Clupeiformies

Clupeiformes
CLASSIFICATION , FAMILY CHARACTERS

Introduction
 Clupeiformes are some of the most economically important fishes in the world's
oceans. They have been widely exploited throughout human history, primarily for
food but also as a source of oil, fertilizer, and animal feed.
 Clupeiformes continue to constitute a large portion of world's commercial fisheries.
Although 186 species are exploited by pelagic fisheries worldwide, 50% of the total
landings in 1997 were represented by only seven species. Among these seven, four
are Clupeiformes: the anchoveta (Engraulis ringens), Atlantic herring (Clupea
harengus), Japanese pilchard (Sardinops melanostictus), and South American
pilchard (Sardinops sagax). Herrings and anchovies constitute approximately 25%
of the total fisheries harvest worldwide

In addition to being heavily utilized by humans, Clupeiformes are an
important component of the broader marine ecosystem.
They serve as food items for larger predatory fishes, sea birds, and
marine mammals. Thus, clupeoids sustain other organisms of
importance to humans through ecosystem interactions

General characters
• Clupeoids are small fusiform (tapering toward each end) fishes with streamlined
bodies that facilitate fast swimming in open water.
• They have dark shading on their backs and bright silvery sides.
• Except for the head, their bodies are completely covered in large scales.
• Most clupeoids lack a lateral line, and only in the deticipitoid herring does this
line extend along the body.
• The fins of clupeoids lack spines.

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• A single dorsal fin is located near the middle of the body, and the tail is forked.
• Many clupeoids have a row of scutes, modified scales that usually have sharp
points towards the rear, along the medial line of the belly.
• The smallest cluepoid is the Sanaga pygmy herring (Thrattidion noctivagus),
measuring only 0.83 in (2.1 cm) in standard length; male wolf herrings
(Chirocentrus spp.) are the largest herring, attaining standard lengths of 39 in
(100 cm).

Distribution
Clupeiformes are widely distributed worldwide between 70°N to 60°S
latitude.
They primarily live in oceans, but some species inhabit coastal margins and
fresh water for at least a portion of their lives
In India the clupeids are restricted distribution between latitude 8°N and
14°N and longitude 75°E and 77°E 3 (Malabar upwelling zone along the
southwest coast of India

Habitat
• Nearly all Clupeiformes are open-water, pelagic species.
• Four-fifths of all species are marine, with habitats ranging from nearshore
littoral zones to nearly 100 mi (160 km) offshore.
• Many are found near the surface at times but often move to deeper waters
during the day.
• Some Clupeiformes live in inland waters or are anadromous, moving inland to
spawn. These species utilize bays, estuaries, marshes, rivers, and freshwater
streams as habitats.
• Landlocked populations have formed as shads, alewives, and herrings moved
into lakes or rivers and became trapped between dams

Schooling and shoaling behaviour
• Clupeiformes are perhaps best recognized for the large schools they form.
Schools may include hundreds or thousands of individuals ranging from the
young to adults, but individuals in a school are usually of similar size.
• Schooling is a form of organization in which behaviour is synchronized;
large numbers of fish may swim parallel to each other in the same direction
with fairly uniform spacing. These synchronized aggregations are believed
to confer swimming efficiency and, most importantly, to enable fishes to
avoid or deter predators. Clupeiformes also congregate in smaller, less-
organized shoals, particularly during spawning seasons

Feeding ecology and diet
• Most Clupeiformes filter feed by straining water through
their long and numerous gill rakers.
• They consume plankton, particularly small crustaceans
and the larval stages of larger crustaceans and fishes.
• Some herrings visually locate and target food particles.
Clupeoid fishes are important prey for larger fishes,
seabirds, and marine mammals

School of sardine swimming across the plankton rich water column keeping their mouth open to
facilitate filter feeding

Reproductive biology
• Clupeiformes produce large numbers of offspring, either through a single seasonal
spawning event or by spawning in seasonal peaks throughout the year.
• Most Clupeiformes spawn in shoals by broadcasting large numbers of small,
buoyant eggs in waters near the surface.
• The eggs and larvae drift passively in currents as they develop.
• Herrings, on the other hand, produce demersal eggs that sink to the bottom, where
they often adhere to the substrate until they hatch. After hatching, larvae become
pelagic

classification
 Two suborders, Clupeoidei and Denticipitoidei, are recognized in the
Clupeiformes.
 The Clupeiodei includes the families Chirocentridae (wolf herrings; 1 genus, 2
species), Clupeidae (herrings, menhadens, pilchards, sardines, shads, and sprats;
5 subfamilies, 56 genera, 214 species), Engraulidae (anchovies;
2 subfamilies, 16 genera, 145 species), and Pristigasteridae (sawbelly herrings;,
2 subfamilies 9 genera, 36 species).
 The Denticipitoidei includes the family Denticipitidae (denticle herring;
1 genus, 1 species).

Suborder :Denticipitoidei
Caudal skeleton of extremely primitive type; small arches
present on 2 centra (bodies of vertebrae) to carry the first 3
hypural bones (fused spines of the vertebrae) of the tail fin.
1 family

Family : Denticipitidae
• The denticle herring (Denticeps clupeoides)
is a small (15-cm-long) species of ray-finned
fish found only in the rivers of Benin,
Nigeria, and Cameroon
• It is related to the herrings, but notable for its
large anal fin and its array of denticles-like
scales under the head, which give it almost a
furry appearance.
• It is the sole living member of the
Family : Denticipitidae

Suborder : Clupeoidei
• Characteristic caudal skeleton: the second hypural bone lacks any
connection with the urostyle (tail support) and is separated from it by
a distinct gap.
• Lateral line pores completely lacking on trunk. Keeled scutes
(projecting scales) usually present along the ventral midline of the
abdomen. Family Pristigasteridae.
• Mouth superior or terminal; abdominal scutes present; anal fin long,
30–92 rays; no notch in third hypural bone of caudal skeleton.
• Primarily marine, some freshwater; Atlantic, Indian, and Pacific
oceans. 9 genera, 34 species.

Family : Clupeidae
 The family currently comprises 54 genera and approximately 200 species
 Body usually fusiform, round to strongly compressed.
 Head without scales; jaw teeth, when present, are small or minute.
 A single dorsal fin, small and near midpoint of body; pelvic fins more or less below
dorsal fin base; dorsal and pelvic fins absent in some species; soft rays only.
 Lateral line spanning a few scales behind the head in some species, missing in others;
scales cycloid (smooth to touch).
 Branchiostegal rays usually 5-10.
 Most feed on small planktonic animals.
 Size range (adults): from 2 to 75 cm.

Subfamily: Clupeinae
Herklotsichthys
The genera comprises of 12 species all around the world
and 2 of them are found in Indian waters
 Opercle smooth
 Gill opening with two fleshy outgrowths
 Pelvic finrays 8 or 9; back blue/green
 Fronto-parietal striae In top of head few, 3 to 8 lower
portion of paddle-shaped 2nd supramaxilla longer than
upper
 No dark spot at origin of dorsal fin

Herklotsichthys punctatus (Spotback
herring)
 No dorsal spines;
 Dorsal soft rays(total): 13-21
 Anal soft rays: 12 - 23.
 Flank silvery, without spots, but a series of
small dark spots on back on either side of
dorsal fin base or behind it.

Herklotsichthys quadrimaculatus (Bluestripe herring)
 Elongate wing-like scales underneath paired pre-
dorsal scales separate it from all other species
except H. koningsbergeri and Herklotsichthys
Species A, which have prominent black spots on
the flank, and Herklotsichthys Species B, which
has dusky tips to dorsal and caudal fins and more
lower gill rakers.
 Flank silvery with an electric blue line preceded
by two orange spots located on each operculum

There are currently 22 recognized species in this
genus
 Opercle smooth
 Pelvic finrays 8 or 9
 Back blue/green
 Fronto parietal striae (on top of head)
usually many, 8 to 19 lower portion of
paddle-shaped 2nd supramaxilla equal to
upper
 Gillrakers more than 40 in fishes over 5 cm
standard length
 Predorsal scales paired and overlapping in
midline
Sardinella

Vertebrae: 45 - 49.
Body sub-cylindrical;
A faint golden spot behind gill opening,
followed by a faint golden mid-lateral line;
A distinct black spot at hind border of gill cover
(absence of pigment).
Sardinella longiceps (Indian oil sardine)

The other species that are found in Indian waters are as follows
Sardinella albella
Sardinella brachysoma
Sardinella fimbriata
Sardinella gibbosa
Sardinella jussieu
Sardinella melanura
Sardinella sindensis

Amblygaster
3 species representing this genera of which 2
are found in Indian coast
 Opercle smooth
 Back blue/green
upper
 Gillrakers 26 to 43
 Predorsal scales forming a well-defined
single median row

Amblygaster clupeoides
 Body fairly elongate, subcylindrical in cross-
section
 Belly rounded, but with a low keel of scutes.
 Head moderate, about 4 times in standard
length; Gill openings with 2 fleshy
outgrowths; gillrakers 26 to 31 on lower limb
of first gill arch.
 Dorsal fin origin at mid-point of body or a
little nearer to caudal fin base
 8 pelvic fin rays;
 Anal fin origin well behind dorsal fin base.
 Predorsal scales forming a single (median)
series. Colour: back blue/green, flanks silvery
and without spots.

Amblygaster leiogaster (Smoothbelly sardinella)
 Body moderately slender, belly rather
rounded, scutes not prominent.
 Distinguished from A. sirm by the absence
of spots along the flank (gold in life, black on
preservation) and fewer lower gill rakers.
 Closely resembles A. clupeoides, which has
fewer lower gill rakers.

Escualosa thoracata
 Opercle smooth
 Back blue/green
upper
 Gill opening smoothly rounded
 Pelvic fin rays 7
 Body creamy white

Sardinops sagax (South American pilchard)
 Vertebrae: 48 - 54.
 Body cylindrical and elongate
 ventral part of operculum with clear cut
bony striae radiating downwards
 Belly rounded with ventral scutes
 Back blue green; flanks white, with 1 to 3
series of dark spots along the middle
 The radiating bony striae on the operculum
distinguish this species from all other
clupeids

Subfamily: Dorosomatinae
 Branchiostegal rays 4 to 8
 scutes present
 Anal fin short (less than 30 rays)
 lower jaw not prominent
 Mouth inferior, lower jaw flared at corners
 last dorsal ray often filamentous
• Last dorsal fin ray filamentous
• Predorsal scales paired .......................... Nematalosa
• Last dorsal fin ray normal
• Predorsal scales forming a single median row ....Anodontostoma

Anodontostoma chacunda (Chacunda gizzard shad)
 Body depth increasing with size of fish, 40 to
70 % standard length in fishes over 10 cm.
 Second supra-maxilla a mere splint.
 Longest gill rakers on lower part of arch less
than corresponding gill filaments.
 Hind edges of scales toothed, the teeth
thinner than the gaps between them; a
median series of pre-dorsal scales.
 A large black spot behind gill opening.

Nematalosa nasu (Bloch's gizzard shad)
 Dorsal soft rays 15-19;
 Dark bluish dorsally, silvery below
 A dark spot behind gill opening.
 Belly with 28 to 32 (usually 30) scutes.
 Anterior arm of pre-operculum with the third
infra-orbital bone immediately above it, no
fleshy gap between;
 Lower jaw strongly flared outward.
 Pectoral axillary scale present; hind edge of
scales distinctly toothed.
 There are about 11 species of this genera
were found all over the world another species
that occur in east coast of India is
Nematalosa galatheae

Subfamily: Alosinae
 Branchiostegal rays 4 to 8
 scutes present
 Mouth terminal, lower jaw not flared outward at corners
 Last dorsal ray not filamentous
 Upper jaw with a distinct notch at centre
• Fronto-parietal striae (on top of head) many, 8 to 14.
• Gillrakers on inner arches distinctly curved outward
• Scales perforated ............................................... Hilsa
• Fronto-parietal striae weakly developed, usually hidden by skin
• gillrakers on inner arches straight
• scales unperforated ……............................ Tenualosa

Hilsa kelee (Kelee shad)
 Dorsal soft rays 16-19
 Top of head with numerous fronto-parietal
striae
 Upper jaw with median notch.
 Gill rakers about 100 to 175, those on inner
arches distinctly curled; outer row of gill
filaments on first arch not more than half
length of gill rakers.
 A series of small triangular scales above axil
of pectoral fin; hind part of body scales
perforated.
 A black spot behind gill opening; usually 10
along flank.

Tenualosa ilisha (Hilsa shad)
 Dorsal soft rays 18-21
 Anal soft rays 18 - 23.
 Belly with 30 to 33 scutes.
 Distinct median notch in upper jaw.
 Gill rakers fine and numerous, about 100 to
250 on lower part of arch.
 Fins hyaline.
 A dark blotch behind gill opening, followed
by a series of small spots along flank in
juveniles.
 Color in life, silver shot with gold and
purple.

Tenualosa toli (Toli shad)
 Belly with 28 to 30 scutes.
 A distinct median notch in upper jaw, which
distinguishes it from other similar clupeids,
except Hilsa kelee.
 Gill rakers fine but not numerous, 60 to 100
on lower part of arch.
 Caudal fin short.
 At most, a dark diffuse mark behind gill
opening, but no other spots on flank.
Total about 5 species of this genera are found all
around the world

Family : Engraulidae
 The 144 species are placed in 17 genera; they are found in the Atlantic, Indian and
Pacific Oceans, and in the Black Sea and the Mediterranean Sea.
 Mouth inferior. Upper jaw produced.
 Jaw teeth well developed to absent.
 Gill rakers on lower limb of first arch 10-50 or more.
 Branchiostegal rays 7-19.
 A silvery stripe down flanks. Body translucent.
 Abdominal scutes present in most Old World anchovies; absent in New World
anchovies, except for one pelvic scute.
 Mostly filter feeding on zooplankton a few piscivorous.
 About 50 cm maximum length; most species below 15 cm.

Engraulis
• Body normal, not tapering to a point,
• Caudal fin forked
• Anal fin not joined to caudal fin
• No scutes before or behind pelvic fin base
(except a single pelvic scute)
• Maxilla tip blunt
• Anal fin origin well behind last dorsal fin
ray
There are about 9 species occurring all
over the world and non of them occurs in
Indian waters

Stolephorus
• Scutes needle-like, present only before pelvic fin base
• Anal fin short, less than 25 fin rays
• Small, slender fishes (usually 4 to 8 cm) with a silvery band on
flanks
• 20 species of this genera are found all
around the world and 8 species are found in
Indian waters

Stolephorus commersonnii (Commerson's anchovy)
 Belly slightly rounded with 0-5 small needle-like pre-
pelvic scutes.
 Maxilla tip pointed, reaching to or a little beyond hind
border of pre-operculum, the latter convex, rounded.
 Small teeth on hyoid bones.
 Isthmus muscle tapering evenly forward.
 Body light transparent fleshy brown with a pair of dark
patches behind occiput, followed by a pair of lines to
dorsal fin origin. Bears a silver stripe on flanks.

Stolephorus indicus (Indian anchovy)
Dorsal soft rays15-17
Anal soft rays 18 - 21.
Belly with 2 to 6 small needle-like pre-pelvic
scutes.
Maxilla tip pointed, reaching to or only just
beyond front border of pre-operculum; hind
border of pre-operculum convex, rounded.
Isthmus muscle tapering evenly forward to hind
border of branchial membrane.
Body light transparent fleshy brown, with silver
stripe down flank; no dark pigment lines on
back between head and dorsal fin.

Thryssa
• Scutes present before and behind pelvic fin base, forming a keel
• Anal fin longer, with more than 25 fin rays
• Larger, more compressed fishes (usually 10 to 14 cm), with a dark
humeral area behind gill opening
About 24 species occurs all around the world and
11 species occurs in India

Thryssa malabarica (Malabar thryssa)
 Belly with 23 to 26 keeled scutes (isthmus to anus).
 Tip of snout about level of upper rim of eye.
 Maxilla moderate; first supra-maxilla small, oval;
 Jaw teeth small.
 A dark blotch behind upper part of gill opening;
small spots on cheek, gill cover, maxilla and paired
fins;
 Gill arches pinky orange, inside of gill cover yellow
and gold; Inner part of anal fin deep yellow, margin
whitish.

Thryssa hamiltonii (Hamilton's thryssa)
 Belly with 23 to 26 keeled scutes from
isthmus to anus.
 Tip of snout above level of eye center,
usually about level of upper rim of eye.
 Maxilla short or moderate; first supra-
maxilla small, oval.
 A dark blotch behind upper part of gill
opening.

Coilia
• Body tapering to a point, caudal fin small, not
forked
• 6 or 7 upper pectoral rays filamentous
• Anal fin very long and joined to caudal fin
There are about 13 species found all over the
world and 4 species of this genera are found in
Indian waters

Coilia neglecta (Neglected grenadier anchovy)
 Anal soft rays 80.
 Body tapering, belly rounded in front of
pelvic fins,with 12 to 15 keeled scutes from
just behind pectoral fin base to anus. Maxilla
almost or just reaching to edge of gill cover.
 Pectoral fin with 6 long filaments; the
branched fin rays longer than those of pelvic
fin

•
Coilia dussumieri (Goldspotted grenadier anchovy)
 Anal soft rays 80.
 Body tapering, belly rounded before pelvic
fins, with 12 to 15 keeled scutes from just
behind pectoral fin base to anus.
 Maxilla short.
 Pectoral fin with 6 long filaments and 9 to 11
(rarely 8) branched fin rays.
 Flanks and belly with golden or pearly spots
(light organs) in rows below scales, also
along isthmus, edge of lower jaw, on cheek
and gill cover.
The other two species includes Coilia ramcarati
and Coilia reynaldi

Family : Spratelloidinae
• Anal fin short (less than 30 rays)
• Lower jaw not prominent
• Mouth terminal, lower jaw not flared outward
at corners
• Last dorsal ray not filamentous
• Upper jaw without a median notch
• Two supramaxillae
• Scutes absent
Even though many species are remarked
none of them are found in Indian waters

Family : chirocentridae
 Body elongate and compressed.
 Jaws with fanglike teeth.
 Intestine with spiral valve.
 Gill rakers 17-22.
 Pyloric caeca absent.
 Scales small.
 Dorsal fin rays 16-18.
 Pectoral fin rays 13-15.
 Branchiostegal rays 6-8.
 Vertebrae 70-74.
 Attains 3.6 m maximum length.
 Ravenous carnivores

Chirocentrus dorab
 Scales numerous, small, usually lost. Silvery
 Back bright blue (fading to grey) flanks bright silver.
 The slightly shorter pectoral fin and the black marking of
the upper part of the dorsal fin are the only satisfactory
characters separating this species from C. nudus .
 Fins spineless
 with numerous dagger-like teeth
Chirocentrus nudus
 Scales numerous, small, usually lost. Silvery
 Back bright blue (fading to grey) flanks
bright silver.

Family : Dussumieriidae
 Differs from Clupeidae in having a single pelvic scute
 Branchiostegal rays fewer (12 to 15) and posterior part of scales
marked with numerous tiny radiating striae.
 Color is iridescent blue with a shiny gold or brass line below
(quickly fading after death)
 Hind margin of tail broadly dark.
 W-shaped pelvic scute; isthmus tapering evenly forward; more
anal fin rays.

These were included as sub family under family cluepidae and later
separated which includes 2 Genera and 9 Species
Dussumieria acuta (Rainbow sardine)
Anal soft rays 14 - 18.
Branchiostegal rays fewer (12 to 15) and
posterior part of scales marked with
numerous tiny radiating striae.
Color is iridescent blue with a shiny gold or
brass line below (quickly fading after death);
hind margin of tail broadly dark. W-shaped
pelvic scute;
Isthmus tapering evenly forward; more anal
fin rays.

Family : Pristigasteridae
 Moderate or sometimes quite large clupeoid fishes (usually to about 20 to 25 cm
standard length), but some South American members (Pellona) up to 50 cm standard
length.
 Body compressed, very deep in some (Pristigaster), more elongate in others with
complete series of scutes along abdomen.
 Mouth terminal or more often with lower jaw projecting beyond upper with 2
supramaxillae, and jaw teeth usually small or minute (but enlarged canines in
Chirocentrodon).
 A toothed hypomaxillary bone present in some genera, joining posterior tip of
premaxilla to blade of maxilla. Tooth plates fused to second basibranchial and to one or
more pairs of hypobranchials, especially the second pair.

 Adipose eyelids with broad vertical opening in middle.
 No spiny rays in dorsal fin;single,short dorsal fin (if present) near midpoint of body.
Dorsal-fin origin (if present) near body midpoint in very deep-bodied forms, but posterior
to midpoint in elongate forms; reduction of dorsal fin occurs in long-bodied
pristigasterids (Opisthopterus, etc.), but only Raconda has completely lost this fin.
Predorsal bones either upright or inclined forward (inclined backward in all clupeids,
except upright in Ramnogaster).
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 Pectoral fins moderate or large in elongate species
 Pelvic fins small, with 6 or 7 fin rays, but absent in some genera
 Anal fin long, with at least 30 fin rays (60 or more in some species).
 Caudal fin forked. No gap between second and third hypural bones (upper and lower
caudal-fin rays separated by this gap in clupeids).
 Scales cycloid, without posterior striations, adherent or easily lost, of moderate size
(about 35 to 55 in lateral series).
 No lateral line.
 Colour: typically blue-green on dorsum and silvery on sides, but generally without
distinctive colour patterns

Subfamily : Pellonulinae
 Toothed hypomaxilla present
 Anal fin short (less than 30 rays);
 Lower jaw not prominent
 Mouth terminal, lower jaw not flared
outward at corners
 Last dorsal ray not filamentous
 Upper jaw without a median notch
 A single (posterior) supra maxilla

The genera ilisa include 16 species The genera Chirocentrodon includes 1 species
• Ilisha africana Chirocentrodon bleekerianus
•Ilisha amazonica
Ilisha compressa The genera Neoopisthopterus includes 2 species
Ilisha elongate Neoopisthopterus cubanus
•Ilisha filigera Neoopisthopterus tropicus
•Ilisha fuerthii
• Ilisha kampeni The genera Pellona includes 6 species
• Ilisha lunula Pellona altamazonica
• Ilisha macrogaster Pellona castelnaeana
•Ilisha megalopteran Pellona dayi
•Ilisha melastoma Pellona flavipinnis
•Ilisha novacula Pellona ditchela
•Ilisha obfuscate Pellona harrower
•Ilisha pristigastroides
•Ilisha sirishai The genera Pliosteostoma includes
•Ilisha striatula Pliosteostoma lutipinnis

Subfamily : Pristigasterinae
 Body moderately deep, compressed belly with a sharp keel of
scutes.
 Head large, lower jaw strongly projecting,
 Eye large
 Dorsal fin short
 Anal fin long.
 Scales with vertical striae overlapping at centre or continuous.

Pristigasterinae includes 4 genera and 12 species
•Odontognathus 3species
• Odontognathus compressus
•Odontognathus mucronatus
•Odontognathus panamensis
•Opisthopterus 6 species
• Opisthopterus dovii
•Opisthopterus effulgens
•Opisthopterus equatorialis
•Opisthopterus macrops
•Opisthopterus tardoore
•Opisthopterus valenciennesi
•Pristigaster 2 species
•Pristigaster cayana
•Pristigaster whiteheadi
•Raconda 1species
•Raconda russeliana

reference
 FAO Fisheries Department. (13 Nov. 2002)
<http://www.fao.org/fi/default.asp>
 FishBase. 8 Aug. 2002 (13 Nov. 2002).
<http://www.fishbase.org/search.cfm>Katherine E. Mills, MS
 A contribution to the classification of Clupeoid fishes P.J.P.
Whitehead
Pages 737-750

Clupeiformies

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