Int. J. Aquat. Biol. (2017) 5(3): 128-192 

ISSN: 2322-5270; P-ISSN: 2383-0956

Journal homepage: www.ij-aquaticbiology.com 
© 2017 Iranian Society of Ichthyology 

Review Article 
Review of the Herrings of Iran (Family Clupeidae) 

  
Brian W. Coad1 

 
Canadian Museum of Nature, Ottawa, Ontario, K1P 6P4 Canada.

 

 

 

 

Article history: 
Received 4 March 2017 

Accepted 5 May 2017 

Available online 2 5 June 2017 

Keywords: Morphology, Biology, Alosa, Clupeonella, Tenualosa, Kilka, Shad. 

Abstract: The systematics, morphology, distribution, biology, economic importance and 
conservation of the herrings (kilkas and shads) of Iran are described, the species are illustrated, and 

a bibliography on these fishes in Iran is provided. There are 9 native species in the genera 

Clupeonella, Alosa and Tenualosa in the Caspian Sea and rivers of southern Iran. 

 

  
Introduction 

The freshwater ichthyofauna of Iran comprises a 

diverse set of families and species. These form 

important elements of the aquatic ecosystem and a 

number of species are of commercial or other 

significance. The literature on these fishes is widely 

scattered, both in time and place. Summaries of the 

morphology and biology of these species were given 

in a website (www.briancoad.com) which is updated 

here for one family, while the relevant section of that 

website is now closed down. Other families will also 

being addressed in a similar fashion. 

 

Family Clupeidae 

Herrings, shads, sardines, sprats, pilchards and 

menhadens are moderate-sized fishes, usually less 

than 25 cm long, found in warmer marine waters 

with some species anadromous or permanent 

freshwater residents. There are about 64 genera and 

about 218 species world-wide (Nelson et al., 2016), 

with 8 species in the Caspian Sea and one commonly 

found in Persian Gulf drainages. Some other, 

primarily marine, species are known to enter rivers 

in southern Iran (e.g. Nematalosa nasus (Bloch, 
1795) and Sardinella sindensis (Day, 1878)) but are 
not dealt with here (Hashemi and Ansary, 2012; 

Jouladeh-Roudbar et al., 2015). The diversity of this 
                                                           
* Corresponding author: Brian W. Coad                                                                                       DOI: https://doi.org/10.22034/ijab.v5i3.282 

E-mail address: bcoad@mus-nature.ca 

family in the Caspian Sea is seen in the number of 

subspecies which have been described, rather than in 

genera. At the species level these are Caspian Sea 

endemics. A study by Pourrafei et al. (2016) based 

on the nuclear gene RAG1 did not support the 

monophyly of Clupeidae but, as an abstract, details 

are lacking. These fishes are dealt with as a single 

family here. 

Curiously, the species and subspecies in the 

Caspian Sea are generally of larger size than their 

relatives in the Black Sea basin. These observations 

are attributed to the variable environment in the 

Caspian Sea over time, with repeated changes in 

salinity and temperature which the fish could not 

avoid. Black, Mediterranean and Atlantic species 

lived under more stable conditions and could, in any 

case, retreat from lowered temperatures for example. 

In addition, the Caspian Sea clupeids lacked the 

competitors which entered the Black Sea from the 

Mediterranean and Atlantic and some (Clupeonella 
spp., Alosa caspia) could occupy the pelagic, 
planktivore niche taken up by other species in the 

Black Sea. There are no other pelagic fish but these 

herrings in the stable salinity areas of the Caspian 

Sea. 

These fishes usually have modified scales on the 

belly forming abdominal scutes with a saw-like 



129 
 

Int. J. Aquat. Biol. (2017) 5(3): 128-192 

 
edge. Most species have two, long, rod-like 

postcleithra. The lateral line is usually absent or on 

only a few scales. Silvery cycloid scales are easily 

detached and are found only on the body. The mouth 

is usually terminal with jaws about equal in length. 

Teeth are small or absent but gill rakers are long and 

numerous for sieving plankton. Fins lack spines and 

there are no barbels. There is no adipose fin. The 

pectoral and pelvic fins have a large axillary scale. 

The caudal fin is deeply forked. The eye is partly 

covered by an adipose eyelid. The flesh is 

particularly oily and is highly nutritional. 

Members of this family often form immense 

schools in surface waters of the ocean and the 

Caspian Sea where they feed on plankton. Schooling 

is an anti-predator device making it difficult for a 

predator to pick out an individual from a tight mass 

of fish. There is also a "sentry effect" where 

awareness is increased by the presence of many fish. 

The school is maintained by a balance between 

visual attraction and lateral line stimulus repulsion. 

Herring can feed on the smaller plankton, less than 

300-400 µm, at night by filter-feeding but during the 

day can also use particulate feeding. In the latter, 

they select larger plankton using the area temporalis, 

a specialised ventro-posterior region of the retina 

which improves vision as herring approach food 

items from slightly below. 

Herring are easily caught and are extremely 

valuable to commercial fisheries. They are the most 

important fishes economically, both as food for man 

and also for many other commercial fish species. 

Wars have been fought over fisheries for herrings. In 

one year, members of the herring family made up 

37.3% of all fish caught in the world. Some are used 

for fish meal, as fertiliser and as an oil source. The 

1994-1995 catch of clupeids in the Iranian Caspian 

was 98.3 tonnes by beach seine and 671.5 tonnes by 

gill nets, a decrease of 200 tonnes in total over the 

previous year's catch (Iranian Fisheries Research and 

Training Organization Newsletter 10: 4-5, 1995) (but 

see below where the catch is much higher – sources 

tend to vary). The Caspian Sea shads accounted for 

about 35% of total inland production in Iran which 

was 117,300 tonnes in 1995 (Bartley and Rana, 

1998). These fish are used in a high value form as 

frozen whole consumer packs, as fish meal for 

poultry and in aquaculture, and in canning (Food and 

Agriculture Organization, Fisheries Department 

1996).  

Catches in the Caspian Sea for 1991 and 1992 

were respectively 13,817 and 21,527 tonnes of kilka 

(Clupeonella spp.) compared with 3,036 tonnes and 
2,692 tonnes of sturgeons and 18,571 and 16,873 

tonnes of bony fishes. The catch reached 51,000 

tonnes in 1994 from none 10 years previously (Food 

and Agriculture Organization, Fisheries Department, 

1996). The kilka catch in Gilan Province in 1997 was 

36,077 tonnes (total catch 39,154 tonnes for the 

province) and in Mazandaran in 1998 was 31,583 

tonnes (total catch 48,027 tonnes). 

The Caspian Environment Programme (1998) 

gives the following catches in tonnes for the Iranian 

Caspian Sea for kilka (Clupeonella spp.) and for 
Alosa pontica (= kessleri), respectively as 1,013 and 
2 (1973), 1,170 and 2 (1974), 1,286 and 4.5 (1975), 

900 and 5.5 (1976), 1,261 and 2 (1977), 771 and 0 

(1978), 836 and 0 (1979), 619 and 0.1 (1980), 1,341 

and 0.4 (1981), 798 and 10.4 (1982), 621 and 1.6 

(1983), 1,517 and 20.3 (1984), 1,828 and 34.8 

(1985), 2,450 and 71.9 (1986), 4,389 and 13 (1987), 

4,700 and 16 (1988), 7,902 and 30 (1989), 8,814 and 

30 (1990), 13,817 and 35 (1991), 21,527 and 35 

(1992), 28,730 and 893 (1993), 51,000 and 720 

(1994), 41,000 and 490 (1995) and 57,000 and 330 

(1996). Abdolmalaki and Psuty (2007) give figures 

over a wide range of years for Iranian coastal catches 

in the southern Caspian Sea with selected 

comparative species as presented in Table 1. 

Overfishing of clupeids in the Caspian Sea and 

stock collapses caused by the invasive ctenophore 

Mnemiopsis leidyi have also had deleterious effects 
on the Caspian seal (Pusa caspica) which fed on 
these fishes (Harkonen et al., 2012). 

A major source for the biology and systematics of 

Caspian clupeids remains Svetovidov (1952), now 

inevitably dated but not yet updated. Whitehead 

(1985), Hoestlandt (1991), Coad (1997) and Coad et 



130 
 

Coad/ Herrings of Iran 

al. (2003) gave brief overviews of Iranian herrings. 

There has been no recent, careful systematic and 

taxonomic study of these species in the Caspian Sea 

basin as a whole and extensive new material was not 

available for examination here.  

Key to Clupeidae: Caspian Sea species have 

numerous nominal subspecies and keys to these may 

be found in Berg (1948-1949) and Svetovidov 

(1952).  

1a. Upper jaw without a median notch, rounded 

when viewed from in front; last two anal fin rays 

enlarged; lower jaw articulation with skull below or 

anterior to posterior eye margin; Caspian Sea 

species; Clupeonella spp. …………………………2 
1b. Upper jaw with a median notch; last two anal fin 

rays not enlarged; lower jaw articulation with skull 

behind posterior eye margin; Caspian Sea and 

Persian Gulf; Alosa and Tenualosa spp. ………….4  
2a. Pectoral fins rounded at tips; head large and 

narrow (interorbital width 15.5% or less of head 

length)…………………………Clupeonella grimmi 
2b. Pectoral fins pointed at tips; head short and wide 

(interorbital width 16% or more of head length)…..3 

3a. Body and belly compressed (body depth about 

21-27% of standard length); keeled belly scales 

evident …………………………Clupeonella caspia 
3b. Body cylindrical and belly rounded (body depth 

16-19% of standard length); keeled belly scales 

weakly developed……...Clupeonella engrauliformis 
4a. Branched pelvic fin rays 7; upper gill rakers not 

overlapping lower gill rakers at angle of first arch; 

Persian Gulf basin………………… Tenualosa ilisha 
4b. Branched pelvic fin rays 8; upper gill rakers 

overlap lower gill rakers at angle of first arch; 

Caspian Sea species; Alosa spp. ………………… 5 
5a. Body deep and compressed; head large and deep, 

wedge-shaped in anterior view; caudal peduncle 

short; pectoral fins long……………………………6 

5b. Body not deep and not compressed; head not 

large and deep, not wedge-shaped in anterior view; 

caudal peduncle not short; pectoral fins short…….8  

6a. Gill rakers on first arch 50 or more, thin and long, 

much longer than gill filaments; teeth weakly 

developed…...…………………………Alosa caspia 
6b. Gill rakers on first arch 48 or less, shorter, equal 

to or somewhat longer than gill filaments; teeth well 

developed …………………………………………7  

7a. Upper and lower profiles of head straight; lower 

jaw protruding and its upper edge straight 

………………………………Alosa saposchnikowii 
7b. Upper and lower profiles of head rounded; jaws 

equal in length and lower jaw has a crescentic upper 

edge………………………… Alosa sphaerocephala 
8a. Gill rakers 49 or less, thick and 

coarse………………………… Alosa braschnikowii 
8b. Gill rakers 57 or more, may be thin and long but 

can be coarse and short……………… Alosa kessleri 
 

Genus Alosa Linck, 1790 
The Caspian Sea species of Alosa were formerly 
placed in the genus Caspialosa Berg, 1915. 
Svetovidov (1952) synonymised Caspialosa with 
Alosa. There are 5 species in Iranian waters and the 
Caspian Sea as a whole but numerous subspecies 

have been described. Alosa species are also found in 
the Black Sea, Mediterranean Sea and Atlantic 

Ocean.  

Often distinguished by gill raker counts which in 

any case overlap, the various subspecies are difficult 

to identify. Morphometric characters are of little help 

and Zamakhaev (1944) points out that some named 

taxa are merely different age groups.  

Alosa species are distinguished from sympatric 
Clupeonella species by larger size (up to 75 cm total 

Table 1. Iranian coastal catches in the southern Caspian Sea with selected comparative species (Abdolmalaki and Psuty, 2007) 

Catch and frequency 
1927-

1936 

1937-

1946 

1947-

1956 

1957-

1966 

1967-

1976 

1977-

1986 

1987-

1996 

1997-

2003 

Total recorded catch (t) 8,959 7,224 4,986 3,262 5,547 5,384 16,903 16,201 

Sturgeon meat + caviar (%) 13.4 8.8 16.3 50.9 40.9 34.2 9.4 5.0 

Rutilus kutum (%) 12.2 43.0 24.9 25.8 17.8 19.8 53.2 45.4 

Rutilus rutilus (%) 20.7 25.5 18.8 0.7 0.8 2.3 5.8 6.1 

Alosa spp. (%) 1.9 6.2 14.7 2.9 0.3 0.2 3.2 3.9 

 



131 
 

Int. J. Aquat. Biol. (2017) 5(3): 128-192 

 

length compared to 20 cm), a large mouth, a black 

spot on the flank behind the operculum and 

sometimes a row of such spots, an elongate scale or 

ala at the upper and lower base of the caudal fin, a 

notch at the mid-line of the upper jaw and by the last 

two anal fin rays not being elongated.  

Caspian Sea species have a laterally compressed 

belly with 29-36 spiny scutes running from the throat 

to the anal fin, the dorsal fin origin is closer to the 

snout tip than the caudal fin base, the dorsal fin lies 

in a groove formed by enlarged scales, scales are 

easily detached, the pelvic fin origin lies below or 

slightly posterior to the dorsal fin origin, teeth are 

usually present on the jaws, roof of the mouth (on the 

palatine bone and always on the vomer bone), and on 

the tongue, the opercular bone is distinctly striated, 

eggs are demersal, semi-pelagic, and lack an oil 

globule, gill rakers highly variable in shape and 

number (18-180), dorsal fin branched rays 11-16, 

anal fin branched rays 10-21, scales in lateral series 

49-60, and vertebrae 43-55.  

Afraei Bandpyi et al. (2004) examined 336 

specimens Alosa species from Mazandaran and 
Golestan provinces and found the distinguishing 

characters presented in Table 2. In addition, Hosseini 

et al. (2011) examined 132 specimens of Alosa 
species from Gilan Province and found the 

distinguishing characters presented in Table 3. The 

two characters used in combination distinguish the 

species. 

The general Farsi name for these fishes is shag 

mahi or zalun (both in Gilaki). Alosa volgensis 

(Berg, 1913) was recorded from Iranian waters by 

Kottelat and Freyhof (2007) but presence needs 

confirmation by specimens. Alosa curensis 
(Suvorov, 1907) is poorly known and described from 

Kyzylagach Bay of Azerbaijan. It may eventually be 

recorded from Iranian waters. These herrings 

migrate from the north Caspian Sea to overwinter in 

the central and southern parts, returning north in the 

spring. 

 

Alosa braschnikowii (Borodin, 1904) 
(Fig. 1) 

Common names: Shagmahi, shagmahi-ye Khazari.  
[Dolkii siyanayn, Agraxan siyanayi, Sara siyanayi, 

irikoz siyanak, hasangulu siyanayi, agbas siyanak, 

all in Azerbaijan; Caspian marine shad, Kurinskaya 

sel'd or Kura herring, poloschataya sel'd or striped 

herring, Agrakhanskaya sel'd or Agrakhan herring, 

bol'sheglazaya sel'd or bigeye herring, dolginskaya 

sel'd or dolginka herring, belogolovaya sel'd or 

whitehead herring, Astrabadskaya sel'd or Astrabad 

herring, sel'd-gonets or driver, zheltospinka or 

yellow-back, Gasankulinskaya sel'd or Gasan-Kuli 

herring, kiselevichevskaya sel'd or Kiselevitch 

herring, Krasnovodskaya sel'd or Krasnovodsk 

herring, vostochnaya sel'd or eastern herring, 

obzhorka or glutton, Sarinskaya sel'd or Sara herring, 

maiskaya sel'd or May herring, Brashnikovskaya 

sel'd or Brashhnikov's shad, all in Russian]. 

Systematics: Originally described as Clupea caspio-
pontica var. Braschnikowii. Reshetnikov et al. 
(1997) revert to the original double "i" ending to the 

Table 2. Distinguishing characters of Alosa species from Mazandaran and Golestan provinces, southern Caspian Sea (Afraei Bandpyi et al., 
2004). 

Species Gill rakers Ratio of eye diameter to total length (%) 

A. braschnikowii 20-40, mean 30.93 2.9-5.82, mean 4.72 
A. caspia 110-125, mean 118.3 5.73-7.46, mean 6.21 
A. pontica (= kessleri) 60-73, mean 66.82 4.27-6.48, mean 5.46 
A. saposchnikowii 20-48, mean 32.83 6.0-9.33, mean 7.3 

 
Table 3. Distinguishing characters of Alosa species from Gilan Province, southern Caspian Sea (Hosseini et al., 20114). 

Species Gill rakers Ratio of eye diameter to head length (%) 

A. braschnikowii 22-49, mean 36.6 15.91-21.88, mean 18.53 
A. caspia 74-128, mean 106.3 19.36-24.3, mean 22.48 
A. kessleri 49-73, mean 58.5 17.75-22.4, mean 20.88 

 



132 
 

Coad/ Herrings of Iran 

specific name although the Catalog of Fishes 

(downloaded 13 January 2017) has a single “i” 

ending. A lectotype from Fort Shevchenko 

(Aleksandrovsk) is in the Zoological Institute, St. 

Petersburg (ZISP 13051) and paralectotypes were 

designated by Svetovidov (1952) (ZISP 13051). 

Clupea caspio-pontica is an unneeded new name 
according to Eschmeyer et al. (1996).   

Alosa braschnikowii was regarded as a subspecies 
of Alosa caspia by some authors. Clupeonella 
leucocephala Berg, 1913 from Sumgait and 
Gyurgenchai, Azerbaijan is a synonym (as 

Caspialosa brashnikovi leucocephalia (sic). it is 
listed as a synonym of C. b. grimmi in Mikhailovsky 
(1941)), as is Caspialosa caspia nigra Kisselevitsh, 
1923 from the Caspian Sea opposite Dzambai (the 

material also included specimens of Alosa 
saposchnikowii) (Whitehead 1985, Eschmeyer et al. 
1996).  

Alosa braschnikowii has several subspecies in the 
Caspian Sea, namely agrachanica (Mikhailovsky, 
1941) (author also spelt Mikhaylovsky or 

Mikhailovskaya; dated 1940 in Eschmeyer et al. 

(1996) but 1941 in the online Catalog of Fishes and 

1941 on the paper itself and in Svetovidov (1952) 

and Berg (1948-1949); species also spelt 

agrakhanika in Berg (1948-1949); Caspialosa 
brashnikovi morpha elata is a synonym according to 
Mikhailovsky (1941)), the Agrakhan herring; 

autumnalis (Berg, 1915), the bigeye herring; 
braschnikowii (Borodin, 1904) (also spelt 
brashnikovi in Svetovidov (1952) and Berg (1948-

1949)), the dolginka herring; grimmi (Borodin, 
1904), the whitehead or Astrabad herring, driver or 

yellow-back; kisselevitschi (Bulgakov, 1926) (spelt 
kisselevitschi on the plate in Bulgakov (1926), 
kisselevitschi in Mikhailovsky (1941), kisselevitshi 
in Svetovidov (1952) and Whitehead (1985) and 

kisselewitschi in Berg (1948-1949)), the Gasan-Kuli 
or Kiselevitch herring; nirchi (Morosov, 1928) 
(author also spelt Morosow in Mikhailovsky (1941) 

and Morozov in Eschmeyer et al. (1996)) (with 

Caspialosa brashnikovi kenderlensis Budamshin, 
1938 from Kendyrli Bay as a synonym in Svetovidov 

(1952) and Berg (1948-1949)), the Krasnovodsk 

herring; orientalis (Mikhailovsky, 1941), the eastern 
herring or glutton; and sarensis (Mikhailovsky, 
1941), the Sara or May herring. Caspialosa 
brashnikovi derzhavini Tarasevich, 1946 described 
from the Caspian Sea near the Apsheron Peninsula, 

Azerbaijan may be another subspecies. Caspialosa 
kiselevitschi morpha elata Morozov, 1928 from the 
Caspian Sea, Krasnovodsk Bay, Turkmenistan is an 

infrasubspecific taxon and its availability and 

validity as a taxon have not been examined 

(Eschmeyer et al., 1996). 

This high number of subspecies is an indication 

of the populational variation of this shad and not all 

subspecies may be valid. A modern revision is 

required to assess this problem. In light of this 

uncertainty and the lack of adequate sample sizes to 

determine which of the subspecies occurs in Iranian 

waters or which taxa are valid, reference is made 

here mostly to the species level. Additionally, it 

Figure 1. Line drawing of Alosa braschnikowii. 



133 
 

Int. J. Aquat. Biol. (2017) 5(3): 128-192 

 
should be noted that hybrids between the various 

subspecies, and between this species and other 

species, do occur to complicate matters even further. 

Jaafari et al. (2013) compared fish from Anzali 

and Sari morphologically and found standard length, 

postorbital length and anal fin base length were 

significantly different. Jafari et al. (2014) used 

microsatellite data to study fish from Golestan 

Province between Gomishan and Miankaleh. Low 

genetic differentiation between the two populations 

could be due to natural migration, genetic diversity 

within populations was only 1% and among 

populations was 99%, and it was considered possible 

that a genetic bottleneck could arise in the future. 

Paknejad et al. (2014) examined morphological 

variation between 6 localities along the whole 

Iranian coast and found populations to be distinct, 

particularly the Tonekabon region. Head shape, eye 

diameter and predorsal, prepelvic and preanal 

lengths were the main characters. Heidari et al. 

(2015a) examined fish from the Miankaleh Gulf for 

morphometric and meristic characters and suggest, 

in this brief abstract, that morphological variations 

should be taken into account in fisheries 

management and stock enhancement. 

The neotype of Caspialosa brashnikovi 
agrachanica was designated by Svetovidov (1952) as 
the specimen described by Berg as Caspialosa 
brashnikovi m. elata taken in front of the Sulak River 
mouth, Agrakhan Bay and housed in the Zoological 

Institute, St. Petersburg under ZISP 7334. However, 

this neotype was not validly designated because 

qualifying conditions 75.3.1, 75.3.4 and 75.3.5 of the 

International Code on Zoological Nomenclature 

were not met (N.G. Bogutskaya, pers. comm. 23 

January 2013). This also applies to the following 6 

taxa. 

The neotype of Caspialosa braschnikovi 
autumnalis was designated by Svetovidov (1952) as 
a specimen 26.9 cm long from the eastern shore of 

the Caspian Sea at Gasan-Kuli (just north of the 

Iranian border in Turkmenistan) caught on 8 April 

1948 and housed under ZISP 31749.  

The neotype of Caspialosa kisselevitschi is also 

from Gasan-Kuli caught on 30 June-1 July 1926 and 

was housed in the Faculty of Zoology, Central-Asian 

State University (Sredne-Aziatskogo Gosudarstven-

nogo Universiteta), Tashkent. 

The neotype of Clupea caspio-pontica var. 
grimmi was designated by Svetovidov (1952) as a 
specimen 34.0 cm long found at Ashur-ade (= 

Ashuradeh) near Astrabad Bay (= Gorgan Bay or 

Khalij-e Gorgan) on 23 April 1903 is under ZISP 

13045.  

The neotype of Caspialosa nirchi as designated by 
Svetovidov (1952) is from the southern part of the 

Caspian Sea opposite North Cheleken Spit and is 

under ZISP 31780.  

The neotype of Caspialosa brashnikovi orientalis 
as designated by Svetovidov (1952) is from the 

southern part of the Caspian Sea opposite Kara-

Ashly and is under ZISP 32187.  

The neotype of Caspialosa brashnikovi sarensis 
from Sara Island is under ZISP 32184 as established 

by Svetovidov (1952).  

Key characters: Characterised by a relatively 

elongate and rounded body likened to a "herring" 

shape, not as deep as in some related species which 

are likened to a "shad" shape. Total gill rakers 18-49 

and short (about equal to gill filaments in length, 

sometimes shorter). Teeth are well-developed in 

both jaws. 

Morphology: Dorsal fin with 3-5 unbranched and 

12-15, mostly 14, branched rays, anal fin with 2-4, 

usually 3, unbranched rays and 10-20, mostly 18, 

branched rays. Scales in lateral series 51-54. Teeth 

are well-developed on the jaws, tongue and roof of 

the mouth. Ghotbi Jokandan et al. (2014) briefly 

distinguish this species from A. saposchnikowii on 
morphometric characters in Gilan. The distance 

between the posterior end of the dorsal fin to the 

anterior caudal peduncle, distance between origins of 

pelvic and anal fins, head length, pectoral fin height 

and ratio of anal fin height to standard length were 

greater than in A. saposchnikowii. Alavi-Yeganeh 
and Razavi (2016a, 2016b) found 26-47 gill rakers in 

50 specimens 15-45 cm long from Iran and number 

increased with total length, although only fish 15-20 



134 
 

Coad/ Herrings of Iran 

cm and 40-45 cm long had significantly different 

counts. Gill raker length, length/width ratio and ratio 

of raker to filament length decreased with increase 

of total length. Equal lengths of rakers and filaments 

were found in fish greater than 25 cm. Evidently, 

some Alosa species may overlap in counts depending 
on size.  

The accompanying table (Table 4) summarises 

characters of the subspecies and is taken from 

Svetovidov (1952) and Mikhailovsky (1941) but 

identification to subspecies should be done with the 

keys from these works. Some of the characters used 

in the keys are not in the table as they do show 

individual variation and are difficult to summarise. 

An example is the nature of the gill raker (thin, thick, 

blunt, pointed, bent, straight, curved, branched, 

broken off, forked, swollen at the tip, etc.); another 

is the degree of protrusion of the lower jaw.  

The subspecies grimmi is quite specialised in 
association with its benthic mode of life, feeding 

mostly on gobies (Gobiidae). It has a unique 

character in the well-developed callus on the tip of 

the lower jaw which adults acquire from rubbing the 

jaw on the sea bed while feeding, gill rakers are low 

in number as fine food is not taken, and the tips are 

broken off, broadened, and split owing to abrasion, 

and the rakers on the lower arch are reduced in 

number so the first raker is far from the tongue base. 

The subspecies nirchi is similar. In contrast, the 
subspecies kisselevitschi has a high gill raker count, 
rakers are pointed and not split at the tips, and the 

first raker is close to the tongue base. This species 

lives in surface waters feeding on Clupeonella, 
Atherina, shrimps, gammarids, and gobies 

(Gobiidae). 

Sexual dimorphism: None reported. 
Colour: The back and top of the head are dark with a 
green or blue tint and may be grey-green. Some 

subspecies are paler in colour with a grey or grass-

green back and pale flanks, nirchi has a whitish blue-
green head, light grey back with a slight greenish 

tint, and lower jaw and pectoral fins light, while 

grimmi is also quite pale with a grey-blue back and 
top of the head and whitish anterior head and 

pectoral fins. There is a dark spot behind the 

operculum but no series of spots along the flank in 

most subspecies, except in rare cases when there may 

be up to 7, occasionally 12-13. The subspecies 

grimmi regularly has a row of diffused, grey spots 
almost merging into a stripe, and nirchi occasionally. 
Pectoral fins are dark on some subspecies 

(braschnikowii, sarensis, and kisselevitschi), pale or 
whitish on the others, although there is confusion in 

the literature over this, perhaps indicative of 

individual variation (cf. sarensis in Mikhailovsky 
(1941)). The back and upper part of the head may 

become a deep black at spawning. The flanks and 

belly are silvery. 

Size: Attains 50.0 cm standard length but average 
lengths are about 27-34 cm. 

Distribution: All the Caspian subspecies are found 

widely distributed in the sea but chiefly in the south 

in winter, moving north to spawn in spring. The 

subspecies sarensis is reported from the Lenkoran 
coast and from southwest of Gasan-Kuli (in 

Turkmenistan just north of the Iranian border), the 

subspecies orientalis from Gorgan or Astrabad Bay, 
autumnalis from coastal waters at Gasan-Kuli, 

Character / Subspecies 
Gill rakers 

(mostly) 

Pectoral fins as % 

body length 

Head as % of 

body length 
Vertebrae 

agrachanica 20-46 (28-33) 13.1-15.6 22.6-25.2 47-54 

autumnalis 21-37 (28-30) 16.4-19.9 26.0-29.2 45-53 

braschnikowii 24-47 (30-33) 14.3-16.7 23.5-26.6 48-55 

grimmi 18-28 (20-22) 12.9-15.2 22.9-26.4 45-52 

kisselevitschi 29-49 (36-40) 13.9-16.8 24.2-26.9 43-53 

nirchi 20-31 (23-26) 10.9-14.7 23.4-26.3 48-52 

orientalis 20-35 (27-32) 13.5-18.0 25.0-27.8 45-53 

sarensis 20-33 (24-27) 14.1-16.2 23.8-26.6 45-53 

 

Table 4. Characters of the subspecies in Alosa braschnikowii based on Svetovidov (1952) and Mikhailovsky (1941). 



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Int. J. Aquat. Biol. (2017) 5(3): 128-192 

 
kisselevitschi from Astara and Gasan-Kuli, and 
grimmi from Astara and Gorgan Bay. Esmaeili et al. 
(2014) record it from the Anzali Wetland or Talab. 

Zoogeography: This species is endemic to the 

Caspian Sea. 

Habitat: In winter, this species moves into deeper 

water towards the Iranian coast. In March, it 

approaches coastal waters (Vetchanin, 1984) 

including brackish waters but does not enter fresh 

water. It never enters rivers in the south of the 

Caspian Sea (Jolodar and Abdoli, 2004). Salinities 

up to 47.6‰ are survived by this species. Spawning 

and feeding grounds are in the north Caspian for 

some populations but others live permanently in the 

south Caspian Sea and are of smaller size. The 

subspecies kisselevitschi, for example, lives off 
Gasan-Kuli in winter at depths below 25 m, not 

migrating or feeding. In March, they move north to 

feed and then return south to spawn but live almost 

entirely as a pelagic species in the southern Caspian 

Sea. Knipovich (1921) reports this species from 

depths of 80-98 m in Iranian waters. The density of 

this species increased from east to west in a 1999-

2001 study in Iranian waters (Afraei et al., 2006). 

Abdoli and Naderi (2009) list it as from the 

southwest, southeast and south-central Caspian Sea 

in Iranian waters. 

Age and growth: Maturity is attained at age 2-5 and 

life span is up to 10 years, although this varies with 

the subspecies. Most south Caspian forms apparently 

mature at age 2 according to Svetovidov (1952). 

Growth rates also vary between subspecies, 

orientalis being one of the slowest growing herrings 
in the Caspian Sea and reaching 10 years of age. The 

catch near Astara of sarensis, for example, is mainly 
4-5 year olds but this too varies with the subspecies 

and also with the year-class strength. Vetchanin 

(1992) reported on grimmi catches from the 
southeastern Caspian where the average length was 

27.8-28.6 cm and the average weight 294-313 g. 

There is a tendency for length and weight to fall in 

catches as the summer progresses, from April to 

July. Length and weight are less in southern, 

compared to northern waters. Afraei et al. (2006) 

found this species to be the largest Alosa in Iranian 
waters on average at 395 mm and 760.3 g. Males 

predominate at 55.8% in Mazandaran and 69.4% in 

Golestan catches. Six age classes were present (1+ to 

6+) with the 2+ class being the most common at 

28.9% and 6+ the rarest at 8.9%. Alavi-Yeganeh et 

al. (2013) found Mazandaran fish were 1-6 years old 

and had a b value of 3.693 showing positive 
allometric growth. Ghotbi-Jokandan et al. (2015) 

examined 147 fish from Anzali, Noor and Torkeman 

and found a b value of 3.1. Mousavi-Sabet et al. 
(2016) gave a b value of 3.24 for Iranian fish from 6 
fishing areas from Astara to Miankaleh. 

Food: Diet in the southeastern Caspian Sea in winter 

comprises 85% Clupeonella engrauliformis with 
some gobies (Neogobius) and shrimps (Vetchanin, 
1984). From March to November the diet is 

dominated by Clupeonella caspia, Atherina boyeri (= 
caspia) and shrimps. Juvenile Liza saliens, 
Syngnathus caspius, molluscs, crabs and higher 
aquatic plants are also recorded along with foreign 

objects such as rice husks, pieces of wood, foil, 

polyethylene, etc. This species is a cannibal. The 

more southerly populations examined favour 

Atherina boyeri (= caspia) and Neogobius species 
and some of these populations favour benthic 

invertebrates. The subspecies grimmi is the most 
benthic one and takes primarily gobies with some 

molluscs as well as Clupeonella. Feeding intensity 
rises sharply after spawning. While some herrings, 

like Alosa pontica (= kessleri), feed poorly on their 
migration, this species feeds intensively on its spring 

migration. Azizov et al. (2015) examined fish from 

the western Caspian Sea north of Iranian Astara in 

Azerbaijan waters and found the main foods were 

kilkas, gobies and shrimps with some zoobenthos. 

Intensive feeding took place in spring (March-April) 

before spawning and in July after spawning. 

The feeding regime altered after the invasion of 

the ctenophore, Mnemiopsis leidyi. A shift was 
observed from 85% Clupeonella engrauliformis to 
65% Atherina boyeri (= caspia). Other fishes were 
also eaten including Clupeonella grimmi, C. caspia, 
Cyprinus carpio, Liza saliens, as well as Palaemon 



136 
 

Coad/ Herrings of Iran 

spp. (Iranian Fisheries Research Organization 

Newsletter 49: 2, 2006). Afraei Bandpei et al. (2012) 

examined 357 fish from beach seines in the south-

central Caspian Sea and found the diet was 

dominated by Atherina boyeri (= caspia) at 58% with 
Gobiidae (26%), Clupeidae (11%), Cyprinidae (3%) 

and Mugilidae (1%). Atherina boyeri (= caspia) was 
the most frequent prey at 78.6% index of relative 

importance, followed by Neogobius fluviatilis (= 
pallasii) at 18.2%. A wider variety of items was fed 
on in November, December and February with 

lowest feeding in October and January. There were 

also differences in diet related to fish size. Increases 

in the gastrosomatic index in February and March 

coincide with the spawning season and energy 

preservation for gonadal development. 

Reproduction: Vetchanin (1984) reports spawning of 

this species in the southeastern Caspian Sea north of 

Iranian waters to begin in early May, continuing to 

July as it is intermittent. The subspecies sarensis 
spawns along the Lenkoran coast from mid-April to 

the end of June. The subspecies orientalis spawns in 
Gorgan Bay from the end of March to the beginning 

of April, spawning schools forming at 17-18°C or 

higher. The subspecies autumnalis spawns at the 
same time off Gasan-Kuli near the Iranian border 

with Turkmenistan. The subspecies grimmi spawns 
in May-June in Gorgan Bay. The subspecies 

kisselevitschi has the latest spawning date, June to 
July and even in August off Gasan-Kuli when 

temperatures exceed 25°C. Spawning takes place in 

shallow water (1.8-5.8 m) in the sea over sand or silt 

bottoms at 15-18°C (some subspecies and 

populations at 20-22°C, others beginning as low as 

11°C), and a salinity of 8-13‰. Fecundity is up to 

178,400 eggs, average 66,000 per fish. There is no 

feeding while spawning. Early maturers, like the 

south Caspian populations, can reproduce up to 7 

times in their life. 

Parasites and predators: The Caspian seal, Pusa 
caspica, is a predator on this species (Krylov, 1984). 
Economic importance: The catch for all species of 

"Caspialosa" in Iran varied between 5,337 kg and 
419,518 kg for the years 1956/1957 to 1961/1962 

(Vladykov, 1964). In the Anzali region, the catch for 

the years 1933/1934 to 1961/1962 varied from 1,553 

kg to 539,710 kg (Vladykov, 1964). The catch has 

been as high as 126,900 centners or 12,690 t in the 

sea as a whole for the type subspecies alone (1 

centner=100 kg (Svetovidov, 1952)), taken chiefly in 

spring. Other subspecies were not fished for as 

extensively although kisselevitschi was the most 
numerous of the south Caspian forms of Alosa 
braschnikowii, forming 70% of the drift net catch. 
Conservation: Reputedly depleted in Iranian waters. 

Kiabi et al. (1999) consider this species to be data 

deficient in the south Caspian Sea basin according to 

IUCN criteria. Criteria include medium numbers, 

medium range (25-75% of water bodies), absent in 

other water bodies in Iran, and present outside the 

Caspian Sea basin. Afraei Bandpei et al. (2012) 

noted that fishing is forbidden in the southern 

Caspian Sea from May to September.  

Sources: Iranian material: CMNFI 1970-0581, 5, 

226.0-245.0 mm standard length, Gilan, Caspian Sea 

near Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 

1979-0431, 1, 297.2 mm standard length, 

Mazandaran, bazaar at Now Shahr (no other locality 

data); CMNFI 1980-0126, 1, 245.8 mm standard 

length Gilan, Caspian Sea near Bandar-e Anzali 

(37º28'N, 49º27'E); CMNFI 1980-0150, 1, 222.4 

mm standard length, Gilan, Safid River estuary 

(37º24'N, 49º58'E).  

Comparative material: BM(NH) 1938.8.2:1, 1, 245.9 

mm standard length, Kazakhstan, Caspian Sea, 

Kaidak Bay (no other locality data); BM(NH) 

1938.8.2:2, 1, ca. 337.5 mm standard length, 

Kazakhstan, Caspian Sea, Kaidak Bay (no other 

locality data); BM(NH) 1939.2.21:17-18, 2, 285.0-

305.2 mm standard length, Caspian Sea (no other 

locality data); BM(NH) 1939.2.21:19-20, 2, 222.9-

273.4 mm standard length, Caspian Sea (no other 

locality data). 

 

Alosa caspia (Eichwald, 1838) 
(Figs. 2, 3) 

Common names: Shagmahi-ye shekambozorg (= big 
belly herring fish), shagmahi-ye chekameh dar, 



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Int. J. Aquat. Biol. (2017) 5(3): 128-192 

 

shagmahi-ye darya-ye khazar (= Caspian Sea herring 

fish), shah mahi (= king fish), zalun (in Gilaki), 

puzanok. [Xazar sisgarini, sara sisgarini in 

Azerbaijan; Kaspiiskii puzanok or Caspian shad, 

severokaspiiskii puzanok or North Caspian shad, 

srednekaspiiskii puzanok or Central Caspian shad, 

il'mennyi puzanok or il'men shad, Enzeliiskii 

puzanok or Enzeli (= Anzali) shad, Sarinskii 

puzanok or Sara shad, Bakinskii puzanok or Baku 

shad, Astrabadskii puzanok or Astrabad shad, all in 

Russian]. 

Systematics: Clupea caspia was originally described 
in Latin from "Hab. in Caspio mari, meridiem 

versus" (Caspian Sea, towards the south). Alosa 
caspia has 3 subspecies in the Caspian Sea basin, 
namely caspia (Eichwald, 1838) (= North Caspian, 
Central Caspian, Caspian or il'men shad); 

knipowitschi (Iljin, 1927) with natio knipowitschi 
(Iljin, 1927) (= Enzeli or Anzali shad) and natio 

saraica (Berg, 1948) (= Sara or Baku shad); and 
persica (Iljin, 1927) (= Astrabad shad). The 
differences between natio knipowitschi and natio 
saraica are small (e.g. gill rakers 122-166 versus 

140-150, both averaging 145; vertebrae 43-49 versus 

45-51, both with mostly 47 or 48; growth differences 

are known, the former grows faster in the first 2 years 

of life but the latter reaches a greater size) and they 

probably have no taxonomic significance being 

simply separate breeding populations. The 

differences between Alosa caspia caspia natio caspia 
(the North or Central Caspian shad) and natio 

aestuarina Berg, 1932 (the il'men shad) were found 
to be based on geography and growth rate 

(Svetovidov, 1952). 

Alosa rossica Kessler, 1870 described from the 
Volga River is a nomen nudum and is this species. 
Other taxa now considered as synonyms of Alosa 
caspia are Caspialosa caspia salina Svetovidov, 1936 
from Mertvyi Kultuk and Kaidak bays in the 

northeast Caspian Sea and Caspialosa caspia 
kaidakensis Kazancheev, 1936 (spelt kajdakensis in 
Svetovidov (1952)) from Kaidak, the latter being in 

any case a synonym of the former subspecies. 

C. caspia m. elongata Berg, 1913 is also a synonym. 
Alosa caspica Yakovlev, 1871 is presumably a 
misspelling. 

Figure 2. Line drawing of Alosa caspia. 

Figure 3. Alosa caspia courtesy of A. Abdoli. 



138 
 

Coad/ Herrings of Iran 

Knipovich (1921) records a species, Caspialosa 
enzeliensis Iljin, from the southern Caspian Sea 
which he places as a subspecies of caspia. I have 
been unable to locate the original description of this 

taxon, which presumably is found in the Anzali 

Mordab (= Talab) of Iran. It is probably an unused 

manuscript name for what Iljin later described as 

knipowitschi. As of 15 July 2007, this scientific 
name returned no hits on Google and as of 12 

December 2016 the only hit was for 

www.briancoad.com.  

The lectotype of Caspialosa knipowitschi is a 
specimen 21.2 cm long from Anzali in Iran caught 

on 15 April 1915 and housed in the Zoological 

Institute, St. Petersburg (ZISP 31892). The lectotype 

of Caspialosa caspia var. persica is a specimen 147.5 
mm long from the Caspian Sea Bay of Asterabad (= 

Gorgan Bay or Khalij-e Gorgan) north of Ashur-ade 

(= Ashuradeh) at 36°53'N, 53°55'E caught on 25 

April 1904 on the Caspian Expedition of 1904 and 

housed in the Zoological Institute, St. Petersburg 

(ZISP 16413). The lectotype of Caspialosa caspia 
knipowitschi n. saraica is from near Sara Island and 
is under ZISP 32183. The lectotype of Caspialosa 
caspia salina is from Mertvyi Kultuk Bay, 10 km 
west of Cape Kizil-kair and is under ZISP 25813. 

These taxa were designated by Svetovidov (1952) as 

none were before or material was not preserved. 

Key characters: Characterised by a relatively deep 

and compressed body likened to a "shad" shape, not 

as elongate and rounded as in some related species 

which are likened to a "herring" shape. Total gill 

rakers 50-180, variously reported as thin or thick, 

long (obviously longer than the gill filaments), and 

forming a convex outline on the lower arch. Teeth 

are poorly developed in both jaws.  

Morphology: Dorsal fin with 3-4 unbranched and 

12-15 branched rays, anal fin with 3-4, usually 3, 

unbranched and 15-20 branched rays. Scales in 

lateral series 49-54. The characters distinguishing 

subspecies all overlap widely and are given in Table 

5 after Svetovidov (1952) and Hoestlandt (1991).  

Sexual dimorphism: Females are longer and weigh 

more than males of the same age. 

Colour: The back is blue-green to dark and the flanks 

silvery. There is a black spot on the flank behind the 

upper operculum margin and sometimes up to 7 

spots extending along the upper flank to a level of 

the rear of the dorsal fin. 

Size: Reaches 28.0 cm standard length for caspia, to 
29.6 cm for knipowitschi, and to 33.8 cm for persica. 
Distribution: Found in the Caspian Sea. The 

subspecies caspia is found mostly in the western half 
of the Caspian Sea basin but is the most widely 

distributed subspecies, found throughout almost the 

whole sea. The subspecies knipowitschi is found in 
the south near Anzali, Astara and the Baku 

Archipelago, near the northern shore of the 

Apsheron Peninsula in autumn with a few reaching 

the Gorgan Bay in fall and winter; natio saraica is 
found north of Astara and spawns near Sara Island, 

natio knipowitschi spawns in the Anzali Wetland. 
The subspecies persica is found in the southeast, near 
Gorgan or Astrabad Bay. Holčík and Oláh (1992) 

report persica from the western basin of the Anzali 
Wetland and this species is reported from the Safid 

River and Anzali Wetland as subspecies persica and 
from the Anzali Wetland as knipowitschi (Abbasi et 
al., 1999). Abdoli and Naderi (2009) list it as from 

the southwest, southeast and south-central Caspian 

Sea, the Anzali Wetland and Gorgan Bay in Iranian 

waters for both knipowitschi and persica. Reported 

Characters / 

Subspecies 

Head length as % 

of body length 

Pectoral length as 

% of body length 

Vertebrae 

(mostly) 
Gill rakers (mostly) 

caspia 25.5-28.1 15.5-18.1 45-52 (49-51) 68-150 (100-140) 

knipowitschi 18.3-24.1* 16.0-19.1 43-51 (47-48) 120-180 (130-160) 

persica 25.6-27.1 16.5-17.7 45-51 (47-49) 50-120 (60-90) 
* The numbers cited in Svetovidov (1952: 256 in the English version) and Hoestlandt (1991: 128) in the keys to 

subspecies do not agree with the numbers on p. 148 and p. 265 respectively in the species descriptions.  

Table 5. Characters of the subspecies in Alosa caspia based on Svetovidov (1952) and Hoestlandt (1991). 



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Int. J. Aquat. Biol. (2017) 5(3): 128-192 

 
from the Siahdarvishan River as A. caspia caspia 
(Abbasi et al., 2007). Esmaeili et al. (2014) record 

A. knipowitschi and A. persica as species from the 
Anzali Wetland but these natio are listed as A. caspia 
in the Catalog of Fishes (downloaded 15 December 

2016). 

Zoogeography: This species is a Caspian Sea 

endemic.  

Habitat: The type subspecies prefers open waters. 
Caspian shad winter at depths of 30-40 m or more 

and prefer temperatures not less than 8-11°C. They 

rise to surface waters in spring, moving north along 

the western shore of the Caspian Sea in waters of 

about 9-11°C according to Kushnarenko (1986) 

while Heckman in Hoestlandt (1991) states that this 

shad begins to migrate at the end of March at 5-6°C 

water temperature with a peak at 9-14°C in mid to 

late-April, ending in early May. Males migrate in 

large numbers at the beginning and end of the 

migration, females in the middle (Pushbarnek, 1987) 

while Heckman in Hoestlandt (1991) states that two 

waves of migration occur, one usually in late April 

at 7.6-10.2°C comprised of over 80% males and the 

second in the first half of May at 10.8-14.0°C 

comprised of over 70% females. The young, which 

hatch in the spring, leave the summer feeding 

grounds before the adults and migrate south before 

October-November. Adults follow as temperatures 

fall. Some populations do not migrate north and 

spend their whole life in the southern Caspian Sea. 

This subspecies will enter fresh waters to spawn in 

addition to spawning in the open Caspian Sea. The 

subspecies knipowitschi prefers water warmer than 
that of all other Caspian Sea clupeids except for 

Alosa caspia persica. Its sea movements are not well 
known but spawning fish favour waters with 

freshwater input and some fish enter rivers so it is 

classified as semi-anadromous. This subspecies was 

common in the Anzali Wetland but is now replaced 

by persica (Holčík and Oláh, 1992). The rise in 
Caspian Sea level since 1977 is gradually returning 

the Wetland to its supposed, natural brackish state 

and may improve the fisheries situation which had 

declined over the last 50 years. Emergent and 

submergent aquatic macrophytes were decreasing 

and Alosa caspia were increasing in numbers since 
1989 (Mandych, 1995). However, the fishery will 

require extensive engineering and management 

innovations to recover. 

It is also reported westwards to Astara and 

eastwards to Gorgan Bay. The winter habitat of 

persica is unknown. It is semi-anadromous and 
remains in the southern Caspian Sea near the shore. 

From spring to fall this subspecies moves northward 

along the eastern Caspian shore towards 

Krasnovodsk Bay and westwards to the Anzali 

Talab.  

Age and growth: Pushbarnek (1987) found shad of 

the type subspecies up to 7 years of age on the 

western coast of the middle Caspian Sea. In the 

spawning population, the predominant sizes and 

weights for males were 16-21 cm and 60-130 g and 

for females 18-23 cm and 70-140 g. Males and 

females usually mature at 2-3 years although most 

spawn for the first time at 3 years. Females grow 

faster than males. Shad may spawn up to four times 

as the period of sexual maturation may continue for 

2-5 years. The age composition of the spawning 

population is dependent on year-class strength. First 

spawners constitute 75.9% of 3-year-olds, 41.7% of 

4-year-olds and 23.5% of 5-year-olds. The Caspian 

shad is a slow-growing species compared to 

A. braschnikowii and A. saposchnikowii, its mean 
length being 21.2 cm compared to 32.2 cm and 25.6 

cm for the two other species respectively (Shubina, 

1981).  

Dmitriev (1947) briefly examined the Anzali, Iran 

population and found 6 age groups but life span is 

noted by Heckman in Hoestlandt (1991) to be up to 

9 years. Maturity is attained as early as 2 years 

although most fish appear to mature later as most 

spawners are 4-5 years old. The subspecies persica is 
the slowest growing of the shad species in the 

Caspian Sea, sexually mature fish being 13-21 cm 

long. Some fish become mature at 2 years of age. 

Life span is up to 8 years. The populations of both 

knipowitschi and persica are small compared to 
caspia. Abbasi and Sabkara (2004) studied 180 fish 



140 
 

Coad/ Herrings of Iran 

from the southeast Caspian Sea coast of Iran and 

found fork length to be 103-232 mm, mean 158.8 

mm, weight 16-130 g, mean 52.2 g and age 2-5 years, 

mean 2.64 years. Afraei et al. (2006) found this 

species to be the smallest Alosa in Iranian waters on 
average at 110 mm and 109 g. Patimar et al. (2011) 

found Iranian fish to reach age 5+ years, with positive 

allometric growth in the southeast Caspian, and 

negative allometric growth in the central and 

southwest Caspian. Females had a b value larger 
than males. von Bertalanffy growth functions were 

variable, different between males and females of 

each area and between sexes from different areas. 

The largest L∞ was in the southeast for males and in 

the central area for females. The highest K value was 
for males and the lowest for females in the southeast 

area. t0 was negative in all areas with lowest value 

for females (-0.531) in the southeast and the highest 

for males (-0.145) in the central area. Alavi-Yeganeh 

et al. (2013) found Mazandaran fish were 2-5 years 

old and had a b value of 3.022. Ghotbi-Jokandan et 
al. (2015) examined 113 fish from Anzali, Noor and 

Torkeman and found a b value of 2.98. Mousavi-
Sabet et al. (2016) gave a b value of 3.12 for Iranian 
fish from 6 fishing areas from Astara to Miankaleh. 

Food: The most intensive feeding period occurs after 

reproduction, beginning in June and the highest 

condition factor is found at the end of this summer 

feeding period. Little food is eaten in winter. 

Temperature (affecting metabolic rate) and 

zooplankton biomass (decreases engender 

competition with C. engrauliformis and other 
planktivores) are important factors governing 

catches of this species (Shubina 1981). Food is 

chiefly copepods, more than 70%, with mysids at 

20%, but some phytoplankton and small fishes are 

taken. Food in rivers after spawning is mostly 

cladocerans and other crustaceans. The above refers 

to the type subspecies; food of the other two 

subspecies is assumed to be similar.  

The southeast Caspian Sea fish studied by Abbasi 

and Sabkara (2004) fed on phytoplankton 

(Rhizosolenia and Sprirogyra) at 4.5%, zooplankton 
(Foraminifera, Copepoda, Cirripedia, Bivalvia 

larvae) at 95.0%, and bony fish larvae and eggs at 

0.5%. The presence of the ctenophore, Mnemiopsis 
leidyi, a food competitor reduced the index of 
fullness and fish growth was reduced. Abdollapour 

Bereya et al. (2007) studied diet in fish from beach 

seines and gill nets in Gilan. 98.0% of the stomach 

contents were zooplankton (ostracods, rhizopods, 

cladocerans, rotatarians, copepods, cirripedes, 

mysids, bivalve larvae and bony fish larvae and 

eggs), 1.8% was phytoplankton (notably 

Rhizosolenia and Spirogyra), and 0.2% was benthic 
items (foraminiferans, sponges, cumaceans, 

amphipods, insect larvae and palaemonids). Acartia 
spp. (copepods) at 83.1%, and Balanus (cypris larvae 
of the cirripede) at 12.9% were the most abundant. 

The zooplankton have declined drastically from 

predation by M. leidyi, the invasive ctenophore, and 
the fish have shown a great reduction in the index of 

fullness and in growth recently. 

Reproduction: Most spawning of the type subspecies 

occurs in the north Caspian Sea near the outflow of 

the major rivers, particularly the Volga, and the fish 

overwinter in the south Caspian, migrating between 

the two areas (Shubina, 1981). This subspecies 

spawns successively, 3 times within a week. Some 

fish enter fresh water to spawn. Spawning takes 

place at the favoured water temperature of 13.8-

24.1°C, with mass spawning at 18-22°C, beginning 

as early as late April or as late as mid-May and 

continuing to mid- or late June. Most eggs are 

released in the upper 3m of the water column. 

Fecundity reaches 41,000 eggs. The eggs are 1.11-

1.38mm when ripe but unfertilised and 1.92-2.91 

mm in diameter when fertilised, and are semi-pelagic 

to demersal. The subspecies knipowitschi spawns in 
the Anzali Talab (and probably the Chemkhaleh 

River to the east of the Safid River) in May and June 

after a spring migration from the sea, leaving in the 

fall. Spawning of the subspecies persica takes place 
in Gorgan Bay and Holčík and Oláh (1992) suspect 

from catches of mature and spent fish that it also 

occurs in the Anzali Talab. 

Parasites and predators: Naem et al. (2002) found the 

monogenean trematode Mazocraes alosae on the 



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gills of this species in the western branch of the Safid 

River. Barzegar et al. (2008) record the digenean eye 

parasite Diplostomum spathaceum from this fish. 
Youssefi et al. (2011) report the digenean 

Pronoprymna ventricosa from fish in the Babol 
River. Barzegar et al. (2012) found Mazocraes 
alosae, D. spathaceum, P. ventricosa (digenean), and 
Hysterothylacium sp. (anisakid). 

The Caspian seal, Pusa caspica, is a predator on 
this species (Krylov, 1984) and it forms a substantial 

part of the diet of Silurus glanis in the Anzali Talab 
(Holčík and Oláh, 1992). Ashoori et al. (2012) found 

that grey herons (Ardea cinerea) in the Siahkeshim 
Protected Area of the Anzali Wetland or Talab ate 

this species. 

Economic importance: The type subspecies was the 

most important subspecies in the herring family in 

the Caspian Sea. It was caught off the coasts of 

Dagestan and Azerbaijan for research purposes and 

comprises 85% of the clupeid catch (Pushbarnek, 

1987), and 80-90% of the Caspian commercial catch 

(Kushnarenko, 1986). During the 1970s, it was only 

2% of the total Caspian fishery production. These 

herrings dominated the commercial catch in the 

Caspian Sea until the 1960s when commercial 

fishing was banned except on the western coast of 

the central Caspian. Many young of other 

commercial species were being killed in the herring 

fishery, entangled in the gill nets used. Soviet 

catches have weighed as much as 75,000 t. This fish 

is fattier than other Caspian Clupeidae, except for 

A. kessleri, up to 18.1% of the body weight. The fat 
content decreases on the spring migration. The catch 

of the subspecies knipowitschi is of minor economic 
importance and had been little exploited when 

Svetovidov (1952) summarised biology, as the age 

of captured fish indicated. About 420 tons (sic, 
possibly tonnes) were caught in the Anzali Talab in 

1933 and 1934, but this may be an error in the report 

by Vladykov (1964) according to Holčík and Oláh 

(1992) although Berg (1948-1949) reports 4,200 

centners for the same period. The fishing season in 

the talab began in mid-April and ended in mid-June 

when spent fish appeared. There appears to be no 

fishery data on the subspecies persica in the sea. 
Holčík and Oláh (1992) report catches of persica, 
which replaced knipowitschi, in the Anzali Talab 
from the end of April to the beginning of June but in 

1990 this comprised only 5 kg. It is regarded as of 

inferior quality in Iran. The Caspian shad is the 

dominant fish catch in the Iranian Caspian, 

comprising 51,000t in 1994 rising from nothing a 

decade earlier (Food and Agriculture Organization, 

Fisheries Department, 1996). Robins et al. (1991) list 

this species as important to North Americans. 

Importance is based on its use as food.  

Ershad Langroudi (2004) investigated the 

durability of polycyclic aromatic hydrocarbons 

(PAH), a carcinogen, in cold smoked fish and found 

relatively large amounts in the skin and flesh of this 

fatty shad compared to cyprinids. The PAH density 

declined with storage time. Nemati et al. (2012) 

determined that the protein hydrolysate produced 

from by-products (head, skin, viscera) using the 

enzyme alcalase was the best, with high nutritional 

value and was useful as a protein source in fish food. 

Soleimani et al. (2012) found the amounts of fat, 

protein, ash and dry matter were higher in this 

species than in C. engrauliformis and it was a rich 
source of omega-3 fatty acid. Salting and drying 

methods were compared. 

Najm et al. (2014) determined the concentration 

of the heavy metals cadmium, chromium and lead in 

fish from the Babol Sar coast and found cadmium 

levels exceeded international standards for human 

consumption although a risk assessment concluded 

the fish were safe for consumers. Sadeghi Bajgiran 

et al. (2016) assessed this species for muscle content 

of nickel, vanadium and zinc and found levels not to 

be dangerous to consumers. 

Conservation: The stocks of this species in the 

Anzali Talab are likely to increase if the lagoon 

becomes more saline (Holčík and Oláh 1992). Kiabi 

et al. (1999) consider this species to be of least 

concern in the south Caspian Sea basin according to 

IUCN criteria. Criteria include abundant in numbers, 

widespread range (75% of water bodies), absent in 

other water bodies in Iran, and absent outside the 



142 
 

Coad/ Herrings of Iran 

Caspian Sea basin. 

Sources: Iranian material: CMNFI 1970-0524, 11, 

58.7-88.9 mm standard length, Gilan, Caspian Sea at 

Bandar-e Anzali (37º28'N, 49º27'E); CMNFI 1970-

0532, 1, 113.0 mm standard length, Gilan, Caspian 

Sea near Bandar-e Anzali (37º28'N, 49º27'E); 

CMNFI 1970-0543A, 1, 85.9 mm standard length, 

Gilan, Caspian Sea at Hasan Kiadeh (37º24'N, 

49º58'E); CMNFI 1970-0586, 1, 77.5 mm standard 

length, Mazandaran, Gorgan Talab at Ashuradeh-ye 

Kuchak (36º50'N, 53º56'E); CMNFI 1970-0587, 2, 

107.4-108.6 mm standard length, Mazandaran, 

Babol Sar (36º43'N, 52º39'E); CMNFI 1971-0343, 1, 

95.5mm standard length, Gilan, Langarud at 

Chamkhaleh (37º13'N, 50º16'E); CMNFI 1979-

0430, 1, 118.0 mm standard length, Mazandaran, 

river east of Now Shahr (36º39'N, 51º31'E); CMNFI 

1979-0431, 7, 120.8-155.1 mm standard length, 

Mazandaran, Now Shahr bazaar (no other locality 

data); CMNFI 1979-0686, 2, 119.7-126.9 mm 

standard length, Gilan, Safid River (37º24'N, 

49º58'E); CMNFI 1980-0146, 2, 106.9-171.8 mm 

standard length, Mazandaran, Gorgan Talab at 

Ashuradeh-ye Kuchak (36º50'N, 53º56'E).  

Comparative material: BM(NH) 1938.8.2:3, 1, 203.8 

mm standard length, Caspian Sea (no other locality 

data); BM(NH) 1939.2.21:22-23, 2, 175.6-179.2 mm 

standard length, Caspian Sea (no other locality data); 

BM(NH) 1954.6.24:5-7, 3, 164.1-189.1 mm 

standard length, Caspian Sea (no other locality data). 

Alosa kessleri (Grimm, 1887) 
(Figs. 4, 5) 

Common names: Shagmahi-ye poshtsiah, shagmahi 
darya-ye siah, shagmahi-ye moohajer or shagmahi-

e-mohajer, zalun (in Gilaki), puzanok. [Volga 

siyanayi, garabel siyanak in Azerbaijan; arkasy gara 

takgas in Turkmenistan; blackback, Caspian 

anadromous shad; chernospinka or black-spined 

herring, chernonosik or blacknose, beshenka, zalom, 

poluzalom, zheleznitsa, veselka, Volzhskaya 

mnogotychinkovaya sel'd or Volga many-rakered 

herring, Volzhskaya sel'd or Volga herring, 

Astrakhanskaya sel'd or Astrakhan herring, all in 

Russian].  

Systematics: Clupea kessleri was originally 
described from the Volga River delta, Astrakhan. 

Clupea pontica (Eichwald, 1838) was originally 
described in Latin from "Hab. in Ponte Euxino prope 

Odessam" (= Black Sea near Odessa). Alosa kessleri 
was formerly considered as a subspecies of 

A. pontica. Alosa pontica then had two subspecies in 
the Caspian Sea, namely kessleri (Grimm, 1887) 
(chernospinka or black-spined herring, chernonosik 

or blacknose, beshenka, zalom, poluzalom, 

zheleznitsa, veselka, blackback), and volgensis 
(Berg, 1913) (Volzhskaya mnogotychinkovaya sel'd 

or Volga many-rakered herring, Volzhskaya sel'd or 

Volga herring, Astrakhanskaya sel'd or Astrakhan 

herring, zheleznitsa, beshenka, veselka). Kottelat 

and Freyhof (2007), Abdoli and Naderi (2009) and 

Figure 4. Line drawing of Alosa kessleri. 



143 
 

Int. J. Aquat. Biol. (2017) 5(3): 128-192 

 

Naseka and Bogutskaya (2009) consider Alosa 
kessleri and A. volgensis to be valid species.  

A lectotype of kessleri, 40.1 cm long, was 
designated from the Volga Delta by L. S. Berg under 

ZISP 15925 (in the Zoological Institute, St. 

Petersburg). A lectotype of volgensis, 34.8 cm long, 
is under ZISP 15926 and is from the Volga River at 

Chernyi Yar (Svetovidov, 1952). A paralectotype of 

kessleri is under ZISP 15922. 
Caspialosa volgensis bergi Tanasiichuk, 1938 

described from the Volga Delta is a synonym of 

Alosa kessleri (Heckman in Hoestlandt, 1991). 
Eschmeyer et al. (1996) give author and date for 

Alosa volgensis bergi as Tanassiychuk, 1940, the 
variation probably being due to transliteration of a 

Russian name and to year of actual publication rather 

than year on the journal.  

Caspialosa kessleri infraspecies volgensis imitans 
Berg, 1948 from the Caspian Sea (see Berg (1948-

1949) for further details) is not available because of 

its infrasubspecific rank (Eschmeyer et al., 1996). 

Clupea caspio-pontica Borodin, 1904 is an unneeded 
new name (Eschmeyer et al., 1996).  

Key characters: Characterised by a relatively 

elongate and rounded body likened to a "herring" 

shape, not as deep and compressed as in some related 

species which are likened to a "shad" shape. Total 

gill rakers 49-158 in the Caspian Sea, with ranges 

about 57-95 in kessleri and 87-158 in volgensis. 
Rakers are usually longer than the gill filaments in 

volgensis, shorter in adult kessleri. Teeth are well 
developed in both jaws in kessleri and can be felt 
with a finger, poorly developed in volgensis such 
that they sometimes cannot be felt.  

Morphology: Dorsal fin with 3-5 unbranched and 

12-16 branched rays, anal fin with 2-4, usually 3, 

unbranched and 15-21 branched rays. Vertebrae 47-

50 in kessleri (also a report of 50-54, both in 
Svetovidov (1952)), 48-54 in volgensis. Pyloric 
caeca 21-62. Scales in lateral series 53-56. Gill 

rakers in adults are thick and often broken off at the 

tip or near the base in kessleri, unbroken in 
volgensis. The tips of the gill rakers may be swollen 
and they are arranged in a straight line. Young fish 

have long and thin gill rakers with strong lateral 

spines. Spines are lost with age.  

Sexual dimorphism: None reported. 

Colour: The overall coloration is dark with a black 

back which has a violet tinge in spring in kessleri, 
light olive-green in volgensis. There is dark, 
sometimes vague, spot on the flank behind the 

operculum and sometimes a series of spots in 

kessleri, but these are absent in volgensis. The 
pectoral fin is black on top. Spawning kessleri 
become grey or grey-green on the back and flanks 

with bronze spots on the operculum and flanks. A 

greenish-yellow circle forms around the eye after 

spawning.  

Size: Reaches 52.0 cm total length and 2.0 kg for 

kessleri, 40.0 cm for volgensis. 
Distribution: Found in the Caspian Sea, entering 

northern rivers to spawn. Abdoli and Naderi (2009) 

list it as from the southwest, southeast and south-

central Caspian Sea in Iranian waters. Esmaeili et al. 

(2014) list this species from the Anzali Wetland or 

Talab. 

Zoogeography: This species is a Caspian Sea 

endemic. 

Habitat: Both subspecies are found in the open sea 

but kessleri ascends rivers much higher than 

Figure 5. Alosa kessleri courtesy of A. Abdoli. 



144 
 

Coad/ Herrings of Iran 

volgensis which spawns in the delta region. Both 
subspecies overwinter in the southern Caspian Sea 

off the Iranian coast and then migrate north to enter 

the Volga and other northern rivers to spawn. The 

subspecies volgensis is absent from the southern 
Caspian in summer. The subspecies kessleri shows a 
greater affinity than volgensis for cold water.  

The subspecies kessleri begins to migrate 
northward in March and April mostly along the 

western shore of the Caspian Sea, beginning to arrive 

in northern waters when temperatures are still below 

5°C, most arriving when temperatures are 6-8°C 

compared to 10-13°C for volgensis. A mass 
migration into the lower Volga takes place in late 

April or early May for both subspecies when water 

temperature reaches 9°C and the peak run begins at 

12-15°C, ending at 22°C. The run of volgensis is 
about 10 days later than that of kessleri and spawning 
takes place earlier as they do not travel as far upriver. 

Speed is up to 70 km/day for kessleri and depends on 
temperature. This fish used to run 2,000 km up the 

Volga River. Sexually immature fish remain in the 

south and do not migrate. Knipovich (1921) reports 

kessleri as deep as 235-300 m in Iranian waters. 
Temperatures up to 25ºC are tolerated.  

Age and growth: Males are sexually mature at 3 

years and females at 4 years, other reports give 4-5 

years for both sexes in kessleri. Many fish die after 
spawning but some survive to spawn two or three 

times. Four and five-year-olds dominate on kessleri 
spawning runs with some older fish also present. 

Females predominate in older fish making the 

spawning run. Life span is between 7 and 8 years. 

Growth of the volgensis subspecies is slower than 
in kessleri, which apparently grows faster than any 
other Caspian clupeid. Life span in volgensis is 7-8 
years with females living longer than males. Most 

spawners are 3-4 years old although in some years, 5 

year old fish are abundant. Males may mature at 2 

years, females later. Most fish spawn again the next 

year after their first time but some may miss a year. 

An individual may spawn up to four times during its 

life.  

Heidari et al. (2015b) examined fish from the 

Tonekabon fishing area and found b values of 3.687 
and 3.571 for total and standard length, indicating 

positive allometric growth. Alavi-Yeganeh et al. 

(2013) found Mazandaran fish were 2-5 years old 

and had a b value of 2.993. Ghotbi-Jokandan et al. 
(2015) examined 103 fish from Anzali, Noor and 

Torkeman and found a b value of 2.88. Mousavi-
Sabet et al. (2016) gave a b value of 3.13 for Iranian 
fish from 6 fishing areas from Astara to Miankaleh. 

Food: Cladocerans are the main food item of young 

kessleri which have a feeding peak at 1800-2200 
hours and another at about 0800 hours. Adults in the 

sea take fishes such as Clupeonella and Atherina 
with some crustaceans and insect larvae. Clupeonella 
caspia makes up 92% of the diet of kessleri in the 
northern Caspian in May, with Sander lucioperca at 
6.6% and gammarids at 1.0%. There is said to be 

little feeding on the spawning run although some fish 

sampled contained cladocerans, copepods, insects, 

bryozoans and fish fry.  

The food of volgensis is similar to the other 
subspecies, taking copepods when young and larger 

items with growth. The main items are copepods, 

mysids, cumaceans, amphipods and small fishes. 

This subspecies feeds on the spawning migration. 

Reproduction: Spawning in kessleri occurs in rivers 
from mid-May to mid-August, either the delta or 

lower reaches when entering in a ripe condition, or 

as much as 500 km upriver when entering in an 

unripe condition. Larger fish have spawning grounds 

further upriver than smaller fish and predominate 

earlier in the run. The spawning grounds in the Volga 

River cover a considerable stretch. Spawning usually 

occurs at 18-20°C between 0300 and 0600 hours or 

from 1600 hours to sunset. Spawning occurs in the 

main channel, over shallow sand banks, or in 

backwaters. Batches of eggs are laid at intervals of 

several days. Eggs are pelagic as in other Caspian 

Alosa and develop as they drift downriver near the 
bottom. At 22.7°C incubation takes about 40 hours. 

The young fish descend in late summer and early 

fall. Fecundity in kessleri reaches 344,000 eggs and 
egg diameter 1.51 mm. Shed eggs are up to 4.1 m in 

diameter. Some fish may return to spawn in total 



145 
 

Int. J. Aquat. Biol. (2017) 5(3): 128-192 

 
three times.  

Spawning of the first batch of eggs in volgensis 
may occur in the sea with the subsequent 2 batches 

at 7-10 day intervals in the delta and river. This takes 

place from mid-May to the beginning of August. Up 

to 281,000 eggs are shed. Peak spawning occurs at 

15-19°C and ends at 25-27°C. Most spawning takes 

place in the evening between the 1600 and 2200 

hours. The young appear in the pre-estuarine area of 

the Volga River in July and towards October begin 

to migrate south.  

Parasites and predators: The Caspian seal, Pusa 
caspica, is a predator on this species (Krylov, 1984), 
larval shad are fed on by other fishes and by various 

invertebrates, and adults by various fishes and birds. 

Economic importance: The subspecies kessleri and 
volgensis were caught on the spawning run with as 
much as 5,750 t being taken annually pre-World War 

II. It is the biggest shad in the Caspian Sea. The 

subspecies kessleri was the most important and 
valuable herring in the Caspian Sea. Early spring 

catches were mostly kessleri but as the run of 
volgensis built up it formed an increasingly 
significant part of the catch, forming as much as 92% 

of the total. The catch of volgensis has declined from 
this period until the 1970s when the fish taken were 

mostly kessleri. The catch of Alosa pontica (= 
kessleri) on the North Caspian fishing grounds in 
1965-1972 has declined to 2-4% of the 1938-1943 

catch. The subspecies volgensis was one of the most 
important Caspian herrings, 23-29% of the total 

catch from 1936-1939, as high as 69,100t in 1939. 

The subspecies kessleri is said to be the tastiest 
Caspian clupeid because of its high fat content, 

averaging 18.9% of weight along the coast of 

Azerbaijan, while in volgensis it was 9.6%. Post-
spawners of kessleri may have a fat content as low 
as 0.5%. Catches are processed as canned, salted and 

pickled fish. Beach seines are used to catch this fish. 

Akhondzadeh Basti et al. (2006) found the 

bacterial pathogen Vibrio haemolyticus in fresh and 
smoked Alosa kessleri. Tavakoli et al. (2012) found 
a frequent contamination rate with Staphylococcus 
aureus and Vibrio parahaemolyticus, human 

pathogens, in fresh and smoked shad and they may 

cause health problems. Robins et al. (1991) list this 

species as important to North Americans. 

Importance is based on its use as food.  

Conservation: Stocks in Iranian waters are said to be 

depleted. The subspecies volgensis was in Category 
I on the "Red List" of the Russian Republic (Pavlov 

et al., 1985). Kiabi et al. (1999) consider this species 

to be data deficient in the south Caspian Sea basin 

according to IUCN criteria. Criteria include 

commercial fishing, numbers unknown, range 

unknown absent in other water bodies in Iran, absent 

outside the Caspian Sea basin.  

Sources: Iranian material: None available. 

Comparative material: BM(NH) 1879.11.14:22-23, 

2, 255.9-259.1 mm standard length, Caspian Sea (no 

other locality data); BM(NH) 1939.2.21:21, 1, 388.6 

mm standard length, Caspian Sea (no other locality 

data). 

 

Alosa saposchnikowii (Grimm, 1885) 
(Figs. 6, 7)  

Common names: Shagmahi-ye cheshmdorosht, 

shagmahi, kilka (incorrectly), herring. [Irikoz 

sisgarin in Azerbaijan; bol'sheglazyi puzanok or 

bigeye shad, Sapozhnikovskii puzanok or 

Saposhnikovi shad, all in Russian].  

Systematics: The lectotype of Clupea 
saposchnikowii from the Volga Delta is in the 
Zoological Institute, St. Petersburg under ZISP 

15921 (Berg 1948-1949; Eschmeyer et al. 1996). 

The name is often spelt saposchnikovi, in error, or 
with a single terminal "i"; Reshetnikov et al. (1997) 

revert to the original spelling of the specific name. 

Caspialosa caspia nigra Kisselevitsh, 1923, in 
part, from the Caspian Sea opposite Dzambai is a 

synonym with a lectotype in the Zoological Institute, 

St. Petersburg (ZISP 15938) (Kisselevitsh is also 

transliterated Kiselevich and Kisselevitz). The 

material also included specimens of Alosa 
braschnikowii (Whitehead, 1985; Eschmeyer et al., 
1996).  

Key characters: Characterised by a relatively deep 

and compressed body likened to a "shad" shape, not 



146 
 

Coad/ Herrings of Iran 

as elongate and rounded as in some related species 

which are likened to a "herring" shape. The upper 

and lower head profiles are straight. The upper edge 

of the lower jaw is straight. Total gill rakers 20-48, 

short (obviously shorter than the gill filaments), and 

thick. Teeth are well developed in both jaws.  

Morphology: Dorsal fin with 3-5, usually 4, 

unbranched rays and 12-15, mostly 13, branched 

rays, anal fin with 2-4, usually 3, unbranched rays 

and 15-21, mostly 18, branched rays. Lateral series 

scales 52-55. Vertebrae 47-53. Pyloric caeca 36-59. 

Sexual dimorphism: None reported.  

Colour: Fish from the southern Caspian Sea are more 

intensively coloured than those from the north. The 

back is violet with a green sheen and the flank has 4 

dark stripes which merge with the dark on the back. 

There is a spot posterior to the operculum, which 

may be absent.  

Size: Reaches 36.0 cm total length and 650 g.  

Distribution: Found mainly in the north Caspian Sea 

and the coast of Dagestan but entering Iranian 

waters. Abdoli and Naderi (2009) list it as from the 

southwest, southeast and south-central Caspian Sea 

in Iranian waters and see below under Sources for 

some specific localities. 

Zoogeography: This species is endemic to the 

Caspian Sea. 

Habitat: This species spends its whole life in the 

Caspian Sea and never enters rivers. It favours colder 

water and is one of the first clupeid species to 

migrate north in spring, principally along the western 

coast. Large fish migrate first. Fish first approach the 

shore of Azerbaijan in mid-March with a mass 

approach from late March to mid-April. It is less 

frequently encountered in the southern part of the 

Caspian Sea, overwintering in the central Caspian 

and only moving south if winters are cold. A Caspian 

Sea Biodiversity Database (from www.caspian 

environment.org) has it at 400-600 m in the southern 

Caspian in cold winters but later states it keeps at 15-

Figure 6. Line drawing of Alosa saposchnikowii. 

Figure 7. Alosa saposchnikowii courtesy of A. Abdoli. 



147 
 

Int. J. Aquat. Biol. (2017) 5(3): 128-192 

 
32 m. Winter temperatures at which this species is 

found are 6-7°C. Depths are 25-32 m in winter, more 

shallow in summer but below 9 m. Knipovich (1921) 

reports this species in a depth range of 52-77 m in 

Iranian waters. It tolerates a range of 3-25°C and 

spawns at salinities of 0.7-11.0‰, although 

preferring 4.0-7.5‰. The Caspian Sea Biodiversity 

Database (from www.caspian environment.org) 

estimates a population of 1.1125 billion fish.  

Age and growth: Life span is about 9 years and 

female lengths and weights exceed those of males 

throughout life. On average, males weigh less than 

half the weight of females since females carry a 

heavy egg load. Growth is most intensive in the first 

two years of life and slows thereafter (Chang, 1972). 

Males mature at age 2 and females at age 3. Ghotbi-

Jokandan et al. (2015) examined 40 fish from Anzali, 

Noor and Torkeman and found a b value of 3.14. 
Mousavi-Sabet et al. (2016) gave a b value of 2.99 
for Iranian fish from 6 fishing areas from Astara to 

Miankaleh. 

Food: A rapacious fish which takes young herrings 

and kilka, Atherina and even Benthophilus 
(Lönnberg, 1900) as well as large crustaceans such 

as mysids and gammarids. It is a cannibal. This shad 

overwinters and feeds in the south Caspian Sea 

(Chang, 1972).  

Reproduction: The spring spawning migration (end 

of April to end of May) enters the north Caspian Sea 

and fish are mostly 15-25 cm in body length. Males 

mature at a younger age than females as evidenced 

by fish 3-4 years old predominating among females 

and fish 2-4 years old among males in the north 

Caspian catch. Spawning takes place in May 

(peaking in the first 10 days) and most fish are 

returning for the second time. Spawning 

temperatures are lower than in Alosa caspia, being 
only 13-14°C although the peak is at 19-20°C. 

Spawning occurs in il'mens, the sea where there is a 

freshwater discharge such as near the Volga River 

mouth, and in the northeastern sea. Females may 

spawn up to 6 times and males up to 5 times (Chang 

1972). Spawning takes place in shallow water at 1-

6m depths. Fecundity is up to 318,852 eggs. The 

young migrate southwards.  

Parasites and predators: Mazandarani et al. (2016) 

found a trematode (Pronoprymna ventricosa) and 
two nematodes (Anisakis simplex and 
Eustrongylides sp.) in the abdominal cavity of fish 
from Behshahr, Mazandaran Province. The latter 

two are pathogenic to humans. The Caspian seal, 

Pusa caspica, is a predator on this species (Krylov 
1984).  

Economic importance: An important commercial 

species in the central and northern Caspian, taken on 

their way to, and on, the spawning grounds. The 

fishery in Azerbaijan during 1937 caught fish on 

average 17cm long and 62g in weight, most fish 

being 2-3 years old. The Caspian catch in the period 

1936-1939 reached a peak of 8,800t annually. Fish 

are caught with beach seines, stationary nets and drift 

nets.  

Conservation: Stocks in Iranian waters are reputed to 

be depleted. Kiabi et al. (1999) consider this species 

to be data deficient in the south Caspian Sea basin 

according to IUCN criteria. Criteria include numbers 

unknown, range unknown, absent in other water 

bodies in Iran, absent outside the Caspian Sea basin. 

Sources: Iranian material: CMNFI 1970-0531, 15, 

49.9-108.7mm standard length, Mazandaran, Larim 

River (36º46'N, 52º58'E); CMNFI 1970-0532, 1, 

137.4mm standard length, Gilan, Caspian Sea near 

Bandar-e Anzali (37º28'N, 49º27'E); CMNFI 1970-

0543A, 2, 78.8-80.2mm standard length, Gilan, 

Caspian Sea at Hasan Kiadeh (37º24'N, 49º58'E); 

CMNFI 1970-0581, 1, 102.1mm standard length, 

Gilan, Caspian Sea near Hasan Kiadeh (37º24'N, 

49º58'E); CMNFI 1979-0788, 3, 96.0-114.9mm 

standard length, Mazandaran at Khadje Nafas 

(37º00'N, 54º07'E); CMNFI 1980-0136, 3, 107.3-

127.6mm standard length, Mazandaran, Fereydun 

Kenar River (36º41'N, 52º29'E); CMNFI 1980-0157, 

2, 96.6-101.1mm standard length, Mazandaran, 

Gorgan River estuary (36º59'N, 53º59'30"E); 

CMNFI 1980-0908, 1, 77.9mm standard length, 

Gilan, Safid River estuary (ca. 37º28'N, ca. 49º54'E). 

Comparative material: BM(NH) 1954.6.24:8-10, 3, 

150.5-177.0mm standard length, Caspian Sea (no 



148 
 

Coad/ Herrings of Iran 

other locality data). 

 

Alosa sphaerocephala (Berg, 1913)  
(Fig. 8) 

Common names: Shagmahi-ye Agrakhan. 

[Kruglogolovyi puzanok or roundheaded shad, 

Agrakhanskii puzanok or Agrakhan shad, both in 

Russian].  

Systematics: The holotype of Clupeonella 
sphaerocephala from Agrakhan Bay, at Tyulenii 
Island, Turali in the northern part of the Caspian Sea 

is in the Zoological Institute, St. Petersburg under 

ZISP 15928 with more than 30 paratypes 

(Eschmeyer et al., 1996). 

Key characters: Characterised by a relatively deep 

and compressed body likened to a "shad" shape, not 

as elongate and rounded as in some related species 

which are likened to a "herring" shape. The upper 

and lower head profiles are obviously rounded. The 

upper edge of the lower jaw is crescent-shaped. Total 

gill rakers 25-45, long (equal to or longer than the 

gill filaments), and thin. Teeth are well developed in 

both jaws.  

Morphology: Dorsal fin with 3-4, usually 4, 

unbranched rays and 13-15 branched rays, anal fin 

with 3-4, usually 3, unbranched rays and 17-20 

branched rays. Vertebrae 47-51.  

Sexual dimorphism: None reported.  

Colour: The back is dark with an olive tint, the tip of 

the snout is occasionally black and the pectoral fins 

are dark. There is a black spot behind the operculum 

and occasionally a row of such spots.  

Size: Reaches 25.0 cm standard length.  

Distribution: Found in the Caspian Sea including 

Iranian waters.  

Zoogeography: This species is endemic to the 

Caspian Sea.  

Habitat: This species does not enter fresh waters. It 

is most common along the eastern shore of the 

northern part of the sea in spring where spawning 

occurs and along the northern shore of the northern 

part of the sea in summer. Knipovich (1921) reports 

this species from Iranian waters in a depth range of 

52-77 m. 

Age and growth: Unknown.  

Food: Unknown, although presumably similar to 

other shads.  

Reproduction: Spawning takes place in the 

northeastern Caspian from mid-May to the end of 

June peaking at 18-20°C, most frequently in a 

salinity of 8-11‰ and in depths around 3-8 m. The 

young move south in late autumn, as late as 

November, the last clupeids to leave this area. 

Fecundity is about 20,000 eggs. 

Parasites and predators: The Caspian seal, Pusa 
caspica, is a predator on this species (Krylov, 1984).  
Economic importance: This species is caught only in 

small numbers.  

Conservation: The status of this species is unknown.  

Sources: Iranian material: None available. 
Comparative material: BM(NH) 1954.6.24:11-13, 3, 

145.6-162.1 mm standard length, Caspian Sea (no 

Figure 8. Line drawing of Alosa sphaerocephala. 



149 
 

Int. J. Aquat. Biol. (2017) 5(3): 128-192 

 
other locality data). 

 

Genus Clupeonella Kessler, 1877 
This genus is found in the Black and Caspian seas 

basins with 5 species, 3 of which are in the Caspian 

Sea and in Iranian waters. Clupeonella species are 
distinguished from sympatric Alosa species by 
smaller size, a small and toothless mouth, adipose 

eyelids are small or rudimentary, no spots on the 

flank, no elongate scales (ala) at the base of the 

caudal fin, no vomerine teeth, the lack of a notch at 

the mid-line of the upper jaw, and by the last two anal 

fin rays being elongated.  

Species in this genus live entirely in the sea, or in 

fresh water, or migrate between the two. Eggs are 

pelagic and have a large oil globule. The general 

Farsi name for these fishes is kilka or kelka, i.e. 

"sprat", although sprat is erroneous according to 

Berg (1948-1949) who uses tyulka for these fishes).  

The three Clupeonella species have been fished in 
modern Iran since December 1971 but the 

commercial catch did not exceed 15,000 tonnes. 

Earlier catches date back only to 1939, with an 

annual catch of about 100t in 1943-1949, exported in 

a marinated form to the Soviet Union (Alam, no 

date). Curiously, the abundance of kilka has long 

been known as Kinneir (1813) records "and herrings 

are in such abundance, that after a storm, the shores 

of Ghilan and Mazanderaun are nearly covered with 
them". Caddy (1984) refers to the kilka fisheries of 

the Iranian Caspian by the scientific name Sprattus 
sprattus but this is an error.  

Caddy (1984) indicated that there were problems 

in marketing and utilizing these fishes in Iran even 

though up to 50,000 t could be caught annually 

(200,000 t elsewhere in the same article). Their best 

use was probably as food for predators such as 

Sander lucioperca, Esox lucius and Salmo caspius. A 
study by Razavi Sayad (1993) suggested a ceiling of 

100,000 t was possible. The Caspian Sea resources 

of kilka are estimated at 800,000 t from which 

340,000 t can be exploited (Abzeeyan, Tehran, 6(8): 

IV, 1995). 

The catch reached 51,000 t in 1994 from none 10 

years previously (Food and Agriculture 

Organization, Fisheries Department, 1996) and was 

36,000 t in 1997-1998 (Irna, 31 March 1998) and 

85,000 t in 1998-1999 (Fazli and Roohi, 2002a). The 

catch for the first 6 months of the Iranian year was 

17,000 t, taken by 70 trawlers and a 10% increase 

over the previous year (Irna, 20 October 1998). 

Fishermen in Gilan caught 50,000t annually in the 

late 1990s (Tehran Times, 5 September 1999). A 

reported catch of 56,000 t was made in 1999-2000, a 

13% increase over the previous year (Irna, 27 March 

2000). A later estimate expects the kilka catch to 

reach 66,000 t by the year 2000 (Abzeeyan, Tehran, 

5(9): IV, 1995).  

Fazli (2006) records that kilka fishing ships 

discharge their catches at three ports, Babol-Sar and 

Amirabad in Mazandaran and Anzali in Gilan. The 

catch decreased from 28,000 t to 19,600 t in 

Mazandaran and from 57,000 to 42,600 t in Gilan 

from 1999 to 2000. The catch per unit effort also 

decreased from 3,900 kg to 2,500 kg over the two 

years. Anchovy kilka dominated the catch but the 

frequency fell from 85-90% to 76% of the catch and 

common kilka sharply increased. Common kilka had 

been caught in spring and summer but in 2000 they 

were taken in all months. The average length of 

anchovy kilka declined from 96.3 mm in 1997 to 

87.3 mm in 2000 and this was also reflected in the 

age structure, 5+ and 6+ fish being rare.  

The presence of the ctenophore, M. leidyi, was 
thought to be damaging stocks (Fazli and Roohi 

2002a; Ahmadpour et al., 2012). Darvishi et al. 

(2004) studied dietary overlap between the 

ctenophore and the anchovy kilka (see below). Barati 

(2009) found that the diet of chicks of the Great 

Cormorant, Phalacrocorax carbo, at Ramsar 
comprised only 2% Clupeidae by frequency, perhaps 

a result of the decline in Clupeonella stocks.  
Fazli (2002) studied kilka catches off Mazandaran 

in 1996-2000. Fishing occurred at night and lasted 

7.78-8.22 hours. The maximum catch at 42.8% was 

taken in October, November and December with a 

minimum catch in June. The least annual catch per 

vessel occurred in 1999-2000 (499,401 kg).  



150 
 

Coad/ Herrings of Iran 

A study utilizing an echo-sounder and a pelagic 

trawler concludes that the maximum biomasses for 

the three Clupeonella species in the southern Caspian 
Sea were in winter (422,300 t) and autumn (326,900 

t) while in summer and spring values were lower at 

275,100 t and 260,800 t, respectively. The 

population consisted of 66.1% anchovy kilka 

(C. engrauliformis), 18.9% bigeye kilka (C. grimmi) 
and 15% common kilka (C. caspia) (Iranian 
Fisheries Research and Training Organization 

Newsletter, 14: 6, 1996). Note that later, the Iranian 

Fisheries Research and Training Organization 

Newsletter (17: 3, 1997) gives kilka biomass in the 

southern Caspian Sea as winter 22,300 t, autumn 

26,900 t, summer 75,100 t and spring 60,800 t, 

presumably lacking the initial digit, and the 

percentages of kilka species in the biomass are also 

wrong. This is corrected in a subsequent newsletter 

(Iranian Fisheries Research and Training 

Organization Newsletter, (18: 43, 1997) but the 
corrected percentage biomasses are given as 66% for 

C. engrauliformis, 19% for C. caspia (as 
C. delicatula) and 15% for C. grimmi. It is unclear 
whether grimmi or caspia is the second most 
important kilka species. Poorgholam et al. (1996) 

gave a stock assessment for kilkas in 1995-1995 

using the hydro-acoustic method.  

Clupeonella engrauliformis dominated the catch 
in Iran at 91.8%, followed by C. grimmi at 6.84% 
and by C. caspia at only 1.35% (Coad 1997). The 2+ 
and 3+ year classes accounted for 69.95% of 

C. engrauliformis, 81.06% of C. grimmi and 80.88% 
of C. caspia catches. Catch rates of kilka on the top 
ranking 17 fishing grounds of 56 studied range from 

800 to 1,200 kg per unit effort per hour while 

traditional grounds have rates of 400-800 kg per unit 

effort per hour. The kilka are caught by attraction to 

lights and netting or pumping the catch into specially 

constructed ships (Fig. 9).  

The kilka fishing fleet of Iran expanded in the 

1980s and 1990s. There were 30 active vessels in 

Mazandaran in 1994, each with a capacity up to 

30tons (sic, probably tonnes here and elsewhere for 
modern catches) (Abzeeyan, Tehran, 4(10): IV, 

1994). The Mazandaran Kilka Cooperative 

Companies Union had 75 boats in 2000 (Tehran 

Times, 31 December 2000). Gilan planned to 

construct 12 fish meal factories each with an annual 

capacity of 1,000 t and 10 kilka canneries also with 

1,000t capacities (Abzeeyan, Tehran, 4(4): III, 

1993). Catches off Gilan alone from April 1994 to 

January 1995 increased 59% compared to the same 

period in 1993-1994, exceeding 20,000 t (Abzeeyan, 

Tehran, 6(1): II, 1995). The catch off Mazandaran 

from March 1994 to March 1995 was 15,400t, an 

increase of 10% over the previous year. About 1,000 

t were processed for human consumption and the rest 

for fishmeal production (Abzeeyan, Tehran, 6(2): V, 

1995). The total kilka catch for Iran has increased to 

45,000 t annually and efforts were being made to 

increase it to 110,000 t (Abzeeyan, Tehran 4(5): IV, 

1993). The catch in 1995 was 32,000 t with 64.7% 

from Mazandaran and 35.3% from Gilan, with the 

maximum catch occurring in April (Abzeeyan, 

Tehran, 7(6): II, 1996). Catches declined from 

95,000 t in 1999 to 15,497 t in 2003 (Sayyad Bourani 

et al., 2008). Annual Soviet catches reached 37,000 t 

in 1956 but this declined to 300-1,500 t by the end of 

the 1970s or 0.2-0.8% of all kinds of tyulka or kilka 

in the Caspian Sea. Turkmenistan harvested 7,660 

and 8,500 t in 1995 and 1996 although previously 

almost 45,000t valued at $22.5 million had been 

taken before equipment deteriorated (http://bisnis. 

doc.gov/isnis/isa/9805fish.htm, downloaded 14 

March 2000). Stocks remained large even though 

kilka were heavily fished. Kohani et al. (2012) 

Figure 9. Conical lift net or kilka net after www.farsnews.com. 



151 
 

Int. J. Aquat. Biol. (2017) 5(3): 128-192 

 

reviewed fishing in Gilan and noted the number of 

vessels used increased from 9 in 1989 to 110 in 1999, 

resulting in the highest catch at 55,900 t. Mis-

management, the invasive ctenophore, pollution and 

temperature changes caused a decline in catches to 

4,603 t in 2009.  

The kilka fisheries were threatened by the comb 

jelly (a ctenophore, M. leidyi), which arrived in the 
Caspian Sea in 1995 in the ballast water of ships and 

spread through the entire sea by the year 2000, 

feeding voraciously on zooplankton. It is now known 

as the "Caspian monster" despite its small size of 

5cm (J. Muir 2001, http://news.bbc.co.uk/2/ 

hi/middle_east/1453117.stm). It doubles in size in 

one day, reaches maturity in two weeks and then 

produces 8,000 young each day. The fisheries 

collapsed by 50% in a few months, catches by one 

fisherman falling from being 3-6 t a night to half a 

tonne. Bilio and Niermann (2004) review the 

interplay of overfishing, pollution and the 

ctenophore invasion on the kilka fisheries of the 

Black and Caspian seas, Kideys et al. (2005) gave an 

overview of ctenophore impacts on fisheries, and 

Roohi et al. (2010) compared the effects on 

phytoplankton, zooplankton, zoobenthos and kilka. 

Ghadirnejad (2003) reported that C. engrauliformis 
originally dominated the kilka catch at 85-90% but 

has dropped to 55% through the impact of the comb 

jelly which has up to 2,285 individuals per cubic 

metre in the southwest Caspian Sea. Roohi et al. 

(2010) stated that Iranian landings of kilka dropped 

about 70% from 69,070 t in 1995-2000 to 23,430 t in 

2001-2006, a loss of at least U.S. $125 million. 

Parafkandeh (2006) gave similar figures for declines 

in catches. 

Fazli et al. (2009a) describe a multi-species 

approach for stock management, allowing for the 

decline of C. engrauliformis and increase in 
C. caspia in Iranian waters through competition with 
the ctenophore. The fisheries may recover somewhat 

after the comb jelly population collapses (Tidwell, 

2001) or if a predator, Beroe ovata, is introduced and 
can survive in the less saline waters of the Caspian 

Sea (J. Muir 2001, http://news.bbc.co.uk/2/hi/ 

middle_east/1453117.stm). Studies indicate it can 

survive the brackish Caspian Sea water, feed on the 

comb jelly and not feed on other plankton (Iranian 

Fisheries Research Organization Newsletter, 36: 35, 

2003). The catch records for the total kilka catch in 

Mazandaran in tonnes is courtesy of F. Darvishi 

(pers. comm. 2003) (Table 6) and shows the drastic 

decline caused by the ctenophore, as well as monthly 

variations in catches. 

The species composition of kilkas changed after 

the introduction of the comb jelly - comparing the 

year 2000 and before with the year 2002 - the 

common kilka changed from about 1-5% to about 

30%, the bigeye from about 10-15% to 0.2% and the 

anchovy kilka from about 85-90% to about 70% 

(Iranian Fisheries Research Organization 

Months/Years 
1998 

(1377) 

1999 

(1378) 

2000 

(1379) 

2001 

(1380) 

2002 

(1381) 
Mean 

March-April (Farvardin) 2,848 2,703 4,644 1,217 876 2,458 

April-May (Ordibehesht) 1,116 607 972 1,422 195 862 

May-June (Khordad) 370 763 1,819 125 158 647 

June-July (Tir) 1,392 919 194 425 444 675 

July-August (Mordad) 2,152 2,306 433 614 249 1,151 

August-September (Shahrivar) 3,117 2,010 581 528 336 1,314 

September-October (Mehr) 3,103 6,184 1,785 432 575 2,416 

October-November (Aban) 4,120 3,468 2,305 3,051 1196 2,828 

November-December  Azar) 3,835 3,410 2,655 993 - 2,723* (2,179) 

December-January (Dey) 2,754 1,735 620 1,082 - 1,548* (1,238) 

January-February (Bahman) 3,968 1,262 2,146 1,586 - 2,241* (1,792) 

February-March (Esfand) 2,815 1,667 1,192 1,903 - 1,894* (1,515) 

Total 31,590 28,034 19,648 13,378 4,029   

* = averaged over 4 and (5) years. 

Table 6. The catch records for the total kilka catch in Mazandaran in tonnes is courtesy of F. Darvishi (pers. comm. 2003). 



152 
 

Coad/ Herrings of Iran 

Newsletter, 36: 2, 2003). The catch per unit effort 

(catch per vessel per fishing night) fell from 4 t to 1 

t. The catch during 1997-1999 of anchovy kilka fell 

from 51,300 t to 491 t and bigeye kilka from 7,600 t 

to 309 t while common kilka rose from 1,500 t to 

24,600 t (Iranian Fisheries Research Organization 

Newsletter, 65: 4, 2011). Parafkandeh Haghighi and 

Kaymaram (2012) found, for the years 2006-2007, 

that the common kilka was the dominant species 

(89.7%) while the anchovy kilka was at only 8.7% 

after previously being the dominant species. This 

was attributed to the comb jelly which occupied the 

anchovy kilka habitat at depths greater than 50 m. 

The total catch of kilkas fell from 95,000 t in 1999 to 

less than 20,000 t in 2007. The fishery moved to 

areas with depths less than 50m, the main reason for 

the change in species composition.  

In 2004, more than 200 fishing boats had been 

forced to stop operations. The kilka stock has been 

reduced from 400,000 t to 80,000t over the past 4 

years and the catch fell by 34,000 t (www. 

iranmania.com, downloaded 4 October 2004). 

Mamedov (2006) gives details of the biology and 

decline of kilkas in Azerbaijan waters. 

The Caspian seal was once a major predator on 

kilkas but the number of seals has declined on the 

Kazakhstan and Iranian coasts from 300,000 to 5,000 

in recent years through DDT pollution, viral 

infections and food shortages (Hashemi, 2001). 

Fazli (2011) evaluated changes in species 

composition, catch and catch-per-unit-effort from 

1961 to 2009. He found a negative correlation 

between long-term sea level changes and total catch 

which significantly declined in 1995 when sea level 

increased to its highest level. The catch-per-unit-

effort before the ctenophore invasion (1996-1999) 

and after (2000-2009) showed significant declines in 

C. caspia and C. engrauliformis while C. grimmi 
increased significantly. Overfishing and environ-

mental factors after the introduction of the 

ctenophore were the major factors in species 

composition changes. Catches are presented in Table 

7. 

Fazli et al. (2013) also examined the stock status 

of Iranian kilkas. The anchovy kilka and all three 

species together showed no overfishing from 1991 to 

2000 but in 2000 the catch-per-unit-effort neared an 

“in danger” region and from 2001 to 2007 stocks 

were overfished. During 2008-2010 the biomass 

increased and the stocks became non-overfished. 

The anchovy kilka fishery generally reduced since 

1998 and collapsed to the lowest level in 2010. For 

the anchovy stock to recover, the catch of all three 

species would have to be restricted during the 

spawning period and an international effort made to 

manage stocks. 

The Food and Agriculture Organization 

(www.fao.org/fishery/facp/IRN/en, downloaded 16 

January 2017) summarised catches in the south 

Caspian Sea (Table 8), showing the relative 

Year 
C. caspia 

catch (t, %) 

C. caspia 

CPUE* 

C. engrauliformis 

catch (t, %) 

C. engrauliformis 

CPUE* 

C. grimmi 

catch (t, %) 

C. grimmi 

CPUE* 

1997 52,246, 86.5 3.35 6,704, 11.1 0.5 1,450, 2.4 0.09 

1998 61,880, 72.8 2.92 18,445, 21.7 0.87 4,675, 5.5 0.22 

1999 67,450, 71.0 3.12 14,535, 15.3 0.67 13,015, 13.7 0.6 

2000 57,486, 73.7 2.26 9,828, 12.6 0.39 10,686, 13.7 0.42 

2001 37,590, 83.2 1.31 2,801, 6.2 0.1 4,789, 10.6 0.17 

2002 17,358, 69.5 0.75 25, 0.1 0 7,592, 30.4 0.33 

2003 7,530, 50.5 0.57 89, 0.6 0.01 7,291, 48.9 0.55 

2004 5,089, 26.9 0.4 227, 1.2 0.02 13,602, 71.9 1.06 

2005 4,249, 18.8 0.31 542, 2.4 0.04 17,808, 78.8 1.3 

2006 1,896, 8.5 0.15 1,048, 4.7 0.08 19,356, 86.8 1.51 

2007 910, 6.0 0.1 379, 2.5 0.04 13,881, 91.5 1.49 

2008 440, 2.7 0.06 277, 1.7 0.04 15,573, 95.6 2.13 

2009 454, 1.8 0.06 302, 1.2 0.04 24,444, 97.0 3.35 

*CPUE = catch-per-unit-effort (tonnes per vessel per night) 

Table 7. The catch after after the introduction of the ctenophore (after Fazli, 2011). 



153 
 

Int. J. Aquat. Biol. (2017) 5(3): 128-192 

 

importance of kilka. 

The by-catch from the kilka fishery on the 

grounds at Anzali comprised 0.2% of the catch 

(Moradinasab et al., 2015). This low by-catch was 

attributed to the use of lantern nets and the positive 

phototropism of kilkas. By-catch species were 

A. caspia, the mullet Mugil cephalus, the cyprinids 
Alburnus chalcoides, Cyprinus carpio and Rutilus 
kutum, and the catfish Silurus glanis.  

Kilka are smoked, salted, canned in sauce and oil 

and marinated according to a traditional recipe and 

seasoned with fruits, herbs and vegetables in Iran 

(Keivany and Nasrollahzadeh, 1990; www.netiran. 

com/business.html, downloaded 31 October 2003). 

Moini and Koochekian (2003) gave details of fish 

sauce production from kilkas using traditional, 

microbial and enzymatic methods, along with taste 

tests. Vacuum packaging of fresh, smoked and salted 

kilka has been investigated in Iran (Annual Report, 

1995-1996, Iranian Fisheries Research and Training 

Organization, Tehran, p. 45-46, 1997) and studies on 

processing kilkas as fish balls have also been carried 

out (Annual Report, 1994-1995, Iranian Fisheries 

Research and Training Organization, Tehran, p. 40, 

1996). Koochekian Sabour and Moini (2009) 

described investigations on using Iranian kilkas to 

produce a fermented fish sauce for marketing in 

Southeast Asian countries. One company markets 

kilka in a clear package which gives the product a 

bright and colourful appearance. Kilka have even 

been made into crackers (Iranian Fisheries Research 

and Training Organization Newsletter, 18: 6, 1997, 

Shojaei 1998). Kilka have also been made into oil as 

a by-product of the fish meal industry (Iranian 

Fisheries Research and Training Organization 

Newsletter, 27: 3, 2001). Omega-3 fatty acids have 

been extracted from kilka oil under laboratory 

conditions (Salmani Joloudar et al. 2009). M. 

Shivazad, H. John Mohammady, A.A. Yousef 

Hakimi and H. Fazaely (http://iman.ut.ac.ir 

/news/agr.htm, downloaded 12 December 2004) 

discuss the use of C. engrauliformis as fish meal in 
animal nutrition and analyse the protein quality and 

Faeed et al. (2006) studied spoilage in kilka meal 

from bacteria and fungi. Nassiri Moghaddam et al. 

(2007) studied commercial samples of kilka fish 

meal and found differences in protein efficiency ratio 

and net protein ratio, attributed to the species used, 

conditions and length of storage and processing 

methods. Agh et al. (2012) showed that replacing 

kilka fish oil with vegetable oil in the diet of rainbow 

trout (Oncorhynchus mykiss) broodstock sign-
ificantly reduced total fecundity and hatching rate, 

and increased larval mortality and deformed larvae 

rate. Jalili et al. (2012) compared diets with 

Clupeonella fish meal and plant protein in various 
proportions on growth of rainbow trout fry and 

found, for example, replacing 50% of the fish meal 

with plant protein did not affect growth nor did a 

complete replacement with poultry by-product and 

plant proteins. 

The Iranian Fisheries Research and Training 

Organization Newsletter (20: 4, 1998) and Rezaei et 

al. (2003) report on methods of transporting kilka in 

cold water and crushed ice to processing factories 

which were better than traditional methods. Salmani 

et al. (2001) recommended chilled sea water for 

preservation for human consumption. Motamed-

zadegan et al. (2009) found that partial hydrolysis of 

fish myofibrillar proteins using papain improves its 

functionality. Motalebi et al. (2010) investigated the 

use of whey protein coating on quality and shelf life 

of kilkas; it can enhance quality and increase frozen 

shelf life in fish stored for up to 4 months. Khanedan 

 2003 2004 2005 2006 2007 2008 2009 2010 2011 2012 2013 2014 

Bony fish 16,573 15,665 21,845 23,802 23,538 20,045.5 18,665 16,601 17,034 16,160 17,146 16,733 

Sturgeon 463 500 416 330 225 178.5 131 94 80 68 56 41 

Kilka 15,497 19,610 22,626 22,303 15,411 16,743 25,483 27,110 20,717 24,086 23,221 22,873 

Total 32,533 35,775 44,887 46,435 39,174 36,967 44,279 43,805 37,831 40,314 40,423 39,647 

 

Table 8. The summarised catches in the south Caspian Sea (Food and Agriculture Organization, 2017) 



154 
 

Coad/ Herrings of Iran 

et al. (2011) found that an edible film of sodium 

alginate on dressed kilka increased shelf life. 

Seyfzadeh et al. (2013) used an edible film of whey 

protein and sodium alginate to coat gutted kilka, 

enhancing quality and shelf life for up to 6 months in 

frozen storage.  

Valipour Meri et al. (2011) showed that Bacillus 
licheniformis can decrease significantly the 
aflotoxins in kilka fishmeal. Khoshkoo (2008) and 

Khoshkoo et al. (2010, 2012) studied deterioration of 

fish protein concentrate made from kilkas caught 

near Anzali in different packaging and temperatures, 

stored in light and darkness. Vacuum packaging and 

storage in darkness at lower temperatures was the 

best method to avoid deterioration. Ebadi and 

Shokrzadeh (2006) examined Clupeonella species in 
Mazandaran for the organochlorine pesticide lindane 

and found levels in muscle tissues to be less than 

FAO and WHO recommended permissible intake 

and so were not a public concern. 

An account on the biology and identification of 

Caspian kilka in Farsi is given by Emadi (1991) and 

Fazli (1990). Fazli and Besharat (1998) and 

Poorgholam et al. (1996) give accounts of biology 

and catches in Iran in Farsi. 

 

Clupeonella caspia Svetovidov, 1941 
(Figs. 10, 11) 

Common names: Rizeh keraye, rizeh kuli, kilka-ye 

ma'muli or kilka-e-maamooli (= common shad). 

[Xazar kilkasi in Azerbaijan; adaty kulke balyk in 

Turkmenistan; Kaspiiskaya tyul'ka or kil'ka (i.e. 

Caspian tyulka or kilka), tyulka, obyknovennaya 

tyul'ka (i.e. common tyulka), all in Russian; common 

kilka, common Caspian kilka, sardelle, Caspian 

sprat]. 

Systematics: Formerly identified as Clupeonella 
cultriventris (Nordmann, 1840), originally described 
from the northern shore of the Black Sea. 

Clupeonella delicatula (Nordmann, 1840), described 
from Odessa market on the Black Sea, is a synonym 

of C. cultriventris and a lectotype is in the Zoological 
Museum St. Petersburg under ZISP 2254 with 

paralectotypes also under ZISP 2254, as designated 

by Svetovidov (1952). Clupeonella delicatula caspia 
Svetovidov, 1941 was considered to be a synonym 

and was described as from the "Caspian Sea, where 

it is met with almost everywhere, from very saline 

parts (Kaydak Bay) to quite fresh. Enters the mouths 

of the Volga and Ural rivers, ascending sometimes 

very far upstream". The holotype is from the Volga 

Delta and is under ZISP 15883 (Svetovidov, 1952). 

Kottelat and Freyhof (2007) considered this 

subspecies to be a distinct species found in the 

Caspian Sea with cultriventris restricted to the Black 
Sea. Reshetnikov et al. (1997) considered 

recognition of this subspecies as questionable. The 

Caspian Sea taxon, Clupeonella caspia, has a 
lectotype, 152 mm long, designated by Svetovidov 

(1952) in the Zoological Institute, St. Petersburg 

(ZISP 15883).  

Clupea cultriventris is spelled cultiventris in some 
parts of Eschmeyer et al. (1996), apparently in error. 

Three syntypes of Clupea cultriventris may be in the 
Muséum National d'Histoire Naturelle, Paris under 

MNHN 3681 (Svetovidov, 1952; Eschmeyer et al., 

Figure 10. Line drawing of Clupeonella caspia. 



155 
 

Int. J. Aquat. Biol. (2017) 5(3): 128-192 

 

1996). 

Clupea cultriventris var. tscharchalensis Borodin, 
1896 from Lake Charkhal in the Ural River basin is 

variously listed as a variety, morpha or a distinct 

species (see Svetovidov (1952); Kottelat and 

Freyhof (2007)). mtDNA studies of fish from 

Mazandaran and from Gilan showed statistically 

significant differences in haplotype frequencies, 

indicating genetically different populations (Lalouei 

et al., 2006). Norouzi et al. (2012, 2013) examined 

microsatellite markers in fish from Anzali collected 

in spring and summer and found evidence for two 

differentiated populations in the southwest Caspian 

Sea. Norouzi et al. (2014) also sampled spring and 

summer populations at Anzali and also at Babol-Sar 

and found significant genetic differentiation among 

seasons and regions.  

Key characters: This species has a moderately deep 

body (21-27% of standard length), a short and wide 

head (interorbital width 16% or more of head 

length), a sharply keeled belly, and pointed pectoral 

fin tips. 

Morphology: The dorsal fin has 3-4 unbranched 

rays, usually 3, followed by 11-14 branched rays and 

the anal fin has 1-3 unbranched rays, usually 3, and 

14-19 branched rays. Scales in lateral series 42-55. 

There are 24-30 belly scutes and 41-62 (rarely to 64), 

usually 51 or more, gill rakers. Vertebrae 40-44 

(rarely to 45) compared to 44-47 in the anchovy kilka 

and usually 46-48 in the bigeye kilka, probably as a 

result of higher water temperatures during 

development compared to other kilka species 

(Prikhod'ko, 1979b).  

Sexual dimorphism: Sexual dimorphism is only 

evident during egg development when the belly of 

females is swollen.  

Colour: The back is blue-green or light-green, the 

flanks silvery and the belly silvery-white or golden-

yellow. Fins are hyaline except the dorsal fin which 

has a central dark but faint stripe and the caudal fin 

which is darkish at the base. The iris is black. 

Size: Reaches 14.5 cm standard length and 19.0 g.  

Distribution: Found in the Caspian Sea, tributary 

rivers and some adjacent lakes. In Iran, it is reported 

from sea and also the confluence of the Pasikhan and 

Pir Bazar rivers of the Anzali Talab and the Anzali 

Talab and its outlets by Holčík and Oláh (1992), and 

from the Safid River and Anzali Talab by Abbasi et 

al. (1999). Abdoli and Naderi (2009) list it as from 

the southwest, southeast and south-central Caspian 

Sea in Iranian waters. Esmaeili et al. (2014) recently 

list it from the Anzali Wetland or Talab. Also in 

Gorgan Bay. 

Zoogeography: This species is a Caspian Sea 

endemic.  

Habitat: The habitat of this species in the Caspian 

Sea is the coastal zone of the sea at depths less than 

100m, more usually less than 50-70 m, over a wide 

range of temperatures (2.6-27.6°C for adults, higher 

for larvae, and possibly lower temperatures since 

they are found under ice and probably over 28°C 

according to some reports), and in fresh and 

Figure 11. Clupeonella caspia courtesy of A. Abdoli. 



156 
 

Coad/ Herrings of Iran 

hypersaline waters (to 36‰). The young can develop 

in water at 16‰. Southern populations live in a more 

saline habitat than northern and central Caspian 

populations which are mostly in fresh water. This 

tyulka may not migrate far but does move between 

summer-winter feeding and spring-early summer 

spawning grounds. Large schools are found 0.5-2.0 

km from shore at depths of 20-25 m on the eastern 

coast of the Caspian Sea, descending deeper if water 

temperatures rise and coming up to about 8 m in 

autumn as temperatures fall. In winter this species is 

found at about 30-40 m deep where the temperature 

range is 7-10°C, warmer than surface waters. Larvae 

and young remain in shallow coastal areas. 

Knipovich (1921) reports a fish from a depth range 

of 235-300 m in Iranian waters but populations at 

these depths are small (Iranian Fisheries Research 

and Training Organization Newsletter, 14: 6, 1996). 

The Caspian Sea Biodiversity Database (from 

www.caspianenvironment.org) states that the largest 

concentrations are found at 3-7‰ with most 

intensive spawning at 2-4‰.  

It is the most widely distributed kilka and with the 

other kilka species the most abundant fish in the 

Caspian Sea (Prikhod'ko, 1979b). Large schools can 

be found by day but these disperse at night. It 

overwinters in the southern Caspian Sea and some 

individuals move north to spawn and feed in April. 

The Caspian Sea Biodiversity Database (from 

www.caspianenvironment.org) estimates the 

population to number 224 billion fish, with 96 billion 

fish in the south Caspian. The south and north 

Caspian Sea stocks are about equal in number after a 

decline in copepod biomass in the north. The relative 

frequency of this species compared to other kilkas 

increased after the invasion of M. leidyi, by more 
than 10% (Fazli et al., 2006).  

The rise in Caspian Sea level since 1977 is 

gradually returning the Anzali Talab to its supposed, 

natural brackish state and may improve the fisheries 

situation which had declined over the last 50 years. 

Emergent and submergent aquatic macrophytes were 

decreasing and such fish as C. cultriventris (= caspia) 
were increasing in numbers since 1989 (Mandych, 

1995). However the fishery will require extensive 

engineering and management innovations to recover. 

Age and growth: Osipov and Kiyashko (2008) found 

that using otoliths gave more reliable estimates than 

using scales for ageing. This Caspian species grows 

faster than its Black Sea relative. Together with the 

sturgeons, this species comprises 82.1% of the fish 

biomass in the Caspian Sea. Condition in this species 

is better in winter because of the summer-autumn 

feeding period after spring spawning compared to 

C. engrauliformis in the Big Kizil-Agach (= 
Bol'shoy Kyzylagach or Imeni Kirova) Bay of 

Azerbaijan (Badalov, 1972). Local populations have 

differing growth regimes depending on the 

productivity of these areas (Prikhod'ko, 1979b) and 

there are great variations on a yearly basis too. 

Southern populations grow faster than northern ones 

in their first year. Females grow somewhat faster 

than males (9.0 g versus 7.3 g average weight along 

the Dagestan coast for example), and life span is 

about 6 years. This species is mature there at 1 year 

and average life span is about 3 years.  

Females dominated the population in Iran and 

sexual maturity was attained usually at age 2 and 2-

4 year olds dominated catches but life span is up to 8 

years (Iranian Fisheries Research and Training 

Organization Newsletter, 14: 6, 1996; Abtahi et al., 

2002). Fazli et al. (2006) found age classes 0+ to 5+ 

in Iranian waters with 0+ to 3+ making up 95% of the 

fish in 1997-1999. In 2000, age classes 0+ and 1+ 

were reduced in numbers and 2+ to 4+ fish comprised 

93.8%. Parafkandeh Haghighi (2010) examined 

Iranian populations in 2006-2007 and found a mean 

length of 102.4 mm, negative allometric growth, a 

mean age of 3.6 years, a male:female sex ratio of 

0.6:1, the growth coefficient (K) was the highest of 

the three species at 0.321, total mortality (Z) was 

1.28/yr, natural mortality (M) was 0.622/yr, fishing 

mortality (F) was 0.658, exploitation rate (E) was 

0.514, and the maximum constant yield for the kilka 

fishery was 14,100 t. Fazli et al. (2013) studied the 

status of stocks in Iranian waters from 1996 to 2011. 

The length range with optimum yield was 80-100 

mm, mega-spawners were larger than 105 mm, the 



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percentage of mature fish increased, optimum size 

declined since 2005, the percentage of mature fish 

decreased from 100% to 58% in 1999 and increased 

to 90% in 2001, and this kilka stock was in better 

condition than the other two species. 

Fazli et al. (2017) found that the biomass of this 

species increased from 22,000 t in 1997 to 112,000 t 

in 2009 and declined to 83,300 t in 2013. Over this 

time span the instantaneous coefficient of fishing 

mortality varied between 0.246/yr and 1.64/yr and 

exploitation rates were 0.327 and 0.764. The 

reference points at F0.1 and F40% were estimated at 
0.92 and 0.8/yr, respectively. The maximum 

sustainable yield (MSY) and fMSY were estimated at 
22,670t and 8,690 vessel x nights (a unit of effort). 

The acceptable biological catch should be restricted 

to 17,500 t. 

Abtahi et al. (2004) examined fish from the 

conical net and light catch at Babol-Sar, Mazandaran 

and found average fork lengths were 69.82 mm, 

83.56 mm, 88.38 mm and 88.43 mm while weights 

were 2.2 g, 4.18 g, 4.77 g and 5.06 g for fishes at 

maturity stages I, II, II and IV. Fazli (2007) and Fazli 

et al. (2007) studied this species from 1995 to 2004 

in Iranian waters, sampled at landing sites at 

Amirabad and Babol-Sar in Mazandaran and Anzali 

in Gilan. Growth parameters were L∞=132 mm, 

K=0.259/yr. t0=-1.285/yr. The instantaneous 

coefficient of natural mortality was 0.506/yr, the 

instantaneous coefficient of total mortality (Z) was 

1.62/yr and the instantaneous coefficient of fishing 

mortality varied over 10 years from 0.125/yr to 

1.487/yr. Annual survival rate (S) was 0.2/yr. The 

exploitation rate was 0.092-0.64 and the acceptable 

biological catch was 7,450 t. Age at first capture was 

2.8 years. Ages ranged from 1 to 7 years with age 

groups 2, 3 and 4 dominating at different periods. 

Mean fork lengths were 59.3, 77.5, 87.4, 97.2, 104.5, 

111.9 and 116.8 mm. Females dominated in each 

month except April, averaging 0.47:1, possibly due 

to differing attraction to lights used in the fishery. 

Biomass increased from 16,000 t in 1995 to more 

than 41,000 t in 2002, declining to less than 28,000 t 

in 2004. The increase was simultaneous with a sharp 

decline in anchovy kilka, changes in zooplankton 

composition and abundance, and especially an 

increase in zooplankton species favoured by this 

kilka. This kilka was overfished. Janbaz and 

Abdolmalaki (2009) found this species comprised 

over 86% of the total catch at Babol-Sar and 

Amirabad in Mazandaran, and age groups were 1+ to 

6+ with age class 4 predominating at 35.3%. von 

Bertalanffy growth parameters were L∞=114.92mm, 

K=0.339/yr, t0=-1.878/yr, Lt=114.92(1-e
0.339 

(t+1.878)), total mortality (Z)=0.84/yr, natural 

mortality (M)=0.41/yr, fishing mortality (F)=0.44/yr 

and exploitation rate (E)=0.51. Karimzadeh et al. 

(2010) and Karimzadeh (2011a) examined fish from 

the Babol-Sar region and calculated growth 

parameters as L∞=143.5mm, K=0.30/yr
-1 and t0=-

1.02/yr, instantaneous coefficient of natural 

mortality was 0.671 yr-1, fishing mortality was 0.849 

yr-1, and the current exploitation rate was estimated 

as 0.55 and this species is now overfished. Akbari 

Miarkolae et al. (2014) examined 175 fish from 

Babol-Sar and found a b value of 2.71 and an average 
condition factor of 2.33 with maximum and 

minimum values in January and June respectively. 

Condition factors were significantly different in all 

months studied and greater in winter than spring. An 

increased mean length and decreased mean weight in 

recent years was attributed to the ctenophore 

invasion limiting access to zooplankton. 

Aliasghari et al. (2012) examined 3,774 fish from 

Mazandaran and found a b value of 2.92, growth 
parameters were L∞=128.7  mm, K=0.41/yr, t0=-

0.59/yr, age groups were 1-6 years, average age was 

3.27 years, 3-year-old fish dominated (45.24%) of 

the catch), male:female sex ratio was 1:0.779, 

survival rate was 0.239/yr, natural mortality was 

0.448/yr, fishing mortality 0.983/yr, and exploitation 

rate was 0.687 indicating overfishing. The mean 

length and weight of fish had increased but mean age 

had decreased in recent years.  

Moradinasab et al. (2012) studied fish caught in 

lift nets in Gilan and found b values of 2.37 in males 
and 2.57 in females (both negatively allometric 

growth) with variations between sexes and seasons. 



158 
 

Coad/ Herrings of Iran 

Khedmati Bazkiaei et al. (2013) studied 547 fish 

from the Gilan coast and found most fish were 4-5 

years old, older fish dominated and fewer younger 

fish were being caught, the growth equation was 

Lt=142.1(1-exp
-0.248(t+1.63)), natural mortality (M) 

was 0.506/yr, fishing mortality (F) was 0.994/yr and 

total mortality (Z) was 1.5/yr.  

Food: Plankton is the main food and copepods 

predominate but diet also includes Cladocera, 

Balanus larvae and clam larvae. The dominant food 
item is the copepod Eurytemora grimmi, particularly 
in winter when plankton biomass is lowered in the 

Bol'shoy Kyzylagach Bay of Azerbaijan. The food 

of the common kilka is more varied than the other 

kilka species simply because of its habitat in shallow 

coastal areas (Badalov, 1972; Prikhod'ko, 1979b). 

Older fish take larger and faster crustaceans and 

consume less food in proportion to body size as they 

grow. The most intensive feeding is in summer and 

autumn, decreasing in winter and during 

reproduction. Food is taken during the day.  

Roushan Tabari et al. (2009) examined fish from 

fishing vessels of Mazandaran and found highest 

feeding activity in April with 280±153 prey items 

per fish weighing 2.9±1.6 mg. Balanus nauplii and 
cypris larvae comprised 93% and Acartia 7% at this 
time with increasing spring temperatures and 

reproduction, but the copepod Acartia biomass 
dominated from October to February. Khedmati 

Bazkiaei et al. (2014) found fish from the Anzali 

shore had copepods, Balanus cypris and larvae, 
cladocerans, and Azolla in their gut, dominated by 
copepods at 94.5% in number. 

Reproduction: Spawning occurs in January-February 

in the southern Caspian, later in the north, mainly in 

depths less than 10m and where salinity is low to 

average for the Caspian Sea (Badalov, 1972; 

Prikhod'ko, 1979b). The largest southern Caspian 

population spawns near the mouths of the Volga and 

Ural rivers (Kozlovsky in Hoestlandt, 1991). 

Spawning is most intensive at 11°C, but occurs at 10-

20°C. Spawning is intermittent and lasts from mid-

April to July.  

Peak spawning in Iranian waters of Mazandaran 

Province is April-May with an average fecundity of 

28,240 eggs (Abtahi et al., 2002). Fazli et al. (2006) 

recorded mass spawning in Iranian waters in April, 

continuing on until August. Eggs are released in 

water 0.5-9.0 m deep at a salinity range of 0.02-15‰, 

perhaps as high as 29.15‰. Fecundity reaches 

60,000 eggs and egg diameter 1.0 mm, 0.48-1.46 mm 

for fertilised eggs. Relative fertility is 4-13 times 

greater than in Alosa species. Holčík and Oláh 
(1992) consider that it may spawn in rivers entering 

the Anzali Talab.  

The studies of Fazli et al. (2006, 2007) showed 

that reproduction started in March, peaked in May 

and finished at the end of August. Half the females 

were mature at 84.3 mm fork length. Aliasghari et al. 

(2012) found from gonadosomatic indices and 

sexual maturity stages that spawning in Iran began in 

February and peaked in May and June. Karimi et al. 

(2013) examined 800 fish from the Anzali coast and 

found batch spawning with group-synchronous 

oocyte development. Maximum gonadosomatic 

indices were found in March-April and May in 

different years suggesting temporal variation in 

spawning. Estimated relative batch fecundity was 

6,718 eggs/g, with a range from 3,646 to 10,198 

eggs. A low level of atresia (4.7%) during spawning 

showed that this species was in optimal condition. 

Female common kilka were in better condition in 

2008 than in 1994-1996. The invasive ctenophore 

M. leidyi did not appear to affect competitively the 
somatic condition of this kilka and overfishing is the 

probable cause of decline in numbers caught. 

Khedmati Bazkiaei et al. (2013) studied fish from the 

Gilan coast and found the highest gonadosomatic 

indices in April-May and the lowest in November-

December.  

Aliasghari and Parafkandeh Haghighi (2013) 

examined 3,774 fish caught in 2010 from 

Mazandaran and found a male:female sex ratio of 

1:0.779, the sex ratio varied between months 

(females appear to be less attracted to fishing lights 

with development of gonads as they cease feeding), 

and spawning began in March and ended in July with 

a peak in May.  



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Parasites and predators: Samples of this species from 

Babol Sar and Bandar Anzali contained the digenean 

parasites P. symmetrica (probably P. ventricosa after 
Youssefi et al. (2011)) and B. cingulata, the 
acanthocephalan C. strumosum, metacercariae of a 
Bucephalus species, and larvae of a Contracaecum 
and an Anisakis species (Iranian Fisheries Research 
and Training Organization Newsletter, 11: 4-5, 1996, 

Annual Report, 1995-1996; Iranian Fisheries 
Research and Training Organization, p. 28, 1997; 
Shamsi and Dalimi, 1996; Shamsi et al., 1998). 

Ghayoumi (2009) used the helminth parasites 

C. strumosum and P. ventricosa as bioindicators of 
the heavy metals lead and cadmium. Ghayoumi et al. 

(2009) found that fish from Babol-Sar harbour 

contained the intestinal helminths C. strumosum, 
P. ventricosa, Contracaecum sp. larvae and 
Raphidascaris sp. larvae, diet being the main factor 
affecting diversity of the parasites. Varshoie et al. 

(2010) record the helminths Pseudopentagramma 
symmetrica (see above), B. cingulata, Mazocreas 
alosae, C. strumosum, Contracaecum sp. and 
Anisakis sp. in this species from Iranian waters. 

Clupeonella species are an important food fish for 
sturgeons (59.4% by weight of Acipenser stellatus 
diet in the Middle Caspian), Sander, herrings 
(Clupeidae) and the Caspian seal (Badalov, 1972; 

Krylov, 1984) as well as Salmo caspius and Stenodus 
leucichthys (Kosarev and Yablonskaya, 1994).  
Economic importance: It is caught by attraction to 

underwater electrical lights (Prikhod'ko, 1979b) and 

in school seines in spring and purse seines in 

summer. In Gilan, fish are caught at 12-43 m depths 

by lift nets equipped with underwater lights 

(Moradinasab et al., 2012). In Iranian waters, this 

species formed only a small proportion (1.35%) of 

the total kilka catch in a study by Razavi Sayad 

(1993) and Fazli et al. (2006) gives values of 1.34%, 

2.5% and 5.5% for the years 1990-91, 1997-98 and 

1998-99, respectively. However, as the anchovy 

kilka catch declined, this species increased from 

13.7% of the total catch in 1999 to 48.9% in 2003 

(Sayyad Bourani et al., 2008).  

Jorjani et al. (2014) detailed the chemical 

composition and fatty acid profile of this species 

from specimens landed at Anzali. This species was 

rich in essential unsaturated fatty acids of the omega-

3 family especially docosahexanoid acid which is 

nutritionally attractive for humans. Catches should 

be used for human consumption rather than as fish 

meal. 

Gasan-kuli or Hasan Kuli is a town in 

Turkmenistan near the Iranian border referred to in 

fishery reports from this area. The catch of Rutilus 
caspicus, C. carpio and Sander marinum was nearly 
1.44 x 104 tonnes with only 1.9% being accounted 

for by C. cultriventris (= caspia). However, by 1972 
the catch of the commercially important species had 

declined to 1.5% and the less desirable Clupeonella 
had increased to 5.73 x 104 t or 98.3% of the catch. 

The causes were reduction in the Atrak runoff 

through irrigation withdrawals, pollution from 

agriculture, overfishing in the sea and the drop in sea 

level. Flows of the Atrak did not reach the sea in 

1984, 1986, 1990 and 1991and spawning of species 

using the lower reaches did not occur (Caspian 

Environmental Programme, 2000). 

Ebadi and Shokrzadeh (2006) examined 

C. delicatula (= caspia) for lindane at Chalus, Babol 
Sar, Khazarabad and Miankaleh but levels detected 

were less than the FAO/WHO recommended 

permissible intake and were no cause for public 

concern. Similar studies on D.D.T. and D.D.E. and 

on chlorobenzilate from the same sites found levels 

were also less than the permissible intake 

(Shokrzadeh and Ebadi, 2005, 2006). Shokrazadeh 

et al. (2009) also found that levels of Lindane in 

dorsal muscle of kilka species were less than 

FAO/WHO recommended intake. Shokrzadeh 

Lamuki et al. (2012) evaluated residues of D.D.T. 

and D.D.A. in fish from localities in Mazandaran and 

found values (0.017-0.025 mg/kg D.D.T. and 0.019-

0.026 mg/kg D.D.A.) in the middle of the range of 

other species, cyprinids and a mullet, and varying 

between localities. Iraj et al. (2014) examined 

concentrations of chromium, copper, nickel and zinc 

in fish from the southern Caspian Sea and found 

levels of chromium and zinc exceeded international 



160 
 

Coad/ Herrings of Iran 

standards. Najm et al. (2014) determined the 

concentration of the heavy metals cadmium, 

chromium and lead in fish from the Babol-Sar coast 

and found cadmium levels exceeded international 

standards for human consumption although a risk 

assessment concluded the fish were safe for 

consumers. 

Ojagh et al. (2004) showed that natural 

antioxidants beta-carotene with ascorbic acid and 

green tea polyphenols were useful in preservation 

during storage with ice powder. Naseri et al. (2005) 

found that a fluorescence detection method was a 

good technique for assessing degradation in steam-

cooked kilka. Naseri et al. (2006) also used 

fluorescence detection to compare the quality of oil 

and brine filling material in canned kilka. Naseri et 

al. (2009) studied chilling times on quality of the 

canned product but deterioration was not well 

shown. Fluorescence detection of interaction 

compounds provided a good technique to assess 

product quality. Naseri et al. (2010) studied lipid 

changes in canned kilka after long-term storage. 

Motalebi and Seyfzadeh (2011) found that a 20% 

whey protein coating on kilka frozen at -18ºC 

maintained quality for 6 months while control 

samples lost their quality after 3 months. Zamani et 

al. (2012) found that belly bursting (enzymatic 

decomposition leading to unmarketable fish) can be 

prevented by trypsin inactivation through 

application of low temperature (4ºC), acidic ph, 

certain metals (aluminium, copper and zinc ions) and 

trypsin inhibitors. Dorvaj et al. (2013) showed that 

protein hydrolysates from this kilka can be used as 

an alternative substrate for the culture of 

microorganisms. Khanipour et al. (2014) found that 

the shelf life of breaded kilka with tempura batter 

was about 3 months at -18ºC. Navidshad (2014) used 

an oil supplement from this fish in combination with 

soybean oil in chicken food to enrich the product. 

Sahari et al. (2014) showed loss of certain vitamins 

in fish stored at -24ºC including A, D and K and 

significant losses in C and E vitamins. Dehbandi et 

al. (2015) found that encapsulated nisin z 

(a polycyclic antibacterial and antioxidant peptide) 

in liposome improved shelf life of kilka surimi. 

Robins et al. (1991) list this species as important to 

North Americans. Importance is based on its use as 

food and as bait.  

Conservation: Stocks on the Iranian coast are said to 

have been depleted but its ecological specialisation 

on zooplankton means there is comparatively little 

competition with other fishes. It is probably not in 

any immediate danger. Kiabi et al. (1999) consider 

this species to be of least concern in the south 

Caspian Sea basin according to IUCN criteria. 

Criteria include commercial fishing, abundant in 

numbers, widespread range (75% of water bodies), 

absent in other water bodies in Iran, and present 

outside the Caspian Sea basin.  

Sources: Iranian material: CMNFI 1970-0531, 14, 

78.0-88.6 mm standard length, Mazandaran, Larim 

River (36º46'N, 52º58'E); CMNFI 1980-0146, 7, 

79.9-96.2 mm standard length, Mazandaran, Gorgan 

Bay at Ashuradeh-ye Kuchak (36º50'N, 53º56'E); 

CMNFI 1993-0146, 3, 80.2-98.2 mm standard 

length, Mazandaran, Gorgan Bay (no other locality 

data); CMNFI 1993-0167, 1, 96.6 mm standard 

length, Mazandaran, Caspian Sea, 10 km offshore 

(ca. 36º49'N, ca. 52º39'E); CMNFI 1993-0168, 3, 

84.9-88.0 mm standard length, Mazandaran, Caspian 

Sea, 10 km offshore (ca. 36º49'N, ca. 52º39'E). 

 

Clupeonella engrauliformis (Borodin, 1904) 
(Fig. 12) 

Common names: Rizeh keraye, kilka-ye anchovy or 

kilka-e-anchovi. [Ancousabanzar kilka in 

Azerbaijan; ancous sekilli kulke balyk in 

Turkmenistan; anchousovidnaya tyul'ka or anchovy-

like tyulka, sardelle or sardel'ka, "sardinka" but 

incorrectly, all in Russian; anchovy kilka, anchovy 

sprat]. 

Systematics: No major synonyms. Originally 

described from Buinak, central part of the Caspian 

Sea. The lectotype is in the Zoological Institute, St. 

Petersburg (ZISP 13860) with paralectotypes as 

established by Svetovidov (1952) (Eschmeyer et al., 

1996). Eschmeyer et al. (1996) gave the date as 1906 

but Reshetnikov et al. (1997) and the online Catalog 



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Int. J. Aquat. Biol. (2017) 5(3): 128-192 

 

of Fishes give 1904.  

Key characters: This species has a slender body 

(body depth 16-19% of standard length), a short and 

wide head (interorbital width 16.0-18.5% of head 

length), a rounded belly with keel scales weakly 

developed, and pointed pectoral fin tips.  

Morphology: Dorsal fin with 3 unbranched and 12-

14 branched rays, anal fin with 3 unbranched and 15-

19 branched rays. Scales in lateral series 45-49. 

Vertebrae 44-47, rarely to 48 compared to 40-45 in 

the common kilka (C. caspia). Gill rakers number 
56-67. Belly scutes 23-31.  

Sexual dimorphism: None reported.  

Colour: The back and head are dark blue with violet, 

green or olive tints. These colours become brighter 

or turn black in dead fish. The fins are hyaline except 

the caudal fin which has a black base and the dorsal 

fin which has a central dark stripe.  

Size: Attains 15.5 cm standard length.  

Distribution: Found in the central and southern 

Caspian Sea, and in Iranian waters the southeast 

Caspian Sea, southwest Caspian Sea and the south-

central Caspian Sea (Kiabi et al., 1999) as well as the 

Anzali Talab, Babol-Sar Beach and Gorgan Bay 

(Armantrout, 1980; Abdoli and Naderi, 2009). 

Zoogeography: This species is endemic to the 

Caspian Sea.  

Habitat: The anchovy kilka, along with other kilkas, 

is the most abundant fish in the Caspian Sea forming 

large concentrations in the central and southern 

Caspian wherever water depth exceeds 30m. The 

anchovy kilka is estimated to be the most numerous 

kilka at about 77% (Ivanov and Katunin, 2001; 

Daskalov and Mamedov, 2007). It is generally found 

in the upper water layers but may descend to 120 m. 

Nearshore areas, inlets and water of salinity below 

8‰ are avoided. They can tolerate a salinity range of 

8-14‰ but the main part of the population is found 

at 10-12‰ (Fazli et al., 2007). Overwintering takes 

place in the southern Caspian and the southern part 

of the central Caspian Sea at 8.5-9.0°C and up to 

13.5°C. Schools extend their range into the central 

and northern Caspian in spring to feed (Prikhod'ko, 

1979b). This species has a hibernation period in the 

south Caspian Sea, a spring migration of part of the 

population to the central Caspian, a feeding period in 

the central and south Caspian and an autumn 

prespawning migration to the south Caspian (Sedov 

and Rychagova, 1983).  

In Iran, larvae are found mostly in surface layers 

at 5-20 m while adults are found in deeper zones. 

Males dominate in winter while females dominate in 

other seasons. The maximum juvenile density (fish 

<75 mm), comprising 36% of the population, is seen 

in the summer (Iranian Fisheries Research and 

Training Organization Newsletter, 20: 7, 1998). 

Jolodar and Abdoli (2004) state it is most abundant 

at 100-150 m. Zare et al. (2017) investigated acoustic 

target strength in Iranian waters for calculating 

biomass, stock assessment and management 

strategies. Mean target strength was correlated with 

indices of female maturity status and depth of fish 

occupancy, but not size. There was a high variability 

in mean target strength estimates despite a small 

range in fish size (3.2 cm). Biological characteristics 

other than size were important drivers of this 

Figure 12. Line drawing of Clupeonella engrauliformis. 



162 
 

Coad/ Herrings of Iran 

variation. 

Age and growth: Abundance of young anchovy 

kilka, and hence future year-class strengths, depends 

on water temperature in autumn (October-

November). Falling water temperatures, in the 

eastern Caspian for example, are caused by 

upwelling which brings nutrients to surface waters 

and promotes growth of plankton on which the kilka 

larvae feed (Prikhod'ko, 1979a). Females are 

somewhat larger than males in the spawning areas. 

Sexual maturity is attained usually at age 2 and 2-4 

year olds dominate catches but life span is up to 8 

years (Iranian Fisheries Research and Training 

Organization Newsletter, 14: 6, 1996). This species 

shows the fastest rate of growth in the genus. Of the 

8 age classes, 0+, 1+, 2+ and 3+ form 99.91% of the 

whole population (Iranian Fisheries Research and 

Training Organization Newsletter, 20: 7, 1998). The 

same study showed that 18.6% of the population 

matures in the first year of life while 81% matures in 

the second. The mean age in coastal areas is 2.9 

years, slightly higher than that in deep zones below 

200 m where 0+ fish are more abundant. The 

Caspian Sea Biodiversity Database (from 

www.caspian environment.org) gives a population 

of up to 293 billion fish in the Caspian Sea.  

Fazli et al. (2007) and Sayyad Bourani et al. 

(2008) studied these kilkas from catches with conical 

liftnets carrying underwater lights in the fisheries of 

Gilan and Mazandaran in the 1995-2004 period. Fish 

were aged using the sagittal otoliths. Length and 

weight ranges were 40-140 mm and 0.4-18.4 g with 

averages of 94.0 mm and 5.7 g (89.2-100.4 mm from 

1999 to 2003 in Sayyad Bourani et al. 2008). The age 

range was 1-7 years (rarely to 8+ years in 

Parafkandeh Haghighi and Kaymaram (2012)). The 

dominant age group varied from age 2 to age 4, 

making up 40.6% to 57.7% of the catch (Fazli et al., 

2007) or 5+ years with 4+-5+ making up 84.6% for 

1999-2003 (Sayyad Bourani et al., 2008). Growth 

was high for the first year of life and then gradually 

decreased. The von Bertalanffy growth equation was 

Lt=148(1-e
-0.238(t+1.340)) (Fazli et al., 2007, and 

following data). The sex ratio varied with season and 

was significantly different from equal at 

male:female=0.78:1 for adults. Females were more 

abundant from January to June and males 

predominated from September to November. 

Condition factors differed significantly between 

years, increasing from 1995 to 1996, being lowest in 

1998 and then increasing to 2004, and between 

months, being lowest in January and February and 

then increasing in March. 50% of fish were mature 

at 84.5 mm fork length. Annual survival rate was 

estimated at 0.32, the instantaneous coefficient of 

total mortality (Z) was 1.14/year, and natural 

mortality was 0.473/year. Age at first capture was 

estimated as 2.92 years. The total biomass declined 

from 186,000 t in 1996 to less than 12,000 t in 2004 

and the exploitation rate for 1995-2004 varied 

between 0.34 and 0.815.  

Fazli (2007) examining fish for the 10-year period 

1995-2004 gave slightly different figures: estimated 

survival rate (S) was 0.174, fishing mortality was 

0.65-2.93/yr, biomass declined from 185,000 t in 

1996 to 8,320 t in 2004, exploitation rate was 0.39-

0.83, maximum sustainable yield (MSY) and fMSY 
were 44,652 t and 18,609 vessel x nights (a unit of 

effort), and the acceptable biological catch was 2,190 

t. Sayyad Bourani et al. (2008) give a K value of 
0.598/year and a L∞ of 110.13 mm. Natural, fishing 

and total mortality coefficients were 0.69, 0.31 and 1 

per year respectively and the sex ratio was 

female:male=68.2:31.8. These latter results for the 

1999-2003 period show how values can change 

when subsets of data are used.  

Karimzadeh et al. (2010) examined fish from the 

Babol-Sar region off Mazandaran and calculated 

growth parameters as L∞=151.9 mm, K=0.28/yr
-1 

and t0=-1.12/yr, instantaneous coefficient of natural 

mortality was 0.633/yr-1 and the current exploitation 

rate was estimated as 0.41. Parafkandeh Haghighi 

(2010) and Parafkandeh Haghighi and Kaymaram 

(2010) studied fish from Anzali and Babol-Sar in 

2006-2007 and found the mean length was 117.8 

mm, mean age was 4.5 (or 4.6) years, a male:female 

sex ratio of 0.52:1, and 80% of fish were in the 4 and 

5 years old age classes. Growth parameters were 



163 
 

Int. J. Aquat. Biol. (2017) 5(3): 128-192 

 
L∞=147.45 mm and t0=-1.73/yr, b=2.56 (negative 
allometric growth), K=0.245/yr, Z=1.067/yr, 

M=0.503/yr and F=0.564/yr and E=0.528/yr. 

Aliasghari et al. (2011) estimated growth parameters 

for fish from Babol-Sar and found an increase in 

mean length and weight over previous years 

attributed to weak stock construction consequent on 

the ctenophore invasion, L∞=160.5 mm, K=0.25/yr
-

1 and t0=-0.92/yr for males and L∞=138.7 mm, 

K=0.55/yr-1 and t0=-0.42/yr for females, b values for 
both sexes were 2.5, males had age groups 1-7 years 

and females 2-7 years, mean age of males was 3.86 

years and 4.58 for females, and four-year-old fish 

predominated at 26.55%. 

Fatemi et al. (2009) examined fish taken from 

commercial vessels in 2007 using lift nets and lights. 

Age structure ranged from 2 to 7 years and was 

dominated by the third year class (38.6%). Back-

calculation methods were validated using otoliths to 

determine lengths. Janbaz et al. (2012) gave values 

for Iran from 2005 to 2007 of K=0.375/year, 

L∞=131.7 mm, instantaneous coefficient of natural 

mortality was 0.49/year, fishing mortality was 

0.51/year and total mortality 1.0/year, and 

exploitation rate was 0.51. Abundance declined over 

the three years from 18.8% to 8.5% and to 6%, catch 

per unit effort declined from 0.3 to 0.1 t, and the 

condition factor declined. Overfishing and the 

competitive ctenophore were the main causes of the 

decline. Fazli et al. (2013) studied the status of stocks 

in Iranian waters from 1996 to 2011. The length 

range with optimum yield was 85-105 mm, mega-

spawners were larger than 100mm, the percentage of 

mature fish increased, optimum size declined since 

2004, the percentage of mature fish were more than 

78% (often more than 95%), and the length structure 

of this kilka stock is a matter for concern. 

Food: Plankton is the main food and copepods 

predominate but diet also includes Cladocera, 

Balanus larvae and clam larvae. The dominant food 
item is the copepod Eurytemora grimmi, particularly 
in winter when plankton biomass is lowered 

(Badalov, 1972). It can make up over 70% of its 

food. This copepod is more characteristic of the diet 

of this kilka compared to the other two species and 

the daily vertical migrations and seasonal 

movements of the copepod are mirrored by the kilka. 

The most abundant fish species in the Caspian 

depends on the most abundant member of the 

crustacean zooplankton (Prikhod'ko, 1979b). This 

species feeds in winter, unlike C. caspia. Intensive 
feeding begins in spring as a preparation for 

spawning (Sedov and Rychagova, 1983). Spawning 

males show a positive response to light and so feed 

during the spawning season, while females do not.  

F. Darvishi (pers. comm. 2003) has demonstrated 

that the this species has a similar feeding niche as the 

exotic ctenophore M. leidyi in Iran and Esmaili Sari 
et al. (2002) also determined that there is a similar 

diet suggesting that a decline in stocks of the fish is 

the result of competition. Darvishi (2003) and 

Darvishi et al. (2004) studied catches of the anchovy 

kilka and the ctenophore in the southern Caspian Sea 

from August 2001 to October 2002. Dietary overlap 

was >89 in Babol-Sar samples and >84 in Nowshahr 

samples using the Schoener Index (presumably 0.89 

and 0.84 where 0 is no dietary overlap and 1 is an 

identical diet). The ctenophore was also feeding on 

fish eggs but the effect of this was less than 

competition for food. Rahimi Bashar and Alipour 

(2009) listed a wide range of phytoplankton and 

zooplankton species in the diet of fish from the 

southwest Caspian Sea. 

Reproduction: Spawning ends in late autumn and 

winter food requirements are higher than in spring-

spawning C. caspia (Badalov, 1972). Areas for 
spawning in this species are extensive. Spawning is 

most intensive in July when temperatures are 13-

24°C and salinity 8-13‰ although the Caspian Sea 

Biodiversity Database (from www.caspian 

environment.org) gives peak spawning (70%) as in 

October-November. Fazli (2006) gives spawning in 

Iran as spring and autumn but mass spawning takes 

place in in autumn. Spawning takes place in the 

central and southern Caspian along both eastern and 

western shores both in coastal regions and the open 

sea from late April to November. Mass spawning 

takes place at depths of 50-200 m and as a result eggs 



164 
 

Coad/ Herrings of Iran 

and larvae are carried over a wide area by the 

Caspian gyral current at these depths (Prikhod'ko, 

1979b). Young hatch mainly in autumn and reach 

4.5-8.0 cm at an age of 8-10 months (Prikhod'ko, 

1979a). Eggs are up to 1.82 mm in diameter and 

fecundity reaches 39,900 eggs.  

In Iran, 80% of the population spawns in autumn and 

the remainder in spring. Accordingly the fishery 

should be closed in October and November (Iranian 

Fisheries Research and Training Organization 

Newsletter, 19: 5, 1998). The subsequent Iranian 

Fisheries Research and Training Organization 

Newsletter (20: 7, 1998) states that 89% of the 

population spawns in autumn with September, at 

68.3%, the major month. Fazli et al. (2007) found 

reproduction to start in June, peaking in October and 

then declining. Janbaz et al. (2012) found an absolute 

fecundity of 12,625 eggs in Iranian waters, more 

than 50% of fish were mature when length exceeded 

92.5 mm, and spawning was in autumn but 

overfishing and the ctenophore invasion pushed the 

kilka to spawn in winter which caused stocks to 

collapse. Year-round spawning occurred as a 

strategy to mitigate these problems. Aliasghari and 

Parafkandeh Haghighi (2013) examined 1,659 fish 

caught in 2010 from Mazandaran and found a 

male:female sex ratio of 0.569:1, the sex ratio varied 

between months (females appear to be less attracted 

to fishing lights with development of gonads as they 

cease feeding), and spawning was from April to 

November with a peak in November. 

Parasites and predators: Samples of this species from 

Babol Sar and Bandar Anzali contain the digenean 

trematode parasites P. symmetrica (probably 
P. ventricosa after Youssefi et al. (2011)) and 
B. cingulata, the acanthocephalan C. strumosum and 
larvae of the nematode Contracaecum sp. (Iranian 
Fisheries Research and Training Organization 

Newsletter, 11: 4-5, 1996; Shamsi et al., 1996b; 

Annual Report, 1995-1996; Iranian Fisheries 
Research and Training Organization, p. 28, 1997; 

Shamsi and Dalimi, 1996; Shamsi et al., 1998a, 

1998b). Ghayoumi (2009) used the helminth 

parasites C. strumosum and P. ventricosa as 

bioindicators of the heavy metals lead and cadmium. 

Ghayoumi et al. (2009) found that fish from Babol-

Sar harbour contained the intestinal helminths 

C. strumosum, P. ventricosa, Contracaecum sp. 
larvae and Raphidascaris sp. larvae, diet being the 
main factor affecting diversity of the parasites. 

Varshoie et al. (2010) record the helminths 

P. symmetrica (see above), B. cingulata, Mazocreas 
alosae, C. strumosum and Contracaecum sp. in this 
species from Iranian waters. 

Clupeonella species are an important food fish for 
sturgeons (59.4% by weight of sevryuga diet in the 

Middle Caspian), Sander (Percidae) and herrings and 
the Caspian seal (Badalov, 1972; Krylov, 1984) as 

well as other fishes. 

Economic importance: This species forms 80-90% 

of the catches of kilkas in former Soviet waters 

(Sedov and Rychagova, 1983) and, as noted above, 

91.8% of catches in an Iranian study (Razavi Sayad, 

1993; Rezaei et al., 2003). High catches are related 

to the larger spawning and foraging range of this 

species compared to other kilkas and to its habitat in 

the Caspian gyre, an area of increased biological 

productivity (Prikhod'ko, 1979b). It is caught in 

former Soviet waters by attraction to underwater 

electrical lights attached to the middle of the mouth 

of a fine-mesh conical net or the sides of a fish pump 

(Ben-Yami, 1976). Fishing is suspended at full 

moons as the fish are dispersed. Both large and small 

individuals are taken by these non-selective methods 

(Prikhod'ko, 1981). Incidental catches include 

Mugilidae (common), and Alosa spp., Atherinidae 
and the cyprinid Pelecus cultratus (all occasional) 
(Ben-Yami, 1976). The catch per unit effort for 

funnel nets and midwater trawls is 2,321 and 1,014 

respectively (Iranian Fisheries Research and 

Training Organization Newsletter, 20: 7, 1998). 

Fazli et al. (2009c) estimated the maximum 

sustainable yield at 44,652 t and 18,609 vessel x 

nights (as a unit of effort). The acceptable biological 

catch was estimated at 2,190 t in 2004 but in 2005 it 

was 4,300 t. In 2008, the catch declined to 220 t 

through overfishing combined with sea level 

changes, invasive species and pollution. In 1999, the 



165 
 

Int. J. Aquat. Biol. (2017) 5(3): 128-192 

 
catch was 67,450 t in Iranian waters. Janbaz et al. 

(2012) recorded a decline in catches from 4,250 t in 

2005 to 924 t in 2007. 

It is regarded as a valuable and cheap food 

resource in Iran where it is canned, made into 

sausages and surimi, and processed as fish meal 

(Shamsi et al., 1996; Moeini, 2002; Shabanpour et 

al., 2002, 2006). Ovissipour et al. (2103) showed that 

protein hydrolysates from this fish had high 

antioxidant activity and amino acid content and low 

heavy metal concentrations. This species has a high 

potential for use in animal and human diets.  

Various studies on its preparation and storage as 

food have been carried out, e.g. Rezaei et al. (2002, 

2003), Rezaeian et al. (2002), Rezaie et al. (2007) 

and Moeini et al. (2009). Shelf life in cold storage at 

-18ºC was 60 days for packed and 30 days for 

unpacked fish and at -30ºC lipid quality was higher. 

Moeeni et al. (2006) determined the proper 

formulation of soup powder and found that shelf life 

of the product was about 60 days. Sarhadi et al. 

(2012) and Sarhaddi et al. (2015) studied the 

proximate composition, amino acid profile, fatty 

acid and mineral composition of this species and 

found it to have good nutritional quality such that 

powdered bone may be used as an additive in food 

industries. Robins et al. (1991) list this species as 

important to North Americans. Importance is based 

on its use as food and as bait.  

Conservation: Prikhod'ko (1981) recommends 

fishing in deeper waters where larger fish are 

concentrated to avoid an excessive take of young fish 

which favour the upper water layers. Stocks in the 

southern Caspian Sea are said to be depleted. Kiabi 

et al. (1999) consider this species to be of least 

concern in the south Caspian Sea basin according to 

IUCN criteria. Criteria include commercial fishing, 

abundant in numbers, widespread range (75% of 

water bodies), absent in other water bodies in Iran, 

absent outside the Caspian Sea basin. Daskalov and 

Mamedov (2007) studied commercial catch data in 

the Caspian Sea generally and found a period of high 

catches from 1991 to 2000 with high spawning-stock 

biomass and relatively good recruitment. Catches 

peaked at 271,400 t, fishing mortality reached 1.8y-1 

in 1999 and overfishing occurred. From 2001 to 

2004, the stock collapsed, recruitment failed in 2001 

and catches fell to 54,300 t in 2005, of which the 

Iranian fishery was about 8%. This was attributed to 

the spread of the ctenophore M. leidyi, with 
contributions from overfishing. Fazli et al. (2007) 

also concur that both overfishing and the invasive 

ctenophore caused the collapse of stocks. The catch 

in Iran declined from 71% of the total kilka catch in 

1999 to 52.5% in 2003 (Sayyad Bourani et al., 2008). 

Janbaz et al. (2012) give figures for the collapse in 

Iran from 4,250 t in 2005 to 924 t in 2007. 

Sources: Iranian material: CMNFI 1993-0167, 1, 

99.5 mm standard length, Mazandaran, Caspian Sea 

(ca. 36º49'N, ca. 52º39'E); CMNFI 1993-0168, 4, 

89.3-107.6 mm standard length, Mazandaran, 

Caspian Sea (ca. 36º49'N, ca. 52º39'E).  

 

Clupeonella grimmi Kessler, 1877  
(Fig. 13) 

Common names: Kilka-ye cheshmdorosht (= bigeye 

kilka). [Irikoz kilka in Azerbaijan; sardelle or 

sardel'ka, bol'sheglazaya tyul'ka or bigeye tyulka, 

bol'sheglazaya kil'ka or bigeye kilka, all in Russian; 

bigeye kilka, southern Caspian sprat].  

Systematics: Clupeonella Grimmi was originally 
described from the central part of the Caspian Sea. 

The lectotype is in the Zoological Institute, St. 

Petersburg under ZISP 10934 as designated by 

Svetovidov (1952). Harengula macrophthalma 
Knipovich, 1921 is a synonym. Four syntypes are in 

the Natural History Museum, London under 

BM(NH) 1897.7.5:41-44 (when examined were 

numbered 42-44, 3 fish, 29.9-33.5 mm standard 

length in poor condition, September 2007), with 

many others apparently in the Zoological Institute, 

St. Petersburg (Eschmeyer et al., 1996).  

Key characters: This species has a moderately 

slender body (17-22% of standard length), a long and 

narrow head (interorbital width 13.0-15.5% of head 

length), a sharply keeled belly, and rounded pectoral 

fin tips.  

Morphology: Dorsal fin unbranched rays 3-4, 



166 
 

Coad/ Herrings of Iran 

usually 3, branched rays 13-15, and anal fin 

unbranched rays 3, branched rays 14-21. There are 

44-49, usually 46-48, vertebrae, more than in the 

other two kilka species and probably a consequence 

of the low water temperature larvae develop in. Belly 

with 26-32 scutes. Gill rakers 42-51.  

The bigeye kilka is adapted to life in deeper water 

having, as its name indicates, big eyes with more rod 

cells and a weaker retina but also more transparent 

body tissues than other kilkas. 

Sexual dimorphism: None reported except size. 

Colour: The back and top of the head are dark.  

Size: Reaches 14.5 cm standard length and 15.2 cm 

fork length (Karimzadeh, 2011b).  

Distribution: Found in the Caspian Sea and 

concentrated in the south including Iranian waters. 

Abdoli and Naderi (2009) list it as from the 

southwest, southeast and south-central Caspian Sea 

in Iranian waters. 

Zoogeography: This species is endemic to the 

Caspian Sea.  

Habitat: The bigeye kilka is found further away from 

the coast than the anchovy kilka at depths over 50-

70 m, down to 450 m, with large schools down to 

130m. It does not enter fresh water or low salinity 

areas, staying well away from the shore. There is a 

daily vertical migration, avoiding sunlight, and 

following food items. Larvae live in water 

temperatures of 5°C. Overwintering occurs in the 

southern Caspian at temperatures of 9-11°C, a 

migration to the central Caspian takes place in 

spring, with a return south in autumn (Prikhod'ko, 

1979b).  

Age and growth: Sexual maturity is attained usually 

at age 2, and 2-4 year olds dominate catches, but life 

span is up to 8 years (Iranian Fisheries Research and 

Training Organization Newsletter, 14: 6, 1996). The 

female is larger than the male at the same age. 

Growth is slower than in C. engrauliformis. Males 
dominate the population (Iranian Fisheries Research 

and Training Organization Newsletter, 14: 6, 1996; 

Fazli et al., 2005) but this study may have sampled 

spawning fish (see below).  

Fazli et al. (2005) examined fish from the main 

landing ports (Babol-Sar, Amirabad and Anzali) and 

found the mean fork length of fish increased from 

95.87 mm in 1997 to 105.0 mm in 2000 but then 

decreased to 102.3 mm afterwards. Over this time 

period, fork length range became wider with 

specimens in the upper length classes representing 

most of the catch. Six age classes were present, 1+ to 

6+ years. During 1998-1999, age classes 1+ to 3+ 

comprised more than 90% of the catch. In 2000, 

there was a decrease in age classes 1+ and 2+ and an 

increase in 3+ to 5+ classes. In 2001, age classes 3+ 

and 4+ decreased and classes 5+ and 6+ increased. 

The relative frequency of the bigeye kilka has 

decreased in recent years as a result of the 

introduction of the ctenophore, M. leidyi, a food 
competitor and predator on kilka eggs and young. 

Janbaz et al. (2016) examined 262 fish from 

Babol-Sar, Amirabad and Anzali stations caught 

between May 2006 and April 2007. The largest 

length-groups were found from December to 

Figure 13. Line drawing of Clupeonella grimmi. 



167 
 

Int. J. Aquat. Biol. (2017) 5(3): 128-192 

 
February and the lowest in April, coinciding with 

spawning and feeding during wintering respectively. 

The male:female sex ratio was 0.33:1 which differed 

significantly from the expected ratio of 1:1. The 

length-weight regression was W= 0.0000744 FL3.14 

for females and W=0.0000341FL3.16 for males, 

indicating isometric growth. Age based on otolith 

readings showed that the population was composed 

of age groups from 2 to 7 years old with rapid growth 

during the second year. Four-year-olds with a mean 

fork length and weight 115.5 mm and 11.9 g were 

the most abundant at 40.1%. The condition factor 

was 0.7-0.8 which varied in different months and 

coincided to the gonad development. The K value 
and L∞ of females was more than that of males. This 

was a rapidly-growing species in the southern parts 

of the Caspian Sea. 

Khorashadizadeh et al. (2006) found fish in the 

Babol-Sar area off Mazandaran to have 5 age classes, 

dominated by the 4+ class. Karimzadeh et al. (2010a) 

examined fish from Babol-Sar and calculated growth 

parameters as L∞=148.6 mm, K=0.46/yr
-1 and t0=-

0.18/yr, instantaneous coefficient of natural 

mortality was 0.881/yr-1 and the current exploitation 

rate was estimated as 0.26. Karimzadeh (2011b) 

determined mortalities for these Babol-Sar fish as 

natural and fishing mortality 0.881 and 0.309/yr 

respectively, annual survival rate 0.304/yr, 

instantaneous coefficient of total mortality 1.19/yr 

and exploitation rate 0.26/yr. Age groups were 2-6 

years with ages 3 and 4 comprising 72% of the catch. 

Average age of females was 4.194 years and males 

3.347 years. Catches decreased from overfishing, 

predators and ecological disturbances. Aliasghari et 

al. (2011) also examined fish from Babol-Sar 

numbering 1,043 specimens. Growth parameters 

were L∞=149.7 mm, K=0.32/yr
-1 and t0=-1.2/yr for 

females and L∞=136.5mm, K=0.65/yr
-1 and t0=-

0.24/yr for males. b values were 3.043 in females and 
3.079 in males, both positively allometric. The fish 

were aged at 2-7 years with mean age 4.09 for 

females and 3.61 for males. Four-year-old females 

dominated at 33.34% and three-year-olds in males at 

47.45%. The male:female sex ratio was 0.656:1. This 

kilka had not been affected by pelagic changes in the 

Caspian Sea since it inhabits deeper waters. In recent 

years the population became older and length and 

weight increased as Karimzadeh (2011b) noted also. 

This was attributed to use of nets with a standard 

mesh, overfishing and change in living depth.  

Fazli (2007) examining fish for the 10-year 

period1995-2004 gave von Bertalanffy parameters 

L∞=144 mm, K=0.265/yr, t0=-1.422/yr, estimated 

survival rate (S) was 0.298, natural mortality (M) 

was 0.46/yr, fishing mortality was 0.46-0.98/yr, 

biomass increased from 36,000 t in 1995 to more 

than 52,000 t in 1998 and declined to less than 5,900t 

in 2001, exploitation rate was 0.3-0.52, and the 

acceptable biological catch was 2,210 t. Fazli et al. 

(2009) examined changes in the population biology 

of this kilka over the period 1995 to 2001, attributed 

to the invasive ctenophore. The overall sex ratio was 

1.65:1 in favour of males, length-weight regressions 

were W=0.00922L2.851 for females and W=0.008021 

L2.907 for males, indicating a negative growth for 

both sexes, growth parameters were L∞=142 mm, 

K=0.28 year-1, and t0=-1.39 years, the instantaneous 

coefficient of natural mortality was 0.460 year-1, and 

the instantaneous coefficient of fishing mortality 

varied between 0.469 and 0.980 year-1. Biomass 

increased from 36,900 t in 1995 to more than 53,500 

t in 1998 but declined to less than 5,900t in 2001. 

This was attributed to overfishing and the 

appearance of the ctenophore, a competitor for 

zooplankton food.  

Parafkandeh Haghighi (2010) studied fish in 

2006-2007 and found the mean length was 119.5 

mm, mean age was 4.6 years, a male:female sex ratio 

of 1.6:1, growth was negative allometric, 

K=0.267/yr, Z=1.015/yr, M=0.537/yr and 

F=0.478/yr and E=0.471/yr. 

Fazli et al. (2013) studied the status of stocks in 

Iranian waters from 1996 to 2011. The length range 

with optimum yield was 90-110 mm, mega-spawners 

were larger than 110 mm, the percentage of mature 

fish increased, optimum size declined since 2002, the 

percentage of mature fish were more than 78% (often 

more than 95%), and the length structure of this kilka 



168 
 

Coad/ Herrings of Iran 

stock is a matter for concern.  

Food: Migratory mysids often predominate in the 

planktonic diet of this species. Fish fry are also eaten. 

Its foods are less diverse than that of other kilkas 

because the variety is less in the deeper waters this 

fish inhabits during the day. The three kilkas share 

the available habitat and its foods, the common kilka 

in shallow, coastal waters, the anchovy kilka in the 

upper layers of the open sea and the bigeye kilka in 

deeper water of the open sea (Badalov, 1972; 

Prikhod'ko, 1979b).  

Reproduction: Spawning is extended, from January 

through to September but is most intense in spring 

and autumn (Prikhod'ko, 1979b). Males predominate 

in the spawning areas, remaining there while females 

leave immediately after spawning. Males are mainly 

at 10-20 m and females at 20-25 m during the 

spawning season. Water temperatures are at 6-13°C 

and salinity 12.6-13.0‰. Fecundity is 28,300 eggs. 

In Iranian waters, mature fish ready to spawn are 

always present in catches in winter and early spring 

(Fazli et al., 2005). Khorashadizadeh et al. (2006) 

found fish in the Babol-Sar area of the Iranian coast 

to have peak spawning in early January. Aliasghari 

and Parafkandeh Haghighi (2013) examined 1,034 

fish caught in 2010 from Mazandaran and found a 

male:female sex ratio of 0.656:1, the sex ratio varied 

between months (females appear to be less attracted 

to fishing lights with development of gonads as they 

cease feeding), and spawning occurred year round 

with a peak in October. 

Parasites and predators: Samples of this species from 

Babol Sar and Bandar Anzali contain the digenean 

parasites P. symmetrica (probably P. ventricosa after 
Youssefi et al. (2011)), B. cingulata, the 
acanthocephalan C. strumosum, Eustrongylides 
excisus, and larvae of a Contracaecum and an 
Anisakis species (Iranian Fisheries Research and 
Training Organization Newsletter, 11: 4-5, 1996, 

Annual Report, 1995-1996; Iranian Fisheries 
Research and Training Organization, p. 28, 1997; 

Shamsi and Dalimi, 1996; Shamsi et al., 1998, 

1998a). Ghayoumi (2009) used the helminth 

parasites C. strumosum and P. ventricosa as 

bioindicators of the heavy metals lead and cadmium. 

Ghayoumi et al. (2009) found that fish from Babol-

Sar harbour contained the intestinal helminths 

C. strumosum, P. ventricosa, Contracaecum sp. 
larvae and Anisakis sp. larvae, diet being the main 
factor affecting diversity of the parasites. Varshoie et 

al. (2010) record the helminths P. symmetrica (see 
above), B. cingulata, Mazocreas alosae, 
C. strumosum, Contracaecum sp. and Anisakis sp.in 
this species from Iranian waters. 

Clupeonella species are an important food fish for 
sturgeons (59.4% by weight of sevryuga (Acipenser 
stellatus) diet in the Middle Caspian), Sander 
(Percidae) and herrings and the Caspian seal. 

Predators consume 590 million kg of the three kilka 

species which themselves are the main consumers of 

zooplankton. Kilkas are a very important element in 

the life of the Caspian Sea (Badalov, 1972; 

Prikhod'ko, 1979b; Krylov, 1984). This species is 

taken to a lesser extent than other Clupeonella 
species because it is relatively sparse.  

Economic importance: The bigeye kilka catch 

amounts to about 70 million kg a year in former 

Soviet waters of the Caspian by means of electric 

light. All three kilka species are caught by using 

underwater electric lights and fish pumps 

(Nikonorov, 1964) but in the case of the bigeye the 

effect is avoidance used to drive it to the bottom 

where it can be caught. Other kilkas are attracted to 

the light but the bigeye is a vertical migrator, 

avoiding sunlight (Prikhod'ko, 1979b). Light-

assisted catches of kilkas damages young shad 

(Alosa) stocks which are an incidental catch 
(Zakharyan and Teruni, 1979). Catches in Iranian 

waters are only 6.84% of the total kilka take (Razavi 

Sayad, 1993). The relative frequency of the bigeye 

kilka in Iranian catches was ranked second after 

anchovy kilka in 1990-1991 at 6.84%, increasing to 

12.6% and 21.7% in 1997 and 1998 and then 

decreasing. The catch in 2008 was 490.8 t with 

maximum catch-per-unit-effort 0.19 t per vessel per 

night in February and minimum 0.03 t in January. 

Omega-3 fatty acids from fish oil of this species 

have been tested as a dietary supplement and were 



169 
 

Int. J. Aquat. Biol. (2017) 5(3): 128-192 

 

found to relieve symptoms of dysmenorrhoea 

(Moghadamnia et al., 2010). 

Conservation: Stocks in Iranian waters are said to be 

depleted. Kiabi et al. (1999) consider this species to 

be of least concern in the south Caspian Sea basin 

according to IUCN criteria. Criteria include 

commercial fishing, abundant in numbers, 

widespread range (75% of water bodies), absent in 

other water bodies in Iran, and absent outside the 

Caspian Sea basin.  

Sources: Iranian material: CMNFI 1993-0167, 1, 

93.0 mm standard length, Mazandaran, Caspian Sea 

(ca. 36º49'N, ca. 52º39'E); CMNFI 1993-0168, 2, 

91.8-94.0 mm standard length, Mazandaran, Caspian 

Sea (ca. 36º49'N, ca. 52º39'E).  

 

Genus Tenualosa Fowler, 1934  
This genus comprises 5 species found from the 

Indian Ocean to Indonesia and China. A single 

species enters rivers of southern Iran. The genus is 

defined by a series of characters listed below under 

Key characters: These fishes form part of local, 

artisanal fisheries throughout their range. 

  

Tenualosa ilisha (Hamilton, 1822)  
(Figs. 14, 15) 

Common names: Sobour, soboor, sobur, sabur, 

zobur, zabur, zamur or zomur, all variants of the 

same word), bari, barak; mahi-ye khor kuchiku (= 

small bone fish, at Abadan from www.abadan. 

com/abadanhistory.html, downloaded 15 March 

1998). [Zoboor, soboor, sbour in Arabic; hilsa, 

Indian shad or river shad; palo, palla or pulla and 

tikki-palwar in Pakistan]. 

Systematics: Clupanodon ilisha was originally 
described from the Ganges estuaries in India. 

Formerly placed in the genus Hilsa Regan, 1917. Al-
Hassan (1982), citing a personal communication 

from a Mr. Al-Abaychi in 1973, suggests that Shatt 

al Arab fish are distinct from those in Pakistan on 

morphometric and meristic grounds but no data have 

been published. Milton and Chenery (2001) used 

genetic and otolith chemistry data that provided 

strong evidence for a distinct stock in Kuwait, 

compared with stocks from India to Sumatra. Al-

Hassan (1999) mentions that people in Basrah can 

distinguish two kinds of sobur, based on taste. One 

is the tastier and pricier Shatt al-Arab form and the 

other is the less desirable estuarine/sea form. This 

has not been confirmed by systematic studies. Jorfi 

et al. (2008, 2009) found differences between 

populations in Iran and Iraq using molecular 

techniques. 

Key characters: This species is distinguished from 

other Indian Ocean clupeids by the upper jaw with a 

median notch, the anal fin ray count being less than 

Figure 14. Line drawing of Tenualosa ilisha. 



170 
 

Coad/ Herrings of Iran 

30 rays, a terminal mouth (lower jaw not prominent 

nor flared at the corners), scales in lateral series are 

not perforated posteriorly, last dorsal fin ray not 

filamentous, weakly developed lines (the fronto-

parietal striae) on top of the head (usually covered by 

skin and not visible), gill rakers on inner arches 

straight not curled, a long head 28-32% of standard 

length, and 30-33 ventral scutes forming a keel along 

the belly, 15-18 being prepelvic and 11-15 

postpelvic (Al-Nasiri and Al-Mukhtar, 1988a, 

1988b; Marammazi et al., 1995).  

Morphology: Dorsal fin with 4-5 unbranched rays 

followed by 14-16 branched rays, anal fin with 2-3 

unbranched rays followed by 16-20 branched rays, 

pectoral fin branched rays 12-15 and pelvic fin 

branched rays 7. Lateral series scales 44-51. Gill 

rakers are fine and numerous, up to about 275 on the 

lower arch.  

Iranian fish examined by Marammazi et al. (1995) 

from the Bahmanshir River in Khuzestan have 30-32 

total scutes along the belly, 16-18 prepelvic scutes, 

13-15 postpelvic scutes, 19-21 dorsal fin rays, 19-24 

anal fin rays, 13-15 pectoral fin rays, 8 pelvic fin rays 

and 44-51 scales. 

Sexual dimorphism: None reported.  

Colour: The back is grey-blue, bluish to green and 

the sides are silvery with golden, purplish or pink 

highlights. The dorsal fin is grey, the caudal fin grey-

blue with a silvery tinge and darkened margin, and 

the anal fin is light blue with some silvery tinges. 

Paired fins are hyaline. The area behind the gill cover 

in young fish and many adults have a dark blotch 

followed by a series of spots or blotches running 

along the upper flank, for a total of 6-7. The blotches 

may take the form of bars. The eye is yellow to red. 

Young have a bronze back, silvery flanks and a 

caudal fin margined in black.  

Size: Attains 60.6 cm total length and 2.49 kg for 

females and 43.0 cm and 0.68 kg for males. A sample 

of 233 moribund fish from the Ashar Canal, a branch 

of the Shatt al Arab, Iraq examined by Al-Nasiri and 

Al-Mukhtar (1988a, 1988b) had a total length range 

of 70-152 mm. Hussain et al. (1994) record fish 

migrating to the Shatt al Arab for breeding at 21-38 

cm for males and 33-43 cm for females. Mature 

females in the Shatt al Arab weighed about 0.5-1.1 

kg (Jabir and Faris, 1989). Fishes from Kuwait 

attained 57.0 cm (Al-Baz and Grove, 1995). Fishes 

from the Arvand, Bahmanshir, Karun and Dez rivers 

of Iran were 12-50 cm long (Marammazi et al., 1998; 

Ghafleh Marammazi et al., 2004). Nasri Tajan 

(2009) gave a maximum length and weight for 344 

fish from the Bushehr coast as 45.1 cm and 953 g. 

Distribution: Reportedly found from the Red Sea and 

Persian Gulf through the Indian subcontinent to the 

Malayan Archipelago in some general works, or 

more narrowly from the Persian Gulf to Myanmar. It 

enters the Shatt al Arab and Tigris River, once as far 

north as Baghdad (Kanazawa, 1955), but the 

northernmost distribution today in Iraq is the Hawr 

Figure 15. Tenualosa ilisha from Kuwait by J. E. Randall after FishBase. 



171 
 

Int. J. Aquat. Biol. (2017) 5(3): 128-192 

 
al Hammar. Before the construction of dams on the 

Euphrates the migration was up to Yaou and 

Meshkhab and up to Qal`at Salih (31°31'N, 47°16'E) 

in the Tigris of Iraq (van den Eelaart, 1954). 

The lower reaches of the Tigris and Euphrates 

rivers were connected by a channel to the Khor Al-

Zubair in Iraq during 1983. As a consequence the 

Khor became oligohaline (at less than 10‰) rather 

than hypersaline (at more than 40‰), becoming an 

estuary with heavy reed growth. The catch of sobour 

in the Khor by 1997 exceeded that in the Shatt al 

Arab and may involve diversion of stocks from the 

original habitat of the Shatt (Hussain, 1997).  

In Iran, it is recorded as far north as the Gargar 

Shoteit on the Dez River (Marammazi, 1994). 

Hussain, Jabir and Yousif (in litt. 1995) record this 

species from the Shatt al Arab in Iraq and the 

Bahmanshir, Hilleh, Jarrahi and Zohreh rivers in 

Iran. Marammazi (1994), Najafpour (1997), 

Marammazi et al. (1998) and Ghafleh Marammazi et 

al. (2004) report this species from the Arvand, 

Bahmanshir, Dez, Karun and Zohreh rivers. It may 

be found in the Hormuz basin but this has not been 

verified with specimens. Esmaeili et al. (2014) 

record it from the Shadegan Wetland and Esmaeili et 

al. (2015) from the Persis basin. Hashemi et al. 

(2014) caught a single specimen from the Shadegan 

Wetland in a year-long survey. Hashemi et al. (2015) 

listed this species as an exotic in the Shadegan 

Wetland but in the sense of a migrant from the sea. 

In the sea, they are found from Bushehr around to 

Kuwait in coastal waters (Blegvad and Løppenthin, 

1944; Hussain et al., in litt. 1995).  

Zoogeography: Al-Hassan (1982) mentions a study 

comparing a population of this species from Basrah, 

Iraq with one from Pakistan and finding significant 

meristic and morphometric differences, perhaps 

indicative of distinct stocks.  

Habitat: Sobour enter the Shatt al Arab in February 

and March during high tides and feed there until the 

fall according to a study by Al-Nasiri and Al-

Mukhtar (1988a, 1988b) working on fish taken from 

the Ashar Canal, Basrah, Iraq. van den Eelaart 

(1954) reported that most fish entered the Shatt al 

Arab in April during the last and first phase of the 

moon and anecdotal reports indicate the end of 

March to be the peak period of entry. They ascended 

into the Hawr al Hammar and from there into the 

Euphrates as well as into the Tigris (van den Eelaart 

1954). Significant numbers were recording as 

entering the recovering Hawr al Hammar in 2005-

2006 (Hussain et al., 2006). Small specimens (50-

100 mm) were observed in the east Hawr al Hammar 

in June 2005 and July 2006 (www.iraqmarshes.org, 

downloaded 29 August 2005, N.A. Hussain in litt. 

2006). In mid-April sobour were found below the 

Yaou and Meshkhab regulators which formed the 

limit of their migration on the Euphrates in the early 

1950s. The limit in the Tigris was beyond Amara. 

The main spawning grounds in the Euphrates were 

probably somewhere between Shinafiya and 

Samawa and in the Tigris between Amara and Qalat 

Saleh. 

The last ones leave the Shatt in July and fry are 

found in the rivers of Iraq at the end of the June. 

Hussain et al. (1994) recorded sobour ascending the 

Shatt al Arab during March with a continuing 

migration upstream through April to July for 

spawning and a return migration to the sea during 

August to October. Al-Hassan (1993) notes that local 

people believe that sobour ascend the Shatt al Arab 

during spring to marshes north of Basrah for 

spawning, suggesting that they are the fluvial 

anadromous type. Al-Hassan (1999) considers they 

migrate to the sea in September-November, when 

they are landed in Kuwait, and they then migrate to 

the Iranian coast during December-January. Males 

and females move upriver in separate groups 

according to Iraqi fishermen (Al-Hassan, 1999). 

Jorfi et al. (2008) suggest, based on molecular 

studies, that a population in the Persian Gulf chooses 

the Karun River for spawning and migrates via the 

Bahmanshir River, while others migrate up the 

Tigris and Euphrates rivers in Iraq via both the 

Bahmanshir and the Arvand rivers. 

Blegvad and Løppenthin (1944) mention this 

species on sale at Khorramshahr on 28-29 April. The 

spawning migration in Iran occurs in spring 



172 
 

Coad/ Herrings of Iran 

(Sharifpour, in litt. 1991). It is only found in the 

Zohreh River in spring and summer (Marammazi, 

1994). Ghobeishavi et al. (2016) found a depression 

is some haematological and immunological 

parameters during the upstream migration in the 

Karun River, possibly a result of hormonal and 

environmental effects. Dastan et al. (2017) described 

gill and kidney tissue changes on the migration into 

the Bahmanshir and Karun rivers, chloride cells 

decreasing in number and area for example. 

They may be found in deep water, over 18 m, or 

in shallows, on their spawning migration. Large 

concentrations of sobour occur below dams blocking 

their migration. Young occur in side branches of the 

Shatt al Arab near food, shelter and the spawning 

grounds (Hussain et al., in litt. 1995).  

This species occurs in river estuaries and coastal 

waters and appears to be restricted to the northern 

end of the Persian Gulf because this is the only part 

with large spawning rivers (Hussain et al., in litt. 

1995). These authors also suggest that an 

anadromous stock from the Shatt al Arab migrates to 

warmer waters off Bushehr during January, February 

and March. At the same time there is a winter decline 

of Kuwaiti stocks. There may also be a marine stock 

inhabiting coastal waters of Kuwait since larvae have 

been found in Kuwait Bay during June and 

November and catches are made in the Bay year 

round.  

Hussain (1997) notes the changing conditions in 

the Khawr az Zubayr, which became oligohaline 

from hypersaline after it was connected to the Tigris-

Euphrates basin by the Shatt al Basrah Canal. In 

1994, fishermen began catching sobour in the Khawr 

az Zubayr and by 1997 the numbers caught exceeded 

the catch in the Shatt al Arab.  

Migrations in the Indus River of Pakistan (Islam 

and Talbot, 1968) may last over 7 months and the 

migration up the Ganges River extends over 1,287 

km. Fish may move as much as 70.8 km in one day 

and may jump out of the water on the migration.  

Age and growth: In the Bahmanshir River, Iran most 

fish are 4-5 years old. The minimum total length and 

age at maturity are 26.2 cm, 200 g and 2 years for 

males and 32.18 cm, 450 g and 3 years for females. 

von Bertalanffy growth parameters in Iranian 

females are L∞=57.78 cm and K=0.282 and in males 

46.37 cm and 0.252 (Marammazi, 1995; Iranian 

Fisheries Research and Training Organization 

Newsletter, 12: 5, 1996, Annual Report, 1995-1996; 
Iranian Fisheries Research and Training 

Organization, p. 53-54, 1997). Nasri Tajan (2009) 

found Bushehr coast fish were 2-4 years old with 

most 2+ years. 

Hashemi et al. (2009) studied fish landed at 

Hendijan and Abadan and recorded L∞ as 42.81 cm, 

K was 0.9, M was 1.37, F was 2.41, Z was 3.78 and 

E was 0.64. Y'/R was 0.048 and B'/R was 0.19, 

exploitation rate (U) was 0.61, annual stock at the 

beginning of the year (P) was 7,615 t, annual 

standing stock (b) was 1,927 t and MSY was 3,642 t. 

The stock was overfished. Hashemi et al. (2009), 

also studying fish at Hendijan and Abadan, recorded 

L∞ as 54.6 cm, K was 0.96, M was 1.34, F was 2.8, 

Z was 4.2, E was 0.68 and t0 was -0.14, and the stock 

was overfished. Hashemi et al. (2010) studied 9,317 

fish from the landings at Abadan and Hendijan. Size 

range was 20-39 cm. The von Bertalanffy growth 

parameters were L∞=43.32 cm, K=0.78yr
-1, Φ' was 

3.16 and t0 was -0.18. Mortality rates were M=1.29 

and Z=4.53, and fishing mortality (F) was 3.24yr-1. 

The exploitation rate (E) was 0.72 and the stock was 

overfished. Values of the sizes where the probability 

of capture was 50% (L50) and 100% (L100) were 22.3 
and 28.5 cm total length respectively. Fish were 

recruited to the fishery at a mean size of L100=22.3 
cm. The relative yield per recruit (Y'/R) was 0.062, 

relative biomass per recruit (B'/R) was 0.12 and 

exploitation rate (U) was 0.76. The values for annual 

catch, total annual stock, standing stock and 

maximum sustainable yield were 4,645t, 6,635.71 t, 

1,433.64 t and 3,274.19 t, respectively. The fishing 

pressure must be reduced from 3.24yr-1 to about 

0.97yr-1for this population to be adequately 

managed. Another study apparently based on the 

same or similar samples (Hashemi Seyed et al., 

2010) found slightly different parameters: L∞=42.81 

cm, K=0.0yr-1, t0=-0.25, Z=3.78, M=1.37, F=2.41, 



173 
 

Int. J. Aquat. Biol. (2017) 5(3): 128-192 

 
E=0.64 and the values for annual catch, annual 

average standing stock and maximum sustainable 

yield were 7,615 t, 1,927 t, and 3,624 t, respectively. 

Hashemi et al. (2010) assessed production in the 

Shadegan Wetland, Khuzestan as 0.73 kg/ha/year. 

Roomiani and Jamili (2011) examined fish landed 

in Iran from a northern Persian Gulf fishery. Growth 

was isometric. Maximum total length was 43 cm and 

weight 949 g. von Bertalanffy growth parameters 

were L∞=42.74 cm total length, K=0.77 and t0=-0.21 

years-1. Total mortality (Z) was 2.55 years-1, natural 

mortality (M) was 0.75 years-1, fishing mortality (F) 

was 1.8 years-1, and exploitation rate (E) was 0.7 

years-1, and parameters indicate overfishing. 

Maximum sustainable yield was calculated to be 

2,653t. Roomiani et al. (2012), presumably working 

with a partially similar dataset, found L∞=42.8 cm, 

K=0.7, Z=3.45, M=1.24, F=2.21 and E=0.64.  

Al-Nasiri and Al-Mukhtar (1988a, 1988b) give a 

length-weight relationship of W=3.9 x 10-6 L3.16 or 

logW=3.16 logL-5.4 for fish aged at 0+ from the 

Ashar Canal at Basrah. The mean condition factor 

was 0.87. Fishes in the Shatt al Arab are in age 

groups 5 to 6 for the period May to August (Hussain 

et al., 1991). In contrast, a later study on the Shatt al 

Arab fish showed there are 5 age groups and the 

second and third age groups dominate in catches 

(Hussain et al., 1994). In this latter study, Shatt al 

Arab fish mature at 25 cm for males and 33 cm for 

females, similar to an Iranian study (see below). The 

length-weight relationship was logW=-4.7074+ 

3.0479 logL for females and logW=-4.5802+3.0193 

logL. Condition factor gradually increased with 

length groups in males, peaking at 32-33 cm 

followed by a sharp decline while females had a 

nearly stable condition factor from 34 to 43cm. 

Mohammed et al. (2001) gave a von Bertalanffy 

growth equation as L∞=60.47 cm and a condition 

factor of 0.32, slower growth than in Indian and 

Bangladesh populations and probably maturing later. 

Amodeo (1956) gives lengths of 25 to 35 cm for 

fish caught in the Shatt al Arab on their spawning 

migration. Young grow rapidly, 4.3 cm in October-

November. Most fish on the migration in the Indus 

River were in age groups 3 and 4. Life span is up to 

an estimated 7 years with maturity as early as 1 year. 

Jawad et al. (2004) found haematocrit level to 

increase with body length up to 40 cm after which it 

decreased, males showed higher levels than females, 

and levels were higher pre-spawning than during 

spawning and increased slightly post-spawning, a 

general correlation with fish activity in Iraqi waters. 

Al-Baz and Grove (1995) studied fish taken from 

Kuwait fish markets. Females dominated the catch, 

male:female ratio being 1:2.4, perhaps because the 

sexes moved in different schools. The smallest 

mature female was 34.4 cm and 50% of the females 

are mature at 41.5 cm. They estimated natural 

mortality (M) based on von Bertalanffy growth 

parameters (L∞ and K) and mean annual water 

temperature as logM=-0.0066 -0.279 logL∞+0.6543 

logK + 0.4634 logT. The length-weight relationship 

was W=0.011L2.983 for males and W=0.007L3.104 for 

females. Growth in the sexes follows different 

patterns. Five age groups were detected using 

otoliths and fish were fully recruited to the fishery at 

3 years of age. von Bertalanffy growth parameters 

were L∞=52.70 cm and condition factor (K)=0.28 

per year while using Allen's method they were 

L∞=52.50 cm and condition factor (K)=0.36 per 

year. Annual total mortality was estimated to be 1.2 

using the K value of 0.36. A fishing mortality was 

calculated to be 0.8 per year.  

Mohamed et al. (2001) studied 7,535 fish from 

landings at Fao, Iraq in 1998-1999 and found growth 

and mortality parameters were L∞=60.47 cm, 

K=0.32, Z=1.28, M=0.62 and F=0.66. The 

exploitation rate (E) was 0.52, a bimodal recruitment 

pattern of unequal strength was observed (peaks in 

March at 54.65% and October at 45.35%), maximum 

yield per recruitment was achieved at Emax=0.63 and 

Lc=25.0 cm. They recommended fishing nets should 

be adjusted to capture fish over 25 cm to avoid 

overfishing. Mohamed and Qasim (2014a) assessed 

the stock in 2012-2013 in Iraqi marine waters close 

to Iran. They found fish were 12.2-48.0 cm total 

length with >93% of fish 26-44 cm. Age span was 

almost 6 years. The b value was 3.2683 (allometric 



174 
 

Coad/ Herrings of Iran 

growth), asymptotic growth (L∞)=61.47 cm, growth 

rate (K)=0.275, annual instantaneous rate of total 

mortality (Z)=1.66, natural mortality (M)=0.55, 

fishing mortality (F)=1.11 and exploitation rate 

(E)=0.67. A bimodal pattern of unequal recruitment 

was observed with April fish being migrant breeders 

and July fish juveniles from the previous year 

recruiting to the adult stock. The maximum yield per 

recruitment was achieved at Emax=0.72 and Lc=27.8 

cm. The stock was overexploited. 

Food: The Ashar Canal study found them to feed on 

phytoplankton such as dinoflagellates and diatoms 

and on zooplankton, mainly copepods, as well as 

their own young. The sieve-like gill rakers are used 

to strain out planktonic organisms without selection. 

Presence of some sand grains indicates that feeding 

can occur on the river bed. Feeding intensity may 

decrease or cease on the spawning migration and is 

very high after spawning. The Bahmanshir fish feed 

principally on copepods and diatoms. Shatt al Arab 

juveniles feed mostly on filamentous algae and 

diatoms with some organic matter, fish eggs and 

zooplankton while adults have empty stomachs on 

the spawning migration (Hussain et al., in litt. 1995). 

In the Indus River, the newly hatched larvae and 

juveniles graze for five to six months in fresh waters 

before they migrate to the sea (www.jang-

group.com/thenews/feb2003-daily/18-02-2003/busi 

ness/b2.htm, downloaded 18 February 2003). The 

prime food in the sea off the Iranian coast at Deylam, 

Bushehr is phytoplankton, principally Bacilliario-

phyta followed by Pyrrophyta. Zooplankton was also 

taken (Nasri Tajan et al., 2008; Nasri Tajan, 2009). 

Reproduction: The spawning migration depends on 

the flood regime of the rivers. Turbid water and fast 

current are probably stimulants to egg deposition. 

The sobour depends on river-edge vegetation for egg 

deposition. Spawning grounds in Iraq are probably 

located near the beginning of the side branches of the 

northern sector of the Shatt al Arab, 120 km from the 

sea (Hussain et al., 1994). This species is 

gonochoristic (Blaber et al., 1997). Males may 

ascend the river before females but females become 

dominant in Indian populations. Males dominate in 

March in the Shatt al Arab and the sex ratio reaches 

equilibrium in the spawning months of May-July 

(elsewhere in the same communication spawning is 

given as June to August) (Hussain et al., 1994; Jawad 

et al., 2004). Spawning may occur more than once in 
a season in India. This has not been demonstrated for 

Iran but could occur. The gonadosomatic index for 

fishes in the Iraqi Shatt al Arab indicates peaks in 

March-May and July-August, suggesting two 

spawnings (Hussain et al., 1991) although a later 

report (Hussain et al., 1994) gives spawning as June 

to July and July to August as evidenced by two 

modes of juveniles found in September. Sex ratio is 

equal during this period. All females entering the 

Shatt al Arab were mature with smallest female 

being 33.0cm long. Males less than 25.0 cm were 

immature, the population reaching 100% maturity at 

31-32cm (Hussain et al., 1994). The Kuwait fish 

studied by Al-Baz and Grove (1995) indicated 

spawning between May and July with a peak in June. 

Fecundity in the Indus River population was 

estimated to be up to 2,917,000 eggs per female, egg 

diameters reached 0.89mm, and the hatching takes 

place in about 23 to 26 hours (www.jang-

group.com/thenews/feb2003-daily/18-02-2003/busi 

ness/b2.htm, downloaded 18 February 2003). 

Estimates for the Hooghly River of India reach 

13,230,500 eggs per female (Al-Hassan, 1993). 

Fecundity in the Shatt al Arab ranges between 

444,960 and 1,616,560 eggs for fish 33.0-41.5 cm 

total length although 2 fish 37.3 and 2 fish 39.0 cm 

total length had a range in egg numbers of 109,000-

233,840, showing that great variations in fecundity 

occur between individuals; possibly some fish had 

partially spawned before capture (Jabir and Faris, 

1989). This latter study gave a relationship between 

absolute fecundity and total length as F=1.3699 

L3.6681 and logF=0.1367 + 3.6681 logL and between 

fecundity and weight F=302.8214 W1.2087 and 

logF=2.4812 + 1.2087 logW. Fecundity increased 

significantly with body weight, ovary weight and 

total length. Relative fecundity (ova/gramme body 

weight) varied from 737 to 1,721, mean 1,216. 

Hatching can occur within one day at an average 



175 
 

Int. J. Aquat. Biol. (2017) 5(3): 128-192 

 
temperature of 23°C. Eggs, larvae and young are 

found on the spawning grounds but with growth the 

young move into estuarine and foreshore areas 

during winter months. Hussain et al. (1994) record 

the appearance of juveniles from the northern Shatt 

al Arab from June to November. Adults return to 

their original habitat in the sea after spawning. There 

is some evidence for freshwater resident populations 

in India which migrate upriver to spawn but do not 

descend to the sea.  

The Bahmanshir fish are thought to spawn from 

April to July. Only adults enter the Bahmanshir 

(Iranian Fisheries Research and Training 

Organization Newsletter, 12: 5, 1996). Absolute 

fecundity of fish from the Arvand, Bahmanshir, Dez 

and Karun rivers ranges from 374,892 to 1,954,144 

eggs for total lengths of 380 to 500 mm, respectively 

and is related to age. Ova with diameters 0.64-0.795 

mm were released spontaneously in a study of this 

fish in Khuzestan Province, in several batches along 

its migration route (Ghafleh Marammazi et al., 

2004). Spawning begins on entry to the Bahmanshir 

and Arvand rivers in Khuzestan in April, continuing 

to September and the end of their migration at the 

cities of Shushtar and Dezful higher upriver. Males 

enter these rivers first in March, followed by females 

in April (Ghafleh Marammazi et al., 2004). Nasri 

Tajan (2009) found Bushehr coast fish were batch 

spawners in April-May. 

Parasites and predators: None reported from Iran 

other than nematode larvae by Ebrahimzadeh and 

Nabawi (1975) for fish from the Karun River. Bannai 

and Muhammad (2016) studied the parasite fauna of 

this shad as it immigrated to Iraqi waters from Fao in 

the estuary of the Shatt al Arab to the Al Hammar 

Marshes. They concluded that these fish do not come 

from the Indian Ocean and are endemic to the 

Persian Gulf (unless parasites are lost on the 

migration from the Indian Ocean). 

Economic importance: The Ashar Canal study cites 

996,308 kg reaching the Ashar fish market from 

October 1975 to June 1977 (see also Sharma (1980)). 

The catch landed at Fao on the Shatt al Arab estuary 

of Iraq was 6,576 t in 1990-1991 (L.A.J. Al-Hassan 

in litt. 1995; however this seems much too high 

although the estimate is from the Food and 

Agriculture Organization). This species forms the 

most important commercial fishery in the Basrah 

region of southern Iraq, average catches being 

491.086, 319.661 and 267.988 t in 1977, 1978 and 

1979 respectively (sic, Jabir and Faris, 1989). There 
is a drift-net and stake-net ("hadra") fishery in the sea 

by Kuwait in Kuwait Bay and around Falaikah Island 

(Al-Baz and Grove, 1995).  

The fishing season on the Tigris-Euphrates is 

March to August with a peak in April, or late April 

to early June (Jabir and Faris, 1989) or to November 

(Ali et al., 1998). van den Eelaart (1954) gave the 

fishing season for this species as March-August 

(peaking in April) in rivers, and March-May 

(peaking in April) in Hawr al Hammar, Iraq. Fish are 

caught at the mouth of the Shatt al Arab as they enter 

the river with stationary gill nets, drifting gill nets, 

and in "mailan" and "odda" traps from March to 

August. The catch averaged 150-180 kg per ten odda 

and in March 1953 the total catch at the mouth of the 

Shatt al Arab was about 25,000 kg (Amodeo, 1956). 

Large fish are only caught in the summer (Al-

Hassan, 1999).  

The catch at Abadan in Iran from February to 

November in 1943 was about 401.42 t and from 

January to June about 336.67 t (Pillay and Rosa, 

1963). This species is seen on markets at Ahvaz, 

Khuzestan in November but these are sea-caught 

fish. Marjan Iran Company was selling 600-800 g 

fish for U.S.$1.40/kg, 800-1,000 g fish for 

U.S.$1.60/kg, 1,000-1,200 g fish for U.S.$1.70/kg, 

and 1,200 g and larger fish for U.S.$1.80/kg in 

August 2003 (http://groups.yahoo.com/groups 

/hilsa/message/25). The catch in Khuzestan Province 

in 2000 was 2,688 t (Ghafleh Marammazi et al., 

2004) and in 2006 was 4,989.83 t (about 15% of 

Khuzestan's total commercial fish landing) 

(Roomiani and Jamili, 2011). The catch in 

Khuzestan Province in 2008 was 4,645 t (Hashemi et 

al., 2010). 

These fish are caught with traps, weirs, gill nets 

and other devices in rivers on the spawning 



176 
 

Coad/ Herrings of Iran 

migration. They are excellent eating until spawning 

occurs after which they lose their flavour. However 

this species has been implicated in clupeotoxic 

poisoning. Hindi et al. (1996a) give the chemical 

composition of flesh of this species as 66.41% 

moisture, 12.12% fat, 18.72% protein and 1.98% 

ash, indicating a valuable food fish characterised as 

fatty. Hindi et al. (1996b) give chemical indices for 

assessing fish freshness according to the month of 

capture and marketing (pH 6.06, total volatile 

nitrogen bases 15.32 mgN/100g fish, thiobarbituric 

acid 1.35 mg, and free fatty acids 1.33%).  

Biogenic and anthropogenic sources were noted 

for the hydrocarbons in this species from the Shatt al 

Arab; n-alkanes attained 31.11 µg/g and 

hydrocarbons 10.91 µg/g, the highest for the fish 

species studied (Al-Saad et al., 1997). The fat 

content of this shad is a factor in these high levels 

(Al-Saad, 1990). Salari and Sadough (2009) 

compared heavy metal (Cd, Pb, Cu, Co, Ni) content 

in muscle, liver and gill tissues of fish from the 

Karun River and found levels less than those 

considered dangerous in Iran. Sadough Niri et al. 

(2010) examined fish from the Arvand River and 

found some of the above listed heavy metals 

exceeded international limits however. 

In Pakistan, the Indus River fishermen number 

between 8,000 and 9,000. Jafri (1994) reviews the 

Indus fishery which had yields up to 2,694mt. It is 

the most important Indo-Pacific shad species. The 

failure of the Indus River fishery in 2003 through 

drought resulted in Iranian fish being flown to 

Pakistan for marketing there at rupees150-400 per 

piece (www.jang-group.com/thenews/feb2003-daily 

/18-02-2003/business/b2.htm, downloaded 18 

February 2003). Robins et al. (1991) list this species 

as important to North Americans. Importance is 

based on its use as food, in aquaculture and in 

textbooks.  

Conservation: Hussain et al. (in litt. 1995) report a 

decline in catches over the previous two decades in 

the Shatt al Arab. Al-Nasiri and Al-Mukhtar (1988a, 

1988b) mention that fish enter the polluted Ashar 

Canal, a side tributary of the Iraqi Shatt al Arab, 

during high tide when waters are diluted. A low tide 

in October resulted in severe oxygen depletion and 

fish suffocated. Das et al. (1977) found samples from 

the Ashar fish market in Basrah to be contaminated 

with hydrocarbons, emitting a kerosene smell and 

being unfit for human consumption. Al-Saad (1990) 

found petroleum hydrocarbon residues to be high in 

Khawr az Zubayr fish at 40.6 μg/g as this species is 

one that accumulates fat. Mohamed and Qasim 

(2014b) found Iraqi landings decreased from 90.2% 

of total landings in 1965-1973, to 52.9% in 1991-

1994, to 41.8% in 1995-1999 and to 30.7% in 2000-

2006. They recommended establishment of closed 

areas to protect spawning such as the Shatt al Arab 

for 30 days in May, catch of small fish up to 23 cm 

in spawning areas banned, reduction of pollution, 

and cooperation within Iraq and with Iran on 

fisheries management. 

Evidently, overfishing and pollution are major 

factors in the conservation of this species, to which 

must be added variations in freshwater flow and 

quality from the marshes and Tigris-Euphrates 

through human processes.  

Sources: Some aspects of the biology of this species 

were based on Pillay and Rosa (1963) and Al-Hassan 

(1993) writing mostly on Indian and Pakistani 

populations. Bhaumik (2015) also reviews studies on 

the biology of this species. Specimens on markets in 

Ahvaz, Khuzestan examined.  

Iranian material: CMNFI 1991-0153, 1, 243.3 mm 

standard length, Khuzestan, Zohreh River (no other 

locality data).  

Comparative material: BM(NH) 1875.1.14:11-13, 3, 

118.8-135.8 mm standard length, Iraq, Tigris River 

(no other locality data); BM(NH) 1920.3.3:178-182, 

6, 103.3-132.4 mm standard length, Iraq, Basra 

(30º30'N, 47º47'E); BM(NH) 1989.1.13:1-3, 3, 53.9-

59.9 mm standard length, Iraq, Khawr az Zubayr (no 

other locality data); BM(NH) 1989.1.13:4-5, 2, 66.6-

69.8 mm standard length, Iraq, Khawr az Zubayr (no 

other locality data). 

 

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Int. J. Aquat. Biol. (2017) 5(3): 128-192 

E-ISSN: 2322-5270; P-ISSN: 2383-0956

Journal homepage: www.ij-aquaticbiology.com 

© 2017 Iranian Society of Ichthyology 

 چکیده فارسی

 

  ماهیان(مروری بر شگ ماهیان ایران )خانواده شگ
 

 برایان دبلیو کد

 .کانادا ، K1P 6P4،انتاریو ،اتاوا، موزه تاریخ طبیعی کانادا
 

  چکیده:

 هاشود، گونهتوصیف می ماهیان ایرانشناسی، اهمیت اقتصادی و وضعیت حفاظتی شگزیست شناسی، پراکنش،در این مقاله سیستماتیک، ریخت

، Clupeonellaهای سز جنبومی اگونه  9گردد. در مجموع تعداد مورد بررسی قرار گرفته و یک مرور منابع در مورد این ماهیان در ایران فراهم می

Alosa  وTenualosa  شود.یافت میهای جنوب ایران در دریای خزر و رودخانه  

 .زالون ،کیلکا ،Alosa، Clupeonella، Tenualosa شناسی، شناسی، زیستریخت :کلمات کلیدی