw ainson, 1838:278 (on Jangarloo o f R ussell, 1803; nom en oblitum ); ? P latygaster parva
S w ainson, 1839:294 (on C lu p ea m otius o f G ray , 1834); P e llo n a d u ssu m ieri V alen c ien n es, 1847:316, pi. 516
(Bombay, M alabar, Coromandel ; Pellona russellii B leeker, 1852:72 (Java, M adura, Pasuruan, Singapore); ? Ilisha
m elastom a: Fow ler, 1941:659 (Kalim antan;
m egaloptera in synon.); Ilisha m egaloptera - W hitehead, 1967:113,
pi. 7c (V alenciennes figure) (type o f dussum ieri; m acrophthalm a rejected); Idem , 1973b:213, fig. 38 (key, synon.,
descr.; filig e ra , x a n th o p te ra , ru ssellii w rongly in cluded ; R am aiyan & W h iteh ead , 1975:191, fig. Id (key, descr.;
Bombay to Singapore, Sarawak to Macao); W ongratana, 1980:201, pis 163, 164 (revision); Whitehead & B auchot, in
press (type o f dussum ieri).
FAO Names:
En - Bigeye ilisha.
D iagnostic Features : Body
rather deep, its depth 28 to 39%
o f sta n d a rd le n g th , b e lly w ith 19 to 23 +8 to 12, to ta l 28 to 35
sw im bladder
(usually 30 to 34) scutes. Eye large, low er ja w strongly projecting;
low er gillrakers 18 to 23. Dorsal fin origin near m idpoint o f body;
anal fin with 38 to 53 finrays, its origin below hind part, o f dorsal fin
base. Sw im bladder w ith a single long tube passing back down right
side o f body above anal fin base. Resem bles L elongata, L filigera,
L m acrogaster and L p ristig a stro id es in th is sw im bladder ch aracter,
but these have more scutes total 34 to 42).
O ther Ilisha species have
either tw in tubes from the hind end o f the sw im bladder (and usually only 25 to 30 scutes) or no posterior tubes
(and more or few er scutes). Pellona species
have a distinct toothed hypo-m axilla. See CLUP Ilish 4, Fishing Area
51, also 57, 71.
G eograp hical
D istrib u tio n
In d ian
O cean
(B o m b a y t o B ay o f B engal and A n d am an co ast o f
Thailand), Java Sea (o ff Java, Singapore). Saraw ak, 40°
‘C o c h in c h in a ’ and Macao specim ens o f R am aiy an &
W hitehead (1975) should be rechecked.
20°
H abitat and Biology : M arine, pelagic, inshore,
b u t a p p a r e n tly o c c u rs a ls o in r iv e r s (G a n g e s at
A lla h a b a d , r iv e r s o f e a s te r n U tta r P r a d e s h - see
W o n g ratan a, 1980), but not in V ellar estuary (Ram aiyan
& W hitehead, 1975); D ay (1875:644) referred to Pellona
m o tiu s ( i f th is w as in d e e d th e p re s e n t species) as 2QO
riverine, “descending as low as the coast” and gave its
size as n o t m u ch o v e r 4 in c h e s , i.e. a b o u t 10 cm
standard length; this would suggest that L m egaloptera
ascends and spawns in rivers. Feeds on fis h e s , crusta- 40°
ceans, am phipods, o ccasio n ally p o ly ch ae tes, tu n icates
and small amounts o f algae and diatoms.
Size :
To 27.5 cm standard length.
Interest to Fisheries : No separate statistics, but it is perhaps the com m onest species o f Ilisha in the
In d ian O cean and m ust m ake a sig n ifican t c o n trib u tio n to a rtisan al catc h es o f clupeoids. C au g h t m ainly by
gillnets and boat seines in India (even found in trawl catches from 30 to 40 m).
Local Names :
INDIA:
Phansa (Calcutta, but probably the same name used for other species o f Ilisha).
Literature : R am aiyan & W hitehead (1975); other identifications in the literature cannot be w holly relied
upon.
272
Ilish a m elastom a (Schneider,
1801)
PRIST Ilish 3
F orm erly C LU P Ilish 3
Clupea m elastom a Schneider, 1801, S y st.Ich th y o l.B loch. :427
(near Coromandel).
Synonym s : Platygaster verticalis Sw ainson, 1838:278 (on D itchoee o f R ussell, 1803); Platygaster indicus
S w ainson,1839:294 (on D itchoee); Pellona ditchoa V alenciennes, 1847:313 (on D itchoee); Pellona m icropus
V a le n c ie n n e s, 1 8 4 7 :3 2 0 ( C o r o m a n d e l c o a st; in p a rt, th e o th e r sp e c im e n b e in g Ilis h a o b fu s c a ta ) ;P e lIo n a
brachysom a B leek er, 1852:22 (Jak arta); W eb er & de B eau fo rt, 1913:87 (on B leek er); Ilish a b rac h y so m a :F ow ler,
1941:653 (Padang, Sum atra); W hitehead et ah, 1966:100, pi. 13, fig. 1 (B leek e r’s fig.) (type o f brachysom a); Ilisha
in d ica:Fow ler, 1941:652 (B om bay, C alicut, Jakarta, large synon.);W hitehead, 1967:116 (type o f ditchoa); Ilisha
m ic ro p u s:W h iteh ead , 1 9 6 7 :1 1 5 ,1 1 6 (ty p e o f m ic ro p u s); Ilish a m elasto m a - F o w ler, 1941:659 (S an d ak an B ay,
K alim antan, large synon.); W hitehead, 1973b:212, fig. 37 (key, synon., refs); R am aiyan & W hitehead, 1975:189,
fig. la-c (key, descr.; Karachi to Hong Kong); W ongratana, 1980:211, pis 175, 176 (revision).
FA O Names :
E n - In d ian ilisha.
Diagnostic Features :
Body
stria e
m o d e ra te ly d e e p , its d e p th 33 to
continuous
42% o f standard length, belly w ith
or
usually 17 to 21 + 8 or 9, total 25
overlapping
to 30 scutes. Eye larg e, low er ja w
projecting; low er g illrak ers 21 to
l í 2 tubes
25. Dorsal fin origin a little before
m idpoint o f body; anal fin w ith 35
sw im bladder
to 48 fin ray s, its origin under hind
p a rt o f d o rsal fin b ase. V e rtic a l
stria e on sc a le s tra v e rs in g w h o le
body cavity
scale or o verlapping across centre
o f scale.
S w im bladder w ith tw o
tubes passing back in the m uscles
on e ith e r sid e o f h a e m a l sp in es.
Resem bles three other In d o -P ac ific
scale
s p e c ie s o f I l is h a in th is sw im ­
b lad d er ch aracter, but th ese have
more gillrakers (L obfuscata) or scale striae not continuous, w ith a distinct gap across centre o f scale (I k am p en i.
L s tria tu la ). O th er Ilisha species have either a single long tube from the hind end o f the swim bladder (and m ore
scutes), or no posterior tubes (and a more slender body or a distinct bony lobe on the shaft o f the m axilla behind
the tip o f the pre-maxilla). Pellona species have a distinct toothed hypo-maxilla. See CLUP Ilish 3, Fishing Area
51, also 57, 71.
273
Geographical Distribution :
Indian
O cean
(M alabar coast to Calcutta), Java Sea (o ff Java), South
C hina Sea (Singapore, G u lf o f Thailand), E ast China Sea
(north to Taiwan Island).
H abitat and B iology : M arine, pelagic, coastal,
but entering estu arie s and presu m ab ly able to to lerate
low ered salinities. F eeds on p lankton (probably sm all
cru stacean s, etc.).
Size :
To 17 cm standard length.
Interest to Fisheries :
C o n trib u tes to
catches by artisanal fish eries. M ostly caught
nets and b o at seines o v er dep th s o f 10 to 15
coasts and in estu arie s, also by cast nets in
(southern India).
clup eo id
by g illm along
estu aries
L ocal N am es : Literature :
L striatu la).
R a m a iy a n & W hitehead (1975 - b rief notes on distribution and biology, but K arachi record is
Ilisha n o v a cu la (V a le n c ie n n e s
1847)
PRIST Ilish 10
Pellona novacula V alenciennes, 1847, H ist.nat.poiss.. 20:319 (Rangoon).
Synonym s : P ello n a sladeni D ay, 1869:623 (Irraw ady R iv er at M andalay); Ilish a sla d en i:Fow ler. 1941:657
(com piled); W hiteh ead , 1971:74 (type o f sla d e n i); Id e m . 19 7 3 b :2 11 (key o nly); W hitehead & Talw ar, 1976:155
(type o f sladeni, listed); Ilisha elongata:W hitehead. 1967:121 (type o f novacula);W ongratana. 1980:199, pis 161,
162 (re v is io n ).
FAO Names :
En - B urm ese R iv er ilisha.
D iagnostic Features : Body rather slender, its depth 22 to 27%
o f standard len g th , belly w ith 24 to 25 + 1 0 or 11, total 34 to 36
scutes. E ye large, low er ja w strongly pro jectin g ; low er g illrak ers 21
sw im bladder
to 23. D orsal fin origin behind m idpont o f body; anal fin w ith 43 to
45 finrays, its origin b elow dorsal fin base. S w im bladder w ith o u t a
body cavity
p o sterio r tube (or tu b es) p assing b ackw ard into the body m uscles.
no tube
R esem b les L sirish ai in th is sw im bladder c h aracter, but th a t species
has a distinct bony lobe on the shaft o f the m axilla behind the tip o f
the pre-m axilla, few er scutes (total 28 or 29), more gillrakers (22 to
26) and a deeper body (depth 32 to 37%).O ther Ilisha species have one or two
thin tubes passing backward from
the sw im blad der, also few er scutes (26 to 30) or a deeper body (27 to 39%).
P ello n a species have a distinct
toothed hypo-m axilla and are deep-bodied;
O pisthopterus species have the anal fin origin well in advance o f the
dorsal fin origin.
274
G eographical
D istrib u tio n
B urm a
rivers
(Irrawaddy at Rangoon and Mandalay; Sittang River).
H abitat and B iology
R iverine, reaching at
least 650 km up the Irraw addy, perhaps more. M ore
data needed.
Size :
To 32 cm standard length.
Interest to Fisheries : Perhaps o f local interest,
but no special fishery.
L o ca l N am es : L iteratu re :
See under Synonyms.
Ilisha obfuscata W ongratana,
PRIST
1983
Ilish
11
Ilisha obfuscata W ongratana, 1983, Jap .J.Ic h th y o l.. 2 9 (4 ):3 9 7 , fig. 14 (Bom bay, Pondicherry; on a syntype o f
Pellona filigera Valenciennes and a paralectotype o f Pellona m icropus Valenciennes, i.e. L m elastom a).
Synonym s : None.
FAO Names : En - H idden ilisha.
Ä SS""**'
ip
D iagnostic Features : Body m oderately deep, its depth 34 or
2 tubes
35% o f stan d ard len g th , b elly w ith 19 or 20 + 8, total 27 or 28
scutes. Eye large, low er ja w projecting; low er gillrakers 27 or 28
sw im bladder
(2 fishes only). D orsal fin origin at about m idpoint o f body; anal
fin with 39 to 42 finrays, its origin under or ju st behind final dorsal
body cavity
fin rays. V ertical striae on scales traversing w hole scale or
overlapping across centre
o f scale.
Sw im bladder w ith two very
short tubes passing back in the m uscles on either side o f haemal
spines (not reaching back beyond vertical from about 10th anal
fin ray base).
R esem bles three other Indo-P acific species in its
paired swim bladder tubes,
but these all have fewer gillrakers (25 or less;
L m elastom a. L striatula. L kam peni').
Other Ilisha species have
fewer gillrakers and either a single long tube from the hind end o f the
swimbladder and
more scutes), or no posterior tubes (and a more slender body or a distinct bony lobe on the shaft o f the m axilla
behind the tip o f the pre-maxilla). Pellona species have a distinct toothed hypo-maxilla.
275
Geographical D istribution : Indian
known from B om bay and Pondicherry).
O cean
(only
mmmmymm
s j î
- y
\
—
-----—
T ----------------
------------------ y
*
S
p
)
H abitat and B iology : M arine, pelagic,
M ore specimens and data needed.
Size : To 7.4
grows larger.
coastal.
>
cm standard length, but probably
Interest to Fisheries
\ y
/
V
/
: No data.
---- .
" S ’
’S
• '
»
L ocal N am es : -
r
L iterature
P robably m ixed w ith records
L filigera and other species o f Indian coasts.
of
^
1
u ■
»
e
*
Rem arks : The only two specim ens know n came
from the type series o f filig era and m ic ro p u s.
The
species is unique among Indo-Pacific Ilisha in its high
gillraker count and very short sw im bladder tubes.
Ilisha pristigastroide
PR IST Ilish 1
(B leeker, 1852)
F o rm e rly C L U P Ilis h 1
Pellona
pristigastroides
Bleeker,
1852,
V erh.batav.G enoot.K unst.W et., 24:20 (Jakarta).
Synonym s : P ellona am blyuropterus B leeker, 1852:21 (Jakarta; W eber & de B eaufort, 1913:90 (Java,
Sum atra, K alim an tan , S in g ap o re); Ilish a a m b ly u ro p te ra :Fowler. 1941:656 (c o m p ile d );
Ilish a p ristig a stro id e s W h ite h e a d e t a l., 1966:93, 94, pi. 11, fig. 3 B leek er fig. (types o f p ristig a stro id e s, am b lu ro p teru s ; W h ite h e a d ,
1973b:211 (key only); W ongratana, 1980:208, pi. 170 (revision).
FA O Names :
E n - Javan ilisha.
D iagnostic Features : Body m oderately slender, its depth 31
or 32% o f standard length, belly with 26 or 27 + 11 or 12, total 37
sw im bladder
to 39 scutes. Eye larg e, low er ja w p ro jectin g ; low er g illrak ers 17
(3 fishes). Dorsal fin origin behind m idpoint o f body; anal fin with
1 tube
45 or 46 finrays, its o rigin b elow fro n t p art o f dorsal fin base.
on right
body
cavity
Swimbladder with a single long tube passing back down right side o f
side
b ody ab o v e an al fin b ase.
R e se m b le s fo u r o th e r In d o -P a c ific
sp e c ie s o f Ilis h a in th is c h a ra c te r, b u t th e y h a v e fe w e r sc u te s
(L m egaloptera)
or
m ore
gillrakers
(L elongata, L filig era,
L m acrogaster). O ther Ilisha species have either tw in tubes from the hind end o f the sw im bladder or no posterior
tubes and more gillrakers and few er scutes). Pellona species have a distinct toothed hypo-m axilla. See CLUP
Ilish 1, Fishing Areas 57, 71.
276
G eographical
D istribution
: Java
Sea
(Jakarta).
Habitat and Biology :
M arine, pelagic, coastal,
apparently entering rivers and brackishw ater (W eber &
de B eaufort, 1913 - as P ellona pristigastroides and
R am blyuropterus).
Size :
To 30 cm standard length.
Interest to Fisheries : No data.
Local Names : IN D O N E S IA : Ik a n puput.
Literature :
See under Synonyms.
Remarks :
W ongratana (1980) was able to exa­
mine only three specimens, but the species seems to be
distinct.
Ilisha sirish ai Seshagiri R ao, 1975
PRIST Ilish 12
Ilisha sirishai Seshagiri Rao, 1975, H ydrobiologia, 47:463, figs 1, 2 (V izagapatnam ,
Synonyms:
F A O Names :
Ilisha
India).
sirishai - W ongratana, 1980:198, pis 159, 160 (revision).
E n - L obejaw ilish a .
D iagnostic Features : Body m oderately
deep, its depth 32 to 37% o f standard length, pre-m ax illa
belly with 18 to 21 + 8 or 9, total usually 28 or
29 scutes.
E ye la rg e , lo w e r ja w stro n g ly
projecting;
shaft o f m ax illa w ith a d istin ct
lobe or flange betw een hind tip o f pre-maxilla
sw im bladder
and blade o f m axilla; low er g illrak ers 22 to
26. D orsal fin orig in near or before m id p o in t
body cavity
o f body; anal fin w ith 39 to 43 finrays, its
origin a little behind dorsal fin base. Swim­
no tube
b la d d e r w ith o u t a p o s te rio r tu b e (o r tu b e s )
bonypassing b a c k w a rd in to th e b o d y m u s c le s .
R e se m b le s I. n o v a c u la in th is sw im b la d d e r
maxilla blade
character, but that species has no lobe on the
m axilla, m ore scutes (to tal 34 to 361, fe w e r
gillrakers 121 to 23) and is more slender (depth 22 to 27%). Other Ilisha species have one or two thin tubes passing
backward from the sw im bladder, also usually more scutes or more g illrak ers or are more slender, and none has a
lobe on the m axilla (L striatula and L melastom a are otherwise very sim ilar to L sirishai). Pellona species have a
distinct toothed hypo-m axilla in p la c e o f the lobe on the maxilla.
i
277
Geographical D istribution : Indian O cean (from
the “G u lf ’ to Bay o f B engal and A ndam an coast o f
Thailand), G ulf o f Thailand (Songkhla Lake).
H abitat and B iology : M arine, pelagic and in­
shore, presum ably able to tolerate some low ering o f
salinity. M ore data needed.
Size :
To 17.2 cm standard length.
Interest to F isheries : C ontributes to
catches o f clupeoids, but no special fishery.
L ocal N am es
Literature :
L indica).
artisanal
:D eh ad rai (1960 - sw im bladder, as
PRIST Ilish 13
Ilsha striatula W o n g ra tan a , 1983
Ilisha
T u tic o rin ).
striatula
W ongratana,
1983,
J a p .J .I c h t h y o l.,
29(4):396,
fig.
13
(K arachi,
Porto
N ovo,
M adras,
Synonyms : None.
FA O Names :
E n - B anded ilisha.
D iagnostic Features :
B ody m odestria e
rately deep, its depth 32 to 39% o f standard
d is c o n tin o u s
length, belly w ith usually 18 to 21 - 7 to 9,
total 26 to 29 scutes. Eye large, low er jaw
projecting! low er g illrak ers 21 to 24. D o r­
sal fin o rig in a little b e fo re m id p o in t o f
sw im bladder
bodyi
anal fin w ith 40 to 47 finrays, its
origin below final dorsal finray bases. V er­
bodv cavitv
tic a l stria e on sc a le s n o t c o n tin u o u s , b u t
w ith a d istin ct gap across centre o f scale.
2 tubes
A faint dark band along flanks. Swimbladder
w ith two tubes passing back in the muscles
on eith er side o f haem al spines. R esem bles
scale
three other In d o -P acific species o f Ilish a in
this sw im bladder ch ara cte r, but th ese have m ore g illr a k e rs (L o b fu s c a ta ),or scale striae that traverse
the scale
(I. m elastom a) or are more slender and have shorter pectoral fins (Lkam peni! pectoral fin 15to 17%
o f standard
length, cf. 18 to 21%). O ther Ilisha species have either a single long tube from the hind end o f the sw im bladder
(and more scutes), or no posterior tubes (and a more slender body or a distinct bony lobe on the shaft o f the
maxilla behind the tip o f the pre-maxilla). Pellona species have a distinct toothed hypo-maxilla.
click for n e x t p a g e
c lic k fo r p r e v io u s p a g e
278
G eographical D istribution :
Indian
O cean
(K a ra c h i to M a d ra s and p e rh a p s n o rth to C a lc u tta );
sp ecim en s from th e n o rth w e ste rn c o asts o f A u stra lia
may be this species.
H abitat and B iology : M arine, pelagic, coastal,
but probably also entering estuaries. M ore data needed.
Size :
To 18 cm standard length.
In terest to F ish eries :
L melastoma.
L ocal N am es
Literature :
P robably cau g h t w ith
:-
See under Synonyms.
Remarks : A lm ost certainly confused in the past
w ith the very sim ilar L melastom a in Indian waters.
Pellona V alenciennes,
P R IS T
1847
F o r m e r ly
P eli
C L U P Peli
P ellona V alenciennes, 1847, H ist.n at.P o iss., 20:300 (type: Pellona orbignyana
V alenciennes = Pristigaster
fla v ip in n is (V a le n c ie n n e s ). N e o s te u s N o rm a n , 1 9 2 3 , A n n .M a g .n a t.H is t., 9 11:17 (ty p e :
P e llo n a d itc h e la
V alenciennes).
D iagnostic F eatures
M oderate or large-sized
m arine, estuarine or freshw ater clupeoid fishes (to about
50 cm standard length), body m oderately deep or elon­
gate, com pressed, w ith a sharp keel o f s c u te s along belly.
Eye fairly large; low er ja w projecting, m outh directed
obliquely upw ard; upper ja w reaching at m ost to eye
centre; a small toothed hypo-m axilla betw een hind tip o f
pre-m axilla and blade o f m axilla (easily detected by
rubbing a finger-nail along the low er edge o f the jaw );
jaw teeth sm all or m inute, usually w ith a distinct gap at
centre o f upp er jaw .
G illrak ers fairly short and thick,
few , 20 to 31 (but anterior gillrakers regress in some
S o u th A m e ric a n P e llo n a o v e r 30 cm sta n d a rd le n g th ).
Dorsal fin at about m idpoint o f body; pelvic fins present,
s m a ll, w ith 6 or 7 finrays, a little or w ell in front o f
dorsal fin origin; anal fin m oderately long, w ith 34 to 43
finrays, its orig in under or ju s t behind dorsal fin base.
S c a le s m o d e ra te , a b o u t 35 to 50 in la te ra l s e rie s. N o
distinctive colour patterns. No tube or tubes at hind end
o f sw im bladder passing b ackw ard into body m uscles (cf.
species o f Ilisha).
P ello n a strongly resem bles Ilish a , but
th e la tte r la c k s a to o th e d h y p o -m a x illa ry b o n e in th e
upper jaw.
Biology, H abitat and D istribution : M arine, pela­
gic and inshore in Indo-Pacific, marine and freshw ater in
N ew W orld (A tlantic and its drainage; absent o ff Pacific
coasts).
Interest to Fisheries :
Species :
pre-m axilla
toothed
m axilla
m axilla
blade
no
postcoelom ic
tube
sw im bladder
body cavity
anus
C o n trib u te to artisan al fish eries.
W o n g ratan a (1980) re c o g n iz e d 2 Indo-Pacific species,
and Whitehead (1973a) 3 N ew World species:
Indo-W est Pacific
P. dayi W ongratana, 1983, India, Indonesia
R ditchela V alenciennes, 1847, E ast A frica to Papua N ew Guinea.
279
N ew W orld
R castelnaeana V alenciennes, 1847, O rinoco, A m azon, freshw ater
P. flavipinnis (V alenciennes, 1837) O rinoco to B uenos A ires, freshw ater
R harroweri (Fow ler, 1917), Panam a to Brazil, marine.
P ellon a
casteln aea n a
V alenciennes, 1847
PRIST Peli 3
Pellona castelnaeana V alenciennes, 1847, H ist.nat.P oiss., 20:306 (A m azon mouth).
Syn onym s :
(P ello n a p o ly p e is. MS nam e o f S tein d ach n er: ty p e N M V 1103 ex J a t u a r a n a ] ; ? P ello n a
altam azonica C ope, 1872:256 (A m byiacu R iver, E cuador: types lost fide B öhlke, 1984:62): FW N A , 1964:417, fig.
104 (Amazon betw een Para and M anàos: low er gillrakers only 12 or 13): Ilisha desuratus N akashim a, 1941:77, fig.
(near Iq u ito s, P eru v ian A m azo n ): P e llo n a c a s te ln a e a n a - W h iteh ead , 1967:108 (types o f ca ste ln a e a n a ): Id em ,
1970:25 (key, separatio n from P. flavipinnis on g illrak ers):
Idem , 1973a:71, fig . 21 (G uianas); W h i t e h e a d &
Bauchot, in press (types o f castelnaeana).
FAO N am es :
E n - A m azon pellona.
D iagnostic Features : Body moderately deep and compressed, its depth about 29 to 34% o f standard length,
belly w ith 23 or 24 + 8 to 11, total 33 or 34 scutes. L ow er ja w p ro jectin g : u p p er ja w w ith a to o th ed hypo
m a x illa ry b o n e b e tw e e n h in d tip o f p re -m a x illa and
low er bulge o f m axilla blade (easily felt w ith fin g er­
nail): lower gillrakers 12 to 14 (in fishes o f 20 to 50 cm
standard length). P elv ic fins p resen t, w ith a d istin ct
axillary scale: anal fin long, its origin under dorsal fin
base, w ith about 34 to 38 finrays. Scales fairly sm all,
a b o u t 70 in lateral series. C losely resem bles P. flav i­
pinnis. w hich has more low er gillrakers (23 to 31) and
m ore p o s t-p e lv ic sc u te s (13 or 14); R h a rro w e ri has
only 5 to 7 po st-p elv ic scutes and no pelvic a x illa ry
scale. Ilisha am azonica lacks a toothed hypo-m axilla.
O th e r sim ilar clupeo id s have a sh o rter anal fin (less
than 30 finrays).
Geographical Distribution :
A m az o n sy stem
(Iquitos on P eruvian A m azon and A m b y iaco s R iv er in
E c u a d o r to M anáos and p e rh a p s P ara), th e G u ia n a s
(L ake A m uku area w here th e A m azo n and E sseq u ib o
systems jo in in w et years).
H abitat and Biology : R iv erin e, apparen tly not
e n te rin g th e sea, a lth o u g h p re s u m a b ly to le ra tin g at
least som e salinity at th e A m azo n m outh. M ore data
needed.
Size :
To at least 47 cm standard length.
Interest to Fisheries : A lm o st certain ly caught
by artisanal fisherm en on the Amazon.
L ocal N am es : -
280
Literature :
See under Synonyms; otherwise not clearly distinguished from R flavipinnis.
Rem arks : M uch confused in the literature w ith the superficially sim ilar P. flavipinnis until separated by
gillraker and scute counts. The ranges o f the two species overlap in the A m azon drainage, at least in the middle
and low er parts, so that previous biological or fishery data may apply to either species.
P ello n a d ay i W o n g ratan a, 1983
PR IST Peli 4
Pellona dayi W ongratana, 1983, Ja p .J .Ic h th y o l., 29(41:395, fig. 12 (P orto N ovo, M adras).
Synonym s : None.
FAO N am es :
En - D ay's pellona.
stria e
discontinuous
D iagnostic Features : Body m oderately deep, com pressed, belly w ith 19 +
8 or 9, total 27 or 28 scu tes. Eye large, low er ja w projecting; upper ja w w ith a
toothed hypo-m axillary bone between hind tip o f pre-m axilla and low er bulge o f
m axilla (easily felt w ith fingernail); low er gillrakers 20 or 21. Dorsal fin origin
near m idpoint o f body; pelvic fins present; anal fin w ith 35 to 41 fin ray s. Scales
w ith upper and low er v ertical striae not m eetin g at centre o f scale.
C losely
resem bles the w idespread P. ditchela, w hich has 22 to 27 low er gillrakers and the
vertical scale striae overlapping at centre o f scale. Species o f Ilisha have no
hypo-m axilla. O ther p ristig aste rid s are m uch m ore slen d er and lack p elvic fins
('O p isth o p te ru s. R a c o n b a l.
scale
Geographical Distribution :
eastern coast o f southern India).
In d ia n ocean (on
Habitat and Biology :
M a rin e , in s h o re and p e r­
haps, like R ditchela, entering estuaries.
Size
To 13.5 cm standard length.
Interest to Fisheries :
catch records o f R ditchela).
N o d a ta (in c lu d e d in
L o ca l N am es
Literature
to P. ditchela.
P resu m ab ly in clu d ed in referen ces
R em arks : N ot easy to separate from R ditchela
on the scale striation character since scales easily lost.
Further specimens may blur the distinction on gillraker
numbers.
281
PRIST Peli 1
Pellona ditchela V alenciennes, 1847
F o rm e rly C L U P Peli 1 and
C LU P P e lló n 2 (erro r)
Pellona ditchela V alenciennes,
1847, H ist.nat.poiss., 20:314 (on D itchelee o f Russell, 1803).
Synonym s : Pellona hoevenii B leeker, 1862:21 (Jakarta); W eber & de B eaufort, 1913:86, fig. 29 (jaws)
(Indonesia); Pellona natalensis G ilchrist & Thom pson, 1908:202 (N atal coast); Pellona ditchela - Fow ler, 1941:648
(S ou th A frica to In d o n esia and the P h ilip p in es; large synon.); W h iteh ead et aL, 1966:91, pi. 11, fig. 2 (B leek er's
fig.) (types o f hoevenii); L osse, 1968:104 (East A frica;synon., descr.); W hitehead, 1973b:209, fig. 35 (key, synon.,
refs); W o n g ratan a, 1980:195, pis 155, 156 (rev isio n );C L O F F A , 1984:48 (A frican fresh w ater refs); SFSA , in press
(South Africa).
F A O Names
En - In d ian pellona.
Cm
jJ
/V .Y Y 'Y
striae
o verlapping
D iagnostic Features
Body m oderately deep, com pressed, belly w ith
usually 18 or 19 + 8 or 9, total 26 to 28 scu tes. Eye large, low er jaw projecting;
upper ja w w ith a to o th ed h y p o -m ax illary bone b etw een hind tip o f p re-m ax illa
and lower bulge o f m axilla blade (easily felt with fingernail); low er gillrakers 22
to 27. Dorsal fin origin near m idpoint o f body; pelvic fins present; anal fin with
34 to 42 finrays. Scales with upper and low er vertical striae slightly overlapping
each other at centre o f scales. Closely resembles R d a y i, which has only 20 or
21 low er gillrakers and the vertical scale striae not m eeting at centre o f scale.
Species o f Ilish a have no h ypo-m axilla.
O th er p ristig a sterid s are m uch m ore
slender and lack p elvic fins (O p isth o p te ru s, R a c o n d a ).
See C LU P P e lló n 2,
Fishing Area 51; also CLUP Peli 1, Fishing Areas 57, 71.
scale
Geographical Distribution : Indian O cean (w est­
ern c o a sts fro m th e G u lf o f O m an to D u rb a n , a lso
M adagascar; coasts o f India and A ndam an Sea, perhaps
o ff eastern Java, and south to W estern A ustralia), South
China Sea (the Philippines), Indonesia to northern A us­
tralia and Papua N ew Guinea.
Habitat and Biology :
M arin e, c o astal, but en ­
tering m angrove swamps and penetrating estuaries into
fresh w ater (e.g. in the P an g an i R iv er, T an zan ia), thus
apparently fully euryhaline.
Size :
To 16 cm standard length.
Interest to F ish eries :
N o specific fishery, but
co n trib u tes to artisan a l catches o f clupeoids.
Only
M alaysian catch reported (2 542 tons in 1983).
Local Names :
E A ST A FR IC A : C haa (g en era),
Simu (Shimoni), Simu koko (M alindi); SOUTH AFRICA:
Indiese pellona.
282
Literature :
Sadasivan (1965 - general biology);
P ello n a flavip in n is (V alen cien n es, 1837)
Losse (1968 - description, habitat in East Africa).
PR IST P eli 5
Pristigaster flavipinnis V alenciennes, 1837, in d'O rbigny, V oy.A m er.m erid., Poiss., A tlas:pl. 10, fig. 2 (figure
only based on Buenos A ires specim ens fide W hitehead, 1967:106); V alenciennes, 1847, ibid ., 5(2):8 (descr.).
Synonyms :
P ello n a o rb ig n y an a V alen cien n es, 1847:302 (Buenos A ires); (Pellona staudingeri, MS name o f
S tein d ach n er; ty p es N M V 1112, 1893 ex Iquitos] (P ello n a m a c ro le p sis, MS nam e S teindachner; type N M V 1101
ex T effé, middle Amazon]; Ilisha c a s t e l n a e a n a :F W N A , 1964:419 based on N orm an, 1923; m isidentified); Pellona
c a s te ln a e a n a :C e rv igón, 1966:130 (O rin o c o m o u th ;
lo w er g illrak ers 25, 2 7 , thus the p resen t species); P ello n a
flavipinnis - R inguelet, A ram buru & A ram buru, 1967:60, fig. le (A rgentina; descr. refs); W hitehead, 1967:106,
107, pi. 7a (original V alenciennes fig.) (types o f flavipinnis, orbignyana); Idem , 1970:26, fig. 2 (gillraker num bers)
(key , d istin c tio n from P. c a ste ln a e a n a ; ‘ty p e s ’ o f sta u d in g e ri, m a c ro le p is); Id e m , 1973a:67, fig. 19 (G uianas);
C ervigón, 1982:211 (O rinoco); W hitehead & B auchot, in press (types o f flav ip in n is, o rb ig n y an a).
FAO N am es
En - Y ello w fin riv e r pellona.
D iagn ostic F eatu res : Body m oderately deep and com pressed, its depth 30 to 37% o f standard length, belly
w ith 20 to 24 + 12 to 14, total 32 to 37 scutes. Low er iaw projecting; upper jaw w ith a toothed hypo-m axillary
bone betw een hind tip o f pre-m axilla and low er bulge ofi
m axilla blade (easily felt w ith fin g ern ail); low er g ill­
rakers 23 to 31 (in fish es o f 10 to 50 cm stan d ard
len g th , w ith the lo w er num bers in th e larg er fishes).
P elvic fins present, w ith a d istin ct ax illary scale; anal
fin long, its origin under dorsal fin base, w ith 38 to 46
finrays. Scales fairly sm all, about 60 in lateral series.
C lo se ly re s e m b le s R c a s te ln a e a n a , w h ic h has fe w e r
low er gillrakers (12 to 14) and few er p o st-p elv ic scutes
(8 to 11);
P. harrow eri has only 5 to 7 p o st-p elv ic
scutes and no pelvic ax illary scale. Ilish a am azonica
lacks a toothed hypo-m axilla. O ther sim ilar clupeoids
have a shorter anal fin (less than 30 finrays).
G eographical D istribution : A rgentina (Rio de la
Plata down to Buenos Aires, Rio P a ra n á ), U ruguay (Rio
U ruguay), Brazil (Rio Tapajoz, A m azon as far up as Rio
Jurua and dow n probably to m outh), the G uianas (S uri­
nam, G uyana), V enezuela and Colom bia (O rinoco from
Rio M anacacias at Puerto G aitan to O rinoco mouth).
Habitat and B iology :
R iv erin e, ap p aren tly not
entering the sea, alth o u g h p resu m ab ly to le ra tin g som e
salinity at river mouths.
Size :
To 50 cm standard length.
Interest to Fisheries :
A lm o s t c e r ta in ly
im portance to artisan a l fisherm en.
of
283
Local Names :
Literature :
ARGENTINA: Lacha, also Lacha pelada, Salaca.
See under Synonyms; otherwise not clearly distinguished from R
castelnaena.
Remarks : See under P. castelnaeana.
P R IST Peli 2
P e llo n a h a r r o w e r i (F ow ler, 1917)
F o rm e rly
Ilisha harroweri Fow ler, 1917, Proc.A cad.nat.Sei.Philad.,
C L U P Peli 2
69:128, fig. 1 (Colon, Panama).
S yn on ym s : Ilis h a n a ra n a n se ta e F o w le r, 1 9 1 1 :2 0 8 , fig. 1 (N e w p o rt, R ho de Is la n d ); N e o ste u s te rn e tz i
N orm an, 1923:595 (Rio de Janeiro); Ilisha argentata H ildebrand, 1923:190, pi. 9 (Fox B ay, Colon, Panam a);
Pristigaster vanderbilti Borodin, 1928:6, fig. 1 Panam a; abnorm al specim en lacking pelvic fins fide H ildebrand,
1964:426; N eosteus m ayrinki Pinto, 1972:2 (B acia do Parnaiba, B razil); Ilisha harrow eri :F W N A , 1964:423, fig.
106 (descr., synon.); Pellona harroweri C ervigón, 1966:128, fig. 54 (V enezuela); W hitehead, 1973a:61, fig. 16
(key, synon., descr., Guianas specimens); Figueiredo & M enezes, 1978:25, fig. 31 (Brazil, synopsis).
E n - A m erican coastal pellona.
F A O N am es:
V.
D iagnostic Features : Body fairly deep and com pressed, its depth about 35 to 42% o f standard length, belly
with 17 to 20 + 5 to 7, total 22 to 26 scutes. Eye larg e, low er ja w pro jectin g ; upper ja w w ith a toothed hypomaxillary bone betw een hind tip o f pre-m axilla and low er bulge o f maxilla blade (easily felt w ith fingernail); lower
gillrakers 23 to 25. Pelvic fins present, w ithout a distinct axillary scale; anal fin long, its origin under middle o f
dorsal fin base, w ith 36 to 42 fin ray s. Scales easily lost. R esem bles P. castelnaeana and R flavipinnis, w hich
have m ore po st-p elv ic scutes (8 to 14) and a distinct pelvic axillary sc a le . Ilisha amazonica has no hypo-maxilla.
Other similar clupeoids have a shorter anal fin (less than 30 finrays). See CLUP Peli 2, Fishing Area 31.
284
G eographical D istribution
W estern A tlantic
(Panam a to T rinidad, the G uianas, entire B razilian
c o a st so u th o f th e A m a z o n to R io G ra n d e do Sul at
about 30°S).
H abitat and B iology : M arine, inshore, along
beaches and down to at least 16 m, entering estuaries,
but perhaps not tolerating very low salinities.
Size :
To about 13 cm standard length.
Interest to Fisheries : E n ters a rtisan al catc h es,
but apparently not in large numbers.
L ocal Names :
VENEZUELA:
Literature : M atsuura (1973
and figured, Brazil).
Sardina.
postlarva described
Rem arks : Pellona harrow eri bears a
resem blance, not to the two other N ew W orld
Pellona, but to R ditchela o f the Indo-W est
differs in having fewer scutes (22 to 26; cf.
and few er pelvic finray s (i 5; cf. i 6).
rem arkable
species o f
Pacific. It
26 to 28)
P lio steo sto m a N orm an, 1923
Pliosteostom a
N orm an,
PR IST Plio
1923,
A nn.M ag.nat.H ist., (9)11:21 (type:
Pristigaster
lutipinnis
Jordan & Gilbert).
D iagnostic F eatures
M oderate-sized m arine clupeoid fishes (to
about 15 cm standard length), body elongate and com pressed, w ith a sharp
keel o f scutes along belly. L ow er ja w strongly projecting; hypo-m axilla
present. Pelvic fins absent; anal fin long, w ith around 50 finrays. The
h y p o -m a x illa d is tin g u is h e s it fro m all o th e r p ris tig a s te rid g e n e ra o f
Pacific coasts (see species for further distinctions).
B iology, H a b ita t and D istrib u tio n : See sp ecies.
^ re m a x ' ^ a
In terest to F ish eries : See species.
Species ! A single species recognized:
P. lutipinnis (Jordan & Gilbert, 1881), eastern central P acific.
to o th ed
h y p o -m ax illa
m a x illa blade
P liosteostom a lu tip in n is (Jordan & Gilbert, 1881)
PR IST Plio
1
Pristigaster lutipinnis Jordan & Gilbert, 1881, P ro c .U .S .n a tn .M u s., 4:340 (M azatlán , M exico).
Synonyms : O p isth o p teru s lu tip in n is:Jordan & E v e rm a n n , 1896:437 (M azatlán ); P lio steo sto m a lu tip in n is N o r m a n , 1923:21 (M azatlán, M e x ic o ); P e te rso n ,
1956:180 (B a rra n c a B a y , C o sta R ic a ); A n o n ., 1976:67, 68
(M exico; key, listed).
F A O Names :
En - Y ello w fin herring.
285
Diagnostic Features ! Body elongate, com pressed, its depth about 25% o f standard length, belly w ith about
27 or 28 scu tes.
L o w e r ja w strongly p ro je c tin g , m outh poin tin g o b liq u ely upw ard; a d istin c t to o th ed hypo
m a x illa ry b o n e b e tw e e n h in d tip o f p re -m a x illa and
low er bulge o f maxilla blade (sm all, but can be felt with
fin g ern ail);
upper ja w not reaching back beyond eye
c e n tre . L ow er g illrak ers around 18 to 20. D o rsal fin
behind m idpoint o f body; pelvic fins absent; anal fin
long, w ith 49 to 51 fin ray s, its o rigin a little before
dorsal fin origin. A silver band along flank. No other
P a c ific c o a st p ris tig a s te rid h as a h y p o -m a x illa ;
in
addition. Ilisha furthii has pelvic fins, species o f Odon­
tognathus have a long upper ja w (reaching back to gili
o p en in g ), and N eo o p isth o p te ru s tro p icu s has a terminal
mouth. R esem bles very closely the P acific species o f
O p isth o p te ru s, w hich have th e anal fin very m uch in
advance o f dorsal fin origin and more anal finrays (55 to
62).
G e o g r a p h i c a l Distribution:
E a s te r n c e n tr a l
Pacific (M azatlán , M exico south to Colombia).
H ab itat and B iology :
data needed.
Size :
M arin e,
coastal.
More
To 14.3 cm standard length.
Interest to F ish eries : Probably little.
Local Names :
Literature :
See under Synonyms.
c lic k for n e x t p a g e
c lic k for p r e v io u s p a g e
286
C hirocentrodon
PR IST Chiro
G ü n th er, 1868
F o rm erly C LU P C hiro
C h iro cen tro d o n G u n th er, 1868, C a t.F ish .B rit.M u s., 7:463 (type: C hirocentrodon taeniatus G ünther = Pellona
b l e e k e r i a n a Poey).
M edipellona Jo rd a n & Seale, 1 9 2 6 , B u ll.M us.co m p.Z o o l.H a rv ., 67(111:417 (ty p e :
P ellona
b leekeriana Poey).
D iagnostic F eatures
Small m arine clupeoid fishes (to about 9 cm
standard length), body elongate and som ehwat com pressed, w ith a sharp keel
o f scutes along belly.
M outh term in al:
no hypo-m ax illa: hind tip o f p re ­
m axilla m eets blade o f m axilla: strong conical te e th in ja w s, th o se at front
canine-like.
P e lv ic fin s p re s e n t:
anal fin lo ng, w ith around 40 finrays.
D istin g u ish ed from all o th er p ristig aste rid s and clu p eid s by the presence o f
canine teeth (but otherwise very closely resembles N eoopisthopterus).
B iolog y,
H ab ita t and
D istrib u tio n
See species.
Interest to F ish eries : See species.
S p ecies :
A s in g le s p e c ie s re c o g n iz ed , the scute count
T om m azi (1964) for his C cladileokae probably being an error:
given
by
C. bleekerianus (Poey, 1867), Atlantic coasts o f Central and South America.
R em arks :
frontal v iew o f m outh
Closely resem bles N eoopisthopterus (see under that genus).
C hirocentrodon bleekerianus (P oey, 1867)
PR IST C hiro 1
F o rm erly C LU P C hiro
Pellona bleekeriana Poey, 1867, R e p e rt.F is ic o -n a t.C u b a , 2:242
1
(M atanzas, Cuba).
Synonym s : C h iro centro d o n taen iatu s G unther, 1868:463 (Jam aica): Ilish a carib b ea M eek & H ild eb ran d,
1923:191, pi. 10, fig. 1 (P a n a m a ): C h iro c e n tro d o n c la d ile o k a e T o m m a z i, 1964:30 ( S a n to s , B razil: key, synon.,
descr.): C ervigón, 1966:127 (Nueva E sp arta and outsid e O rinoco d elta , V en ezu ela): W h iteh ead , 1973a:80, figs 2629 (jaw s) (T rinidad, O rinoco m outh, G uyana, S urinam ): F ig u eired o & M enezes, 1978:26, fig. 32 (B razil): U yeno,
M atsuura & Fujii, 1983:90, fig. (colour photo) (Surinam ).
F A O Names :
En - D o g to o th herring.
D iagn ostic F eatu res :
B ody elo n g ate, m oderately com pressed, its d epth about 20 to 27% o f standard
length, belly w ith 16 or 17 (rarely 19) + 9 to 11, total 26 to 28 (rarely 30) scutes. M outh term inal, lower ja w not
strongly projecting: strong con ical te eth in both ja w s, continued as large and small teeth along maxilla blade,
large and canine-like at front o f both jaw s: lower gillrakers 14 to 17. Dorsal fin origin behind m idpoint o f body:
pelvic fins present: anal fin m oderately long, w ith 38 to 44 fin ray s, its origin below or in front o f dorsal fin origin.
C losely resem bles N eo o p isth o p teru s cubanus in general appearance and general shape o f head and jaw s, but that
species lacks canine-like teeth and has no pelvic fins. No other sim ilar species has canine-like teeth. See CLUP
Chiro 1, Fishing Area 31.
287
G eographical D istribution
A tlantic coasts o f
C e n tra l and S o u th A m e ric a (C u b a , H a iti, Ja m a ic a ,
Puerto Rico, Trinidad; also from Panam a and coasts o f
V enezuela, including o ff Orinoco m outh, Guyana, Suri­
nam south to U batuba, near Santos, Brazil).
Habitat and B iology : M arine and coastal, appa­
rently down to about 60 m, but also occurring in lagoons
and o ff riv er m ouths (e.g. the O rin o co ), p erh ap s en ­
terin g w ater o f low ered salinity. P ro b ab ly spaw ns in
the sea.
Size :
To 9 cm standard length.
In terest to F ish eries : None.
L ocal N am es : L iterature : H ildebrand (i.e. FW N A, 1964 - note
on spawning);
C e rv ig d n (1 9 6 6 - h ab itat, fish eries),
M atsuura (1973 - juveniles described and illustrated).
120°
N eoopisthopterus
100°
80°
60°
PRIST Neop
Hildebrand, 1948
N eoopisthopterus
Hildebrand).
H ildebrand,
1948,
Smithson, mi sc. C olin s,
110(9):6 (ty p e :
D iagnostic Features : Small m arine clupeoid fishes (to about 9 cm
standard len g th ), body elo n g ate and som ew hat co m p ressed , w ith a sharp
keel o f scu tes along belly. M outh term in al; no hyp o -m ax illa; hind tip o f
pre-m axilla m eets blade o f m axilla; fine or m inute te eth in jaw s. P elvic
fins absent;
anal fin lo n g , w ith a b o u t 40 to 50 finrays.
M ost closely
resem bles C h iro cen tro d o n in all resp ects, but th a t has c a n in e-lik e teeth
and pelvic fins present. All other similar pristigasterids either have pelvic
fins or possess a toothed hypo-m axillary bone betw een hind tip o f pre­
maxilla and lower bulge o f maxilla blade (P lio ste o sto m a ).
B iology, H ab ita t and
D istrib u tio n
M arine
and
co astal,
approaching river m ouths, thus perhaps tolerating low ered salinities.
A tlantic and Pacific sides o f Central and South America.
Interest to F ish eries : L ittle o r none.
Species :
A u th o rs h a v e
gillraker and anal finrays counts:
40°
O dontognathus
tro p icu s
p re -m a x illa
no gap
m ax illa blade
re co g n ized an A tlan tic and a P acific species, d istin g u ish ed alm ost solely on
N. cubanus H ildebrand, 1948, w estern central A tlantic
N . tro p ic u s (H ildebrand, 1946), eastern central P acific.
Remarks : N eoopisthopterus hardly differs from Chirocentrodon in all but the absence o f canine-like teeth
and pelvic fins, but these characters occur consistently in all adults o f the Pacific £L tro p ic u s and p resu m ab ly do
so also in the A tlantic 15L cubanus (o v erlap in g the northern range o f C h iro ce n tro d o n ), w h ich seem s to be know n
only from small specim ens.
288
PRIST N eop 1
N eoopisthopterus cubanus Hildebrand, 1948
N eo o p isth o p teru s cubanus Hildebrand, 1948, Sm ithson, mi se. C o lin s, 110(9): V, figs 3-4 (Havana, Cuba).
Synon ym s :
N eo o p isth o p te ru s cubanus - FWNA, 1964:436, fig. 112 (Havana, Cuba).
F A O N am es:
En - Cuban longfin herring.
'< (•
D iagnostic Features : B ody elongate, m oderately com pressed, its depth about 17 to 22% o f standard
length, belly with 23 to 28 s c u te s . M outh term inal, low er jaw not strongly projecting; hind tip o f m axilla meets
low er bulge o f m axilla blade and is o v erlapped by it;
teeth very sm all or m inute; lo w er g illrak ers 17 or 18.
D orsal fin w ell behind m id p o in t o f body; pelvic fins
absent; anal fin long, w ith 39 to 43 fin ray s, its origin
before dorsal fin origin. The P acific FT tropicus has
m ore gillrakers (18 to 21) and more anal finrays (43 to
48). C losely resem b les C h iro cen tro d o n b leek e rian u s in
g e n e ra l fo rm and sh a p e o f h e a d an d ja w s , b u t th a t
species has canine-lik e te e th and pelvic fins.
Sim ilar
p ristig asterid s have the low er ja w strongly p rojecting;
similar clupeids have less than 30 anal finrays.
G eographical D istribution :
W e ste rn
A tlantic (Cuba; probably more widespread).
Habitat and Biology :
data needed.
M a rin e ,
c o a sta l.
c e n tra l
M o re
Size : U nknow n, but probably to about the same
size as FT tropicus, i.e. 9 cm standard length.
Interest to F ish eries : Probably none.
L o ca l N am es : L itera tu re : Remarks :
A p p ears to be uncom m on;
m ore
specimens may blur its distinction from FT tropicus.
N eoop isthop terus tr o p ic u s (H ild eb ran d , 1946)
PRIST N eop 2
O d o n to g n ath u s tro p ic u s H ildebrand, 1946, B u ll.U .S .n atn .M u s., (189):94, fig. 19 (Puerto Pizarro, Peru, also
P anam a B ay).
Synonyms:
N e o o p is th o p te ru s tro p ic u s - FW N A , 1964:436 (com pared w ith K
(C hira Island flats and Barranca Bay, G ulf o f N ico y a, C osta Rica).
F A O Names :
E n - T ropical longfin h e rrin g .
cu b an a); Peterson, 1956:184
289
D iagnostic Features : B ody elongate, m oderately com pressed, its depth about 25 to 27% o f standard
length, belly with around 26 scu tes. M outh term inal, low er ja w not strongly projecting; hind tip o f m axilla meets
low er bulge o f m axilla blade and is overlapped by it;
teeth very small or m inute; low er gillrakers 18 to 21.
D orsal fin well behind m idpoint o f body; pelvic fins
absent; anal fin long, w ith 43 to 48 finrays, its origin
before dorsal fin origin. A silver band along flank. The
A tlan tic 15k cubanus has few er g illrak ers (17 or 18) and
fe w e r a n al finrays (39 to 43). Sim ilar pristigasterids
have a toothed hypo-m axillary bone betw een hind tip o f
pre-m axilla and lower bulge o f m axilla (Pliosteostom a
lutipinnis) or low er ja w prom inent and m outh directed
obliquely upward (O dontognathus, O pisthopterus). Sim i­
lar clupeids have less than 30 anal finrays.
G eograp hical
D istrib u tion
E astern
central
Pacific (R io M ayo, G u lf o f C alifornia, C osta R ica,
Panama Bay and south to G ulf o f Guayaquil, Peru).
H a b ita t
and
B io lo g y
M a rin e ,
c o a stal,
occurring o ff river mouths (e.g. Rio M ayo) thus perhaps
tolerating low ered salinities. Feeds on planktonic crus­
taceans. A n extended spaw ning period o ff Costa R ica
(Peterson, 1956).
Size :
To 9 cm standard length.
Interest to F ish eries : Probably little or none.
Local Names :
Literature :
and m aturity).
P eterson (1956
- b rief notes on food
290
O pisth op terus Gili, 1861
PRIST
Form erly
O p is th o p te ru s G ili, 1 8 6 1 , P ro c .A c a d .n a t.S c i.P h ila d .:38 (type:
gaster tard o o re C uvier).
P ristigaster tartoor
D iagnostic F eatures
M oderate-sized m arine or estuarine
clupeoid fishes (to about 20 cm standard length), body rather elongate
and strongly co m pressed , belly w ith a sharp keel o f scu tes. E ye
large; low er ja w p ro jectin g , m outh d irected o b liq u ely upw ard; upper
ja w short, not reach in g back beyond ab o u t eye cen tre; no to o th ed
hypo-m axilla betw een hind tip o f p re-m ax illa and blade o f m axilla;
ja w teeth sm all, usu ally w ith a d istin c t gap at centre o f upper jaw ;
lower gillrakers 17 to 28. Dorsal fin small, its origin much behind
m idpoint o f body; pelvic fins ab sen t; anal fin long, m ore th an 50
finrays, its o rigin w ell before th a t o f dorsal fin. Scales easily lost,
about 45 to 60 in lateral series. The long anal fin and elongate body
easily d istin g u ish es this genus from o th er p ristig a ste rid s ex cep t the
In d o -P acific Raconda (w hich lacks a dorsal fin) and the N ew W orld
O d o n to g n ath u s
(m axilla long, to or beyond gili o pening), P lio ste o ­
stoma (toothed hypo-m axilla present) and N eoopisthopterus (less than
50 anal finrays, tip o f pre-m axilla reaches to blade o f m axilla w ithout
a ligam entous gap).
CLU P
V alenciennes
Opis
Opis
Pri sti -
p re-m ax illa
no
h y p o -m a x illa
m ax illa blade
Biology, H abitat and D istribution : C oastal, m arine and estuarine fishes, probably schooling, able to
tolerate at least brackish conditions. Found in tropical w aters o f the Indo-W est Pacific and in the eastern central
Pacific (but not along the A tlantic coasts o f the Americas).
Interest to Fisheries :
A small contribution to clupeoid catches, but no special fisheries.
Species : The 2 Indo-Pacific species were revised by W ongratana
study o f the 4 New World species:
(1980), but there has been no com parable
Eastern Central Pacific
O.
O.
O.
O.
dovii (Günther, 1868), California to Peru
effulgens (Regan, 1903), Ecuador
equitorialis H ildebrand, 1946, Honduras to Peru
macrops (Günther, 1866), Panama.
In d o -P a cific
O. tardoore (Cuvier, 1829), northern Indian Ocean to Indonesia
O. valenciennesi Bleeker, 1872, China to Indonesia.
Rem arks : The separation o f Indo-Pacific from eastern Pacific species w ithout any interm ediate A tlantic
species is a unique distributional pattern am ongst clupeoids. The two Indo-Pacific species have been well studied,
but the four Pacific species have not been revised since N orm an (1923); they have been separated on gillraker
counts and positions o f dorsal and anal fin origins, but much more material should be studied before O. effulgens
and O. macrops can be considered distinct from the apparently w ide-ranging O. dovii and (A e q u i t o r i a l i s .
PRIST Opis 3
O pisthopterus dov ii (G unther, 1868)
Pristigaster dovii G ünther,
argenteus G ü n th er, 1866).
1868,
C at.F ish.B rit.M us.,7:4611
(Panam a;
replacem ent
name
for
Pristigaster
Synonyms :
P r is tig a s te r a rg e n te u s G ü n th er, 1866:603 (P a n a m a ; preo ccu p ied by P ristig a ste r argenteus
S chinz, 1822 = P ristig a ste r cay an a);
M eek & H ild e b ran d 1 9 23 :1 9 2 , pi. 15, fig. 1 (la b e lle d O d o n to g n ath u s
pan am en sis) (C ham e P o in t, B a lb o a tid e p o o ls and P a n a m a M a rk e t); P e te rso n , 1956:181 (G u lf o f N ic o y a , C o sta
R ica) ; C hirichigno, 1962:4, fig. 2 (p hoto) (P uerto P izarro ,P ta M a i P elo , P eru); Cobo & M assay, 1969:7 (E cuador,
listed).
F A O Names :
En - D o v e’s longfin herring.
291
D iagnostic Features : Body elongate and strongly com pressed, its depth about 30% o f standard length,
belly w ith about 29 scu tes. L o w er ja w p ro jectin g , m outh p ointing o b liquely upw ard; no hy p o -m ax illa; low er
g illrak ers 15 to 19. P ecto ral fin a little lo n g er than
head; dorsal fin origin nearer to caudal fin base than to
vertical from pectoral fin base by 114 eye d iam eters (or
behind m idpoint o f body by 2 and 1/4 eye d iam eters);
anal fin long, w ith 55 to 62 fin ray s, its o rig in m uch
nearer to snout tip than to caudal fin base. C losely
resem bles O p isth o p teru s e q u ito ria lis, w hich has m ore
g illrak ers (21 to 25) and a s ilv e r b an d on fla n k , and
O. effulgens, w hich has the anal fin origin equidistant
betw een front m argin o f eye and caudal fin base (also,
only 15 g illrak ers); O. m acrops has the dorsal fin origin
eq u id istan t betw een caudal fin base and p ecto ral fin
base. Species o f O dontognathus have a long upper jaw
(to g ili c o v e r;
cf. to b e lo w ey e ).
P lio steo sto m a
lu tip in n is has a d istin ct to o th ed h y p o -m ax illa betw een
the hind tip o f the pre-m axilla and the low er bulge o f
the m axilla blade. N eo o p isth o p teru s tro p icu s has 50 or
fe w e r a n a l finrays. O th e r c lu p e o id s are m o re d e e p ­
bodied or the m outh is term inal or inferior.
G eographical D istribution
Pacific coasts o f
central and South A m erica (R io M uerto in G u lf o f
C a lifo rn ia so u th w a rd to P a n a m a B ay ;
a lso G u lf o f
G u a y a q u il, P e ru ).
H abitat and B iology : M arine, coastal, perhaps
entering w ater o f low ered salinity. M ore data needed.
Size : To 19.6 cm standard length.
Interest to Fisheries :
fisheries.
Local Nam es
P ro b ab ly enters artisan al
: ECUADOR:
O pisthopterus effulgens
Chaparra.
PRIST Opis 4
(R egan, 1903)
P ristig aster (O p isth o p te ru s) e ffu lg e n s R e g a n , 1903, A n n .M ag , n at.H i s t., (7)12:621 (Rio V aqueira, northw est
E c u a d o r).
Synonym s : O pisthopterus effulgens
FAO N am es:
- Norman, 1923:13 (the type only).
En - V aqueira longfin herring.
292
D iagnostic Features :
B ody e lo n g ate and stro n g ly co m p ressed , its d epth ab o u t 26 to 30% o f standard
length, belly w ith about 29 scutes. Low er ja w projecting, m outh pointing obliquely upw ard; no hypo-m axilla;
low er gillrakers about 15. Pectoral fin a little longer than head; dorsal fin origin nearer to caudal fin base than
to vertical from pectoral fin base by 1 eye diam ter (or behind m idpoint o f body by 2 and 3/4 eye diam eters); anal
fin long, w ith 65 fin ray s, its origin about equidistant betw een caudal fin base and front m argin o f eye. Other
Pacific species o f O pisthopterus have the anal fin origin
nearer to the snout than to the caudal fin base, also
m ore lo w er g illra k e rs (17 to 25). Species o f O d o n to ­
gnathus have a long upper ja w (to gili cover; cf. to
below eye).
Pliosteostom a lutipinnis has a distinct
toothed hypo-m axillary bone betw een the hind tip o f
the pre-m axilla and the low er bulge o f the maxilla.
N eoopisthopterus tropicus has 50 or few er anal fin ray s.
O ther clupeoids are more deep-bodied or the m outh is
term inal or inferior.
G eographical D istribution
P acific coasts (or
rivers) o f Central A m erica (know n only from the Rio
Vaqueira, northwest Ecuador).
H abitat and B iology
P erhaps riverine, or
merely entering rivers or estuaries from the sea. More
specimens and data needed.
Size :
To 20.5 cm standard length.
Interest to
fisheries.
Fisheries
: Probably
enters
artisanal
Local Names :
Literature :
Rem arks : The low gillraker count and rather
posterior anal fin origin seem to be distinctive, but
more specimens needed.
Opisthopterus
equitorialis
O pisthopterus
G u ay aq u il, Peru).
Hildebrand,
equitorialis
1946
H ildebrand,
Synonym s :
O pisthopterus equitorialis
1969:7 (Ecuador, listed).
F A O Names :
PR IST O pis 5
1946,
B ull.U .S .natn.M us., (189):93, fig. 18 (Puerto Pizarro, G u lf o f
- Peterson,
En- E quitorial longfin h errin g .
1956:182 (G ulf o f N icoya, Costa Rica); Cobo & M assay,
293
D iagnostic F eatures :
B ody elo n g ate and strongly co m p ressed , its d ep th ab o u t 28 to 30% o f stan d ard
length, belly w ith 26 to 28 scu tes. L o w er ja w p ro jectin g , m outh p ointing o b liquely upw ard; no h y p o-m axilla;
low er gillrakers 21 to 25; dorsal fin origin nearer to
caudal fin base than to vertical from pectoral fin base
by about 2 eye diam eters (or behind m idpoint o f body by
about 3 eye diam eters); anal fin long, w ith 56 to 62
fin ray s, its o rig in m u ch n e a re r to sn o u t tip th a n to
caudal fin base. O ther Pacific species o f O pisthopterus
have 20 or few er low er gillrakers. O dontognathus
panamensis has a long upper ja w (to gili cover; cf. to
below eye). P lio s te o s to m a lu tip in n is has a d is tin c t
too thed h ypo-m axillary bone b etw een the hind tip o f
th e p re -m a x illa and th e lo w e r b u lg e o f th e m a x illa
blade. N e o o p is th o p te ru s tro p ic u s has 50 or fewer anal
finrays. O ther clu p eo id s are m ore deep -b o d ied or the
m outh is term inal or inferior.
G eographical D istribution
Pacific coasts o f
Central and South A m erica (G u lf o f Fonseca, Flonduras
to G ulf o f Guayaquil, Peru).
Habitat and Biology :
M arin e, c o astal, perhaps
entering w ater o f low ered salinity. M ore data needed.
Size :
To 14.6 cm standard length.
Interest to
fisheries.
Fisheries
: Probably
enters
artisanal
Local Names :
Opisthopterus
m a cro p s (G ü n th er, 1866)
PR IST
O pis 6
P ristig aster m icrops G ü n th er, 1866, P ro c .z o o l.S oc.L o n d .:603 (P anam a).
S yn on ym s : O p isth o p te ru s m acro p s - N o rm an , 1923:11 (P an am a); M eek & H ild e b ra n d , 1923:14 (P an am a,
compiled); H ilbebrand, 1946:94 (com pared with O. equitorialis, m eristics, etc.).
F A O Names :
E n - B igeyed longfin h e r r in g .
294
D iagnostic Features :
B ody m oderately deep and strongly co m p ressed , its d epth a b o u t 35% o f standard
length, belly w ith about 28 scu tes. L o w er ja w p ro jectin g , m outh p ointing o b liquely upw ard; no h y p o-m axilla;
low er gillrakers 19 or 20. Pectoral fin a little longer
than head; dorsal fin origin about equidistant betw een
caudal fin base and vertical from pectoral fin base (or 1
and 1/4 eye diam eters behind m idpoint o f body); anal
fin long, w ith about 60 finrays, its origin nearer to
snout tip than to caudal fin base. O ther Pacific species
o f O pisthopterus have the dorsal fin. base nearer to the
caudal fin base than to the pectoral fin base, also more
gillrakers in O. equitorialis (21 to 25). P liosteostom a
lutipinnis has a
distinct toothed hypo-m axilla betw een
the hind tip o f the pre-m axilla and the low er bulge o f
the m axilla blade. N eoopisthopterus tropicus has 50 or
fe w e r an al finrays.
O th er clu p eo id s are m ore d e e p ­
bodied or the m outh is term inal or inferior.
G eographical D istribution
Central A m erica (Panam a Bay).
d ata
H abitat
needed.
Size :
and
B iology
Pacific
: M arine,
coasts
coastal.
of
M ore
To 17 cm standard length.
Interest to Fisheries :
fisheries.
P ro b ab ly enters artisanal
L ocal N am es :
L iteratu re
See under Synonyms.
R em arks : A pparently distinct from the other
three P acific species because o f the rath er advanced
dorsal fin, but more specimens needed.
O p isth o p teru s ta rd o ore (C uvier, 1829)
P R IST O pis 1
F orm erly C LU P O pis 1
P ris tig a s te r ta rd o o re C u v ie r,
p a tn a m , India).
1829, R égne anim al, 2nd ed .,2:381 (o n T a rto o r e o f R u s s e ll, 1 8 0 3 , V z a g a -
Synonym s : P ristig a ste r e lo n g a ta , S w ain so n , 1838:278 (on T arto o re); P ristig a ste r in d icu s S w ain son,
183 9:294 (on T a rto o re ); P ris tig a s te r ta rto o r V a le n c ie n n e s, 1 8 4 7 :3 2 8 (P o n d ic h e rry , M a la b a r); O p isth o p te ru s
m acrognathus B leek er, 1866:25 (Jak arta); O p isth o p teru s tartu s Z ugm ayer, 1913:9 (O m an); O p isth o p teru s tardoore
295
- F ow ler, 1941:663 (C a lic u t, In d ia and P a d a n g , S u m a tra ;
Jav a, but Flong K ong sp ecim en s w ere perhaps
O. v a le n c ie n n e si); W h iteh ead et ah, 1966:104 pi. 13, fig. 3 (B leek er's fig.) (key, types o f m a c ro g n ath u s); W h iteh ead,
1973b:215, fig. 39 (key, synon., refs); W ongratana, 1980:216, pis 179, 180 (revision).
F A O Names :
E n - T a rd o o re .
D iagnostic Features : Body elongate and strongly com pressed, its
depth 27 to 33% o f standard length, belly concave in front, w ith 29 to 35
scutes.
M outh po in tin g o b liq u ely upw ard;
low er g illrak ers 22 to 28,
increasing w ith size o f fish. Pectoral fin w ith 12 to 14 finrays, its length
21 to 26% o f sta n d a rd le n g th (u s u a lly a b o u t e q u a l to h ead le n g th or
greater); dorsal fin small, well behind m idpoint o f body; pelvic fins absent;
anal fin long, with 51 to 63 fin ray s, its origin well before dorsal fin origin.
Scales in lateral series 46 to 51. Closely resembles ÇT valenciennesi, which
is more slender (depth 24 to 29% o f standard length), and has a shorter
pectoral fin w ith more fin ray s (length 14 to 17% o f standard length, thus
sh o rter than head; 15 to 17 fin ray s). R aco n d a ru sselian a is su p erficially
similar, but lacks a dorsal fin and has more anal finrays (81 to 93 finrays).
O ther p ristig asterid s have p elvic fins and a sh o rter anal fin. See C LU P
Opis 1, Fishing Area 51, also Fishing Areas 57, 71.
pectoral fin equal or
longer than head
G eographical D istribution : Indian O cean (from
the G u lf o f O m an to at least M adras, perhaps to the
north and along the coasts o f B urm a, certainly at w
Penang), S in g a p o r e a n d J a v a S e a ( S u m a tr a ) , J a v a ,
K alim antan).
20°
H abitat and B iology : M arine, close to shore,
also entering estuaries (e.g. A luhaluh on B arito River,
K alim antan). Feeds on m ysids, P seudodiaptom us and
copepod eggs, also prawns and other small crustaceans,
bivalve eggs and larvae, am phipods and small fishes.
S p aw n s in la te F e b ru a ry o r e a rly M arc h to Ju ly or
A ugust (around K arw ar, India).
Size :
To 18 cm standard length.
^
20
°
40„
Interest to Fisheries : No special fishery, but is
caught w ith other clupeoids in shore seines (e.g. at
K arw ar from M ay to Septem ber); also caught w ith lift
nets and trawls in shallow waters.
Local Names :
INDIA: T ardoor, T artoor; IN D O N E SIA : K e n tu t (Jak arta).
Literature : John (1951 - ju veniles); R adhakishnan (1961, 1967, 1968a, b - general biology, m aturation and
spawning, racial variation); B asheerudin & N ayar (1962 - juveniles); Bensam (1968 - embryos and larvae).
296
O pisthopterus valen cien n esi
Bleeker. 1872
P R IS T O p is 2
O p isth o p teru s v alen cien n esi B leek er, 1 8 7 2 , A tia s ic h th y o l.In d .N e e rla n d ., 6:124 (replacem ent name for
O pisthopterus tartoor o f previous page, preoccupied by P ristigaster tartoor V alenciennes = O pisthopterus
tardoore, the other species in the genus: Jakarta, Singapore).
Synonyms : O p isth o p teru s ta rto o r B leek er, 1872:124 (seeabove) ; O pisth o p te r u s ta r d o o re :C hu, T c h an g &
C h en , 1 9 6 3 :1 0 6 , fig 80 (C h in a ): O p is th p te ru s v a le n c ie n n e s i - F o w le r, 1 9 4 1 :6 6 4 (c o m p ile d : k e y , sy n o n .):
W hitehead et a h , 1966:106,
pi. 14, fig. l(B leek er’s fig.) (key, types o f Q. valenciennesi): W ongratana, 1980:215,
pis 177, 178 (revision).
F A O Names :
E n - S lender ta rd o o r.
D iagnostic Features : Body very elongate and strongly com pressed,
its depth 24 to 29% o f standard length, belly concave in front, w ith 30 to
33 scutes. M outh pointing obliquely upw ard: low er gillrakers 23 to 25.
Pectoral fin w ith 15 to 17 fin ray s, its length 14 to 17% o f standard length
(thus distinctly shorter than head): dorsal fin small, well behind m idpoint
o f body: pelvic fins absent: anal fin long, with 54 to 65 fin ray s, its origin
w ell before dorsal fin origin. Scales in lateral series 51 to 56. C losely
resem bles CF tardoo re, w hich has a deeper bod
y (27 to 33% o f standard
len g th ) and a longer pectoral fin w ith few er fin ray s (length 21 to 26% o f
standard len g th , th u s ab ou t equal to head len g th or lo n g er:
12 to 14
finrays). Raconda russeliana is superficially similar, but lacks a dorsal fin
and has m ore anal finrays (81 to 93 finrays). O th e r p ris tig a s te rid s have
pelvic fins and a shorter anal fin.
G eograph ical D istrib u tion : Indo-W est P a c if ic
(Java Sea, S ingapore north to E ast C hina Sea at
F oochow ).
H abitat and B iology : A long shores and in
estuaries: probably sim ilar to O. tard o o re, but m ore
specimens and data needed.
Size :
head lo n g e r th an
p ecto ra l fin
40"
2o"
To 20 cm standard length.
Interest to Fisheries : Probably
artisanal fisheries for clupeoids.
contributes
to
L ocal
N am es
IN D O N E S IA :
L ip ira m
(if
O. m a c ro g n ath u s o f W eb er & de B e a u fo rt, 1 9 13:96 is
th is sp ecies).
40 °
60 °
80 °
100 °
120°
140 °
160 °
180 °
c lic k for n e x t p a g e
c lic k fo r p r e v io u s p a g e
297
P R IS T O d o n t
O d on togn ath u s L acep è d e , 1800
F o rm e rly CLU P O d o n t
O d o n to g n ath u s L ace p èd e , 1800, H is t.n a t.P o is s .. 2:220 (type: O dontognathus
bolus Schneider, 1801, Syst.Ichthyol.B loch.:556 (type:
Odontognathus mucronatus
D iagnostic Features
M oderate-sized m arine, estuarine
and freshw ater clupeoids (to about 17 cm standard length), body
elongate and highly com pressed, w ith a sharp keel o f scutes
along belly. Eye fairly large: low er ja w som ewhat projecting,
m outh directed obliquely upward: upper ja w long, reaching to
or beyond gili opening: no to o th ed hyp o -m ax illa b etw een hind
tip o f pre-m axilla and low er bulge o f m axilla blade: ja w teeth
sm all or m inute, w ith a d istin c t to o th le ss n otch at cen tre o f
upper jaw . G illrakers short and slender, 18 to 26 on low er part
o f a rc h .
D o rsa l fin sm a ll, w e ll b e h in d m id p o in t o f bo d y :
pectoral fins large: pelvic fins absent: anal fin very long, with
55 to 85 fin ra y s its o rig in w e ll in a d v a n c e o f d o rsa l fin.
Closely resem bles Pacific O pisthopterus, in w hich the upper jaw
barely reaches to eye centre, as also in N eoopisthopterus, w hich
has a shorter anal fin (50 fin ray s or lesss). C h iro c e n tro d o n has
canine teeth and other clupeoids are deeper-bodied.
m ucronatus L acepède).
Lacepede).
no
\
hypo-m axilla
gili
opening
tip o fm axilla
B iology, H abitat and D istribution : M arine and inshore, but penetrating 10 km or more up rivers in
cases. A tlantic and Pacific coasts and drainage o f South and Central America.
Species :
G n ath o ­
some
A ccording to W hitehead (1973a:73), there are 3 species:
A tlan tic
Ö com pressus M eek & H ildebrand, 1923, Panam a to the Guianas
O. m ucronatus L acepede, 1800, Trinidad to Brazil
P acific
O. panam ensis (Steindachner, 1876), Honduras to Panama Bay.
O dontognathus com pressus
Meek & H ildebrand, 1923
P R IS T O d o n t 1
F o rm erly C LU P O dont 1
O dontognathus com pressus M eek & H ildebrand, 1923, Pubis Field M us.nat.H ist.. 15( 1): 194 (Colón, Panam a).
Synonyms : O d o n to g n ath u s m u c ro n a tu s:V alen c ien n es. 1 8 47:91, fig. 611 (C ay en n e, S urinam : m isid e n tifie d ):
O dontognathus com pressus - F W N A ,1964:433, fig. 110, 111 (Panam a, G u lf o f V enezuela): C ervigón, 1966:130
(V enezuela, com piled): W hitehead, 1973a:78, fig. 25 (T rinidad): W hitehead & B auchot, in press (m ucronatus o f
Valenciennes).
FAO Names :
E n - C arib b ea n longfin herring.
298
D iagnostic Features :
Body long and com pressed, with
an uninterrupted keel o f 24 to 28 scutes, the outer edges o f the
sc u te s d is tin c tly se rra te d .
U p p e r ja w b ro a d at m id p o in t,
tapering posteriorly, the tip reaching or surpassing gili opening.
P ectoral fins large; p elvic fins absent; dorsal fin rath er sm all,
set far back on body;
anal fin lo n g , w ith 55 to 61 finrays.
R esem bles Q. m ucronatus (w hich it overlaps in Trinidad and
C ay en n e), w hich has sm ooth edged scutes w ith a gap b elow
p e c to ra l fin b a se , and m o re an al finrays (70 to 85); no o th e r
clupeoid in the area has so m any anal finrays or such a long
uno er iaw (N eo o p isth o p te ru s, C h iro cen tro d o n ).
See CLUP
Odont 1, Fishing Area 31.
Geographical Distribution :
A tlan tic coasts and
perhap s e s t u a r ie s o f C e n tr a l a n d S o u th A m e ric a
(P an am a, coasts o f V en ezu ela, T rin id ad , S urinam or
C ayenne - th e m u c ro n a tu s fig u re d by V a le n c ie n n e s ,
1847).
ed8e o f scutes serrated
40-
H abitat and B iology : Coastal, inshore, probably
entering estuaries, but more data needed.
Size :
To at least 13 cm
Interest to F ish eries :
artisanal catches.
standard length.
P ro b ab ly
20°
co n trib u tes
to
o°
L ocal N am es : -
20°
120°
100°
80°
60°
40°
PRIST O dont 2
O dontognathus m ucronatus L a c e p è d e , 1800
F o rm erly C LU P O dont 2
O d o n to g n ath u s m u cro n atu s L a c e p è d e, 1800, H ist.nat.poiss.. 2:220, pi. 7, fig. 2 (Cayenne).
Synonyms : O dontognathus aculeatus Schinz, 1822:300
1964:431, fig . 109 Trirnidad; C e r v ig ó n , 1 9 6 6 :1 3 1 , fig. 55
m u c r o n a tu s ); Id em . 1973a:73, figs 22-24 (G uayana, S urinam
fig. 30 (B razil, com piled); U yeno, M atsuura & F u jii, 1983:94,
in press (type o f m ucronatus).
FAO Names : E n - G uiana longfin herring.
(on L acep èd e); O dontognathus m ucronatus - F W N A ,
( o f f O rin o c o d e lta ; W h ite h e a d , 1 9 6 7 :1 0 4 (ty p e o f
coasts and riv ers); F igueiredo & M enezes, 1978:25,
fig. (colour photo) (Surinam ); W hitehead & B au ch o t,
299
D iagnostic F eatures : Body long and strongly com ­
pressed, w ith a keel o f scutes along belly, but interrupted below
pectoral fin base; scutes 7 or 8, then a gap, followed by 10 to
13, the o u ter edges o f scutes sm ooth and not serrated . U pper
jaw broad at midpoint, tapering posteriorly, the tip reaching or
surpassing gili opening; no h y po-m axilla. P ectoral fins large;
pelvic fins absent; dorsal fin very small, set far back on body;
anal fin long, with 70 to 85 finrays. R esem b les Cf c o m p ressu s,
w hich has serrated o u ter edges to scutes w ith no gap b e lo w
pectoral fin base and only 52 to 62 anal finrays;
no oth er
elongate clupeoid in the area has so many anal fin ray s or such a
long upper jaw (N eoopisthopterus. C hirocentrodon).
See CLUP
Odont 2, Fishing Area 31
gap in scutes
Geographical Distribution :
A tla n tic
coasts and
riv e rs o f S o u th A m e ric a (T rin id a d and th e G u ia n a s;
th e n no re c o rd s u n til so u th e rn B ra z il, fro m S e rg ip e
south to about Rio de Janeiro).
H abitat and B iology : O ff coasts, traw led at
down to 40 m, also close to shore, o ff river m ouths, in
estuaries and some distance up rivers in freshw ater
(e.g. up C oppenam e R iver near W ilh elm in a M ts,
Surinam). Possibly spawns in rivers.
Size :
To 15, perhaps to 17 cm standard length.
In terest to F ish eries :
N o special fishery, but
contributes to clupeoid catches in coastal waters.
Local Names : SURINAM:
M atsuura & F ujii, 1983).
Literature :
G iana-nishin (Uyeno,
See under Synonyms.
Remarks : The extension o f the range to include
southern B razil is based on specim ens in the M useu de
Zoología, Sâo Paulo.
T» 0 ^ 7 S Cf, ...
300
PRIST O dont 3
Odontognathus p a n a m en sis (Steindachner, 1876)
Pristigaster ( O dontognathus) panamensis
1876, Ichthvol.B eitr., (5):24 (repeat).
Steindachner,
1876,
Sber.Akad. W iss. W ien,
74:72
(Panam a);
Idem ,
Synonyms : O dontognathus panamensis - Meek & H ildebrand, 1923:194, but the illustration is O pisthopterus
dovii (Cham e Point, Panam a); N orm an, 1923:16 (Panam a); P eterso n ,1956:183 (G u lf o f N ic o y a , C osta Rica);
Whitehead, 1971:27, fig. 3a, b (type o f panam ensis: key).
FAO Names : En - Panam a longfin h errin g .
D iagnostic Features :
Body long and compressed, with
an uninterrupted keel o f about 29 or 30 scutes, the outer edges
o f the
scutes distinctly serrated. U pper jaw broad at midpoint,
tapering p osteriorly , the tip reaching or surpassing the gili
opening;
no hypo-m axilla.
P ecto ral fins large; pelvic fins
absent;
dorsal fin rather small, set far back on body; anal fin
long, w ith 61 to 68 finrays. R esem b les th e A tla n tic Cf com ­
pressus , w hich has only 55 to 61 anal finrays and 24 to 28
scutes;
also very sim ilar to species o f O p isth o p te ru s, w hich
have a short upper jaw , not reaching beyond eye. Pliosteostom a
has a small toothed hypo-m axillary bone betw een the hind tip o f
the pre-m axilla and the low er bulge o f the m axilla blade. O ther
clupeoids have a shorter anal fin.
o f scutes serrated
G eographical D istribution : Pacific coasts and
perhpas estuaries o f central A m erica (G ulf o f Fonseca,
Honduras to Panama; possibly it extends further south).
H abitat and B iology : M arine, inshore, perhaps
like its congeners entering estuaries. Feeds on small
crustaceans and fishes (Peterson, 1956).
Size :
40°
To 17 cm standard length.
20°
Interest to Fisheries :
ries, but not in great numbers.
E n te rs a rtisa n a l f is h e ­
L ocal N am es : Literature :
Peterson (1956
o°
- maturity, food).
20 °
120 °
1 00 °
80 °
60 °
40°
301
Pristigaster C uvier, 1816
PR IST
P rist
Pristigaster Cuvier, 1816, Règne anim al, 1st ed., 2:176 (d e s c r., no species named); Id e m , ib id .. 4:pl. 10, fig.
3 (presum ed type figured, but not named = Pristigaster cayana C uvier, 1829); Idem , 1829, Règne anim al, 2nd ed.,
2:321 (P. tardoore and P. cay a n u s listed); Id e m . 1830, ibid., 3:pl. 12, fig. 3 (repeat o f 1st ed., species still not
nam ed) (the problem o f the type species is discussed by W hitehead, 1967:100, 101).
Pristogaster Sw ainson, 1838,
N a t.H is t.A n im ., 1:266, fig. 57 (type: not in d ica ted , but figure =R cay an a
C u v ier); Idem . 1839, ib id .. 2:294 (P.
indicu s, on R u sse ll’s T artoore = C lu pea m elasto m a S ch n eid er, a species o f
Ilish a , and R m artii A g assiz = R
cay an a).
D iagnostic Features : R elatively small freshw ater c l u ­
peoid fishes (to 15 cm standard length) characterized by the
enorm ously deep body and strongly convex belly profile. M outh
directed nearly upw ard; no toothed hypo-m axilla. Pelvic fins
usually absent; anal fin long, w ith more than 40 finrays. No
other clupeoid has such a deep body (at m ost to about 40% o f
standard length; cf. over 50% in Pristigaster).
Biology, H abitat and D istribution : R iverine, but appa­
re n tly reaching to m outh o f A m azon and thus perhaps able to
tolerate some salinity. K nown only from the A m azon drainage.
Interest to Fisheries : Little or none.
Species : Until recently, only a single species has been re c o g n iz e d , but there is some evidence (see Remarks
under R cayana) that a second species may be present:
R cayana C uvier, 1829, A m azon drainage.
P ristig a ste r
caya n a
C uvier, 1829
P ristig aster
cayanus C uvier, 1829 (F ebruary or M arch), R ègne an im a l, 2nd. ed.,
A m erican seas); Id e m . 1830, ib id .. 4 :p l. 10, fig .3 (m e re ly a s P r is tig a s te r ).
PR IST P rist 1
2:321 (fo o tn o te, nam e only,
Synonym s
P ristigaster argenteus Schinz, 1822:300 (on C u v ier’s fig.; nom en o blitum ); P ristigaster
lichtensteinii Jarocki, 1822:332, fig. 3 (on C uvier’s fig., reproduced; for Jarocki’s Polished, o f the Regne anim al,
see W h iteh ead , 1982b; nom en o b litu m ); P ristig a ste r tria n g u la ris S tark , 1828:408 (on P ristig a ste r fig. in C u vier,
1816:pl. 10, fig. 3; nomen oblitum ); Pristigaster martii A gassiz, in Spix & A gassiz, 1829 (M ay or June):55, pi. 24a
(Amazon); ju st post-dates cayanus by a m onth or so);
Pristigaster am ericanus G uerin-M eneville, 1844:33, pi. 57,
fig. 3 (A tlantic coast o f South A m erica); Pristigaster phaeton V alenciennes, 1847:338 (A m azon); Pristigaster
cayana FW N A , 1964:428, fig. 108 (synopsis); W hitehead, 1967:100, 102 (types o f cay an a, phaeton): W hitehead &
M yers, 1971:487 (validity o f m artii); W hitehead, 1973a:85 (triangularis a nom en oblitum ; synon.); W hitehead &
B auchot, in press (types o f cayana, phaeton).
FAO Names : E n - A m azo n h atch e t herring.
302
D iagnostic Features : Body extrem ely deep and com pressed, its depth 50 to 60% o f standard length, belly
profile strongly convex (alm ost circular from throat to anus), w ith 30 to 33 scutes. Eye large; low er jaw
projecting, m outh directed alm ost vertically upw ard; no toothed hypo-m axillary bone betw een hind end o f p r e ­
m axilla and low er bulge o f m axilla blade. Pectoral fins high on body; pelvic fins usually absent, but present in
some (see R em arks below); dorsal fin before m idpoint o f body; anal fin long, its origin behind dorsal fin base, with
45 to 54 fin ra y s upper c a u d a l fin lobe ending in a filam ent in larger fishes. The deep body distinguishes it from
other A m azon clupeoids (P e llo n a . Ilisha, etc.). Superficially, it resembles the deep-bodied gasteropelecine fishes,
but these have a keel on the chest and no distinct bony scutes.
G eographical D istribution
A m azon drainage
(from perhaps m outh o f R io B ra n co at G o ia ç u , Rio
Solim öes, Rio Ju ru á and as far as the R io U cayali in
Peru fide F o w ler, 1940:16).
H abitat and B iology
R iverine, perhaps en ­
tering slightly saline w aters near m outh o f A m azon (but
this part o f its range not well docum ented). Compared
w ith th e ‘f l y in g ’ c h a r a c o id f is h e s (G a ste ro p e le c u s,
T h o raco ch arax , etc.), the v ertically aligned scute arm s
in P ristigaster and the rather small pectoral muscles
sh o w th a t flig h t by v ib ra tio n o f th e p e c to ra l fin s is
a lm o st c e rta in ly im p o ssib le ;
sin ce P ris tig a s te r and
T h o ra c o c h a ra x h av e b e e n c a u g h t to g e th e r (e.g. at
P a r a n á do M ocam bo near P a r i n ti n s ) , is m im icry
involved?
Size : To 14.4 cm standard length, usually about
7 to 10 cm.
20
"
Interest to Fisheries : L ittle or none.
Local Nam es :
B RA ZIL: A p a p a .
40°
Literature :
See under Synonyms.
Rem arks :
P o s s ib ly th e r e a re tw o s p e c ie s ,
ju d g in g from specim en s in the M useu de Z oología, Sâo
Paulo:
A.
P elvic fins ab sen t, low er g illrak ers 22 to 24 (u su ally 23);
probably throughout (5 fishes)
B.
Pelvic fins present (i 4 finrays), lower gillrakers 19 or 20 (usually 20); Rio Solim öes (3 fishes).
R acon d a
A m a z o n fro m R io Ju ru á to M anáus, but
P R IS T Racon
G ray, 1831
R a c o n d a G ra y , 1 8 3 1 , Z o o l.M isc e lla n y . 1:9 (type: R aco n d a ru sselian a Gray).
A p tery g ia G ray, 1835 (20
February),
Illustr.Ind.Zool.H ardw icke. 2:pl. 92, fig. 1 (type: A p te ry g ia ra m c a ra te G ra y , n o t M y stu s ra m c a ra ti
H am ilton-Buchanan = C oilia, an engraulidid).
D iagnostic Features : M edium -sized marine or estuarine clupeoid fishes (to perhaps not more than 20 cm
standard length) w ith elongate and compressed bodies. No dorsal fin; no pelvic fins; anal fin longer than in any
other pristigasterid genus, its base more than h alf standard length. The lack o f a dorsal fin and the very long anal
fin easily distinguish Raconda from O pisthopterus, the m ost sim ilar genus in the Indo-Pacific region.
B iology,
H a b ita t
In terest
to
S p ecies :
and
D istr ib u tio n
F ish eries
: See species.
: See species.
A single species:
R. russeliana G ray, 1831, eastern Indian O cean to Java Sea.
303
PRIST R acon 1
R aconda russeliana Gray, 1831
Raconda russeliana G ray, 1831, Z ool.M iscellany, 1:9 (Sangar Roads, India).
Synonym s :
A p te r y g ia ra m c a ra te G ra y , 1835:pl. 92, fig. 1 (Sangar R o c k s ) ;
A p te ry g ia h a m ilto n i
V a le n c ie n n e s , 1847:333 (o n r a m c a ra te , not on T hryssa h a m ilto n ii G ray = an anch o v y );
F o w le r, 1941:665
(com piled); W hiteh ead , 1973b:216, fig. 4 (key, synon., refs); W ongratana, 1980:217, pis 181, 182 (revision).
FA O Names
En - R aconda.
D iagnostic Features : Body elongate, w ith strongly convex low er profile bearing a sharp keel o f 33 to 38
s c u te s. Lower jaw prom inent, mouth pointing obliquely upward; upper jaw short, not reaching beyond eye centre.
Low er gillrakers 23 to 27. D orsal fin absent; pelvic fins absent; anal fin very long, w ith 81 to 93 finrays, its
origin well before m idpoint o f body. A dark spot behind gili opening. M ost closely resem bles species o f
O p isthopterns, w hich have a small doral fin and a slighlty shorter anal fin (51 to 65 fin ray s, not more than h alf
body le n g th ). Other pristigasterids are deeper bodied, have a well developed dorsal fin and a shorter anal fin.
G eographical D istribution :
Indian Ocean (eastern
coasts o f India, possibly also coasts o f Burm a), Singapore
and Java Sea (south coast o f K alim antan, also B agan Api
Api, Sumatra).
Habitat and Biology :
M arine, or p erh ap s m ore
com m only estuarin e (e.g. A lu h alu h on the B arito R iv er,
K alim antan). Feeds m ainly on praw ns (esp ecially A c ete s),
also copepods. More specimens and data needed.
Size :
To 19 cm standard length.
Interest to F isheries : Probably slight
species seems to be uncom m on in catches.
L ocal N am es :
L iteratu re :
length-w eight).
IN D IA :
since
the
R aconda.
V e rg h e se (1 9 6 2 - food, fecu n d ity ,
Remarks :
R ed u ctio n o f the d orsal fin o ccu rs in
other long-bodied pristigasterids (O pisthopterus, etc.), but
only Raconda has lost this fin.