Review Article Review of the Herrings of Iran (Family Clupeidae)
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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. Abstract: The systematics, morphology, distribution, biology, economic importance and Article history: Received 4 March 2017 conservation of the herrings (kilkas and shads) of Iran are described, the species are illustrated, and Accepted 5 May 2017 a bibliography on these fishes in Iran is provided. There are 9 native species in the genera Available online 25 June 2017 Clupeonella , Alosa and Tenualosa in the Caspian Sea and rivers of southern Iran. Keywords: Morphology, Biology, Alosa, Clupeonella, Tenualosa, Kilka, Shad. Introduction family in the Caspian Sea is seen in the number of The freshwater ichthyofauna of Iran comprises a subspecies which have been described, rather than in diverse set of families and species. These form genera. At the species level these are Caspian Sea important elements of the aquatic ecosystem and a endemics. A study by Pourrafei et al. (2016) based number of species are of commercial or other on the nuclear gene RAG1 did not support the significance. The literature on these fishes is widely monophyly of Clupeidae but, as an abstract, details scattered, both in time and place. Summaries of the are lacking. These fishes are dealt with as a single morphology and biology of these species were given family here. in a website (www.briancoad.com) which is updated Curiously, the species and subspecies in the here for one family, while the relevant section of that Caspian Sea are generally of larger size than their website is now closed down. Other families will also relatives in the Black Sea basin. These observations being addressed in a similar fashion. are attributed to the variable environment in the Caspian Sea over time, with repeated changes in Family Clupeidae salinity and temperature which the fish could not Herrings, shads, sardines, sprats, pilchards and avoid. Black, Mediterranean and Atlantic species menhadens are moderate-sized fishes, usually less lived under more stable conditions and could, in any than 25 cm long, found in warmer marine waters case, retreat from lowered temperatures for example. with some species anadromous or permanent In addition, the Caspian Sea clupeids lacked the freshwater residents. There are about 64 genera and competitors which entered the Black Sea from the about 218 species world-wide (Nelson et al., 2016), Mediterranean and Atlantic and some (Clupeonella with 8 species in the Caspian Sea and one commonly spp., Alosa caspia) could occupy the pelagic, found in Persian Gulf drainages. Some other, planktivore niche taken up by other species in the primarily marine, species are known to enter rivers Black Sea. There are no other pelagic fish but these in southern Iran (e.g. Nematalosa nasus (Bloch, herrings in the stable salinity areas of the Caspian 1795) and Sardinella sindensis (Day, 1878)) but are Sea. not dealt with here (Hashemi and Ansary, 2012; These fishes usually have modified scales on the Jouladeh-Roudbar et al., 2015). The diversity of this belly forming abdominal scutes with a saw-like * Corresponding author: Brian W. Coad E-mail address: [email protected] Int. J. Aquat. Biol. (2017) 5(3): 128-192 129 edge. Most species have two, long, rod-like was 117,300 tonnes in 1995 (Bartley and Rana, postcleithra. The lateral line is usually absent or on 1998). These fish are used in a high value form as only a few scales. Silvery cycloid scales are easily frozen whole consumer packs, as fish meal for detached and are found only on the body. The mouth poultry and in aquaculture, and in canning (Food and is usually terminal with jaws about equal in length. Agriculture Organization, Fisheries Department Teeth are small or absent but gill rakers are long and 1996). numerous for sieving plankton. Fins lack spines and Catches in the Caspian Sea for 1991 and 1992 there are no barbels. There is no adipose fin. The were respectively 13,817 and 21,527 tonnes of kilka pectoral and pelvic fins have a large axillary scale. (Clupeonella spp.) compared with 3,036 tonnes and The caudal fin is deeply forked. The eye is partly 2,692 tonnes of sturgeons and 18,571 and 16,873 covered by an adipose eyelid. The flesh is tonnes of bony fishes. The catch reached 51,000 particularly oily and is highly nutritional. tonnes in 1994 from none 10 years previously (Food Members of this family often form immense and Agriculture Organization, Fisheries Department, schools in surface waters of the ocean and the 1996). The kilka catch in Gilan Province in 1997 was Caspian Sea where they feed on plankton. Schooling 36,077 tonnes (total catch 39,154 tonnes for the is an anti-predator device making it difficult for a province) and in Mazandaran in 1998 was 31,583 predator to pick out an individual from a tight mass tonnes (total catch 48,027 tonnes). of fish. There is also a "sentry effect" where The Caspian Environment Programme (1998) awareness is increased by the presence of many fish. gives the following catches in tonnes for the Iranian The school is maintained by a balance between Caspian Sea for kilka (Clupeonella spp.) and for visual attraction and lateral line stimulus repulsion. Alosa pontica (= kessleri), respectively as 1,013 and Herring can feed on the smaller plankton, less than 2 (1973), 1,170 and 2 (1974), 1,286 and 4.5 (1975), 300-400 µm, at night by filter-feeding but during the 900 and 5.5 (1976), 1,261 and 2 (1977), 771 and 0 day can also use particulate feeding. In the latter, (1978), 836 and 0 (1979), 619 and 0.1 (1980), 1,341 they select larger plankton using the area temporalis, and 0.4 (1981), 798 and 10.4 (1982), 621 and 1.6 a specialised ventro-posterior region of the retina (1983), 1,517 and 20.3 (1984), 1,828 and 34.8 which improves vision as herring approach food (1985), 2,450 and 71.9 (1986), 4,389 and 13 (1987), items from slightly below. 4,700 and 16 (1988), 7,902 and 30 (1989), 8,814 and Herring are easily caught and are extremely 30 (1990), 13,817 and 35 (1991), 21,527 and 35 valuable to commercial fisheries. They are the most (1992), 28,730 and 893 (1993), 51,000 and 720 important fishes economically, both as food for man (1994), 41,000 and 490 (1995) and 57,000 and 330 and also for many other commercial fish species. (1996). Abdolmalaki and Psuty (2007) give figures Wars have been fought over fisheries for herrings. In over a wide range of years for Iranian coastal catches one year, members of the herring family made up in the southern Caspian Sea with selected 37.3% of all fish caught in the world. Some are used comparative species as presented in Table 1. for fish meal, as fertiliser and as an oil source. The Overfishing of clupeids in the Caspian Sea and 1994-1995 catch of clupeids in the Iranian Caspian stock collapses caused by the invasive ctenophore was 98.3 tonnes by beach seine and 671.5 tonnes by Mnemiopsis leidyi have also had deleterious effects gill nets, a decrease of 200 tonnes in total over the on the Caspian seal (Pusa caspica) which fed on previous year's catch (Iranian Fisheries Research and these fishes (Harkonen et al., 2012). Training Organization Newsletter 10: 4-5, 1995) (but A major source for the biology and systematics of see below where the catch is much higher – sources Caspian clupeids remains Svetovidov (1952), now tend to vary). The Caspian Sea shads accounted for inevitably dated but not yet updated. Whitehead about 35% of total inland production in Iran which (1985), Hoestlandt (1991), Coad (1997) and Coad et 130 Coad/ Herrings of Iran Table 1. Iranian coastal catches in the southern Caspian Sea with selected comparative species (Abdolmalaki and Psuty, 2007) 1927- 1937- 1947- 1957- 1967- 1977- 1987- 1997- Catch and frequency 1936 1946 1956 1966 1976 1986 1996 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 al. (2003) gave brief overviews of Iranian herrings. short; pectoral fins long……………………………6 There has been no recent, careful systematic and 5b. Body not deep and not compressed; head not taxonomic study of these species in the Caspian Sea large and deep, not wedge-shaped in anterior view; basin as a whole and extensive new material was not caudal peduncle not short; pectoral fins short…….8 available for examination here. 6a. Gill rakers on first arch 50 or more, thin and long, Key to Clupeidae: Caspian Sea species have much longer than gill filaments; teeth weakly numerous nominal subspecies and keys to these may developed…...…………………………Alosa caspia be found in Berg (1948-1949) and Svetovidov 6b. Gill rakers on first arch 48 or less, shorter, equal (1952). to or somewhat longer than gill filaments; teeth well 1a.